Pachythonina, Seraphim & Freitas & Kaminski, 2018

Dolibaina, Diego Rodrigo, Dias, Fernando Maia Silva, Mielke, Olaf Hermann Hendrik & Casagrande, Mirna Martins, 2024, An overview of the Brazilian Atlantic Forest species of Pachythone Bates (Lepidoptera: Riodinidae), with the description of a new species and an updated checklist of Pachythonina, Zootaxa 5523 (1), pp. 35-48 : 42-46

Taxonomy of Pachythonina

Family- and genus-group taxonomy: Before Seraphim et al. (2018), all species now included in Pachythonina Seraphim, Freitas & Kaminski, 2018 ( Nymphidiini ) were previously included in the incertae sedis section of four forewing radial-veined Riodininae ( Harvey 1987; Callaghan & Lamas 2004). However, due to incomplete taxon sampling, only the genera Pachythone , Pseudonymphidia Callaghan, 1985, Pixus Callaghan, 1985 and Roeberella Strand, 1932 were originally included in the subtribe ( Seraphim et al. 2018). Of these, Pixus and Roeberella are now regarded as junior subjective synonyms of Pachythone ( Hall 2018) and as a subgenus of Apodemia Felder & Felder, 1865 (Emesidini) ( Zhang et al. 2020), respectively. All but two species previously combined with Roeberella were combined with Pachythone ( Zhang et al. 2020) . The genera Lamphiotes Callaghan, 1982 and Machaya Hall & Willmott, 1995 were regarded as junior subjective synonyms of Pachythone by Hall (2018) and Zhang et al. (2021), respectively, and are logically also included in Pachythonina . The genus Minstrellus Hall, 2007 , widely recognized as morphologically related to Pachythone in the “ Pachythone cluster of genera” ( Hall 2007) or even implicitly included in Pachythonina by some authors (e.g., Espeland et al. 2015; Seraphim et al. 2018; Hall 2018; Kaminski et al. 2020, Mota et al. 2020), is here formally included in Pachythonina . Pseudonymphidia was down-ranked to a subgenus of Pachythone ( Zhang et al. 2020) , a result questioned by Pérez-Lachaud et al. (2021), who returned it to its original genus rank. Pseudonymphidia is a replacement name proposed by Callaghan (1985) to the junior homonym Pseudonympha Callaghan, 1982 (not Pseudonympha Wallengren, 1857 ( Lepidoptera : Nymphalidae )). The subtribe was erected based only on molecular data, but the few species in this group with known immature stages present “armored” carnivorous larvae ( Seraphim et al. 2018; Mota et al. 2020; Pérez-Lachaud et al. 2021). Additionally, Hall (2018) comprehensively examined morphologic characters of species of Pachythone and related genera and confirmed that these species almost certainly did not belong in any of the previously established subtribes of Nymphidiini .

Species-group taxonomy: Bates (1868) described Pachythone to include five species newly described by him ( P. erebia Bates, 1868 , P. mimula Bates, 1868 , P. distigma, Bates, 1868 , P. lateritia Bates, 1868 , and P. xanthe Bates, 1868 ); P. erebia was subsequently designated as the type species by Scudder (1875). Four species were added to the genus in the late XIX century, by Hewitson (1873) ( P. pasicles , P. palades , and P. philonis Hewitson, 1873 ) and Godman & Salvin (1878) ( P. gigas Godman & Salvin, 1878 ); and further five species or subspecies in early XX century, by Kaye (1904) ( Cricosoma latertia coccineata Kaye, 1904 ), Stichel (1911, 1926) ( P. thaumaria Stichel, 1911 , P. ignifer Stichel, 1911 , P. conspersa ) and Schaus (1913) ( P. nigriciliata Schaus, 1913 ). Stichel (1911, 1930) provided an overview of the genus and divided Pachythone into three “cohorts”: the Erebiiformes, the Distigmiformes, and the Lateritiformes, based on characters of wing color and pattern. In the next fifty years, only a few species were described in the genus (i.e. P. robusta Lathy, 1932 , Pachytone [sic] lateritia f. radiata Rebillard, 1958, P. barcanti Trite, 1968 ). Only in the late XX and early XXI centuries, further species or subspecies were again described in Pachythone : P. lateritia gourda Gallard, 1998 , P. analuciae Hall, Furtado & DeVries, 1999 , P. sumare Callaghan, 1999 , P. crepusculi , and P. erebia thalassina Vargas & Salazar, 2018 .

