Potential Invasive Pests of Agricultural Crops

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1 3 CABI INVASIVES Series 3 3 CABI INVASIVES Series Potential Invasive Pests of Agricultural Crops edited by Jorge E. Peña Invasive arthropods cause significant damage in agricultural crops and natural environments across the globe. Potentially threatened regions need to be prepared to prevent new pests from becoming established. Therefore, information on pest identity, host range, geographical distribution and biology, and tools for detection and identification are all essential to researchers and regulatory personnel. Potential Invasive Pests of Agricultural Crops focuses on the species most recently determined to be invasive pests of agricultural crops in temperate subtropical and tropical areas and on potential invaders, discussing their spread, biology and control. This book is an essential resource for research entomologists, international, national and local regulatory agencies, integrated pest management specialists and researchers and students in pest management and crop protection. Related titles Invasive Plant Ecology and Management: Linking Processes to Practice CABI Invasives Series No.2 Edited by T.A.Monaco and R.A.Sheley 2012 c.216 pages ISBN Invasive Alien Plants: An Ecological Appraisal for the Indian Subcontinent Edited by J.R.Bhatt, J.S.Singh, S.P.Singh, R.S. Tripathi and R.K.Kohli CABI Invasives Series No pages ISBN Insect Pests in Tropical Forestry F.R. Wylie and M.R. Speight PB pages ISBN HB pages ISBN Invasive Plant Ecology in Natural and Agricultural Systems Modular Text Series B.D. Booth, S.D. Murphy and C.J. Swanton pages ISBN CABI INVASIVES Series Potential Invasive Pests of Agricultural Crops Peña Potential Invasive Pests of Agricultural Crops edited by Jorge E. Peña For further information on these titles and other publications, see our website at CABI Nosworthy Way Wallingford Oxfordshire OX10 8DE UK CABI 38 Chauncey Street Suite 1002 Boston, MA USA Space for bar code with ISBN included

2 7 Tecia solanivora Povolny (Lepidoptera: Gelechiidae), an invasive pest of potatoes Solanum tuberosum L. in the Northern Andes Daniel Carrillo 1 * and Edison Torrado-Leon 2 1 Department of Entomology and Nematology, Tropical Research and Education Center, University of Florida, Homestead, Florida 33031, USA; 2 Facultad de Agronomia, Universidad Nacional de Colombia, Bogota D.C., Colombia 7.1 Introduction Tecia solanivora Povolny (Lepidoptera: Gelechiidae) is a key pest of potato (Solanum tuberosum L.) tubers that recently invaded some cultivation areas in South America and the Canary Islands, with disastrous effects on the potato industry in these areas. The putative area of origin of T. solanivora is Guatemala, whence its common name (Guatemalan potato moth) is derived, probably ranging from the Isthmus of Tehuantepec in Mexico to Northern Honduras and El Salvador (Puillandre et al., 2008). It was first described in 1973 and reported as a pest of potato in Guatemala, Costa Rica, Panama, Honduras and Salvador (Povolny, 1973; Niño, 2004). In 1983, it was introduced to Venezuela (Tachira State) through importation of infested potato tubers (variety Atzimba) from Costa Rica (Salazar and Torres, 1986); it rapidly became the most damaging insect pest of potato in the Venezuelan Andes. It was detected in Colombia in 1985, in the northeastern state of Norte de Santander (Arias et al., 1996), which borders Tachira. By 1990 it had reached the major cultivation areas in the center of the country, spreading to all potato cultivation areas of Colombia thereafter (Arias et al., 1996). In 1996 it reached Ecuador (Pollet, 2001); and in 1999, the Canary Islands, Spain (EPPO, 2005), where it arrived through illegal importation of infested potato tubers, possibly from South America (Puillandre et al., 2008). In all cases, the introduction of T. solanivora to new geographical areas was attributed to movement of infested tubers, and has resulted in population explosions that have significantly harmed potato production, often devastating potato crops in the invaded areas (Arias et al., 1996; Pollet, 2001; Torres et al., 1997). 7.2 Related Genera and their Distribution In addition to T. solanivora, two other Gelechiid species, Phthorimaea operculella (Zeller) and Symmetrischema tangolias (Gyen), are considered potato pests in various areas of the world and are collectively referred as the potato tuber moth complex. Whereas there are similarities in their life history, the three species differ in their distribution, degree of polyphagy and the parts of the potato plant utilized. In contrast to T. solanivora 126 CAB International Potential Invasive Pests of Agricultural Crops (ed. J. Peña) Pena_Ch07.indd 126 1/11/ :29:42 AM

