Banasa maculata, Campos & Garbelotto, 2010

Campos, Luiz Alexandre, Grazia, Jocelia, Garbelotto, Thereza De Almeida, Bianchi, Filipe Michels & Lanzarini, Nara Coral, 2010, A new South American species of Banasa Stål (Hemiptera: Heteroptera: Pentatomidae: Pentatominae): from egg to adult, Zootaxa 2559, pp. 47-57 : 48-56

Banasa maculata sp. nov. Campos & Garbelotto

( Figs 1–13 )

Etymology. Specific epithet referring to the irregular macula on anterior half of pronotum.

Type locality. Brazil, Criciúma (-28.685, -49.360).

Holotype male ( Figs 1, 2 ), BRAZIL: Santa Catarina: Criciúma, Morro do Céu , 10.v.2007, Remus, G. [- 28.685, -49.360] (MZSP).

Paratypes. BRAZIL: Santa Catarina: Criciúma, Parque José Milanese , 1 male, 15.iii.2007, Frassetto, J. [- 28.690, -49.431] ( UFRG) ; 4 females and 1 male, S.A.T.C., 18.x.2007, Bianchi, F.M. [-28.707, -49.409] ( MZSP) ; 2 females, same data as previous ( UFRS) ; 6 females and 5 males, laboratory reared, 03.xii.2007, Bianchi, F.M. ( MCNZ) ; 3 females, laboratory reared, 05.xii.2007, Bianchi, F.M. ( MZSP) ; 2 females and 4 males, laboratory reared, 10.xii.2007, Bianchi, F.M. ( MZSP) ; 4 females, laboratory reared, 14.xii.2007, Bianchi, F.M. ( UFRS) ; 1 female, laboratory reared, 17.xii.2007, Bianchi, F.M. ( MCNZ) ; 1 female, laboratory reared, 18.xii.2007, Bianchi, F.M. ( MCNZ) ; 1 female, laboratory reared, 19.xii.2007, Bianchi, F.M. ( MCNZ) ; 2 males, laboratory reared, 03.i.2008, Bianchi, F.M. ( UFRS) ; 7 females and 5 males, laboratory reared, 09.i.2008, Bianchi, F.M. ( UFRS) ; 1 female, laboratory reared, 11.i.2008, Bianchi, F.M. ( MCNZ) ; 1 female, laboratory reared, 14.i.2008, Bianchi, F.M. ( MCNZ) ; 1 male, laboratory reared, 18.i.2008, Bianchi, F.M. ( MCNZ) ; 1 male, laboratory reared, 23.i.2008, Bianchi, F.M. ( MCNZ) .

Description. Dorsal color greenish brown with a large, irregular ivory macula on anterior half of pronotum ( Fig. 1 ); posterior half of pronotum and disk of scutellum green in live specimens ( Fig. 3 ). Head usually ivory, varying from light brown to copper in some specimens. Dorsal punctures brown. Ventral surface ( Fig. 2 ) sordid ivory with pale, concolorous punctures.

Head. Wider across eyes than long. Punctures sparsely distributed on juga, tylus and vertex, denser on base of head and near the eyes, forming dark areas. Almost all punctures on juga forming a line along median margins. Length of juga and tylus subequal; lateral margins of juga lined in dark green. Antennal segment I, basal third of IV and basal half of V ivory; II and basal 2/3 of III brown to greenish light brown; apical third of III, 2/3 of IV and half of V dark brown. Proportion of antennal segments: I<II<III<IV≈V. R o s t r u m n o t surpassing metacoxae; apex black.

