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⎪10⎪
Advances in Crop Research
edited by Jürgen Kroschel
Integrated Pest Management for the

Potato Tuber Moth,
Phthorimaea operculella (Zeller)
- A Potato Pest of Global Importance
Jürgen Kroschel and

Lawrence A. Lacey
Result of a symposium held in Boise, Idaho on August 21, 2006 as part of

the Sixth World Potato Congress jointly organized by USDA-ARS,
Wapato, WA, USA and The International Potato Center, Lima, Peru
I
Preface
Within the series “Tropical Agriculture” and its subseries “Tropical
Agriculture – Advances in Crop Research”, this is the fourth publication
related to the potato tuber moth, Phthorimaea operculella. The previous
publications focused on the development of a country-specific integrated pest
management program (see Tropical Agriculture No. 8), on pest population
dynamics studies and management issues in different agroecological zones (see
No. 11), and on the improved use of microbial insecticides for the potato tuber
moth (see No. 13).
The potato tuber moth, which originated in tropical mountainous regions of South
America, is the most economically important pest of potato in developing
countries and occasionally it also affects other Solanaceae crops like tobacco,
tomato or eggplant. Today, it has achieved a worldwide distribution and is a pest
reported from more than 90 countries including countries in temperate regions. In
warm and dry areas, potato tuber moth infestations and losses are especially
severe and it is predicted that the moth will further expand its current distribution
and pest severity under climate change. Hence, effective tools for forecasting its
future spread and for managing the pest in potato fields and stores with less
negative impacts of pesticides on humans and the environment are urgently
needed and implemented.
The present publication is the result of a symposium on “Integrated Pest

Management for the Potato Tuber Moth - A Potato Pest of Global
Proportions”, which was held as part of the 6th World Potato Congress, 20-24
August 2006, Boise, Idaho. The principal objective of this symposium was to
bring together scientists from all over the world working on potato tuber moth and
to share and summarize the “state-of-science” on the potato tuber moth problem
and its management as part of an integrated pest management program. The book
is divided in three chapters: (i) Pest status and forecasting, (ii) Biocontrol and
use of biorationals, and (iii) Integrated Pest Management.
The first book chapter describes in two papers the invasion of the potato tuber
moth into the Pacific Northwest and new modeling tools linked with GIS to
forecast regional and seasonal distribution of the potato tuber moth. We are proud
of providing along with this book publication a software package (called Insect
Life Cycle Modeling, ILCYM) that facilitates the development of pest insect
phenology models and allows for pest risk mapping of a specific pest. An alpha
version of this computer program, which includes the potato tuber moth model, is
enclosed to this book. The second chapter provides an overview on the state-of-
the-art of microbial control and the limitations and possibilities of using the potato
tuber moth granulovirus (PoGV) for field applications. Classical biological
control of the potato tuber moth in New Zealand clearly shows the need for the
implementation of an integrated system to make naturalized parasitoids fully
effective. Further, the biopesticide Spinosad and attracticides are proposed as
II Preface
promising new tools for potato tuber moth management. Chapter three focus on
the use of different control practices in an integrated approach highlighting the
importance of cultural and biological management practices, and the need for an
integration of several pest issues in potato to achieve adoption of integrated pest
management by farmers. Finally, potato tuber moth resistant potatoes may play an
increasing role for farmers in developing countries in the future; an approach to
commercialize resistant potato is described for South Africa.
It is hoped that the book will benefit researchers and extension workers of
international and national agricultural research and extension institutes involved
in potato production to better cope with potato tuber moth infestations in the
future.
Prof. Dr. Jürgen Kroschel Dr. Lawrence A. Lacey

International Potato Center, Lima, Agricultural Research Service,
Peru USDA, Wapato, WA, USA
III
Acknowledgements
We are grateful to the organizers of the 6th World Potato Congress for providing the
opportunity to conduct our potato tuber moth symposium. We thank Andy Jensen of
the Washington State Potato Commission for his encouragement and suggestions.
Further, we sincerely thank the authors for their contributions to the proceedings.
Finally, we are grateful to Sonia Santivañez for her great editorial support without
which the publication would have not been possible.
IV
Table of Contents
Preface………………………………………………………….…………………………………….I
Acknowledgements……………………………………………………….…………….….III
Table of Contents…………………………………………………………………….……….IV
Chapter I
Pest status and forecasting…………………………………………….7
Ocurrence of the potato tuber moth, in the Columbia Basin of

Oregon and Washington…………………………………………….………………………9
SILVIA I. RONDON, SANDRA J. DEBANO, GEORGE H. CLOUGH,
PHILIP B. HAMM AND ANDREW JENSEN
Regional and seasonal forecasting of the potato tuber moth

using a temperature-driven phenology model linked with
geographic information systems…………………………………………….………15
MARC SPORLEDER, REINHARD SIMON, HENRY JUAREZ AND
JÜRGEN KROSCHEL
Chapter II
Biocontrol and use of biorationals………………………………….31
An overview of microbial control of the potato tuber moth…………..33

LAWRENCE A. LACEY AND STEVEN P. ARTHURS
The potato tuber moth granulovirus (PoGV): use, limitations

and possibilities for field applications……………………………………………..49
MARC SPORLEDER AND JÜRGEN KROSCHEL
Biological control of potato tuber moth, by Apanteles

subandinus Blanchard in New Zealand…………………………………………..73
TIM J. B. HERMAN
Spinosad: a new biopesticide for Integrated Pest

Management of the potato tuber moth in Tunisia………………….………81
KHAMASSY NOURI AND ISSAM ARFAOUI
V
Laboratory experiments towards the development of an

attract-and-kill strategy for the potato tuber moth complex…………89
JÜRGEN KROSCHEL AND OCTAVIO ZEGARRA
Chapter III
Integrated Pest Management……………………………………..…99
Reducing potato tuber moth damage with cultural practices

and pesticide treatments…………………………………………………….…………101
GEORGE H. CLOUGH, SANDRA J. DEBANO AND PHILIP B. HAMM
Integrated Pest Management dealing with potato tuber moth

and all other pests in Australian potato crops………………………………111
PAUL HORNE AND JESSICA PAGE
Integrated Pest Management of potato tuber moth in New

Zealand………………………………………………………………………………………….119
TIM J. B. HERMAN
Integrated Pest Management of potato tuber moth in India……….127

RAVINDER S. CHANDEL, VIRENDER K. CHANDLA AND
ISHWAR D. GARG
Commercialization of potato tuber moth resistant potatoes in

South Africa……………………………………………………………………………………139
DAVE S. DOUCHES, JOHANN A. BRINK, HECTOR QUEMADA,
WALTER L. PETT, MUFFY KOCH, DIEDRICH VISSER, KARIN MAREDIA
AND KELLY ZARKA
Chapter I
Pest status and forecasting

9
Occurrence of the potato tuber moth, in the

Columbia Basin of Oregon and Washington
Silvia I. Rondon1, Sandra J. DeBano1, George H. Clough, Philip
B. Hamm and Andrew 2Jensen
1
Oregon State University, Hermiston Agricultural Research and Extension
Center, Hermiston OR, USA
2
Washington State Potato Commission, Moses Lake, WA, USA
Abstract
The potato tuber moth (PTM), Phthorimaea operculella (Zeller), is one of the most
significant constraints to potato (Solanum tuberosum L.) productivity worldwide.
Commonly found tropical and subtropical regions, PTM larvae mine leaves, stems,
petioles, and excavate tunnels throughout potato tubers. Although PTM was recorded
in California as early as 1856, the first report of PTM in the Pacific Northwest
occurred in 1913 in Washington. No further evidence of PTM in the Pacific Northwest
existed until 2000 and 2001, when tubers suspected to have been damaged by PTM
were found in Oregon. The objective of this report is to present the status of PTM in
the Columbia Basin. Since 2004, a pheromone trapping network has been deployed in
the Columbia Basin of Oregon (35 traps) and Washington (189 traps) to assess the
distribution, population numbers, and document the spread of PTM. The liners in the
traps were changed once a week and lures were changed monthly. PTM numbers vary
highly from field to field and from area to area. Therefore, it is suggested that control
management recommendations be based on field–specific information. Adequate
control of PTM is critical. Control methods used in other areas may or may not be
effective under Columbia Basin conditions. Thus, control of this pest in the area will
require a novel integrated approach.
Introduction
The potato tuber moth (PTM), Phthorimaea operculella (Zeller), is a cosmopolitan
pest of solanaceous crops, including potatoes (Solanum tuberosum L.). The pest is
difficult to control and over the years farmers have relied extensively on the use of
broad spectrum pesticides. PTM larvae mine leaves, stems, petioles, and excavate
tunnels throughout the potato tubers, making this pest difficult to control.
The history of PTM in the Pacific Northwest (PNW) dates back to the mid-1800s
when PTM was recorded in California as early as 1856 (GRAFT, 1917). In the early
1900s, the presence of PTM was reported in Washington state in the Seattle, Auburn,
and Yakima areas (CHITTENDEN, 1913). Additional information of the presence of
PTM in the Pacific Northwest dates back to 1959 (Boise, Idaho) and 1972 (Corvallis,
Oregon). This information is based on samples deposited in entomological museums
In: Kroschel J and L Lacey (eds.) Integrated Pest Management for the Potato Tuber Moth,
Phthorimaea operculella Zeller – a Potato Pest of Global Importance. Tropical Agriculture 20,
Advances in Crop Research 10. Margraf Publishers, Weikersheim, Germany, 9-13.
10 Ocurrence of PTM in the Columbia Basin
but not based on written reports. No further evidence of PTM in the Pacific Northwest
existed until 2000 and 2001, when tubers suspected to have been damaged by PTM
were found in Oregon. However, PTM was not a major concern for growers in the
Columbia Basin potato production region until 2002 when a field with severe tuber
damage was documented in northeastern Oregon, and was not considered an issue in
Washington until the following year. By 2004, large numbers of the insect were
confirmed from pheromone traps placed in Umatilla and Morrow counties (Oregon),
the southern most region of the Columbia Basin potato production area, and from
southern Washington. Idaho, one of the major potato production areas in the United
States, confirmed the occurrence of the insect in 2005.
Economic losses increased substantially in 2004 and 2005 due to range expansion,
tuber damage, and the cost of control measures. In two years this insect’s range
expanded 140 miles north of Oregon into Washington, including Benton, Franklin,
Adams, Grant, and Lincoln counties. In Oregon, 2006, PTM had spread beyond
Umatilla and Morrow counties, to western, central, and eastern Oregon potato
production areas including Washington, Multnomah, Jefferson, Crook, Klamath,
Union, Baker, and Malheur counties, although no damage has been reported in any of
these areas (Figure 1). In addition, PTM has been confirmed in at least three counties
in western Idaho (Canyon, Payette, and Elmore counties); however, only adults have
been observed and no foliar or tuber damage has been detected. Adequate control of
PTM is critical because larvae infest tubers, rendering them unmarketable regardless of
whether they are fresh marketed or processed (there is a zero tolerance for the presence
of PTM larvae in raw processing product because they are classified as foreign
material). Since this region stores large quantities of potatoes, additional losses are
likely from infested tubers that rot in storage or from infestation of additional tubers
that may occur early in storage facilities.
Figure 1. Potato tuber moth in Oregon. Counties in which it has become established are marked.
PTM has been reported previously in tropical and subtropical countries in South,
Central and North America, Africa, Australia, and Asia (FLINT, 1986; KROSCHEL and
Ocurrence of PTM in the Columbia Basin 11
KOCH, 1994; LARRAIN, 2003). In the United States, PTM has been reported in
California, Arizona, Florida, Texas, Maryland, Virginia (RADCLIFFE, 1982), Colorado,
North Carolina, South Carolina, New York, North Dakota (CHITTENDEN, 1913),
Oregon, Washington (JENSEN et al., 2005; RONDON et al., 2006) and Idaho (RONDON
et al., 2007). Besides potato, PTM has also been reported to infest solanaceous plants
such as tomato, pepper, eggplant, tobacco, and nightshade (FENEMORE, 1988). In the
Pacific Northwest, PTM has only been found infesting potatoes.
The objective of this report is to present the status of PTM in the Columbia Basin
of Oregon and Washington.
Materials and Methods

Since 2004, a pheromone trapping network has been deployed in the Columbia Basin
of Oregon (35 traps) and Washington (189 traps) to assess the distribution, population
numbers, and document the spread of PTM. The pheromones used in the traps are
concentrated quantities of the female sex attractant; therefore the traps attract only
males, although some other insects, including other male moths in the family
Gelechiidae (relatives of PTM) can be trapped in the sticky liners. In the center of each
sticky trap liner was placed a rubber septum containing the pheromone. The liners in
the traps were changed once a week for easy viewing of moths and lures were changed
monthly.
Results
Oregon. In 2004, the average number of PTM per trap per week in the lower
Columbia Basin in Oregon, based on 35 traps, steadily increased from May through
December. The highest numbers of moths trapped per week per trap was in December
(82 moths per trap per week) (Figure 2). The lowest average number of PTM trapped
was in January and February. An increase was observed during May, April, and
August, with a maximum number of 53 moths per trap per week in early April and 49
in mid-August, respectively. The PTM population steadily decreased from September
throughout December 2005. In 2006, from January through May, PTM numbers were
not detected (0 moths per trap per week); however, by the end of June numbers
increased (5 moths per trap per week) with a peak on early October (26 moths per trap
per week) (Figure 2).
Washington. PTM pheromone-baited traps were set out in Washington in late

March, 2005 and mid May in 2004. The total number of moth traps deployed in 2004
was 62, while in 2005 the number was 189. Approximately 120 of these traps were
deployed by cooperators (ConAgra and Simplot). PTM traps were left in place and
monitored weekly through the fourth weekend in November, 2004 and the first
weekend of December in 2005, respectively. In 2005, traps were left in place through
mid-January, when most liners were retrieved without replacement.
By the end of the 2005 trapping season, PTM had been caught throughout
Washington’s potato production region in the Columbia Basin. Numbers were high
12 Ocurrence of PTM in the Columbia Basin
enough to warrant control south of Othello, and generally were much lower north of
Othello. Traps as far north as Wilbur and Coulee City (about 47° 37" North) recorded
moths in 2005.The numbers were high enough and consistent enough to suggest that
the moth had successfully established reproductive populations in these northern areas
that year. In 2006 the trapping network showed extremely reduced populations of PTM
throughout Washington. Most traps were at or near zero through July, but populations
were starting to build in late July and early August in some areas. However, through
July 2006 no PTM had been trapped north of Othello.
Nu mber of PT M males per trap
90
80
70
60
50
40
30
20
10
0
Jan F eb Mar A pr M ay Ju n Jul A ug Se pt Oct No v D ec
2 0 0 4 Av g . o f 3 5 2005 A vg. of 40 2 0 0 6 A v r. o f 3 5
Figure 2. Population dynamics of the potato tuber moth in the lower Columbia Basin, Umatilla and
Morrow counties, Oregon
For updated information on PTM trapping in Oregon visit
http://oregonstate.edu/Dept/hermiston/TrapReports.php (OSU, HAREC), and in
Washington, www.potatoes.com/research.cfm (Washington State Potato Commission).
Discussion
PTM has been found throughout the Columbia Basin of Oregon and southeastern
Washington. Moths may be captured in pheromone traps year round but highest
numbers are generally found in early spring and late summer. The importance of moths
trapped in early spring to the following summer infestation is unknown.
Growers in areas impacted by this insect are encouraged to monitor insect numbers
using pheromone traps. Treatment levels have not been established for Oregon and
Washington; however in California, which has faced PTM damage since the late
1800’s, a treatment threshold of 15-20 moths per trap per night is considered the
general threshold level. However, PTM numbers vary highly from field to field and
from area to area; thus, it is suggested that control management recommendations be
based on field–specific information. Washington reports the trap catch data for PTM
on www.potatoes.com/research.cfm. This website reporting system is map-based, and
was developed specifically for the PTM project. The map is presented as a set of 5x5
Ocurrence of PTM in the Columbia Basin 13
mile grids, each grid containing the data for the traps located in it. These individual
grid squares are linked to the charts of trap catch in that specific grid.
The lack of PTM trapped north of Othello through July in 2006 suggests the region
may not be suitable for establishment of resident populations of PTM. Low numbers in
2006 could be explained as the result of heavy chemical application during the 2005
growing season in combination with cold winter temperatures.
Acknowledgements
We would like to extend our gratitude to summer students and growers cooperators.
Thanks to Aymeric Goyer and Lerry Lacey from USDA-Prosser and Wapato,
respectively, for their comments and editorial contribution. This research was funded
by Oregon and Washington State Potato Commission.
References
CHITTENDEN FH 1913. The potato tuber moth. Bulletin USDA 557, pp 1-7
FENEMORE PG 1988. Host-plant location and selection by adult potato moth
Phthorimaea operculella (Lep. Gelechiidae) a review. Journal of Insect
Physiology 34: 175-177
FLINT M 1986. Integrated pest management for potatoes in the Western United
States. Pub. 3316. Univ. Calif. pp 1-146.
GRAFT JE 1917. The potato tuber worm. US Department of Agriculture,
Washington DC. Bulletin 4217: 56.
KROSCHEL J and W KOCH 1994. Studies on the population dynamics of the potato
tuber moth (Phthorimaea operculella Zell. (Lep., Gelechiidae)) in the Republic
of Yemen. Journal of Applied Entomology 118: 327-341.
LARRAIN P 2003. Plagas de la papa y su manejo. Instituto de Investigaciones
Agropecuarias. Ministerio de Agricultura. Colección Libros INIA No 9,
Intihuasi, Chile. pp 110.
JENSEN A, DEBANO S, DAVID N, MARTIN M and D BATCHELOR 2005. Tuber Moth
Survey, April 2005. Potato Progress 5 (6).
RADCLIFFE EB 1982. Insect pest potato. Annual Review of Entomology. 27: 173-
204.
RONDON SI, HAMM PB and A JENSEN 2006. Population dynamics of the potato tuber
moth in eastern Washington/Oregon. Potato Progress 6 (9):1-2.
RONDON SI, DEBANO SJ, CLOUGH GH, HAMM PB, JENSEN A, SCHEIBER A, ALVAREZ JM,
THORTHON M, BARBOUR J and M DOGRAMACI. 2007. Biology and management of
the potato tuberworm in the Pacific Northwest. A Pacific Northwest Extension
publication of Oregon State University, University of Idaho, and Washington
State University
http://oregonstate.edu/potatoes/ipm/insects/PTW_PNW_594.pdf
14
15
Regional and seasonal forecasting of the

potato tuber moth using a temperature-driven
phenology model linked with geographic
information systems
Marc Sporleder, Reinhard Simon, Henry Juarez and Jürgen
Kroschel
International Potato Center (CIP), Apartado 1558, Lima 12, Peru
Abstract
The potato tuber moth (PTM), Phthorimaea operculella (Zeller) is an economically
important and highly invasive potato pest of global proportions. Temperature-
dependent phenology models are valuable for understanding pest population
dynamics and implementing IPM strategies in different agro-ecological zones. This
paper demonstrates the use of a phenology model developed for PTM linked with
geographic information systems (DIVA-GIS) for mapping population growth
potentials according to real-time or climatic average air temperature data as a tool for
pest risk assessments in different agro-ecological regions and to support the
development of management strategies. The likelihood of population survival
throughout the year, average numbers of generations and an activity index was
computed worldwide using interpolated daily minimum and maximum temperatures
at a spatial resolution of a square kilometer (WorldClim data). Alternatively,
simulations were based upon point temperature series from weather stations (1994-
2004, WMO, Resolution 40) to compute the within year distribution of risk for these
specific areas. In the latter case, life-table parameters (intrinsic rate of population
increase, mean generation time, net reproduction rate, immature survival etc.) were
computed and visualised over time. All simulations were achieved on a daily time
scale that allowed for generating summaries of population phenology and
development within a defined period (i.e. cropping cycle). The approach used for the
simulations of age-stage structured populations is described briefly. Possible uses and
limitations of the presented model linked with GIS (pest risk mapping) in research
and as a decision-aiding tool in Integrated Pest Management and policy (quarantine
pest risk) are discussed.
Phthorimaea operculella Zeller – a Potato Pest of Global Importance. Tropical Agriculture
20, Advances in Crop Research 10. Margraf Publishers, Weikersheim, Germany, 15-30.
16 Regional and seasonal forecasting of PTM
Introduction
The potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera:
Gelechiidae), which probably originated in tropical mountainous regions of South
America (GRAF, 1917), has become a cosmopolitan pest of potatoes and other
solanaceous crops. Today, the moth has been reported from more than 90 countries
and occurs in almost all tropical and subtropical potato production zones where it
causes significant crop losses (CISNEROS and GREGORY, 1994; KROSCHEL and
SPORLEDER, 2006). PTM is considered the most damaging potato pest in the
developing world (RADCLIFFE, 1982). While population development is historically
limited by the 10°C annual isotherm in both the southern and northern hemisphere,
where cold and long winters generally restrict its development, the moth can still be
of economic significance in temperate regions, including New Zealand, southern
Europe (Italy and Spain) and in the United States.
The capacity of PTM to adapt to a wide range of climatic conditions in very
different agro-ecologies was possibly developed in its native habitat in the eastern
Andes where the moth had originally experienced wide daily and seasonal climatic
fluctuations (TRIVEDI et al., 1994). The distribution of insects and other
poikilothermic animals is largely determined by climate (ANDREWARTHA and BIRCH,
1954). Development of organisms, which cannot internally regulate their own
temperature, depends principally on the temperature to which they are exposed to in
the environment. Climate also extensively determines the distribution of host plants,
and thus indirectly influences the distribution of pest insects.
The biology and ecology of PTM have been extensively studied in different parts
of the world in the field and laboratory (FENEMORE, 1977; FOOT, 1979; BRIESE, 1980;
1986; CHI, 1988; CHI and GETZ, 1988; KROSCHEL and KOCH, 1994; TRIVEDI et al.,
1994; KELLER, 2003). Temperature-dependent phenology models for PTM were
developed (ROUX and BAUMGÄRTNER, 1995; SPORLEDER et al., 2004), which could
provide information of the moth’s population growth potential in a given potato
production area or region according to temperature conditions. This is important
information for both assessing the establishment potential of the pest and pest
management. Linking pest insect phenology models with Geographic Information
Systems (GIS) allows for visualizing predicted pest potentials for specific countries
or regions (pest risk mapping) while accounting for seasonal (within year) and intra-
year variation.
The purpose of this study was to implement one of the temperature-driven
phenology models for PTM in a GIS environment and to predict the moths’
population growth potentials in potato growing areas worldwide. Temporal variations
in population growth due to seasonal temperature fluctuations were simulated for a
single location as an example. How far this computer-aided simulation approach
might help to improve the current understanding of quantitative pest biology and how
it could be used as a decision-aiding tool for pest management in specific agro-
ecologies will be discussed briefly.
Regional and seasonal forecasting of PTM 17

Phenology model
The phenology model developed by SPORLEDER et al. (2004) for PTM, which was
validated by comparing model outputs with life-tables derived from field studies
conducted in different agro-ecological zones (KELLER, 2003), was used in this study.
Functions and parameters used in the model to describe development, mortality and
reproduction are presented in Table 1.
Table 1. Parameter values of the functions used to describe simulation of P. operculella

development, mortality and reproduction. Forms of the functions are demonstrated in
right-side figures (see SPORLEDER et al., 2004) for further description of model terms)
Development rate [SHARPE and DEMICHELE (1977) model]
Life-stage RHO25 HA TL HL
Dev.rate
Egg 0.767 5276.7 297.9 -30463
Larva 0.317 6126.3 303.3 -24558
Temp
Pupa 0.664 -133.8 304.9 -27966
Distribution function of development [lognormal CDF]
Development
Slope
Egg 35.52
Larva 23.34
0 1
Pupa 20.26 normalized age
Mortality [second-order polynomial function]

a1 a2 a3
Mortality
Egg 0.0003 -0.0177 0.3598

Larva 0.0027 -0.1404 2.0457
Pupa 0.0017 -0.0828 1.0613 Temperature
Total eggs per female [second-order polynomial

function]
Oviposition
a1 a2 a3
-1.029 42.5037 -269 Temperature
Proportion of progeny production with age [cumulative

P ro p o rtio n o f
gamma function]
p ro g en y
α β
1.1977 5.4859
0 normalized age 1
RHO25=development rate at 25ºC; HA, TL, and HL represent kinetic parameters of rate-controlling
enzymes.
Climate data
The phenology model requires daily maximum and minimum temperatures as
minimal inputs and may additionally be parameterised to incorporate rainfall.
Assessment of the pest indices on a worldwide scale simulation (risk maps) was
based on WorldClim data. WorldClim is a set of global climate layers (grids) with a
spatial resolution of 30 arc seconds (~1 km2, downloadable at
http://www.worldclim.org). The data are described in HIJMANS et al. (2005).
However, for compiling the maps presented in this paper, the data were used at a
lower resolution of 10 min (18.6×18.6=344 km2 at the equator) derived through
aggregation. WordClim provides monthly aggregated climate variables. Because this
raises substantial problems of temporal scale, daily maximum and minimum
temperatures were interpolated for each grid before simulation. While more
computationally complex than interpolating point phenology results, this method is
considered to provide more flexible and robust results.
For simulating within-year variations of pest population growth, real time daily
minimum and maximum temperature data from a meteorological station in
Cochabamba, Bolivia, were used (WMO data, Resolution 40). The data are available
for the years 1994 to 2004 (downloadable at http://www.ncdc.noaa.gov/cgi-
bin/res40.pl?page=gsod.html).
Software
A program in a Java environment was developed to carry out the grid processing
geographical and phenological simulations and the calculations of pest risk indices.
For visualizing pest risk potentials (risk maps), components of DIVA-GIS (Version
6.0.2) were coupled with the program.
Phenological and geographical modeling

A 15 min time step length was chosen for the model. Model calculations were based
on daily maximum and minimum air temperatures; either real point-based weather
station records or interpolated grid-point data. Temperature in each 15 min time step
was calculated using a cosine function for half-day temperature predictions. The
equation used for the first half-day was:
Ti =
(Max − Min ) × cos⎛ π × (i + 0.5) ⎞ + (Min + Max )
⎜ ⎟
2 ⎝ 48 ⎠ 2
in which Ti is the temperature (in ºC) of time step i ( i = 1, 2, 3, …48), and Min and
Max are daily minimum and maximum temperatures. The calculation was then
repeated to obtain Ti for the second half-day employing the minimum temperature,
Min, of the following day in the equation.
The phenological simulation was executed in two phases. First, the program
calculated stage-specific daily development rates, mortality rates (for egg, larva,
pupa), adult senescence rates (for females and males) and total fecundity per female
according to temperature starting from Julian day 1 (January 1st) to Julian day 365
(December 31st). From these results the following parameter estimates were
generated for each Julian day:
a) Generation length in days (T)
T = (1/dEgg) + (1/dLarva) + (1/dPupa) + (1/sFemale) * 0.163
where dx is the stage-specific development rate of immature life-stages, s is the
senescence rate of females and 0.163 is the normalized age of females until 50%
oviposition.
b) Net reproduction rate (R0) = fecundity * immature survival / 2
c) Intrinsic rate of increase (rm) = ln(R0) / T
d) Finite rate of increase (λ) = exp(rm)
e) Doubling time (Dt) = ln(2) / rm
These data were plotted against Julian day to visualize variability in pest
potentials due to seasonal climate variation (only one example is given in this paper).
In the second phase, from these model outputs three indices for pest risk, as defined
below, were calculated.
Establishment risk index (survival risk)

This index identifies the areas in which the pest may survive. The index is 1 when a
certain proportion of all immature life stages of the pest survive throughout the year.
Otherwise, the number of days in which a single stage would not survive are counted
and divided by 365. These ratios, x, for the egg, larval and pupal stage are employed
in the following equation:
Establishment Risk Index = (1-xEgg) × (1-xLarva) × (1-xPupa)
Generation index
This index estimates the mean number of generations that may be produced within a
given year. The generation index was computed by averaging the sum of estimated
generation lengths, Tx, calculated for each Julian day, x, as demonstrated in the
following equation:
Generation Index = ∑ 365 / Tx

365
where 365 is the number of Julian days per year and Tx is the predicted generation
lengths in days at Julian dayx (x = 1,…, 365). The number of generations per year
does not give conclusive information about population growth potentials.
Development times decline with increasing temperatures, thereby increasing the
speed of population build-up; however, at extremely high temperatures, immature
survival and oviposition also declines so that extremly high generation numbers per
year may result in lower population increase with time.
Activity index
This index is explicitly related to the finite rate of population increase, which takes
the whole life history of the pest into consideration. The index was computed by
employing the following equation:
Index = log Π λx
where λx is the finite rate of increase at Julian dayx (x = 1,…, 365). For example, an
index value of 4 would illustrate a potential population increase by a factor of 10,000
within one year (all other population limiting factors, including food availability, etc.,
are neglected). Alternatively, the index could be related to the population doubling
time, Dt, because the parameter has the same biological significance. In addition to
the establishment risk index, this index illustrates the pest severity and spread
potential.
Results and discussion

Global indices of risk
Establishment risk (survival) index

Figure 1 globally indicates the regions where PTM populations theoretically could
establish according to temperature conditions. The map can be presented for potato
production areas only (Figure 1B) or for the whole world (Figure 1A).
The latter map is reasonable when PTM host crops other than potatoes, e.g.
tomato (Lycopersicon esculentum Mill.), are cultivated in the region, increasing the
risk of pest establishment. Regions where the index is 1 indicate that a certain
proportion of the PTM population is always expected to survive, i.e. complete the life
cycle, throughout the year. In these regions, the long-term likelihood of getting
established when accidentally introduced is highest. The whole tropical and
subtropical regions fall within this characteristic; however, population increase may
be very variable throughout the year and between different zones. In zones where the
index deviates from the maximum number of 1 the likelihood of long-term
establishment is considerably reduced.
PTM has been reported from regions where the map indicates a survival index
between 0.6 and 0.85, but populations never became established. Economically
significant outbreaks occurred occasionally in cooler regions, including New Zealand
(PEACOCK and WORNER, 2006), southern Europe (Italy, Spain) (ARNONE et al., 1988:
PUCCI et al., 2003), and in the southwestern USA (California) (SHELTON and
WYMAN, 1979), regions that are assigned to an index between 0.85 and 0.99;
however, in those regions the pest never reached a significant status in the long term.
In Korea, the index of >0.6 coincided approximately with the isotherm of –8ºC
minimum temperature in January, restricting the distribution of PTM (CHOE and
PARK, 1980). However, PTM has also been reported from areas that were assigned
with an index <0.6, as for example in the Ukraine (BAKLANOVA et al., 1994).
Recently, the outbreak of PTM in the Northwest USA (Oregon, Washington)

received much attention. The pest was first reported in that region in 2003 and again
during 2004 and 2005. However, in 2006, presentation of the pest was marginal. It
seems that it is possible to achieve eradication of the pest in regions of such low
survival indices. With the exception of large cull piles, no other pest refuges, such as
favourable potato stores, exist for winter survival.
Generation index
Multiple generations of PTM developing within a cropping cycle and during potato
storage periods obviously have the potential to cause considerably more damage than
a single one. As the generation index, i.e. the average number of generations per year
increases, the damage potential increases as well (Figure 2). Therefore the index is
strongly correlated with the activity index explained in the following section.
However, at high temperature (>30ºC) mean generation time still decreases while
population increase may be reduced (even negative) because the intrinsic rate of
population increase is negatively affected by increased mortality and reduced
reproduction rates (see northern Sudan).
Globally simulated generation indices (Figure 2) gave reasonable predictions
when compared to generation numbers reported in the literature. For example, 13
generations per year were observed at temperatures of 20.5 to 33.0ºC (52-66.0% RH)
in Saudi Arabia, while 9 generations occurred during the first 6 months, 4 in the last 3
months and no generations were completed between early August and late October
when temperatures were highest (EL-ATROUZY and AWAAD, 1986). However, most
researchers report on maximum numbers of generations per year because they
experimentally assess generation times from egg to first progeny (1st generation),
which is used to start a new cycle (2nd generation). The number of cycles completed
within one year represents the maximum number of generations produced per year.
For example, by using this methodology, 18 generations were reported from
Bangladesh (KABIR, 1994), 10 generation from Egypt (ABDEL-WAHAB et al., 1987;
ABUL-NASR et al., 1971), 9 from Yemen (KROSCHEL and KOCH, 1994). KELLER
(2003) determined 3-4 generations in Huancayo, Peru (3300 m a.s.l.), 7 generations in
Arequipa, Peru (1440 m a.s.l.), and 12 generations in San Ramon, Peru (770 m a.s.l.)
per year, restarting each generation cycle when about 20% of progeny was produced.
In reality, it is impossible to assess mean generation times throughout the year in
fluctuating temperatures through rearing experiments; large variation in development
times of individuals and large oviposition periods always result in an age structured
generation and multiple cohort rearing would be not feasible; oviposition of eggs of
the 3rd generation would be already mixed with the eggs of the 4th generation when
the first generation was started with eggs of the same age.
The phenology model takes these stochastic components into account, calculating
the average number of generations, which is approximately the time from eggs to
50% oviposition of eggs of the following generation. Hence, when using literature
data for comparing model results, this aspect needs to be considered. The number of
generations per year or cropping cycle is an important index when management of
insect resistance development against insecticides or Genetically Modified