3 Tecia solanivora Povolny, an Invasive Pest of Potatoes 127 which originated in Central America, both the potato tuberworm and the Andean potato moth (P. operculella and S. tangolias, respectively) likely originated in the mountainous region of South America (Sporleder et al., 2004; Sporleder, 2008; Medina et al., 2010). P. operculella has been a highly successful invasive species currently considered a cosmopolitan potato pest, whereas S. tangolias has a more restricted distribution in Bolivia, Peru, Ecuador, Colombia and Australia, where it was accidentally introduced (Osmelak, 1987; Griepink, 1995; Pollet et al., 2003b; Sporleder, 2008). Co-occurrence of the three potato tuber moth species has been observed only in southern Colombia and Ecuador (Dangles et al., 2008). Although little is known about the interaction of the three species, Dangles et al. (2008) studied their physiological responses to constant temperatures and found that S. tangolias was more cold tolerant, while T. solanivora had the highest growth rates at warmer temperatures. Surprisingly, the tuber moth species with the widest distribution, P. operculella, showed the poorest performance across the range of temperatures. The most important difference among the species of the potato moth complex can be found in their feeding habits. T. solanivora is specialist on potato tubers, whereas P. operculella and S. tangolias also feed on potato leaves and stems (Rondon et al., 2007; Sporleder, 2008). Moreover, P. operculella has at least 60 reported alternate hosts (Das and Raman, 1994) and S. tangolias is known to feed on other solanaceous plants (Osmelak, 1987; Sporleder, 2008). 7.3 Biology and Life History T. solanivora has adapted to various mountainous ecosystems distributed throughout a wide altitudinal range. In the Northern Andes, T. solanivora occurs at medium and high altitudes between 1600 and 3400 m above sea level (ASL) (Torres et al., 1997). Areas affected by T. solanivora in Costa Rica are located in medium altitudes between 1300 and 2300 m ASL (Povolny, 1973); in the Canary Islands, the highest damage occurs at low altitudes between 500 and 600 m ASL (EPPO, 2005). Studies conducted throughout an altitudinal range suggest a negative relationship between altitude and population densities of the pest (Barreto et al., 2003; Gallegos, 2002). However, management practices and climatic variables are regarded as the key factors regulating population dynamics of T. solanivora in different areas of invasion (Notz, 1996; Nústez et al., 1999; Barreto et al., 2003; Torres, 1998; López-Ávila and Espitia, 2000). 7.4 Life Cycle The life cycle duration of T. solanivora can vary greatly, depending on the temperature. Notz (1996) studied the development and reproduction of T. solanivora at a range of constant temperatures and concluded that the life cycle can last from approximately 42 days at 25 C to almost 200 days at 10 C. Thus, the number of generations per annum can vary from ten at 25 C to two at 10 C. Eggs are laid individually or in small groups on the soil surface near the plant stem, or on uncovered tubers; seldom on the leaves or stems (Barreto, 2005). In stored potatoes, eggs are laid on or near the tuber eyes and on the sisal storage bags. Eggs are semi-spherical in shape and measure approximately 0.5 mm in diameter, which makes them almost imperceptible to the naked eye. Their coloration is white but they turn progressively yellow as they mature (Fig ). This stage can last from 5 to 25 days, depending on the environmental temperature (Notz, 1996). Upon emergence, neonate larvae actively search for potato tubers, either burrowing into the soil or locating exposed tubers in storage conditions. Larvae enter tubers through the potato eyes or skin leaving imperceptible entrance holes, and start feeding and making galleries inside the tuber. The larval stage is composed of 4 instars that usually develop inside a single tuber; feeding starts in the outer layers (cortex and vascular ring), and advances into the inner structures (medulla) (Hilje, 1994). The first instar larvae are approximately 1.5 mm long and hyaline in appearance (Fig ); as they grow, larvae acquire a bluish-green coloration and reach approximately 16 mm by the last instar (Torres, 1998). The larval stage can last from approximately 18 to 80 days depending on the temperature (Notz, 1996). Mature larvae exhibit a pink coloration in the dorsum but retain the blue-green coloration ventrally (Fig ). At this point larvae cease Pena_Ch07.indd 127 1/11/ :29:43 AM

