Morphological notes on Mamestra brassicae (Lepidoptera Noctuidae)

Sannino ed Espinosa

Morphological notes on Mamestra brassicae 13

Morphological notes on Mamestra brassicae (Lepidoptera Noctuidae) Luigi Sannino, Bruno Espinosa Summary The morphology of the Mamestra brassicae (Linnaeus, 1758) was studied, by means of laboratory and field observations, at the Tobacco Research Institute in Scafati (Salerno, Southern Italy). Adult has the forewing blackish-gray with the reniform stigma conspicuous, encircled by white points. Typical is the big black slightly curved and pointed spine present on the foretibiae. Egg is subspherical with a flat base and chorion sculpturing composed of 34-40 longitudinal ribs of which 14-19 reach the micropylar area. Newly-born larva is white-pale yellow with bright black head. Anal shield with some peculiar characteristics: it is sclerotized along margins and in correspondence of the D1 setae. Mature larva is quite variable in background colour, ranging from pale green to olive green-black and greyish with pink shades. On the back, except for the green forms, couples of linear, dark, subdorsal spots stand out. In the ocellar position a characteristic subcircular, yellow spot is evident. Pupa is pale brown lucent when newly formed and purplish-brown just before adult emergence. Cremaster is prominent, finely rugose, bearing two long, straight, subparallel spines, ending as a pinhead.

Luigi Sannino, Sezione di Biologia e Difesa dell’Istituto Sperimentale per il Tabacco, Scafati. Bruno Espinosa, Dipartimento di Entomologia e Zoologia Agraria dell’Università di Napoli “Federico II”, Portici

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Riassunto. Note morfologiche sulla Mamestra brassicae (Lepidoptera Noctuidae) Allo scopo di facilitare il riconoscimento della Mamestra brassicae (Linnaeus, 1758) nei vari stadi del ciclo, è stato intrapreso uno studio realizzato, mediante osservazioni di campo e di laboratorio, presso l’Istituto Sperimentale per il Tabacco di Scafati (SA). L’adulto ha le ali anteriori grigio-brunastre su cui risalta la macchia reniforme cerchiata da punti bianchi. Tipico è il grosso uncino nero presente all’apice delle tibie anteriori. L’uovo, subsferico, a base piana (larghezza mm 0,60-0,65; altezza mm 0,35-0,40), presenta il corion con 34-40 coste verticali, di cui 14-19 raggiungono l’area micropilare, intersecate da 15-20 strie trasversali sottili ed irregolari. Biancopaglierino appena deposto, appare cinerino a fine incubazione con due macchie grigio scuro nella regione polare. La larva neonata è di colore paglierino chiaro traslucido con la capsula cefalica nero lucido. Sul meso- e metatorace risaltano i pinnacoli neri delle setole D2 perché molto allungati longitudinalmente. La placca sopranale è di aspetto particolare, sclerificata solo lungo il bordo e in corrispondenza delle setole D1; il suddetto margine, più o meno ispessito, si presenta sempre interrotto posteriormente (tra le setole D2) ed a volte anche presso gli angoli laterali. La larva matura ha il colore di fondo molto variabile, dal verde pisello al bruno olivaceo fino al grigiastro con sfumature rosacee. Sul dorso si evidenziano, tranne che negli individui verdi, tipiche coppie di macchie lineari scure che sul VII e VIII urite diventano più marcate e di forma subtriangolare. In particolare sull’VIII urite queste si fondono posteriormente formando un caratteristico disegno a ferro di cavallo squadrato. Il capo è verdastro negli individui verdi, castano nelle altre forme con reticolatura più scura. Carattere diagnostico di rilievo è la macchia subcircolare giallo carico in posizione ocellare.

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La crisalide è castana lucente appena formata, marrone purpurea a fine ninfosi. Punteggiatura profonda e densa sulla fascia basale degli uriti VVII. Cremaster prominente, finemente rugoso, con due spine lunghe, diritte, subparallele, capitate.