Organisms (GMO) is of interest. With increasing generation numbers, i.e. shorter life
cycles, the risk that resistance develops in an insect species against a pesticide or that
an insect species breaks the resistance of GMO crops increases.
Activity index
Activity indices for PTM, specifying the potential population increase within a year
for potato production areas worldwide, are presented in Figure 3. An index of 4
indicates a 10,000-fold (=104) potential increase of population numbers within one
year. Each increase of the index by 1 indicates a 10-fold higher increase rate.
However, the rate of population increase might oscillate considerably within a year.
For example, in high-elevation mountainous regions of the Andes (>3300 m a.s.l.) the
index is around 1 to 4, which is very similar to indices calculated for the northwestern
USA (Oregon, Washington). In both regions, average yearly temperatures may be
quite similar (e.g., about 12.5ºC in Huancayo, Peru and 12.3ºC in Hermiston,
Oregon). However, due to relatively low within-year temperature fluctuations in
Huancayo, populations could increase with finite rates between 1 and 1.04. In
Hermiston, due to larger within year temperature fluctuation and lower within-day
temperature fluctuations, the finite rate of increases may fluctuate within the year
from 0.98 (negative increase) to 1.15; with highest increase rates during the warm
cropping season. Hence, in temperate climates, the pest activity can be higher during
the cropping season as compared to tropical mountainous regions. On the other hand,
in tropical dry areas where potatoes are grown during the cooler winter month (e.g.
Northern Africa) the potential rate of population increase would lower during the
cropping season. Heavy rain (>5 mm/day) may additionally reduce pest activity
(FOOT, 1979; WHITESIDE, 1980; KROSCHEL and KOCH, 1994; KELLER, 2003).
Accounting for within-year variation

The model can be used for a detailed analysis of within-year variation of pest
population increase. Figure 4 shows an example for such an analysis by plotting
simulated life-table parameters against Julian days. Average minimum and maximum
temperature data from a meteorological station in Cochabamba, Bolivia (1994-2004;
WMO data) were used for the simulation.
Maximum temperature fluctuates between 25 and 30ºC within the year, while
minimum temperatures drop from about 14ºC during the summer months (December
– March) to >2ºC during winter months (June – August). Potatoes are grown from
November to March/April. During the total cropping season, PTM populations
potentially might increase with a finite rate of about 1.08 (A); accordingly
populations double within less than 10 days (Figure 4F). Survival of immature life-
stages is about 45% (D) and it would take an average of about 45 days to complete
one generation (C). Flight activity might be expected to be high during the cropping
season; each female adult reproduces about 30 to 35 females during the cropping
season, while this number is less than 10 during the cooler months (B). This analysis
considers the effects of temperature only, and ignores other factors like rain, food
availability, parasitoids, etc. Hence, the results should be considered as the potential
population growth parameters in a given region or zone. However, in combination
with field observation, this model’s outputs contribute to a better interpretation of
PTM population dynamics.
Figure 1. Survival index of P. operculella simulated using interpolated daily minimum and
maximum temperature data globally (A) and for the potato production areas only (B). The
index is 1 where all immature life-stages develop and survive throughout the year. An
index less than 1 indicates that individuals of at least one immature life-stage do not
survive at least one day within the year (see explanation in the text). It can be considered
that P. operculella is barely established in regions where the index is lower than 0.85.
Figure 2. Activity indices for P. operculella simulated globally for potato production areas using
interpolated daily minimum and maximum temperature data. The index specifies
population increase potentials within a year. An index of 4 indicates a 10,000-fold
(104) potential increase of population numbers within one year. The index may be very
different for different time periods within the year. Alternatively, the index can be
calculated for the potato-cropping season only or for specific time-spans of the year.
Figure 3. Generation index for P. operculella simulated globally using interpolated daily
minimum and maximum temperature data for the potato production areas. The index
shows mean numbers of generations developing within a year. P. operculella
produces overlapping generations and hence heterogeneous populations, comprising
all life-stages, typical at any time of the year. Note that due to variation in
development times and extending oviposition periods the maximum number of
generations might differ largely from the mean number of generations determined in
this map.
A B
1.15 40
net reproduction rate (♀/♀)

35
finite rate of increase
1.10
30
1.05 25
20
1.00 15
10
0.95
5
0.90 0
0 100 200 300 0 100 200 300
Julian day Julian day
C D
90 1.0
80 0.9
generation time (days)
70 0.8
inmature survival
60 0.7
0.6
50
0.5
40
0.4
30 0.3
20 0.2
10 0.1
0 0.0
0 100 200 300 0 100 200 300
E F
35 30
30
doubling time (days)
25
temperature (ºC)
25
20
20
15
15
10
10
5 5
0 0
0 100 200 300 0 100 200 300
Figure 4. Within year variation of life-table parameters for P. operculella simulated for
Cochabamba, Bolivia, using mean daily minimum and maximum temperature data
from a meteorological station (1994-2004; WMO data). Figures A, B, C, D, and F
show the fluctuations of population life-tables parameters, i.e. finite rate of increase,
net reproduction rate, mean generation time, immature survival, and population
doubling time, as examples. Figure E indicates daily mean maximum (blackdots) and
minimum (open dots) temperatures; bars above the x-axis indicate the potato-cropping
season.
General outlook
Climate is one of the major factors limiting the distribution of pest species. Assessing
the potential for an organism in a given area can be done in several ways by using
different methodologies. Computer-aided tools such as CLIMEX (PEACOCK and
WORNER, 2006; WILMOT SENARATNE et al., 2006) and BIOCLIM (KOHLMANN et al.,
1988; STEINBAUER et al., 2002) seek out the potential exploitation of non-indigenous
invasive species to areas by looking for climatic similarity to the climatic
characteristic of the region where the species originates. These programs use match-
climate functions, which compare the long-term meteorological data for each of a
selected location where the species is absent with the location of origin (or locations
where the species prevails) (SUTHERST and MAYWALD, 1991). The approach
presented in this paper is to develop a full species climatic response model
(phenology model) based on laboratory assessments for a pest species that allows the
simulation of populations and produces life-table parameters according to real or
interpolated temperature data for a given region and time. The current model for PTM
species takes temperature as the only influencing variable on insect phenology
because these effects are well assessed through laboratory experiments. However,
further effects of biotic or abiotic factors should be quantified in future research and
included in the model.
The simulation approach can be used to examine the distribution of PTM species
for a variety of purposes determining the climatic/temperature mechanisms that limit
the geographical distribution of the pest species and to identify the climatic
conditions that favour population growth. Further species-specific limiting factors
and stress indices should be determined and made available for each location; i.e.
potato vegetation periods, rain intensity, parasitoids, and the limits of extreme
temperatures. Gradients of suitability of habitats for PTM in terms of temperature can
be identified and related to the spatial abundance of host species.
In its present state, the model provides information about the potential pest
population growth under a given temperature regime, which is useful for
understanding quantitative pest biology. An important issue in integrated pest
management (IPM) research is to evaluate the potential effects of certain pest
management strategies. If models for specific parasitoids are available, they can be
coupled with the existing PTM model (“two-species-interaction model”) and used to
predict potential release areas according to climatic (temperature) conditions. For this
purpose, CIP currently develops phenology models for some prospective PTM-
specific parasitoid species. As another example, the potential of entomopathogenic
microorganisms used in IPM strategies can be evaluated. Several entomopathogens
attack only specific stages of the pest, and within a given stage individuals of a
specific age only (for example young larvae). The PTM model provides essential
information about the pest insect population age-stage structure, which makes it
possible to include a mortality factor in the model that is specific to individuals of a
certain stage and age class within the PTM population. An example how such a
mortality factor might be coupled with the PTM model is discussed in a further paper
presented in this book for the baculovirus (PoGV) – PTM system by SPORLEDER and
KROSCHEL (2008). However, this pathogen-pest interaction model still does not
simulate pathogen epizootics in the field over time in its totality because several
influencing factors are not known at present (virus stability, horizontal and vertical
virus transmission, etc.). Knowledge of such factors and their effects on population
development in the field are essential for realistic model results and need to be
assessed in additional experiments. Nevertheless, this modelling is valuable in

exploring the factors that could be important for using entomopathogens in pest
management strategies and might help to define hypothesis for field experiments
conducted in different agroecologies under a variety of climatic conditions. As a last
example, the PTM model might be used to forecast the potential changes in the
distributions and activities of PTM in response to global warming.
Thus, the presented model might be used by scientists and IPM practitioners as a
tool that assists country-specific pest risk assessments for PTM and adaptation
planning, and for improving pest management strategies. Policy makers might use the
information to improve national pest management and quarantine programs.
The approach used to develop and implement the P. operculella model as
presented in this paper can be principally applied for other insect species. CIP is
currently developing a software package (called Insect Life Cycle Modeling-ILCYM)
that facilitates the development of further pest insect phenology models and allows
for pest risk mapping of a specific pest under study as described in this paper. An
alpha version of this computer program in enclosed to this proceeding. The “open
source” package consists of two main modules; a) the “Model Builder”, which
facilitates researchers to develop pest insect phenology models based on data from
temperature experiments of a specific pest species - this module also provides life-
table analysis tools and instruments for validating developed models – and b) the
“risk mapping” environment, which couples developed models with GIS tools for
spatial simulation of the pest’s activity. The PTM model as presented in this paper is
already available in this version. For further information on installation and use of the
software see the manual on the CD enclosed here. For follow-up versions and up-
dates of the software please examine the CIP homepage (www.cipotato.org) or
contact one of the authors of the present paper.
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Chapter II
Biocontrol and use of biorationals

Regional and seasonal forecasting of PTM
33
An overview of microbial control of the potato

tuber moth
Lawrence A. Lacey1 and Steven P. Arthurs2
1
Yakima Agricultural Research Laboratory, USDA-ARS, 5230 Konnowac Pass
Road Wapato, WA 98951 USA
2
Mid-Florida Research and Education Center, UF-IFAS, Apopka, FL 32703, USA
Abstract
Naturally occurring insect specific pathogens and inundatively applied biopesticides
can significantly contribute to control of the potato tuber moth (PTM). The most
researched and practically used for control of PTM are a granulovirus and the
bacterium Bacillus thuringiensis Berliner (Bt). The PTM granulovirus (PoGV) has the
potential to play a significant role in the integrated management of PTM in stored
tubers and in field crops. Similarly, Bt has been successfully used against PTM
infestations in both field crops and stored tubers. PoGV and Bt are safe to application
personnel and the food supply and do not affect beneficial insects and other non-
target organisms. Other natural insecticides include a biofumigant fungus (Muscodor
albus), botanicals, sex pheromones and physical measures for PTM control in stored
tubers. The implementation of biopesticides will ultimately depend on an increased
awareness of their attributes by growers and the public, which will be the main
drivers for their use and commercialization.
Introduction
A diverse spectrum of insect pathogens (bacteria, fungi, viruses, protozoa, and
nematodes) infects and kills pests of virtually every crop. Several of these agents have
been developed as microbial pesticides (LACEY et al., 2001; KAYA and LACEY, 2007),
some of which have been used to control key insect pests of potato including potato
tuber moth (PTM), Phthorimaea operculella (Zeller) (VON ARX et al., 1987;
HAMILTON and MACDONALD, 1990; RAMAN, 1994; CLOUTIER et al., 1995; KROSCHEL
et al., 1996b; LACEY et al., 1999; SPORLEDER, 2003; WRAIGHT et al., 2007).
Substantial effort has gone into the development of certain microbial agents for PTM
control in several countries worldwide. Until recently, only limited attention has been
paid to biopesticides for PTM control in the United States. In this overview we will
present information regarding the development and potential of a virus, a bacterium,
fungi, and entomopathogenic nematodes for control of PTM.
34 Microbial control of PTM
Insect pathogens
Virus
A granulovirus (PoGV) that attacks PTM larvae has accompanied the moth from its
South American center of origin to most countries, where it has become established
(including the United States). Extensive surveys confirm the presence of the virus in
PTM populations in the Andean potato growing areas of South America (ALCÁZAR et
al., 1991, 1992a). Several isolates and their origins are summarized by SPORLEDER,
2003). The virus has been reported from Africa (BROODRYK and PRETORIUS, 1974;
LAARIF et al., 2003), the Middle East (KROSCHEL and KOCH, 1994), Asia (ZEDDAM et
al., 1999; SETIAWATI et al., 1999), Australia (REED, 1969; BRIESE, 1981) and North
America (HUNTER et al., 1975).
The name of the virus is derived from its granular appearance under high
magnification (Figure 1A). Each granule contains a single viral rod (Figure 1B). After
the granules, also known as occlusion bodies (OB), are ingested by PTM larvae, they
dissolve in the alkaline midgut freeing the viral rods which attach to and pass through
the membrane of the midgut epithelial cells. From there they invade a variety of host
cells and produce hundreds of millions of rod-containing OBs per larva. The larval fat
cells are the predominant site of virus production. Ultimately, infected larvae die and
become a source of inoculum for infection of other PTM larvae. REED (1971)
reported on the effect of virus concentration, temperature and larval age on the
progression of disease in PTM larvae. Most larvae die within 2-3 weeks of ingesting
virus, but very high dosages of PoGV can cause death by toxicosis within 48 hours.
SPORLEDER (2003) assessed the infectivity and virulence of 14 geographical
isolates of PoGV and found a wide range of activity covering several orders of
magnitude. VICKERS et al. (1991) demonstrated minor differences among 8
geographically diverse PoGV isolates using restriction endonuclease DNA analysis.
Three distinct, but closely related genotypes of PoGV were revealed and the Peruvian
isolate was readily distinguishable from granuloviruses of 5 other insect hosts
(VICKERS et al., 1991). Bioassays of 3 PoGV isolates from Indonesia revealed similar
biological properties (ZEDDAM et al., 1999). The restriction pattern of the Indonesian
Wonsosobo isolate varied only slightly from other PoGV isolates from different
regions of the world (ZEDDAM et al., 1999). KROSCHEL et al. (1996a) reported
similarity between an isolate from Yemen and a Peruvian isolate. In contrast, LERY et
al. (1998) demonstrated considerable genetic heterogeneity between a Tunisian
isolate and isolates of PoGV from other regions.
Like most granuloviruses, PoGV has a fairly specific host range. Only PTM and
certain other species in the same family (Gelechiidae) are infected by the virus. Tecia
solanivora Povolny is susceptible to PoGV, but at lower levels than PTM (ZEDDAM et
al., 2003). Although PoGV could be isolated from Symmetrischema tangolias (Gyen)
(ANGELES and ALCÁZAR, 1996), it does not appear to affect this species (J. Kroschel
and M. Sporleder, personal communication). POKHARKAR and KURHADE (1999)
reported no infectivity to 11 other lepidopteran species.
Microbial control of PTM 35
1A 1B
Figure 1. Occlusion bodies of the potato tuber moth granulovirus (PoGV). 1A Intact PoGV
OBs. Scanning electron micrograph courtesy of the International Potato Center,
Lima, Peru. 1B Cross and longitudinal sections of PoGV OBs. Transmission electron
micrograph courtesy of Darlene Hoffmann, USDA-ARS, Parlier, CA.
The natural incidence of PoGV in PTM populations has been documented in
Australia and Yemen (BRIESE, 1981; KROSCHEL and KOCH, 1994; KROSCHEL, 1995)
where PTM larvae were infected at low frequencies. KROSCHEL and KOCH (1994)
also observed that the majority of PTM mortality in Yemen was caused by braconid
and ichneumonid parasitoids. KROSCHEL et al. (1996b) surmised that parasitoids were
slightly inhibited by application of 5x1013 PoGV OBs/hectare, but not by application
of one tenth that amount of the virus. Parasitoid larvae ostensibly continued
development within live virus-infected PTM larvae. Given the opportunity for
different natural enemies to control PTM, interactions between PTM parasitoids and
PoGV warrant further attention in potato agroecosystems. Compatibility of
parasitoids and entomopathogenic virus of other Lepidoptera are reported by BROOKS
(1993) and BEGON et al. (1999).
Parasitoids are better suited for exploiting uninfected hosts, particularly in cryptic
habitats, because of their abilities to search, whereas most pathogens, such as PoGV,
must wait for chance encounters. According to BEGON et al. (1999), one of the most
important aspects to consider in the integration of pathogens and parasitoids is the
stage of the host that is attacked. The fact that PoGV normally infects neonate larvae,
while many parasitoids of PTM attack eggs and older larvae, could enhance combined
control. Interaction of PoGV and natural enemies is not limited to parasitoids.
MATTHIESSEN and SPRINGETT (1973) made interesting observations on a bird, the
silvereye (Zosterops gouldi (Bonaparte)), which is a predator of PTM and a potential
vector of PoGV in Australia.
VON ARX and GEBHARDT (1990) studied the survival of PTM from egg to adult
after exposure to 0.2, 0.02, and 0.002 PoGV infected larvae or “larval equivalents”
(LE) per kg of tubers. Survival and the intrinsic rate of increase of PTM were
significantly reduced after exposure to the two highest concentrations, but not at
0.002 LE. It is evident from the above study and the natural incidence of PoGV in
PTM populations, that in order to provide effective control of PTM populations (i.e.
to reverse population increases and reduce damage caused by larvae in plants and
tubers), inundative applications (e.g. through spraying) of the virus will be required.
Application of PoGV for control of field populations of PTM has been somewhat
limited and the results have been variable. REED (1971) and REED and SPRINGETT
(1971) conducted the first field trials of PoGV in Australia and found that an early
application of virus (6275 LE/hectare) could achieve effective control. They also
observed that PoGV readily spread into untreated areas. REED (1971) concluded that
virus reached leaf mining larvae through the stomata and that wind and birds were
responsible for spreading the virus. SALAH and AALBU (1992) tested a PoGV
suspension and powder under field conditions in Tunisia. Virus was applied to the
surface of the soil in potato fields only incidentally reaching the plants. Field
infestation of tubers by PTM was reduced by up to 73%. KURHADE and POKHARKAR
(1997) reported that PoGV at 5.5 x 1011 OBs/hectare plus endosulfan (0.035%)
provided effective control of PTM, resulting in the lowest tuber infestation (6.9%)
compared to other insecticidal treatments. SALAH et al. (1994) tested a combination of
Bacillus thuringiensis (Bt), PoGV and extra irrigation for integrated control of PTM
in Tunisian field trials. In some cases, the integrated controls proved to be more
efficacious than conventional insecticides. Field evaluations of PoGV in Yemen were
reported by KROSCHEL (1995) and KROSCHEL et al. (1996b) where applications of
5x1013 PoGV OBs/hectare of potato plants resulted in significant control of PTM.
Typical symptoms (milky white coloration and reduced vitality) were observed in
larvae 11 days after treatment and 70% mortality was noted 19 days after treatment.
Ultimately, virus treatments resulted in up to 82.5% mortality of PTM.
Field evaluations of PoGV and Bt subspecies kurstaki (Btk) were conducted in
heavily infested Russett Burbank potatoes in Washington State, USA (ARTHURS et
al., 2008a). Five weekly applications of PoGV (1013 OB/ha), Btk (1.12 kg/ha) or their
combination (weekly alternations at the same rates) per larval generation did not
reduce initial damage (% mined leaves) in the 1st generation, although >90% of
treated larvae were virus-infected and failed to pupate. Season-long treatments
resulted in 76.3% reduction in mined leaves and 96.3% reduction of PTM larvae after
the 2nd generation. Btk was slightly less effective than PoGV (91.6% larval reduction
in 2nd generation). The PoGV/Btk rotation was more effective than Btk alone (97.1%
larval reduction 2nd generation). In addition to effects of PoGV strain, methods of
application, and concentration on efficacy, inactivation of the virus by the ultraviolet
(UV) radiation in sunlight can rapidly reduce the amount of inoculum available to
larvae under field conditions (KROSCHEL et al., 1996a; SPORLEDER et al., 2001;
SPORLEDER, 2003; ARTHURS et al., 2008a). Different preparations of PoGV were
investigated by KROSCHEL et al. (1996a) for their efficacy and persistence on leaves
and tubers in the field. They calculated a half life of PoGV on tubers exposed to the
sun to be 1.3 days. Mortalities of first instar larvae ranged from 43-49% when fed
vegetation collected 2 days after treatment. Only 19.4-25.8% of larvae died when fed
on foliage collected 8 days after virus application. A variety of adjuvants that have
been used to protect other baculoviruses from UV inactivation were reviewed by
BURGES and JONES (1998). SPORLEDER (2003) investigated formulations that included
dyes, optical brighteners, antioxidants, insect host derived materials for protection of
PoGV from UV inactivation. He noted that an optical brightener (Tinopal) and certain
antioxidants protected the infectivity of irradiated virus. However, PoGV-infected
larvae macerated in water were superior to other preparations in protecting the virus
from UV irradiation (KROSCHEL and KOCH, 1996; SPORLEDER, 2003).
PoGV has been reported by several researchers to provide good protection of
treated tubers, especially in non-refrigerated storage. PoGV was tested successfully
on stored tubers in several Andean countries (Peru, Ecuador, Bolivia, and Colombia),
where PTM and the potato are believed to have originated (ALCÁZAR et al., 1992b;
CIP, 1992; ZEDDAM et al., 2003). The virus has also been evaluated in tuber stores in
the Middle East, Northern Africa, Asia, and North America (AMONKAR et al., 1979;
HAMILTON and MACDONALD, 1990; ISLAM et al., 1990; ALI, 1991; DAS et al., 1992;
SETIAWATI et al., 1999; ARTHURS et al., 2008b). Bioassays conducted at 25˚C by
ARTHURS et al. (2008b) showed that PoGV suspended in water or mixed with carriers
(talcum, sand, diatomaceous earth and kaolin clay), was highly effective in
controlling neonate larvae in stored tubers. Because the virus is not exposed to UV
degradation, protection of tubers in storage may last several months.
The potential for development of resistance to PoGV in PTM larvae has been
presented by BRIESE and MENDE (1981, 1983) and SPORLEDER (2003). BRIESE and
MENDE (1981) noted an 11.6 fold difference in susceptibilities to PoGV among 16
field populations of PTM in Australia. BRIESE and MENDE (1983) also observed a 140
fold increase in the LD50 after serial exposure of susceptible PTM larvae to PoGV
over 6 generations. Similar observations were made by SPORLEDER (2003). PTM
larvae that survived exposure to virus concentrations that produced 50, 75, and 90%
mortality in the parent susceptible population were highly resistant to the virus after
12 generations. A single backcross with the susceptible population did not decrease
the level of resistance indicating that the gene or genes responsible for resistance may
be dominant. Based on the above information, resistance management should be
incorporated in PoGV control programs.
Methods for the in vivo production of PoGV are presented by REED and
SPRINGETT (1971), CIP (1992), KROSCHEL et al. (1996b), SPORLEDER (2003) and
others. Basically, the method employs the mass production of PTM followed by
infection of neonate larvae on tubers pre-treated by submersion in an aqueous
suspension of triturated PoGV-infected larvae. Alternatively, PTM eggs can be
dipped in PoGV suspensions (SPORLEDER et al., 2005). Larvae consume virus directly
upon exiting the egg and are provided tubers in which to develop. SPORLEDER (2003)
presented information on the effect of temperature, initial virus concentration, larval
age and density on the yield of OBs. The number of virus infected larvae increased
with increasing virus concentration with an optimal concentration of 109 OBs/ml of
suspension. The optimal temperature and larval density for virus production was 25°C
and 2 grams of potato/larva, respectively. POKHARKAR and KURHADE (1999) also
reported 25°C as optimal for virus production. Another production method involves
spraying of virus suspensions onto infested potato plants in the field, collecting
infested foliage after larvae become diseased, and separating infected larvae using
exposure to heat (MATTHIESSEN et al., 1978). LERY et al. (1997) and SUDEEP et al.
(2005) reported on the establishment of PTM cell lines and demonstrated their utility
for in vitro production of PoGV. The virus has been commercially produced in Peru,
Bolivia, Egypt, and Tunisia. Based on PoGV’s safety, potential for resistance
management for chemical pesticides, and minimal impact on beneficial non-target

organisms, it appears to be a ready made component for incorporation into IPM
programs and warrants further development and commercialization.
Bacteria
The only bacterium that has been evaluated for PTM control is Bt. It is a naturally
occurring bacterium that produces crystal toxins (stomach poisons), which cause
disease in insects through the lysis of midgut epithelial cells (BEEGLE and
YAMAMOTO, 1992; GARCZYNSKI and SIEGEL, 2007). Insecticides based on Bt toxins
are the most widely used of microbial pesticides and are commercially produced for
use against a broad range of pests including Coleoptera (beetles), Diptera (flies) and
Lepidoptera (caterpillars), including species that attack potato (KRIEG et al., 1983;
HAMILTON and MACDONALD, 1990; KROSCHEL and KOCH, 1996; LACEY et al., 1999;
WRAIGHT and RAMOS, 2005; ARTHURS et al., 2008a, 2008b). Bt is considered ideal
for pest management because of its specificity to pests and safety to humans and
natural enemies of many crop pests. Typical agricultural formulations of Bt include
wettable powders, liquid concentrates, dusts, baits, and others, and have been
marketed under trade names such as Acrobe, Bactospeine, Certan, Dipel, Javelin,
Leptox, Novabac, Thuricide and Victory. In order to be effective, Bt must be eaten by
the feeding stage (larvae); it is ineffective against adult insects. Death can occur
within a few hours to a few weeks of Bt application, depending on the insect species,
age and the amount of Bt ingested. Although there are several different strains of Bt,
each with specific toxicity to particular types of insects, Btk is the most commonly
used against lepidopterous insects. Natural incidences of Bt were noted within PTM’s
native range in Bolivia (HERNANDEZ et al., 2005). Several strains were isolated from
agricultural soils, warehouses, and tubers infested with PTM. Moreover, some of the
isolates were shown to have equal or even greater toxicity compared with a standard
commercial strain (HD-1), suggesting more effective indigenous strains of Bt could
be developed for PTM control.
Bt has been widely tested to control PTM infestations under laboratory, field and
storage conditions. At least 2 crystal proteins of Bt have activity against PTM, e.g.
Cry1A(b) and Cry1B. Under laboratory conditions, PTM larvae are susceptible at
differing degrees to various Bt subspecies including kurstaki, thuringiensis, tolworthi,
galleriae, kenyae and aizawai, although the lethal concentration (LC50) required
increases with larval age (SALAMA et al.,1995a). For example, Btk (Thuricide HP)
applied at 200 mg/kg potatoes reduced PTM survival from egg to adult emergence to
0.4%, compared with PoGV (0.8-34.7% depending on dosage) or controls (32.5%)
(VON ARX and GEBHARDT, 1990). In other laboratory studies, dust formulations of Bt
(5000 IU/mg), along with permethrin (0.1%), prothiofos (1%) and rotenone (2.4%)
gave good protection of potato tubers against PTM infestations and were more
effective at controlling existing infestations compared with 1% chlorpyrifos
(HAMILTON and MACDONALD, 1990). In greenhouse and laboratory studies where Bt
was applied to the soil to protect seedlings or tubers in pots, it retained its potency for
up to 60 days (AMONKAR et al., 1979).
Bt has also been reported effective against PTM infestations under field
conditions. However, repeated applications are required because Bt is degraded by
UV light, and rain washes it onto the soil (SALAMA et al., 1995b). Three consecutive
applications of Bt (Bio-T) at 8 day intervals were required to control PTM in an
infested tomato crop in Israel (BROZA and SNEH, 1994). A high application volume
(500 l/ha) was used to bring the active ingredient into the tunnels in the leaves where
young larvae were mining. In field plot tests in India, foliar application of Bt
(Thuricide at 2-5 kg/ha) at 15-day intervals beginning 60 days after planting, were
almost as effective at controlling PTM infestations as parathion and carbaryl applied
to the soil surface and resulted in average tuber yields of 9.3-10.7 tonnes/ha,
compared with 6.7 tonnes without insecticides (AWATE and NAIK, 1979).
In many parts of the world, Bt and other non-chemical methods have been
evaluated for post harvest control of PTM, which is a serious pest in traditional, non-
refrigerated potato stores. In Yemen, KROSCHEL and KOCH (1996) evaluated a range
of low risk pesticides to protect tubers against PTM in storage. Bt mixed with fine
sand and dusted on tubers was completely effective when applied before PTM eggs
were deposited, but also controlled 96% of larvae that were already inside tubers. In
Egypt, another Bt preparation (Dipel 2X at 0.3% concentration) was reported to be
effective in storage, eliminating a PTM infestation compared with 100% infestation in
untreated controls 60 days after treatment (FARRAG, 1998). In Tunisia, an integrated
control approach comprising Bt applied at the beginning of the storage period in
combination with cultural control (early harvest) eliminated the reliance on parathion
sprays (VON ARX et al., 1987). In cases when tubers had a high initial infestation
(over 20%), Bt was replaced with a synthetic pyrethroid (permethrin). In tests in
Indonesia, tubers treated with Btk (Thuricide at 2g/liter) caused 79% larval mortality
after 4 months of storage compared with 58% mortality of larvae on foliage in a
screenhouse (SETIAWATI et al., 1999). In Peru, RAMAN et al. (1987) found Btk (Dipel)
effective in reducing feeding damage in storage when applied as a dust formulation.
Vegetable oil (1-2%) also reduced egg hatch but was phytotoxic, resulting in high
levels of tuber rotting. However, in other studies, Bt subsp. thuringiensis (0.2%
Bactospeine WP 16000 IU/mg) was reported ineffective at protecting tubers in
storage, resulting in as much tuber damage as untreated controls (DAS et al., 1992).
Other research suggests Bt may be improved through formulation with plant extracts
containing insecticidal properties. For example, extracts of Atropa belladonna (L.)
and Hyoscyamus niger L. and Solanum nigrum L. plants reportedly decreased the
LC50 of Bt against PTM from 82 ug/ml to 43, 31 and 40 ug/ml, respectively
(SABBOUR and ISMAIL, 2002).
Emerging research suggests new ways in which Bt toxins might be used to control
PTM. Studies have demonstrated the protective effects of Bt-modified potato lines on
foliage and tubers from PTM damage in field and storage conditions (DAVIDSON et
al., 2004; JANSENS et al., 1995; WESTEDT et al., 1998). Although planting transgenic
potatoes may provide very effective PTM control options for growers, public
acceptance of genetically modified food crops will be the ultimate deciding factor for
widespread use of Bt potatoes (see DOUCHES et al., 2008).
Entomopathogenic Nematodes (EPNs)

