Symmachia atlantica Dias, Dolibaina & Mielke, 2020

Dias, Fernando Maia Silva, Dolibaina, Diego Rodrigo, Mielke, Olaf Hermann Hendrik & Casagrande, Mirna Martins, 2020, Taxonomic review of the “ Probetoriformes ” species group of Symmachia Hübner [1819] (Lepidoptera: Riodinidae), with the description of two new species from the Brazilian Atlantic Forest, Zootaxa 4780 (3), pp. 471-507 : 491-494

Symmachia atlantica Dias, Dolibaina & Mielke , sp. nov.

( Figs 17–20 , 43 , 52 , 61 , 71 , 80 , 93–94 , 111–112 , 127 , 137 , 142 , 152 , 154 ) http:// urn:lsid:zoobank.org:act:CF90518B-A0FE-47B2-A3FB-B6005B6EDADC

Symmachia praxila [misidentification]; Zikán, 1928. Ent. Rundsch. 45(2), p. 7.

Symmachia praxila var.? [misidentification]; Zikán, 1928. Ent. Rund. 45(2), p. 7.

Symmachia probetor ? [sic] [misidentification]; Kesselring & Ebert, [1982]. Rev. nord. Biol. 2(1/2), p. 117.

Symmachia probetor probetor [misidentification]; D’Abrera, 1994. Butt. Neotrop. Reg. IV. Riod., p. 1044, fig. 13, female dorsal] ( Brazil)—Palo Jr., 2017. Borb. Bras. 1, p. 593, fig.[2] (female in situ).

Symmachia probetor [misidentification]; Brown & Freitas, 2000. Bol. Mus. Biol. Mello Leitão N. s., 11/12, p. 110.

Diagnosis. Symmachia atlantica is sympatric with S. praxila and S. uirassu ( Figs 152–153 ). Males are most similar to S. probetor , S. belti , S. championi , and S. estellina ( Figs 17–18 ) and females are quite similar to most species of the “Probetoriformes” species group ( Figs 19–20 ). Males with head and thorax as in S. probetor ; FW with a moderate bulge; bulged area as 45% of the FW length; end of bulge aligned with the origin of M 2 ( Figs 17 , 61 ); FWD black, with a dark green sheen covering most of the wing, with a translucent white and creamy sickle-shaped spot between the discal cell and the costal margin, and a small round whitish spot in the postdiscal area at the costal margin in R 2 –R 3 ( Figs 17 , 61 ); FWD dark green sheen absent roughly on the submedial areas from the inner margin to the discal cell and along the outer margin; HWD black, red along the costal margin and a dark green sheen patch near the tornus ( Fig. 18 ); abdomen color pattern as in S. probetor ; CAS of the tergites 4 and 5 as long as 1/6 of the tergites’ length and as wide as half the tergites’ width, projected anteriorly, CAS scales pale beige ( Figs 111–112 ); valva larger than in S. probetor ; aedeagus thin and short; vesica with two bands of cornuti ( Figs 127 , 137 ). Females with thorax as in S. probetor ; FW moderately bulged ( Figs 18 , 71 ); FWD and FWV submarginal reddish orange band on the submarginal along the outer margin broken beyond M 2 –M 3, not reaching the inner margin at the tornus ( Figs 19–20 , 71 ); FWD dark brown, with a translucent white and creamy sickle-shaped spot between the discal cell and the costal margin and speckled with faint whitish and reddish orange spots ( Fig. 71 ); HWD submarginal band reddish orange and wavy ( Fig. 19 ); FWV and HWV spots well developed ( Figs 20 , 80 ); HWV submarginal band well developed and wavy and; HWV marginal band well developed, outlining a number of ocelli ( Fig. 80 ); sterigma trapezoidal with a triangular median bulge at the ostium; ductus bursae membranous near the ostium; bursa copulatrix rather stretched; signum absent ( Fig. 142 ).

