SciELO - Scientific Electronic Library Online

vol.42 número3Mapas de beneficio y de rentabilidad en la agricultura de precisiónResistencia de portainjertos de vid utilizados en Chile a los nematodos de la raíz Meloidogyne ethiopica, M. hapla y M. javanica índice de autoresíndice de materiabúsqueda de artículos
Home Pagelista alfabética de revistas  

Servicios Personalizados




Links relacionados

  • En proceso de indezaciónCitado por Google
  • No hay articulos similaresSimilares en SciELO
  • En proceso de indezaciónSimilares en Google


Ciencia e investigación agraria

versión On-line ISSN 0718-1620

Cienc. Inv. Agr. vol.42 no.3 Santiago dic. 2015 



Body mass and wing geometric morphology of the codling moth (Lepidoptera: Tortricidae) according to sex, location and host plant in the region of Maule, Chile

Tamaño corporal y morfometría geométrica de la polilla de la manzana (Lepidoptera: Tortricidae) según sexo, localidad y planta hospedera en la Región del Maule, Chile


Fabiola Torres1, Marcela A. Rodríguez1,2, Blas Lavandero3, and Eduardo Fuentes-Contreras1

1 Millennium Nucleus Center in Molecular Ecology and Evolutionary Applications in the Agroecosystems (CEM), Facultad de Ciencias Agrarias, Universidad de Talca. Casilla 747, Talca, Chile. 
2 Departamento de Zoología, Facultad de Ciencias Naturales y Oceanógraficas, Universidad de Concepción,
Casilla 160C, Concepción, Chile.
3 Millennium Nucleus Center in Molecular Ecology and Evolutionary Applications in the Agroecosystems (CEM), Instituto de Ciencias Biológicas, Universidad de Talca. Casilla 747, Talca, Chile.
Corresponding author:


F. Torres, M.A. Rodríguez, B. Lavandero, and E. Fuentes-Contreras. 2015. Body mass and wing geometric morphology of the codling moth (Lepidoptera: Tortricidae) according to sex, location and host plant in the region of Maule, Chile. Cien. Inv. Agr. 42(3): 397-406. Codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), is the key pest of apple and walnut production worldwide. Among other variables, successful management of this pest is dependent on adult dispersal at the local scale. Body mass and wing geometric morphology were evaluated on female and male codling moth adults collected from apple and walnut trees in three localities of the Region of Maule. Codling moth adults were obtained by rearing diapausing larvae collected from the field. Newly emerged adults were killed, and the total dry weight, thorax weight and abdomen weight were calculated. The wings were cleared and mounted for morphometric analyses: 15 and 11 characteristics of the fore and hind wings, respectively, were measured, including the meeting points of the veins and attaching points to the margins. In addition, wing length, area, loading, and aspect ratio were evaluated. As previously reported, females showed larger thorax, abdomen and total body mass, with longer, larger, and wider wings than males but with similar wing loading as males. The geometric morphology analysis also confirmed a sexual dimorphism in the fore and hind wings’ size and shape. Significant differences in body mass and wing loading were found between localities, with morphological differences detected in the fore and hind wing shape but not in size. Host plant (apple and walnut) did not influence body mass, wing size or shape variables. Potential consequences of body mass and wing morphology differences on the dispersal and management of this major pest are discussed.

Key words: Apple, aspect ratio, Cydia pomonella, landmark-analysis, MorphoJ, walnut, wing loading