EPNs are insect-specific parasites in the genera Steinernema (Steinernematidae) and
Heterorhabditis (Heterorhabditidae) that are associated in an obligate manner with
symbiotic bacteria (Xenorhabdis spp. and Photorhabdis spp., respectively) which are
responsible for rapidly killing host insects. After entering a host insect, the infective
juveniles (IJs) of EPNs release their symbiotic bacteria. In addition to killing the host,
the bacteria digest host tissues, producing antibiotics to protect the host cadaver from
saprophytes and scavengers. When host nutrients are depleted, IJs are produced and
begin leaving the host insect. This stage is capable of immediately infecting a new
host or may persist for months in the absence of a host.
Research conducted on EPNs over the past five decades has demonstrated their
potential as biological control agents of a wide variety of insect pests (GREWAL et al.,
2005; GEORGIS et al., 2006). EPNs have been commercially developed for control of
several economically important insect species. However, their use for control of PTM
has only recently been investigated. Research conducted on EPNs and PTM by
LACEY et al. (unpublished data) demonstrated high mortality of larvae entering or
walking on EPN-treated soil. In simulated habitats, application of Steinernema feltiae
(Filipjev) at 104, 105 or 106 IJs/m2 resulted in 44, 83 and 95% mortality, respectively
in larvae that emerged from potatoes and migrated over nematode-treated soil.
Mortality of pupae was considerably less than that of larvae.
Fungi
Entomopathogenic fungi are effective microbial control agents of several insect pests
(GOETTEL et al., 2005), including some key pests of potato (LACEY et al., 1999;
WRAIGHT and RAMOS, 2005). However, there is limited research on the feasibility of
using fungi for PTM control. Laboratory studies on Metarhizium anisopliae (Metsch.)
Sorokin and Beauveria bassiana (Bals.) Vuillemin indicate potential for control of
larvae, particularly younger larvae (HAFEZ et al., 1997; SEWIFY et al., 2000). HAFEZ
et al. (1997) also demonstrated activity of B. bassiana against prepupae, pupae and
adult PTM. SEWIFY et al. (2000) reported that the combination of M. anisopliae and
PoGV resulted in synergistic larval control when a high concentration of the fungus
was used with a low concentration of the virus.
The fungus Muscodor albus Worapong, Strobel and Hess produces a mixture of
volatile organic chemicals that are active against a broad range of plant and human
pathogenic fungi and bacteria. Several researchers have demonstrated the activity of
M. albus against a variety of fungi, bacteria, and plant parasitic nematodes that cause
rot and plant disease (WORAPONG et al., 2001; STINSON et al., 2003; MERCIER and
SMILANICK, 2005; RIGA et al., 2008), but the insecticidal activity of this fungus has
only recently been investigated. In studies conducted by LACEY and NEVEN (2006),
PTM adults and neonate larvae were exposed to volatiles generated by M. albus
mycelia for 72 hours in hermetically sealed chambers at 25°C. Mean percent
mortalities in adult moths exposed to 15 and 30g of fungal formulation were 84.6 and
90.6%, respectively. (LACEY et al., 2008) demonstrated that development to the pupal
stage of PTM that were exposed as neonate larvae to 15 or 30 g of formulated M.

albus mycelia was reduced by 61.8 and 72.8%, respectively relative to controls. Five
day-old larvae in infested tubers that were exposed to 30 g of fungal formulation
responded with 84, 95 and nearly 100% mortality after 3, 7, and 10 days exposure.
Conclusions
Natural enemies including parasites, predators and pathogens can exert substantial
control of PTM populations, especially when little or no insecticide is used
(MATTHIESSEN and SPRINGETT, 1973; BRIESE, 1981; KROSCHEL and KOCH, 1994;
COLL et al., 2000). It is likely that no single natural enemy species will provide stand
alone control, but together they may help to regulate PTM in a complementary
manner throughout the growing season, in the various stages of the life cycle and at
various population densities. In addition to pest control, insect-specific pathogens
offer a variety of other benefits including safety for applicators, other natural
enemies, the environment and human food supply (LAIRD et al., 1990; HOKKANEN
and HAJEK, 2003). The integration of insect-specific pathogens and nematodes into
the potato agroecosystem will depend on their compatibility with other control agents,
including pesticides and the effect of environmental conditions on their infectivity
and persistence. Their successful utilization will require selection of effective
pathogen strains and development of formulations to improve storage, application and
persistence as well as careful timing of applications and a better understanding of how
they will fit into potato production systems. Their implementation will ultimately
depend on an increased awareness of their attributes by growers and the public
(LACEY et al., 2001).
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49
The potato tuber moth granulovirus (PoGV):

use, limitations and possibilities for field
applications
Marc Sporleder and Jürgen Kroschel
Abstract
A specific granulovirus (PoGV, Baculoviridae) infecting the potato tuber moth
(PTM), Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), is a prime
candidate for biological control. Given the controversy concerning the use of
chemical insecticides, PoGV has become an integral part of IPM approaches for PTM
in several developing countries that allow species-specific, narrow-spectrum
applications. This entomopathogen has been successfully applied to protect stored
potatoes and may also replace the application of chemical insecticides to growing
potatoes. Earlier experiments using the virus in the field were very promising but the
high number of infected larvae needed for efficient control hindered its subsequent
use in growing potatoes until today. Baculoviruses act very differently compared to
chemical pesticides and for their efficient use a comprehensive understanding of the
biology and ecology of the virus and its host is crucial. This paper describes several
attributes of PoGV by reviewing recent research results and outlines implications and
possible directions for further improving the strategies of using PoGV in the field.
The important aspects discussed are a) virus field persistence, b) the slope of the
virus dose-host mortality relationship, and c) changing host susceptibility to the virus
with larval age. In addition, modelling studies revealed that it is not possible to kill
many individuals of the pest population with an initial application and that
application of low dosages in frequent intervals during the vegetation season have a
much higher probability of depressing PTM populations over the longer term. We
propose this strategy that takes advantage of the ability of baculoviruses to replicate
and produce additional inoculum in the field. This strategy could suppress host
populations long-term with significantly reduced costs.
Introduction
Due to the many problems associated with the extensive use of chemical pesticides to
control the potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera:
Gelechidae), the use of entomophatogens in Integrated Pest Management (IPM)
strategies is of growing interest. One prime candidate of microbial biocontrol agents
50 The PTM granulovirus (PoGV)
is the naturally occurring granulovirus infecting the potato tuber moth (PoGV,
Baculoviridae). The virus, which is endemic to PTM, has been isolated from larvae in
various parts of the world (REED, 1969; BROODRYK and PRETORIUS, 1974; HUNTER et
al., 1975; ALCÁZAR et al., 1991; KROSCHEL et al., 1996b; SETIAWATI et al., 1999;
ZEDDAM et al., 1999). Possibly the virus accompanied the pest in its spread
throughout the world.
The use of PoGV as a biocontrol agent has been extensively studied in several
regions in the field and in storage (ALCÁZAR et al., 1992; BEN SALAH and AALBU,
1992; KROSCHEL et al., 1996b; MATTHIESSEN et al., 1978; REED and SPRINGETT,
1971; ARTHURS et al., 2008). The use of PoGV for controlling PTM in farmers’
rustic potato storerooms was very successful (ALCAZAR et al., 1992; ALI, 1991;
AMONKAR et al., 1979; DAS et al., 1992; ISLAM et al., 1990; SETIAWATI et al., 1999).
In Peru, the International Potato Center (CIP) initiated research on PoGV in the mid-
1980s and introduced the virus as an alternative to pesticides in their global IPM
program (ALCÁZAR et al., 1992; GELERNTER and TRUMBLE, 1999; RAMAN et al.,
1987). A simple mass-production and dust-formulation of the virus was developed
for protecting stored potatoes, and cottage-type enterprises were launched in Peru,
Bolivia, Colombia, Egypt, and Tunisia with optimization of virus production, quality
control and profitability (FUGLIE et al., 1993; WINTERS and FANO, 1997). Virus-
infected larvae are ground and mixed with talc at a rate of 20 larvae per kg talc in one
liter of water. The dried product was applied at a rate of 5 kg per ton of stored
potatoes. Such applications have provided high levels of control (usually >95%) and
significantly reduced tuber damage (ALCÁZAR et al., 1992; ALCÁZAR and RAMAN,
1992; DAS et al., 1998). Presently, PoGV is used in several countries for protecting
stored potatoes, but its current low cost propagation and formulation technique still
limits its potential application. Besides the common PTM, P. operculella, PoGV
infects the Guatemalan PTM species, Tecia solanivora (Povolny), but not the Andean
PTM species Symmetrischema tangolias (Gyen). The latter species is becoming
economically more important in the Andean region that also limits the use of PoGV
in that region.
Earlier field experiments in different countries, though the results were variable in
some cases, showed that virus applications might successfully suppress the host
population. In some cases applications gave similar results to chemical insecticide
applications. Applications resulted in a large production of secondary inoculum and
the virus could spread extensively to untreated areas (KROSCHEL et al., 1996b; REED,
1971; REED and SPRINGETT, 1971), and subsequent in-store infestation failed to
develop (BEN SALAH and AALBU, 1992). However, the high amount of virus-infected
larvae needed for field applications is a limiting factor. In addition, studies of PTM
field populations have, in some cases, revealed natural PoGV incidence levels as high
as 35–40% (KROSCHEL, 1995; LAARIF et al., 2003). Several authors have shown that
the infestation of potato tubers at harvest can be significantly reduced by effectively
controlling PTM on the foliage during the growing season (RADCLIFFE, 1982;
KROSCHEL, 1995; ARTHURS et al., 2008). Such observations encourage the
exploration of the microbial control potential of PoGV from the standpoint of
inundative augmentation in growing potato crops.
The PTM granulovirus (PoGV) 51
Baculoviruses, comprising two groups of virus, nuclearpolyhedrovirus (NPV) and

granulovirus (GV), are promising biological alternatives to chemical pesticides that
have been promoted for more than half a century. They are regarded as safe and
selective bioinsecticides, restricted to invertebrates, and have been used worldwide
against insect pests, mainly Lepidoptera (HUBER, 1986; FUXA, 1991; CUNNINGHAM,
1995). Some of their characteristics, such as specificity and safeness to non-target
organisms (GRÖNER, 1986; 1990), make them desirable agents in IPM programs,
specifically for species-specific, narrow spectrum insecticidal applications. In the
field, they replicate and persist in the environment and may suppress host populations
longer after application compared to chemical insecticides. However, their use as
microbial insecticides has not met their potential and only a few baculoviruses have
been commercially successful or have been used for large-scale insect pest
management. The main restriction against baculoviruses being developed as
biopesticides is their lower performance compared to chemical pesticides. The
problems associated with baculoviruses can be summarized as follows:
Baculoviruses differ fundamentally from chemicals, because they must be
ingested to cause infection, thereby killing the insects more slowly than chemicals
and they attack only certain stages of the host. Hence, the timing of applications is
important for successful control. To give appropriate recommendations to farmers in
different agro-ecological zones, the effect of the virus on the host as well as the life-
cycle of the pest must be well known. Furthermore, baculoviruses are vulnerable to
solar radiation, especially to the ultraviolet light wavelength of the solar light
spectrum, leading to rapid inactivation after application.
Technical and economic difficulties in mass production and poor virus yield and
insufficient quality control in commercial products might lead to unreliable field
results and virus supply. The limited host range of baculovirues might be a
disadvantage when a complex of pest species needs to be controlled. Strategies to
counteract some of the limitations of baculoviruses, especially their slow-killing
activity and inactivation due to UV, have been investigated and found promising.
UV-screens might be added to the formulation to increase the field stability of the
virus. Chemical or biological substances added to virus formulations and genetic
engineering of the viruses themselves to express insect toxins or hormones are
strategies to enhance the viral activity and increase the speed of killing. However, the
latter strategies are leading to reduced virus replication in the field.
For applying baculoviruses successfully as biopesticides it is important to
understand the nature of baculoviruses and to explore the possibilities to develop new
methods for their application. This paper presents recent research results on PoGV,
discusses advantages and disadvantages for virus field use, and outlines implications
and possible directions for further improving the strategies using baculoviruses for
pest control. The important aspects discussed are a) the PoGV field stability and
possibilities of using UV screens for improving virus stability, b) the slope of the
virus concentration-host mortality curve and its implication for justifying field
dosages, and c) changing host susceptibility to the virus with larval age. The results
on PoGV ecology were combined with theoretical pest population/phenology
modelling studies that might help, adjusting for appropriate management of the virus
differentiating between conditions in different agroecological regions.
Virus field stability (UV-stability)

The ultra-violet spectrum of solar radiation is considered to be the major
environmental factor affecting the field persistence of baculoviruses. Most destructive
is radiation in the UV-B region (280–315 nm); however, the UV-A (315–400 nm)
may also be critical in baculovirus deactivation (SHAPIRO, 1995). In this chapter we
discuss a) the kinetics of the inactivation of PoGV (inactivation curve), b) the
retention of activity in the field, and c) the possibility of protecting viruses from solar
radiation through formulation and ultraviolet adjuvants.
Kinetics of the photo-inactivation

Recent experiments by CIP indicated that PoGV degradation does not follow a
simple exponential curve (SPORLEDER, 2003, and unpublished data). In these
experiments, PoGV has been plated out and dried in petri dishes and then exposed to
natural radiation in different altitudes in the Peruvian Andes. Petri dishes were placed
on a special board, which allowed a continuous adjustment to the sun (Figure 1). A
spectroradiometer and a pyranometer were used to monitor energy of specific wave
lengths and total radiation energy during exposure. In different time intervals of
several minutes, depending on the radiation intensity, one dish was removed from
exposure and stored at 4°C for later use in bioassays. In the laboratory, the virus was
resuspended from the petri dishes in distilled water and the biological activity
determined in bioassays. Suspensions from a non-radiated petri dish in each exposure
experiment served to establish the reference probit line that was used to extrapolate
surviving virus granules of irradiated virus particles.
The two-component Hiatt model (HIATT, 1964) described well the relationship
between the accumulated amount of radiation energy (joules/m2) and the virus
activity retention rate (Figure 2). The model proposes that the inactivation is bi-
segmented with an initial steep decline followed by a gradual decay. The function of
the model is as follows:
activity rate = (1 − γ ) ⋅ e − (k1 + k 2 )⋅E + γ ⋅ e − k 2 ⋅E (1)
where E is the energy sum (joules/m2), k1 and k2 are constants estimated by the model
representing the first order velocities for photo-inactivation assuming the existence of
two viable states for a virus particle; particles of the second state have a greatly
reduced probability of infection, which is represented by the activity coefficient γ. In
other words, Hiatt’s model is based on the assumption that irradiated virus particles
may break into a second form with reduced sensitivity to radiation but with reduced
probability of infection.
Figure 1. Experimental set-up for studying the inactivation curve of PoGV. On the white board
petri dishes with dried virus deposits are exposed to the sun. The table could be adjusted
to the angle of the sun with the back crank. A spectroradiometer (lower left corner on the
board) and a pyranometer were placed on the table connected to a mobile logging station
(left side) to monitor total radiation energy and energy of specific wave lengths during
exposure. Petri dishes were removed in specific intervals of time.
In Figure 2, the energy sum, E, was calculated by accumulation of joules/m2
calculated from the measurements of the pyranometer (visible light, 400–1100 nm,
Figure a) and by using the energy in the erythermal band of 300-320 nm measured by
the spectroradiometer (Figure b). The model explained 66.6% and 65.8% of the
variation within the total sample, respectively. Considering energy in the 400-1100
nm band, the constant k1 had a value of 2.86×10-6, signifying that an energy of
242×103 joules/m2 alters 50% of PoGV particles from the first to the second viable
state (half-life energy, E½, which is calculated by the equation: E½=ln[k1]/2), and the
constant k2 a value of 4.96×10-7, signifying an half-life energy of 1.4×106 joules/m2 to
completely inactivate virus particles of either state one or two; i.e. due to radiation
energy virus particles are converted from the first to the second state 5.76 times faster
than the radiation energy needed to completely inactivate virus particles of either
state one or two. The probability of infection of the second state, i.e. the activity
coefficient, γ, was 0.65%. Consequently, the inactivation curve bends when the
original viral activity is reduced by more than 99%. The same figure for the energy in
the erythermal band of 300-320 nm are k1=0.086 (E½=8.05 joules/m2), approximately
4.4 times higher then k2=0.02 (E½=35.4 joules/m2), while particles of the second state
had a probability of infection of γ=2.36%.
Alternatively to the energy measured by the pyranometer or the energy in the
300-320 nm bands, energy of specific wave bands might be used to assess virus
inactivation curves. However, changes in the light spectrum are expected to alter the
speed of virus inactivation. Using the energy of a specific wave band of the light
spectra may provide only a reasonable approximation of the specific sunlight energy.
The data generated through this study are not enough to confirm the effects of
specific light spectra on PoGV inactivation; for such an assessment more data with
PoGV exposure to sunlight under a variety of changing light spectra need to be

generated or the virus needs to be exposed to a specific wave length of interest only.
In the study on baculovirus photo sensitivity, researchers focused mainly on the
UV-B (280-315 nm) spectra by using artificial light; however, it might be important
to also consider higher wavelength in future research. In the field, the UV-B portion
for example, might be less hazardous than UV-A wavelengths (315–400 nm),
because UV-B is more easily absorbed by various particles in the atmosphere (cloud
cover) and does not penetrate as deeply into the leaf tissue (epidermis) of plants as
UV-A. UV-A wavelengths reach the earth’s surface in quantities approximately eight
times greater than UV-B.
Several other authors (GLOGER, 1993; JONES et al., 1993; HUBER and LÜDCKE,
1996; MCGUIRE et al., 2000) observed similar bi-segmented inactivation curves for
baculoviruses, and the Hiatt model might provide a reasonable explanation of the
kinetics of virus inactivation. It can be concluded that even if initial inactivation of
the virus due to sunlight is relatively fast, a certain degree of activity remains that
might provide a source of infection long after virus applications and eventually cause
long-season suppression of the pest population through release of secondary
inoculums.
Retention of activity in the field

At the plant level the impact of solar radiation is greatly influenced by the crop
architecture and aspectual position of the virus on the crop. A proportion of the virus
particles is better protected and survives longer through shading by foliage, or
because the particles are hidden in crevices of the plant’s surface. However, deposits
of baculoviruses on foliage in a field environment retain activity for relatively short
periods, usually less than two weeks (JAQUES, 1975), but the half-life varies greatly
from as little as 10 hours to 10 days (reviewed by ENTWISTLE and EVANS, 1985;
YOUNG and YEARIAN, 1986). Viral activity can be completely lost in less than 24 h,
but mean half-life generally has varied from 2 to 5 days (JAQUES, 1985). KROSCHEL et
al. (1996a) studied the field persistence of PoGV in the Yemen highlands (>2300 m
a.s.l.) and observed a half-life of 1.3 days.
As discussed in the previous section, PoGV degradation does not follow a simple
exponential curve. SPORLEDER (2003) conducted PoGV field stability experiments
during two different times in the year in Oxapampa, Peru (1800 m a.s.l.), that showed
that initial inactivation is relatively fast, but the inactivation curve was curtailed when
about 90-95% of the viruses were inactivated (Figure 3). During the first experiment
(November) daily solar radiation was relatively constant during the course of
experiment (>800 Wm2 at noon). After the first day post application >90% of the
virus was inactivated with a half-life time, t½, of 0.22 days, while thereafter half-life
time increased to 2.24 days (Figure 3a). During the second experiment (December)
solar radiation was variable; radiation was high during the first day (>800 Wm2 at
noon), but low during the 2nd to 4th days because of clouds (<450 Wm2), and again
high during the 5th and 6th days (>800 Wm2 at noon) post application. Virus
inactivation during the first day was as fast as in the first experiment (t½=0.22 days),
while during the 2nd to 4th days, inactivation was minimal (t½=2.8 – 9.1 days).
Thereafter (5th and 6th days) the inactivation process had become more rapid again
with half-life times of 0.33 and 0.42 days, respectively, so that finally less then 0.2%
of the viral activity remained after the 6th day post application compared to 1.5% in
the first experiment.
A 2
400-1100 nm
1
E ½ = 242 x 103 joules
0
-1 E ½ = 1.4 x 106 joules
log-activity
-2
-3
-4
-5
-6
-7
0 2 4 6 8
6
accumulated energy (x 10 joules)
B 2
300-320 nm
1
E ½ = 8.1 joules
0
log-activity
-1
E ½ = 35.4 joules
-2
-3
-4
-5
-6
0 50 100 150 200 250 300
accumulated energy (joules)
Figure 2. Bisegmented inactivation curves of PoGV in response to exposure to natural sunlight

during ten series of exposure in La Molina, Peru, and in different altitudes in the
Peruvian Andes (data are pooled), in response to total energy recorded by a
pyranometer (400-1100 nm) (A) and to energy in the 300-320 nm erythermal band
recorded by the spectroradiometer (B). Lines represent predicted log-activities by the
two-component Hiatt model (equation 1) fitted to the data; scattered lines are 95%
confidence limits; energy that causes 50% inactivation (E½) either from state one to
state two or from state two to total inactivation are indicated. Virus particles of the
second state have a reduced probability of infection (0.65% in A and 2.36% in B).
Irregular baculovirus degradation in the field that differs from a simple

exponential curve can be explained by the following reasons: a) the inactivation
curve for solar inactivation of baculoviruses is bi-segmented, explained by the
possibility that there are two viable states for virus particles (see previous section), b)
a portion of virus might be well protected from solar degradation in crevices of the
potato foliage, and c) the intensity of solar radiation may fluctuate naturally, causing
further irregular decreases in the inactivation curve.
0.5
activity rate = (1 − 0 .083 ) ⋅ e − (3 .5 + 0 .31 ) t + 0 .083 ⋅ e − 0.31 t
A
0
-0.5 t½ = 0.22
log activity
-1
t½ = 2.24
-1.5
-2
-2.5
-3
0 1 2 3 4 5 6 7
time of exposure (days)
B 0.5
0
t ½ = 0.29
-0.5
t ½ = 2.78
t ½ = 9.14
log activity
-1
-1.5 t ½ = 0.33
-2
t ½ = 0.42
-2.5
-3
0 1 2 3 4 5 6 7
time of exposure (days)
Figure 3. Inactivation of PoGV due to solar radiation after its application to potato foliage in
Oxapampa (1814 m a.s.l.), Peru, in November (A) and December (B) 2000. Solid line in
A: bi-segmented inactivation curve fitted to the observed data; soft scattered line in B:
smoothed inactivation trend line; daily half-life times (t½ in days) are indicated. Scattered
lines represent the bi-segmented inactivation curve for virus exposed as dried deposits in
petri dishes to the sun. In November solar radiation was relatively constant during the
course of the experiment (>800 Wm2 at noon) while in December solar radiation was
variable (1st day: >800 Wm2 at noon, 2nd to 4th days: <450 Wm2 at noon, 5th and 6th
days: >800 Wm2 at noon). Source: SPORLEDER (2003)
However, this does not explain why the activity decreased again from the 4th to
the 6th day of exposure in the second experiment as rapidly as in the beginning of the
experiment, resulting in low virus activity retention at the end of the experiment,
while overall radiation was higher during the first experiment. HUBER and LÜDCKE
(1996) found that virus surviving the first irradiation pace of fast inactivation were as
sensitive to irradiation as un-irradiated virus when irradiation was intermittent for
several days. Such an effect may also play a role in the field, as an explanation of the
re-increased speed of virus inactivation in the second experiment. The existence and
the impact of such effects in the field must be further addressed in future research.
As a conclusion, it can be expected that initial inactivation of the field-applied

virus due to sunlight is relatively fast; however, a certain degree of activity might
remain for long periods that might provide a source of infestation long after virus
applications, causing a certain degree of long-season suppression of the pest
population.
Use of UV adjuvants
Research in formulation development for baculoviruses is still a developing subject
area. Many substances have been tested as sun screening agents to extend viral
activity in the field, while emphasis has been placed upon substances absorbing both
the UV-B and UV-A portion of the solar spectrum as well as upon antioxidants or
radical scavengers (reviewed by BURGES, 1998). Certain substances, especially
optical brighteners of the stilbene group, have enhanced baculovirus activity
(reviewed by SHAPIRO, 1995).
Several efforts were made to protect PoGV through formulation and ultraviolet
adjuvants (SPORLEDER, 2003). Different dyes (Congo Red, folic acid, the optical
brightener Tinopal LPW, the cosmetic sunscreen Uvitex), and several antioxidants
were examined in experiments exposing dried virus deposits mixed with the
adjuvants to natural sunlight. Additionally, the impact of the substances on viral
activity itself was assessed. Selective adsorbents (dyes) had no significant effect on
PoGV stability. Congo Red was even detrimental to viral activity. Optical brighteners
(Tinopal) increased viral activity 8 to 134-fold but did not increase viral stability. For
the optical brightener, higher mortalities obtained from irradiated virus compared to
PoGV alone were attributed to enhanced viral activity rather than to increased virus
persistence. Several antioxidants, which act as free radical scavengers, increased
virus stability 2 to 3-fold without influencing virus activity. However, PoGV in host-
derived material (macerated larvae) was best protected against radiation
(approximately 6-fold increased stability compared to purified PoGV) (Table 1).
Certain optical brighteners of the stilbene group (especially Tinopal LPW) have
enhanced viral activity and reduced the time taken to kill the host in various virus-
host systems (SHAPIRO and ROBERTSON, 1992; SHAPIRO and DOUGHERTY, 1994;
WEBB et al., 1996; FARRAR and RIDGWAY, 1997; LI and OTVOS, 1999; ZOU and
YOUNG, 1996). SPORLEDER (2003) confirmed enhancement of PoGV activity, while
PTM larvae died earlier. Extended activity in the field is desirable in baculovirus
formulations; however, if the larvae die at an early stage, the advantage of
baculoviruses, in that they replicate in the field so that the pathogen persists in the
environment with the possibility to suppress the host population long after its
application, is limited. Only high mortalities due to virus infection in a later larval
stage result in a subsequently greater quantity of virus in the pest environment.
Because potato plants tolerate some PTM damage, fast killing of PTM larvae in the
field is not as important as for protecting stored potatoes. The objective of PoGV
applications in the field should be more to suppress PTM populations over the long-
season, which might be achieved through infected larvae that carry great quantities of
virus into the environment, combined with efficient horizontal transmission.
Furthermore, optical brighteners are hazardous to the environment and can therefore
not be recommended for baculovirus formulations.
Virus inactivation by UV light can be attributed to a direct effect on DNA or
generation of highly reactive radicals, which are responsible for damaging cellular
DNA. Therefore, inhibition of virus inactivation may be possible by the use of
antioxidants or oxidative enzymes. IGNOFFO and GARCIA (1994) tested several
antioxidants and oxidative enzymes, which all provided some level of protection.
Radical scavenging may provide protection to baculoviruses, but its economical
feasibility must be evaluated in the future.
Table 1. Effect of several UV adjuvants on PoGV activity and UV-stability.
UV adjuvants Effect on PoGV activity Effect on PoGV stability
(LC50) (t½)
Selective absorbents (dyes)

Congo Red reduced activity no effect
Folic acid Neutral no effect
UVITEX increased variability no sig. effect
Optical brightners
Tinopal increased activity no effect (?)
(8-134 fold)
Antioxidants (free radical scavengers)

Propyl gallate not clear ~3-fold increased
Phenylthiocarbamide ~5-fold increased ~2-fold increased
Ascorbic acid Neutral ~0-2-fold increased
Peroxidase Neutral ~2-fold increased
Superoxide dismutase Neutral ~6- fold increased
Host derived material

Macerated larvae ~6-fold increased
Source: compiled from SPORLEDER (2003).
Slope of the mortality-concentration curve

Knowledge of the relationship between pathogen concentration (or dose) and the host
response (mortality) is essential for giving recommendations for field applications.
The slope of the Probit-regression is especially important for economic determination
and optimization of field dosages and the interpretation of field responses. The slope
of probit lines in granulovirus – host insect systems seldom exceeds a value of one.
The slope for PoGV and its host PTM derived from field and laboratory experiments
varies around 0.6 (SPORLEDER, 2003; SPORLEDER et al., 2005). In contrast to
chemical insecticides, which show steeper slopes in dose-effect relationships, PoGV
applications of doses causing over 95% mortality in the field are probably not
economically feasible. For example, an application of PoGV at a dose of 5×1013
occlusion bodies (OB)/ha in 500 litres of water as used by KROSCHEL et al. (1996b)
corresponds with approximately 10,000 virus-infected larvae/ha (one larval
equivalent equals approximately with 5×109 OB/larva). Using Probit-regression
curves derived from subsequent field experiments that dose would result in
approximately 85% mortality of neonate PTM larvae. In order to increase the efficacy
of the application to 95% or even 99% mortality a 10- fold and a 100-fold increase of
the dosage would be necessary, respectively. On the other hand, 10 or 100-fold
reduced rates would still result in 66% and 43% mortalities (see Figure 4 for visual
explanation). This gives options to use PoGV as a relatively inexpensive partial
suppression agent in potato fields through the use of low dosages/ha. In such an
approach the virus could be applied at short intervals, depending on the pest
population growth potential in different agroecological zones. Specific treatment
thresholds for such an approach still need to be determined.
A 7 B 100%
y = 0.6x + 1.23 80%
Probit mortality
6
mortality
60%
5
40%
4 20%
3 0%
0 2 4 6 8 10 0 2 4 6 8 10
log concentration (granules/ml) log concentration (granules/ml)
Figure 4. Concentration-mortality relationship between PoGV and its host P. operculella (neonate
larvae); (A) shows the Probit regression line with an average slope of 0.65 derived from
several leaf-disc assays, and in (B) probit mortalities are retransformed into percentage
mortalities. In (B) it is illustrated that for increasing mortality responses from
approximately 85% to appr. 95% a ten-fold increase of PoGV is necessary while a 10-
fold decreased rate produces a mortality of about 65%. For economic adjustment of field
rates it may be beneficial to apply PoGV in several intervals at lower rates instead of
targeting highest mortalities with one single application.
Changing susceptibility to PoGV with larval age

Baculoviruses attack only certain stages of their host, in particular the larval stage,
while the susceptibility to the virus decreases with larval age. Even though the older
and larger larvae consume more food, thereby ingesting more virus particles, virus
deposits on the foliage lose their efficacy against older larvae.
SPORLEDER et al. (2007) studied the relationship between mortality response
(LC50) of PTM to PoGV and larval age (larval weight) and found a linear relationship
between log LC50 and log larval weight. It could be demonstrated that the LC50 of
nine-day-old larvae, i.e. larvae that completed about 65% of their larval development
(113 DD >13.4°C), were 1,000-fold increased compared to neonate larvae. Increased
resistance in host larvae against baculoviruses with larval age is well documented for
several baculovirus-host insect systems (see review by EVANS, 1986) and has been
mostly attributed to increased body weight; however, ENGELHARD and VOLKMAN
(1995) showed that active sloughing of the midgut cells also affects developmental
resistance and might be more important than increased biomass. Since body weight is
difficult to use for predicting field responses and because increased resistance in
larvae might be due to systemic reasons (VOLKMAN, 1997; HOOVER et al., 2000;
MONOBRULLAH and NAGATA, 2000), SPORLEDER et al. (2007) used physiological
larval age as the explanatory variable for predicting virus effects in PTM. The
relationship between virus activity and larval age is illustrated in Figure a. The
activity curve curtails when the larvae passed about 30% of their total development
time. Thereafter, the activity decrease is quite drastic. Three-fold increased resistance
was observed in larvae that completed 37% of their development compared to
neonate larvae, while susceptibility to the virus was 30-fold, 110-fold and >1000-fold
decreased at accumulated development ratios of 0.45, 0.55, and 0.67, respectively.
Larvae that completed over 75% of their development were virtually resistant to
PoGV.
This knowledge can help quantify virus treatment effects on PTM field
populations for a given age structure of PTM at the time of the treatment. The stage-
age structure of pest populations, especially of pests having overlapping generations,
is hardly known and difficult to assess. However, phenology models developed for
PTM can be used to estimate the population structure and growth potentials for a
given temperature regime. Applying the model developed by SPORLEDER et al. (2004)
by using a constant temperature of 24°C, larvae would comprise about 41% of the
total population while eggs, pupae, and adults comprise 53%, 4% and 2%,
respectively when the age-stage distribution has stabilized (Figure b). The population
increases with a intrinsic rate of natural increase, rm, of 0.1106. The age-distribution
of larvae population alone is illustrated in Figure c.
Applying the virus-mortality response curve discussed in the previous section
(Figure 4) and considering reduced susceptibility to the virus with increasing larval
age, the proportion of individuals that might be successfully infected through virus
applications can be predicted. Considering an application of 5×1013 OB/ha in 500
litres of water (108 OB/ml), as used by KROSCHEL (1995) - which would infect 85%
of neonate larvae-, a proportion of 63% of the larval population, i.e. about 26% of the
total PTM population, would be successfully infected by PoGV (Figure C). The low
slope of the mortality response curve provides a proportional slide decrease in
mortality rate for a given reduced field dosage so that a relatively small decrease in
dosage provides no significant effect in response. A 5-fold reduced field dosage (1013
OB/ha, 73% mortality in neonate larvae) would still infect 51% of the larvae, which
is about 21% of the total population.
This illustrates how this knowledge can be used for interpreting potential field
responses and particularly for refining application parameters for optimizing the use
of virus inoculum. In the case of PTM, the dosage necessary to achieve more than
90% mortality (in neonate larvae) may be very high and difficult to achieve in
practice. Considering that it is not possible to kill all target pest stages, it may be
more cost effective to use lower dosages that still produce acceptable levels of
mortality. The information about the slope of the mortality regression line, decreasing
susceptibility of larvae with increasing age, and the age-stage distribution of target
pest populations can be used to estimate trade-offs between costs and achievable
mortality. However, since baculovirus recycles in infected larvae producing
additional inoculum in the field, the final effect on population development could be
higher than expected from the applied dosage as calculated here. The impact of this
secondary inoculum on target pest populations in the field has not been addressed in
the research on PoGV but needs to be studied for determining lower limits for
adequate virus field dosages.A further limiting factor in this modeling study is that
real populations fluctuate during the season also due to other reasons than
temperature (i.e. food availability, precipitation, etc.) and consequently the age-stage
structure of PTM populations also changes. Therefore, the stable age-stage structure
resulting from the model might not represent the real population structure in every
environment at any time of the season. A method to estimate the population structure
in potato fields might be helpful to improve the prediction.
Simulating PoGV field performance