Description. Male. Head ( Figs 43 , 52 ): mostly covered by compact black scales, with some scattered orangeyellowish scales on the frons and bordering the compound eyes and base of antenna; antenna black, each antennomer with proximal white scales, antennal club formed by the last 10–12 antennomeri, tip yellowish; labial palpus dark brown, short, not extending beyond middle of frons, first segment as long as half of the second segment, third segment about 1/6 as long as second.

Thorax: ventrally dark brown; prothorax dorsally with yellowish scales; mesothorax dorsally black, with a dark bluish sheen; metathorax dorsally with long, dark brown scales; legs mostly dark brown, each segment ventrally with distal white scales.

FW, size and shape ( Figs 17 , 61 ): average size 1.16 cm (1.1–1.3 cm; n=33); roughly triangular; costal margin sinuous, moderately bulged in the basal area; apex angled; outer margin slightly convex; tornus angled; inner margin almost straight.

FWD ( Figs 17 , 61 ): mostly black with iridescent dark blue and dark green scales, and translucent white and creamy spots; dark blue sheen on the costal margin bulge and on the basal area; dark green sheen roughly on the basal, postdiscal and submarginal areas; sickle-shaped spot formed by a translucent white spot on the discal cell, and creamy spots between the costal margin and Sc–R; and a round whitish spot in the postdiscal area at the costal margin in R 2 –R 3; fringe black.

FWV ( Fig. 18 ): mostly dark brown, suffused from tornus with white scales along the outer and innermargins; sickle-shaped spot and postdiscal spots as in DFW.

HW, shape ( Fig. 17 ): roughly triangular; costal margin straight; apex rounded; outer margin straight; tornus angled; inner margin slightly convex.

HWD ( Fig. 17 ): mostly black with a scarlet red band along the costal margin reaching the upper half of the discal cell and M 2; dark green sheen in the postdiscal and submarginal areas near the tornus, from about CuA 1 to 1A; dark brown along the inner margin roughly beyond 2A; fringe black.

HWV ( Fig. 18 ): mostly dark brown, with faint whitish spots, and submarginal and marginal bands; whitish spots in a more or less checkered pattern from the basal to the postdiscal area; submarginal and marginal bands outlining one lenticular ocellus per space between M 1 and 1A.

Abdomen ( Figs 18 , 93–94 , 111–112 ): dorsally black, with dark blue sheen; laterally and ventrally dark brown, segments with distal white scales; tergites 4 and 5 with a not divided band of CAS on the posterior margin, as long as 1/6 of the tergites’ length and as wide as half the tergites’ width, projected anteriorly and scales pale beige.

Genitalia ( Figs 127 , 137 ): tegumen rectangular; membranous area between the uncus and the tegumen roughly triangular; uncus rectangular, slightly bilobed posteriorly; gnathos C-shaped, ventral part about one third larger than the dorsal, connected to the tegumen by a small dorsal projection; ventral arm of the tegumen thin and obliquely projected, connected ventrally to the dorsal arm of the saccus; anterior projection of the saccus as wide as long and slightly bilobed anteriorly; valva roughly triangular in lateral view, connected ventrally to the saccus and the pedicel, and dorsally to the opposite valva; pedicel posteriorly sclerotized and keeled, connected to the middle of the aedeagus; aedeagus roughly cylindrical, wider anteriorly; vesica with two bands of small pointed cornuti of similar size; subscaphium sclerotized.

Female. Head: mostly covered by compact yellow scales, including gena and base of antenna; labial palpus yellow, shape and size as in males.

Thorax ( Figs 19–20 ): patagium yellow; dorsally black with three pairs of orange rounded spots, two on mesothorax, the third more elongated on metathorax; ventrally yellow; legs most yellow.

Forewing, size and shape ( Figs 19 , 71 ): average size 1.28 cm (1.1–1.4 cm; n=13); similar to the male, but outer margin more convex; tornus obtuse.