F. Torres, M.A. Rodríguez, B. Lavandero y E. Fuentes-Contreras. 2015. Tamaño corporal y morfometría geométrica de la polilla de la manzana (Lepidoptera: Tortricidae) según sexo, localidad y planta hospedera en la Región del Maule, Chile. Cien. Inv. Agr. 42(3): 397-406. La polilla de la manzana, Cydia pomonella (L.) (Lepidoptera: Tortricidae), es la plaga clave de manzanos y nogales alrededor del mundo. Entre otras variables, el manejo exitoso de esta plaga depende de la dispersión de los adultos en la escala local. Se evaluó el peso corporal y la morfología geométrica de las alas de hembras y machos adultos de la polilla de la manzana, Cydia pomonella (L.) (Lepidoptera: Tortricidae), provenientes de manzano y nogal en tres localidades de la Región del Maule, Chile. A partir de larvas en diapausa recolectadas en el campo se obtuvieron adultos de la polilla de la manzana en laboratorio. Los adultos recién emergidos fueron muertos y se calculó el peso seco total, peso del tórax y peso del abdomen. Las alas fueron aclaradas y montadas para su análisis morfométrico, el cual incluyó 15 hitos para las alas delanteras y once para las alas posteriores, respectivamente, incluyendo puntos de unión de las venas y puntos de unión vena/margen. Además, se evaluó el largo, área, carga alar y razón de aspecto de las alas. Como se ha reportado previamente, encontramos que las hembras mostraron mayor peso de tórax, abdomen y cuerpo completo, con alas más largas, grandes y anchas que los machos. Sin embargo, tales diferencias no resultaron en diferencias de la carga alar entre sexos. El análisis de morfometría geométrica también confirmó un dimorfismo sexual en el tamaño y forma de las alas anteriores y posteriores. Se encontraron diferencias significativas en el peso corporal y la carga alar entre localidades, con diferencias en la morfología geométrica de la forma, pero no del tamaño en las alas anteriores y posteriores. La planta hospedera (manzano y nogal) no influenció ninguna de las variables de peso corporal, tamaño o forma de las alas analizadas en los adultos de la polilla de la manzana. Se discuten las posibles consecuencias de las diferencias en peso corporal y morfología alar según sexo y localidad sobre la dispersión y manejo de esta plaga primaria.

Palabras clave: Carga alar, Cydia pomonella, hito, manzano, MorphoJ, nogal, razón de aspecto.



Codling moth, Cydia pomonella L. (Lepidoptera: Tortricidae), is the major pest of pome fruits (apple, pear, and quince) and walnut worldwide (Barnes, 1991). The control of this pest has been based on regular sprays of synthetic insecticides; in some areas, mating disruption and granulovirus applications are also key components of management programs (Witzgall et al., 2008; Weddle et al., 2009). The spatial population dynamics of the codling moth can affect these control strategies in several ways (Mazzi and Dorn, 2012). Landscape, orchard and tree canopy structure influence codling moth dispersal at the local scale and thus its spatial population dynamics (Stoeckli et al, 2008; Ricci et al, 2009, 2011). The selection of insecticide-resistant individuals in orchards under regular insecticide sprays could be reduced by immigration and gene flow of susceptible individuals from unmanaged areas (Basoalto et al., 2010, Fuentes-Contreras et al, 2014). Furthermore, a decrease in mating disruption efficacy is well-documented as a consequence of the immigration of mated females from unmanaged areas, especially in orchard borders, as well as from high density populations or small orchard sizes with high perimeter-area relationship (Witzgall et al, 2008).

Some studies in codling moth have found an association between insecticide resistance, body size, and sexual dimorphism because females have higher levels of resistance compared with males, which usually are of smaller body size (Varela et al., 1993; Sauphanor et al., 2000; Reyes et al, 2004, 2015; Fuentes-Contreras et al., 2007). Body size has also been related to dispersal capability in the codling moth: in the same population, larger females tend to be more fecund, live longer, and disperse less than smaller females (Gu et al, 2006). Flight distance was also negatively correlated with body weight and wing size in the codling moth, suggesting a trade-off between fitness and dispersal for this species (Schumacher et al, 1997a). Sex dimorphism in the codling moth is apparent in the body size, wing size, and wing shape, as detected using wing geometric morphology analyses (Khaghaninia et al., 2011a), and factors such as altitude and geographic origin affect wing morphometrics (Khaghaninia et al, 2011b).