4 128 D. Carrillo and E. Torrado-Leon Egg 1 Adult 2 N eonate ar l va Pupa 3 Lar va Fig Life cycle of Tecia solanivora: 1. Egg; 2. Neonate larva; 3. Larva; 4. Pupa; 5. Adult. feeding, leave the tubers, and seek protected places to construct a silken cocoon covered with soil or dirt. Pupation occurs in the soil or dead plant tissue, and seldom inside the tubers; in potato storage, pupae can be found on tubers, packing sacks, walls, floors and crevices in the storage facility (López-Ávila and Espitia, 2000). During the pupal stage, differentiation between sexes starts to be conspicuous, as female pupae tend to be larger and heavier than male pupae. Moreover, females have the genital opening in the 8th abdominal segment and males in the 9th, which could be used as a diagnostic character to enable sex differentiation (Rincón and LópezÁvila, 2004). The pupa has a brown-reddish coloration that becomes darker and shiny with time (Fig ). This stage can last from 10 to 90 days depending on the environmental temperature (Notz, 1996). Adults are dull-colored moths with a narrow body and brownish-gray wings with small brown or black longitudinal markings (Figs and 7.2). Females are larger and have a broader abdomen Pena_Ch07.indd 128 Fig Tecia solanivora mating. than males. Female moths can be differentiated by the presence of three dark stigmas in each wing, while males have only two (Povolny, 1973; LópezÁvila and Espitia, 2000). Adults are active at dusk and remain hidden in dark and protected areas during daylight. T. solanivora is a sexually reproducing species in which females require a single mate to remain fertile for the majority of their adult life (Torres et al., 1997; Rincón and Garcia, 1/11/ :29:43 AM

5 Tecia solanivora Povolny, an Invasive Pest of Potatoes 2007) (Fig. 7.2). Adult longevity can vary from 10 to 25 days depending on the temperature. According to Notz (1996) the higher oviposition rates and longer fecundity periods occur at 15 C. 7.5 Sampling, Monitoring Techniques and Action Thresholds The most widely used method for monitoring T. solanivora populations is trapping adult males using a synthetic sexual pheromone (Nesbitt et al., 1995; Bosa et al., 2005) as bait inside water or delta traps. This method has been used in population dynamics studies, for the establishment of action thresholds and in surveillance programs in places with risk of invasion. Various studies using pheromone trapping identified key factors that regulate the population dynamics of T. solanivora. Factors such as the presence of residues from previous crops or other sources of infestation, (i.e., storage facilities, abandoned crops), together with pest-management practices, may influence the dynamics of this pest at a local level (Niño, 2004). However, a marked influence of abiotic factors on the dynamics of T. solanivora has been observed in the different areas. Most studies suggest that precipitation is a key factor regulating population growth of T. solanivora. High pest-population densities population explosions were often associated with dry periods, whereas populations were usually low in periods of high precipitation (Palacios, 2002; Barragan et al., 2002; Alvarado et al., 1993; Torres, 1998; Rodríguez and Lépiz, 1989; Rodríguez et al., 1988). Moreover, these Fig Damage to potato tubers caused by Tecia solanivora. Pena_Ch07.indd studies suggest that population dynamics of T. solanivora are related to the phenological stage of the crop. In general, adult activity (captures) is relatively low during the initial vegetative stages of the crop and increases when potato plants start the tuberization process. In other words, adult activity and oviposition increase when the larval food substrate (tubers) becomes available, and remain high until tubers reach physiological maturity (Torres, 1998; Galindo and Española, 2004). This information has been key to design management tactics in the potato production systems affected by this pest. For instance, Pollet et al. (2003a) proposed a model to predict tuber damage in an early crop stage, based on pheromone trapping and climatic data measured during the first