Numerous attempts have been made to theoretically model virus epizootics and the
potential of entomopathogens, including viruses, on host populations in the field
(ANDERSON and MAY, 1980; 1981). Most of these models do not produce realistic
dynamics and infection levels because too many factors are involved in pest
population development and virus infection; however, they are valuable in exploring
the factors that could be important for pest management strategies using
entomopathogens.
The temperature driven phenology model developed for PTM provides
information about the pest population age-stage structure, so that mortality effects
due to PoGV, according to the mortality response curve and the decreased
susceptibility to virus with larval age, can be calculated for each age-class. The model
was used to simulate population development and effects of virus applications of
different rates and intervals. For simplification the model was run at a constant
temperature of 24°C. It was furthermore assumed that virus treatments have no effect
on the egg stage, which might be possible when the virus is deposited on the egg
surface so that neonates take up a lethal dose when consuming the chorion, and that
virus deposits are stable for one day only. Rate of spread of infection is a relevant
topic for realistic simulation of virus effects on field population; however, because
data on this topic are missing, this factor was also ignored.
At 24°C, larvae would comprise about 41% of the total population. A PoGV
application of 5×1013 granules/ha in 500 litres of water as used by KROSCHEL (1995)
would kill about 85% of neonate larvae; but would kill significantly reduced
proportions of older larvae (see Figure 5). A single application of such a field dosage
would not reduce populations long-term, but would delay population increase for
approximately 3-4 days (Figure 6), thereby causing irregularities in the population
development curve due to alteration in the population stage structure.
A 1
0.1
activity
0.01
0.001
0.0001
0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0
physiological age (accumulated development rate)
B
0.12
0.10 Egg
distribution rate
0.08
0.06
0.04
Larva
Pupa
0.02
Adult
0.00
0 10 20 30 40
age (days)
C 16%
14%
12%
distribution rate
10%
8%
6%
4%
2%
0%
0.0 0.2 0.4 0.6 0.8 1.0
physiological age (acc. dev. rate)
Figure 5. Effects of PoGV applications on P. operculella field populations. (A)

Relationship between the activity ratio of PoGV and physiological age of
P. operculella by using daily development rate summation (50% of larvae
complete their development when the value reaches 1). The line represents
prediction of the activity ratio (LC50-neonates/ LC50-in age x). (B) Stable age-
stage distribution of P. operculella at 24ºC. (C) Age distribution of P.
operculella larvae (bold line) and predicted portion killed by PoGV when
dosages of 5×1013 OB/ha (grey line) and 1013 OB/ha (scattered line) are
applied.
A second application at the same rate, seven days after the first application, which
targets forthcoming developing larvae that were not affected by the previous virus
application, again delayed population increase almost to the same extent as the first
application. Due to the additional proportion of larvae killed due to the second virus
application the oscillation of the population curve is heavier as after a single
application. The second application almost doubles the effect of the first application.
Alternatively, in a further scenario, the application of 5×1013 granules/ha was split
into 5 application of 1013 granules/ha in weekly intervals. A single application of 5-
times reduced application rates compared to the first scenario is expected to cause
about 73% mortality in neonate larvae and correspondingly reduced mortalities in
older age classes. The continuous elimination of larvae resulted in flatter population
development during the period of application as in the scenario with two applications
of 5×1013 granules/ha. Each application reduced the population by about 20-25%.
10
applications 1 - 10
9
natural increase
8
5x "10^13 OB/ha"
7
10x "10^13 OB/ha"
6
ln N
2x "5x10^13 OB/ha"
5
4 r m = 0.1106 1x "5x10^13 OB/ha"
0
0 10 20 30 40 50 60 70 80
Time (days)
Figure 6. Simulated natural populations increase of P. operculella at 24ºC (bold line) and effects
of PoGV applications on field populations. Four scenarios are simulated: [1] a single
application of 5×1013 OB/ha (bold scattered line), [2] two applications of 5×1013
OB/ha with the second application seven days after the first (timid scattered line), [3] a
series of 5 applications of 1013 OB/ha in 7-day intervals (grey solid line), and [4] a
series of 10 applications of 1013 OB/ha in 7-day intervals (light grey solid line).
After the fifth application, the population again increased almost linearly on a
log-scale, with a rate of increase of about 0.1104, which is essentially the same as for
population development without virus effects. Due to the permanent reduction of the
larval population over a period of 5 weeks, which is almost the time span required for
development of one PTM generation at 24°C (≈35 days), subsequent total population
increase did not show the oscillations as after a single application of a higher rate. As
a result, total population increase was delayed by about 10-11 days. Continuation of
the virus application at this rate for another 5 weeks again delayed population
increase for another 10 days. These simulation results are illustrated in Figure 6.
In reality the effect of virus applications can be expected to be higher than
simulated here because transmission of virus within the field population was
neglected. However, these results show that repeated applications of low rates at
relatively short intervals (7 days, depending on temperature) are more likely to lead
to long-term reduction of pest populations than a single application of a higher rate.
Under cooler conditions, where development of PTM populations is slower, fewer
applications and wider application intervals might suffice to successfully reduce pest
populations. Horizontal virus transmission within the population from living virus-
infected larva to another can be expected to be infrequent; the larvae are not
cannibalistic, the virus does not appear to be transmitted via contaminated faecal
material, larvae feed in their individual mines and do not come in contact with each
other habitually, and bioassays conducted with individually versus batch-reared
larvae have produced nearly identical results (Sporleder, unpublished). However,
horizontal transmission occurs from virus-died larva (cadavers) to new up-coming
L1-larvae; infected larvae release new virus inoculum when they die that might be
ingested by other larvae increasing the infection level in the field. Because more
cadavers of infected larvae are produced when low dosages of virus are repeatedly
applied compared to single applications of a high rate, it can be expected that
transmission of virus in the field is also more efficient using this approach. The
application itself also adds to the final cost when using virus in the field and farmers
would prefer a single application for controlling the pest instead of repeated
applications. However, the approach of repeated low-virus dosages might be
especially suitable for farmers in developing countries, in situations where farm
families produce potatoes on limited areas of land and are lacking monetary resources
instead of workforce to increase their production.
Summary and outlook

Given the controversy concerning the use of chemical insecticides, biopesticides
based on specific microorganisms, including the endemic granulovirus infecting
PTM, have become an integral part of IPM approaches for PTM in several
developing countries. PoGV has proved to be very effective in protecting stored
potatoes and may also replace the application of chemical insecticides to growing
potatoes. Earlier experiments using the virus in the field were very promising;
however, several limitations to PoGV (i.e. high numbers of larvae needed for field
applications, inappropriate formulation, low field persistence, etc.) compared to
cheaper chemical pesticides prevented the virus from being used on a wide-scale
basis in growing potatoes today.
However, there is no doubt that microbial biopesticides can play a key role in
controlling pests and make a significant contribution to reducing chemical inputs.
Since the PTM infestation in stored potatoes can be prevented through reduced PTM
populations in the field there is still considerable interest in using PoGV as a highly
specific biological control agent for PTM in growing potato fields. The advantage of
using baculoviruses is that they leave no toxic residues, create little or no
environmental pollution, and are compatible with many IPM practices, parasitoids
and predators. A further advantage of baculoviruses is that they replicate in their
hosts and persist in the environment due to horizontal and vertical transmission,
which may cause long-term suppression of pest populations.
The mode of action of baculoviruses is very different compared to that of
chemical pesticides. For efficient use of baculovirus and its integration in pest control
strategies, especially for developing long-term control strategies, a comprehensive
understanding of the biology and ecology of the virus and its host is crucial. For
entomopathogenic viruses, the ecology of baculoviruses is by far the best studied
(CORY and MYERS, 2003); however, for defining field application systems and spray
regimes, each baculovirus-host system needs to be studied individually. This paper
discusses the biological properties of PoGV and explores the possibilities for
improving current application methods. The advantages, disadvantages and
possibilities can be summarized as follows:
a) Low virus stability in the field: After application, the virus is rapidly
inactivated due to solar radiation; depending on radiation energy half-life times can
be shorter than one day (but during cloudy days, inactivation is significantly
reduced). KROSCHEL et al. (1996a) considered the low field persistence as the main
limiting factor for applying PoGV in the field that makes a second application
necessary. However, there is evidence that PoGV degradation does not follow a
simple exponential curve. A certain portion of the applied virus particles might
remain active for longer periods, either because they are shaded by the foliage or
protected in cavities. Furthermore, according to the theory of two viable states of
virus particles discussed above, it might be possible that a certain proportion of
activity remains in virus particles of the second state for much longer periods. Due to
increased half-life of particles in the second state the inactivation curve curtails when
about 95% of the activity is lost. Therefore, a certain portion of virus particles remain
active for long periods of time after application and may contribute to long-term
suppression of the pest and further dissemination of the pathogen. PoGV particles
released from or recycled in infected cadavers will persist for longer periods of time
than purified virus because they are protected at a certain level against UV.
Formulations that might protect the virus from solar inactivation have been
considered as a key approach for increasing the efficacy of PoGV in the field.
Antioxidants added at rates of 1% to the virus spray suspension increased virus
stability 2 to 3-fold while selective adsorbents (dyes) had no significant effect on the
stability of PoGV. UV protection through formulation may not result in substantial
returns because it implies significant additional costs to the final product. The use of
dyes might imply that the final product becomes hazardous to the environment.
Optimizing application dates and the spray dosages to increase the level of infection
that result in a subsequent load of virus in the field (cadavers) that produces a
secondary wave of infection seems more viable for increased PoGV efficacy then
adding UV screens to the formulation.
b) Low slope of the mortality-dose regression line: As for many other
baculoviruses, the slope of the mortality-dose relationship for the PoGV-PTM system
is relatively low. This implies that a proportionately low increase in mortality rate
will be achieved for a given rise in dose so that even a high increase of the dose
provides no significant increase in response. Although high mortalities are difficult to
achieve, the low slope provides the advantage that field doses might be decreased
without significant reduced mortality responses. In practice, it will be difficult to
achieve mortalities over 90% in neonate larvae because the number of infected
larvae/ha would be too high to be economically feasible (probably over 10,000 LE/ha
would be necessary), while moderate mortalities might be achieved with extremely

reduced numbers of larvae (1,000 or even 100 larvae only).
c) Decreasing susceptibility with larval age: PoGV affects the larval stage only
and within the larval stage resistance in larvae increases significantly with age.
Therefore, PoGV applications must be directed against 1st instar larvae, as well as
against the eggs (emerging larvae may take up a lethal dose with consumption of the
egg chorion as long as the virus was deposited on the egg surface).
These aspects make it clear that a knock-down effect on the PTM population is
hard to achieve by the use of PoGV in the field. It needs to be accepted that it is not
possible to kill all individuals of the pest population with an initial application, even
if virus stability would be increased through formulations with UV screens.
Improving PoGV stability through formulation should not be considered as the first
solution to improving virus efficacy in the field. A more cost effective approach
could be to use significantly lower dosages, which still give acceptable mortalities,
thereby applying the virus more often during the cropping season for frequently
targeting new developing young larvae. As with other modeling studies on microbial
control agents (ANDERSON, 1982) the modeling results presented here indicated that
for long-term control of the pest population and for inoculative augmentation,
subsequent applications causing moderate infection in the host population may be
better than a single hit with greater virulence.
The modelling applied here does not include the effects of secondary inoculum;
however, in earlier experiments using the virus in the field, all authors (KROSCHEL et
al., 1996b; REED, 1971; REED and SPRINGETT, 1971) reported that PoGV applications
resulted in a large production of secondary inoculum and extensive spread of the
virus to neighbouring potato fields. Therefore, there is scope for increasing the level
of virus infection in PTM field populations resulting in long term reduction of PTM
populations below the damage threshold by applying the proposed strategy. However,
the impact of secondary inoculum of PoGV and the beneficial role of the soil as an
important reservoir of the virus on subsequent PTM generations has not yet been
studied in detail. Considerable impact could be expected, because PoGV applications
to the soil surface before harvest reduced tuber infestation at harvest by 73%, and
subsequent in-store infestation failed to develop (BEN SALAH and AALBU, 1992).
Further field research should address PoGV persistence and transmission to allow
for more extended modelling that could help in optimising the PoGV application
strategy. Also more knowledge is necessary in order to understand migration of PTM
into growing potato fields and changes in the population structure during the potato
growing period. In subtropical regions, PTM may produce >9 overlapping
generations per year and populations thus comprise individuals of mixed-ages. In
regions with lower temperatures and shorter growing seasons, however, only 2 to 3
generations are possible and populations increase most rapidly during the period of
maximum vegetative growth of the potato. In these regions, the age structure of the
population is expected to be more homogeneous, and this may allow for more
efficient targeting of young larvae.
d) Host specificity: As is for most baculoviruses, and especially the
granuloviruses, PoGV is highly host specific. It is restricted to P. operculella, which
is the primary host, but might also cause infection in another PTM species, i.e. T.
solanivora. Its host-specificity is desirable to support an overall IPM program,
especially when an ecologically sensitive area has to be treated. It makes selective
control possible, leaving the rest of the fauna, including beneficial arthropods present
in an ecosystem, undisturbed. Therefore, PoGV is a very promising candidate to
control PTM in regions where the pest is the only significant pest of potatoes, i.e.
North Africa, the Middle East, Central and Southeast Asia.
Acknowledgements
The research presented here have been funded by the Bundesministerium für
wirtschaftliche Zusammenarbeit und Entwicklung, BMZ (German Federal Ministry
for Economic Cooperation and Development), Germany.
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72
73
Biological control of potato tuber moth by

Apanteles subandinus Blanchard in New
Zealand
Tim J. B. Herman
Fruitfed Supplies, P.O. Box 322, Hastings, New Zealand
Abstract
In New Zealand the control of the potato tuber moth (PTM), Phthorimaea operculella
(Zeller) is dominated by regular applications of broad spectrum insecticides through
the peak of the growing season and cultural control strategies, such as well structured
hills. A larval parasitoid, Apanteles subandinus Blanchard (Hymenoptera,
Braconidae), was introduced into New Zealand in the 1960s and has become the key
parasitoid of PTM. However, the species has not been considered to have a major
controlling influence on PTM populations. The objective of our study was to
investigate parasitism rates in order to find out about the relative importance of A.
subandinus in an IPM program for PTM. Potato leaves infested with PTM larvae
were collected from commercial crops under different pesticide programs as well as
from untreated potato plots. Samples were collected in three regions and over 5
potato growing seasons. PTM larvae collected were reared on potato tubers to
determine parasitism rates. A. subandinus was the dominant parasitoid recovered
from the PTM larvae collected. In addition, Diadegma semiclausum (Hellén)
(Hymenoptera, Braconidae), a parasitoid of diamondback moth (Plutella xylostella
L.), was often reared in low numbers from PTM and Trigonospila brevifacies
(Hardy) (Diptera, Tachinidae), a parasitoid of light brown apple moth Epiphyus
postvittana (Walker) was occasionally recovered. Parasitism of PTM larvae in potato
foliage of commercial crops fluctuated within and between seasons and between
regions, ranging from less than 10% to more than 80%, and averaged 39.8% (37.9-
41.2%) in the three regions surveyed. Parasitism was higher on the research stations
without insecticide applications, averaging 85%. This suggests that insecticide
applications had a greater impact on parasitoid populations than on PTM host
populations and that the high parasitism potential of A. subandinus should be
integrated into an IPM program for PTM.
Introduction
The potato tuber moth (PTM), Phthorimaea operculella (Zeller), is an important pest
of potatoes in New Zealand because of its high reproductive potential and the
inability of insecticides to prevent tuber infestation (FOOT, 1979). PTM populations
peak about the height of summer, in the period January to March. The timing and
74 Biological control of PTM by Apanteles subandinus
height of the peak is very much dependent on hot temperatures and dry conditions.
Control of this pest is dominated by applications of broad spectrum insecticides at
10-14 day intervals, from late December to March when PTM pressure is highest and
cultural controls, e.g. well formed hills and use of irrigation, embedded in standard
crop management practices (HERMAN, 1997). Since PTM is an introduced pest to
New Zealand, classical biological control was considered with the introduction of 17
parasitoid species over a six year period in the 1960s. The importation and release of
the parasitoids was somewhat irregular and suboptimal with shipments of parasitoids
being in poor condition on arrival and only limited rearing being done before the
parasitoids were released (FERGUSON, 1989). Out of the 17 species, only Apanteles
subandinus Blanchard (Hymenoptera, Braconidae) was successfully established and
spread within New Zealand (FOOT, 1979). FOOT (1979) assessed parasitism in PTM
infested tubers over a ten month period from late summer to early spring, outside the
period when PTM populations peak. Parasitism rates of overwintering PTM larvae
averaged 45% and it was concluded that this may delay the PTM population growth
at the beginning of the potato season by about 3 to 4 weeks (one generation). Since
then no detailed study of the impact of A. subandinus on PTM populations and its
contribution to PTM control have been carried out. The aim of the research presented
here was to assess the parasitism of PTM larvae by A. subandinus and other
associated parasitoids in potato crops in order to determine the potential efficacy and
contribution of parasitoids to an overall IPM program for PTM.

PTM population dynamics in potato were studied over five growing seasons in three
regions of the North Island (HERMAN, 2008, this volume). As part of this research,
PTM larvae occupying foliar mines were collected from 3 to 6 commercial,
insecticide-treated potato crops in each region and each season and taken back to the
laboratory for rearing and determination of percent parasitism. In addition, PTM
larvae were also collected from research plots of the Crop & Food Research stations
in Pukekohe and Hawke’s Bay over three consecutive growing seasons. The plots
were part of a wider PTM IPM research project and received no insecticide
applications. Variation in PTM infestation over the three seasons and between
seasons did not allow collection of the same number of PTM larvae each week, to
determine parasitism rates. Therefore parasitism rates were sometimes derived from
less than 10 larvae per week. Potato leaves with occupied mines were placed in small
(75 ml) vented plastic containers along with a small piece of potato tuber (for
supplementary food) and some sawdust as a pupation substrate. The samples were
kept in incubators or on laboratory benches (variable temperatures) until a moth or
wasp emerged. Parasitoids were identified and percent parasitism determined per
sampling date. Because a high proportion of up to 50% of the collected PTM larvae
died from unknown reasons and independently from regions and seasons, the
parasitism rate was calculated as percent of parasitoids developed from all insects
reared from each sample.
Biological control of PTM by Apanteles subandinus 75
Results
Apanteles subandinus was the most dominant parasitoid in all three regions and the
five seasons surveyed followed by Diadegma semiclausum (Hellén) (Hymenoptera,
Braconidae) and Trigonospila brevifacies (Hardy) (Diptera, Tachinidae), two other
parasitoids recovered from the PTM larvae. D. semiclausum was originally
introduced for biological control of the diamondback moth Plutella xylostella L. It
was collected in small numbers most of the time and reached only a higher parasitism
rate of 24% at one location. T. brevifacies, which was introduced for biological
control of light brown apple moth Epiphyus postvittana (Walker), was only
occasionally reared from PTM larvae.
In Matamata, parasitism varied from 64%, 44% and 24.3% over the three seasons
from 1996 to 1998, respectively (Table 1), with an average of 40% parasitism per
year. In 1996/97 parasitism was initially less than 20% in early February, but
increased to between 70% and 80% in late March to early April as the crop matured.
Most of the PTM samples collected in 1997/98 in Matamata were of less than 10
larvae. At sampling dates with collections of more than 10 larvae parasitism ranged
between 40% and 76%. In the 1998/99 season parasitism did not exceed 37% in
collections of more than 10 larvae.
In Opiki an average parasitism of 42.9% per year was achieved, however, a high
variation between the 1997/98 and 1998/99 seasons was observed with parasitism of
57.4% and 28.4%, respectively (Table 2). In 1997/98 season, parasitism fluctuated
around 58% through February and peaked at 86% in late March, before declining to
16% in May. In the following season (1998/99) parasitism was low for the most of
the season with values less than 25% and only in late March-early April parasitism of
58% was achieved.
In Hawke’s Bay, the third region, PTM larval collections were generally very
low. A mean parasitism over the two seasons of 45.2% was recorded (Table 3),
32.8% in 1999/2000 and 58.8% in 2000/2001. In both seasons parasitism peaked in
March.
Parasitism of PTM was found to be much higher on research plots on research
stations in Pukekohe and Hawke’s Bay, where no insecticides were applied, with
parasitism averaging 85% over three years (Table 4). In Pukekohe during the 1996/97
season parasitism was between 80 and 100% from February to mid March and
dropped to 75% in late March and 29% in early April. Mean parasitism over one
season was 84.3%. At Lawn Road research station, Hawke’s Bay, only four
collections of PTM larvae were made in the 1999/2000 and 2000/01 season.
Parasitism averaged 89.2% and 82.8% in each season, respectively.
In general, in all regions, parasitism was dominated by A. subandinus and D.
semiclausum was collected only in small numbers. However, one exception was
found in Opiki in 1998/99 where parasitism was dominated by D. semiclausum from
late January to early March (Figure 1). A. subandinus was not reared from collected
PTM larvae until late February and did not become the dominant parasitoid until late
March. A. subandinus remained dominant until the foliage died.
Table 1. Percent parasitism of PTM larvae in foliage collected from potato crops in Matamata,
Waikato, over three seasons.
Season
96/97 97/98 98/99
Month/week No. PTM Parasitism No. PTM Parasitism No. PTM parasitism
larvae (%) larvae (%) larvae (%)
Dec 2 7 14.3
3 8 37.5
4 20 20.0
Jan 1 4 50.0
2 8 50.0 2 100.0
3 2 0.0
4 23 39.1 8 12.5
Feb 1 24 16.7 58 75.9 56 2.6
2 47 2.1 51 43.1
3 14 42.9 13 69.2 360 9.7
4 2 100.0 8 37.5
Mar 1 3 66.7 1 0.0 511 36.6
2 7 100.0 1 0.0
3 1 0.0
4 20 80.0 12 66.7
Apr 1 38 76.3 4 25.0
2 40 70.0 37 3.6
3 15 86.7
4 10 60.0 66 5.4
Mean* 20 63.8 13.8 44.0 148.6 24.3
*Parasitism values of zero were omitted from the calculation. Blank spaces indicate that no larvae have been
collected at this week.
Table 2. Percent parasitism of PTM larvae in foliage collected from potato crops in Opiki,
Manawatu, over two seasons.
Season
97/98 98/99
Month/week No. PTM larvae Parasitism (%) No. PTM larvae Parasitism (%)
Jan 2 15 0.0 1 0.0
3 15 46.7 3 0.0
4 6 16.7
Feb 1 12 25.0
2 101 56.4 4 25.0
3 36 66.7 32 15.6
4 71 52.1 29 13.8
Mar 1 133 24.8
2 11 81.8 274 17.9
3 16 87.5 388 25.0
4 70 85.7 170 57.6
Apr 1 214 58.4
2 33 24.2 247 32.8
3
4 63 15.9
Mean* 43.1 57.4 116.4 28.4
* Parasitism values of zero were omitted from the calculation. Blank spaces indicate that no larvae
have been collected at this week.
Table 3. Percent parasitism of PTM larvae in foliage collected from potato crops in Hawke’s Bay
over two seasons.
Season
99/00 00/01
Month/week No. PTM larvae Parasitism (%) No. PTM larvae Parasitism (%)
Jan 2 1 0.0
3 2 50.0 32 78.1
4 16 18.8 8 50.0
Feb 1 31 19.4 6 33.3
2 6 0.0 17 52.9
3 5 0.0
4 9 55.6
Mar 1 5 60.0 5 80.0
2 8 37.5
3 13 30.8
4 29 27.6
Apr 1 7 14.3
2 7 14.3
3 10 0.0
Mean* 10.6 32.8 13.6 58.7
*Parasitism values of zero were omitted from the calculation. Blank spaces indicate that no larvae
have been collected at this week.
Table 4. Percent parasitism of PTM larvae in foliage collected from potato crops on two research
stations over two seasons.
Pukekohe RS Lawn Rd RS, Hawke’s Bay
96/97 99/00 00/01
Month/week No. PTM Parasitism No. PTM Parasitism No. PTM Parasitism
larvae (%) larvae (%) larvae (%)
Jan 3 70 89.5
4 68 85.0
Feb 1 2 100.0
2 18 83.3 97 62.9
3 85 90.6
4 11 81.8 70 81.4
Mar 1 13 100.0
2 13 92.3 69 91.3 97 93.8
3 17 94.1
4 24 75.0
Apr 1 38 28.9 114 96.5
2
3
4
May 1 74 87.5
Mean* 24.6 82.9 81.8 89.2 83.0 82.8
*Parasitism values of zero were omitted from the calculation. Blank spaces indicate that no
larvae have been collected at this week.
60
% parasitism
A. subandinus
D. semiclausum
percent parsitised
40
20
0
12-Jan 26-Jan 9-Feb 23-Feb 9-Mar 23-Mar 6-Apr
Figure 1. Parasitism of PTM larvae in foliage collected from potato crops in Opiki, Manawatu,
during 1998/99, by Apanteles subandinus and Diadegma semiclausum.
Discussion
Biological control of PTM by the introduced parasitoid A. subandinus in New
Zealand has previously been considered to have little economic impact on
populations of the pest (FOOT, 1979; SANKARAN and GIRLING, 1980; FERGUSON,
1989). However, an assessment of PTM parasitism in New Zealand was only made
once from infested tubers during the period from late summer through to early spring
(FOOT, 1979). Additionally, parasitism in tubers and foliage might be quite different
because larvae are more protected in tubers (FERGUSON, 1989). BRIESE (1981) found
that A. subandinus and other parasitoids readily attacked PTM larvae in tubers in the
laboratory, but achieved less than 40% parasitism in tubers in the field, compared to
up to 90% parasitism in the foliage and concluded that searching behavior and host
accessibility in the field could be a reason for this. KEERATI-KASIKORN (1981)
reported that A. subandinus locates hosts in a random manner and found only 0.8% A.
subandinus parasitism in tubers in the field. Studies by KROSCHEL and KOCH (1994)
also showed for the larval parasitoid Diadegma molliplum Holmgren and the egg
parasitoid Chelonus phthorimaea Gahan that parasitism is higher in foliage but that
parasitism also takes place in tubers with a parasitism of up to 36% and 14% for these
two parasitoids.
A. subandinus has been well studied in Australia. It established across a wide
range of climatic regions from Tasmania (temperate, maritime) to tropical
Queensland, and was considered the most wide-ranging and effective of the three
PTM parasitoids introduced to Australia, with parasitism of 70% or more of PTM
larvae (CALLAN, 1972). Laboratory studies in California by CARDONA and OATMAN
(1975) found that A. subandinus prefers low relative humidity. This could be
supported by KFIR (1981) who reported that in South Africa A. subandinus takes
advantage of its greater fertility and longevity under low humidity conditions to be
the dominant parasitoid early in the growing season. Later in the season other
parasitoids become dominant. Similar observations have been reported by HORNE
(1990) from Victoria, Australia, where A. subandinus also dominates only early in the
season.
Our findings suggest that insecticide applications could be a reason for the erratic
performance of A. subandinus in New Zealand. A survey of New Zealand potato
growers showed that the broad spectrum insecticides methamidophos and lambda-
cyhalothrin are the most commonly used products for PTM control (HERMAN, 1997).
HORNE (1990) reported that a single application of methamidophos suppressed PTM
parasitism but did not completely suppress moth infestation. An additional three
insecticide applications effectively reduced parasitism to zero. The impact of
insecticide applications on parasitism was very apparent in Matamata in the 1998/99
season which was warm and dry and therefore ideal for PTM population growth. The
growers responded by increasing the number of insecticide applications which
resulted in the lowest parasitism found over three consecutive years.
Classical biological control of PTM in potato has been successfully implemented
in various countries through the introduction and naturalization of mainly three
species including A. subandinus. In Zimbabwe, releases have been successful and
PTM was eliminated as a significant potato pest (MITCHELL, 1978). In Australia,
although a successful naturalization of the parasitoids was achieved, their failure to
control PTM essentially related to the continued use of broad spectrum insecticides.
The efficacy of introduced parasitoids in Australia first became obvious after a full
IPM program for all potato pests was established with reduced or zero applications of
insecticides in potato crops (HORNE and PAGE, 2008). Our studies suggest that A.
subandinus has a great potential to reduce PTM infestations in potato crops in New
Zealand if the parasitoid is integrated into an IPM program, which encourages its
population growth for effectively parasitizing PTM larvae.
References
Australian Entomological Society 20: 319-326.
CALLAN E 1972. Changing status of the parasites of potato tuber moth
Phthorimaea operculella (Lepidoptera, Gelechiidae) in Australia.
Entomophaga 19: 97-101.
CARDONA C and ER OATMAN 1975. Biology and physical ecology of Apanteles
subandinus Blanchard (Hymenoptera: Braconidae), with notes on
temperature responses of Apanteles scutellaris Museback and its host, the
potato tuberworm. Hilgardia 43: 1-51.
FERGUSON AM 1989. Phthorimaea operculella (Zeller), potato tuber moth
(Lepidoptera, Gelechiidae). In: Cameron PJ, Hill RL, Bain J and WP Thomas
(eds.). A Review of Biological Control of Invertebrate Pests and Weeds in
New Zealand. Technical Communication 10, CAB International, Wallingford,
UK, p 119–128.