Few other studies suggest the existence of different apple and walnut races of the codling moth (Cisneros and Barnes, 1974; Phillips and Barnes, 1975), although genetic studies have found little genetic differentiation between apple and walnut populations (Buès et al. 1995; Thaler et al., 2008; Franck and Timm, 2010). To our knowledge, no studies on body size or wing morphology have been performed with codling moth from different host plants, although such differences could be important for the insecticide resistance and dispersal capabilities of this pest species.

Our aim was to evaluate whether body size and several wing morphometric variables of codling moth populations differ between sex, locality and host plant in the Region of Maule, Chile. This information could shed light on the patterns of insecticide resistance and codling moth dispersal in central Chile, which could contribute to a more successful management of this pest under rotation of insecticides with different modes of action between codling moth generations.

Materials and methods

Insect sampling and rearing

Cardboard traps were strapped around trunks in unsprayed backyard apple and walnut trees at the end of the summer in the Region of Maule (February 2013), Chile. Apple and walnut trees, usually growing together, were sampled in each of these three localities: Las Cruces (35°45’43,11"S, 71°62’10,66"W), Orilla de Maule (35°63’ 39,70’’S, 71°41’89,07"W), and Yerbas Buenas (35°74’64,07"S, 71°58’57,01’’W). One apple and one walnut tree were sampled in Orilla de Maule and Yerbas Buenas, whereas six apple and two walnut trees were sampled in Las Cruces. The cardboard traps were removed in autumn (April 2013) and brought back to the laboratory to obtain fifth instar larvae in diapause. A minimum of fifty larvae were sexed and maintained for three months at 10 ± 1 °C and a 12:12 (L:D) photoperiod to complete diapause. Larvae were then transferred to 24 ± 1 °C and a 16:8 (L:D) photoperiod to promote diapause break and to obtain adults. Less than 1-day old adults were killed to prevent wing damage caused by flight inside small containers and were preserved in 95% ethanol for further analyses. A sample size of ten individuals (N = 10) for each sex, location and host plant combination was used for body mass and wing morphometric measurements.

Body mass evaluation and wing mounting

Adult insects were removed from the alcohol and dried at 45 °C for 48 h to evaluate their total dry mass (m), expressed in mg, using a microbalance (Mettler Toledo XS, Greifensee, Switzerland). After weighing, the fore and hind wings, head, and legs were carefully removed to weigh the thorax (t) and abdomen (a) separately, as described by Norberg and Leimar (2002). The right pair of wings was then transferred to 70% ethanol to induce a partial rehydration for two hours. The wings were placed in a sodium hypochlorite solution (5%) for five minutes, and the scales were removed with a fine brush. The wings were then cleared with a potassium hydroxide solution (5%), and then glacial acetic acid solution (5%) was added to stop the alkaline reaction. The wings were gently washed with eugenol and left in 95% alcohol overnight. Finally, the wings were mounted on a slide/coverslip with Neo-Mount media (Merck Millipore, Darmstadt, Germany).

Wing morphology and geometric morphometric measurements

Geometric morphometrics of the wings were calculated using landmark analysis (Bookstein, 1986) to address the possible influence of sex, location and host plant on wing size and shape. Fifteen and eleven landmarks were defined for the fore and hind wings, respectively, which correspond to the meeting points of the veins and attaching points of the veins and wing margins (Type I landmarks), as described for this species by Khaghaninia et al. (2011a,b). Images with 10X magnification were taken with the software Optika Vision Pro using a stereoscopic microscope with a digital camera (SMZ 2, Optikam Pro5, Optika, Pontenarica, Italy). Two-dimensional coordinates of the landmarks were obtained with TpsDIG 2 software (Rohlf, 2006). Coordinate data were transformed to shape variables and partial warp scores with MorphoJ 1.03a software (Klingenberg, 2011).

Dry body mass in relation to wing size and area was used to estimate the wing loading and aspect ratio, as defined by Betts and Wootton (1988). Wing morphology measurements were performed as follows: i) wing length (R) was the maximum length (mm) from forewing base to apex, ii) wing area (S) was the total area (mm2) of both fore and hind wings, iii) wing loading (Pw) was the ratio between total dry mass and wing area mS-1 (mg mm-2), and iv) aspect ratio (AR) was the ratio between four times the square of the wing length and wing area (4R2S-1).

