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 probetor ( Stoll, 1782 ) ( Figs 1–4 , 39 , 48 , 57 , 67 , 76 , 85–86 , 103–104 , 123 , 133 , 138 , 151 , 156 ) Papilio probetor Stoll, 1782 , in Cramer. Uitland. Kap. 4 , p. 209, 251; pl. 390, fig. I (d); Surinam—Stoll, 1782, in Cramer. Uitland. Kap. 4, Essai , p. 14— Stichel, 1910 . Gen. Ins. 112A , p. 230. Papilio probator [ sic ]; Jung, 1791. Alphab. Verz. 2 , p. 132. Symmachia probetrix [ sic ]; Hübner, [1819]. Verz. bek. Schmett ., p. 26— Westwood, 1851 , in Doubleday. Gen. Diurn. Lep. 2 , p. 445—Hemming, 1967. Bull. brit. Mus. nat. Hist. Suppl. 9, p. 421— Callaghan & Lamas, 2004 , in Lamas, in Heppner. Atlas Neotrop. Lepid. 5A , p. 157 (emend.). Erycina probetor ; Godart, [1824], in Latreille & Godart. Enc. méth. 9 , p. 590—Verloren, 1837. Cat. syst. Lep. Cramer , p. 145, 210. Symmachia probetor ; Doubleday, 1847 . List Lep. Brit. Mus. 2 , p. 8—Bates, 1868. J. Linn. Soc. 9(40) , p. 438—Herrich- Schäffer, 1868. Corr.-Bl. zool.-min. Ver. Regensburg 22(8/9) , p. 122—Kirby, 1871. Cat. diurn. Lep., p. 314—Scud- der, 1875. Proc. Am. Acad. Arts Sci. 10(2), p. 272. syn. : probetrix —Kirby, 1879. Cat. Coll. Diurn. Lep. Hewitson , p. 198—Staudinger, 1888, in Staudinger & Schatz. Exot. Schmett. 1(20) , p. 253-254—Mengel, 1905. Cat. Erycin. , p. 82— Stichel, 1910 . Berl. ent. Ztschr. 55, p. 81—Stichel, 1924. Deutsch. ent. Zschr. , p. 416— Stichel, 1930 . Lep. Cat. 40 , p. 507—Seitz, 1931. Ent. Rund. 48(24), p. 260; syn. : astiama, belti , championi [“Formas individuales”]—Hemming, 1967. Bull. brit. Mus. nat. Hist., Suppl. 9, p. 421—Lamas, 1994, in Forster et al. RAP Work. Pap. 6, p. 168— Hall & Willmott, 1996 . Lambill. 96(4) , p. 639, 645, 650—Robbins et al. , 1996, in Wilson & Sandoval. Manu, p. 236. - Murray, 2000. J. Res. Lep. 35, p. 59—Salazar & Constantino, 2000. SHILAP Revta. lepid. 28(112), p. 376 [ in part ]— Hall & Harvey, 2002 . Zool. J. Linn. Soc. 136 , 183, 188, 196, fig. 6F (concealed androconial scales); syn . : belti , astiama — Hall & Harvey, 2007 . Trop. Lep. Res. 16(1-2) , p. 7— Gallard, 2008 , in Lacomme & Manil. Lep. Guyane . Tome 3 Rhopal. 2 , p. 27-32, 39, 40, fig. 5 p. 31, fig. [1] (male dorsal), [2] (female ventral, schematic illustration); pl. 1 fig. 3 (male in situ ), pl. 2 fig. 3 (male dorsal, female dorsal and ventral)— Gallard & Attal, 2009 . Lambill. 106(2) , p. 165—Salazar et al. , 2010. Bol. Cient. Mus. Hist. Nat. U. Caldas 14(1) , p. 172— Dolibaina et al. , 2012 . Insecta Mundi 249, p. 4-7; figs 7-8 (male dorsal and ventral), fig. 39 (male forewing dorsal), fig. 42 (male genitalia)—Nielsen & Salazar, 2015. Bol. Cient. Mus. Hist. Nat. U. Caldas 18(2) , p. 221— Gallard, 2017 . Les Riod. Guyane , p. 75, 165; fig. 10A (female ventral, schematic illustration); pl. 15 fig. 4 (male dorsal, female dorsal and ventral)— Sambhu & Nankishore, 2018 . Zoot. 4371(1), p. 162. Symmachia probetor probetor ; Stichel, 1910 . Gen. Ins. 112A , p. 234, 420—Stichel, 1923. Zeit. wiss. Ins.-Biol. 18 , p. 283— Stichel, 1924. Deutsch. ent. Zschr.. , p. 417— Stichel, 1930 . Lep. Cat. 40 , p. 507—Kaye, 1940. Trans. R. Ent. Soc. Lond. 90(21) , 557— Callaghan & Lamas, 2004 , in Lamas (ed.), in Heppner. Atlas Neotrop. Lepid. 5A , p. 157; syn : astiama, belti , praxila , probetrix — Gallard, 2008 , in Lacomme & Manil. Lep. Guyane . Tome 3 Rhopal. 2 , p. 29—Palo Jr., 2017. Borb. Bras. 1 , p. 715. Symmachia probetor form astiama Seitz, 1913 . Gross-Schmett. Erde 5 , pl. 128g [3] (male dorsal)— Seitz, 1917 , p. 682. Symmachia probetor belti form astiama ; Stichel, 1924. Deutsch. Ent. Zschr.. , p. 417— Stichel, 1930 . Lep. Cat. 40 , p. 508. Symmachia nr. praxila ; Emmel & Austin 1990. Trop. Lep. Res. 1(1) , p. 9. Symmachia astiama ; Brévignon & Gallard, 1998 . Lambill. 