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 Author Dias, Fernando Maia Silva Author Dolibaina, Diego Rodrigo Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, PO Box 19020, 81.531 - 980, Curitiba, Paraná, Brazil. & dirodrido @ gmail. com; https: // orcid. org / 0000 - 0001 - 5258 - 3787 Author Mielke, Olaf Hermann Hendrik Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, PO Box 19020, 81.531 - 980, Curitiba, Paraná, Brazil. & omhesp @ ufpr. br; https: // orcid. org / 0000 - 0003 - 3655 - 4606 Author Casagrande, Mirna Martins Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, PO Box 19020, 81.531 - 980, Curitiba, Paraná, Brazil. & mibras @ ufpr. br; https: // orcid. org / 0000 - 0002 - 6076 - 8463 text Zootaxa 2020 2020-05-26 4780 3 471 507 journal article 21852 10.11646/zootaxa.4780.3.3 70713ddc-4f67-4f0a-8625-41aa37abb017 1175-5326 3855256 CA5DA614-D448-4D95-B271-DDC8D56A37F60 Symmachia estellina Gallard, 2008 ( Figs 13–16 , 42 , 51 , 60 , 70 , 79 , 91–92 , 109–110 , 126 , 136 , 141 , 152 , 156 ) Symmachia probetor [ misidentification ]; Staudinger, 1887, in Staudinger & Schatz. Exot. Schmett. 1(18) , pl. 91— Seitz, 1913 Gross-Schmett. Erde 5, pl. 128g [4] (male dorsal)— Seitz, 1917 . Gross-Schmett. Erde 5 , p. 682—Seitz, 1931. Ent. Rundsch. 48(24), p. 260—Salazar & Constantino, 2000. SHILAP Revta. lepid. 28(112), p. 376 [ in part ], fig. 7 (female dorsal). Symmachia hippea [ misidentification ]; Seitz, 1913 . Gross-Schmett. Erde 5, pl. 128h[4] (female dorsal). Symmachia calliste [ misidentification ]; Seitz, 1917 . Gross-Schmett. Erde 5 , p. 681 [ in part ]. Symmachia probator [ sic ] [ misidentification ]; Lewis, 1973. Butt. World , p. 243, pl. 79, fig. 3 (male dorsal). Symmachia juratrix [ misidentification ]; D’Abrera, 1994 . Butt. Neotrop. Reg. IV. Riod., p. 1044, fig. [6] (female ventral). Symmachia probetor probetor [ misidentification ]; Barcant, 1970. Butt. Trinid. Tob., p. 206; pl. 27, fig. 19 (male dorsal)— D’Abrera, 1994 . Butt. Neotrop. Reg. IV. Riod., p. 1044, fig. [12] (male dorsal)— Brévignon & Gallard, 1998 . Lambill. 98(3) , p. 308, 316, 317; figs 31-34 (male and female, dorsal and ventral); fig. 1- 2 p. 320 (male genitalia). - Emery et al. , 2006. Rev. Bras. Ent. 50(1) , p. 89. Symmachia estellina Gallard, 2008 , in Lacomme & Manil. Lep. Guyane . Tome 3 Rhopal. 2 , p. 27, 30-32, 39, 40; figs 1- 4 p. 31 ( holotype male and allotype female, dorsal and ventral), (female ventral, schematic illustration); pl. 1, fig. 2 (male in situ ), pl. 2, fig. 2 (male dorsal, female dorsal and ventral)— Dolibaina et al. , 2012 . Insecta Mundi 249, p. 4; figs 9-10 (male dorsal and ventral), 40 (forewing dorsal), 43 (male genitalia)—Salik et al ., 2014. SHILAP Revta. lepid. 