FWD ( Figs 19 , 71 ): mostly brown with yellowish and reddish orange spots; dark blue sheen on the costal margin bulge and on the basal area; discal cell with three reddish orange spots sometimes fused and forming a larger spot; sickle-shaped spot formed by two elements, one semi-hyaline yellowish spot in the discal cell and a narrower nonhyaline yellowish spot between discal cell and costa; postdiscal area with five yellowish creamy spots near costa, one in R 2 –R 3, R 3 –R 4+5, R 4+5 –M 1 (the biggest) and two smaller in M 1 –M 2; five reddish orange spots between CuA 2 and inner margin; postdiscal area with four to five faint yellow spots between M 2 and 2A; submarginal reddish orange band large and continuous in R 4+5 –M 3, then thin and discontinuous, rarely reaching tornus; fringe dark brown.

FWV ( Fig. 20 ): mostly dark brown, whitish along 2A; spots as in upper side but larger and light yellow, and an additional postdiscal spot just after the end of discal cell in M 2 –M 3; submarginal band as in upper side but orange and thinner.

Hind wing, shape: as in male, but longer.

HWD ( Fig. 19 ): mostly dark brown with faint yellow to reddish spots; submarginal reddish orange band very thin, continuous and slightly wavy, from M 1 to tornus; fringe dark brown.

HWV ( Figs 20 , 80 ): dark brown with several somewhat rectangular yellowish cream spots in basal, discal and postdiscal areas; anal area almost completely yellow; submarginal yellow band enclosing five dark brown ocelli along outer margin (one by space between veins), the larger one in CuA 2 –2A; fringe dark brown.

Abdomen ( Figs 19–20 ): dark brown with thin yellow rings at the distal margin of each segment, longer ventrally.

Genitalia ( Fig. 142 ): sterigma trapezoid, about four times as wide as long, poorly sclerotized, with a small trapezoid central smooth area surrounded by a more or less membranous zone; ductus bursae membranous, curved just after ostium bursae, as long as half of bursa copulatrix; corpus bursae moderately large, membranous, usually without signum (n=7), but sometimes with a small triangular signum at its first third (n=2); papilla analis wider than long, area with setae restricted to its distal third, and with a small rounded lateral projection on its upper portion.

Etymology. The name of this species alludes to its occurrence in the Atlantic Forest biome (“Mata Atlântica”) along the northeastern, southeastern and southern coasts of Brazil. The name is treated as a noun in apposition.

Distribution. Widespread in the ombrophilous dense forest of the Atlantic Forest biome of Brazil (Paraíba, Pernambuco, Alagoas, Bahia, Espírito Santo, Minas Gerais (Palo Jr. 2017), Rio de Janeiro, São Paulo, Paraná and Santa Catarina) ( Figs 152 , 154 ), occurring in an altitudinal range between sea level to 1200 m. This species is likely to occur in other states of Brazil within the Atlantic Forest biome as Sergipe, Rio Grande do Norte and Ceará.

Comments. Symmachia atlantica is described based on the male holotype, the allotype, and further 45 males and 16 females paratypes from Brazilian Atlantic Forests. This species is certainly the species tentatively identified by Kesselring & Ebert (1982) and misidentified by Brown & Freitas (2000) as S. probetor , as S. atlantica is the only species occurring in their study area likely to be confused with the allopatric S. estellina . Specimens misidentified as S. praxila and S. praxila var? [sic] recorded by Zikán (1928) probably refers to females of S. atlantica , as can be inferred by specimens of S. atlantica collected and identified by him from Itatiaia, Rio de Janeiro, Brazil, deposited at the IOC. Female specimens were associated with male specimens based on the HWV pattern of spots along the outer margin and the co-occurrence of several male and female specimens caught at the same localities and dates.

Males of this species are found in hilltops along the Brazilian coast area ( Fig. 154 ) from 10:30 h. where they rest in small trees up to three meters and eventually perform short flights resting again in the same spots. This behavior was noted for the holotype captured at a 600 m hilltop in Morro Alto, Morretes, Paraná, and in a series of individuals caught in higher elevation hilltops at Curva do Violão , Santa Teresa , Espírito Santo (about 890 m) and RPPN Serra Bonita , Camacan , Bahia (920 m) ( Fig. 154 ), where no more than three males were seen in a same day .