Key words: Mamestra brassicae (L), Morfology, Tobacco

Introduction The cabbage moth, Mamestra brassicae (Linnaeus, 1758), is one of our common species. Larvae are frequent throughout the autumn on wild and cultivated plants, especially cabbages and other Brassica spp. A great many researchers have studied the insect biology from different viewpoints, particularly in Europe (for a list of references see Kurir, 1978; Sannino & Espinosa, 1998). As far as the morphology is concerned, although there exists the work of Melis (1936, sub Polia pisi L.), still some particulars need to be better known. The purpose of the present work is to allow a rapid recognition of the species in its different stages and deepen some aspects not sufficiently described yet.

Material and methods Observations were conducted at the Tobacco Research Institute in Scafati (province of Salerno, Southern Italy), on a M. brassicae stock culture, initially composed of the larvae collected in the field and larvae from eggs laid by adults captured alive by means of a light trap installed in the abovementioned Institute. Eggs were placed in petri dishes (diameter 10 cm) on a semi-synthetic diet according to Sannino et al. (1995), and followed until the hatching. The newly hatched larvae were kept on this diet for the first 24 hours, in order to reduce the mortality rate, then were transferred to plastic boxes (50 x 15 x 20 cm) with a soil layer at the bottom and covered with gauze. Tobacco leaves, mixed with cabbage, Plantago lanceolata L. and Sonchus oleraceus L., served as food for larvae. The newly formed pupae

were in part removed, to record the morphological characteristics and partly left in the respective boxes until adults emerged. The latter were caged, in promiscuous groups of about ten individuals, on tobacco plants and fed ad libitum with a 10 % honey solution until death. Observations on the external morphology of the different stages and relative chromatic changes were made using the following samples: 70 adults captured in different years; 50 eggs collected immediately after laying (from different females) and observed every 12 hours, till hatching; 60 larvae singly reared until maturity phase; 30 pupae removed from the insect-rearing boxes (in different periods of the year) immediately after formation and followed until adult emergence. All observations were conducted on living specimens obtained both from insectaries and from field collecting. When necessary, some specimens were sectioned and mounted on slide. For the description of the single larval instars, 30 larvae per each of the two generations, collected just after hatching, were placed individually in plastic boxes (4 x 4 cm, commercially available from Color-Dec Italy under the name “rearing tray”), at the bottom of which the above-mentioned semi-synthetic diet or tobacco leaves were placed. Every 24 h, each larva was transferred to a new box and the one used the previous day examined to the searching of the possible exuvia left by the newmoulted larva and in particular the head capsule, the finding of the which was considered the probate sign of the occurred moult. Observations lasted until pupation of all larvae. Sexing of the pupae was made according to well-known external sexual characters present on

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Morphological notes on Mamestra brassicae 15

Fig. I. - Mamestra brassicae (Linnaeus) – 1. Egg, dorsal view. 2. SEM picture of the micropylar rosette. Fig. I. Mamestra brassicae (Linnaeus) – 1. uovo, visto dall’alto. 2. Rosetta micropilare (foto al SEM). the ventral side of the last abdominal segments (Sannino et al., 1987), while adult sexing was done by examination of the frenulum or antennal conformation. All measurements given, expressed in millimetres, refer to the greatest distance along each axis. Some measurements are given as the average ± SE (standard error). The markings on the body of the larva are presented according to the terminology used by Sannino et al. (1995), the setae according to the nomenclature suggested by Hinton (1946). The description of the surface structure of the egg (sculpturing) is based upon the nomenclature proposed by Döring (1955). Part of the examined specimens was prepared adults pinned and dried, immature stages in ethyl alcohol (80 %) or in KAAD solution slightly modified (see Sannino et al., 1995) - and deposited in the Experimental Tobacco Institute collection.