Thorax. Pronotum wider than long; disk green to brown with an irregular macula on anterior half between the humeral angles; macula usually ivory, sometimes concolorous and inconspicuous; anterior margin concave, lined by punctures; posterior margin sinuous; cicatrices almost inconspicuous, concolorous, and without internal punctures. Scutellum longer than wide at base; green to brown; postfrenal lobe sometimes with lighter spots; apex usually with ivory macula. Coria brown with lighter spots, longer than scutellum and reaching VI connexival segment. Membrane of hemelytra infuscate, with eight or nine parallel veins. Pleura sordid ivory; punctures shallow and concolorous; meso- and metapleura with 1+1 black spots; evaporative area concolorous or slightly darker than pleura, occupying less than half of mesopleura and almost 2/3 of metapleura; ostiolar ruga long. Prosternum and metasternum flat, mesosternum carinate; lateral margins not elevated.

Abdomen. Connexival segments yellowish brown to greenish brown; punctures small and concolorous; basal and apical angle of each segment bearing a black spot that continues ventrally. Ventral surface sordid ivory; shallow, sparse, and concolorous punctures on lateral 1/3. Abdominal tubercle conical and short, not surpassing metacoxae. Spiracles concolorous or light brown.

Male. Measurements (n=8). Body length 7.61±0.46 (6.90–8.25); abdominal width 4.26±0.27 (3.90–4.65); head length 1.37±0.15 (1.05–1.50), width 1.95±0.11 (1.80–2.10); length of antennal segments: I 0.40±0.03 (0.35–0.47), II 0.60±0.04 (0.55–0.66), III 0.94±0.06 (0.86–1.01), IV 1.20±0.10 (1.01–1.33), V 1.28±0.08 (1.17–1.37); pronotum length 1.58±0.11 (1.35–1.65), width 4.39±0.27 (3.90–4.65); scutellum length 3.19±0.30 (2.55–3.45), width 2.74±0.16 (2.55–2.85).

Genitalia. Pygophore narrowing at base; posterolateral angles rounded from dorsal view bearing a stout tooth or cusp dorsally projected ( Fig. 4 , t). Dorsal rim shallowly concave lateral to proctiger and slightly projected over its base ( Fig. 4 , dr). Proctiger subtriangular, narrowing from base to apex ( Figs 4, 6 , p). Ventral rim concave with a small median impression ( Figs 4, 5 , vr). Parameres spatulate, dorsally projected on each side of proctiger ( Figs 4, 6, 7 , par). Phallus simple with a bilobed vesica; ductus seminis distalis short, not attaining apex of vesica ( Figs 12,13 , dsd, v); conjunctiva partially covering vesica from lateral and ventral views ( Figs 13, 14 , cj).

Female. Measurements (n=9). Body length 8.40±0.67 (7.14–9.18); abdominal width 4.69±0.30 (4.08– 5.00); head length 1.26±0.19 (0.82–1.43), width 2.12±0.09 (1.94–2.24); length of antennal segments: I 0.40±0.04 (0.36–0.46), II 0.69±0.15 (0.51–0.92), III 0.94±0.08 (0.82–1.02), IV 1.20±0.12 (1.02–1.33), V 1.22±0.07 (1.12–1.38); pronotum length 1.64±0.12 (1.43–1.84), width 4.77±0.41 (4.08–5.30); scutellum length 3.39±0.41 (2.96–4.18), width 2.82±0.34 (2.14–3.26).

Genitalia. Gonocoxites 8 ( Fig. 15 , gc8) wider than long, separated at medial margin; posterior margin deeply excavated in middle 1/3, subrectilinear in lateral 2/3. Laterotergites 8 ( Fig. 15 , la8) wider than long; posterior margin convex, lined in dark brown to black. Gonapophyses 8 emarginated covering the base of gonocoxites 9, the latter trapezoidal ( Fig. 15 , gc9, tr). Laterotergites 9 little surpassing the band uniting laterotergites 8; apex narrowly rounded; segment X quadrangular ( Figs 15, 16 , la9, p). Gonapophyses 9 with 1+1 semicircular sclerotized areas close to elliptical ring sclerites (chitinellipsen, Dupuis 1955) ( Fig. 16 , g9, tg9, ch). Thickening of the vaginal intima elongate ( Fig. 16 , tvi). Ductus receptaculi short before the vesicular area; medial duct of the vesicular area enlarged at base and apex ( Fig. 16 , dre); pars intermedialis with two distinct cylindrical regions, posterior region nearly twice width of anterior ( Fig. 16 , pi). Anterior and posterior annular flanges directed toward ductus receptaculi and capsula seminalis, respectively; capsula seminalis ovoid, twice length of pars intermedialis, constricted near base and without projections ( Fig. 16 , aaf, cs, paf).