(Lepidoptera, Gelechiidae) at Pukekohe. New Zealand Journal of Zoology 6:
623-636.
HERMAN TJ 1997. Integrated pest management for potato tuber moth, a
technology for business growth project. CropInfo Report No. 443. New
Zealand Institute for Crop & Food Research. Lincoln, New Zealand. 35p.
HERMAN TJ 2008. Integrated Pest Management of Potato Tuber Moth in New
Zealand. In: Kroschel J and L Lacey (eds) Integrated Pest Management for
the Potato Tuber Moth, Phthorimaea operculella Zeller – a Potato Pest of
Global Importance. Tropical Agriculture 20, Advances in Crop Research 10.
Margraf Publishers, Weikersheim, Germany. 119-126.
HORNE PA 1990. The influence of introduced parasitoids on the potato tuber
moth, Phthorimaea operculella (Lepidoptera, Gelechiidae) in Victoria,
Australia. Bulletin of Entomological Research 80: 159-163.
HORNE PA and J PAGE 2008. Integrated Pest Management dealing with the
potato tuber moth and all other pests in Australian potato crops. In:
Kroschel J and L Lacey (eds) Integrated Pest Management for the Potato
Tuber Moth, Phthorimaea operculella Zeller – a Potato Pest of Global
Importance. Tropical Agriculture 20, Advances in Crop Research 10. Margraf
Publishers, Weikersheim, Germany 111-117.
KEERATI-KASIKORN M 1981. Initial studies towards the implementation of a pest
management system for the control of potato tuber moth (Phthorimaea
operculella Zeller) in south-eastern Queensland. PhD thesis. University of
Queensland.
KFIR R 1981. Fertility of the polyembryonic parasite Copidosoma koehleri,
effect of humidities on the life length and relative abundance as compared
with that of Apanteles subandinus in potato tuber moth. Annals of Applied
Biology 99: 225-230.
KROSCHEL J and K KOCH 1994. Studies on the population dynamics of the potato
tuber moth (Phthorimaea operculella Zeller) (Lepidoptera, Gelechiidae) in
the Republic of Yemen. Journal of Applied Entomology 118: 327-341.
MITCHELL BL 1978. The biological control of potato tuber moth Phthorimaea
operculella (Zeller) in Rhodesia. Rhodesia Agricultural Journal 75(3): 55-58.
SANKARAN T and GIRLING DJ 1980. The current status of biological control of the
potato tuber moth. Biocontrol News and Information 1 (3): 207-211.
81
Spinosad: a new biopesticide for Integrated

Pest Management of the potato tuber moth in
Tunisia
Khamassy Nouri and Issam Arfaoui
Institut National de la Recherche Agronomique de Tunis (INRAT), Potato
Research Program, Tunisia
Abstract
In Tunisia, conventional chemical pesticides used to limit the potato tuber moth
(PTM), Phthorimaea operculella (Zeller), attacks in rustic potato stores have shown a
decrease in efficacy in the last years. Biological treatments such Bacillus
thuringiensis subsp. kurstaki, while effective, are limited due to their high costs and
the technical skills required by the user for its application. Recently, a new
biopesticide based on Spinosad, a product derived from Sacchropolyspora spinosa
(Mertz and Yao), was evaluated as a part of an integrated PTM management program
included in the Improved Non-Refrigerated Potato Storage System. During 2003 to
2005, Spinosad efficacy was compared to the insecticide Deltamethrine. Additional
tests under high infestation pressure were undertaken to determine Spinosad efficacy
depending on storage duration and potato cultivar. Spinosad showed better efficacy
than Deltamethrine. After 3 months of storage, PTM infestation in Spinosad-treated
tubers reached only 5% compared to 80% and 90% in the Deltamethrine and control
treatment, respectively. Efficacy of Spinosad was similar when applied on different
potato cultivars. Further, a 30-month old product was almost effective as a new
product. The adoption of Spinosad in integrated PTM management in Tunisia may
provide more sustainable control of this pest.
Introduction
The potato tuber moth (PTM), Phthorimaea operculella (Zeller), is one of the worst
pests of potato in warm climate regions. PTM damages are of great concern both in
potato fields and storages. In North Africa, while PTM is active throughout the year,
activity peaks are observed from the end of April to early August (HANAFI, 1999).
Besides potato, the larvae feed on other Solanaceae plants such as tomato
(Lycopersicon esculentum Mill.), eggplant (Solanum melongena L.) and pepper
(Capsicum annuum L.) (GILBOA and PODOLER, 1994).
In Tunisia, PTM threat concerns mainly the spring potato crop. Damages occur in
the field particularly prior to harvest and also during tuber storage. Because of an
82 Spinosad a new biopesticide for PTM
increase in potato production and the lack of refrigerated facilities, around 60 to 70%
of the spring potato production is stored in rustic farm stores. Many farmers simply
heap their potatoes in the shade of some trees and cover them with a thick layer of
potato haulms (vines). Unlike cold stores, such low cost systems have no control on
storage temperature and humidity levels. Moreover, there is a high risk of tuber
damage by PTM followed by a severe rotting of tubers. In the past years, farmers
using rustic stores have controlled PTM with broad spectrum chemical pesticides. In
addition to being costly, many pesticides were misused due to farmer’s lack of
knowledge of the correct use of chemical pesticides. Hence, the overuse of some
insecticides, particularly Pyrethoids based on Deltamethrine, has decreased their
effectiveness against PTM infestations. Using higher doses has not solved the
observed decrease in efficacy. Some unofficial records even mentioned extreme cases
where farmers had used other pesticides like carbamates and organophosphates to
overcome Deltamethrine in-effectiveness. In addition to the environment harm, such
pesticides are not labelled to be used on potato tubers.
Regarding this situation, studies were designed to emphasize an integrated
management of PTM including:
1) The enhancement of the existing rustic storage system: a new system named the
Improved Non-Refrigerated Potato Storage System (INRPSS) was proposed to
potato farmers. It consists of a modified store built with on-farm materials. Store
specifications and components enable better control of inside temperature and
humidity levels. It also provides better protection against post-harvest PTM
infestation (KHAMASSY et al., 2002).
2) The introduction of new pesticides that are more effective and less harmful to the
environment. This step has major importance for the success of the INRPSS.
Unless tubers are carefully sorted, the PTM threat may also come from inside the
store. Since large quantities of tubers are stored, perfect sorting is not possible.
Thus relying on an effective and safe insecticide would be necessary to manage
any sudden PTM infestation development.
For this purpose, a new biopesticide, named TRACER was studied. It is based on
Spinosad, a product derived from the bacterium Sacchropolyspora spinosa (Mertz
and Yao). A series of experiments, including on-farm trials, were carried out during
2003-2005 to evaluate Spinosad efficacy against PTM infestation. The current paper
summarizes the main results obtained from these trials.
Material and Methods

On-farm experiments
Spinosad on-farm experiments were carried out during the period 2003 to 2005 in the
regions of Nabel, Bizerte and Jendouba, Tunisia. Each year, 30 farmers applied the
biopesticide TRACER SC240 (Spinosad concentration: 240 g/l) on their stored
potatoes. In all cases, the same treatment steps were followed: newly harvested potato
tubers were first kept in a pre-storage area for 2 weeks. Tubers were then rigorously
Spinosad a new biopesticide for PTM 83
sorted to eliminate any traces of rot or PTM attack. Tubers were then dipped into a
TRACER mixture diluted 1/10,000; 300 litres of this mixture are required to treat one
tonne of potato. Once dried, treated tubers were stored in the Improved Non-
Refrigerated Store (INRS). Tuber piles were about 1.10 m high and divided into four
sections for easy control during the storage period (June to August).
Storage periods averaged 90 days from mid-June to mid-September. PTM
infestation was measured every 2 weeks. Each measurement consisted of four
randomized samples of 100 tubers taken from the store. Because infestation by first
instar larvae are difficult to detect visually, it had been agreed to consider an initial
infestation rate of 2% for all the stores.
Since the potato stores were for commercial purpose, including a control
treatment was not possible in most cases. Thus, a series of reference trials was
established to confirm farmer results. Spinosad efficacy was compared with other
commonly used pesticides: Deltamethrine, Bacillus thuringiensis Berliner subsp.
kurstaki (Btk), carbamates and organophosphate pesticides. Five reference
experiments were carried out in 5 stores at the Institut National de la Rechere
Agronomique de Tunis (INRAT). In each experiment, Spinosad was compared to
another pesticide. Treatments (Spinosad, other pesticide, and control) were randomly
assigned to 3 separate potato piles in each store. According to pesticide formulation,
tubers were treated as described in Table 1. The evaluation of treatment efficacy
followed the methodology described for the on-farm experiments.
Table 1. Specification of the pesticides used in the reference experiments
Commercial Treatment Treatment
Pesticide Formulation Concentration
name technique concentration
10ml/hl; 3hl
Tracer SC Suspension Mixture
Spinosad 240g/l of mixture/1
240 concentrate dipping
ton of potato
100ml/hl; 3hl
Delta Emulsifiable Mixture
Deltamethrine 25g/l of mixture/1
Decis concentrate dipping
ton of potato
Bacillus Dustable 3 Kg/t of
Bactospeine 1% Powdering
thuringiensis powder potato
150g/hl; 3hl
Wetting Mixture
Methomyl Lannate 25 25% of mixture/1
powder dipping
ton of potato
100ml/hl; 3hl
Chlopyriphos- Emulsifiable Mixture
Dursban 4e 480g/l of mixture/1
ethyl concentrate dipping
ton of potato
Laboratory experiments
To confirm Spinosad efficacy, tests under controlled conditions, including artificial
PTM infestation, were conducted at the INRAT laboratories. Tests were conducted
with the standard cultivars Spunta and Atlas.
In a first experiment, Spinosad biopesticide (S) efficacy was compared with a

Deltamethrine pesticide (D). A completely randomized split plot design was used for
the experiment with pesticide type (Spinosad, Deltamethrine, control) as the main
treatment and cultivar (Spunta and Atlas) as the secondary treatment. Three replicates
of 100 tubers per treatment were used for the trials. Potato tuber treatments and
pesticide concentrations were as described in the on-farm experiments (Table 1).
Treated tubers were stored in open boxes that allow free access by PTM. PTM
infested and non-treated tubers were regularly added to the different treatments. An
insect-proof net was used to cover the boxes and to ensure a high level of PTM
population. Store temperature was kept around 27 to 30°C during the storage period
to enhance PTM activity and cycles. Infested tubers were counted after 30 days and
after 60 days of storage.
In another series of experiments conducted during 2004 and 2005, the effect of
pesticide ageing on Spinosad efficacy was studied. New Spinosad biopesticide (S1)
was compared to one-year-old Spinosad (S2). The experimental protocol used for the
evaluation was the same as the in the first experiment except that the (S2) treatment
was substituted for the (D) treatment.
Statistical analysis
Collected data from on-farm and laboratory experiments were compiled and analysed
with SPSS 10.0 statistical software. Retrieved means were compared using One-Way
ANOVA procedure. Post Hoc range tests and pairwise multiple comparisons were
done to determine significant differences between the mean values at a significant
level of P≤ 0.05%.
Results
On-farm experiments
Depending on the farmer’s marketing decision, storage periods varied during the
three years of the trials. In 2003, due to high yields and consequent low potato prices,
most farmers had chosen to store longer. The storage period was then from 100 to
120 days. However, in 2005, low yields of the spring crops and increased
consumption demand had inflated potato prices. That is why most farmers preferred
to sell their stocks after only 2 months of storage. Only a few maintained their potato
stocks for 3 months or more.
Regardless of annual variation, low PTM infestation rates were measured during
2003 to 2005. After 90 days of storage, mean tuber infestation varied from 5 to 7.7%
(Table 2). No sudden variations were observed between monthly recorded PTM
infestation rates. Rarely more than one or two superficial attacks per tuber were
found. Observed attack levels had little effect on the overall stock’s marketable value
depreciation.
Table 2. Monthly average PTM infestation rates (%) in the Improved Non-Refrigerated Potato
Stores and treated with Spinosad (Tracer SC 240) recorded during 2003 to 2005 in the
states of Nabel, Jendouba and Bizerte, Tunisia.
Year 2003 2004 2005
Storage Period (d) 30 60 90 30 60 90 30 60 90
Nabel 4.1 5 5.4 4.6 5.8 5 3.8 5.5 --
Jendouba 4.7 6.2 7.7 3 5 6 3.2 4.5 --
Bizerte 3.5 5 5.9 3 4.2 5.7 3 4.3 --
In the reference experiments, Spinosad (Tracer SC 240) efficacy against PTM

attacks was significantly greater than of the deltamethrine pesticides, such as Delta
and Decis (Figure 1a, b). After a storage period of 45 days, tuber infestation in the
Delta treatments increased to about 20%. PTM infestation were not significantly
different between Delta treated tubers and the control. In both treatments infestation
increased up to 80 and 90% after 90 days of storage. In comparison, PTM infestation
in the Spinosad treatment reached only about 5% at the end of the storage period. The
product Decis showed a higher efficacy than Delta when it was compared with
Spinosad (Figure 1b). However, after 90 days of storage, about 55% of the Decis-
treated tubers were infested. In the Spinosad treatment, tuber infestation was again
low with less than 5%.
Treatments with Bactospeine, another biological insecticide based on Btk,
showed similar efficacy to that of Spinosad (Figure 1c). Slight but not significant
differences were observed after 75 and 90 days of storage. At the end of the storage
period, PTM infestation reached about 90% in the control, and 11% and 5% in the
Btk and Spinosad treatment, respectively.
Other pesticides such as Dursban (chlopyriphos-ethyl) and Lannate (methomyl)
showed also a high efficacy controlling PTM which were with less 10% similar to
Spinosad (Figure 1d, e).
Laboratory experiments
In the 2004 experiments, Spinosad efficacy against PTM attacks was significantly
higher than Deltamethrine (Table 3). After 30 days of storage, over 78% of Spunta
tubers treated by Deltamethrine were infested. Control Spunta tubers showed similar
infestation rates while infestation was less than 10% with Spinosad. There was no
significant difference observed in treatment efficacy between the two potato varieties
used. In the potato variety Atlas PTM infestation was less than 4% using Spinosad.
No significant difference was found between Deltamethrine treatment and the control
(respectively 40.7% and 36%). One month later, PTM infestation rates were still
significantly lower with Spinosad compared to Deltamethrine. Highest infestation
records were about 16.7% for Spinosad-treated tubers and 98% for Deltamethrine.
In the 2005 experiments, PTM activity and infestation was lower despite the
controlled conditions (Table 3). After 30 days of storage tuber infestation was similar
for Spinosad and Atlas Deltamethrine-treated tubers. Spunta tubers treated with
Deltamethrine showed a higher infestation (19.3%). After 60 days of storage, Spunta
tubers reached a PTM-infestation of about 11% in the Spinosad treatment while
infestation exceeded 45% in the Deltamethrine treatment. The efficacy of the

pesticide treatments was similar in the potato variety Atlas.
The new and old Spinosad pesticide (S1 and S2) showed only minor differences
30 days after treatment in 2004 experiments (Table 4). In the 2005 experiments, no
differences were found among all treatments either 30 or 60 days after storage.
100
90 b 100
c
80
b 90
b 80
PTM Infestation (%)
PTM Infestation (%)

70 c
b 70
b
60 60 c
50 b
b 50
b
40
b b 40
30 30
20 b b 20 bb
ab
10
a a a a a 10 a a a a
0 0
15 30 45 60 75 90 15 30 45 60 75 90
Storage Period Storage Period
(Days) TRACER DECIS Control (Days)
TRACER DELTA Control
100
90 b
80
b
PTM Infestation (%)
70
60
50
b
40 b
30
20
b
a a a a a
10
a a a a a
0
15 30 45 60 75 90
Storage Period
TRACER BACTOSPEINE Control (Days)
100 100
90 90
80 80
b
PTM Infestation (%)
PTM Infestation (%)
70 70
60 60
b
50 b 50 b
b
40
b 40
b
30 30
b b
20 20
10
a
a
a
a aa aa a
a 10 aa aa aa aa
0 0
15 30 45 60 75 90 15 30 45 60 75 90
Storage Period Storage Period
TRACER LANNATE Control TRACER DURSBAN Control (Days)
(Days)
Figure 1. Tracer SC 240 (Spinosad) 2003 reference experiments as compared to other commonly
used pesticides: (A): Tracer compared to Delta (Deltamethrine); (B): Tracer compared to
Decis (Deltamethrine); (C): Tracer compared to Bactospeine (Bacillus thuringiensis
subsp. kurstaki); (D): Tracer compared to Lannate (Methomyl, not recommended to be
used on potato tubers); (E): Tracer compared to Dursban (Chlopyriphos-ethyl, not
recommended to be used on potato tubers). Different smaller case letters represent
significant subsets at P≤ 0.05%.
Table 3. Effects of Spinosad (S) and Deltamethrine (D) treatments on potato tuber moth infestation
for cultivars Spunta and Atlas compared to untreated tubers (C), INRAT 2004-2005.
2004 PTM infestation (%)
Storage period 30 days 60 days
Treatment S D C S D C
Spunta 08.00a1 78.66b 65.33b 16.67a 98.00b 100b
Atlas 03.66a 40.66c 36.00c 09.33a 79.33c 100b
Treatment S D C S D C
Spunta 08.00a 19.33b 31.33c 11.33a 46.00b 94.66d
Atlas 04.00a 07.33a 37.33c 10.00a 41.33b 82.00c
1
Different smaller case letters represent significant subsets at P≤ 0.05%.
Table 4. Effects of a new Spinosad (S1) and 30-month old Spinosad (S2) treatments on potato tuber
moth infestation for cultivars Spunta and Atlas compared to untreated tubers (C), INRAT
2004 and 2005.
Treatment S1 S2 C S1 S2 C
1
Spunta 8.33ab 16b 59.66c 14.66ab 21.33ab 100c
Atlas 4.66a 12ab 46.00d 11.33a 18.00ab 100c
Treatment S1 S2 C S1 S2 C
Spunta 06.00a 04.00a 41.00b 11.00a 10.66a 89.66b
Atlas 04.00a 05.33a 36.66b 08.33a 08.66a 85.00b
1
Different smaller case letters represent significant subsets at P≤ 0.05%.
Discussion
According to the presented results, effectiveness of Deltamethrine pesticides to
control PTM tuber infestation in Tunisia seemed to have decreased. Depending on the
commercial formulation, Deltamethrine efficacy lasts only 3 to 4 weeks
(ANONYMOUS, 2006). It may be the reason why PTM infestation observed after 1
month and more had increased suddenly in our experiments. Deltamethrine is widely
used to protect other crops besides potato. Since it was introduced in Tunisia for more
than 20 years, its overuse may have caused a certain level of PTM resistance to this
pesticide. However, such an assumption is still unclear and needs additional research.
Bacillus thuringiensis appears to be a good biological PTM control option during
storage as earlier shown by KROSCHEL and KOCH (1996). Despite that, on-farm
surveys revealed that it is not largely used in Tunisia (KHAMASSY, unpublished).
Most farmers preferred dipping treatments rather than powdering which require more
labour and skill (treatment layer by layer). Unlike dipping, powder treatment may not
cover the whole tuber surface when large quantities of potato are treated.
Chlopyriphos-ethyl and methomyl pesticides also showed good efficacy for

control of PTM. However, such pesticides are not recommended to be used on potato
tubers because of their high toxicity and long residual effects.
Studying the effect of pesticide ageing on its effectiveness may provide some
clues regarding its adoption. In Tunisia, most farmers keep their unused pesticides
(generally unsealed) to be used on the next crop. This may cause a decline in
pesticide effectiveness. Regarding the experiment conducted with old Spinosad
pesticide, no definitive conclusion can be made concerning the effect of product
ageing on Spinosad efficacy. A 30-month old Spinosad seemed to be as efficient as a
newly produced one. Further trials are warranted to clarify this hypothesis.
The adoption of Spinosad in integrated PTM management may provide a more
sustainable control of this pest in potato stores. Favourable for farmers and key for its
quick acceptance are very likely its application by dipping and at low concentrations.
Actually Spinosad is the main biopesticide used against PTM in the INRPSS to limit
quantitative and qualitative potato losses during summer period. Future research
would concern possible interactions of Spinosad with fungicides and anti-sprouting
products. Further, additional efficient PTM control methods are required that can be
alternated with Spinosad. This would enhance a more sustainable and long-term PTM
control.
Conclusion
Spinosad has shown high efficacy in controlling PTM infestations and is suggested
for its use in integrated PTM management in Tunisia.
References
ANONYMUS 2006. Guide Phytosanitaire de la Tunisie. 3rd Edition. 287p.
GILBOA S AND H PODOLER 1994. Population dynamics of the potato tuber moth on
processing tomatoes in Israel. Entomologia Experimentalis et Applicata 72:
197-206.
KROSCHEL J AND W KOCH 1996. Studies on the use of chemicals, botanicals and
Bacillus thuringiensis in the management of the potato tuber moth in potato
stores. Crop Protection 15: 197-203.
HANAFI A 1999. Integrated pest management of potato tuber moth in field and
storage. pp211-214. in: Proceedings of the 14th Triennial Conference of the
European Association for Potato Research. Sorrento, Italy. May 1999, 2-7th.
749p.
KHAMASSY N, TEMIME AB AND I ARFAOUI 2002. Improved potato storage system.
Abstracts of the 15th Triennial Conference of the European Association for
Potato Research. July 2002, 14-19th. Hamburg, Germany. pp 175.
89
Laboratory experiments towards the

development of an attract-and-kill strategy for
the potato tuber moth complex
Jürgen Kroschel and Octavio Zegarra
Abstract
Since sex pheromones have been identified for the common potato tuber moth
(PTM), Phthorimaea operculella (Zeller) and Andean potato tuber moth (APTM),
Symmetrischema tangolias (Gyen), they have served as ideal tools to monitor flight
activity but have not proved effective as a means of control. The objective of our
studies was to test the pheromones in an attract-and-kill strategy. Pure pheromones
(0.05%) and the contact insecticide Cyfluthrin were the active ingredients, formulated
with plant oils and U.V. absorbers. In lab bioassays, Cyfluthrin, at a concentration of
0.5% resulted in the highest and fastest killing of both moth species after 24 and 48
hrs, with a mortality of 58 and 84% for PTM and 53 and 68% for APTM,
respectively. At this insecticide concentration, 100% mortality was achieved after
three to four days. In contrast, control males survived for 13 days. Mixtures of
pheromones of the two species gave better or equal control, indicating no interference
of the pheromones in one formulation. At 20ºC, stability of the formulation was
achieved for a period of a minimum of 36 and 29 days for the two moth species
without any reduction in efficacy. The practical use of the attract-and-kill strategy is
discussed.
Introduction
The common potato tuber moth (PTM), Phthorimaea operculella (Zeller) and the
Andean potato tuber moth (APTM), Symmetrischema tangolias (Gyen) are important
pests of potato (Solanum tuberosum L.) and other Solanceae crops. PTM has a
worldwide distribution and is considered the most damaging potato pest in the
developing world (KROSCHEL and SPORLEDER, 2006; CISNEROS and GREGORY, 1994).
It had been a key pest in Andean potato production but was replaced in many
growing areas by the APTM (KELLER, 2003). The damaging life-stage of both
species is when larvae attack leaves, petioles, stems and tubers of potatoes. For the
control of the PTM IPM strategies have been developed and proposed for several
regions (e.g., Republic of Yemen, KROSCHEL, 1995; Egypt, LAGNAOUI and EL-
BEDEWY, 1997; India, CHANDEL et al., this volume; Australia, HORNE and PAGE, this
volume).
90 Development of an attract-and-kill strategy for PTM
Sex pheromones have been identified for both moth species (P. operculella:
PERSOONS et al., 1975; S. tangolias: GRIEPINK et al., 1995). Since then, they have
been deployed for mass trapping (RAMAN, 1984; RAMAN, 1988), monitoring the
flight activity of PTM/APTM males and studying population dynamics and seasonal
moth population trends to tuber infestations at harvest (RAMAN, 1982; KROSCHEL and
KOCH 1994; KELLER, 2003). Sex-pheromones used in traps in the field compete with
natural sources released by females. Successes of any trapping will therefore depend
on the proportion of males attracted by the trap and therefore removed from the
mating population. RAMAN (1988) demonstrated that densely deployed pheromone
traps at a density of 45 traps per hectare reduced potato tuber infestation by PTM
below levels of insecticide treatments. However, for practical and economic reasons,
the numbers of traps employed per unit agricultural area have always been a limiting
factor for the use and adoption of a mass trapping technology in pest management.
For this reason, the attract-and-kill or attracticide strategy has received attention and
has been successfully used for controlling orchard pests such as the codling moth
Cydia pomonella L. (HOFER and BRASSEL, 1992; CHARMILLOT et al., 1997). This
strategy consists of an insecticide-pheromone co-formulation by which the male
moths are attracted by the pheromone and killed through contact with the insecticide.
The application of this approach was further developed by the Bayer AG, Germany,
and has led to the novel attracticide product Appeal (LÖSEL et al., 2000; EBBINGHAUS
et al., 2001).
We tested the hypothesis that the attract-and-kill strategy could also be effective
in controlling the PTM and APTM as important pests of potato and other
Solanaceous field crops. The objective of our studies was to evaluate the efficacy of
different pheromone-insecticide co-formulations and their stability under laboratory
conditions.
Insect origin and rearing

Male moths of PTM and APTM used in this study were obtained from a laboratory
colony maintained at the International Potato Center (CIP), Lima, Peru. The PTM
rearing started with moths collected in 1984 in La Molina, Peru. Periodic
introgressions of wild-caught moths from the same locality and Peruvian Highlands,
Huancayo (3,400 m asl), have been made over the years. For APTM, rearing started
in January 2006 with a new collection from the Huancayo region. The rearing was
carried out at a room temperature of 23-25°C for PTM and 18-20°C for APTM,
respectively, 60-80% relative humidity and natural photoperiod. Eggs were placed in
plastic containers (30×20×7.5 cm) containing potato tubers (cv. Peruanita) as food
placed on sand or carton as pupation site for PTM or APTM. Pupae were kept
individually in small plastic cups to separate emerging adults by sex. Newly hatched
males were kept for 48 hrs prior to their use in the experiments to allow them to reach
Development of an attract-and-kill strategy for PTM 91
full sexual maturity. They had free access to a 2% honey solution during their life
time.
Attract-and-kill chemical composition

Pure pheromones (P. operculella: trans-4, cis-7- tridecadien-1-ol-acetate and trans-4,
cis-7, cis-10 tridecantrien-1-ol-acetate; S. tangolias: (E)-3-tetradecen-1-o-acetate and
(E, Z)-3,7-tetradecadien-1-ol-acetate) (Pherobank, The Netherlands) and the
insecticide Baythroid TM 525 SL (Cyfluthrin 25 g/l) (Bayer Crop Science, Chile)
were the active ingredients employed in all experiments. The pheromones were used
in one concentration (0.05%). For the preparation of the formulation we followed the
protocol of LÖSEL et al. (2000). The active ingredients were formulated with castor
oil (50%) and ultraviolet absorbers (UVINIL M40 and UVINIL 539T; BASF,
Germany, both at 7.5%) stabilized by the nonionic emulsifying agent Cremophor CO
40 (BASF, Germany, at 7.5%) and Aerosil 200 V (Degusa Pharma, at 7.0%) for
incrementing the fluidity of the product. Each of the two pheromones was formulated
individually and in combination. The components were mixed in a water bath at
60°C, and stored at 4°C prior to use.
Bioassays
Testing optimal insecticide concentration and efficacy

The first experiments were carried out to determine the most effective insecticide
concentration and the efficacy of the attract-and-kill formulation to kill male adults of
PTM and APTM over time. The experiments consisted of seven treatments that
included five insecticide concentrations (0.03, 0.06, 0.125, 0.25 and 0.5% of
Cyfluthrin) and one control without insecticide and without pheromone. Each of the
two pheromones was tested separately and in combination in the same manner.
Wooden cages (75×50×60 cm) covered with nylon gauze were installed at constant
room temperatures of 20ºC, at 65% rel. humidity and a photoperiod of 16:8 hrs
day/night time. The pheromone formulation was dropped on a Petri dish (100-μl
droplet) placed in one corner of the cage. In each of the cages 20 male moths of the
same age were released. The experiments for PTM and APTM were carried out at
different times, except when the pheromones of both species were tested in one
formulation. In this case, 20 male moths of both species were released together in the
cages. Survival of male moths was monitored every day until all moths in all
treatments died. No replications were included for the different treatments but the
complete experiment was repeated three times.
Stability of the attract & kill formulation
This experiment was carried out to understand the stability, durability and efficacy of
the attract-and-kill formulation over time, which consisted of an insecticide
concentration of 0.5% Cyfluthrin and a pheromone concentration of 0.05%. The
experiment was carried out in wooden cages under the same conditions as described
above for PTM/APTM. At the beginning of the experiment one 100-μl droplet of the
formulation of each of the two species was applied on a Petri dish and placed in one
corner of the test cages; at the time all PTM/APTM males had been killed by the
attract-and-kill formulation a new set of 20 male adults was released per cage. This
was repeated over a period of 36 and 29 days for PTM and APTM, respectively. A
combined formulation of the two pheromones was not tested in this experiment. For
each of the two species, each experiment had four replications and was repeated four
times.
Statistical data analysis was carried out using SAS® statistical software (SAS, 2005).
Prior of analysis of variance (ANOVA), tests for normal distribution and
homogeneity of variances were carried out using Shapiro-Wilks-W-Test. Data of
experiments conducted three or four times and showing homogenous variances were
pooled and analyzed as combined experiments. Mean mortality responses were
adjusted by Abott’s formula (ABOTT, 1925). Significant differences between the
mean values were determined by the Tukey’s Honest-Significant-Difference-Test
(HSD) for multiple means comparison at a significance level P≤0.05.
Results
Speed of mortality
Cyfluthrin at 0.5% resulted in the highest and fastest killing of both moth species
after 24 and 48 hrs with a mean mortality of 60.2% and 81.2%, respectively, testing
both species alone or in mixtures (Table 1).
Table 1. Abbott corrected mortality values for males of the potato tuber moth Phthorimaea
operculella, Andean potato tuber moth Symmetrischema tangolias, or mixtures of both
after 24 and 48 hrs of exposure to an attract-and-kill formulation with different cyfluthrin
concentrations.
Pheromone Concentration of cyfluthrin (%)
/species 0.5 0.25 0.125 0.06 0.03
tested
24 48 24 48 24 48 24 48 24 48
PTM 57.9a 84.2a 47.4b 63.2b 38.8b 63.2b 26.3c 47.4c 21.1c 31.6c
APTM 52.6a 68.4a 15.6b 42.1b 26.3b 31.6b 26.3b 31.6b 0 0
1
PTM and
70a 93a 48b 83a 30b 62b 18c 36c 15 c 28c
APTM
Mean 60.2 81.2 37.0 62.8 31.7 52.3 23.5 38.3 12.0 19.7
Each value represents the mean of pooled data from three repetitions of the experiment. In each
row, means with the same letter are not significantly different at P≤ 0.05 after Tukey’s HSD-Test.
1
The pheromones of both species were tested together in one formulation.
In comparison, the Cyfluthrin concentrations 0.25%, 0.125%, 0.065% and
0.035% reached only mean mortalities of 62.8%, 52.3%, 38.3%, and 19.7%,
respectively, after 48 hrs. The pheromone-insecticide formulation was more effective
for PTM than for APTM, however, mixtures of the two species gave equal or better
control, indicating no interference of the pheromones in one (Table 1).
Another series of experiments confirmed that the Cyfluthrin concentration of
0.5% killed all males rapidly after 24hrs with a mean mortality of 67.5 and 70% for
all three attract-and-kill formulations (Figure 1, 2). After 3 and 4 days, respectively, a
100% kill of all males was observed. In contrast, the males in the control treatments
stayed alive until day 13 after the start of the experiments. Mixtures of the two
pheromones in one formulation gave equal control (Figure 2).
P. operculella
a a
100 a (100)
90 (94.7)
Mortality (%)
80 a (85)
70
60 (70)
50
40
30
20 b b b b
10
0
1 2 3 4 5 6 7 8 9 10 11 12 13 14
S. tangolias
a (100)
100 a
90
Mortality (%)
(89.5)
80 a
70
(68.4)
60
50
40
30
20 b b b
10
0
1 2 3 4 5 6 7 8 9 10 11 12 13
Figure 1. Mortality of potato tuber moth (Phthorimaea operculella) and Andean potato tuber moth
(Symmetrischema tangolias) males over time in cages after the application of one drop of
the attract-and-kill test formulation (containing 0.05% of the sex pheromone and 0.5%
Cyfluthrin). Values in brackets indicate Abbott corrected mortality of pooled data from
three experiments. Vertical bars indicate the standard error of the means. Means with the
same letter for each evaluation date are not significantly different at P≤ 0.05 after Tukey’s
HSD-Test.
P. operculella + S. tangolias
a a
100 a (100)
Mortality (%) 90
80 a (95)
70 (87.5)
60 (67.5)
50
40
30 b
b
20 b
b
10
0
1 2 3 4 5 6 7 8 9 10 11 12 13 14
Time after exposure (days)
Attract-and-Kill Control
Figure 2. Combined mortality of potato tuber moth (Phthorimaea operculella) and Andean potato
tuber moth (Symmetrischema tangolias) males over time in cages after the application of
one drop of a combined P. operculella/S. tangolias attract-and-kill test formulation
(containing 0.05% of both sex pheromones and 0.5% Cyfluthrin). Values in brackets
indicate Abbott corrected mortality of pooled data from three experiments. Vertical bars
indicate the standard error of the means. Means with the same letter for each evaluation
date are not significantly different at P≤ 0.05 after Tukey’s HSD-Test.
Stability of the formulation