Description of the stages Adult Forewing blackish-gray. Reniform stigma conspicuous, encircled by white points (usually denser on the concave margin); orbicular stigma rounded and claviform stigma short, both with blackish margins. Postmedian line angleous and split near the costal margin. Subterminal line white split at the anterior half. Lateral margin of the wing delimited by the subterminal white line and by a series of little black spots. Hindwing light brownish-pale black, darker along the (latero-posterior) outer margin. Thorax with a slight dorsal crest. Antennae simple, filiform in both sexes, finely ciliate in the male. Typical of the species (and of the genus) is the big black slightly curved and pointed spine present at the apex of the foretibiae. Frenulum and retinaculum typical of the Noctuidae: the former consists of a single stout bristle in the male and

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Fig. II. - Mamestra brassicae (Linnaeus), newly-born larva – 1.Thoracic segments. 2. Ab1, dorsal view. 3. Ab5, lateral view. 4. Two different patterns of the prothoracic shield, dorsal view. Fig. II. - Mamestra brassicae (Linnaeus), larva neonata– 1.Segmenti toracici. 2. Ab1, visto dall’alto. 3. Ab5, visto di profilo. 4. Differenti conformazioni della placca protoracica, viste dall’alto.

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three finer bristles in the female, the latter is composed of a strong membranous hook in the male and a tuft of specialized scales in the female. Wingspan 38-49 mm (mean 43 ± 0.2; No.=70), body length 15-22 mm (mean 18 ± 0.1). Females are on average slightly bigger than males. A detailed description of the imago is provided by Melis (1936). The male and female genitalia are described in Pierce (1967a, 1967b) and Berio (1985).

Morphological notes on Mamestra brassicae 17

Egg It belongs to the upright type. Subspherical, slightly pressed and with a flat base, measuring 0.60-0.65 mm in width and 0.35-0.40 mm in height. Chorion surface sculptured with 34-40 longitudinal ribs – fairly standing out and with the superior apices non prominent, of which 14-19 reach the micropylar area – intersected by 15-20 finer, irregular, transverse striae (Fig. I, 1).

Fig. III. - Mamestra brassicae (Linnaeus), larva of second instar – 1. Ab1, dorsal view. 2. Ab5, lateral view. 3. Prothoracic shield. 4. Anal shield. Fig. III - Mamestra brassicae (Linnaeus), larva di seconda età – 1. Ab1, visto dall’alto. 2. Ab5, visto di profilo. 3. Placca protoracica. 4. Placca sopranale.

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Fig. IV. - Mamestra brassicae (Linnaeus), mature larva – 1. Head capsule, frontal view. 2. Ocellar area. 3. Right mandible, oral surface. (T, inner tooth). Fig. IV. - Mamestra brassicae (Linnaeus), larva matura – 1. Capo, visto frontalmente. 2. Area ocellare. 3. Mandibola destra, vista dal lato interno. (T, dente interno). Micropylar rosette slightly exceeds in height the level of the superior apices of the longitudinal ribs. It is composed of 11-14 elliptical cells, a little elongated radially and of various sizes (Fig. I, 2). White-pale yellow when laid, turning paleyellow, showing an irregular polar spot and an above-equatorial ring line of rust colour as the embryo develops. At incubation end it becomes of an uniform ash colour with two dark grey spots in the polar area, corresponding to the head and prothoracic shield of the larva. Infertile eggs slowly

go backwards, remaining more or less of the initial colour. On the egg stage, Döring (1955) provides some information. Newly-born larva Body white-pale yellow, translucent, from which the inner reddish-yellow organs stand out; in onetwo days body coloration turns yellowish and opaque. Head bright black, with stemmata pale and mandibles yellowish, darker at the tip. Pinacula

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black rounded with setae concolorous. On the mesoand metathorax, pinacula of D2 setae stand out because very longitudinally elongated (Fig. II, 1). Pinacula of the first 4-5 abdominal segments very big with respect to the following (Fig. II, 2). Spiracles present, as usual, on prothorax and on abdominal segments 1-8 (A1-8), subcircular and similar in size except for those on prothorax and A8, distinctively bigger then the others. Prothoracic shield subtrapezoidal (Fig II, 1), slightly paler than the head, with setae arranged as