Statistical analyses

Significant differences between body dry mass (t, a, and m) and wing morphology variables (S, R, Pw and AR) were performed with a generalized linear model (GLM) using the package glm in R (R Development Core Team, 2014). A gamma error distribution with a logarithmic link function was used. Sex, host plant, location and their interactions were considered as fixed effects. Model selection was performed according to Crawley (2013), using the package car in R. In all cases, we selected the additive model because the interaction model was not significantly better and showed the lowest value of the Akaike information criterion (AIC). Contrasts between pairs of treatments were calculated using the multcomp package in R following Bretz et al. (2011).

Procrustes superimposition removed the variation not related to shape, and the residuals were analyzed with ANOVA using MorphoJ 1.03a software (Klingenberg, 2011) to evaluate the decomposition of the geometric morphological variation between individuais and independent variables (sex, host plant and location). Furthermore, canonicai variation and discriminant function analyses were performed with the same software.


Body mass and wing morphology

Thorax (t), abdomen (a) and total dry body mass (m) showed significant differences between sex (t: x2 = 40.4, df = 1, P0.001; a: x2 = 39.9, df = 1, P<0.001; m: x2 = 50.0, df = 1, P0.001) and locality (t: x2 = 6.6, df = 2, P0.05; a: x2 = 30.4, df = 2, P0.001; m: x2 = 21.7, df = 2, P0.001), but not between host plants (t: x2 = 0.2, df = 1, P = 0.67; a: x2 = 0.3, df = 1, P = 0.56; m: x2 = 0.5, df = 1, P = 0.47) (Table 1). Females had higher t, a and m than males (Table 1). Regardless of host plant and sex, adults from Orilla de Maule has significantly heavier t, a and m than those obtained from Las Cruces and Yerbas Buenas (Table 1).


Table 1. Mean ± SE dry mass (mg) of adult codling moth thorax (t), abdomen (a) and full
body (m) from males and females obtained from different host plants and localities in the
Maule Region, Chile.

1Sample size N = 10.
2Small letters indicate significant differences between localities in each column.
Capital letters indicate significant differences between sex columns.


Regardless of locality or host plant, females had significantly larger R and S than males (R: x2 = 48.4, df = 1, P<0.001; S: x2 = 59.8, df = 1, P0.001) (Table 2). Significant differences in Pwere found between localities (x2 = 41.6, df = 2, P0.001), but not between sex (x2 = 3.2, df = 1, P = 0.07) or host plant (x2 = 0.04, df = 1, P = 0.84). Pw was significantly higher in wings from Orilla de Maule than in wings from Las Cruces and Yerbas Buenas (Table 2). Finally, AR was significantly higher in males than in females (x2 = 14.9, df = 1, P<0.001), without significant differences by locality (x2 = 0.7, df = 2, P = 0.72) or host plant (x2 = 0.05, df = 1, P = 0.84).


Table 2. Mean db SE wing length (R) (mm), area (S) (mm2), loading (P ) (mg mm-2),
and aspect ratio (AR) from males and females obtained from different host
plants and localities in the Maulé Región, Chile.

1Sample size N = 10.
2Small letters indicate significant differences between localities in each column.
Capital letters indicate significant differences between sex columns.