Comments. Banasa maculata sp. nov. fits within the “ cuspidata group” of Banasa ( Thomas & Yonke 1985) having the presence of a tooth or cusp at each posterolateral angle of pygophore. The new species keys with B. varians Stål , known from Rio de Janeiro ( Brazil), in the key to South American species of Banasa ( Thomas & Yonke 1990) . Banasa maculata can be distinguished by having a more robust, barrel-shaped pygophore instead of being subcylindrical and uniformly narrowed anteriorly as in B. varians ( Figs 7, 8 ); a triangular male proctiger instead of the trapezoidal one of B. varians ( Figs 7, 9 ); and female gonocoxites 8 more deeply excavated.

Life history and morphology of the immatures

Egg ( Figs 17–18 , 24–29). Reddish brown, barrel-shaped, operculum round and convex ( Figs 17, 18 ). Chorion surface, in S.E.M., reticulated ( Fig. 24 ), with deep polygonal cells surrounded by irregular projected rims that converge in somewhat truncate rough masses ( Figs 24, 26, 27 ). Polygonal cells smaller near aero-micropylar processes ( Figs 27, 28 ). Sculpture pattern altered where eggs are attached to each other in egg mass ( Fig. 26 ). Eclosion line well delimited, without depressions, bearing small irregular projections similar to truncate masses ( Fig. 25 ). Cells smaller and much more irregular at operculum. Height: 0.97±0.05 (0.98–1.01); width: 0.84 ± 0.03 (0.80–0.90). Aero-micropylar processes white, clavate, and with apical opening visible in S.E.M. ( Fig. 29 ), mean number of 34 ± 2.91 (28–38). One day before hatching, eyes visible as darkened areas under chorion; ruptor ovis visible only at eclosion.

First instar ( Fig 19 , Table 1 ). Body round and convex. General color predominantly red after hatching, dark brown to black after one hour. Head round in frontal view, declivent; tylus round at apex, slightly longer and darker than juga; margin of juga black. Eyes black, ocelli absent. Antennae four–segmented, dark brown, apical 2/3 of segment IV black; covered by short setae, denser at segment IV; segment I to III cylindrical, segment IV fusiform. Proportion of antennal segments: I<II>III<IV; segment IV long or longer than segments II and III together. Rostrum reaching metacoxae, black at base and apex. Thorax darker on disk; lateral margins depressed, pale, and somewhat translucent. Legs with short setae, denser at tibiae and tarsi. Coxae and femora darker than tibiae and tarsi, except for black base of tibiae and apex of second tarsomere. Tarsi two-segmented with pulvillus and a pair of claws. Abdomen light brown; lateral margins depressed. Five dorsal plates present, the anterior three bearing 1+1 openings of scent glands; lateral plates subtriangular. Spiracles on abdominal sternites II to VIII; 1+1 trichobothria from sternites III to VII.

Second instar ( Fig. 20 , Table 1 ). Body semicircular and convex; body elongated feeding for a few days. Black punctures on dorsal head, thorax, and abdominal plates. General color initially as first instar, then during the development white streaks appear on abdomen; pseudo-sutures brown. Juga and tylus subequal in length, former slightly deflected. Antennae as in first instar, but segment IV shorter than segments II and III combined. Rostrum black, surpassing metacoxae. Thorax dark brown, except for lighter sternum. Coxae and femora dark brown, latter with apical light brown ring; tibiae and tarsi light brown, apex of second tarsomere darker. Abdominal plates dark brown; white macula between all median plates and 1+1 laterad between second and third median plates. Abdominal sternites III to VII with 2+2 trichobothria. Other characters as described for first instar.