At 20ºC and over a period of 36 and 29 days for PTM and APTM, respectively, no
reduction in the stability and efficacy of the attract-and-kill formulation and the time
needed to kill 100% of the males could be observed (Figure 3). At all evaluation
times, 40% of male adults were killed after 24 hours and 100% mortality was
observed after 4 days.
Discussion
The attract-and-kill strategy is consistent with the aims of IPM. The specificity of the
sex pheromone ensures that only the target species are affected, avoiding deleterious
effects on beneficial and other non-target organisms often associated with broadcast
spray-applications of conventional insecticides (EBBINGHAUS et al., 2001). The
insecticide Cyfluthrin was highly effective at a concentration of 0.5%. Compared to
APTM, PTM responded with a faster death rate after 48 hrs at all concentrations
tested. However, after three and four days, mortality of males of both species was 90
and 100%, respectively, neither the pheromones of the two species were individually
or combined used in the attract-and-kill formulation. At the same concentration of
Cyfluthrin, LÖSEL et al. (2000) achieved 100% mortality after bringing adults of the
codling moth Cydia pomonella directly into contact with 100-μl droplets of the
attract-and-kill test formulation, which had the same composition as in our
experiments. In contrast, the same authors reported mortalities of only 40% to 68% in
cage trials with 4% Cyfluthrin and varying concentrations of the C. pomonella
pheromone. Compared to PTM, both the codling moth and the APTM are relatively
stronger and larger insects and perhaps for this reason are more tolerant to the contact
insecticide.
abc ab ab ab ab ab ab a bc c
100
Mortality (%)
80
60
Error!
40
20
0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39
Evaluation (days)
a a a a a a a a
100
Mortality (%)
80
60
40
20
0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31
Evaluation (days)
Figure 3. Efficacy and stability of the attract-and-kill formulation at controlled conditions of 20 ºC

without UV exposure to kill (A) potato tuber moth, Phthorimaea operculella, and (B)
Andean potato tuber moth, Symmetrischema tangolias, males over a period of 36 and 29
days, respectively. One-droplet (100-μl) of the attract-and-kill formulation was applied at
day one of the experiments. Each time all PTM/APTM males had been killed a new set of
20 male adults were released per cage. Mean mortality values are pooled data from four
experiments. Vertical bars indicate the standard error of the means. Means with the same
letter are not significantly different at P≤ 0.05 after Tukey’s HSD-Test, and indicate
differences in time until 100% mortality was achieved.
CHARMILLOT et al. (1997) demonstrated that a density of one hundred 50-μl

attract-and-kill sources, consisting of 0.16% of the codling moth pheromone and
6.0% permethrin, halved the codling moth activity while 5000 50-μl attract-and-kill
sources reduced the male population by 90%. EBBINGHAUS et al. (2001) achieved
efficient control by using 4000 to 6000 100-μl droplets. Taking these application
rates into consideration only 1.25-2.5 g of a.i. of Cyfluthrin would be needed per
hectare which is 15 to 30-fold less than the recommended dosage of Baythroid for
controlling lepidopteran pests. Experiences with the codling moth in wind tunnel
experiments have shown that a pheromone concentration of 0.065% gave best results
in attracting males (LÖSEL et al., 2000). In our experiments we tested a concentration
of 0.05% of the PTM and APTM pheromone, which provided good results under
laboratory conditions. The optimal pheromone concentration might be very species
specific. Hence, additional tests are warranted to evaluate different pheromone
concentrations compared to those emitted by females of both species in order to
optimize male catches and mortality under field conditions. The stability of the
formulation was tested over a period of 29 and 36 days for the PTM and APTM,
respectively, without showing any reduction in efficacy. This is promising, but field
evaluations will be important, taking higher temperature and the influence of UV
light into consideration.
Conclusions
The first results of laboratory experiments to develop an attract-and-kill strategy for
two species of the potato tuber moth complex are promising and comparable to other
Lepidoptera species, for which this strategy could be successfully applied under field
conditions. Field experiments under different environments, field sizes and moth
densities will be required to approve its applicability under practical conditions. A
further potential application of this technology can be found in protecting potato
tubers in storage from PTM and APTM losses.
References
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Journal of Economic Entomology 18, 265-267.
CHARMILLOT P, PASQUIER D, SCALCO A and D HOFER 1997. Lutte contre le
carpocapse Cydia pomonella L. par un procédé attracticide. Revue Suisse de
Viticulture, Arboriculture, Horticulture 29:111-117.
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EBBINGHAUS D, LÖSEL P, ROMEIS J, CIANCIULLI–TELLER M, LEUSCH H, OLSZAK R,
PLUCIENNIK Z and J SCHERKENBECK 2001. Appeal: efficacy and mode of action of
attract and kill for codling moth control. Pheromones for Insect Control in
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and identification of sex pheromone of Symmetrischema tangolias (Gyen)
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Potato Conference, February 7-9, 2006, Moses Lake, Washington, 85-94.
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moth in Egypt. CIP Circular 22/23: 6-7.
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field performance of an attracticide against the codling moth, Cydia
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LÖSEL P, PENNERS G, POTTING R, EBBINGHAUS D, ELBERT A and J SCHERKENBECK 2000.
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kill strategy for the control of the codling moth, Cydia pomonella.
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PERSOONS C, VOERMAN S, VERWIEL P, RITTER F, NOOYEN W and A MINKS 1976. Sex
pheromone of the potato tuberworm moth Phthorimaea operculella:
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RAMAN K 1988. Control of potato tuber moth Phthorimaea operculella with sex
pheromones in Peru. Agriculture, Ecosystems and Environment 21: 85-99.
RAMAN K 1984. Evaluation of a synthetic sex pheromone funnel trap for potato
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98
Chapter III
Integrated Pest Management

101
Reducing potato tuber moth damage with

cultural practices and pesticide treatments
George H. Clough, Sandra J. DeBano, and Philip B. Hamm
Oregon State University, Hermiston Agricultural Research and Extension
Center, Hermiston, OR 97838
Abstract
Cultural practices and insecticide treatments and combinations were evaluated for
effect on foliar damage and tuber infestation by potato tuber moth (PTM),
Phthorimaea operculella (Zeller), a new pest in the Columbia basin of eastern Oregon
and Washington. A range of intervals between initial application of several
insecticides and vine-kill were tested to determine how early to implement a program
to control PTM tuber damage. Esfenvalerate, methamidophos, and methomyl were
applied at recommended intervals, with programs beginning 28 to 5 days prior to
vine-kill. Indoxacarb was applied post vine-kill as a chemigation treatment. All
insecticide treatments significantly reduced tuber damage as compared to the
untreated control, but there was no apparent advantage to beginning control efforts
earlier than later in the season. Another study examined the effects of green foliage
and increased soil moisture on PTM tuber damage. Russet Norkotah and Russet
Burbank plants were allowed to naturally senesce or were chemically defoliated.
They received either no irrigation or were irrigated by center-pivot with 0.25 cm
water daily from vine-kill until harvest 2 weeks after vine-kill. Tuber damage was
similar between varieties, but daily irrigation following vine-kill reduced tuber
damage, and chemical vine-kill tended to reduce tuber damage as compared to natural
senescence. Also, 5 desiccants with rates of vine-killing ranging from hours to weeks
were examined, with and without weekly application of methamidophos beginning 21
days before complete desiccation. Insecticide application reduced foliar and tuber
damage with or without chemical vine desiccation. With insecticide and the
faster-acting desiccant, less tuber damage was observed. However, tuber damage with
the slowest acting desiccant was less without insecticide application.
Introduction
The potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera:
Gelechiidae) is one of the most significant constraints to potato (Solanum tuberosum
L.) productivity worldwide. It is a cosmopolitan, oligophagous pest of solanaceous
crops commonly found in tropical and subtropical regions. In the U.S., PTM has been
found in California, Arizona, New Mexico, Utah, and as far north as Maryland and
102 Use of cultural practices and pesticides treatments
Virginia. It was first detected in Umatilla County (Hermiston), Oregon, in 2002, with
a handful of reports of minor damage.
In 2003, several fields were lost due to PTM, resulting in an estimated economic
loss of about $2 million USD. Economic losses increased substantially in 2004 and
2005, due to increased PTM densities in areas already infested, range expansion,
tuber damage, and the cost of control measures. In two years, its range has expanded
225 km north of Hermiston, Oregon, to the area of Wilbur, Washington. This is
apparently the northern most distribution of PTM in the world (highest latitude north
or south). Recent reports confirm PTM in western, central and eastern Oregon,
including Klamath Falls, Union County; northeastern Washington and in at least two
counties in western Idaho. In all sites, distribution and population information is
incomplete at this time (RONDON et al., 2007).
Several field trials were initiated in 2005 to evaluate the potential to reduce PTM
damage through cultural practices, including timing of pesticide application
programs, application of post vine-kill irrigation with and without chemical
desiccation, and rate of vine-kill through use of various desiccants with and without
insecticide application.
Pesticide Timing Trial

To address the question of when a PTM program should be implemented to control
tuber damage, Asana (esfenvalerate), Monitor 4 (methamidophos), and Lannate LV
(methomyl) were applied, beginning at different intervals before vine-kill at growth
stage 80 (tuber maturation). In a second trial, Avaunt (indoxacarb) was applied at
vine-kill and one week later (Table 1). Plot size was 9.1 m long and eight rows wide
(6.9 m). ‘Ranger Russet’ potatoes were planted on March 31, under center pivot
irrigation. Normal commercial production practices were followed. Pre vine-kill
insecticide treatments were applied in 187 l·ha-1 with a tractor-mounted boom sprayer.
The defoliant Enquik was applied similarly on September 21 in 374 l·ha-1.
Avaunt treatments were applied with a chemigation simulator in 0.35 cm water.
One hundred tubers in the center portion of the inner two rows of each plot were
sampled 15 days after the desiccant application.
Irrigation/Senescence Trial
This study was designed to determine the effects of green foliage and soil moisture on
PTM tuber damage in early and late season potatoes. The three variables examined
were 1) variety, 2) irrigation practices after vine-kill, and 3) senescence. The two
varieties used were ‘Russet Norkotah’ (early) and ‘Russet Burbank’ (late). Potatoes in
the irrigated treatment were irrigated with 0.25 cm water by center pivot every day at
3:00 PM., from vine-kill until harvest. Potatoes in the non-irrigated treatment were
not watered after vine-kill. Half the plots were allowed to naturally senesce and half
were chemically defoliated with Enquik applied in 374 l·ha-1 as previously described.
Use of cultural practices and pesticides treatments 103
Plots were 8 rows wide and 9.1 m long, with each plot separated from the adjacent
plot by 6.1 m at the ends and 6 rows (5.2 m) on the sides to ensure irrigation was
properly applied at the plot level. One hundred tubers in the center portion of the
inner two rows were sampled 14 days after Enquik application.
Table 1. Potato tuber moth insecticide timing trial treatments, Hermiston, Oregon, 2005.
Application
Rate Interval Number
Pre vine-kill treatments
Asana (esfenvalerate) 0.55 l·ha-1 7 days 5
Lannate LV (methomyl) 2.63 l·ha-1 5 days 6
Monitor 4 (methamidophos) 2.04 l·ha-1 7 days 5
Post vine-kill treatments

Avaunt (indoxacarb) 0.22 l·ha-1 7 days 2
Avaunt (indoxacarb) 0.37 l·ha-1 7 days 2
Desiccant Trial
Five desiccants, which ranged from acting within hours to weeks, were examined
with and without weekly application of Monitor 4 beginning 21 days before vine-kill
(Table 2). Plot size was 9.1 m long and eight rows wide. ‘Ranger Russet’ potatoes
were planted on April 20. Normal commercial production practices were followed.
Monitor 4 and desiccants were applied as previously described, except that Enquik
was applied in 374 l·ha-1. One hundred tubers in the center portion of the inner two
rows of each plot were sampled 1 and 15 days after the last desiccant application
(September 20).
Table 2. Desiccant and insecticide treatments, potato tuber moth trial, Hermiston, Oregon, 2005.
Rate Activity* Application date
Desiccant
Rely (glufosinate-
3.50 l·ha-1 14 days 09-07
ammonium)
Aim (carfentrazone) 0.26 l·ha-1 10 days 09-11
ET (pyraflufen ethyl) 0.40 l·ha-1 7 days 09-14
-1
Reglone (diquat) 2.33 l.ha 2 days 09-19
-1
Enquik (monocarbamide 187 l·ha 1 day 09-20
dihydrogen sulfate)
Insecticide
Monitor 2.04 l·ha-1 - 08-31, 09-07
09-14, 09-20
*Estimates provided by manufacturer’s representatives.
Parameters used in the evaluation

Potatoes were stored at ambient temperature (21°C) for 10 to 14 days after harvest to
increase the likelihood of detecting PTM damage. Tubers were processed with a
French fry cutter and examined throughout for the presence or absence of PTM
damage.
Data were evaluated as the percent tubers damaged by PTM. Data were subjected to
analysis of variance by SAS GLM procedures, with mean separation by Duncans
multiple range test.
Results
All pre vine-kill insecticide treatments significantly reduced tuber damage as
compared to the control (F=3.9, p=0.004) (Figure 1). Also, Avaunt applied at
desiccation and one week later at the 0.22 l·ha-1 rate reduced tuber damage from
5.25% in the untreated control to 3.0%, and application at 0.37 l·ha-1 further reduced
tuber damage to 1.5% (data not shown).
Figure 1. Effect of timing of pesticide application on PTM tuber damage. Numbers following
insecticide indicate days prior to vine-kill applications began. Lines indicate the direction
of effect expected if there was an advantage to controlling all season.
PTM tuber damage did not differ significantly between ‘Russet Burbank’ and ‘Russet
Norkotah’ potatoes (F=0.007, p=0.94). Chemical defoliation tended to decrease PTM
tuber damage as compared to natural senescence (9.3 to 6.5%) but the difference was
not statistically significant (F=1.5, p=0.23). Daily irrigation following vine-kill
reduced percent tuber damage (F=3.7, p=0.06) as compared to no irrigation (Figure
2). There was no significant difference in the percentage of rotten potatoes from
irrigated and non-irrigated plots, indicating that this level of irrigation does not
increase fungal or bacterial disease.
Figure 2. Effect of irrigation after vine-kill on PTM tuber damage in ‘Russet Burbank’ and ‘Russet
Norkotah’ potatoes. Irrigation reduced the percent tuber damage at P=0.06
Desiccant Trial
The main effect of desiccant was not significant (F=1.75, p=0.135), but Monitor
application reduced PTM tuber damage from 3.81 to 1.85% (F=17.5, p=0.01). The
average percent tubers damaged increased from 2.58% at vine-kill to 3.08% at 2
weeks following vine-kill, but the difference was not significant (F=1.14, p=0.29).
There was a slight interaction between these three variables (F=2.07, p=0.08). At
vine-kill, the effect of desiccant was not significant (Table 3). Two weeks following
vine-kill, there were significant effects of Monitor 4 application and desiccant, and an
interaction between these two main effects (F=5.3, p=0.001). In almost every case,
application of Monitor 4 decreased PTM damage. The only exception occurred when
the application of Rely + Monitor resulted in increased PTM tuber damage compared
to Rely alone (Figure 3, 4). This is believed to be an anomaly.
Table 3. Effect of desiccant and insecticide on PTM tuber damage at vine-kill and 14 days after,
Hermiston, Oregon, 2005.
Tuber damage
Vine-kill Vine-kill +14
Desiccant1 Per cent (%)
Rely 0.63 3.00 b
Aim 3.13 5.00 a
ET 2.88 2.50 b
Reglone 4.00 2.00 b
Enquik 2.63 3.13 b
Control 2.25 3.88 ab
P-value 0.21 0.02
Insecticide2
Monitor 1.58 2.13
None 3.58 4.04
P-value 0.01 0.001
1
Mean of 800 tubers/treatment/sample time. 2Mean of 2,400 tubers/treatment/sample time. Means
followed by different letters significantly different at P≤ 0.05 (Duncans multiple range test).
Figure 3. Effect of desiccant with and without insecticide on PTM tuber damaged at vine-kill.
Figure 4. Effect of desiccant with and without insecticide on PTM tuber damage 14 days after vine-
kill.
Discussion

Numerous studies have found that various pesticides can effectively control PTM
damage to tubers (ABDEL-SALAM et al., 1972; BACON, 1960), but other studies have
found chemical control to be either of variable success (BACON et al., 1972; FOOT,
1974) or non-effective (AKHADE et al., 1970). Little has been published regarding
timing of insecticide application. KELLER (2003) suggested initiation of application at
“growth stage 70”, about 3 weeks prior to harvest. GODFREY and HAVILAND (2003)
state that “insecticides applied at vine-kill do not reduce tuber moth damage.”
The data from this trial indicate that there is no apparent advantage in beginning
control efforts earlier in the season versus later; there was no trend for PTM tuber
damage to increase when the beginning of the pesticide application program was
delayed. The lines on Figure 1 illustrate the effect expected if there was an advantage
to early control. Weekly application beginning 4 weeks prior to vine-kill was no more
effective than a single application one week before vine-kill. All the products tested
were effective. Also, application of insecticide at and following vine-kill also reduced
tuber moth infestation. These results are contrary to the recommendations of KELLER
(2003) and GODFREY and HAVILAND (2003), and indicate that the most critical time
for initiation of control is immediately prior to and at vine-kill.
In greenhouse studies, LANGFORD (1933) reported that maintaining moist soil reduced
tuber infestation to 5% as compared to 80% for dry soil. SHELTON and WYMAN (1979)
found that sprinkler irrigation for 4 weeks prior to vine-kill reduced tuber infestation
by 80% as compared to furrow irrigation. They attributed the effect to a reduction in
soil cracking, which decreased tuber accessibility by adults and larvae. They did not
investigate post vine-kill irrigation. FOOT (1979) also reported that increased soil
moisture, most effective with sprinkler irrigation, reduced the ability of adults to
oviposit on tubers, and of larvae to access tubers from the soil surface. He also
demonstrated that larval survival on the surface of buried tubers was inversely related
to soil moisture. Although our results were not as dramatic as those cited above,
application of light irrigation reduced tuber damage by 44%. These results are
promising, and the study will be repeated.
Chemical desiccation reduced the percent tubers damaged from 10.0% to 5.6%
and 8.6% to 7.4% for Russet Norkotah and Russet Burbank, respectively. At the time
of chemical desiccation, the Russet Burbank variety was rapidly senescing due to
early die-off, with 35% green foliage, while the Russet Norkotah had 70% green
foliage. This may have increased the variability associated with the data.
Desiccant Trial
FOOT (1974) demonstrated that mechanical defoliation 1 week prior to harvest
resulted in increased tuber infestation, as did delaying harvest for 2-4 weeks. Tuber
infestation did not increase with chemical desiccation with paraquat, which caused
rapid vine death. He did not examine the effect of slower acting desiccants.
The data from the desiccant trial support field observations that the most critical
time for controlling PTM tuber damage begins at vine-kill, and that the rate of vine-
dying is not an important factor, as long as there are green vines in the field.
Conclussion
The most critical time for initiation of chemical control is immediately prior to and at
vine-kill. Rate of vine desiccation did not impact PTM damage. Light, daily irrigation
beginning at vine-kill and continuing until harvest is another effective non-chemical
method of reducing damage from the PTM.
References
ABDEL-SALAM AM, ASSEM AM AND KH YOUSEF 1972. Studies on potato pests in
U.A.R. III. Effect of some insecticides on the potato tuber worm
(Phthorimaea operculella) in the field and storage. Journal of Applied
AKHADE MN, TIDKE PM AND MB PATKAR 1970. Control of potato tuber moth
(Gnorimoschema operculella Zell.) in Deccan plateau through insecticides
and depth of planting. Indian Journal of Agricultural Sciences 40: 1071-6.
BACON OG 1960. Control of the potato tuber worm in potatoes. Journal of
economic entomology 53: 868-71.
BACON OG, MCCALLEY NF, RILEY WD AND RH JAMES 1972. Insecticides for control of
potato tuber worm and green peach aphid on potatoes in California.
American Potato Journal. 49: 291-295.
(Lepidoptera: Gelechiidae) at Pukekohe. New Zealand Journal of Zoology 6:

623-636.
FOOT MA 1974. Field assessment of several insecticides against the potato tuber
moth Phthorimaea operculella (Zell.) at Pukukohe. New Zealand Journal of
Experimental Agriculture 2: 191-7.
GODFREY LD AND DR HAVILAND 2003. UC pest management guidelines. Potato
tuberworm. In: UC IPM Online. Statewide Integrated Pest Management
Program. UC ANR Publication 3463.
http://www.ipm.ucdavis.edu/PMG/r607300211.html
cropping systems of different agro-ecological zones. In: Kroschel, J (ed.)
Tropical Agriculture 11– Advances in Crop Research 1, Margraf Verlag,
Weikersheim, Germany. 153pp.
LANGFORD GS 1933. Observations on cultural practices for the control of potato
tuberworm Phthorimaea operculella (Zeller). Journal of Economic
RONDON SI, DEBANO SJ, CLOUGH GH, HAMM PB, JENSEN A, SCHEIBER A, ALVAREZ JM,
THORTHON M, BARBOUR J and M DOGRAMACI. 2007. Biology and management of
the potato tuberworm in the Pacific Northwest. A Pacific Northwest
Extension publication of Oregon State University, University of Idaho, and
Washington State University
http://oregonstate.edu/potatoes/ipm/insects/PTW_PNW_594.pdf
SHELTON AM AND JA WYMAN 1979. Potato tuber worm damage to potato grown
under different irrigation and cultural practices. Journal of Economic
110
111
Integrated Pest Management dealing with

potato tuber moth and all other pests in
Australian potato crops
Paul Horne and Jessica Page
IPM Technologies Pty Ltd, PO Box 560, Hurstbridge, Australia 3099
Abstract
The potato tuber moth (PTM), Phthorimaea operculella (Zeller), is a serious pest of
potatoes and other solanaceous crops in Australia. The pest status varies in different
regions of Australia and related factors include insecticide resistance and the level of
other pests (including other Lepidoptera) associated with potatoes. Growers in several
States of Australia have implemented Integrated Pest Management (IPM) strategies
with the assistance of IPM Technologies Pty Ltd and this has involved monitoring
crops for all pests and making recommendations for their control. This paper reviews
the approach that we have taken with regard to IPM for PTM in general and to IPM
in potato crops in particular. This is not a review of potato tuber moth control in
Australia but is a report on how we have implemented an IPM approach in potato
crops in Australia.
Introduction
Our work on potato tuber moth (PTM), Phthorimaea operculella (Zeller), began in
1986 when organochlorine insecticides (including DDT and dieldrin) were still
permitted for use on some potato crops in Australia, and use of organophosphates and
synthetic pyrethroid insecticides were standard practice. All organochlorine
insecticides except endosulfan were withdrawn from all agricultural use in Australia
in 1987 and potato farmers were looking for new control measures. This change in
the availability of pesticides allowed us to begin research into alternative control
options, including IPM rather than simply looking for substitute chemicals.
For PTM, which is the most important pest in potato in Australia, extensive
literature on its biology and biological control was available from Australia (CALLAN,
1974; BRIESE, 1981) as well as of other countries. We studied the relative importance
of different species of PTM parasitoids and it could be shown that parasitism varies
according to location (HORNE, 1990). IPM requires an integrated set of control
options for all pests, both major and minor, that are encountered in any crop. Potato
crops are no different and there are beneficial species that occur in potato crops
worldwide, cultural options that are available and also selective pesticide options.
112 IPM of PTM in Australia
The objective of our studies was the development of an IPM system for all potato
pests. We describe here how this IPM approach to potato pests was developed for
Australian potato cropping systems.

Our implementation of IPM has been mainly in Victoria, in southern Australia, where
we are based. We have monitored crops for farmers at weekly intervals from crop
emergence to senescence or in some cases until harvest. However, we have helped
potato farmers all over Australia to successfully implement an IPM strategy on their
farms.
As in any IPM strategy, the primary controls are biological and cultural,
supported by selective pesticides, only when necessary. Broad-spectrum insecticides
such as synthetic pyrethroids, organophosphates, and others must be avoided as foliar
sprays. The major pests that are encountered in Australian potato crops are PTM, the
polyphagous cotton bollworm (Helicoverpa armigera Hubner), green peach aphid
(Myzus persicae (Sulzer)), potato aphid (Macrosiphum euphorbiae (Thomas)), onion
thrips (Thrips tabaci Lindeman), tomato thrips (Frankliniella schultzei (Trybom)),
western flower thrips (F. occidentalis Pergande), whitefringed weevil (Naupactus
leucoloma (Boheman)) (all exotic species to Australia), looper caterpillars
(Chrysodeixus argentifera (Guenee) and C. eriosoma (Doubleday)) and potato
wireworm (Hapatesus hirtus Candeze), but there are many other minor, local or
infrequent pests. Potato wireworm in Australia is different from species of potato
wireworm elsewhere in the world and has a very different life-history (HORNE and
HORNE, 1991a). A range of native and introduced natural enemies of these pests are
used in IPM strategies. The most important of these are native predators: damsel bugs
(Nabis kinbergii Reuter), pentatomid bugs (Oechalia schellembergii (Guerin-
Meneville), brown lacewings (Micromus tasmaniae Walker), ladybird beetles
(Coccinella transversalis Fabricius, Hippodamia variegata (Goeze) and Harmonia
conformis (Boisduval) and red and blue beetles (Dicranolaius bellulus (Guerin-
Meneville)). There are many parasitoids of the pests, and these include native species
that attack the native pests, and also introduced species such as Orgilus lepidus
Muesebeck, Apanteles subandinus Blanchard and Copidosoma koehleri Annecke and
Mynhardt that parasitise PTM.
Cultural controls are equally as important as the biological controls and include
soil preparation and management, irrigation, location of plantings, seed source and
variety selection. Weekly monitoring of potato crops by professional entomologists
allows decisions to be made on the relative level of pests to beneficials and the trend
in their numbers. Actions are not triggered by pre-determined thresholds. Selective
bioinsecticides such as Bacillus thuringiensis spp. kurstaki (Btk) are sometimes used,
but the main control of pests is achieved by beneficial insects and cultural means.
Growers successfully applying IPM include those producing certified seed, crisping,
processing, ware and organic crops. Monitoring required looking for all pests and all
beneficials, and making a judgment as to the likelihood of economic loss if no further
action was taken. Trends of parasitoid and predator increase rather than absolute
IPM of PTM in Australia 113
thresholds are used. If damage was considered likely then action to be taken did not
only include chemical options. Cultural controls such as rolling, watering, and
harvesting early or hilling were more important options proposed than spraying
insecticides. A major cultural control that we needed to implement was a good “hill”
made of fine soil (not cloddy soil), which provided good soil cover over developing
tubers. Soil management is a critical component of our IPM strategy.
The approach we took required monitoring for all pests that occurred during the
life of the crops and recommending actions that would control that pest without
interfering with control of other pests. That is, applying actions that would not kill
biological control agents for other potato pests. All monitoring and advice reported
here concerns commercial farms, not plot trials or small trial results within
commercial crops. They include seed, crisping, processing and ware production
crops.
Results and Discussion

When we commenced research on PTM in 1986 there was almost total reliance on
chemical insecticides as the basis for control of all pests. This appears to be the
current situation in other countries such as the USA and UK now. The potato industry
had very little knowledge of biological control agents and how these could potentially
be used in an IPM strategy. Although there was interest from the potato industry there
was also scepticism and a need for both scientific information and on-farm
demonstration of IPM. This situation is one that we have encountered in a range of
horticultural and broad-acre cropping industries and our approach to dealing with it is
by simultaneously gathering entomological information and providing commercial
demonstrations of IPM (HORNE et al., 2008).
To deal with PTM there was little information available about the level of control
that parasitoids could have in PTM infestations of potato crops in Australia. We knew
that three species of parasitoids had been successfully established in Australia
(CALLAN, 1974; BRIESE, 1981) but we found that no potato growers knew about these
species or that they could be utilized in commercial crops. BRIESE (1981) provided
information about the relative abundance of parasitoids of PTM around Australia but
did not take into account the fact that regular use of broad-spectrum insecticides was
standard practice, the role of predators or the possibilities of an IPM approach, and
really considered only a classical biological control approach.
If we were going to tell potato farmers that parasitoids were reliable control tools
to help with the control of PTM then we obviously needed to know which of the three
species were most important or abundant in the different regions of Australia. Firstly,
it was necessary to find out how well established the parasitoids of PTM were in
different locations and what level of control was being achieved. To collect such
information we needed growers to collaborate and agree not to use broad-spectrum
insecticides on certain paddocks. In order to give growers more confidence that their
crop would not be destroyed despite the fact that they were withholding their normal
applications of insecticides we provided regular information on what was happening
in their crops. Essentially we provided limited information on key pests (usually that
they were not present in significant numbers) but also on what we considered to be
key beneficial species (parasitoids of PTM and aphids).
We found that this combination of entomological research and working with
potato growers provided a very useful collaboration. We were able to show that
parasitoids of PTM and aphids were present in all areas where potatoes were grown
and that they exerted considerable control pressure on these two key pests. The main
parasitoid recovered from samples was the larval parasitoid O. lepidus, followed in
importance by another larval parasitoid A. subandinus. The egg-larval polyembryonic
parasitoid C. koehleri was found from most locations but was far less abundant than
the larval parasitoids (HORNE, 1990).
The involvement of growers in this research meant that they were suddenly aware
that there were beneficial insects in their crops as well as pest species and that they
needed to be aware of the impact of pesticides that they applied. This encouraged
them to avoid the use of routine applications of insecticides and instead to use them
strategically. Once this change in attitude happened then we had the opportunity to
help growers base their use of insecticides on the results of monitoring. Importantly,
the decision on insecticide use was now based on the results showing the levels of
beneficial species and not just pest numbers. We do not use rigid threshold levels but
rather relate pest and beneficial numbers over time and make decisions about the
likelihood of economic losses. These decisions are influenced by many factors
including soil management, time of harvest, variety, rainfall and age of plants.
Now that growers were aware that beneficial insects were in their crops they (and
us) began asking for more information about them. Could they release them into their
crops? How long did they take to breed? How did they find the caterpillars? To
answer these questions, we conducted a series of laboratory and field studies.
The high number of beneficial wasps produced from a single parasitised
caterpillar led us to look at the potential of inundative releases of the polyembryonic
species C. koehleri. We undertook studies on temperature-development as there
would be a need for such information if mass-rearing of wasps was required (HORNE
and HORNE, 1991b). Inundative releases of C. koehleri were trialled but we found this
approach to be both ineffective and unnecessary given the high level of naturally
occurring parasitism available (HORNE, 1993a). We found that the level of parasitism
of PTM was often miscalculated by entomologists and so we presented data showing
how to sample and also accurately assess the level of parasitism of PTM (HORNE,
1993b). The method we propose involves taking a series of leaf-mine samples and
relating percent parasitism to the time of emergence (cohort of PTM) rather than the
date that the sample was taken. We suspect that the problem of incorrectly assessing
parasitism still occurs in many studies.
Field studies identified O. lepidus as the most important parasitoid in potato crops
in Australia. We therefore decided to investigate the host-finding abilities of this
parasitoid to improve our ability to use them within an IPM strategy (KELLER and
HORNE, 1993).
In 1996 we created IPM Technologies Pty Ltd and ceased to be government
employees. One significant aspect of this change was that we began to monitor potato
crops for potato growers and advise on what pesticides if any were required. The
legal liability of recommending “No Insecticide Required” was a significant issue,

but we knew the potential of beneficial species. The potato growers who had been
collaborators in our research now largely became our clients wanting to implement
IPM and reduce pesticide use on their crops. The results for the growers were often
massive, with sometimes a total avoidance of insecticides over a period of 10 years,
compared to 7 insecticides per crop previous to our involvement (O’SULLIVAN and
HORNE, 2000). The growers’ involvement in previous commercial trials was critical
in giving them the motivation and the confidence to implement an IPM approach on
their whole farm. Suddenly, we needed to be aware of all pesticide inputs not only for
PTM management and we realised that we had totally ignored the potential biological
control provided by predators (as well as parasitoids) and that the cultural controls of
hilling and irrigation were at least as important as parasitoids of PTM. We now had to
deal with all pests of potatoes in a compatible way, not just promote the parasitoids of
PTM. This was entirely possible, as there are IPM compatible options for all aphids,
caterpillar and other pests of potatoes in Australia (HORNE, 2000). Table 1 provides
an example of the integration of several options in a compatible way. To assist
growers and others in recognizing the beneficial species as well as the pests, a
guidebook was prepared which was distributed to all potato growers in Australia
(HORNE et al., 2002).
Table 1. An example of a typical IPM strategy for potatoes integrating several control options.
Pest Beneficial Cultural Control Pesticide support
Soil management Spray after
PTM 3 parasitoids
Irrigation senescence
Green Peach Aphid Parasitoids
Weed control Pirimicarb
and other aphids Ladybird beetles
Damsel bugs Nucleopoly
Cotton bollworm -
Parasitoids hedrovirus
Damsel bugs
Loopers - Btk
Parasitoids
Avoid infested Insecticide before
Whitefringed weevil Carabidae
areas planting
Avoid infested Insecticide before
Potato Wireworm Carabidae
areas planting
In Australia there is a problem with tomato spotted wilt virus in potatoes

vectored by onion thrips, tomato thrips and western flower thrips. Control of these
pests without recourse to broad-spectrum insecticides is essential to overall control of
pests, including PTM (HORNE and WILSON, 2000). Obviously, broad-spectrum foliar
sprays of insecticides targeting thrips will have massive detrimental effects on the
biological control of PTM, aphids and aphid vectored diseases. Growers using IPM
understand that they have beneficial species that help to control key pests and also
avoid the development of secondary pests. Our observations showed that broad-
spectrum insecticides applied to control pests such as caterpillars can cause aphid
flare (Figure 1). We believe that this is a general principle that can be extrapolated to
many situations.
100
90
% of leaves with aphids

