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
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Banasa maculata sp. nov. Campos & Garbelotto
Etymology. Specific epithet referring to the irregular macula on anterior half of pronotum.
Paratypes. BRAZIL: Santa Catarina: Criciúma, Parque José Milanese , 1 male, 15.iii. 2007, Frassetto, J. [- 28.690, - 49.431] ( UFRG)GoogleMaps ; 4 females and 1 male, S.A.T.C., 18.x. 2007, Bianchi, F.M. [- 28.707, - 49.409] ( MZSP)GoogleMaps ; 2 females, same data as previous ( UFRS)GoogleMaps ; 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. 1View FIGURES 1, 2); posterior half of pronotum and disk of scutellum green in live specimens ( Fig. 3View FIGURE 3). Head usually ivory, varying from light brown to copper in some specimens. Dorsal punctures brown. Ventral surface ( Fig. 2View FIGURES 1, 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. 4View FIGURES 4 – 16, t). Dorsal rim shallowly concave lateral to proctiger and slightly projected over its base ( Fig. 4View FIGURES 4 – 16, dr). Proctiger subtriangular, narrowing from base to apex ( Figs 4, 6View FIGURES 4 – 16, p). Ventral rim concave with a small median impression ( Figs 4, 5View FIGURES 4 – 16, vr). Parameres spatulate, dorsally projected on each side of proctiger ( Figs 4, 6, 7View FIGURES 4 – 16, par). Phallus simple with a bilobed vesica; ductus seminis distalis short, not attaining apex of vesica ( Figs 12,13View FIGURES 4 – 16, dsd, v); conjunctiva partially covering vesica from lateral and ventral views ( Figs 13, 14View FIGURES 4 – 16, 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. 15View FIGURES 4 – 16, gc 8) wider than long, separated at medial margin; posterior margin deeply excavated in middle 1 / 3, subrectilinear in lateral 2 / 3. Laterotergites 8 ( Fig. 15View FIGURES 4 – 16, la 8) 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. 15View FIGURES 4 – 16, gc 9, tr). Laterotergites 9 little surpassing the band uniting laterotergites 8; apex narrowly rounded; segment X quadrangular ( Figs 15, 16View FIGURES 4 – 16, la 9, p). Gonapophyses 9 with 1 + 1 semicircular sclerotized areas close to elliptical ring sclerites (chitinellipsen, Dupuis 1955) ( Fig. 16View FIGURES 4 – 16, g 9, tg 9, ch). Thickening of the vaginal intima elongate ( Fig. 16View FIGURES 4 – 16, tvi). Ductus receptaculi short before the vesicular area; medial duct of the vesicular area enlarged at base and apex ( Fig. 16View FIGURES 4 – 16, dre); pars intermedialis with two distinct cylindrical regions, posterior region nearly twice width of anterior ( Fig. 16View FIGURES 4 – 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. 16View FIGURES 4 – 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, 8View FIGURES 4 – 16); a triangular male proctiger instead of the trapezoidal one of B. varians ( Figs 7, 9View FIGURES 4 – 16); and female gonocoxites 8 more deeply excavated.
Life history and morphology of the immatures
Egg ( Figs 17 –18View FIGURES 17–23, 24– 29). Reddish brown, barrel-shaped, operculum round and convex ( Figs 17, 18View FIGURES 17–23). Chorion surface, in S.E.M., reticulated ( Fig. 24View FIGURES 24 – 29), with deep polygonal cells surrounded by irregular projected rims that converge in somewhat truncate rough masses ( Figs 24, 26, 27View FIGURES 24 – 29). Polygonal cells smaller near aero-micropylar processes ( Figs 27, 28View FIGURES 24 – 29). Sculpture pattern altered where eggs are attached to each other in egg mass ( Fig. 26View FIGURES 24 – 29). Eclosion line well delimited, without depressions, bearing small irregular projections similar to truncate masses ( Fig. 25View FIGURES 24 – 29). 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. 29View FIGURES 24 – 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 19View FIGURES 17–23, 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. 20View FIGURES 17–23, 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. 21View FIGURES 17–23, 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 22View FIGURES 17–23, 30, Table 1). Body oval, general color varying from brown to green ( Fig. 30View FIGURE 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. 23View FIGURES 17–23, 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).
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. 2009 a), 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. 2009 a), 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 .
|2nd instar 1.72 ± 0.14 (1.56–1.95)|
|0.42 ± 0.13 (0.23–0.72)|
|1.49 ± 0.08 (1.37–1.60)|
|0.94 ± 0.06 (0.82–1.01)|
|1.34 ± 0.11 (1.17–1.46)|
|0.41 ± 0.02 (0.39–0.45)|
|0.62 ± 0.07 (0.51–0.78)|
|1.07 ± 0.06 (0.98–1.17)||1.49 ± 0.10 (1.37–1.60)|
|1.52 ± 0.05 (1.44–1.60)||1.99 ± 0.12 (1.85–2.24)|
|1.05 ± 0.15 (0.78–1.27)|
|1.45 ± 0.10 (1.35–1.58)|
|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|
Brazil, Sao Paulo, Sao Paulo, Museu de Zoologia da Universidade de Sao Paulo
Brazil, Rio Grande do Sul, Porto Alegre, Museu de Ciencias Naturais da Fundacao Zoo-Botanica do Rio Grande do Sul
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