Wing geometric morphology

Procrustes ANOVA showed significant effects of sex on wing size (fore wings MS = 1273003, F(1,118) = 54.7, P0.001; hind wings MS = 983139, F(1,118) = 58.8, P0.001) but not for host plant or location. Similarly, significant effects on wing shape were found for sex (fore wings MS = 0.00034, F(26,3028) = 3.5, P0.001; hind wings MS = 0.00644, F(18,2124) = 23.1, P0.001) and location (fore wings Ms = 0.00024, F(52,3042) = 2.5, P0.001; hind wings MS = 0.00081, F(36,2102) = 2.5, P0.001) but not for host plant. Canonical variation analysis showed that the first and second canonical variates accounted for 29.3% and 18.8% of the variance in the fore wings, respectively, and 50.0% and 18.5% of the variance in the hind wings, respectively. Discriminant function analysis showed significant differences between sex (fore wings Mahalanobis distance = 2.21, t2 = 146.2, P0.001; hind wings Mahalanobis distance = 2.62, t2 = 205.5, P0.001) and between some locations, as follows: Las Cruces and Orilla de Maule (fore wings Mahalanobis distance = 1.80, t2 = 64.5, P = 0.05; hind wings Mahalanobis distance = 0.99, t2 = 19.6, P = 0.63), Las Cruces and Yerbas Buenas (fore wings Mahalanobis distance = 1.51, t2 = 45.4, P = 0.29; hind wings Mahalanobis distance = 1.74, t2 = 60.7, P0.01), Orilla de Maule and Yerbas Buenas (fore wings Mahalanobis distance = 1.36, t2 = 37.1, P = 0.52; hind wings Mahalanobis distance = 1.69, t2 = 57.2, P0.01). Finally, discriminant function analysis showed non-significant differences between host plants (fore wings Mahalanobis distance = 1.33, t= 52.7, P = 0.06; hind wings Mahalanobis distance = 0.94, t2 = 26.4, P = 0.24).


Our results support previous reports of sexual dimorphism in body mass and wing morphology in the codling moth (Schumacher et al, 1997a; Gu et al, 2006), with females showing larger thorax, abdomen and body mass than males. Larger body size is related to higher detoxification capacity, which confers resistance towards insecticides (Reyes et al, 2004, 2015; Fuentes-Contreras et al, 2007). Therefore, females with larger body size have higher levels of insecticide resistance than do males with a smaller body size (Varela et al, 1993; Sauphanor et al, 2000; Reyes et al, 2004, 2015; Fuentes-Contreras et al, 2007).

In our study, codling moth females have longer (higher R), larger (higher S), and wider (lower AR) wings than males. However, heavier females with larger, longer and wider wings have a similar wing loading (Pw) as males. The wing shape of females is associated with slow and agile flight in restricted space and dense vegetation, whereas the smaller and more slender wings of males is related to extensive flight in open spaces (Betts and Wootton, 1998). The similar wing loading between sexes is indicative of similar flight speeds (Betts and Wootton, 1998). Wing geometric morphology analyses consistently detected larger wings for females, with females exhibiting broader wings than males, as previously reported by Khaghaninia et al. (2011a,b).

Differences in body mass and wing morphology between sexes are indicative of different dispersal capabilities. Regardless of sex, adult flight in wind-tunnel experiments has shown a negative correlation with body weight (Schumacher et al, 1997a). However, no major differences in flight capacity between males and females were found in another study (Schumacher et al, 1997b). Flight activity was related with the circadian rhythm, age and mating status rather than sex (Schumacher et al, 1997b; Keil et al, 2001; Gu et al, 2006). Additionally, flight activity was higher in mated than virgin females and increased during the egg-laying period; therefore, little dispersal occurs before mating (Schumacher et al, 1997b). Both sexes showed higher activity at dusk, with males reaching flight activity peaks at a younger age than females (Keil et al, 2001). Treatment with the juvenile hormone mimic insecticide fenoxycarb stimulated locomotor activity and provoked a marked activity peak at dawn in both virgin and mated females (Keil et al, 2001). Therefore, morphological traits are important to explain dispersal, and the physiological and life history attributes are of major relevance (Roff and Fairbairn, 2007).