Type material. Male holotype of Symmachia atlantica with the following labels: / HOLOTYPUS / HOLOTY- PUS Symmachia atlantica Dias, Dolibaina & Mielke det. 2018/ DZ 36.642 / B[ar]C[oded]-DZ[ UP] / 22-IX-2015 MORRO ALTO, MORRETES, PARANÁ, BRASIL, MIELKE, DOLIBAINA & SIEWERT LEG./ ( DZUP). Female allotype of Symmachia atlantica with the following labels: / ALLOTYPUS / ALLOTYPUS Symmachia atlantica Dias, Dolibaina & Mielke det. 2019/ DZ 44.063 / ALEXANDRA, PARANÁ, BRASIL, 27-IV-1968 [Mielke leg.]/ ( DZUP).

Paratypes. BRAZIL: Alagoas —São José da Lage (Usina Serra Grande), 300 m, 1♀, 16–17.III.1997, Mielke & Casagrande leg., OM 46.964 ( OM) . Bahia —Camacan, 600 m, 1♂, I.1992, V . O. Becker leg., OM 64.586 ( OM); ( RPPN Serra Bonita ), 920 m , 7♂, 3–10.XII.2016, Mielke, Carneiro, Dias, Dolibaina & Santos leg., DZ 36.612, DZ 43.834, DZ 43.894, DZ 43.934, DZ 43.994, DZ 44.024, DZ 44.054 ( DZUP) . Espírito Santo —São Mateus, 1♂, X.1985, C. Elias leg., DZ 43.864 ( DZUP) . Santa Teresa, 2♀, 5.XII.1966, C. & C. T . Elias leg., DZ 44.003*, DZ 44.007* ( DZUP); ( Curva do Violão ), 890 m , 2♂, 11–12. V .2019, Dolibaina & Lorenzoni leg., DD 589, DD 590 (DD), 1♂, 29–30. VI .2019, Dolibaina leg., DD 562 (DD). Domingos Martins (Pedra Azul), 1200 m, 1♂, 28.I.2003, Mielke & Casagrande leg., OM 59.400 ( OM) . Paraná — Antonina (Cacatu) , 20 m , 1♂, 1♀, 25.IV.1967, Mielke leg., OM 61.814 , OM 61.940 ( OM) . Guaraqueçaba (Tagaçaba), 1♂, 17.IV.1971, Moure & Mielke leg., DZ 43.904 ( DZUP) . Rio de Janeiro —Petrópolis (Independência), 900 m, 1♀, 21. V .1936, P. Gagarin leg., DZ 44.093* ( DZUP); 1♂, 8. VI .1939, P. Gagarin leg., DZ 44.044 ( DZUP). Teresópolis (Barreira), 1♀, 21.VII.1955, ex.-coll. Gagarin, DZ 43.831 ( DZUP) ; 1♀, 20.X.1955, ex.-coll. Gagarin, DZ 43.963 ( DZUP) . Santa Catarina —Joinville, 1♂, Mielke & Miers leg., OM 67.595 ( OM) ; 1♂, ex. coll. E. May, ( MZUSP) ; 1♂, IV, ex. coll. Gagarin, DZ 43.874 ( DZUP) ; 1♂, VIII.1940, B. Pohl leg. ( MZUSP) ; 1♀, IV.1941, B. Pohl leg. ( MZUSP) ; 1♂, 23. V .1942, Rank ex-coll. ( MGCL) ; 1♂, IV.1944, B. Pohl leg. ( MZUSP); 200 m , 2♂, 15.X.1968, Miers leg., DZ 43.854, DZ 43.963 ( DZUP) ; 1♂, 27.X.1968, Mielke leg., DZ 21.824 ( DZUP) ; 3♂, 20.IV.1969, Mielke & Miers leg., DZ 43.319, DZ 43.954, DZ 44.064 ( DZUP) ; 1♀, 19.X.1969, Miers leg., OM 67.651 ( OM) ; 1♂, 31.I.1971, Mielke & Miers leg., DZ 43.984 ( DZUP) ; 1♂, 4.II.1973, Mielke leg., DZ 43.844 ( DZUP) ; 1♀, 14.III.1973, Miers leg., DZ 43.339 ( DZUP) ; 1♂, 9.III.1974 ( MGCL) ; 2♂, 15.I.1977, Mielke & Miers leg., DZ 43.884, DZ 43.934 ( DZUP) ; 2♂, 26.II.1977 ( MGCL) ; 1♀, 30.III.1979, Miers leg., DZ 44.043 ( DZUP) ; 1♀, 26.X.1980, Miers leg., DZ 44.073 ( DZUP) ; 1♀, 2.IX.1982, Mielke leg., DZ 44.083 ( DZUP) ; 1♀, 15.IX.1982, Miers leg., DZ 44.033 ( DZUP) ; 1♂, 20.X.1984, Miers leg. ( MGCL) ; 1♂, 29.XII.1993, Mielke & Miers leg., OM 37.790 ( OM) ; 6♂, 1.IX.1997, Mielke & Miers leg., OM 45.820 , OM 45.828 , OM 45.836 , OM 45.844 , OM 45.852 , OM 45.860 ( OM) . São Paulo —Cananéia (Ariri), 1♀, 26.IV.1980, Miers leg., DZ 43.993 ( DZUP) . Indiana, 1♂, 1♀, IV.1944, B. Pohl leg. ( MZUSP) .

