Interciencia ISSN: 0378-1844 [email protected] Asociación Interciencia Venezuela

Azevedo Pereira, Alexandre Igor; Vinha Zanuncio, Virginia; Simões Lorenzon, Alexandre; Bolognani, Helder; Vique Fernandes, Bianca; Hendrik Mielke, Olaf Hermann; Serrão, José Eduardo; Cola Zanuncio, José BIOLOGICAL AND MORPHOLOGICAL CHARACTERISTICS OF Hylesia paulex (: ) FED WITH Eucalyptus urophylla (MYRTACEAE) Interciencia, vol. 34, núm. 9, septiembre, 2009, pp. 645-649 Asociación Interciencia Caracas, Venezuela

Available in: http://www.redalyc.org/articulo.oa?id=33913149008

How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Biological and Morphological CHARACTERISTICS of Hylesia paulex (Lepidoptera: Saturniidae) Fed with Eucalyptus urophylla (Myrtaceae)

Alexandre Igor Azevedo Pereira, Virginia Vinha Zanuncio, Alexandre Simões Lorenzon, Hélder Bolognani, Bianca Vique Fernandes, Olaf Hermann Hendrik Mielke, José Eduardo Serrão and José Cola Zanuncio

SUMMARY

Hylesia spp. (Saturniidae), important for public health due ±14.52% viability, and the duration of the larval stage reached to urticating bristles of their larvae and/or adults, are collected in 67.83 ±0.84 days, with seven instars. The gregarious behavior is different habitats with potential to defoliate plants in agriculture important to maintain a high survival rate during this stage for and forest systems. Biological and morphological characteristics of H. paulex. The pre-pupa and pupa stages of this lasted 3.87 Hylesia paulex (Dognin, 1922) (Lepidoptera: Saturniidae) fed with ±0.16 and 21.67 ±0.79 days, respectively. The longevity of females Eucalyptus urophylla (Myrtaceae) were studied in the laboratory. of this was longer (4.50 ±0.54 days) than that of males Recently-emerged caterpillars of this species were separated in 10 (2.80 ±0.44 days). The mean number of eggs per egg mass was plastic pots (500ml) with 30 of them per pot, and fed daily with 132.3. Moths of H. paulex completed its life cycle, satisfactorily, fresh leaves of E. urophylla until they reached adult stage. The egg with leaves of E. urophylla in the laboratory. incubation period of H. paulex was 32.00 ±1.19 days with 65.32

Introduction belong to more than one of ing substances and provoke agricultural and forest pests these categories as Saturniidae burns in mammals (Fornés and (Borges et al., 2003; Freitas et can be of medi­ moths do. Individuals of this Hernandéz, 2001; Gouveia et al., 2005; Specht et al., 2006a) cal, agricultural or veterinary family have defense structures al., 2005; Lorini et al., 2007). and social importance as food importance, but few groups with bristles that inject urticat­ They also have potential as for humans (Ande, 2003).

KEYWORDS / Gregarism / / Lepidopterism / / Received: 02/25/2008. Modified: 08/24/2009. Accepted: 08/25/2009.

Alexandre Igor Azevedo Pereira. Alexandre Simões Lorenzon. Olaf Hermann Hendrik Mielke. José Cola Zanuncio. M.Sc. Agronomist, Universidade Fede­ Agronomist and M.Sc. student, Doctor in Entomology, Uni­ in Entomology, USP, Brazil. ral da Paraíba, Brazil. M.S. in UFV, Brazil. e-mail: aleloren­ versidade Federal do Paraná Ph.D. in Entomology, Uni­ Entomology and Doctoral stu­ [email protected] (UFPR), Brazil. Professor, versity of British Columbia, dent, Universidade Federal de Helder Bolognani. Researcher, V UFPR, Brazil. Canada. Professor. UFV, Bra­ Viçosa (UFV), Brazil. e-mail: & M Florestal, Brazil. e-mail: José Eduardo Serrão. Doctor zil. Address: Departamento [email protected] [email protected] in Biological Sciences, Uni­ de Biologia /BIOA­ Virginia Vinha Zanuncio. M.D., Bianca Vique Fernandes. Forest versidade de São Paulo, USP, GRO, UFV. CEP 36.571-000; UFV, Brazil. Current Position: Engineer, V & M Florestal, Brasil. Professor, UFV, Brazil. Viçosa, MG, Brasil. e-mail: Phisician, Universidade Federal e-mail: bianca.fernandes@vm­ e-mail: [email protected] [email protected] de Juiz de Fora, Brazil. e-mail: tubes.com.br e-mail: vivizanuncio@yahoo. com.br