months of the plant cycle. Moreover, pheromone trapping has been used to establish nominal action thresholds that vary according to climatic variables and crop management. Based on experience, researchers have established thresholds from 50 to 100 adult males per week per trap (when using 4 traps per hectare) as an indicator for a pesticide application. In some cases, however, adult trapping does not correlate with tuber damage, finding low damage and high adult captures (Gallegos et al., 2002) and vice versa. In addition, pheromone trapping has been used as a key element of a preventive surveillance network established in Peru due to the high risk of introduction to this area (Oyola, 2002) and also to detect presence of the pest in storage facilities and other places. 7.6 Damage The larval stage of T. solanivora is the only stage that causes direct damage to potato plants. Larvae feed and develop inside potato tubers, excavating galleries and leaving frass inside them, which allows the introduction of fungi and bacteria (Fig. 7.3). Thus, larval feeding renders potato tubers unmarketable and useless for animal feeding purposes. Furthermore, additional resources are required to dispose of infested tubers, as they become a source of infestation for future crops or stored potatoes. Under favorable climatic conditions and poor management practices T. solanivora can cause complete yield loss in field and/or storage conditions (Vargas et al., 2004; Arias, 1996; EPPO, 2005). 1/11/ :29:44 AM

6 130 D. Carrillo and E. Torrado-Leon 7.7 Control Tactics Management of T. solanivora requires the combination of multiple control methods both in field and storage conditions. In places where this pest is present, or threatening to invade, several research centers and institutions have proposed management alternatives including chemical, biological, autocidal, cultural and regulatory control tactics Chemical control The effectiveness of chemical control practices in field conditions depends on the target and the timing of the applications. Most of the immature stages of T. solanivora develop inside the tuber and are out of reach of conventional (non-systemic) pesticide applications. Thus, to prevent the larvae from entering the tubers, applications should be directed to the base of the plant, targeting eggs and adults, the only stages exposed. Moreover, applications should be made only during the tuber formation phase. For instance, early pesticide applications during the vegetative stages of the crop are unnecessary, as plants do not have tubers and therefore larval development is not possible. When tubers are formed and the larvae are already inside, applications do not reduce the damage. However, despite the recommendations made by research centers and extension agents, chemical control is often conducted on calendar-based applications (Niño, 2004). In Colombia farmers rely on intensive pesticide use. This is unfortunate, since experiments conducted by Gallegos et al. (2002) suggested that cultural control measures are more efficient in controlling T. solanivora infestations than the application of several chemical pesticides. Commonly used pesticides against T. solanivora are neurotoxins, including: organophosphates (Clorpirifos, Profenofos, Triclorphon) (Torres, 1998) pyrethroids (Permethrin) (Arevalo and Castro, 2003) and carbamates (López-Ávila and Barreto, 2004). Formulations for field application use water as a carrier, and dust is used as a carrier on pesticides to protect potato tubers that will be used as seed. In addition to chemically synthesized pesticides, plant-derived essential oils have also been shown to control T. solanivora in stored potatoes (Salazar and Betancourth, 2009; Ramirez et al., 2010) Autocidal Studies have demonstrated that the reproduction of T. solanivora can be altered by disrupting the pheromone-mediated communication between sexes (Bosa et al., 2005). The sexual pheromone produced by females to attract males, which can be synthesized and used for monitoring purposes, is a mixture of three chemical compounds in a specific ratio (Nesbitt et al., 1985; Bosa et al., 2005). When the ratio of the three components is modified, attraction of males by calling females is suppressed. Since T. solanivora only reproduces sexually, mating disruption could potentially reduce its populations. According to Bosa et al. (2006, 2008) this technique has potential for reducing populations of the pest both in storage and field conditions