98(3) , p. 308, 316, 318; figs 27-30 (male and female, dorsal and ventral); fig. 3- 4 p. 320 (male genitalia)— Hall & Harvey, 2002 . Zool. J. Linn. Soc. 136 , p. 196; syn. : belti — Gallard, 2008 , in Lacomme & Manil. Lep. Guyane . Tome 3 Rhopal. 2 , p. 29-30. Symmachia probator [ sic ]; Moreno et al. , 1998. Marip. Ecuad. , p. 157. Diagnosis. Symmachia probetor is sympatric throughout its distribution with S. estellina , S. divisora , S. falcistriga and S. meyi ( Figs 151–153 ). Males are most similar to S. belti , S. championi , S. estellina and S. atlantica ( Figs 1–2 ), but females are quite similar to most species of the “Probetoriformes” species group ( Figs 3–4 ). Males of S. probetor with frons covered by compact, black mixed with orange scales; patagia yellow; thorax black dorsally including tegulae; FW (average size: 1.03 cm ; 0.9–1.2 cm ; n=15) only slightly bulged; bulged area about 50% of the FW length; end of bulge aligned to the end of M 2 ( Fig. 57 ); FWD black, with a dark green sheen covering most of the wing, without any spots along the costal margin ( Figs 1 , 57 ); dark green sheen absent roughly on the submedial areas from the inner margin to the discal cell and along the outer margin ( Fig. 57 ); HWD black, with a red band along the costal margin and a dark green sheen patch near the tornus ( Fig. 1 ); abdomen uniformly black including tip, paler ventrally, with a whitish stripe on the posterior margin of each sternum ( Figs 85–86 ); CAS of the tergites 4 and 5 as long as 1/7 of the tergites’ length and as wide as half the tergites’ width, projected anteriorly, CAS scales pale beige ( Figs 103–104 ); aedeagus thinner and longer than in other species of the “Probetorifomes” species group ( Figs 123 , 133 ); vesica without cornutus ( Figs 123 , 133 ). Females can be distinguished by the thorax reddish-spotted dorsally; FW (average size: 1.05 cm ; n=1) slightly bulged ( Fig. 67 ); FWD and FWV submarginal reddish orange bands along the costal margin not extending beyond M 2 –M 3 ( Figs 3 , 67 ); FWD and HWD mostly dark brown, speckled with very faint whitish and reddish orange spots ( Figs 3 , 67 ); HWD submarginal band very faint; FWV and HWV spots well-developed ( Figs 4 , 76 ); HWV submarginal band wavy and outlining a number of ocelli with the marginal band ( Figs 4 , 76 ); sterigma trapezoidal with a triangular median bulge at the ostium; ductus bursae with a sclerotized ring near the ostium, bursa copulatrix and signum unknown (broken in the single available specimen for dissection) ( Fig. 138 ). Distribution. Widespread in the Amazon basin, the Guianas and Trinidad , usually in low to mid-elevation, occurring from about sea level to 1230 m in the eastern slopes of the Andes, in Brazil ( Acre , Rondônia ), Peru , Colombia , Ecuador , Guyana , Surinam ( type locality), French Guiana and Trinidad ( Fig. 151 ). It probably occurs also in Bolivia and Venezuela . 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 21–38. Species of Symmachia belonging to the “Probetoriformes” species group. 21–24. S. praxila . 21–22. Male from Joinville, Santa Catarina, Brazil (DZ 43.995 (DZUP)). 23–24. Female from Rio Natal, São Bento do Sul, Santa Catarina, Brazil (OM 67.541). 25–26. S. uirassu Dolibaina, Dias & Casagrande sp. nov. male holotype (DZUP). 27–30. S. divisora . 27–28. Male from Parque Nacional da Serra do Divisor, Mâncio Lima, Acre, Brazil (DZ 43.479 (DZUP)). 