Morphological notes on Mamestra brassicae 19

in the mature larva (see later). Anal shield is peculiarly sclerotized only along the margin and in correspondence of the D1 setae; the aforesaid margin, more or less thickened, is always posteriorly interrupted (between the D2 setae) and sometimes also near the lateral angles. Setae arranged as in the mature larva (Fig. II, 4). Prolegs, present on A3-6 and A10, have the same background colour; the first two pairs (A3-4) are less developed than the others and show some dark areas on the external side (Fig. II, 3). Legs

Fig. V. - Mamestra brassicae (Linnaeus), mature larva – 1. Ab5, dorsal view. 2. Ab5, lateral view. 3. Ab8, dorsal view. (S, spiracle). Fig. V. - Mamestra brassicae (Linnaeus), larva matura – 1. Ab5, visto dall’alto. 2. Ab5, visto di lato. 3. Ab8, visto dall’alto. (S, stigma).

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Fig. VI. - Mamestra brassicae (Linnaeus), pupa – 1. Dorsal sculpture on Ab5. 2. Ventral view. 3. Abdominal spiracle. 4. Cremaster. (A, antenna; MX, maxilla; L1, prothoracic leg; L2, mesothoracic leg; L3, metathoracic leg; FW, forewing or mesothoracic wing). Fig. VI. - Mamestra brassicae (Linnaeus), crisalide – 1. Punteggiatura dorsale su Ab5. 2. Visione ventrale. 3. Stigma addominale. 4. Cremaster. (A, antenna; MX, maxilloteca; L1, podoteca anteriore; L2, podoteca mediana; L3, podoteca posteriore; FW, pteroteca anteriore).

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greyish, translucent. 1 Body length 1.7-1.8 mm (mean 1.74 ± 0.03) ; head capsule width 0.32-0.35 mm. Further features are described and illustrated in Melis (1936). Larva of second instar Body background colour greenish-yellow, slightly greyish on the back where the median and subdorsal lines are visible because paler. Pinacula black surrounded by little subcircular dark areas, bearing black setae (Fig. III, 1). For the first time there appear the L3 seta (above the base of the prolegs) and the SV2 seta (on the plant of the prolegs) (Fig. III, 2). Head yellow-pale with tonofibrillary spots darker and points of the setae inserting dark or black; stemmata black, mandibles brown. Prothoracic shield of the same background colour with the point of the setae inserting black (Fig. III, 3). Anal shield concolorous with the ground; margins little evident and points of the setae inserting dark grey (Fig. III, 4). Legs greyish, translucent. Prolegs with the dark spot on the external side paler than the previous instar; prolegs on A3-4 still less developed than the others. 1 Body length 4-5 mm (mean 4.6 ± 0.02) ; head capsule width 0.40-0.45 mm. Larva of third instar On the ivory or greenish-white background colour, the median dorsal line, the subdorsal lines, and the lowerspiracular line yellowish-white are visible. Pinacula dark or black less evident than the second instar. Head capsule pale translucent on which stand out the black stemmata. Mandibles with the distal part darker. Spiracles subcircular, white with black peritreme. Prothoracic shield pale yellow with margins more defined. Legs pale translucent. Prolegs concolorous with background. 1 Body length 7-10 mm (mean 8.9 ± 0.4) ; head capsule width 0.7-0.8 mm. Larva of fourth instar Background colour greenish-grey, paler on the ventral side. Are distinctively visible: a) the median dorsal line, thin, whitish, fragmented and slightly black margined; b) the subdorsal lines and the lowerspiracular band (which often incorporates also 1

