Life Cycle of Agrotis Malefida (Lepidoptera: Noctuidae): a Diapausing Cutworm
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ZOOLOGIA 30 (4): 371–378, August, 2013 http://dx.doi.org/10.1590/S1984-46702013000400002 Life cycle of Agrotis malefida (Lepidoptera: Noctuidae): a diapausing cutworm Alexandre Specht1,5, Andrés O. Angulo2, Tania S. Olivares2, Edegar Fronza3, Vânia F. Roque-Specht4, Eduardo Valduga3, Francine Albrecht3, Graziela Poletto3 & Neiva M. Barros3 1 Laboratório de Entomologia, Embrapa Cerrados. Caixa Postal 08223, 73310-970 Planaltina, DF, Brazil. 2 Departamento de Zoología, Facultad de Ciencias Naturales y Oceanográficas, Universidad Concepción. Casilla 160-C, Concepción, Chile. 3 Instituto de Biotecnologia, Universidade de Caxias do Sul. Caixa Postal 1352, 95070-560 Caxias do Sul, RS, Brazil. 4 Faculdade UnB Planaltina. Área Universitária 1, Vila Nossa Senhora de Fátima, 73300-000 Planaltina, DF, Brazil. 5 Corresponding author. E-mail: [email protected] ABSTRACT. This study describes the life cycle of Agrotis malefida Guenée, 1852 (Noctuidae: Noctuinae) under laboratory conditions. The insects were reared in a controlled environment (25 ± 1°C, 70 ± 10% RH and 14 hours photo phase) and observed daily. The larvae were fed Greene’s artificial diet and adults were offered a 10% sucrose solution. The viability and duration of immature stages were assessed. The experiment initiated with 2,410 eggs. Larvae were isolated shortly after hatching. Longevity, pre-, post- and oviposition, fecundity and fertility of 13 adult couples were also evaluated. The viability of eggs, larvae, pupae and pre-pupae was 96.72, 91.25, 78.37 and 95.26%, respectively. The average duration of egg, larva, pre-pupa, pupa and adult was 7.93, 54.26, 61.61, 37.43 and 12.85 days, respectively. The immature stage of A. malefida lasted an average of 161.29 days, ranging from 102 to 227 days. The life cycle of A. malefida is much longer than that of congeners. The mean fecundity was 1,696.77 eggs and fertility 1,641.15 larvae per female. Under the conditions in which the study was conducted, the biotic potential of A. malefida was of 606,666.59 individuals/female/ year. The results also indicated that this species goes through larval (pre-pupae) and pupal diapause. KEY WORDS. Bionomy; caterpillar; development; diapause. Agrotis Ochsenheimer, 1816 (Noctuidae: Noctuinae: Larvae of A. malefida are polyphagous, attacking various Noctuini) (LAFONTAINE & SCHMIDT 2010) is a cosmopolitan genus herbaceous, native and cultivated plants, including economi- with 317 species (POOLE 1989). All Agrotis larvae are voracious cally relevant crops such as alfalfa, barley, bean, beet, cabbage, and are able to feed on a variety of hosts (KITCHING & RAWLINS cauliflower, clover, coffee, collard, corn, cotton, cowpea, cu- 1998, POGUE 2006). They have underground habits, coming to cumber, flax, garlic, lettuce, melon, oat, onion, pea, pepper, the surface at night to feed. While feeding, they cut the stems potato, rice, rye, sorghum, soybean, squash, strawberry, sun- of their food plants, causing severe damage and rapid crop loss. flower, Swiss chard, tobacco, tomato and wheat (SILVA et al. 1968, When disturbed, Agrotis larvae roll up in a very characteristic VILLATA et al. 1988, COTTO et al. 1995, RIZZO et al. 1995, PASTRANA behavior (BAUDINO 2004, LAFONTAINE 2004, ANGULO et al. 2008). 2004, SPECHT et al. 2004, BAUDINO & VILLARREAL 2007, ANGULO et Agrotis malefida Guenée, 1852 has been recorded from al. 2008). Larvae often build a subterranean tunnel which is most of the Americas (PASTRANA & HERNANDES 1979, RIZZO et al. used as a hiding place during the day, and as a place to eat the 1995, LAFONTAINE 2004) and is likely to occur worldwide (ANGULO host plants cut during the night (LAFONTAINE 2004). & QUEZADA 1975). The species has been widely documented in The economic relevance of A. malefida has motivated the United States and Mexico (LAFONTAINE 2004, POGUE 2006), various studies on the morphology of the adult (ANGULO & Chile (ANGULO & QUEZADA 1975, ANGULO et al. 2008), Argentina QUEZADA 1975) and of immature stages (RIZZO et al. 1995, ANGULO (MARGHERITIS & RIZZO 1965, IGARZÁBAL et al. 1994, RIZZO et al. 1995, et al. 2008), along with biological studies (VILLATA et al. 1988, BAUDINO 2004, PASTRANA 2004), Brazil (SILVA et al. 1968, SPECHT & RIZZO et al. 1995, BAUDINO 2004, BAUDINO & VILLARREAL 2007). CORSEUIL 1996, 2002, SPECHT et al. 2004) and Uruguay Despite the morphological similarities of the adults and the (BENTANCOURT & SCATONI 2006). Larvae of A. malefida are known sympatry of Agrotis ipsilon (Hufnagel, 1766) and A. malefida in in North America as Pale-Sided Cutworms; in Latin America as the American continent (ANGULO & QUEZADA 1975), previous “gusano áspero”, “oruga áspera” or “oruga cortadora áspera” studies indicate several biological differences, especially in the (LAFONTAINE 2004); and in Brazil as “lagarta-rosca” (SILVA et al. immature stages of A. malefida (VILLATA et al. 1988, RIZZO et al. 1968). 