SEP 2009, VOL. 34 Nº 9 0378-1844/09/09/645-05 $ 3.00/0 645 CARACTERÍSTICAS Biológicas y Morfológicas de Hylesia paulex (Lepidoptera: Saturniidae) alimentadas con Eucalyptus urophylla (Myrtaceae) Alexandre Igor Azevedo Pereira, Virginia Vinha Zanuncio, Alexandre Simões Lorenzon, Hélder Bolognani, Bianca Vique Fernandes, Olaf Hermann Hendrik Mielke, José Eduardo Serrão y José Cola Zanuncio RESUMEN

Hylesia spp. (Saturnidae), una polilla importante en salud pública huevos de H. paulex fue de 32 ±1,19 días con un 65,32 ±14,52% de debido a las cerdas urticantes de las larvas y/o de los adultos, son viabilidad y duración del estado larval de 67,83 ±0,84 días, con sie- recolectadas en diferentes ambientes con potencial de desfoliación de te instares. El comportamiento gregario es importante para mantener plantas tanto en agricultura como en sistemas forestales. Caracte- una alta tasa de supervivencia durante estos estados en H. paulex. rísticas biológicas y morfológicas de Hylesia paulex (Dognin, 1922) Los estados de pre-pupa y pupa del insecto duraron 3,87 ±0,16 y (Lepidoptera: Saturniidae) alimentados con Eucalyptus urophylla fue- 21,67 ±0,79 días, respectivamente. La longevidad de las hembras de ron estudiados en laboratorio. Larvas recién emergidas de esa espe- la especie fue de 4,50 ±0,54 días y para machos de 2,80 ±0,44 días. cie fueron separadas en vasos plásticos (500 ml) con 30 individuos El número medio de huevos por masa de huevos fue de 132,3. La por vaso y alimentados diariamente con hojas frescas de E. urophy­ polilla H. paulex completa su ciclo de vida, satisfactoriamente, con lla hasta alcanzar el estado adulto. El periodo de incubación de los hojas de E. urophylla en laboratorio.

CARACTERÍSTICAS Biológicas e Morfológicas da Hylesia paulex (Lepidoptera: Saturniidae) alimentadas com Eucalyptus urophylla (Myrtaceae) Alexandre Igor Azevedo Pereira, Virginia Vinha Zanuncio, Alexandre Simões Lorenzon, Hélder Bolognani, Bianca Vique Fernandes, Olaf Hermann Hendrik Mielke, José Eduardo Serrão e José Cola Zanuncio RESUMO

Hylesia spp. (Saturniidae), mariposas importantes para a saúde ovos de H. paulex foi de 32,00 ±1,19 dias com 65,32 ±14,52% pública devido às cerdas urticantes de suas larvas e/ou adultos, de viabilidade e duração do estágio larval de 67,83 ±0,84 dias, são coletadas em diferentes hábitats com potencial de desfolhar com sete estádios. O comportamento gregário é importante para plantas em sistemas agrícolas e florestais. Parâmetros biológicos manter a alta taxa de sobrevivência durante esses estádios para e morfológicos de Hylesia paulex (Dognin, 1922) (Lepidoptera: H. paulex. Os estágios de pré-pupa e pupa desse inseto duraram Saturniidae) alimentada com Eucalyptus urophylla (Myrtaceae) 3,87 ±0,16 e 21,67 ±0,79 dias, respectivamente. A longevida- foram estudados em laboratório. Lagartas recém-emergidas dessa de das fêmeas dessa espécie foi maior (4,50 ±0,54 dias) que a espécie foram separadas em 10 potes plásticos (500ml) com 30 dos machos (2,80 ±0,44 dias). O número de ovos/postura foi de delas por pote e alimentadas, diariamente, com folhas frescas de 132,3. Mariposas de H. paulex completaram seu ciclo de vida, E. urophylla até o estágio adulto. O período de incubação dos satisfatoriamente, com folhas de E. urophylla em laboratório.