Figures

FIGURE 152. Geographical distribution of Symmachia estellina and S. atlantica Dias, Dolibaina & Casagrande sp. nov. Asterisks indicate records from the literature.

FIGURES 1–20. Species of Symmachia belonging to the “Probetoriformes” species group. 1–4. S. probetor. 1–2. Male from French Guiana (SF). 3–4. Female from French Guiana (DZ 40.291 (DZUP)). 5–8. S. belti stat. rev.. 5–6. Male from Potrerillos, Chiriqui, Panamá (USNM). 7–8. Female from 6 miles N of El Llano, Panamá, Panamá (USNM). 9–12. S. championi stat. rest.. 9–10. Male from Santa Tecla, El Salvador (DZ 23.224 (DZUP)). 11–12. Female from San Salvador, El Salvador (USNM). 13–16. S. estellina. 13–14. Male from Santo Antônio do Tauá, Pará, Brazil (DZ 43.499 (DZUP)). 15–16. Female from 4 Km Tena-Puno road, Napo, Ecuador (USNM). 17–20. S. atlantica Dias, Dolibaina & Mielke sp. nov.. 17–18. Male holotype (DZ 36.642 (DZUP)). 19–20. Female allotype (DZ 44.063 (DZUP)).

FIGURES 39–47. Heads of the Symmachia belonging to the “Probetoriformes” species group in anterior view. 39. S. probetor. 40. S. belti stat. rev. 41. S. championi stat. rest. 42. S. estellina. 43. S. atlantica Dias, Dolibaina & Mielke sp. nov.. 44. S. praxila. 45. S. uirassu Dolibaina, Dias & Casagrande sp. nov.. 46. S. divisora. 47. S. falcistriga. Scale bar = 5 mm.

FIGURES 48–56. Heads of the Symmachia belonging to the “Probetoriformes” species group in lateral view. 48. S. probetor. 49. S. belti stat. rev.. 50. S. championi stat. rest. 51. S. estellina. 52. S. atlantica Dias, Dolibaina & Mielke sp. nov.. 53. S. praxila. 54. S. uirassu Dolibaina, Dias & Casagrande sp. nov.. 55. S. divisora. 56. S. falcistriga. Scale bar = 5 mm.