80
70
60
50
Insecticide
40 application
30
20
10
0
/1 99
8 98 9 9 9
5 5 /1 9 / 19 9 /199 /1 99
11 /1 12/ 1 1 /1 4 2 /1 3 3 /15
Evaluation date
Figure 1. Aphid (Macrosiphum euphorbiae) development following an insecticide application

(Permethrin, Amburh).
Conclusion
One of the main issues that we had to deal with in Australia was the integration of
several pest issues (e.g. PTM, aphids and looper caterpillars) and the recognition that
an inappropriate spray for one would disrupt control of other pests. This apparently
simple issue is a stumbling block for IPM adoption in a range of horticultural and
broad-acre crops in which we work.
It is apparent from other papers presented at this symposium that IPM in potato
crops (e.g. for PTM) is not possible for many locations (including the USA) because
of the continued use of broad-spectrum insecticides for some other pests. It simply is
not possible for potato growers to expect that they can use an IPM approach and at
the same time using a broad-spectrum insecticide spray for any pest. It has to be
realized that the control of all pests need to be considered and that the control of
some pests can disrupt the biological control of other pests.
The result of the research described here has been that Australian growers can
now implement an effective IPM strategy that includes PTM as one of the key pests
(HORNE et al., 1999; HORNE, 2000; O’SULLIVAN and HORNE, 2000). We believe that
the model of IPM in potato crops that we have developed in Australia could be
applied in any country. The advantages include reduced use of insecticides and
avoidance of secondary pests.
Acknowledgements
We would like to thank the many potato growers who have collaborated with us for
many years. In addition we would like to acknowledge the support of AusVeg and
Horticulture Australia Limited (HAL) (formerly HRDC) with several projects on
potato pests. Peter O’Sullivan was particularly important in obtaining initial
commercial observations. We also thank Dr. Jürgen Kroschel for the invitation to the
Symposium and for his encouragement and assistance in the preparation of this paper.
References
Australian Entomological Society 20: 319–326.
CALLAN EM 1974. Changing status of parasites of potato tuber moth Phthorimaea
operculella (Lepidoptera: Gelechiidae) in Australia. Entomophaga 19: 97–
101.
HORNE PA 2000. Integrated Pest Management (IPM) in use-overview.
Proceeding of the Potatoes 2000 Conference, Adelaide pp. 89-92.
HORNE PA 1993a. The potential of inundative releases as part of an IPM
strategy. 5th Applied Entomological Research Conference. Canberra, May
1992. Pest Control and Sustainable Agriculture, pp. 118-120.
HORNE PA 1993b. Sampling for the potato moth (Phthorimaea operculella) and
its parasitoids. Australian Journal of Experimental Agriculture 33: 31-96.
HORNE PA 1990. The influence of introduced parasitoids on potato moth
Phthorimaea operculella (Zeller) in Victoria, Australia. Bulletin of
HORNE PA AND JA HORNE 1991a. The biology and control of Hapatesus hirtus
Candeze (Coleoptera: Elateridae) in Victoria. Australian Journal of
Agricultural Research 42: 827-34.
HORNE PA AND JA HORNE 1991B. The effect of temperature and host density on
the development and survival of Copidosoma koehleri (Hymenoptera:
Encyrtidae). Entomologia Experimentalis et Applicata 59: 289-292.
HORNE PA AND C WILSON 2000. Thrips and tomato spotted wilt virus–on the
increase: situation report. Proceeding of the Potatoes 2000 Conference, pp.
103 – 106.
HORNE PA, DE BOER R AND DJ CRAWFORD 2002. Insects and diseases of potato
crops. Melbourne University Press. 80 pp.
HORNE PA, PAGE J AND C NICHOLSON 2008. When will IPM strategies be adopted?
An example of development and implementation of IPM strategies in
cropping systems in Victoria. Australian Journal of Experimental Agriculture,
Special Issue, in press.
HORNE PA, RAE J, HENDERSON A AND R SPOONER-HART 1999. Awareness and
adoption of IPM by Australian potato growers. Plant Protection Quarterly 14:
139-142.
KELLER MA AND PA HORNE 1993. Sources of host location cues for the parasitic
wasp Orgilus lepidus (Braconidae). Australian Journal of Zoology 41: 335-41.
O’SULLIVAN P AND PA HORNE 2000. Using Integrated Pest Management (IPM) on
farm. Proceeding of the Potatoes 2000 Conference, Adelaide pp. 93-96.
118
119
Integrated Pest Management of potato tuber

moth in New Zealand
Tim J. B. Herman
Fruitfed Supplies, PO Box 322, Hastings, New Zealand
Abstract
The potato tuber moth (PTM), Phthorimaea operculella (Zeller), is an annual pest of
potato crops grown in the North Island of New Zealand. Its control is dominated by
cultural strategies and regular applications of insecticides. Five years of research was
conducted in commercial crops of three potato growing regions of New Zealand with
the aim to develop an Integrated Pest Management (IPM) program for this very
important potato pest. Pheromone traps were used to monitor moth flight activity and
larval populations in infested foliage and tubers were regularly surveyed during the
growing season. A positive correlation was found between pheromone trap catches
and larval populations in the foliage with peaks in larval populations occurring after
peaks in the pheromone trap catches. No clear relationship could be established
between the foliar larval populations and tuber infestations of PTM larvae at harvest,
negating efforts to develop action thresholds to optimize insecticide use. It was
proposed that a risk assessment model could be developed using a scoring system to
assess factors contributing to tuber infestation.
Introduction
Potato (Solanum tuberosum L.) is grown throughout New Zealand, from subtropical
regions in the north to temperate regions of the south. Over 500,000 metric tonnes of
potato is produced on 11,000 ha each year with an approximate value of $NZ342
million. The potato tuber moth (PTM), Phthorimaea operculella (Zeller)
(Lepidoptera, Gelechiidae), is an important potato pest because of its high
reproductive potential and the inability of insecticides to prevent tuber infestation
(FOOT, 1979). In most of the North Island, PTM is an annual pest causing major crop
losses, but is only an occasional pest in the South Island.
PTM flight activity peaks in mid-summer, in the period January through March.
The timing and extent of the population growth is favoured through hot dry
conditions. Control of PTM is dominated by regular applications of broad-spectrum
insecticides during the peak of population growth but cultural control methods such
as planting depth and hilling-up, are also an integral part of PTM management
(HERMAN, 1997). For monitoring PTM, pheromones in delta-shaped sticky traps have
been recommended in commercial crops (HERMAN et al., 2005). The research
reported here had the aim of developing Integrated Pest Management strategies (IPM)
20, Advances in Crop Research 10. Margraf Publishers, Weikersheim, Germany, 119-126
120 IPM of PTM in New Zealand
for PTM in New Zealand potato crops to reduce the reliance on frequent insecticide
applications.

PTM populations in potato foliage and tubers were studied over five growing seasons
in three regions of the North Island to define the relationship between PTM
populations and tuber damage and develop action thresholds to time insecticide
applications. Research sites were located in commercial crops in Waikato and
Manawatu, for the first three seasons and in Hawke’s Bay in the later two seasons.
The climate in the Waikato region tends towards subtropical with relatively warm,
humid summers while the climate in the Manawatu can also be humid, but is cooler
on average. The summer climate in Hawke’s Bay is warm and dry.
In the first two seasons, research sites were single large plot of potatoes in
commercial crops. The plots were under normal grower management, but without
insecticide applications. PTM larval populations were sampled weekly through the
vegetative growth by visually assessing foliage of 40 individual potato plants within
the plots and recording the number of occupied PTM mines per plant. Flights of male
PTM moths were monitored by placing a pheromone trap in each plot and recording
weekly moth catches. Tuber sampling started after foliage senescence. At each
sample forty plants were randomly dug up and the tubers were counted, graded
(green or white) and assessed for PTM mining. Tuber samples were taken at two-
week intervals until harvest or, when crops were ground stored into winter, until
cooler damper winter weather started.
In the third season, crops were split in half, one half received standard grower
management (regular insecticides) and the other was under IPM management with
insecticides only being applied when they were deemed necessary. A pheromone trap
was located in both management regimes and PTM populations in the foliage and
tubers were assessed as described previously. The final two season’s research was
conducted in another region, Hawke’s Bay, to further test and validate the IPM
program.
For each data series (crop-year combination) the correlation between numbers
found in foliage and numbers caught on traps was calculated with various time-lags.
A repeated measures analysis of variance was then done on the correlation
coefficients with region x year as the between-sites factor and time-lag as the within-
sites factor, using GenStat (Version 8, VSN International Ltd, Hemel Hempstead,
UK, 2006).
Results
Sampling
The weekly scouting of potato foliage for PTM larvae was validated for accuracy by
destructively sampling a subset of potato plants in the lab each week in the first
IPM of PTM in New Zealand 121
season of research. However, there was little difference in the number of PTM larvae
found by each method (data not presented).
Pheromone trap catches and larval populations

Over the three seasons and across all regions, a consistent result that was recorded
was the relationship between moth catches in the pheromone traps and larval
infestations in the foliage; two examples are presented for two regions, Manawatu
(SW coast, North Island) and Waikato (N, North Island) in the 1997/98 season
(Figure 1 and 2, respectively). In Opiki, Manawatu, a small peak in larvae found in
the foliage in late December/early January coincided with a peak of moth catches in
the pheromone trap (Figure 1). A clearer trend was found in the following generation,
in early February, when larval numbers in the foliage were at their maximum. A third
peak in larval numbers in the foliage was observed late in the growth of the crop, in
late March, which was not clearly linked to a peak in moth catches, but the latter
appeared to be asynchronous as there were many small peaks over that time.
Similar trends were seen at Matamata, Waikato (Figure 2), where a small peak of
larval numbers and moth catches in late December lead to larger peaks in the
following generation in early February and lower peaks in larval numbers and
asynchronous moth catches through the late summer and autumn months.
A clear correlation (p<0.05) between larval numbers in the foliage and moth
catches in pheromone traps was established (Figure 3). The correlations for moth
catches and larval numbers peaking in the same week and for moth catch peaking the
week prior to a peak in larval numbers averaged 0.5 and were significantly different
to zero (0= no correlation).
moth catch
larval mines
30 6
moth catch per day
larvae per plant
20 4
10 2
0 0
D J F M A M J
Figure 1. Catch of male potato tuber moths in pheromone traps and larval populations in the foliage
of potato crops in Opiki, Manawatu, during the 1997/98 season (mean of six crops).
30 0.9
moth catch
larval mines
20 0.6
moth catch per day
larvae per plant

10 0.3
0 0.0
D J F M A M
Figure 2. Catch of male potato tuber moths in pheromone traps and larval populations in the foliage
of potato crops in Matamata, Waikato, during the 1997/98 season (mean of six crops).
Average correlation between larvae per plant and
0.80 Matamata 97/98

Matamata 96/97
0.60 Opiki 98/99
Opiki 97/98
0.40 Opiki 96/97
0.20
moth catch.
0.00
-0.20
-0.40 LSD
-0.60
-0.80
in same 1 week 2 weeks 3 weeks 4 weeks
week before before before before
Figure 3. Correlation between the number of potato tuber moth larvae per plant and the catch of
potato tuber moths in potato crops in Matamata (Waikato) and Opiki (Manawatu) over
three seasons (Points between the dotted lines are not significantly different to zero).
Larval populations and tuber damage

Samples of tubers were harvested in each of the crops at fortnightly intervals, from
haulm death (natural or sprayed off) to harvest. The crops that were sampled in any
one season had different planting dates so it was possible to look at the PTM pressure
on the potato tubers over a longer time period each season.
There was no relationship between populations of PTM larvae in the foliage of
potato crops and subsequent infestation of marketable tubers of the crops (Table 1
and 2). More important was the planting and subsequent harvest date. The earliest
crop (1) had the smallest infestation of larvae in the foliage with 0.28 larvae per plant,
but then developed the highest level of white tuber infestation prior to harvest (Table
1). Conversely, two of the later crops (4 and 6) had the highest level of foliar
infestation but the lowest level of marketable tuber infestation. A similar result was
seen in the crops studied in the following season (Table 2). At this time the crops
were split into two management regimes – IPM and conventional. In the IPM crops,
the growers agreed to apply insecticides after discussion with the science team but
they were able to continue their normal pest management program on the
conventional side of the crop. A key difference to the previous season was no PTM
larvae were found in the foliage of most of the crops with one exception in one of the
conventional managed crops (Table 2). However, as observed in the previous season,
the percentage of infested tubers was again higher in the crops planted and harvested
earlier.
Table 1. Peak number of PTM larvae per plant in foliage and percent infested white tubers in
commercial potato crops in the 1997/98 season, Matamata, Waikato.
Crop
1 2 3 4 5 6
Peak #larvae/plant
0.28 0.6 0.93 1.38 0.6 1.28
in foliage
27 Jan 3.5
9 Feb 8.0
16 Feb 0.6
% 23 Feb 16.6
larvae 5 Mar 3.4 1.5
infested 16 Mar 22.6 2.7
white 23 Mar 4.1 0.6
tubers 6 Apr 0.3
14 Apr 1.4 0.2
24 Apr 0.9 0
10 May 0.3 0 0.9
Table 2. Peak number of PTM larvae per plant in foliage and percent infested white tubers in potato
crops split into two management regimes – IPM and conventional (Conv.), in the 1998/99
season, Matamata, Waikato.
1 2 3
IPM Conv. IPM Conv. IPM Conv.
Peak #larvae/plant
0 0.05 0 0 0 0
in foliage
% 22 Jan 0 0 0 0
larvae 5 Feb 0.9 0.4 0 1.1
infested 19 Feb 12.6 2.8 2.7 7.2
white 5 Mar 17.6 17.4 17.6
tubers 12 Mar 0 0
19 Mar 19.1 12.2
26 Mar 0 0
10 Apr 0.3 0
23 Apr 0 0.3
Discussion
Non-destructive, whole plant scouting was shown to be a suitable method for
monitoring potato tuber moth larvae in potato foliage as a consistent proportion of the
tuber moth larval population was found using this method. Whole plant scouting has
been previously used in PTM studies in New Zealand (FOOT, 1979) and in other
studies (LAL, 1987; LAL, 1989; SHELTON and WYMAN, 1979a; VON ARX et al., 1987;
YATHOM 1968).
Not all planned experiments could be used in the final evaluation. In some fields
no foliage infestations were found and it was noticed that farmers made at least two
insecticide applications to IPM crops without any recommendation. All the risks
associated with PTM infestation of tubers rested with the growers and they did not
have enough confidence in the unproven IPM program. Gaining the trust of growers
is an issue IPM faces in all parts of the world. Further, in one crop centre pivot
irrigation (applied weekly) suppressed the PTM larval population within the crop and
infestations were only found around the crop margin where irrigation coverage was
lower, showing that regular irrigation is an effective control strategy.
The correlation analysis of peaks in moth catches against larval numbers in the
foliage showed significant relationships for peaks in the same week and the following
week. Our results are supported by those of LAL (1989) who found a positive, linear
correlation between moth catches and foliar larval counts in the same week. He
concluded that pheromone traps could be used to predict foliar damage, negating the
need for ‘cumbersome, time consuming’ larval counts. SHELTON and WYMAN
(1979b) also found a positive, linear correlation but this was between moth catch and
foliar larval counts the following week. KEERATI-KASIKORN (1981) found a positive
correlation between moth catch and larval counts 3 weeks later. However, the
research presented here shows that foliar infestations are not necessarily an important
factor in tuber infestations.
The data presented in tables 1 and 2 suggest that for crops senescing during the
hottest summer months growers have a four week harvest window between foliage
death and possible economic losses. This is supported by other research conducted in
New Zealand (FOOT, 1976) and overseas (SHELTON and WYMAN, 1979b; VON ARX et
al., 1990; SIDDIG 1988). FOOT (1976) reported that a 2 to 4 week delay in harvest
resulted in increased levels of tuber infestation and concluded that adults are attracted
to and will oviposit in potato crops in the absence of foliage. SHELTON AND WYMAN
(1979b) found that most tuber damage occurred after vines senesced and irrigation
stopped. VON ARX et al. (1990) reported that delaying harvest by 17 days would
result in 1.7 to 4% more tuber damage and SIDDIG (1988) showed that a delay in
harvest substantially increased infestation and reduced yield.
A key factor in tuber infestation after harvest is soil cracking, opening the hills,
exposing tubers for female moths to lay eggs on. Therefore one important
recommendation that can be made is not to ground store the tubers, but to harvest
them immediately after foliage senescence (KROSCHEL, 1995). For the crops in New
Zealand, this seems to be different. We observed the highest infestation four weeks
after foliage senescing during the peak of PTM populations in February and March.
After this period, crops were much less likely to suffer tuber damage as temperature
dropped in the autumn and soil moisture increased through rainfall.
It was often found in this research that crops with bad tuber infestations were
sometimes relatively free of foliar infestations. As a result it was concluded from the
research presented in this paper that conventional IPM action thresholds, based on
infestations in the foliage were not applicable to control PTM infestations of potato
tubers.
It was concluded from the research presented in this paper that conventional IPM
action thresholds were not applicable for PTM control in potatoes. But it was
identified that there is a definite period during the growing season when potato tubers
were most at risk of infestation by PTM and that the crops had to be at an advanced
stage of growth, namely having their foliage dead or dying. Given that there is a
window during the season when the risk of PTM infesting tubers is greatest (height of
summer temperature/dryness) as well as a window in the growth of a crop of potatoes
where tubers are most likely to incur PTM damage (tuber bulking through to harvest).
It was proposed that a risk assessment model could be developed to optimise PTM
control. The model would use a scoring system to assess factors contributing to tuber
infestation. The role of pheromone traps in this system would be to indicate what the
population is doing. Because they are a good indicator of generations cycling through
the season, insecticides can be targeted at peaks in the generations so they have
maximum effect. If the risk is not too high, a grower may chose to delay an
insecticide application to allow biological control agents to work (HERMAN, 2008). If
there is a severe risk of tuber damage a grower cannot afford to delay applying an
insecticide.
Acknowledgements
The assistance of a large number of growers, technicians and casual workers are
acknowledged. Duncan Hedderley, N.Z. Institute of Crop & Food Research, did the
correlation analysis. Funding from the NZ Vegetable & Potato Growers Federation,
Technology NZ, and AGMARDT is gratefully acknowledged.
References
FOOT MA 1976. Cultural practices in relation to infestation of potato crops by
the potato tuber moth. II. Effect of seed depth, re-moulding, pre-harvest
defoliation and delayed harvest. New Zealand Journal of Experimental
Agriculture 4: 121-124.
(Lepidoptera, Gelechiidae), at Pukekohe. New Zealand Journal of Zoology 6:
623-636.
HERMAN TJ 1997. Integrated pest management for potato tuber moth, a
Technology for business growth project. Crop Info Rpt No. 443. New Zealand
Institute for Crop & Food Research. Lincoln, New Zealand. 35p
HERMAN TJ 2008. Biological control of potato tuber moth by Apanteles

subandinus in New Zealand. In: Kroschel J and L Lacey (eds.) Integrated
Pest Management for the Potato Tuber Moth, Phthorimaea operculella Zeller
– a Potato Pest of Global Importance. Tropical Agriculture 20, Advances in
Crop Research 10. Margraf Publishers, Weikersheim, Germany, 73-80.
HERMAN TJ, CLEARWATER JR and CM TRIGGS 2005. Impact of pheromone trap
design, placement and pheromone blend on catch of potato tuber moth.
New Zealand Plant Protection 58: 219-223.
KEERATI-KASIKORN M 1981. Initial studies towards the implementation of a pest
management system for the control of potato tuber moth (Phthorimaea
operculella Zeller) in south-eastern Queensland. PhD thesis. University of
Queensland.
with special references to the integrated biological control of the potato
tuber moth (Phthorimaea operculella Zeller). Tropical Agriculture 8, Margraf
LAL L 1987. Relationships between pheromone catches of adult moths, foliar
larval populations and plant infestations by potato tuberworm in the field.
Tropical Pest Management 35: 157-159.
LAL L 1989. Winter survival of the potato tuber moth, Phthorimaea operculella
(Zeller), in potato fields in India. Crop Research 27: 111-117.
SHELTON AM and JA WYMAN 1979a. Potato tuber worm damage to potatoes
under different irrigation and cultural practices. Journal of Economic
SHELTON AM and JA WYMAN 1979b. Time of tuber infestation and relationships
between pheromone catches of adult moths, foliar larval populations and
tuber damage by the potato tuber worm. Journal of Economic Entomology
72: 599-601.
SIDDIG SA 1988. Cultural means of controlling potato tuber moth (Phthorimaea
operculella Zell) and improvement of potato yield and quality in the Sudan.
Acta Horticulturae 218: 281-287.
VON ARX R, GOUEDER J, CHEIKH M and A BEN TEMIME 1987. Integrated control of
potato tuber moth Phthorimaea operculella (Zeller) in Tunisia. Insect Science
and its Application 8: 989-994.
VON ARX R, ROUX O and J BAUTGÄRTNER 1990. Tuber infestation by potato tuber
moth, Phthorimaea operculella (Zeller), potato harvest in relation to farmers’
practices. Agriculture, Ecosystems and Environment 31: 277-292.
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in Israel in the spring. Israel Journal of Agricultural Research 18: 89-90.
127
Integrated Pest Management of potato tuber

moth in India
Ravinder S. Chandel2, Virender K. Chandla1 and Ishwar D.
Garg1
1
Central Potato Research Institute, Shimla, Himachal Pradesh 171001 INDIA
2
Department of Entomology, CSK HP Agricultural University, Palampur (HP)
176062 INDIA
Abstract
The potato tuber moth (PTM), Phthorimaea operculella (Zeller), is one of the most
destructive potato pest in the field and store in India. In north western hills, the pest
completed seven generations annually and overwintered as full grown larvae. A
single generation was completed in about three weeks, when diurnal fluctuations in
temperature were 23-24 °C. Moth population was low (1-2 moths/trap/week) during
February-March, but increased quickly in April (15 to 20 months/trap/week).
Temperature was positively correlated (r=0.3662-0.7598), whereas relative humidity
(RH) was negatively correlated (r=-0.4505-0.6340) to its development. Many cultural
practices have been found to minimize tuber damage. Tubers sustained higher
infestation (5.3-8.0%) at a planting depth of 5cm as compared to deeper planting
below 10 cm (1.0-1.3%). Soil moisture markedly affected tuber infestation, with 3.5-
5.0% and 1.2-1.7% in dry soil and wet fields, respectively. Tubers were heavily
damaged in the field, particularly if harvest was delayed. In storage, the use of the
PTM granulovirus at a rate of 350 larval equivalents (LE) per kg talc powder (2.5 g
per kg tubers) gave complete control against PTM for two months.
Introduction
Potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera,
Gelechiidae), an introduced species, has become the most wide spread and
destructive insect pest of potato in field and store in India (CHANDEL and CHANDLA,
2005). Although the larvae do some feeding on potato foliage and stems, the
economic importance of this pest is in the damage it causes in tubers. The PTM is
distributed over a wide range of agro-ecological zones, and its bioecology has been
studied extensively in different parts of the world (CHI and GETZ, 1988; KROSCHEL
and KOCH, 1994; TRIVEDI et al., 1994). In India, it has been found to cause 30-70%
infestation in country stores (CHANDEL et al., 2005). Most infestations originate from
volunteer potato plants or tubers and from cull piles. In mid Himalayan region, severe
PTM infestations have been commonly associated with spring-planted potatoes,
128 IPM of PTM in India
produced specifically for processing purposes. Control measures in the field using
insecticides are not very effective (CHANDEL and CHANDLA, 2005). If tubers have
already become infested in the field, the results in storage can be disastrous as the
temperature in country stores is favourable to insect development and multiplication.
Pheromone traps could be useful tools in determining the suitable time for the
application of insecticides to reduce PTM damage and pesticide use in fields (LAL,
1993). Cultural practices associated with crop production can make the environment
less favourable for the survival, growth, or reproduction of PTM (MOAWAD et al.,
1997). The possibility of using cultural practices as a control method has been
investigated under field conditions (FOOT, 1976; SHELTON and WYMAN, 1979; ALI,
1993). The objective of this work, therefore, was to discover means of preventing
access of PTM to tubers by manipulating various cultural practices in field for
establishment of good initial storage conditions to decrease the likelihood of post
harvest losses due to PTM infestation.

The experiments were carried out at farmer’s fields and stores in Kangra Valley of
Himachal Pradesh during 1997-2004 at a latitude and longitude of 32o6'N and 76o3'E,
respectively, and an altitude of 1100 m above sea level.
PTM population dynamics

The seasonal dispersal movement of moth population using sex attractant pheromone
traps was studied at Palampur and Nagrota. When the potato shoots began to emerge,
one trap was installed at each site and maintained throughout the season. The
pheromone traps were placed on the ground just above the crop canopy in the center
of field. The pheromone capsules were replaced every 6 weeks and data on moth
catches were recorded weekly.
The studies on the biology were conducted under laboratory conditions at
Palampur. Moths were kept in plastic jars (13x10 cm) covered with double layered
black muslin cloth for egg laying. The newly hatched larvae were transferred to fresh
potato leaves and day to day observations on growth and development of various
stages were recorded.
Field Experiments
Effect of planting depth, soil moisture, dehaulming (vine-kill) and harvesting time on
PTM field infestation was studied in Malan area of Kangra valley. The field
experiments were conducted on spring-planted potatoes grown in uniform silty clay
loam soil. The variety was Kufri Jyoti and plot size was 12 m2. The spacing was 20
cm and 60 cm between plants and rows, respectively. Recommended rates of
fertilizers, and time and frequency of ridging were used, but insecticides were not
applied at all.
IPM of PTM in India 129
The effect of planting depth on PTM infestation was studied during the year
1998-1999. Medium sized cut tubers (25-30 g) were planted at 5 cm and 10 cm
during mid January. Each treatment was repeated 15 times. The percentage of
infested tubers was calculated at harvesting on plot basis. To record the effect of soil
moisture on PTM infestation, irrigation intervals were regulated. The field was
prepared and divided into 30 small plots of 12 m2. Treatment started four weeks after
planting. The plots were irrigated using furrow irrigation and the water was applied
until its level reached 15-20 cm up the ridges. Half set of plots (n=15) were irrigated
three times at 30, 50 and 80 days after planting. In remaining plots (n=15), irrigation
was applied 8 times at 30, 40, 50, 60, 70, 80, 90, and 100 days after planting. At
harvesting, percentage of PTM infestation was assessed in each plot.
The dehaulming effect on PTM infestation at harvesting was studied at three sites
during 2004-2005. The crop was planted in mid January in plots of 12 m2 and
harvested in mid May. At maturity, vines were cut and removed from fields, 10 days
before harvesting in half set of plots (n=15 plots). Equal numbers of plots were
maintained without dehaulming and direct harvesting was done. The component of
harvest time x PTM infestation was tested repeatedly in on-farm trials during 1999
and 2000. The crop was harvested at weekly interval starting from May 1, 8, 15, 21
and 28 during both years. At harvesting, healthy and infested tubers were separated
and per cent PTM infestation was calculated. For multiple regression analysis, the
PTM infestation was taken as dependent variable (Y), while dates of harvest (X) were
considered as independent variable. The non-linear polynomial equation was found to
be the best fit on the basis of goodness of fit (R2).
Storage Experiments
To obtain an accurate picture of losses under farmer’s storage condition, damage was
assessed regularly. A total of 62 stores were evaluated over a period of two years.
The stores were visited frequently between May-June and the information on
traditional storage methods was collected and analysed to find their correlations with
PTM infestations. In 2003, five Indian potato cultivars were evaluated against the
PTM under farmer storage conditions at Malan. The experiment was conducted in a
single store. Each variety was replicated three times; each replicate consisted of about
20 kg tubers stored in a bamboo basket.
The effectiveness of wheat straw and PTM-specific granulovirus (PoGV) was
investigated in stored potatoes over a period of 3 months during 2004. Treatments
consisted of 60 kg potato divided into 3 replications for every treatment and placed
inside the rustic stores at ambient temperature. Wheat straw was used as a complete
layer of about 1 cm over stored potatoes. PoGV dust at 350 larval equivalents (LE)
per kg of talc powder was mixed with tubers in plastic bags (50 g/20 kg tubers)
before storing tubers in bamboo baskets.
Results
PTM population dynamics

PTM was active from March-October in Kangra valley, with peak activities from
April to end of July. In the Northwestern hills, the pest completed seven generations.
Under laboratory condition, female moths laid 72-127 eggs during her life time and
the maximum number of eggs was deposited 3-4 days after emergence. Eggs hatched
in 4-6 days and ranged between 80.0- 93.6%. Larvae have four distinct instars and
each successive instar lasted for 4 to 6, 2 to 6, 2 to 6 and 3 to 7 days, respectively.
Under normal conditions, larvae became fully grown in 12-24 days. In winter, fourth
instar larva of 7th generation hibernated for 80-84 days during December-March. The
pupal stage lasted for 5-15 days. The longevity of female moths (9 to 14 days) was
more than of male moths (7-13 days). Sex ratio (M:F) of 1:1.2 showed slight biasness
for females.
During the summer, PTM completed a generation in 22 to 25 days, when mean
temperature ranged about 23-24oC and this temperature was found to be the most
favourable for this species. During the cool season from February till March,
population remained low (1-2 months/trap/week) and PTM did not cause significant
damage to the crop. However, populations developed quickly from the beginning of
April (15-20 moths/trap/week).
The simple correlation with mean temperature was found to be positive and
correlation coefficients were 0.3662 and 0.5612 for Palampur and Nagrota,
respectively. However, the relative humidity (RH) showed a negative correlation in
both the regions (r=-0.4505 to -0.6340). The simple linear regression analysis
between temperature and PTM population indicated gradual increase in population
with rise in temperature at Palampur and explained around 44% variation in moth
population. However, at Nagrota the data were less consistent (Figure 1) and the
results were less conclusive (with low value of R2=0.2586). The regression equation
calculated for RH indicated that with rise in relative humidity the PTM population
decreased considerably (Figure 1), although the values of R2 were low (0.1483-
0.2955).
Field Experiments
During both the years, PTM infestation decreased with deeper planting of potato
tubers. Tuber infestation varied between 5.3-8.0% and 1.0-5.3% at a planting depth of
5 cm and 10 cm, respectively, with significant differences in 1998 (Table 1).
Soil moisture markedly affected tuber infestation, but was not significantly
different between treatments. In dry soils, PTM infestation varied from 3.5-5.0%,
whereas in wet plots average damage ranged from 1.2-1.7% (Table 2).
Removal of senescing plants (dehaulming) and their destruction reduced tuber
infestation at harvest. The field infestation of PTM ranged from 7.5 to 10.3% without
removal of plants compared to 2.8 to 3.5% with removal 10 days before harvest
(Table 3). However, the differences in PTM infestation were not significant.
Palampur Nagrota
200 120
y = 4.8475x - 50.589 y = 2.5168x - 23.526
100
Moth population
150 R2 = 0.4415 R2 = 0.2586
Moth population
80
100
60
50 40
20
0
0 5 10 15 20 25 30 0
-50 0 5 10 15 20 25 30
Temperature °C Mean Temperature °C
Palampur Nagrota
180 120
160 y = -0.9317x + 61.544
100
140
R2 = 0.2955
Moth population
Moth population
120 80
100
60
80
y = -0.6267x + 61.128 40
60
R2 = 0.1483
40 20
20
0
0
-20 0 20 40 60 80 0 20 40 60 80
-20
Relative humidity (%) Relative humidity (%)
Figure 1. Regression analysis of the effect of temperature (°C) and relative humidity (%) on potato
tuber moth catches in pheromone traps in the Kangra Valley, India.
Table 1. Potato tuber moth infestation of potato cultivar ‘Kufri Jyoti’ planted at two depths.
PTM incidence at harvest (%) during
Tuber depth (cm)
1998 1999
1
5 cm 8.0a ± 2.81 5.3a ± 3.2
≥10 cm 1.3b ± 1.1 1.0a ± 0.9
1
Different smaller case letters represent significant subsets at P≤ 0.05.
Table 2. Effect of soil moisture on potato tuber moth infestation at harvest, 1999-2000.
% PTM infestation during
Treatment
1999 2000
Dry soil (3 irrigations) 5.0 ±2.04 3.5 ±1.36
Wet soil (8 irrigations) 1.5 ±1.23 1.2 ± 0.7
Treatments were not significantly different from one another at P≤ 0.05.
Table 3. Effect of dehaulming on potato tuber moth infestation at harvesting.