The majority of the stud­ maire, 2002; Camargo, 2007), Material and Methods and transferred to Petri dishes ies on the Saturniidae fam­ except for research on Hylesia (9.0×1.5cm) and kept under ac­ ily (Rodríguez-Morales et al., nanus (Walker) by Santos et The study was carried out climatized chambers under the 2005; Salomon et al., 2005; al. (1996), H. nigricans (Berg) at the Laboratory of Biological same conditions as the pupa Iserhard et al., 2007; Lund­ by Specht et al. (2006a) and Control, Institute of Applied stage. Groups of 30 newly- berg et al., 2007) refer to its H. metapyrrha (Walker) by Biotechnology to Agriculture hatched caterpillars of H. paulex capacity to cause dermatologic Specht et al. (2007). However, (BIOAGRO), Federal University were put in ten 500ml plastic irritations as for Hylesia spp. other studies should be per­ of Viçosa (UFV), Viçosa, Minas pots, with a screened cover in (Saturniidae: Hemileucinae). formed to obtain information Gerais, Brazil. the center, and fed daily with They are small to medium size on the behavior, life cycle and Pupae of Hylesia paulex fresh leaves of E. urophylla. moths, found from Mexico to adaptability of other species (Dognin, 1922) (Lepidoptera: The petiole of each eucalyptus Argentina (Lemaire, 2002). of that genus to specific hosts. Saturniidae) were collected in leaf was wrapped in a moist­ The occurrence of these moths Such data is important to pre­ a Eucalyptus spp. (Myrtaceae) ened cotton wad to avoid drying has also been recorded in pop­ vent lepidopterism and erucism plantation, in an area of savan­ and inserted in a plastic tube of ulation outbreaks in Brazil cases (dermatitis provoked by nah of Minas Gerais State, Bra­ odontological type anesthetic in agriculture (Camargo and moths and caterpillars, respec­ zil, and individualized in Petri (2.5ml) with distilled water. Becker, 1999; Glasser et al., tively) (Rodríguez-Morales dishes (9.0×1.5cm) at 25 ±2oC The eucalyptus leaves were 1993; Camargo, 2007; Iserhard et al., 2005) and to develop temperature, 60 ±10% relative washed in running water and et al., 2007; Moreira et al., combat strategies against them humidity and 12h photoperiod. bathed in sodium hypochlo­ 2007) and forest monocultures (Salomon et al., 2005). Recently-emerged H. pau­ rite solution (10%) before be­ (Zanuncio et al., 1998, 2001, The present study had as a lex adults were maintained ing supplied to H. paulex, and 2006). main goal the study the biolog­ in screened wood cages renewed every 48h. The pots The genus Hylesia presents ical and morphological aspects 60×60×60cm) under field con­ were cleaned daily and substi­ ~110 species (Lemaire, 2002) of H. paulex, under laboratory ditions, ~30 pairs per cage, in tuted when necessary, while the but the biology of most of conditions, fed with Eucalyptus order to obtain egg masses that material used by the caterpillars them is poorly studied (Le­ urophylla (Myrtaceae) leaves. were collected on dry cotton to weave its pre-pupa and pupa