29–30. Female from Fazenda Rancho Grande, Cacaulândia, Rondônia, Brazil (MGCL). 31–34. S. falcistriga . 31–32. Male from Parque Nacional da Serra do Divisor, Mâncio Lima, Acre, Brazil (DZ 43.509 (DZUP)). 33–34. Female, same locality as male (DZ 43.965 (DZUP)). 35–38. S. meyi stat. nov. . 35–36. Male from Chemin de la Sourçe, Cacao, French Guiana (SF). 37–38. Female from Route de l’Este, French Guiana (SF). Comments. Papilio probetor Stoll, 1782 was described based an unknown number of males collected in Surinam , from Stoll’s personal collection (“Celui-ci avec la Fig. C, D, est de ma Collection”) ( Stoll 1782 ). Even though a number of specimens probably used by Stoll in his species descriptions were located at the NHMUK and NBC, the origin and fate of his personal collection remain unknown and it has been assumed to be lost or destroyed (Vane- Wright 1975; Chainey 2005 ). Therefore, a neotype is here designated so as to reduce present and future uncertainty to a minimum and to make clear the taxonomic status of S. probetor , as distinguished here from its most similar congeners. The neotype , collected in the same ecoregion (Guianan lowland moist forests sensu Olson et al. 2001 ) and relatively near to the original type locality in Surinam , compares closely with Stoll’s illustration ( Stoll 1782 , pl. 390, fig. I). To ensure recognition of the specimen designated as neotype , the specimen deposited at the DZUP will be labeled accordingly. Symmachia probetrix Hübner, [1819] was introduced as a replacement for the name Papilio probetor Stoll, 1782 , an unjustified emendation not in prevailing usage ( ICZN 1999 , Art. 33.2). Therefore, it is a junior objective synonym of S. probetor . Symmachia probetor form astiama Seitz, 1913 description is based on the illustration of a male in dorsal view, while the text was published later ( Seitz 1917 ) and mentions the name as a “form. nov.” (even though clearly denoting a subspecific entity) based on an unknown number of male specimens from “ Guiana and the Amazon”. Callaghan & Lamas (2004) follow Griffin (1936) and recognize the date of the publication of the plate 128 with the description as 1914, when the plate was received at the British Museum ( i.e. 27.I.1914 ; Griffin 1936: 251 ). Nevertheless, the correct date of publication of the plate 218 is 1913, as acknowledged by Stichel (1930) . Seitz (1913 ; 1917 ) correctly recognized two different sympatric phenotypes, one with one white spot at the costa on the FWD (later described as S. estellina by Gallard (2008)) , and another, without such spot ( S. probetor ). However, he wrongly described a new species to the already described phenotype without the white spot. Therefore, S. probetor form astiama is a junior subjective synonym of S. probetor . A male specimen of S. probetor form astiama deposited at the SMF from Seitz’s type series is here designated lectotype so as to reduce present and future uncertainty to a minimum. Gallard (2008 ; 2017 ) reports that S. probetor flies, in French Guiana , during end of the morning from 10:00– 11:00 h, visiting flowers in January, June and December. Similar to other species of the genus, the species mark territory on hilltops and light gaps, flying fast and erratically two or three meters from the ground, frequently perching on the underside of leaves ( Gallard 2008 , 2017 ; Dolibaina et al. 2012 ; DRD and FMSD, pers. obs.;). In Mâncio Lima, Acre , Brazil , the species was observed earlier in the morning, between 9:00–9:30 h (DRD, pers. obs.). Type material. Male neotype of Papilio probetor