42(166), p. 278— Gallard, 2017 . Riod. Guyane , p. 75, 165; fig.10B (female ventral, schematic illustration); pl. 15 ( in situ ), fig. G (male dorsal), pl. 15, fig. 5 (male dorsal, female dorsal and ventral)— Sambhu & Nankishore, 2018 . Zootaxa. 4371(1), p. 162. Symmachia estellina estellina ; Palo Jr., 2017. Borb. Bras. 1 , p. 716. Diagnosis. Symmachia estellina is sympatric with S. probetor , S. divisora and S. falcistriga ( Figs 151–153 ). Males are most similar to S. probetor , S. belti , S. championi , and S. atlantica ( Figs 13–14 ) and females are quite similar to most species of the “Probetoriformes” species group ( Figs 15–16 ). Males with head and thorax as in S. probetor ; FW (average size: 1.11 cm ; 1.1–1.15 cm ; n=19) moderately bulged, apex narrower than in other similar species; bulged area as 40% of the FW length; end of bulge aligned to the origin of M 2 ( Figs 13 , 60 ); 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 ( Figs 13 , 60 ); 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 spot along the costal margin and a dark green sheen patch near the tornus ( Fig. 13 ); abdomen color pattern as in S. probetor ; CAS of the tergites 4 and 5 as long as 1/6 to 1/7 of the tergites’ length and as wide as half the tergites’ width, projected anteriorly, CAS scales pale beige ( Figs 109–110 ); aedeagus thicker than in S. probetor and rather short; vesica with two bands of cornuti ( Figs 126 , 136 ). Females with thorax as in S. probetor ; FW (average size: 1.1 cm ; 1.1–1.15 cm ; n=8) moderately bulged ( Figs 15 , 70 ); FWD and FWV submarginal reddish orange band on the submarginal area along the outer margin, continuous from the costal margin, near the apex, to the inner margin at the tornus ( Figs 15–16 , 70 ); FWD mostly dark brown, with a weakly developed translucent white and creamy sickle-shaped spot between the discal cell and the costal margin ( Fig. 70 ); FWD and HWD speckled with faint whitish and reddish orange spots; HWD submarginal band reddish orange ( Fig. 15 ); FWV and HWV spots well developed ( Figs 16 , 79 ); HWV submarginal band well developed and straight and marginal band faint, not outlining ocelli ( Fig. 79 ); sterigma trapezoidal with a triangular median bulge at the ostium; ductus bursae bulged, but membranous near the ostium; bursa copulatrix rather stretched; signum pointed ( Fig. 141 ). 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 113–122. Fourth (left) and fifith (right) tergites of males of Symmachia belonging to the “Probetoriformes” species group. 113–114. S. praxila . 115–116. S. uirassu Dolibaina, Dias & Casagrande sp. nov. . 117–118. S. divisora . 119–120. S. falcistriga . 