State of dehaulming before % PTM infestation during
harvesting 2004 2005
Yes (10days before digging) 2.8a ±1.15 3.5a ±1.41
No (directly harvested) 7.5a ±1.58 10.3a ±2.69
Treatments were not significantly different from one another at P≤ 0.05.
PTM infestation increased sharply near crop maturity. Data recorded during 2004
and 2005 demonstrated that, on May 1, PTM infestation varied from 1.9 to 2.4%
during 2004 and 2005, respectively. There was a gradual increase in PTM infestation
up to third week of May (19.3 to 24.9%) during both years. Regression analysis
revealed that PTM infestation peaked around May 25 during both the years (Figure
2). The rate of increase of PTM infestation followed an increasing trend up to May
12. Thereafter, infestation increased but at a lower rate.
2004 2005
30 25
y = -0.0025x 3 + 300.68x 2 - 1E+07x + 2E+11 3 2
y = -0.002x + 241.22x - 1E+07x + 1E+11
25 R2 = 0.9986 R2 = 0.9954
20
% infestation
20
15
% infestation
15
10
10
5 5
0 0
27-Apr 2-May 7-May 12-May 17-May 22-May 27-May 1-Jun 27-Apr 2-May 7-May 12-May 17-May 22-May 27-May 1-Jun
Harvesting dates Harvesting dates
Figure 2. Effect of harvest date on potato tuber moth infestation in spring potato, Kangra Valley,
India.
Management of PTM in storage

Farmer’s storage practices were found to increase the PTM tuber infestation. Most of
the farmers simply store potato under low cost rustic unrefrigerated conditions with
temperature ranging between 25 and 30oC. Further, the storages are not physically
protected enough to hinder moths from entering stores. Hardly any wire mesh screens
on windows were installed. The doors were kept open particularly at dusk to provide
adequate storage ventilation. In the stores surveyed for PTM infestation after one
month of storage in June, the mean PTM infestation varied between 12.5 and 13.6%
and the rotting was 6.0 to 9.3%. No significant differences in infestation were found
between the varieties (Table 4).
Table 4. Influence of Indian potato cultivars on potato tuber moth infestation.
Kufri Cultivars Uninfested tubers (%) Infested1 (%) Rottage (%)
Jyoti 78.8 12.7 8.5
Chandermukhi 80.4 13.6 6.0
Giriraj 78.3 12.45 9.25
Chipsona – I 79.7 13.3 7.0
Chipsona – II 80.5 12.75 6.75
1
No significant differences of potato tuber moth infestation at P≤ 0.05 were found among the
potato cultivars.
Fewer storage losses were observed where good storage conditions were
maintained. Tuber inspection and removal of infested tubers at periodic intervals
were important factors affecting PTM infestation in storage. More than 50% of the
farmers are monitoring potato storage conditions for rotting and PTM infestation. The
potato stores were opened every 3 to 4 weeks and the tubers with black spots were
sorted out. This process is locally called palta. The tubers which were at the top are
rotated down during inspection. It was observed that rotting was positively correlated
with infestation (r=0.43).
Almost 50% of farmers relied heavily on insecticide application to tubers at the
beginning of the storage period mainly using methyl parathion or fenvalerate which
are easily available in the market (Figure 3). Some farmers mixed synthetic
pyrethroids like cypermethrin or deltamethrin with soil at low doses and broadcasted
this insecticide dust on tubers. Plant products like dry leaves of Ageratum
houstonianum Mill., Lantana camara L. and Eucalyptus terticornis Sm. are also
commonly used. It was observed that 20.0 to 28.5% of the farmers covered their
stored potatoes with either of the plant products as a preventive measure against
PTM. On the basis of 62 stores surveyed during 2002 to 2004, the mean PTM
infestation was 17.6 to 28.0%. Covering tubers with about 1 cm layer of wheat straw
afforded significant control in stored potatoes. After 60 days of storage, the incidence
of PTM infestation was less than 1.0%. Similarly, PoGV (350 LE/kg powder) applied
as a dust gave complete protection against PTM for two months (Figure 4). In
controls, there was 26.0% PTM infestation as compared to 0.8% in wheat straw
treatment after 60 days of treatment. There was a gradual increase in infestation, and
after 90 days 53.0% infestation was observed in controls, however, in treatments the
infestation varied from 4.1 to 6.5% only.
60
Farmers PTM control practices
% and mean PTM infected of
48 46.7 2002
50 2003
42.8
stored tubers (%)
2004
40
26.7 28.5 28.5
30
20 21.8
17.6
20
10
0
Insecticides Plant Products PTM Infestation
Figure 3. Percentage of farmers using insecticides and/or leaves of different plants to control the
potato tuber moth, and incidence of infestation in potato stores, Kangra Valley, India.
102
Reduction of PTM damage (%)
100
100
PoG V
98 9 6 .9
W h e a t s tr a w
96
94
9 2 .2
92
90
88 8 7 .7
86
84
82
80
60 days 90 days
Figure 4. Efficacy of PoGV and wheat straw to control the potato tuber moth in storage as
evaluated after 60 and 90 days, respectively.
Discussion
The number of PTM generations produced in a year is mainly dependent upon the
agro-climatic conditions prevailing in a particular region. The number of generations
found in the present study for the Kangra valley is in accordance to observations of
SHAHEEN (1983) for Egypt and BOZKURT and GENC (1994) for Turkey. GUBBIAH and
THONTADARYA (1977) reported 13 generations per year in South India and in
Bangladesh 18 generations are completed (Kabir, 1994). Developmental times of egg,
larva and pupa reported here do not differ substantially from results obtained by
CHAUHAN and VERMA (1990) in India and KABIR (1994) in Bangladesh. The
temperature in the present studies fluctuated from 17.7 to 24.1oC. Under these
temperature conditions the development period of larvae was more sensitive
compared to eggs and pupae. There exists considerable differences in total fecundity
per female and the results are in agreement with the observations of earlier workers
from India, USSR and Bangladesh (GLEZ, 1989; CHANDLA, 1986; KABIR, 1994).
The development of the adult moth population exhibited the same pattern at both
the research sites in the Kangra Valley. Early in the potato season, the PTM
population increased slowly until the end of February, but with a temperature
increase thereafter the PTM population increased rapidly until harvest. TRIVEDI et al.
(1994) demonstrated that the development time from egg to adult varied considerably
in relation to the prevailing temperature requiring 35 days at 17.7oC and 22 days at
24.1oC, respectively. Such a pattern of population build up is similar to those reported
by CHANDEL et al. (2001) and LAL (1989) in India. The regression analysis indicated
a strong relationship between PTM activities, as monitored by pheromone traps, and
temperature. This information is important to understand peak activity periods and to
design suitable management strategies to reduce PTM infestation.
The results suggest that significant reduction in PTM infestation in the field could
be achieved by deep planting of tubers, followed by a proper hilling. Both deep
planting and hilling up kept the tubers beyond easy access of PTM moths as also
demonstrated ALI (1993) and SILESHI and TERIESSA (2001). The adult moths cannot
easily reach tubers covered with more than 5 cm of soil, unless it is deeply cracked.
FOOT (1976) also demonstrated that PTM infestation is inversely proportional to
planting depth and number of hilling operations. Similarly, soil moisture caused
considerable reduction in PTM infestation at harvest. The PTM infestation was
directly proportional to the length of irrigation intervals, which is in agreement with
the findings of Shelton and WYMAN (1979). Maintaining regular irrigation until
harvest reduced soil cracking and thus prevented the exposure of tubers to PTM
female moths (HANAFI, 1999). Furrow irrigation is a common practice in parts of
Himachal Pradesh. With furrow irrigation, furrows and hills should be well
maintained to prevent wash out and exposure of tubers.
Removal of senescing vines has been found to affect the level of infestation as it
affects the exposure of tubers. As the vines wilt, PTM larvae leave the vines and
infest the tubers underneath. In Ethopia, harvesting tubers before 70% of the
vegetative growth dried up has been recommended to minimize exposure and
infestation (SILESHI and TERIESSA, 2001). PTM infestation increases sharply near
crop maturity. Leaving potatoes in the ground until the skin is properly hardened,
improves the tuber storage quality, but it delays the harvest and increases the chances
of tuber exposure to moth infestation. The study results suggest a potato harvest in
early May to prevent tuber damage. The importance of timely harvest has been also
demonstrated by VON ARX et al. (1987) and HANAFI (1999) in North Africa.
Management of potato tuber moth in storage

There is a typical price rise in potato wholesale between July and August and hence
70 to 80% of the spring potato production in Himachal Pradesh is held in rustic
stores. PTM, introduced from fields to stores at harvest, poses a substantial threat to
the stored potatoes through out the summer storage period due to high temperatures.
The stores are low cost and non-refrigerated generally without any screens on
windows to prevent adults from gaining access to stores.
Ambient climatic conditions are favorable to store potatoes for 2 to 3 months, but
these conditions are also ideal for PTM development. In our survey, none of the
stores was without PTM and the average tuber infestation was 30%. Low infestation
was related to good storage conditions like wire mesh on windows and removal of
infested tubers during storage. Periodic inspection is recommended at an interval of 2
to 3 weeks. The young larvae are difficult to see and their damage is inconspicuous
(CHANDEL et al., 2001), hence sorting becomes difficult at an early stage. The
damage becomes conspicuous as the larval stage advances and larvae are fully grown
in 12 to 13 days.
The level of infestation did not differ significantly between cultivars despite
differences in tuber shape, skin characteristics and dry matter contents. The dry
matter in these varieties ranged from 21 to 25% (GAUR et al., 1999). The overall loss
due to PTM damage and subsequent tuber rotting was estimated at 19.5 to 21.6%.
More than 50% of the farmers applied various PTM control interventions in
storage. Dry leaves of different plants used to cover tubers reduced the infestation
considerably. They have dual functions of forming a physical barrier to the egg
laying moths and also acting as repellent (KROSCHEL, 1995). The wheat straw used in
this study acted as a mechanical barrier for PTM adults to reach tubers.
Biological control using PoGV was very effective in preventing tuber infestation
over a period of 60 days. MAOWAD et al. (1997) used PoGV in Egypt at a rate of 80
infected larvae per ton of tubers and achieved a 75% reduction of PTM infestation
after two months of storage. Earlier, a substantial amount of successful testing of
PoGV has been conducted in the Andean countries and the Middle East, and PoGV
has been commercially produced in Peru, Bolivia, Egypt, and Tunisia using low cost
facilities for propagation (see review by LACEY and ARTHURS, 2008, in this volume).
Hence, the production technology used in other countries could be used to develop
PoGV as a bioinsecticide for India.
Conclusions
PTM is a serious potato pest in the Kangra Valley of the Himachal Pradesh, in India.
During the potato growing season the PTM population increases rapidly causing
sever tuber damage at harvest. The study showed that cultural practices can
significantly contribute to reduction of tuber infestation at harvest. This is crucial to
reduce initial infestation of stored potato and population build-up in storage. The
application of physical barriers like wheat straw to cover tubers helps preventing
tuber damage but should be combined with direct control tools. Further research into
the use and adoption of PoGV for PTM storage management is suggested.
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139
Commercialization of potato tuber moth

resistant potatoes in South Africa
Dave S. Douches1, Johann A. Brink3, Hector Quemada3,
Walter Pett2, Muffy Koch4, Diedrich Visser5, Karin Maredia3
and Kelly Zarka1
1
Department Of Crop And Soil Sciences; 2Department Of Entomology;
3
Institute Of International Agriculture, Michigan State University, East Lansing,
MI 48824; 4AGBIOS, Box 475, Merrickville, ON, K0G 1N0, Canada; 5ARC-
Roodeplaat, Vegetable And Ornamental Plant Institute, Private Bag X293,
Pretoria, 0001, SOUTH AFRICA
Abstract
The potato tuber moth (PTM), Phthorimaea operculella (Zeller) is a major pest
problem facing potato farmers in developing countries. Currently, the primary means
to control the PTM and avoid major crop losses is the use of chemical pesticides.
Michigan State University, funded by the U.S. Agency for International
Development through its Agricultural Biotechnology Support Project, initiated
biotechnology research on the development of PTM resistant varieties in 1992. A
Bacillus thuringiensis Berliner (Bt)-cry1Ia1 gene, was obtained from ICI Seeds (now
Syngenta seed company) and successfully introduced into several potato varieties,
including Spunta. Transgenic lines were shown to have complete efficacy against
PTM. This Bt-potato will be one of the first public sector developed products to
reach farmers in developing countries and will serve as a model for the public sector
deployment of insect resistant transgenic crops. The commercialization project
includes six components: 1) Product development, 2) Regulatory file development,
3) Obtaining freedom to operate on intellectual property/proprietary technologies and
establishing licensing relationships, 4) Marketing and technology delivery, 5)
Documentation of socio-economic benefits, and 6) Public communication. The
expected benefits of this Bt-potato to farmers and end-users will be increased
marketable yield, improved quality, reduced storage losses, reduced post-harvest
losses and reduced human exposure to pesticides.
Introduction
The Agricultural Biotechnology Support Project (ABSP), funded by the U.S. Agency
for International Development (USAID), was established in 1991 as the premier US-
sponsored agricultural biotechnology program designed to assist developing countries
in accessing and using biotechnology to address local agricultural constraints. ABSP
worked in Asia, the Middle East, Latin America and Africa to develop a number of
140 Commercialization of PTM resistant potatoes
crops with improved agronomic traits. ABSP, managed and implemented through
Michigan State University (MSU) in collaboration with other US universities and the
private sector, has integrated research, product development, and policy/regulatory
development to assist developing countries in accessing and generating
biotechnology and in establishing a regulatory framework for production of biotech
crops (Figure 1). ABSP has also been successful in assisting with the development of
appropriate policy frameworks in developing countries through technical assistance
in regulatory development and intellectual property rights.
networking
research ABSP policy
management
Figure 1. ABSP’s integrated approach to biotechnology development and transfer.
Our project team believes that the tools of biotechnology, including tissue culture,
molecular markers, molecular diagnostics, and genetic engineering, can be applied
safely and effectively to address crop production constraints in developing countries.
Managed properly and safely, biotechnology can play an important role in improving
food and forage crop production, and thus enhancing food and feed security.
Scientific solutions to improve crop productivity, where biotechnology can play an
important role, can empower the rural sector, by boosting food production, enhancing
income for the small farmer and improving their nutritional security. This project has
focused on developing and commercializing a Bt-potato variety to manage resistance
to potato tuber moth (PTM), Phthorimaea operculella (Zeller). The project goals are
to provide safe, economic control of PTM via Bt-based host plant resistance. With
this development, farmers may be able to reduce application of chemical insecticides
in the field and should be able to reduce tuber breakdown in storage due to PTM
feeding.
The cultivated potato, Solanum tuberosum L., is one of the world's major food
crops following rice (Oriza Sativa L.), wheat (Triticum aestivum L.) and maize (Zea
mays L.) in importance (ROSS, 1986). The potato is widely grown over many
latitudes and elevations in 130 of the world's 167 independent countries. PTM is one
Commercialization of PTM resistant potatoes 141
of the most serious insect pests of potatoes worldwide. It is of greatest importance in

sub-tropical and tropical latitudes, including the southern and southwestern US.
Insecticide use is the most common means of PTM control in both field and storage.
Twelve to twenty insecticide applications may be used to control PTM during the
growing season, and in storage, three to four insecticide sprays may be applied in
storage for PTM control, with the last application within one week of marketing
(MADKOUR et al., 1999).
Bt-potato project
The introduction of the Bacillus thuringiensis Berliner (Bt) protein gene via genetic
engineering offers a form of plant resistance against PTM. The high dose expression
of Bt in crop plants offers an ecologically sound means to control specific crop insect
pests (MCGAUGHEY and WHALON, 1992). We focused on utilizing the codon-
modified Bt-cry1Ia1 gene from the Syngenta Company (formally ICI Seeds). ICI
Seeds was an ABSP partner that developed the Bt-cry1Ia1 gene initially targeted for
corn transformation (TAILOR et al., 1992). Through a research agreement, ICI Seeds
extended the use of this gene for transformation into potato. Use of the Bt-cry1Ia1
gene provided a means to achieve our research goal of high expression of Bt in potato
(Figure 2). Transformations in potato with a codon-modified Bt-cry1Ia1 gene
(effective against both lepidopteran and coleopteran insects) have produced high
levels of Bt expression with 80 to 100% insect mortality in detached leaf laboratory
bioassays (LI et al., 1999; WESTEDT et al., 1998; MOHAMMED et al., 2000).
This research has led to germplasm that has commercial potential in the U.S. and
abroad (Egypt, South Africa, Argentina, Mexico and Indonesia) (MOHAMMED et al.,
2000; DOUCHES et al., 2002). With transgenic material in hand, the MSU potato
research team was able to distribute the Bt-potatoes and establish field tests in Egypt
(DOUCHES et al., 2004) and South Africa, and to conduct greenhouse tests in
Indonesia and Peru (LAGNAOUI et al., 2001) (Figure 3). A field trial planted in 1997
was the first transgenic field trial ever conducted in Egypt. Since then, yearly trials
were conducted in Egypt until 2001 and then in South Africa from 2001 to the
present. Potential target countries for Bt-potato commercialization include Egypt,
South Africa, Indonesia, India and Mexico.
South Africa was selected as the target country for product commercialization for
various reasons. First, PTM is an important constraint in South Africa (VISSER, 2005)
and product efficacy of the Bt-potato was demonstrated in contained field trials.
Secondly, the commercial potato industry in South Africa (Potatoes South Africa) is
well developed and it, along with the Agriculture Research Council (ARC), has had
previous in-country experience with genetically modified (GM) potato research. To
complement the potato industry, there is a well established enabling environment for
commercialization of GM crops – cotton (Gossypium barbadense L.), maize (Zea
Mays L.) and soybean (Glycine max L.) have been commercialized. A functioning
biosafety regulatory framework exists in South Africa.
oriV
Km (E.coli)
RB
Nos
11292 NptII pSPUD 3764

15057 bp
T
CaMV35S
Bt-cry1Ia1 LB
BamHI T
7528 BamHI
Figure 2. The Bt-cry1IA1 vector construct (pSPUD5) used in Agrobacterium tumefaciens
transformations with Spunta. The pBI121 binary vector contains the Bt-cry1IA1 gene in
place of the gus gene.
Michigan
State
University
Egypt Indonesia
(Field trial) (Laboratory trial)
Peru
(Greenhouse trial)
South Africa
(Field trial)
Figure 3. International testing of Bt potatoes developed through USAID.
The consortium for commercializing the Bt-potato project includes multi-

disciplinary partners from three continents. In North America the team members
include primarily MSU and AGBIOS (Canada). The International Potato Center
(CIP) in Peru is also a project partner and the main in-country partner in South Africa
is the Agricultural Research Council (ARC).
Technology development and proof of concept were established many years

before the Bt-potato project reached South Africa. Consequently, product
development has been a major thrust of the project in South Africa (Figure 4). In the
US, agronomic trials of the Bt-potato lines have been conducted since 1994. The
results demonstrate that Spunta-G2 has equivalent yield, type, tuber size distribution,
specific gravity and internal defects frequency as the non-transgenic Spunta. PTM
efficacy trials were also conducted from 2004 to 2006 in Washington. Spunta-G2
foliage had no mining damage from the adults and the tubers were free of mining
damage in post harvest assays (L. LACEY unpublished data). PTM field and storage
trials have given comparable efficacy (MOHAMMED et al., 2000). Multi-location
agronomic and PTM efficacy trials (field and storage) have been conducted at six
sites in South Africa since 2001. Complete control of the PTM was found at all
locations in all years as seen at Ceres, South Africa (Figure 5). There was no
infestation in the field when examining the foliage and tubers of Spunta-G2 and no
infestation was found in tubers stored up to 6 months.
Figure 4. Transgenic product development overview.
To move beyond the first phase of research and have greater impact, the cry1Ia1
gene construct was used to transform two popular South African potato cultivars. We
currently have produced Bt-potato lines of these cultivars, confirmed by PCR and
Southern analysis. In addition, we have used Spunta-G2 as a parent in our breeding
program at MSU. Currently there are 15 selections, which are Bt-positive and PTM
resistant that are derived from crosses with late blight resistant parents (data not
shown). Based on existing GM crop approvals, these Spunta-G2 derived progeny are
likely to be approved under a general release approval for Spunta-G2, when this is
obtained in South Africa.
With field testing in place in target areas, we are currently addressing the
intellectual property rights (IPR) of the Bt-potatoes, food safety and socioeconomic
issues that are specific to South Africa’s regulatory approval requirements (MORRIS
and KOCH, 2002).
14
Number of Living 12
10
Larvae/Stem
8
6
4
2
0
Spunt BP1 G2 G3
7-Jan 22-Jan
Figure 5. Number of potato tuber moth (PTM), Phthorimaea operculella larvae per stem in field
trials in non-transgenic (Spunta, BP1) and transgenic (Spunta-G2 and Spunta-G3) potato
varieties. Ceres, South Africa.
Biosafety, food safety and IPR regulatory issues

associated with Bt-potatoes
The safe and legal application of biotechnology requires appropriate and functioning
biosafety and IPR systems. Biosafety encompasses policies and procedures to ensure
environmental, food and feed safety of applications of biotechnology. The IPR and
biosafety policies must be in place both at national and institutional levels to ensure
safe and legal exchange of technology. Appropriate IPR agreements giving freedom
to operate (FTO) must be in place before proprietary biotechnology products can be
utilized.
Considering the importance of the biosafety, food safety, and IPR issues
associated with Bt-potatoes, we have from the beginning of the project put concerted
efforts into building capacity in these areas in collaborating countries. Our team
views biosafety and IPR as integral parts of the technology transfer process and has
always ensured that all the legal and biosafety requirements of recipient countries are
met before any technology is exchanged. The technology transfer process has been
carried out in a safe and legal manner. The integrated approach to technology transfer
includes the following steps:
1) Preparation of biosafety permit applications.
2) Review by the National Biosafety Committee of the recipient country.
3) Development of Material Transfer Agreements (MTA) encompassing both IPR
and Biosafety elements.
4) Issuance of U.S. Phytosanitary Certificates for importation of material into
recipient countries.
5) Import permit applications - Plant Quarantine Department of the recipient

country.
6) Transfer and storage of regulated plant materials.
7) Greenhouse and field tests in local environments, under permits for contained
use or for contained field trials and the supervision of biosafety officers.
These project activities have significantly contributed to capacity building in public

sector biosafety and IPR, and have helped strengthen the national biosafety and IPR
frameworks in Egypt and Indonesia.
Product commercialization approach

There are many other components of the Bt-potato project besides product
development that must be addressed for the project to succeed. Commercialization of
a GM product requires a comprehensive approach as outlined in Figure 6. To support
an application for commercial release of the Bt-potato, a regulatory file containing
extensive safety data must be generated. Components include protein expression,
toxicology evaluations, substantial equivalence testing and socioeconomic studies. A
marketing strategy must be developed in consultation with the local potato industry.
To have commercial impact, we also need a product delivery mechanism that must be
coordinated with the potato industry and potato seed growers. It is imperative for
Freedom to Operate (FTO) to be granted by the main IP holders through the
development and establishment of valid licensing agreements. By developing a
stewardship program potential liability issues can be identified and addressed before
actual commercialization of the product. Effective and regular communication with
the industry producers and consumers must occur so that transparent and
scientifically sound data are made available in a timely manner. We have had to work
closely with the South African potato industry and other stakeholders to address
consumer acceptance strategies. Post-approval strategies addressing product
stewardship and regulatory compliance (including post-market monitoring) must also
be defined.
Figure 6. Product commercialization approach.

Lessons learned
Over the course of the Bt potato project, management aspects such as budgets,
personnel, technology, regulations, etc., changed over time. Our commercialization
consortium learned to adjust to these changes. Along the way valuable lessons were
learned:
- Communication between collaborating scientists is very important. Cultural
differences can limit or impede communication.
- Training of developing country scientists is valuable. Training helps with
effective collaboration and establishing good communication. In addition, long-
term training was more valuable and effective than short-term training.
- The research program goals must provide clear actions and must address social,
economic, and ethical issues surrounding the research to gain confidence of all
stakeholders.
- Food safety issues of Bt-crops became larger than anticipated as the research
progressed.
- World-wide anti-GM issues developed during the project and limited progress
towards commercialization. GM plants have become more regulated rather than
less regulated.
- GM crops became a trade barrier during the project and limited
commercialization options due to fear of market loss in Europe.
- Close linkage between lab and field research is needed to ensure progress
towards commercialization.
- Vector construction is a continuous process, not a final step. Many transgenic
lines must be generated and this process needs to be an on-going objective in the
work plan.
- IPR and FTO on proprietary technology are a changing target.
- Biosafety and IPR issues are as important as research when commercialization is
the final goal.
- Research drives the development of policies. Transgenic plants must be available
to provide a reason for establishing field testing and food safety regulations.
Why is the Bt-potato project important?

The PTM resistant potato will be one of the first public sector-developed products to
seek approval and deployment in a developing country. This product has the potential
to have a major economic impact if released in other developing countries such as
Indonesia and Bangladesh, where PTM is a serious problem. With proper commercial
development, this product will become a tool available to all potato farmers in South
Africa. This product will demonstrate the feasibility of efforts led by the public
sector, and developing country institutions, to make biotechnology products available
in Africa. Furthermore, it will demonstrate the value of developing country
involvement in generating safety assessment data, from the standpoint of scientific
contribution and reduced cost. Finally, the Bt-potato project provides important
capacity building opportunities for public sector institutions with regard to the
comprehensive approach that is required to commercialize transgenic crops. The

expertise and know-how to commercialize transgenic products has been, almost
exclusively, the domain of private companies. We intend to publish the information
we have gained during this project, for the benefit of public-sector institutions.
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Integrated Pest Management for Potato Tubermoth

Book · August 2008

Lawrence A Lacey Jürgen Kroschel

The user has requested enhancement of the downloaded file.

Integrated Pest Management for the

Jürgen Kroschel and

Result of a symposium held in Boise, Idaho on August 21, 2006 as part of

The present publication is the result of a symposium on “Integrated Pest

Prof. Dr. Jürgen Kroschel Dr. Lawrence A. Lacey

Ocurrence of the potato tuber moth, in the Columbia Basin of

Regional and seasonal forecasting of the potato tuber moth

An overview of microbial control of the potato tuber moth…………..33

The potato tuber moth granulovirus (PoGV): use, limitations

Biological control of potato tuber moth, by Apanteles

Spinosad: a new biopesticide for Integrated Pest

Laboratory experiments towards the development of an

Reducing potato tuber moth damage with cultural practices

Integrated Pest Management dealing with potato tuber moth

Integrated Pest Management of potato tuber moth in New

Integrated Pest Management of potato tuber moth in India……….127

Commercialization of potato tuber moth resistant potatoes in

Pest status and forecasting

Occurrence of the potato tuber moth, in the

Materials and Methods

Washington. PTM pheromone-baited traps were set out in Washington in late

Regional and seasonal forecasting of the

Materials and Methods

Table 1. Parameter values of the functions used to describe simulation of P. operculella

Development rate [SHARPE and DEMICHELE (1977) model]

Mortality [second-order polynomial function]

Egg 0.0003 -0.0177 0.3598

Total eggs per female [second-order polynomial

-1.029 42.5037 -269 Temperature

Proportion of progeny production with age [cumulative

Phenological and geographical modeling

Establishment risk index (survival risk)

Generation Index = ∑ 365 / Tx

Results and discussion

Establishment risk (survival) index

Recently, the outbreak of PTM in the Northwest USA (Oregon, Washington)

insect resistance development against insecticides or Genetically Modified

Accounting for within-year variation

net reproduction rate (♀/♀)

assessed in additional experiments. Nevertheless, this modelling is valuable in

BRIESE DT 1980. Characterisation of a laboratory stain of the potato moth,

KROSCHEL J and M SPORLEDER 2006. Ecological approaches to Integrated Pest

WILMOT SENARATNE K, PALMER W and R SUTHERST 2006. Use of CLIMEX modelling

Biocontrol and use of biorationals

An overview of microbial control of the potato

management for chemical pesticides, and minimal impact on beneficial non-target

Entomopathogenic Nematodes (EPNs)

stage of PTM that were exposed as neonate larvae to 15 or 30 g of formulated M.

ANGELES I AND J ALCAZAR 1996. Susceptibilidad de la polilla Symmetrischema

CIP. 1992. Biological control of potato tuber moth using Phthorimaea

HAMILTON JT AND JA MACDONALD 1990. Control of potato moth, Phthorimaea

using granulosis virus: propagation and effectiveness of the virus in field

operculella (Zeller) (Lepidoptera: Gelechiidae). Journal of Biological Control

combinations on the potato tuber moth Phthorimaea operculella (Zeller)

WRAIGHT SP, SPORLEDER M, POPRAWSKI TJ AND LA LACEY 2007. Application and

The potato tuber moth granulovirus (PoGV):

Baculoviruses, comprising two groups of virus, nuclearpolyhedrovirus (NPV) and

Virus field stability (UV-stability)