1995, BAUDINO 2004, BAUDINO & VILLARREAL 2007). In view of the © 2013 Sociedade Brasileira de Zoologia | www.sbzoologia.org.br | All rights reserved. 372 A. Specht et al. importance of the species, and its large intraspecific variabil- SILVEIRA NETO et al. (1976) and the following parameters were de- ity (VILLATA et al. 1988, RIZZO et al. 1995), we have endeavored termined: net reproductive rate (Ro), mean generation time (T), to describe the biological parameters and to estimate the bi- natural intrinsic growth rate (rm), and infinite growth rate (⌳). otic potential of A. malefida in the laboratory, using individual Monthly collections of A. malefida moths were made observations on thousands of specimens. within the CARVI (29°08’99"S; 51°31’39"W, 675 m a.s.l.), from January 2007 to December 2008, four nights during each new MATERIAL AND METHODS moon, using light traps as described in SPECHT et al. (2005), in order to obtain information on the period of adult occurrence. Our laboratory rearing experiment initiated with 13 The temporal and morphometric parameters were ana- couples obtained from a mass rearing colony maintained by lyzed using descriptive statistics to calculate the means and stan- the Laboratório de Biologia, Campus Universitário da Região dard errors. When necessary, the means were compared using a dos Vinhedos (CARVI), Universidade de Caxias do Sul (UCS). t-test assuming unequal variances, at a significance level of 95%. The mass rearing itself originated from individuals collected Voucher specimens (10 couples) were deposited in the in the Vale dos Vinhedos, Bento Gonçalves, state of Rio Grande collection of the CARVI-UCS. do Sul. Our experiments were conducted in a controlled envi- ronment (25 ± 1°C, 70 ± 10% UR and 14 hours photo phase), RESULTS with daily observations. Each adult couple (n = 13) was maintained inside a cy- Adult stage lindrical plastic container (10 cm diameter, 15 cm height). In The longevity of A. malefida did not significantly differ order to stimulate oviposition, strings of filter-paper of the same between the sexes (p = 0.076), and represented 7.38% of the length as the plastic vial were inserted into each container and developmental cycle (Fig. 1, Table I). Under the conditions of attached using a plastic film cover. The moths were fed an aque- this study, adults of A. malefida reach sexual maturation soon ous sucrose solution (10%), made available in hydrophilic cot- after emergence (Table II). The egg laying period occurs from ton inside a 10 mL glass Erlenmeyer flask. The following the second to the tenth day after sexual maturation, and the parameters were evaluated: fecundity (number of eggs per fe- highest fecundity was observed between the seventh and the male), fertility (number of hatched larvae per female), longev- tenth day (Fig. 2). The fecundity of A. malefida varied greatly ity and duration of pre-oviposition, post-oviposition and between individuals resulting in approximately 1,700 eggs per oviposition periods. female (Table II). During the egg stage, we assessed the viability and dura- tion of the incubation period of five random ovipositions from each couple (including first and last ovipositions), totaling Table I. Survival and duration of the life cycle of Agrotis malefida during different stages of development, under controlled 2,410 eggs. Each egg mass was individually placed in a Petri conditions (25 ± 1°C, 70 ± 10% RH and 14 hours photo phase). dish, where it remained until the larvae hatched. The bottom Developmental Survival Duration Inicial-final N Range of each Petri dish was lined using filter paper moistened with stage (%) (days) distilled water, in order to avoid desiccation. Adult 26 – 12.85 ± 3.78 5-22 After hatching, neonate larvae were separately placed in Egg 2,410-2,331 96.72 7.93 ± 1.22 6-10 150 ml plastic cups, covered with a lid, and fed an artificial diet Larvae 2,331-2,127 91.25 54.25 ± 6.39 44-77 (GREENE et al. 1976), previously used successfully by RIZZO et al. Pre-pupae 2,127-1,667 78.37 61.61 ± 17.56 10-131 (1995) for the same species. We evaluated the survival and the Pupae 1,667-1,588 95.26 37.43 ± 4.13 11-99 duration of the larval period of 2,331 individuals. The pre-pu- Total immature 1,588 – 161.29 ± 17.35 102-227 pal period was considered separately, and its onset was recorded Total – 65.89 174.08 – when the larva stopped feeding and decreased in size. We added expanded vermiculite, moistened with distilled water, to each cup, in order to allow the pre-pupae to build a pupal chamber. Table II. Mean fecundity and duration of the pre-, post-, and Pupae were kept in the same containers as pre-pupae. oviposition periods (days) of Agrotis malefida reared under controlled conditions (25 ± 1°C, 70 ± 10% RH and 14 hours photo On the second day after pupation, when the cuticle had hard- phase). (N = 13 couples).