121–122. S. meyi stat. nov. . Scale bar = 1 mm. Distribution. Widespread in the Amazon basin, Suriname , Guyana , French Guiana and Trinidad , extending to the western areas of the Atlantic Forest, usually in low to mid-elevation, occurring from about sea level to 1100 m in the eastern slopes of the Andes and in Brazil ( Rondônia , Acre , Amazonas, Pará, Ceará , Distrito Federal, Goiás , Mato Grosso , São Paulo and Paraná ). It may occur in Venezuela , Bolivia , Paraguay , and northeastern Argentina and in other Brazilian states. Comments. Symmachia estellina was described based on 38 specimens : the holotype and the allotype , caught in copula, and further 16 males and 20 females paratypes . While the holotype and the allotype are deposited at the MNHN, the paratypes are deposited at the JYG and LCB collections, both of which include additional specimens of S. estellina and multiple species of Riodinidae from French Guiana ( Brévignon & Gallard 1998 ; Gallard 2008 ; Gallard 2017 ). The Jean-Yves Gallard collection (JYG) was recently acquired by the MGCL. As noted above, this species was previously considered as a variation of or misidentified with S. probetor by multiple authors, including Seitz (1913 ; 1917 ) and Brévignon & Gallard (1998) . Only a decade after, Gallard (2008) realized their mistake, described S. estellina , and recognized S. astiama as a junior subjective synonym of S. probetor . Symmachia estellina seems to be the most abundant species of the species group in the Amazon basin and the Guianas, based on the number of specimens in collections and the reports by Seitz (1917) , Brévignon & Gallard (1998) , and Gallard (2008 ; 2017 ). Gallard (2017) reports the species visiting flowers, and, although sympatric with S. probetor in French Guiana , both species do not occur in the exact same place of the forest. Dolibaina et al. (2012) mentioned this species competing for territory with S. probetor at 14:00 h about five meters from the ground, in a small patch of sun of a hilltop at Serra do Divisor National Park, Acre , Brazil . In 2018, the species was also observed in the morning, between 9:30–10:00 h (DRD, pers. obs.). In the relict of Atlantic Forest at the Pico Alto, Guaramiranga, Ceará , Brazil , males of S. estellina are commonly found resting on small shrubs with no more than three meters high, between 10:00–13:00 h; females are infrequent but are found at the same places and time of activity of males (DRD, pers. obs.). Type material. Holotype male of Symmachia estellina Gallard, 2008 with the following labels: / HOLOTYPE / Macouria, [ Cayenne arrondissement,] GUYANE [ French Guiana ], 2 sept[ember]. [19]86, Hermier leg[it] /accouplée [in copula] / no 261 [male sex symbol] J[ean-]Y[ves] GALLARD/ S. estellina [male sex symbol] n[ew].sp[ecies]. J[ean]-.Y[ves]. GALLARD sept[ember]. 2008. ALF [ Association des Lépidoptéristes de France ]/ [voucher number MNHN-EL-H0000455] ( MNHN ) . Allotype female with the following labels: / ALLOTYPE /Macouria, [ Cayenne arrondissement,] GUYANE [ French Guiana ], 2 sept[ember]. [19]86, Hermier leg[it] /accouplée [in copula] / no 261 [female sex symbol]/ S. estellina [female sex symbol] n[ew].sp[ecies]. J[ean]-.Y[ves]. GALLARD sept[ember]. 