ESTIMATE of Alabama Argillacea (HÜBNER) (LEPIDOPTERA: NOCTUIDAE) DEVELOPMENT with NONLINEAR MODELS
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ESTIMATE OF Alabama argillacea DEVELOPMENT WITH NONLINEAR MODELS 589 ESTIMATE OF Alabama argillacea (HÜBNER) (LEPIDOPTERA: NOCTUIDAE) DEVELOPMENT WITH NONLINEAR MODELS MEDEIROS, R. S.,1 RAMALHO, F. S.,2 ZANUNCIO, J. C.1 and SERRÃO, J. E.3 1Departamento de Biologia Animal, Universidade Federal de Viçosa, CEP 36571-000, Viçosa, Minas Gerais, Brazil 2Embrapa Algodão, Rua Osvaldo Cruz, 1143, C.P. 174, CEP 58107-720, Campina Grande, Paraíba, Brazil 3Departamento de Biologia Geral, Universidade Federal de Viçosa, CEP 36571-000, Viçosa, Minas Gerais, Brazil Correspondence to: José Eduardo Serrão, Departamento de Biologia Geral, Universidade Federal de Viçosa, CEP 36571-000, Viçosa, Minas Gerais, Brazil, e-mail: [email protected] Received July 2, 2002 – Accepted August 27, 2002 – Distributed November 30, 2003 (With 1 figure) ABSTRACT The objective of this work was to evaluate which nonlinear model [Davidson (1942, 1944), Stinner et al. (1974), Sharpe & DeMichele (1977), and Lactin et al. (1995)] best describes the relationship be- tween developmental rates of the different instars and stages of Alabama argillacea (Hübner) (Lepi- doptera: Noctuidae), and temperature. A. argillacea larvae were fed with cotton leaves (Gossypium hirsutum L., race latifolium Hutch., cultivar CNPA 7H) at constant temperatures of 20, 23, 25, 28, 30, 33, and 35oC; relative humidity of 60 ± 10%; and photoperiod of 14:10 L:D. Low R2 values obtained with Davidson (0.0001 to 0.1179) and Stinner et al. (0.0099 to 0.8296) models indicated a poor fit of their data for A. argillacea. However, high R2 values of Sharpe & DeMichele (0.9677 to 0.9997) and Lactin et al. (0.9684 to 0.9997) models indicated a better fit for estimating A. argillacea development. Key words: cotton leafworm, developmental rate, Gossypium hirsutum, temperature. RESUMO Estimativa do desenvolvimento de Alabama argillacea (Hübner) (Lepidoptera: Noctuidae) por meio de modelos não lineares O objetivo deste estudo foi avaliar qual modelo não linear [Davidson (1942, 1944), Stinner et al. (1974), Sharpe & DeMichele (1977) e Lactin et al. (1995)] descreve melhor a relação entre as taxas de desenvolvimento dos diferentes ínstares e fases de Alabama argillacea (Hübner) e a temperatura. As lagartas de A. argillacea foram alimentadas com folhas de algodoeiro (Gossypium hirsutum L., raça latifolium Hutch., cultivar CNPA 7H) e submetidas às temperaturas constantes de 20, 23, 25, 28, 30, 33 e 35oC, umidade relativa de 60 ± 10% e fotoperíodo de 14 horas. Os baixos valores de R2 obtidos para os modelos de Davidson (0,0001 a 0,1179) e de Stinner et al. (0,0099 a 0,8296) indicaram que eles não são adequados para estimar o desenvolvimento de A. argillacea em função da temperatura. Entretanto, os altos valores de R2 dos modelos de Sharpe & DeMichele (0,9677 a 0,9997) e de Lactin et al. (0.9684 a 0.9997) indicaram que os mesmos estimam, adequadamente, o tempo de desenvol- vimento de A. argillacea em função da temperatura. Palavras-chave: curuquerê-do-algodoeiro, taxa de desenvolvimento, algodoeiro, temperatura. Braz. J. Biol., 63(4): 589-598, 2003 590 MEDEIROS, R. S. et al. INTRODUCTION using logistic equations. Stinner et al. (1974) described the temperature effect on developmental rate as a Alabama argillacea (Hübner) (Lepidoptera: modified sigmoid equation that results in a Noctuidae) is a native species of Central and South symmetrical curve at higher temperatures. Sharpe & America found in most areas where cotton is DeMichele (1977) formulated a complex biophysical cultivated, from South Canada to northern Argentina model, later modified by Schoolfield et al. (1981), (Carvalho, 1981). This species occurs during the which describes a nonlinear response of developmental whole cultivation period of cotton plants, with rate of insects exposed to low and high temperatures increasing populations as the cycle progresses. This as well as a linear response at intermediate insect defoliates cotton plants from the lower to the temperatures (Wagner et al., 1984). Lactin et al. (1995) upper parts, with most damage occurring during the modified the nonlinear model of Logan et al. (1976) last three instars (Bellettini et al., 1999). A. argillacea by eliminating the parameter y and introducing is the main cotton defoliator pest, greatly impacting parameter intercepts l, which allowed estimation of cotton plant productivity in Argentina, Brazil, the developmental threshold. Colombia, Mexico, Nicaragua, Paraguay, Peru, and Because linear models are not very precise the USA (Almestar et al., 1977; Falcon & Daryl, sources of information on developmental rate 1977; Cies, 1978; Alvarez & Sanchez, 1982; inhibition at extreme temperatures, the purpose of Nyffeler et al., 1987; Michel, 1994; Ramalho, 1994; this research was to determine which nonlinear model Lobos, 1999). A. argillacea is a lesser pest in South- (Davidson, 1942, 1944; Stinner et al., 1974; Sharpe Central Brazil but, except for the State of Bahia, & DeMichele, 1977; Lactin et al., 1995) better in the Northeast this pest can cause damage during describes the temperature effect on the developmental the whole cotton crop cycle (Gravena & Cunha, rate of A. argillacea. 