Biological control The most developed and efficient biological control tactic against T. solanivora is the use of viral pathogens (Table 7.1). Initially, a granulovirus isolated from P. operculella, and formulated as a biopesticide by the International Potato Center (Lima, Peru), was considered promising to control T. solanivora under storage conditions (Sporleder, 2004). More recently, some Colombian granulovirus isolates found infecting populations of T. solanivora were selected for developing a biopesticide, as they were more pathogenic than Peruvian isolates (Espinel-Correal et al., 2010; Gomez et al., 2009). Parasitoids of P. operculella were also evaluated for potential as biological control agents of T. solanivora (Navas et al., 1986; Torres, 1998). The egg parasitoid Copidosoma koehleri (Hymenoptera: Encyrtidaeidae) was introduced to Venezuela in 1994 from Peru and Bolivia but showed poor development and low parasitism rates on T. solanivora (Torres, 1998) (Table 7.1). A Colombian native parasitoid, Trichogramma lopezandinensis (Hymenoptera: Trichogrammatidae), was tested for control of T. solanivora and showed potential to be used under storage and field conditions (Rincón and López-Ávila, 1999; Rubio et al., 2004). In addition, the native entomopathogenic nematode Steinernema feltiae has shown some potential because of its ability to actively search Pena_Ch07.indd 130 1/11/ :29:44 AM

7 Tecia solanivora Povolny, an Invasive Pest of Potatoes 131 Table 7.1. Natural enemies reported in association with T. solanivora in the different countries where this pest is present. Type of natural enemy Scientific name Location Reference Egg parasitoids Chelonus phthorimaea Gahan Guatemala Navas et al., 1986 Copidosoma koehleri Blanchard Venezuela Torres, 1998 Trichogramma lopezandinensis Sarmiento Colombia Rincón and López- Ávila, 1999 Trichogramma aff. pretiosum Colombia Osorio et al., 2001 Larval parasitoids Apanteles sp. Colombia Osorio et al., 2001 Predators (eggs and Buchananiella contigua Colombia Osorio et al., 2001 larvae) (Buchanan-White) Predators (eggs) Lyctocoris campestris (Fabricius) Colombia Osorio et al., 2001 Parasites (nematodes) Steinernema feltiae Filipjev Colombia, Saenz, 2003 Venezuela Torres, 1998 Pathogens Granuloviruses Ecuador, Colombia, Venezuela Zeddam et al., 2003 Espinel-Correal et al., 2010 Niño and Notz, 2000 for T. solanivora larvae inside potato tubers; however, methodologies for field applications still need to be developed (Saenz, 2003). Finally, several entomopathogenic fungi have been tested against T. solanivora, showing low control (Feris et al., 2002). Under natural conditions a number of natural enemy species have been reported in association with T. solanivora (Table 7.1). However, little is known about their influence in the population dynamics of this pest. For instance, Pollet et al. (2003a) suggested that natural enemies were almost non-existent in Ecuador. In contrast, studies conducted by Osorio et al. (2001) reported various parasitoids, predators and entomopathogens associated with T. solanivora in different areas of Colombia. The higher number of natural enemies reported in Colombia could be related to the longer time of establishment of the pest in this country. However, very few natural enemies have been reported in Venezuela where T. solanivora has been established for a longer period of time (Torres, 1998), and only one natural enemy has been reported from its native range in Guatemala (Navas et al., 1986). The higher number of natural enemies found in Colombia could be explained by the sampling methodology that included placement of sentinel eggs in the field and collection of potato tubers from different locations, which resulted in significantly more findings than when only direct observation was used (Osorio et al., 2001). However, the potential of most of the reported natural enemies of T. solanivora is still unknown, and requires further investigation Host plant resistance Controlled infestation experiments have shown that commercial potato varieties in Venezuela (Niño, 2004) and Colombia (Bejarano et al., 1999; Leiva and Echeverria, 1999) are susceptible to the attack of T. solanivora. An alternative that has been tested experimentally is the production of transgenic potato plants that express Bacillus thuringiensis (Bt) crystalline insecticidal proteins, achieving efficient control of T. solanivora (Valderrama et al., 2007; Rivera and López-Ávila, 2010) Cultural control Cultural practices recommended for T. solanivora are often the same as those recommended for managing P. operculella. They are designed to destroy sources of infestation of the pest, create unfavorable conditions for pest development and interrupt the pest s life cycle. Pena_Ch07.indd 131 1/11/ :29:44 AM