Recently, several taxonomic acts greatly modified and increased the number of species in Pachythone : P. bicolor (Godman & Salvin, 1886) , transferred from Pheles Herrich-Schäffer, [1853] (Riodinini) by Dias et al. (2015); P. rubigo ( Bates, 1868) , transferred from Nymphidium (Nymphiidini: Nymphidiina) ( Callaghan & Lamas 2004); P. strati (Kaye, 1925) and its junior subjective synonym Nymphidium corculum Stichel, 1929 (type species of Pixus , by original designation), transferred from Nymphidium and Pixus by Hall (2018), respectively; P. velazquezi (Beutelspacher, 1976) (type species of Lamphiotes , by original designation), transferred from Lamphiotes by Hall (2018); P. obstinata (Hall & Willmott, 1995) (type species of Machaya , by original designation), P. watkinsi ( D’Abrera, 1994) , P. aenigmatica (Rodríguez, Salazar & Constantino, 2011) , directly or implicitly transferred from Machaya by Zhang et al. (2020); and P. lencates , P. flocculus (Brévignon & Gallard, 1993) , P. floccus (Brévignon, 2013) , P. heberti (Jauffret & Jauffret, 2007) and P. marajoara (Jauffret & Jauffret, 2007) , transferred from Roeberella by Zhang et al. (2021).

Pachythone barcanti was recognized as junior subjective synonym of P. erebia erebia , and P. ignifer and P. nigriciliata as subspecies of P. gigas by Callaghan & Lamas (2004); Pachitone [sic] lateritia bourda Gallard, 1998, was recognized as junior subjective synonym of P. lateritia lateritia , and P. mimula and P. thaumaria were recognized as synonyms (although he confused junior and senior synonyms) and as a subspecies of P. xanthe by Gallard (2017); Echenais ochracea Lathy, 1932 , E. striata Lathy, 1932 and E. trinitatis Lathy, 1932 were transferred from Adelotypa Warren, 1895 ( Nymphidiini : Nymphidiina) to Pachythone and recognized as junior subjective synonym of P. conspersa by Hall (2018). Here, one new species is added to the genus ( P. gagarini Dolibaina & Dias , sp. nov.) and P. palades (syn. nov.) is recognized as a junior subjective synonym of P. pasicles .

Emesis clearista Butler, 1871 (type species of Pseudonympha Callaghan, 1982 and Pseudonymphidia, by original designation), Lemonias agave Godman & Salvin, 1886 and Anatole leucogonia Stichel, 1911 were transferred from Pseudonymphidia to Pachythone (Pseudonymphidia) by Zhang et al. (2021); Anatole leucogonia Stichel, 1911 is long recognized as subspecies of Pachythone (Pseudonymphidia) agave ( Callaghan & Lamas 2004) . However, Pérez-Lachaud et al. (2021) returned Pseudonymphidia to the genus rank until further studies addressing the taxonomic status of the group are conducted.

All four species included in Minstrellus ( M. grandis ( Callaghan, 1999) , type species by original designation), P. nivosa (Stichel, 1929) , M. emphatica (Stichel, 1911) and its junior subjective synonym Ematurgina albovata Stichel, 1929 ( Callaghan & Lamas 2004), and M. leucotopus (Stichel, 1911) were originally described in Ematurgina Röber, 1903 ( Nymphidiini : Lemoniadina), but were previously combined with Zelotaea Bates, 1868 ( Nymphidiini : Nymphidiina) ( Hall 2007).

