Volume 9 Number 3 29 May 2021

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

ISSN 2643-4776 (print) / ISSN 2643-4806 (online)

Genomics-guided refinement of butterfly taxonomy

Jing Zhang2?, Qian Cong”, Jinhui Shen23, Paul A. Opler’ and Nick V. Grishin'?*

"Howard Hughes Medical Institute, Departments of *Biophysics and *Biochemistry, and “Eugene McDermott Center for

Human Growth & Development, University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX

75390-9050, USA; °Department of Agricultural Biology, Colorado State University, Fort Collins, CO 80523-1177, USA.

“Corresponding author: grishin@chop.swmed.edu

ABSTRACT. Continuing with comparative genomic exploration of worldwide butterfly fauna, we use all protein-

coding genes as they are retrieved from the whole genome shotgun sequences for phylogeny construction. Analysis of these

genome-scale phylogenies projected onto the taxonomic classification and the knowledge about butterfly phenotypes suggests

further refinements of butterfly taxonomy that are presented here. As a general rule, we assign most prominent clades of

similar genetic differentiation to the same taxonomic rank, and use criteria based on relative population diversification and the

extent of gene exchange for species delimitation. As a result, 7 tribes, 4 subtribes, 14 genera, and 9 subgenera are proposed as

new, 1.e., in subfamily Pierinae Swainson, 1820: Calopierini Grishin, trib. n. (type genus Ca/opieris Aurivillius, 1898); in

subfamily Riodininae Grote, 1895: Callistiumini Grishin, trib. n. (type genus Callistium Stichel, 1911); in subfamily

Nymphalinae Rafinesque, 1815: Pycinini Grishin, trib. n. (type genus Pycina Doubleday 1849), Rhinopalpini Grishin, trib. n.

(type genus Rhinopalpa C. & R. Felder 1860), Kallimoidini Grishin, trib. n. (type genus Kallimoides Shirézu & Nakanishi

1984), Vanessulini Grishin, trib. n. (type genus Vanessula Dewitz 1887), and Doleschalliaini Grishin, trib. n. (type genus

Doleschallia C. & R. Felder 1860); in tribe Mesosemiini Bates, 1859: Eunogyrina Grishin, subtrib. n. (type genus Eunogyra

Westwood, 1851); in tribe Satyrini Boisduval, 1833: Callerebiina Grishin, subtrib. n. (type genus Callerebia Butler, 1867),

Gyrocheilina Grishin, subtrib. n. (type genus Gyrocheilus Butler, 1867), and Calistina Grishin, subtrib. n. (type genus Calisto

Hubner, [1823]); in subfamily Euselasiinae Kirby, 1871: Pe/o/asia Grishin, gen. n. (type species Eurygona pelor Hewitson,

[1853]), Myselasia Grishin, gen. n. (type species Eurygona mys Herrich-Schaffer, [1853]), Eurylasia Grishin, gen. n. (type

species Eurygona euryone Hewitson, 1856), Maculasia Grishin, gen. n. (type species Euselasia albomaculiga Callaghan,

1999), and Eugelasia Grishin, gen. n. (type species Eurygona eugeon Hewitson, 1856); in subtribe Mesosemiina Bates, 1859:

Ectosemia Grishin, gen. n. (type species Papilio eumene Cramer, 1776) and Endosemia Grishin, gen. n. (type species Papilio

ulrica Cramer, 1777); in tribe Symmachiini Reuter, 1896: Tigria Grishin, gen. n. (type species Mesene xypete Hewitson, 1870)

and Asymma Grishin, gen. n. (type species Symmachia virgatula Stichel, 1910); in tribe Riodinini Grote, 1895: Putridivora

Grishin, gen. n. (type species Charis argyrea Bates, 1868), Chadia Grishin, gen. n. (type species Charis cadytis Hewitson,

1866), Inkana Grishin, gen. n. (type species Charis incoides Schaus, 1902), and Oco Grishin, gen. n. (type species Symmachia

ocellata Hewitson, 1867); in subtribe Zabuellina Seraphim, Freitas & Kaminski, 2018: Teenie Grishin, gen. n. (type species

Calydna tinea Bates, 1868); Boreographium Grishin, subgen. n. (type species Papilio marcellus Cramer, 1777, parent genus

Eurytides Hubner, [1821]), Esperourus Grishin, subgen. n. (type species Papilio esperanza Beutelspacher, 1975, parent genus