8 132 D. Carrillo and E. Torrado-Leon Cultural practices proposed to reduce field infestations control T. solanivora include: 1. Good soil preparation to physically destroy immature stages present in the soil. Adequate soil preparation should also avoid formation of soil clumps that serve as refuges and oviposition sites for adult moths. 2. Using certified, pest-free seeds (tubers) previously treated with formulated granulovirus or chemical insecticides formulated as dusts. 3. Increasing planting depth by 15 cm to reduce the possibility of larvae reaching the tubers. 4. Increasing hilling height and frequency to protect tubers from ovipositing females and to reduce the possibility of larvae reaching the tubers. 5. Avoiding leftover potatoes that could serve as sources of infestation for the next planting season. Cultural practices recommended to prevent infestations in potato storage include: 1. Storing tubers immediately after harvesting to avoid unnecessary exposure to ovipositing adult female moths in the field. 2. Washing tubers after harvesting; oviposition is significantly reduced on washed tubers (Vargas et al., 2004). 3. Disinfecting storage facility. 4. Avoiding reuse of packing material. 5. Allowing indirect light inside the storage facility, as adults are more active under dark conditions. 6. If possible, keeping store facility at temperatures below 8 C; T. solanivora cannot develop or reproduce below this temperature (Notz, 1996). 7.8 Regulatory Quarantine Methods Studies suggest that T. solanivora could become established in various potato cultivation areas of South America and other parts of the world (Shaub et al., 2009). The introduction of this pest to Peru, the site of origin and where the highest diversity of potato varieties is found (Spooner et al., 2005), could have disastrous consequences. Restrictions to potato tuber movement and the implementation of surveillance networks in critical areas, together with extension programs, have helped to prevent the introduction of T. solanivora to this country (Naccha and Villar, 2005). The spread of T. solanivora to other parts of the word will depend largely on the preventive measures that the different potato-growing regions can implement to reduce the chances of an introduction. The European and Mediterranean Plant Protection Organization included T. solanivora in the list of species considered a threat to European agriculture, and recommended its regulation as a quarantine pest (EPPO, 2005). No major actions have been taken in other parts of the world. 7.9 Conclusion The negative consequences experienced in areas affected by T. solanivora indicate the importance not only of preventing its introduction to new areas, but also of being prepared to respond to an eventual invasion. Scientists from various research centers and institutions have studied the biology and life history of this pest, and tested an array of management tactics for it. As a result, several alternatives to reduce the damage it causes are now available. However, the large-scale adoption of these management tactics in an integrated manner has not been achieved in the different areas. Unfortunately, due to the risk of high losses, lack of collaborative management, insufficient extension programs and the involvement of other important pest problems, potato growers tend to rely on the application of chemical pesticides as their only management option. Efforts are needed to increase the adoption of other tactics and to establish integrated pest management programs at regional levels References Alvarado, J., Ortega, C. and Acevedo, J. (1993) Evaluación de la densidad de trampas de feromona en la captura de la polilla Centroamericana de la papa. Revista Latinoamericana de la Papa 1, Arévalo, A. and Castro, R. (2003) Evaluación postregistro de los insecticidas con licencia de uso para controlar la polilla guatemalteca de la papa Tecia solanivora (Povolny 1973) (Lepidoptera: Gelechiidae) en Colombia. In: Memorias II Taller Nacional, Tecia Solanivora. Centro Virtual de la Cadena Alimentaria de la Papa, Bogotá, Colombia, pp Pena_Ch07.indd 132 1/11/ :29:44 AM

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