Therefore, Pachythonina now comprises 31 species and six subspecies in three genera: Pachythone , with 25 species and five subspecies, Pseudonymphidia, with two species and one subspecies, and Minstrellus , with four species.

Checklist of Pachythonina

PACHYTHONINA Seraphim, Freitas & Kaminski, 2018 (31 spp., 6 sspp.)

PACHYTHONE Bates, 1868 (25 spp., 5 sspp.)

= Lamphiotes Callaghan, 1982

= Machaya Hall & Willmott, 1995

= Pixus Callaghan, 1982

1) erebia Bates, 1868

a) erebia Bates, 1868

= barcanti Tite, 1968

b) thalassina Vargas & Salazar, 2018 ( Pachytone [sic])

2) robusta Lathy, 1932

3) pasicles Hewitson, (1873)

= palades Hewitson, (1873) , syn. nov.

4) philonis Hewitson, (1873)

5) distigma Bates, 1868

6) analuciae Hall, Furtado & DeVries, 1999

7) lateritia Bates, 1868

a) lateritia Bates, 1868

=f. radiata Rebillard, 1958 ( Pachytone [sic])

= bourda Gallard, 1998 (Pachitone [sic])

b) coccineata ( Kaye, 1904) ( Cricosoma )

8) xanthe Bates, 1868

a) xanthe Bates, 1868

b) mimula Bates, 1868

= thaumaria Stichel, 1911

9) gigas Godman & Salvin, 1878

a) gigas Godman & Salvin, 1878

b) ignifer Stichel, 1911

c) nigriciliata Schaus, 1913

10) obstinata (Hall & Willmott, 1995) ( Machaya )

11) watkinsi ( D’Abrera, 1994) ( Ematurgina )

12) aenigmatica (Rodríguez, Salazar & Constantino, 2011) ( Machaya ) 13) bicolor (Godman & Salvin, 1886) ( Lepricornis )

14) conspersa Stichel, 1926

= ochracea (Lathy, 1932) ( Echenais )

= striata (Lathy, 1932) ( Echenais )

= trinitatis (Lathy, 1932) ( Echenais )

15) sumare Callaghan, 1999

16) rubigo ( Bates, 1868) ( Nymphidium )

17) crepusculi Gallard & Fernandez, 2017

18) strati (Kaye, 1925) ( Nymphidium )

= corculum (Stichel, 1929) ( Nymphidium )

19) velazquezi (Beutelspacher, 1976) ( Nymula )

20) lencates (Hewitson, 1875) ( Lemonias )

= petronia ( Schaus, 1913) ( Echenais )

21) flocculus (Brévignon & Gallard, 1993) ( Roeberella ) 22) floccus (Brévignon, 2013) ( Roeberella )

23) heberti (Jauffret & Jauffret, 2007) ( Roeberella )

24) marajoara (Jauffret & Jauffret, 2007) ( Roeberella ) 25) gagarini Dolibaina & Dias , sp. nov.

PSEUDONYMPHIDIA Callaghan, 1985, repl. name (2 spp., 1 ssp.)

= Pseudonympha Callaghan, 1982 , preocc. (Wallengren, 1857 ( Nymphalidae ))

1) agave (Godman & Salvin, 1886) ( Lemonias )

a) agave (Godman & Salvin, 1886) ( Lemonias )

b) leucogonia (Stichel, 1911) ( Anatole )

2) clearista (Butler, 1871) ( Emesis )

MINSTRELLUS Hall, 2007 (4 spp.)

1) grandis ( Callaghan, 1999) ( Ematurgina )

2) nivosa (Stichel, 1929) (Ematurgia [sic])

3) emphatica (Stichel, 1911) ( Ematurgina )

= albovata Stichel, 1929 ( Ematurgina )

4) leucotopus (Stichel, 1911) ( Ematurgina )