Third instar ( Fig. 21 , Table 1 ). Body semicircular to oval; general color variable, head and thorax predominantly light brown, in some individuals head and thorax black, similar to second instar; abdominal color pattern as in second instar after development of white streaks. Head and pronotum declivent. Posterior margin of head dark brown; two pairs of dark brown maculae, 1+1 projecting towards the base of tylus and 1+1 close to eyes. Area between eyes and macula unpunctured. Tylus and juga dark brown, latter outlined in red. Rostrum as in first instar but black only at apex. Thorax light brown, with contrasting dark brown maculae and median line; margins depressed and serrated. Maculae and punctures more conspicuous in lighter colored individuals. Prothorax trapezoidal; mesothorax rectangular, posterior margin concave, showing early developing scutellum. Abdomen as in second instar with reddish pseudo–sutures. Other characters as described for second instar.

Fourth instar ( Figs 22 , 30, Table 1 ). Body oval, general color varying from brown to green ( Fig. 30 ); maculae and punctures as in previous instar. Juga deflected, slightly surpassing tylus. Antennal segment II with reddish dorsal line and apex; segment III reddish. Posterior margin of mesothorax sinuous, wing pads slightly developed; developing scutellum and wing pads reaching posterior margin of metathorax. Other characters as described for previous instar.

Fifth instar ( Fig. 23 , Table 1 ). Body oval, general color variable as in fourth instar. Maculae and punctures as in third and fourth instars. Ocelli present. Mesothorax more developed; posterior margin of scutellum surpassing middle of abdominal segment II; wing pads well developed, attaining or surpassing posterior margin of abdominal segment III. Lateral margins of metathorax punctured. Lateral plates of abdomen brown, semicircular. Ventral abdomen punctured in red. Other characters as described for previous instar.

Biology. A total of 774 eggs of 66 egg clutches were studied. Modal number of eggs per clutch was 12, mean number was 11.73. Sex ratio was 0.48 (or 0.93 male: 1 female). Mean development time from egg to adult was 37.60 ± 13.24 days; shortest and longest development times were observed respectively for first and fifth instars ( Table 2 ). Mortality was similar from first to fourth instars and higher in fifth instar ( Table 2 ).

Comments

Morphology of the immatures. Eggs of Banasa maculata sp. nov. differ from other species by color and structure of chorion. In the new species, eggs are reddish brown, whereas they are pale yellow in B. centralis and B. zeteki ( Brailovsky et al. 1992) and brown in B. calva and B. dimiata ( DeCoursey 1963) . The chorion was described as granulated in B. dimiata and B. calva ( DeCoursey 1963) and punctured in B. zeteki ( Brailovsky et al. 1992) . A reticulated chorion similar to B. maculata sp. nov. was described for B. centralis ( Brailovsky et al. 1992) . These previous descriptions of the structure of the chorion, however, were made under light microscopy and should be considered with caution until S.E.M. studies are available. Eggs of B. maculata sp. nov., in S.E.M., show a reticulate pattern that resembles Chinavia longicorialis (Breddin) ( Matesco et al. 2009a), but with deeper cells and more developed rims. Nymphs of B. maculata sp. nov. can be distinguished from other species of Banasa mainly by the color of abdomen, although there are similarities with first and second instars of B. calva ( DeCoursey 1963) . Color variation observed in fourth and fifth instars of B. maculata sp. nov. was found in B. calva and B. dimiata ( DeCoursey 1963) , as well several Neotropical pentatomids; such variation may be regarded as a result of both heredity and diet ( Campos et al. 2007).