1991). The introduction of another cotton pest in Brazil, Anthonomus grandis Boheman (Coleoptera: MATERIAL AND METHODS Curculionidae), complicated pest control in this crop, This research was developed at the Biological resulting in a significant reduction in the area Control Unit (UCB)/Embrapa Algodão, in Campina cultivated (Ramalho et al., 1989). New areas for Grande, State of Paraíba, Brazil. Specimens of A. agriculture in Brazil such as the savannah regions argillacea were collected in Touros, State of Rio in the States of Mato Grosso do Sul and Goiás have Grande do Norte, and maintained in BOD at constant added the importance of A. argillacea as a cotton temperatures of 20, 23, 25, 28, 30, 33, and 35oC, crop pest. Besides, the appearance of populations relative humidity of 60 ± 10%, and photoperiod of resistant to insecticides has been increasing problems 14:10 L:D. Larvae of this species were fed with with pest control in this crop. cotton plant leaves (Gossypium hirsutum L., race The relationship between insect developmental latifolium Hutch., cultivar CNPA 7H) while adults rate and temperature represents an important received a solution of honey (20%) and distilled ecological variable for modelling population water (80%). dynamics of these organisms. Linear models were Newly emerged adults were used to form 15 the first developed for insects (Howe, 1967) but the pairs of A. argillacea, with five of them per PVC lack of linearity of insect developmental rate at low cage (Medeiros, 1997). First egg clutches of A. and high temperatures suggests that such models argillacea were placed in Petri dishes (9.0 x 1.5 cm) are inadequate to describe this parameter for these with a moist cotton ball, and observed daily to de- organisms. Since early 1980 this problem has led termine their incubation period and viability at the to increasing interest in developing nonlinear same temperatures and humidity conditions used phenological models in integrated pest management for A. argillacea pairs. programs (Wagner et al., 1984). The fifty-first instar of A. argillacea larvae Nonlinear models (Logan et al., 1976) have were individualized in plastic cups (250 ml) for each been elaborated for several insect species in certain treatment. A cylindrical plastic tube (2.5 cm) with circumstances. Davidson (1942, 1944) described the distilled water was inserted in a circular hole in the insect developmental rate as a function of temperature cover of each plastic cup. One fresh cotton leaf was Braz. J. Biol., 63(4): 589-598, 2003 ESTIMATE OF Alabama argillacea DEVELOPMENT WITH NONLINEAR MODELS 591 removed from apical parts of plants and used daily The parameters a, b, and k were estimated with as food for A. argillacea larvae. The dorsal surface the regression nonlinear model of Marquardt using of these larvae was marked with dye (Day-Glo PROC NLIN (Sas Institute Inc., 2000). This method Colour Corp) to facilitate observation of ecdyses. is used to determine the minimum square of the Everyday, results were recorded and cotton leaves parameters estimated with this model. substituted. Mean developmental rate of egg, instars, 2) sigmoid equation of Stinner et al. (1974): prepupae, and pupae of A. argillacea at different c temperatures was estimated with the formula: rT( ) = , (1+ ek12+kT' ) ïïìüéùn íýêúå ln(din) /, r(Te) =1.0/ îþïïëûi=1 where r(T) is the mean developmental rate at tem- o x k1 + k2Tmax perature T ( C); c, (1/Tmax) (e ) (asymptote); where r(T) is the mean developmental rate at tem- k and k , empirical constants; and T’ = T, for T < o 1 2 perature T ( C); di, individual observations of de- T and T’ = 2 x T – T, for T > T . velopment period in days; and n, number of max max max The parameters c, k1, and k2 were estimated observations. with Marquardt’s method using PROC NLIN (Sas This method is recommended by Logan et al. Institute Inc., 2000). (1976) to account for linearity in the transformation of development period to developmental rate. 3) biophysical model of Sharpe & DeMichele Developmental rate is the reciprocal of (1977), modified by Schoolfield et al. (1981): development period in days and represented by values from 0 to 1. These rates are used in development models éùæö æöT êúæöH ç÷1 where data are added daily. Organism development RHO exp êúA 25 ç÷ç÷ç÷1 is completed when the sum of their daily developmental èø298.15 êúèøR ç÷298.15 - êúèøT rate reaches value 1 (Curry & Feldman, 1987). rT( ) = ëû éùéù Therefore, the integral of the developmental rate æHHLHöæ11öæöæö11 1+expêç÷ç-÷ú+-exp êúç÷ç÷ function along time (as in the models of Davidson, ëêèRTøèLHTøûúëûêúèøRTTèø 1942, 1944; Stinner et al., 1974; Sharpe & DeMichele, 1977; Lactin et al., 1995) can be used to simulate the where r(T) is the mean developmental rate at tem- development of an organism submitted to different perature T (oK); R, universal gas constant (1.987 cal –1 –1 o temperatures.