FIGURES 57–66. Forewings of males of Symmachia belonging to the “Probetoriformes” species group in dorsal view. 57. S. probetor. 58. S. belti stat. rev. 59. S. championi stat. rest. 60. S. estellina. 61. S. atlantica Dias, Dolibaina & Mielke sp. nov.. 62. S. praxila. 63. S. uirassu Dolibaina, Dias & Casagrande sp. nov.. 64. S. divisora. 65. S. falcistriga. 66. S. meyi stat. nov.

FIGURES 67–75. Forewings of females of Symmachia belonging to the “Probetoriformes” species group in dorsal view. 67. S. probetor. 68. S. belti stat. rev. 69. S. championi stat. rest. 70. S. estellina. 71. S. atlantica Dias, Dolibaina & Mielke sp. nov.. 72. S. praxila. 73. S. divisora. 74. S. falcistriga. 75. S. meyi stat. nov.

FIGURES 76–84. Hindwings of females of Symmachia belonging to the “Probetoriformes” species group in ventral view. 76. S. probetor. 77. S. belti stat. rev. 78. S. championi stat. rest. 79. S. estellina. 80. S. atlantica Dias, Dolibaina & Mielke sp. nov.. 81. S. praxila. 82. S. divisora. 83. S. falcistriga. 84. S. meyi stat. nov.

FIGURES 85–94. Abdomens of males of Symmachia belonging to the “Probetoriformes” species group in dorsal (left) and lateral (right) views. 85–86. S. probetor. 87–88. S. belti stat. rev. 89–90. S. championi stat. rest. 91–92. S. estellina. 93–94. S. atlantica Dias, Dolibaina & Mielke sp. nov. Scale bar = 5 mm.

FIGURES 103–112. Fourth (left) and fifth (right) tergites of males of Symmachia belonging to the “Probetoriformes” species group. 103–104. S. probetor. 105–106. S. belti stat. rev. 107–108. S. championi stat. rest. 109–110. S. estellina. 111–112. S. atlantica Dias, Dolibaina & Mielke sp. nov. Scale bar = 1 mm.

FIGURES 123–127. Male genitalia of Symmachia belonging to the “Probetoriformes” species group in lateral view, uncus in dorsal view and anterior projection of saccus in ventral view. 123. S. probetor. 124. S. belti stat. rev. 125. S. championi stat. rest. 126. S. estellina. 127. S. atlantica Dias, Dolibaina & Mielke sp. nov. Scale bar = 1 mm.

FIGURES 133–137. Male genitalia of Symmachia belonging to the “Probetoriformes” species group in ventral-posterior view. 133. S. probetor. 134. S. belti stat. rev. 135. S. championi stat. rest. 136. S. estellina. 137. S. atlantica Dias, Dolibaina & Mielke sp. nov. Scale bar = 1 mm.

FIGURES 138–146. Female genitalia of Symmachia belonging to the “Probetoriformes” species group in ventral view and papilla analis in lateral view. 138. S. probetor. 139. S. belti stat. rev. 140. S. championi stat. rest. 141. S. estellina. 142. S. atlantica Dias, Dolibaina & Mielke sp. nov. 143. S. praxila. 144. S. uirassu Dolibaina, Dias & Casagrande sp. nov.. 145. S. divisora. 146. S. falcistriga. Scale bar = 1 mm.

FIGURES 154–156. Habitat of some species of Symmachia belonging to the “Probetoriformes” species group. 154. Hilltop at the Bromélias trail, Reserva Serra Bonita, Camacan, Bahia, Brazil, the arrow indicates the exact location where males of S. atlantica Dias, Dolibaina & Casagrande sp. nov. were collected. 155. Opposite view of the previous figure, arrow indicates the exact location where males of S. uirassu Dolibaina, Dias & Casagrande sp. nov. were collected. 156. DRD on a tree at the lightgap of the hilltop in Parque Nacional da Serra do Divisor, Mâncio Lima, Acre, Brazil where males of S. probetor, S. estellina, S. divisora and S. falcistriga were collected (photo by Eduardo Carneiro).