646 SEP 2009, VOL. 34 Nº 9 (T5) caterpillars per pot. Leaves females laid unfertile eggs that of E. urophylla were offered were not covered by hairs, as ad libitum every 24h and the reported (Gardiner, 1967) for development and survival of the Periphoba hircia (Cramer) (Lep­ insects, as well as the weight idoptera: Saturniidae). of male and female pupae were The larval stage of H. paulex registered. lasted 67.83 ±0.84 days with Adult specimens of H. paulex seven instars (Table I). Females were deposited in the Depart­ of Automeris randa (Druce) ment of Zoology, Federal Uni­ (Lepidoptera: Saturniidae) and versity of Paraná, Paraná State, H. nigricans had an additional Brazil, and in the Museum of instar to those of males (Tuskes, Entomology and Laboratory of 1985; Specht et al., 2006a), Biological Control of BIOAGRO, which can be related to the sex UFV, Brazil. dimorphism of these species, but this was not observed for Results and Discussion H. paulex. The caterpillars of this species fed on the egg co­ The eggs of H. paulex have rium during the first hours of an oval shape, round extremities, life, when they showed slow flat in their long axis, clear beige movements. Afterwards, they color, are deposited in layers and abandoned the egg mass and measure 1.71 ±0.07mm long by moved towards the E. urophylla 0.61 ±0.04mm wide (Figure 1a). leaves inside the pots. Eucalyp- The values of the incubation pe­ tus spp. defoliators may prefer riod (32.00 ±1.19 days) and egg younger leaves mainly in their viability (65.32 ±14.52%) were initial stages because they pose intermediate between those of a lower physical impediment by 21.8 ±0.6 days and 16.2 ±8.1% being more turgid (Steinbauer for H. nanus (Walker), 90.83 and Matsuki, 2004). However, ±4.23 days and 78.28 ±5.84% younger H. paulex larvae did not for H. nigricans (Berg), and 31.8 follow this pattern and preferred ±5.8 days and 80.9 ±20.97% older E. urophylla leaves than for Lonomia obliqua (Walker) younger ones, independently of (Lepidoptera: Saturni­idae) re­ instar, as found for most species ported by Santos et al. (1996), of this plant in Brazil. Figure 1. a: Clutch (bar: 2mm), b: eggs (bar: 10mm), c: third instar larvae (bar: 5mm), d: seventh instar larva (bar: 10mm), e: irritant hairs (bar: 2mm), f: pupa Specht et al. (2006a), and Lori­ Recently-emerged H. paulex (bar: 10mm), g: adult female (bar: 10mm), and h: adult male (bar: 8mm) of Hyl- ni et al. (2004), respectively. caterpillars presented a black esia paulex (Lepidoptera: Saturniidae) fed with Eucalyptus urophylla (Myrtaceae) Egg masses of H. paulex were head and a clear orange ventral leaves, kept at 25 ±2oC, 60 ±10% relative humidity and 12h photoperiod. always covered by urticating abdomen area, while its dorsal hairs (Figure 1b) removed from surface was clear brown (cara­ the lateral-areas of the abdomen mel) or clear ash with yellowish stages shelter (branches and with water to test if this substra­ of the females, which indicates thorn tufts. During development remains of leaves) was main­ tum is preferred by H. paulex a possible defensive adaptation the ventral area turns whitish tained. females to lay their eggs. to avoid parasitism and/or pre­ and the dorsal one darkens (Fig­ The duration and number of Egg masses were collected dation (Rodríguez et al., 2004). ure 1c). Fully developed cater­ specimens at each instar were from each cage and placed in However, non-mated H. paulex pillars were 4.01 ±0.27cm long, obtained by measuring the head Petri dishes (9.0×1.5cm). The capsule of H. paulex with a number of eggs, incubation pe­ stereoscopic microscope and riod, viability of eggs, pre-ovi­ Table I micrometric ocular microscope. position period, number of eggs Duration of each instar, width of the head The duration of the pre-pupa per egg mass, and longevity of capsule and growth rate of Hylesia paulex and pupa periods, the weight of males and females were record­ REARED with Eucalyptus urophylla leaves, pupae and the sex rate of this ed. Female specimens were dis­ in the laboratory at 25 ±2ºC, 60 ±10% relative insect were also recorded. sected to determine the number humidity and a 12h photoperiod Thirty pairs of H. paulex of ova kept in the abdomen. Stage Duration (days) * Width of the head Growth rate were placed immediately after its A second experiment was capsule (mm) * emergency in five 30×30×30cm carried out to test the hypoth­ screened cages with wood bot­ esis that the gregarious behav­ I 12.00 ±0.00 0.60 ±0.00 — tom and glass cover under field ior is important for survival of II 10.10 ±0.05 0.80 ±0.01 1.33 conditions, to study the longev­ H. paulex in immature stages. III 5.90 ±0.05 1.11 ±0.00 1.39 ity and some reproductive pa­ Third instar caterpillars per IV 8.77 ±0.20 1.47 ±0.01 1.32 rameters of this insect. Branches treatment were maintained in V 7.37 ±0.21 1.95 ±0.01 1.33 of E. urophylla were supplied plastic pots (500ml) with densi­ VI 9.57 ±0.30 2.47 ±0.02 1.27 VII 14.10 ±0.73 3.52 ±0.03 1.42 in the interior of these cages ties of: one (treatment T1), five wrapped in cotton wads soaked (T2), 10 (T3), 20 (T4) or 30 * Average ±standard error.