Morphological notes on Mamestra brassicae 21

spiracles) whitish and with clear margins; c) the upper-spiracular fascia slightly darker than the ground coloration. Pinacula inconspicuous, with black papilla. Setae pale. Spiracles white with black peritreme. Head, shields, legs and prolegs characteristics do not differ from those of the third instar. 1 Body length 11.5–14.3 mm (mean 13.4 ± 0.2) ; head capsule width 1.4-1.5 mm. Larva of fifth instar It shows chromatic characteristics similar to those described for the fourth instar, except for the head which is yellowish-pale brown with brown maculation. In this and in the previous instars, the colours and patterns always lose intensity prior each moult. 1 Body length 15.6-20.0 mm (mean 18.5 ± 0.3) ; head capsule width 2.1-2.3 mm. Larva of sixth instar Body integument smooth. Variation in ground colour is considerable, from pale green to olive green-black and greyish with pink shades. Head hypognathous, greenish in the green forms. In the other forms, it is pale brown with darker reticulation concentrated mainly at the upper half and in correspondence of the P2 setae (Fig. IV, 1). There are also two linear brown spots along the adfrontal sutures. In the ocellar position, a characteristic subcircular, yellow spot is quite evident. In addition, two brown or black little spots are present between the antennae and the mandibles. These are pale brown, distally darkened, with an evident tooth on the inner side (Fig. IV, 3). On the back, except for the green forms, couples of linear, dark, subdorsal spots stand out (Fig. V, 1, 2). The ones present on A7-8 are more marked, subtriangular in shape and usually on the A8 are joined posteriorly by a dorsal transverse band forming a characteristic horseshoe drawing (Fig. V, 3). Even in another Noctuidae species, the Paranoctua comes (Hübner), [1813], a similar pattern is present, but in this case the body background colour ranges between greyishochreous and blackish-grey. Lowerspiracular fascia, yellow in the pale individuals, tends to yellow

Measures of the body length and head capsule width were taken at the beginning of each instar. The number of observed individuals for each instar was 40.

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shaded with orange-pink in the dark forms. Ventral side, prolegs included, green or yellowish-green, and in any case always paler than the dorsal side. Pinacula inconspicuous, are concolorous with the body integument and dark papilla. Spiracles elliptical, white with black peritreme, usually incorporated in black spots. Those on the prothorax and A8 are distinctively bigger than the remaining. Prothoracic shield subtrapezoidal and anal shield subtriangular, both a little darker than the ground coloration and with the typical chaetotaxy of the Noctuidae. Legs greenish in green forms and pale brown in the dark ones. Prolegs concolorous with the ventral areas (Fig. V, 2). Crochets arranged in an uniordinal mesoseries, in number of 22-25, 23-27, 25-28, 29-31 and 3034, on A3-6 and A10 prolegs, respectively. At initial instar, larva is 23.4 ± 0.3 mm (range 21.1-25.3; No.=40) long and 2.6 ± 0.09 mm (r. 2.52.7; No.=40) wide; at full-growth, length and width of the body, and width of the head capsule are 39.8 ± 0.04 mm (r. 37.1-41.8; No.=40), 5.8 ± 0.2 mm (r. 5.3-6.2; No.=40) and 3.1 ± 0.1 mm (r. 3.0-3.3; No.=40), respectively. A description of the mature larva was made by

Melis (1936), Beck (1960), Merzeewskaja (1967). Pupa Pale brown lucent when newly formed, turning to purplish-brown just before adult emergence. Cuticle glossy, slightly rugose on head, thorax and appendages. Sculpture – constituted of small pits – weak on the first eight abdominal segments (except for the apical margins), deep and dense on the cephalic margins of the A5-7 (Fig. VI, 1). Maxillae and meta-thoracic legs reach the caudal margin of the wing lobes; the meso-thoracic legs and the antennae, of equal length, reach two thirds of the distance elapsing between the apices of the prothoracic legs and the maxillae (Fig. VI, 2). Spiracles elliptical, very narrow (ratio length/ width about 4/1) not elevated on the cuticle (Fig. VI, 3). Cremaster prominent, finely rugose, bearing two long, straight, subparallel spines, ending as a pinhead (Fig. VI, 4). Genital and anal openings are typical of the Noctuidae. Body length 19.5 ± 0.2 mm (range 17.0-21.6 No.=30), body width (across the thorax) 6.4 ± 0.1 mm (r. 5.8-7.2; N0.=30). For additional details on the pupal characteristics cf. Khotko (1968), Sannino et al. (1992), Patocka (1996).