Biology. Some generalizations have been made concerning duration of nymphal instars of Pentatomidae , such as the first instar is shortest and the fifth longest, each succeeding instar is longer than preceding one, and second to fourth instars are nearly equal length ( DeCoursey & Esselbaugh 1962). In B. maculata sp. nov., however, the third instar is longer than the fourth; and similar conditions have been described for Neotropical pentatomines Oebalus ypsilongriseus (DeGeer) ( Vecchio & Grazia 1993) , Oebalus poecilus (Dallas) ( Greve et al. 2003) , Chinavia obstinata (Stål) ( Matesco et al. 2003) , Chinavia longicorialis (Breddin) ( Matesco et al. 2009a), and Euschistus hansi Grazia (Martins & Campos 2006) , whose second instar is longer than the third. Further investigations of other species are needed to confirm whether these relative development times described constitute exceptions in the Pentatominae .

Figures

FIGURE 3. Banasa maculata sp. nov., general aspect of live specimen showing greenish coloration.

FIGURE 30. Banasa maculata sp. nov., general aspect of live nymphs of 4th instar showing variation in color pattern.

FIGURES 1, 2. Banasa maculata sp. nov., holotype male, respectively dorsal and ventral.

FIGURES 4 – 16. Genitalia of Banasa maculata sp. nov. and Banasa varians Stål. Figs 4 – 6. B. maculata, male pygophore respectively dorsal, ventral, and posterior. Figs 7 – 9. B. varians, male pygophore respectively dorsal, ventral, and posterior. Figs 10, 11. B. maculata, male left paramere, lateral and mesial. Figs 12 – 14. B. maculata, male phallus respectively dorsal, ventral and lateral. Fig. 15. B. maculata, female genital plates, ventral. Fig. 16. B. maculata, female receptaculum seminis and ausenwand, ventral. (aaf—anterior annular flange; ch—chitinellipsen; cj—conjunctiva; cscapsula seminalis; dr—dorsal rim; dre—ductus receptaculi; dsd—ductus seminis distalis; g 8 — gonapophyses 8; g 9 — gonapophyses 9; gc 8 — gonocoxites 8; gc 9 — gonocoxites 9; la 8 — laterotergites 8; la 9 — laterotergites 9; p—proctiger; paf—posterior annular flange; par—paramere; pi—pars intermedialis; t—tooth; tg 9 — thickening of gonapophyses 9; tvi—thickening of vaginal intima; v—vesica; vr—ventral rim; X—tenth segment).

FIGURES 17–23. Banasa maculata sp. nov., egg and nymphs. Figs 17, 18. Egg lateral and apical. Figs 19–23. Nymphs, respectively 1st to 5th instars, dorsal. Scale bars = 1mm.

FIGURES 24 – 29. Banasa maculata sp. nov., egg in S. E. M .. Fig. 24. Lateral wall and operculum with aero-micropylar processes by the eclosion line. Fig. 25. Detail of the operculum showing the eclosion line. Fig. 26. Detail of the lateral wall showing part of contact area with neighboring egg (arrow). Fig. 27. Detail of the lateral wall showing area with smaller cells near aero-micropylar processes (arrow). Fig. 28. Detail of the lateral wall showing cells (asterisk) and truncate rough masses (arrow). Fig. 29. Aero-micropylar process with a apical hole (arrow). (amp – aero-micropylar process, el – eclosion line, lw – lateral wall, op – operculum).

Tables

TABLE 1. Measurements in millimeters (mean ± standard deviation, minimum – maximum) of morphometric parameters of nymphs of Banasa maculata sp. nov. reared under controlled conditions. (AL, AW—abdominal length and width; HL, HW—head length and width; ID—interocular distance; I, II, III, IV—1st to 4th antennal segments; RL— rostrum length; THL, THW—thorax length and width; TL—total length).