Stoll, 1782 , here designated, with the following labels: / NEOTYPUS / NEOTYPUS Papilio probetor Stoll, 1782 Dias, Dolibaina, Mielke & Casagrande det. 2019 /Montagne des Chevaux[, Montsinéry-Tonnegrande, Cayenne arrondissement] ( R [ou]te. de Cayenne à Regina [Route N2]) 9 novembre [november] 2003 Coll[ection]. Diringer /DZ 21.834 / ( DZUP ). Male lectotype of Symmachia probetor form astiama Seitz, 1913 , here designated, with the following labels: /Coll[ection]. A[dalbert]. Seitz / SYNTYPE ? Symmachia probetor f. astiama Seitz G. Lamas 2000 / Surinam V .-IX. Fruhstorfer/ ( SMF ), and the following labels will be added: / LECTOTYPUS / LECTOTYPUS Symmachia probetor f. astiama Seitz, 1913 Dias, Dolibaina, Mielke & Casagrande det. 2019 /. Examined material. BRAZIL : Acre— Mâncio Lima ( Parque Nacional da Serra do Divisor , Porção Norte ), 455 m , 1♂ , 10–21.IX.2011 , Dolibaina & Moura leg ., DZ 21.483* ( DZUP ) ; 8♂ , 20–27. VI .2013 , Mielke , Casa- grande, Carneiro , Dias & Dolibaina leg ., DZ 43.349, DZ 43.853, DZ 43.883, DZ 43.893, DZ 43.903, DZ 43.923, DZ 43.933, DZ 44.013* ( DZUP ) ; 4♂ , 23–30.VIII.2014 , Mielke , Casagrande , Carneiro , Dias , Dolibaina , Siewert & Salik leg. , DZ 36.602, DZ 43.953*, DZ 43.973, DZ 44.053 ( DZUP ) ; 2♂ , 7-10,13. X.2018 , Mielke , Casagrande , Dolibaina & Medeiros leg. , DZ 40.780, DZ 40.790 ( DZUP ) . Rondônia — Cacaulândia ( Fazenda Rancho Grande ), 1♀ , 21.XI.1991 , G. T . Austin leg. ( MGCL ) . ECUADOR : Napo — El Capricho (28 Km Tena-Puyo Road ), 800 m , 1♂ , 5.I.2007 , R . Busby & D. H. Ahrenholz leg. ( USNM ) ; 4 Km Tena-Pano road Km 4, 550 m , 1♀ , 24.IX.1990 , D. H. Ahrenholz leg. ( USNM *) ; Tena-Loreto road Km 49, 1350 m, 1♂ , X.2005 , I. Aldas & D. H. Ahrenholz leg. ( USNM ) ; Jatun Sacha Biological Reserve , 450 m , 3♂ , 11.XI.1992 , D. H. Ahrenholz leg. ( USNM ) . Pastaza — Shell (9 Km S ), 1000 m , 1♂ , 17.IX.1990 , 1♂ , 18.IX.1990 , 3♂ , 26.IX.1990 , all D. H. Ahrenholz leg . ( USNM *) ; Puyo-Arahuno road Km 37, 850 m , 2♂ , 10.I.2006 , D. H. Ahrenholz leg. ( USNM ) . Sucumbíos — Lumbaqui , ( Cerro Lumbaqui Norte ), 950 m , 1♂ , 24.II.2001 , D. H. Ahrenholz leg. ( USNM ) ; 1♂ , 27.II.2001 , D. H. Ahrenholz leg . ( USNM ); ( Lumbaqui - Puerto Libre rd. Km 7) , 900 m , 1♂ , 24.II.2005 , R . Busby & D. H. Ahrenholz leg. ( USNM ) ; 800 m , 1♂ , II.2008 , M. Simon leg. ( MGCL ) . FRENCH GUIANA : Cayenne — Cayenne ( Cayenne to Régina , Montagne des Chevaux ), 1♂ *, 9.XI.2003 , Diringer leg. , neotype designed above ( DZUP ) . Roura ( Galion ), 1♂ , 7.VII.1990 , C. Brévignon leg. ( USNM *) ; 2♂ , 15.XII.1987 , 1♂ , 8.X.1988 , 1♂ , 11.VII.1989 , 3♂ 11.VIII.1989 , 2♂ , 13. VI .1990 , 1♂ , 23. VI .1990 , C. Brévignon leg. ( LCB ); ( Road to Kaw , km 7) 1♂ 13.I.2018 , 11:00 h, 1♀ 7.I.2012 , 1♀ , 8.I.2012 , 11:00 h, C. Brévignon leg. ( LCB ); ( Road to Kaw , km 16) 1♀ 25.I.2009 , 13:00 h h, C. Brévignon leg. ( LCB ) . Montsinéry-Tonnégrande , 10 m , 1♂ , M. Benmesbah leg. , DD 422 (DD) . Sinnamary , 1♀ , 7.II.1992 , J.-Y. Gallard leg. ( USNM *) ; 1♂ 4.I.1991 , C. Brevignon leg. ( LCB ) . GUYANA : Potaro-Siparuni — Iwokrama Rainforest Reserve ( Iwokrama Mountain ), 810–1040 m , 2♂ , 28.III–1.IV.2001 , S. Fratello leg. ( USNM ) . Upper Takutu-Upper Essequibo — Acarai Mountains (ridge Sipu River), 830–1230 m , 5♂ , 6–9.XI.2000 , S. Fratello et al. leg. ( USNM *) . Kanuku Mountains ( Nappi Mountain ), 900–1100 m , 1♂ , 21.II–10.III.1999 , S. Fratello et al. leg. ( USNM ) . Two Hat Mountain ( East Kanuku , South Rupununi , South slope summit), 750 m , 3♂ , 23–28.IX.2000 , S. Fratello et al. leg. ( USNM ) .