SEP 2009, VOL. 34 Nº 9 647 head of a brilliant Benrey, 1997; Reader can be explained by the accen­ red color, ash to and Hochuli, 2003). tuated sexual dimorphism of this black tegument, Where they stayed species. The pupa remained pro­ pronounced clear during the day, H. tected, externally, by the same yellow tufts of paulex from the late protection structure as in the thorns and small instars wove a shel­ pre-pupa stage. The sex was black stripes in ter in the form of a differentiated in pupae examin­ the lateral parts sack with a mixture ing with a magnifying glass (Figure 1d). The of silk threads and (10×) the ventral part of the last caterpillars of H. remnants of branch­ abdominal segment. paulex presented a es, leaves and feces, The adults of H. paulex strong aggregation which may represent a emerged from the larger width with few individu­ kind of defensive be­ of the pupa, with a sex rate of als dispersed from havior of this genus 0.59. The wings were usually the group during (Santos et al., 1996; fully extended after a few hours the first instars. Specht et al., 2006a). from emergence. Some adults From the second At night, they aban­ stayed immobile while others to sixth instars doned the shelter to walked slowly to the bottom of they moved in feed. The caterpillars the cages, in a similar behavior single lines, dur­ of H. paulex present­ (Manley, 1993) as in Automeris Figure 2. Accumulated mortality (%) of Hylesia paulex reared ing the night, in a with Eucalyptus urophylla leaves from the third instar in 500ml ed a strong perception io (Fabricius) (Lepidoptera: Sat­ similar pattern as pots at different densities: one (T1), five (T2), 10 (T3), 20 (T4), or to sound and physi­ urniidae). Adults that did not that reported for 30 (T5) caterpillars, kept at 25 ±2oC, 60 ±10% relative humidity cal contacts (Hogue, spread their wings after 24h did H. lineata (Druce) and 12h photoperiod. 1972). When upset, not mate. The egg masses of by Fitzgerald and they regurgitated a H. paulex were laid preferen­ Pescador-Rubio dark liquid, arched the tially in the superior part of the (2002), for Dirphia avicula infection when 30 of them were posterior part of the body and cages, without a defined sched­ (Walker) (Lepidoptera: Saturnii­ reared per pot (T5), mainly in they contorted wildly to defend ule, while mating always took dae) by Zanuncio et al. (1994), the last two instars, as also de­ themselves. The thorns at the place at night. The eucalyptus and for Euselasia eucerus (Lepi­ scribed for H. metabus (Cramer) back of H. paulex caterpillars leaves inside the cages were not doptera: Riodinidae) by Zanun­ (Lepidoptera: Saturniidae) by have bunches, in pairs (Figure used as substratum for oviposi­ cio et al. (2008). Osborn et al. (2002). This may 1e), but without erucism, differ­ tion, indicating that females of In the second experiment have occurred due to horizontal ing from other Saturniidae as H. paulex prefer to lay eggs (Figure 2), the third instar transmission of pathogens, a H. metabus (Rodríguez-Morales on the trunk and branches of H. paulex caterpillars placed negative characteristic of the et al., 2005), Lonomia obliqua E. urophylla, as it is frequently individually in 500ml pots gregarious behavior in insects Walker (Lorini and Corseuil, observed in the field. The pre- (treatment T1) did not reach (Cocroft, 2001). Caterpillars 2001) and Automeris illustris oviposition period of this insect the fourth instar, while none without infection completed sat­ Walker (Specht et al., 2006b). was 2.1 days, with 132 eggs per of those in treatment T2 (five isfactorily the whole life cycle. This characteristic facilitated the egg mass and 112 ova kept in caterpillars per pot) reached the The weight of pupae of each sex manipulation of H. paulex in the the abdomen per female. The pupa stage. The accumulated (Table II) did not differ between laboratory and can be useful for individualization of H. paulex mortality of caterpillars was the treatments T3, T4 and T5 manual control of this species pairs in the cages might have lower in T3 and T4 (10 and 20 (F= 0.54; P<0.05), suggesting in the field. propitiated a lower fertilization caterpillars per pot) than in the that the survival of H. paulex is The pre-pupa stage of H. and, consequently, larger reten­ T5 (30 per pot) in the maximum more affected by its gregarious paulex lasted 3.87 ±0.16 days, tion of ova in the abdomen. The age of 50 days.The intraspecific behavior during its development with a considerable reduction retention of ova in the abdomen competition for food in this last than by nutrition conditions. of the corporal length, feeding was also found in other spe­ treatment may be higher due The hypothesis that gregarious arrest and search for a place to cies of this family, and it can to the limited space. Besides, behavior benefits H. paulex was build a cocoon, mainly with silk be related to the polyandry of most caterpillars of H. paulex accepted, as found for other threads, remnants of branches, females (Gardiner, 1967; Wolfe, presented symptoms of bacterial Lepidoptera species (Denno and leaves and feces. Several indi­ 1998) or to the conditions of the viduals of H. paulex pu­ present study. Table II pate together in a single The longevity of H. paulex Weight* of pupae from different numbers substratum mass, also males was shorter (2.80 ±0.44 of caterpillars of Hylesia paulex reared with suggesting aggregation in days) than that of females (4.50 Eucalyptus urophylla LEAV E S this stage as shown for ±0.54 days). Males and females the Hemileucinae H. acu- did not feed during the adult Sex Treatments ta Druce (Wolfe, 1998). stage and, for this reason, the T1 T2 T3 T4 T5 The pupa stage of H. higher longevity of females can Male — — 294.34 ±15.15 aA 288.18 ±10.31 aA 286.73 ±9.48 aA paulex (Figure 1f) lasted be related to the need of finding Female — — 392.29 ±19.42 aB 411.83 ±11.25 aB 397.48 ±22.53 aB 21.67 ±0.79 days, with partners for mating. Adults of females being heavier and H. paulex present different char­ * Weight in mg; average ±standard error. Means followed by a common lower case longer (385.75 ±12.69mg acteristics among sexes, such as letter (lines) or capital letter (columns) do not differ (Tukey test; P=0.05). T1: one caterpillar, T2: two caterpillars, T3: 10 caterpillars, T4: 20 caterpillars, and and 1.90 ±0.89cm) than the size of the abdomen, span T5: 30 caterpillars per 500ml pot, kept at 25 ±2oC, 60 ±10% relative humidity and a males (248.17 ±7.32mg and color of the wings, and an­ 12h photoperiod. and 1.30 ±0.78cm), which tennae type, as in other Saturnii­