Table 1. – Results of measurements (mm) of the body length and head capsule width of Mamestra brassicae (L.) larva. Case of 5 instars. Tabella 1. – Lunghezza del corpo e largnezza della capsula cefalica (mm) della larva di Mamestra brassicae (L). Caso di 5 età. No.

Body length

Head capsule width

Larval instar observations

mean ± SE

1st instar

15

1.74 ± 0.04

2nd instar

mean ± SE

range

1.7 - 1.8

0.35 ± 0.02

0.32 - 0.37

12

4.30 ± 0.05

3.9 - 4.5

0.50 ± 0.02

0.45 - 0.53

3 instar

13

7.8 ± 0.4

7.1 - 9.0

0.74 ± 0.02

0.69 - 0.81

4th instar

15

13.3 ± 0.5

11.7 - 14.2

1.45 ± 0.03

1.40 - 1.52

5th instar

16

22.5 ± 0.6

17.8 - 24.2

2.33 ± 0.01

2.0

Mature larva

12

39.0 ± 0.3

37.3 - 40.5

3.10 ± 0.02

2.95 - 3.20

rd

7

range

th

- 2.50

Measurements were taken at the beginning of each instar except 5 where they were taken at the end (mature larva) as well

Morphological notes on Mamestra brassicae 23

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Table 2. – Results of measurements (mm) of the body length and head capsule width of Mamestra brassicae (L.) larva. Case of 7 instars. Tabella 2. – Lunghezza del corpo e largnezza della capsula cefalica (mm) della larva di Mamestra brassicae (L). Caso di 7 età. No.

Body length

Head capsule width

Larval instar observations

mean ± SE

st

mean ± SE

range

20

1.74 ± 0.04

nd

1.7 - 1.8

0.35 ± 0.03

0.32 - 0.36

2 instar

15

4.30 ± 0.05

3.9 - 4.5

0.50 ± 0.02

0.45 - 0.53

3rd instar

14

7.8 ± 0.4

7.1 - 8.6

0.75 ± 0.03

0.70 - 0.80

4th instar

12

12.6 ± 0.4

11.9 - 13.0

1.45 ± 0.04

1.39 - 1.53

5th instar

16

17.5 ± 0.3

16.2 - 18.5

2.20 ± 0.02

2.10 - 2.30

6th instar

12

21.5 ± 0.4

19.4 - 22.2

2.80 ± 0.02

2.72 - 2.90

7 instar

10

26.7 ± 0.3

24.9 - 28.2

3.00 ± 0.01

2.90 - 3.10

Mature larva

9

40.5 ± 0.2

38.8 - 41.5

3.35 ± 0.01

3.30 - 3.40

1 instar

th

range

th

Measurements were taken at the beginning of each instar except 7 where they were taken at the end (mature larva) as well

Comments The adult chromatic variability observed in M. brassicae, was very low with respect to many other noctuid species, although several forms (six, according to Kurir, 1978) were so far recorded (Tutt, 1891, 1892; Bergmann, 1954). In the newly-born larva stage, a peculiar character is represented by the margin of the anal shield, which is sclerotized along the margin and in correspondence of the D1 setae. In the mature larva characteristic is the yellow subcircular spot present in the ocellar position. Concerning the considerable ground chromatic variation observed in this instar, no relation with the rearing conditions was noticed. In fact, neither the colour of food nor the high population density have influenced the last instar coloration. Majerus (1983a, 1983b) analysed the causes and effects of the complex larval polymorphism in Phlogophora meticulosa (L.) (Lepidoptera Noctuidae). According to the author, the high larval colour variability present in this species is exclusively induced by hereditary factors in the last instars,

while it is controlled by the colour of the food ingested in the first, second and part of the third instar. In our experiment the number of larval instars ranged between five and seven being six the far most frequent. The measurements of the body length and head capsule width of M. brassicae larvae, have been reported in the course of the text, while the ones related to the less common cases of five and seven instars, are schematically represented in tables 1 and 2.