1st instar TL 1.06 ± 0.04 (1.01–1.13)	2nd instar 1.72 ± 0.14 (1.56–1.95)	3rd instar 2.50 ± 0.14 (2.32–2.73)	4th instar 3.70 ± 0.44 (3.08–4.45)	5th instar 5.27 ± 0.49 (4.22–5.80)
HL 0.24 ± 0.05 (0.17–0.33)	0.35 ± 0.05 (0.29–0.41)	0.42 ± 0.13 (0.23–0.72)	0.59 ± 0.07 (0.51–0.74)	1.03 ± 0.09 (0.88–1.13)
HW 0.65 ± 0.04 (0.56–0.68)	0.89 ± 0.02 (0.86–0.94)	1.17 ± 0.05 (1.11–1.27)	1.49 ± 0.08 (1.37–1.60)	1.76 ± 0.27 (1.32–2.14)
ID 0.50 ± 0.02 (0.47–0.53)	0.63 ± 0.07 (0.43–0.72)	0.83 ± 0.05 (0.78–0.94)	0.94 ± 0.06 (0.82–1.01)	1.12 ± 0.17 (0.69–1.26)
RL 0.65 ± 0.07 (0.54–0.73)	1.34 ± 0.11 (1.17–1.46)	1.55 ± 0.06 (1.46–1.68)	1.95 ± 0.10 (1.79–2.11)	2.42 ± 0.13 (2.21–2.58)
I 0.09 ± 0.01 (0.08–0.11)	0.17 ± 0.02 (0.14–0.20)	0.18 ± 0.02 (0.16–0.21)	0.21 ± 0.03 (0.16–0.25)	0.36 ± 0.06 (0.25–0.48)
II 0.16 ± 0.01 (0.16–0.19)	0.32 ± 0.02 (0.29–0.35)	0.41 ± 0.02 (0.39–0.45)	0.63 ± 0.05 (0.57–0.70)	0.99 ± 0.05 (0.88–1.03)
III 0.13 ± 0.02 (0.11–0.16)	0.24 ± 0.03 (0.20–0.27)	0.33 ± 0.03 (0.27–0.39)	0.52 ± 0.05 (0.45–0.59)	0.91 ± 0.03 (0.85–0.95)
IV 0.33 ± 0.03 (0.29–0.39)	0.50 ± 0.07 (0.45–0.53)	0.62 ± 0.07 (0.51–0.78)	0.82 ± 0.03 (0.74–0.86)	1.08 ± 0.03 (1.03–1.13)
THL 0.39 ± 0.02 (0.36–0.42)	0.80 ± 0.03 (0.76–0.84)	1.07 ± 0.06 (0.98–1.17)	1.49 ± 0.10 (1.37–1.60)	2.24 ± 0.15 (2.02–2.52)
THW 1.02 ± 0.04 (0.93–1.09)	1.52 ± 0.05 (1.44–1.60)	1.99 ± 0.12 (1.85–2.24)	2.75 ± 0.18 (2.38–2.93)	3.73 ± 0.13 (3.53–3.91)
AL 0.40 ± 0.04 (0.31–0.47)	0.63 ± 0.17 (0.45–0.90)	1.05 ± 0.15 (0.78–1.27)	1.72 ± 0.37 (1.13–2.34)	2.53 ± 0.52 (1.51–3.21)
AW 1.01 ± 0.05 (0.93–1.15)	1.45 ± 0.10 (1.35–1.58)	2.00 ± 0.18 (1.77–2.24)	2.99 ± 0.28 (2.61–3.39)	3.99 ± 0.32 (3.47–4.28)

TABLE 2. Duration (in days) and relative mortality (%) of eggs and nymphs of Banasa maculata sp. nov. reared under controlled conditions. (SD—standard deviation).

	Duration (days) (mean ± SD)	Variation interval (days)	Mortality (%)
Egg	5.17 ± 1.46	3–7	11.12
1st instar	4.10 ± 1.27	3–7	26.44
2nd instar	6.53 ± 1.88	3–13	14.99
3rd instar	7.16 ± 3.85	3–16	23.88
4th instar	6.32 ± 2.10	2–17	12.76
5th instar	8.37 ± 2.67	3–14	45.95