Regardless of sex and host plant, adults from Orilla de Maule were heavier and had higher Pthan adults from Las Cruces and Yerbas Buenas. However, no differences in R, S or AR were found between locations. The geometric morphology analyses revealed significant differences in the shape of the fore and hind wings between locations, which were accounted for by marginal differences in the fore wings between Orilla de Maule and Las Cruces and larger differences in the hind wings from Yerbas Buenas in comparison with Las Cruces and Orilla de Maule. Previous studies on wing geometric morphology of codling moth populations from different localities in North-West Iran (Khaghaninia et al, 2011a,b) found that abiotic conditions, such as wind velocity at different altitudes, were associated with significant changes in wing size and shape. Slender and longer wings were found in codling moth adults from higher altitudes (Khaghaninia et al, 2011a,b). All locations included in our study were in the central valley of the Maule River, less than 50 km apart, at altitudes between 100 and 200 masl. At the small geographic scale of our study, it is less likely that abiotic factors explain the differences in body mass or wing shape. We propose that biotic factors, such as host plant condition and codling moth population density, which might vary at smaller spatial scales, explain the changes in body mass and wing size and shape observed between locations. For instance, water availability for home-grown trees could be higher or the codling moth population density lower in Orilla de Maule than in Las Cruces and Yerbas Buenas, which could improve the availability of fruits as resources for the development of the codling moth and therefore allow a higher body mass and wing size in adults from this location in relation to others with lower fruit availability or higher codling moth population density (Ferro and Hardwood, 1973; Howell, 1991).

Previous studies detected behavioral differences between codling moth from apple and walnut trees in California (Cisneros and Barnes, 1974; Phillips and Barnes, 1975). Because no influence of the host plant species on body mass or wing morphology was found, our results did not support a phenotypic differentiation between apple and walnut populations in the Maule Region of central Chile. Walnut is often regarded as a secondary host for codling moth in comparison with apple because of the high content of secondary metabolites (e.g., juglone) that the walnut fruit contains (Piskorski and Dorn, 2011). Effective metabolism of juglone was found in codling moth larvae feeding on walnut, although a longer developmental period and a male biased sex ratio were observed when reared in this host plant (Piskorski and Dorn, 2011). In our study, we used adults obtained from apple and walnut trees; therefore, we selected those individuals who successfully developed on those host plants.

Our results confirm previous studies showing differences in body mass and wing morphology depending on codling moth sex and location but did not find differences between apple and walnut as host plants. Small individuals from home-grown trees should have high insecticide susceptibility and dispersal capabilities. The scenario of insecticide-susceptible adults migrating to managed orchards has been proposed as source-sink population dynamics that could explain the low levels of insecticide resistance found in codling moth in central Chile (Fuentes et al, 2014). Further studies could evaluate whether such differences based on sex and location in body mass and wing morphology could affect the dispersal of this species and subsequently influence its management through the use of insecticides and mating disruption.


This work was funded by Fondo de Desarrollo Científico y Tecnológico (FONDECYT) grant 1120374 to EFC and BL, and Programa de Atracción e Inserción de Capital Humano Avanzado en la Academia (CONICYT-PAI) 7912010027 to MR.



Barnes, M.M. 1991. Codling moth occurrence, host race formation, and damage. In: Van der Geest L.P.S. and H.H. Evenhuis (eds.). Tortricid pests: their biology, natural enemies, and control. Elsevier, Amsterdam. p. 313-327.         [ Links ]

Basoalto, E., M. Miranda, A.L. Knight, and E. Fuentes-Contreras. 2010. Landscape analysis of adult codling moth (Lepidoptera: Tortricidae) distribution and dispersal within typical agroecosystems dominated by apple production in central Chile. Environ. Entomol. 39: 1399-1408.         [ Links ]

Betts, C.R., and R.J. Wootton. 1988. Wing shape and flight behaviour in butterflies (Lepidoptera:

Papilionoidea and Hesperioidea): a preliminary analysis. J. Exp. Biol. 138: 271-288.

Bookstein, F.L. 1986. Size and shape spaces for landmark data in two dimensions. Statist. Sci. 1: 181-222.         [ Links ]

Bretz, F., T. Hothorn, and P. Westfall. 2011. Multiple comparisons using R. CRC Press, Boca Ratón, 182 pp.         [ Links ]

Bùes, R., J.F. Toubon, and H.S. Poitout. 1995. Variabilité écophysiologique et enzymatique de Cydia pomonella L. en fonction del’origine géographique et de la plante hôte. Agronomie 15: 221-231.