References Beck H. - 1960 - Die Larvalsystematik der Eulen (Noctuidae) - Akademie Verlag, Berlin: 406 pp. Bergmann A. - 1954 - Die Grosschmetterlinge Mitteldeutschlands. 4. Eulen. - Urania-Verlag, Jena: 1060 pp. Berio E. - 1985 - Fauna d’Italia 22. Lepidoptera Noctuidae. I. - Ed. Calderini, Bologna: 970 pp. Döring E. - 1955 - Zur morphologie der schmetterlingseier Akademie-Verlag, Berlin: 154 pp. Hinton H. E. - 1946 - On the homology and nomenclature of the setae of lepidopterous larvae, with some notes on

24 the phylogeny of the Lepidoptera. - Trans. R. Ent. Soc. London 97: 1-37. Khotko E. I. - 1968 - A key to the Noctuid pupae. - Minsk: 126 pp. Kurir A. - 1978 - Noctuidae, Eulen. In: Schwenke W., “Die Forstschädlinge Europas” 3: 266-313. - Parey, Hamburg. Majerus M. E. N. - 1983a - Larval colour variation in Phlogophora meticulosa (L.) (Lep.: Noctuidae). Part I: the variation, and its control in instars 1-3. Proc.Trans. Br. Ent. Nat. Hist. Soc. 16: 34-49. Majerus M. E. N. - 1983b - Larval colour variation in Phlogophora meticulosa (L.). Part II: genetic control in instars 3-5. - Proc. Trans. Br. Ent. Nat. Hist. Soc. 16: 63-76. Melis A. – 1936 - Contributo alla conoscenza morfo-biologica della Polia (Mamestra) pisi L. - Redia 22: 1-31. Merzeewskaja O. I. - 1967 - The larvae of Noctuidae, their biology and morfology (with keys). - Academy of Science, Byelorussia, Sci. Tech., Minsk: 452 pp. Patocka J. – 1996 – Die puppen der mitteleuropäischen eulen: Cuculliinae, Heliothinae, Ipimorphinae, Hadeninae, Agrotinae. (Lepidoptera: Noctuidae). – Nova Supplementa Entomol., Berlin, 8 (1996): 1-232. Pierce F .N. – 1967a – The genitalia of the group Noctuidae of the Lepidoptera of the British Islands. An account of the morphology of the male clasping organs. – E. W.

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Classey, Feltham: 88 pp. Pierce F. N. – 1967b – The genitalia of the group Noctuidae of the Lepidoptera of the British Islands. An account of the morphology of the female reproductory organs. – E. W. Classey, Hampton: 66 pp. Sannino L., Balbiani A. & Espinosa B. - 1987 - Osservazioni morfo-biologiche su alcune specie del genere Spodoptera (Lepidoptera: Noctuidae) e rapporti di parassitismo con la coltura del tabacco in Italia. - Inf. Fitop. 11: 29-40. Sannino L., Balbiani A. & Espinosa B. – 1992 - Morfologia comparativa negli stadi preimmaginali di lepidotteri del tabacco. 1. Bozzoli e crisalidi. (Parte seconda: Crisalidi). – Mem. Soc. ent. ital., Genova, 70 (2) (1991): 251-266. Sannino L., Espinosa B. & Balbiani A. - 1995 - Morfologia comparativa negli stadi preimmaginali di lepidotteri del tabacco. 2. Larve. - Boll. Lab. Ent. agr. “Filippo Silvestri” 50 (1993): 23-91. Sannino L. & Espinosa B. – 1998 – Ciclo biologico di Mamestra brassicae e danni alle colture ortive in Campania. – Inf. Fitop. 5: 59-67. Tutt J. W. - 1891 - The British Noctuae and their varieties. Vol. I - Swan, Sonneschein & Co., London: 164 pp. Tutt J. W. - 1892 - The British Noctuae and their varieties. Vol. IV - Swan, Sonneschein & Co., London: 140 pp.