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Zanuncio JC, Gudes RNC, Zanuncio edge about the biological and Osborn F, Berlioz L, Vitelli-Flores J, Gardiner BOC (1967) The life his­ TV, Fabres AS (2001) Species morphological aspects of H. Monsalve W, Dorta B, Lemoine tory of Periphoba hircia (Sat­ richness and abundance of de­ paulex are also useful for pub­ VR (2002) Pathogenic effects urniidae) with a note on distri­ foliating Lepidoptera associ­ of bacteria isolated from larvae lic health inspectors to develop bution and larval variation. J. of ated with Eucalyptus grandis of Hylesia metabus Crammer measures to avoid dermatologi­ Lepidopt. Soc. 21: 198-204. in Brazil and their response (Lepidoptera: Saturniidae). J. to plant age. Austral Ecol. 26: cal damages by Hylesia moths Glasser CM, Cardoso JL, Carré­ Invert. 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HB, Dietrich CP, Gremskia morphology of abdominal se­ Bras. Ciênc. 81: 61-66. and Fundação de Amparo à W, Veiga SS (2005) Identifica­ tae from male and female of Zanuncio TV, Zanuncio JC, Meira Pesquisa do Estado de Minas tion and partial characterisation Hylesia metabus adults (Lepi­ IA, Araújo FS (1994) Caracte­ of hyaluronidases in Lonomia Gerais (FAPEMIG), Brazil. doptera: Saturniidae) and their rização das fases larval e adulta obliqua venom. Toxicon 45: function. Fla. Entomol. 87: de Dirphia avicula (Lepidopte­ 403-410. REFERENCES 30-36. ra: Saturniidae) em folhas de Hogue CL (1972) Protective func­ Rodríguez-Morales AJ, Arria M, Eucalyptus urophylla. Árvore. tion of sound perception and Rojas-Mirabal J, Borges E, 18: 153-158. Ande AT (2003) The lipid profile of gregariousness in Hylesia larvae the pallid emperor moth Cirina Benítez JA, Herrera M, Villa­ Zanuncio TV, Zanuncio JC, Miranda (Saturniidae: Hemileucinae). 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