Cisneros, F.H., and M.M. Barnes. 1974. Contribution to the biological and ecological characterization of apple and walnut host races of codling moth, Laspeyresia pomonella (L.): moth longevity and oviposition capacity. Environ. Entomol. 3: 402-406.         [ Links ]

Crawley, M.J. 2013. The R book. 2nd Edition, Wiley, Chichester. 1076 pp.         [ Links ]

Ferro, D.N., and R.F. Hardwood. 1973. Intraspecific larval competition by the codling moth, Laspeyresia pomonella. Environ. Entomol. 2: 783-789.         [ Links ]

Fuentes-Contreras, E., E. Basoalto, P. Franck, B. Lavandero, A.L. Knight, and C.C. Ramírez. 2014. Measuring local genetic variability in populations of codling moth (Lepidoptera: Tortricidae) across an unmanaged and commercial orchard interface. Environ. Entomol. 43: 520-527.         [ Links ]

Fuentes-Contreras, E., M. Reyes, W. Barros, and B. Sauphanor. 2007. Evaluation of azinphosmethyl resistance and activity of detoxifying enzymes in codling moth (Lepidoptera: Tortricidae) from central Chile. J Econ. Entomol. 100: 551-556.         [ Links ]

Franck, P, and A.E. Timm. 2010. Population genetic structure of Cydia pomonella: a review and case study comparing spatiotemporal variation. J. Appl. Entomol. 134: 191-200.         [ Links ]

Gu, H.N., J. Hughes, and S. Dorn. 2006. Trade-off between mobility and fitness in Cydia pomonella L. (Lepidoptera: Tortricidae). Ecol. Entomol. 31: 68-74.         [ Links ]

Howell, J.F. 1991. Reproductive biology. In: Van der Geest L.P.S. and H.H. Evenhuis (eds.). Tortricid pests: their biology, natural enemies, and control. Elsevier, Amsterdam. p. 157-174.         [ Links ]

Keil, S., H.N. Gu, and S. Dorn. 2001. Diel patterns of locomotor activity in Cydia pomonella: age and sex related differences and effects of insect hormone mimics. Physiol. Entomol. 26: 306-314.         [ Links ]

Khaghaninia, S., S.A. Mohammadi, A.L. Sarafrazi, K.H.I. Nejad, and R. Zahiri. 2011a. Thin-plate spline analysis of diversity in field populations of codling moth, Cydia pomonella L. (Lep., Tortricidae) in North West of Iran. J Agric. Sci. Technol. 5: 35-41.         [ Links ]

Khaghaninia, S., S.A. Mohammadi, A.L. Sarafrazi, K.H.I. Nejad, and R. Zahiri. 2011b. Geometric morphometric study on geographic dimorphism of coding moth Cydia pomonella (Lepidoptera, Tortricidae) from North West of Iran. Vestn. Zool. 45: e-20-e-28.         [ Links ]

Klingenberg, C.P. 2011. MorphoJ: an integrated software package for geometric morphometrics. Mol. Ecol. Resour. 11: 353-357.         [ Links ]

Mazzi, D., and S. Dorn. 2012. Movement of insect pests in agricultural landscapes. Ann. Appl. Biol. 160: 97-113.         [ Links ]

Norberg, U., and O. Leimar. 2002. Spatial and temporal variation in flight morphology in the butterfly Melitaea cinxia (Lepidoptera: Nymphalidae). Biol. J. Linnean Soc. 77: 445-453.         [ Links ]

Phillips, P.H., and M.M. Barnes. 1975. Host race formation among sympatric apple, walnut and plum populations of the codling moth, Laspeyresia pomonella. Ann. Entomol. Soc. Amer. 68: 1053-1060.         [ Links ]

Piskorski, R., and S. Dorn. 2011. How the oligophage codling moth Cydia pomonella survives on wal-nut despite its secondary metabolite juglone. J. Insect Physiol. 57: 744-750.         [ Links ]