2008 A.L.F. [ Association des Lépidoptéristes de France ]/ [voucher number MNHN-EL-H0000456] ( MNHN ) . Examined material. COLOMBIA : Norte de Santander— Cucuta , 1♂ , Schaus coll. ( USNM *) . Tolima — Chaparral , 800 m , 1♂ , 9.XI.1974 , S. R . & L. M. Steinhauser leg. ( MGCL ) . ECUADOR : Napo — 4 Km Tena-Pano road, 550 m , 1♂ , 27.IX.1990 , D. H. Ahrenholz leg. ( USNM ) . High hill NW Misahualli Lodge , 610 m , 1♂ , 18.X.2000 , D. H. Ahrenholz leg. ( USNM *) . Sucumbíos — Lumbaqui , 700 m , 1♂ , XII.2005 , M. Simon leg. ( MGCL *) ; 800 m , 1♂ , XII.2005 , M. Simon leg. ( MGCL *); ( Cerro Lumbaqui Norte ) , 950 m , 2♂ , 24.II.2001 , D. H. Ahrenholz leg. ( USNM ) ; 980 m , 1♂ , 2.I.2002 , D. H. Ahrenholz leg . ( USNM ) , 2♂ , 3.I.2002 , D. H. Ahrenholz leg . ( USNM ) , 1♂ , 8.II.2008 , D. H. Ahrenholz leg . ( USNM ) . PERU : Cusco — Cosñipata Valley , Quebrada Quitacálzon , 1100 m , 1♂ , 27.I.2013 , O. Mielke leg. , OM 73.668 ( OM ) . Loreto — Pucallpa , 2♂ , 8.XI.1960 , Jae leg. ( MGCL *) . BRAZIL : Acre— Mâncio Lima ( Parque Nacional da Serra do Divisor , Porção Norte ), 455 m , 2♂ , 10–21.IX.2011 , Dolibaina & Moura leg ., DZ 21.774*, DZ 43.983 ( DZUP ) ; 5♂ , 20–27. VI .2013 , Mielke , Casagrande , Carneiro , Dias & Do- libaina leg ., DZ 43.843, DZ 43.944, DZ 43.974, DZ 44.004, DZ 44.034 ( DZUP ) ; 4♂ , 23–30.VIII.2014 , Miel- ke, Casagrande , Carneiro , Dias , Dolibaina , Siewert & Salik leg. , DZ 36.632, DZ 43.914, DZ 44.014, DZ 44.074 ( DZUP ) ; 2♂ , 7-10,13. X.2018 , Mielke , Casagrande , Dolibaina & Medeiros leg. , DZ 40.800, DZ 40.810 ( DZUP ) . Amazonas — Alvarães ( Nogueira-Alvarães ), 40 m , 1♀ , 7.XII.1987 , Mielke & Casagrande leg. , DZ 43.489 ( DZUP ) . Ceará — Guaramiranga , 900 m , 4♂ , 1♀ , 7–11.I.2012 , Dolibaina & Leite leg. , DZ 23.214, DD 412, DD 438, DD 439, DD 414 ( DZUP , DD); ( Pista de Pouso ), 920 m , 2♂ , 14.VII.2012 , Dolibaina & Lima leg ., DD 431, DD 432; (Pico Alto), 1090 m , 5♂ , 1♀ , 15.VII.2012 , Dolibaina & Lima leg. , DD 413, DD 433, DD 434, DD 435, DD 436, DD 437, 3♂ , 18.I.2016 , Dolibaina leg ., DD 440, DD 441, DD 442, 2♂ , 26.XI.2016 , Dolibaina leg ., DD 415, DD 416, 4♂ , 12.VII.2019 , Dolibaina leg. , DD 585, DD 586, DD 587, DD 588 (DD). Goiás — Cocalzinho de Goiás , 1♂ , 29. V .2006 , Emery leg ., ex-coll. Emery , DZ 43.863 ( DZUP ) . Mato Grosso — Cáceres ( Rio Vermelho , Cel. Rio Branco ), 400 m , 1♀ , 2.VII.1972 , Mielke & Brown leg. , DZ 43.913 ( DZUP ) . Pará — Santo Antônio do Tauá ( RPPN Klagesi ), 40 m , 1♀ , 8.VIII.2003 , P. Jauffret leg. , DZ 43.873 ( DZUP ) ; 1♂ , 17.VII.2004 , P. Jauffret leg. , DZ 43.493 ( DZUP ) ; 1♂ , 1♀ , 4.XI.2004 , P. Jauffret leg. , DZ 43.499, DZ 44.023 ( DZUP ) ; 1♂ , 29.XI.2004 , Mielke , Casagrande & P. Jauffret leg. , DZ 21.814 ( DZUP ) ; 1♂ , 8.XI.2008 , J. Jauffret leg. , DD 417 (DD); 1♀ , 9.XI.2008 , J. Jauffret leg. , DD 418 (DD); 1♀ , 13.XI.2008 , J. Jauffret leg. , DD 419 (DD). Paraná — Diamante do Norte ( Estação Ecológica do Caiuá ) , 1♂ , 16. V .2009 , Dolibaina & Carneiro leg. , DZ 21.804 ( DZUP ) . Rondônia — Cacaulândia ( Fazenda Rancho Grande ) , 1♀ , 6.XI.1989 , G. T . Austin leg. ( MGCL *) ; 1♀ , 9.XI.1989 , J. P. Brock leg . ( MGCL ) ; 1♂ , XI.1991 , V . O. Becker leg ., OM 64.075 ( OM ) ; 1♂ , 1♀ , 14.XI.1991 , G. T . Austin leg. ( MGCL ) ; 1♀ , 18.XI.1991 , G. T . Austin leg . ( MGCL ) ; 1♀ , 19.XI.1991 , J. P. Brock leg . ( MGCL ) ; 1♀ , 13.XI.1992 , D. L. Eiler leg . ( MGCL ) ; 1♀ , 1.VIII.1993 , O. Gomes leg . ( MGCL ) ; 1♀ , 4.IX.1993 , O. Gomes leg . ( MGCL *) ; 1♀ , 13.IV.1994 , O. Gomes leg . ( MGCL ) ; 1♀ , 14.VII.1994 , G. T . Austin leg . ( MGCL ) ; 1♂ , 16.XI.1994 , G. T . Austin leg . ( MGCL ) ; 1♂ , 1. VI .1995 , O. Gomes leg . ( MGCL *) ; 1♂ , 24. VI .1995 , O. Gomes leg . ( MGCL ) ; 1♀ , 1.VII.1995 , O. Gomes leg. ( MGCL ) ; 1♂ , 8.XI.1996 , J. P. Brock leg . ( MGCL *) . Presidente Médici ( Morro Embratel ) , 1♀ , 9.VII.1991 , Mielke & Miers leg. , OM 61.926 ( OM ) . Vilhena , 1♂ , 22. VI .1978 , D. Gifford leg. , ex-coll. Gifford , DZ 23.234 ( DZUP ) . São Paulo — Pereira Barreto , 300m , 1♀ , 4. VI .1972 , H. Ebert leg ., ex-coll. Ebert , DZ 44.086 ( DZUP ) . Poloni , 1♀ , 8. V .1985 , 1♀ , 29. V .1985 , 1♂ , 23. V .1986 , 1♂ , 5. V .1987 , 1♂ , 30. V .1987 , F. M. Pinto leg. , ex-coll. F. M. Pinto ( MZUSP ) . FRENCH GUIANA : Cayenne —Roura (Road to Kaw , km 5) , 1♂ , 30.XII.2011 , M. Benmesbah leg. , DD 420 (DD); (Road to Kaw, km 6), 1♂ , 12.II.2011 , M. Benmesbah leg ., DD 421 (DD); (Road to Kaw, km 7), 6.II.2011 , 12:00 h, 1♀ , 17.II.20112, 13:00 h, 2♀ , 24.XII.2011 , 13:00 h, 2♀ 7.I.2012 , 10:00 h, 1♂ , 7.I.2012 , 12:00 h, 2♂ , 7.I.2012 , 13:00 h, 1♂ , 7.I.2012 , 14:00 h, 1♀ , 3.I.2014 , 1♂ , 17.I.2016 , 13:00 h, 2♂ , 11.I.2018 , 1♀ , 27.I.2018 , C. Brévignon leg. ( LCB ); ( Road to Kaw , km 16) , 1♀ , 24.XII.2009 , C. Brévignon leg. ( LCB ); ( Road to Kaw , km 21) , 1♀ , 3.XI.2006 , C. Brévignon leg. ( LCB ) . Maripasoula , 1♂ , 1♀ , 4. VI .1987 , 1♀ , 3. VI .1987 , C. Brévignon leg. ( LCB ) . Sinnamary , 1♀ , 3.I.1990 , 1♀ , 25.XII.1990 , 1♂ , 30.XII.1990 , 1♂ , 3.I.1991 , 1♂ , 1♀ , 10.II.1991 , 1♂ , 17.II.1991 , C. Brévignon leg. ( LCB ) . Macouria (Larivot) , 1♂ , 1♀ , 10.II.1990 , C. Brévignon leg. ( LCB ) . Matoury ( Mount Matoury ) , 1♂ , 22.XI.2009 ,, 1♂ , 18.I.2014 , 12:00 h, C. Brévignon leg. ( LCB ) . TRINIDAD : no locality , 1♂ , II.1922 ( MGCL *) ; 1♂ , R . Dick leg. ( MGCL ) . Sans Souci — Sangre Grande , 1♂ , V .1976 , F. C. Dick leg . ( MGCL ) ; 1♂ , VII.1978 , F. C. Dick leg . ( MGCL ) ; 1♂ , VIII.1979 , F. C. Dick leg . ( MGCL ) . NO COUNTRY: Amazon, Igaripe, 1♂ , A. Hall leg. ( MGCL ) .