R Development Core Team. 2014. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available online at (Website accessed: october, 6th, 2014).         [ Links ]

Reyes, M., J.C. Bouvier, T. Boivin, E. Fuentes-Contreras, and B. Sauphanor. 2004. Susceptibilidad a insecticidas y actividad enzimática de Cydia pomonella L. (Lepidoptera: Tortricidae) proveniente de tres huertos de manzano de la Región del Maule. Agric. Téc. (Chile) 64: 229-237.         [ Links ]

Reyes, M., W. Barros-Parada, C.C. Ramírez, and E. Fuentes-Contreras. 2015. Organophosphate resistance and its main mechanism in populations of Cydia pomonella (Lepidoptera: Tortricidae) from central Chile. J. Econ. Entomol. 108: 277-285.         [ Links ]

Ricci, B., P. Franck, J.F. Toubon, J.C. Bouvier, B. Sauphanor, and C. Lavigne. 2009. The influence of landscape on insect pest dynamics: a case study in southeastern France. Landscape Ecol. 24: 337-349.         [ Links ]

Ricci, B., P. Franck, J.C. Bouvier, D. Casado, and C. Lavigne. 2011. Effects of hedgerow characteristics on intra-orchard distribution of larval codling moth. Agr. Ecosyst. Environ. 140: 395-400.         [ Links ]

Roff, D.A., and D.J. Fairbairn. 2007. The evolution and genetics of migration in insects. BioScience 57: 155-164.         [ Links ]

Rohlf, FJ. 2006. TpsDig2. Ecology and Evolution. SUNY Stony Brook, NY, USA.         [ Links ]

Sauphanor, B., V Brosse, J.C. Bouvier, P. Speich, A. Micoud, and C. Martinet. 2000. Monitoring resistance to diflubenzuron and deltamethrin in French codling moth populations (Cydia pomonella). Pest Manag. Sci. 56: 74-82.         [ Links ]

Schumacher, P., D.C. Weber, C. Hagger, and S. Dorn. 1997a. Heritability of flight distance for Cydia pomonella. Entomol. Exp. Appl. 85: 169-175.         [ Links ]

Schumacher, P., A. Weyeneth, D.C. Weber, and S. Dorn. 1997b. Long flights in Cydia pomonella L. (Lepidoptera: Tortricidae) measured by a flight mill: influence of sex, mated status and age. Physiol. Entomol. 22: 149-160.         [ Links ]

Stoeckli, S., K. Mody, and S. Dorn. 2008. Influence of canopy aspect and height on codling moth (Lepidoptera: Tortricidae) larval infestation in apple, and relationship between infestation and fruit size. J. Econ. Entomol. 101: 81-89.         [ Links ]

Thaler, R., A. Brandstatter, A. Meraner, M. Chabicovski, W. Parson, R. Zelger, J. Dalla Via, and R. Dallinger. 2008. Molecular phylogeny and population structure of the codling moth (Cydia pomonella) in Central Europe: II. AFLP analysis reflects human-aided local adaptation of a global pest species. Mol. Phyl. Evol. 48: 838-849.         [ Links ]

Varela, L.G., S.C. Welter, V.P. Jones, J.F. Brunner, and H. Riedl. 1993. Monitoring and character-ization of insecticide resistance in codling moth (Lepidoptera, Tortricidae) in four Western States. J. Econ. Entomol. 86: 1-10.         [ Links ]

Weddle, P.W., S.C. Welter, and D. Thomson. 2009 History of IPM in California pears-50 years of pesticide use and the transition to biologically intensive IPM. Pest Manag. Sci. 65: 1287-1292.         [ Links ]

Witzgall, P., L. Stelinski, L. Gut, and D. Thomson. 2008. Codling moth management and chemical ecology. Annu. Rev. Entomol. 53: 503-522.         [ Links ]


Received March 15, 2015.
Accepted July 24, 2015.


Creative Commons License Todo el contenido de esta revista, excepto dónde está identificado, está bajo una Licencia Creative Commons