Pterourus Scopoli, 1777), Hyppasonia Grishin, subgen. n. (type species Papilio hyppason Cramer, 1775, parent genus

Heraclides Hubner, [1819]), Sisvmbria Grishin, subgen. n. (type species Pieris sisvmbrii Boisduval, 1852, parent genus Pontia

[Fabricius], 1807), Greenie Grishin, subgen. n. (type species Thecla sheridonii |sic| Edwards, 1877, parent genus Callophrys

Billberg, 1820), Magda Grishin, subgen. n. (type species Erebia magdalena Strecker, 1880, parent genus Erebia Dalman,

1816), and in genus Eresia Boisduval, 1836: Notilia Grishin, subgen. n. (type species Eresia orthia Hewitson, 1864), Levinata

Grishin, subgen. n. (type species Eresia levina Hewitson, 1872), and /thra Grishin, subgen. n. (type species Phyciodes ithra

Kirby, 1900). Furthermore, we resurrect 6 genera, change the rank of 36 currently used genera to subgenus, synonymize 3

subtribes, 42 genera or subgenera, assign 3 genera to tribes and subtribes, and transfer 34 additional species to genera different

from those these taxa are presently assigned to, present evidence to support 7 taxa as species instead of subspecies, and 1 taxon

as a Subspecies instead of species. Namely, the following taxa are valid genera: Terias Swainson, 1821 (not in Eurema Hubner,

[1819]), Erythia Hubner, [1819] and Marmessus Hubner, [1819] (not in Euselasia Hubner, [1819]), Eucorna Strand, 1932 (not

in Voltinia Stichel, 1910), Cremna Doubleday, 1847 (not in Napaea Hubner, [1819]), and Hallonympha Penz & DeVries, 2006

(not in Zabuella Stichel, 1911). The following taxa are best treated as subgenera: Zegris Boisduval, 1836 of Anthocharis

Boisduval, Rambur, [Duménil] & Graslin, [1833]; Baltia Moore, 1878 and Pontieuchloia Verity, 1929 of Pontia [Fabricius],

1807; Phrissura Butler, 1870 of Appias Hubner, [1819]; Saletara Distant, 1885 of Catophaga Hubner, 1819; Leodonta Butler,

1870 of Pereute Herrich-Schaffer, 1867; Takashia M. Okano & T. Okano, 1985 of Polycaena Staudinger, 1886; Corrachia

Schaus, 1913 of Styx Staudinger, 1876; Jonotus Hall, 2005 and Voltinia Stichel, 1910 of Cremna Doubleday, 1847;

Hermathena Hewitson, 1874 of Ithomiola C. & R. Felder, 1865; Lucillella Strand, 1932 of Esthemopsis C. & R. Felder, 1865;

Mesenopsis Godman & Salvin, 1886 and Xenandra C. & R. Felder, 1865 of Symmachia Hubner, [1819]; Pirascca J. Hall &

Willmott, 1996 of Prerographium Stichel, 1910; Imelda Hewitson, 1870 of Echenais Hubner, [1819]; Calicosama J. Hall &

Harvey, 2001 of Behemothia Hall, 2000; Polygrapha Staudinger, 1887 and Fountainea Rydon, 1971 of Anaea Hubner, [1819];

Siderone Hubner, [1823] and Phantos Dias, 2018 of Zaretis Hubner, [1819]; Harsiesis Fruhstorfer, 1911 of Platypthima

Rothschild & Jordan, 1905; Vila Kirby, 1871 of Biblis Fabricius, 1807; Diaethria Billberg, 1820 and Perisama Doubleday,

1849 of Callicore Hubner, [1819]; Antigonis C. Felder, 1861 of Haematera Doubleday, 1849; Asterope Hubner, [1819], Nica

Hubner, [1826], Peria Kirby, 1871, and Callicorina Smart, 1976 of Temenis Hubner, [1819]; Anthanassa Scudder, 1875,

Castilia Higgins, 1981, Telenassa Higgins, 1981, Dagon Higgins, 1981, and Janatella Higgins, 1981 of Eresia Boisduval,

1836; and Wallengrenia Berg, 1897 of Polites Scudder, 1872. The following taxa are junior subjective synonyms: Maniolina

Grote, 1897 of Erebiina Tutt, 1896; Melanargiina Wheeler, 1903 of Satyrina Boisduval, 1833; Phyciodina Higgins, 1981 of

Melitaeina Herrich-Schaffer, 1843; Cunizza Grote, 1900 of Hesperocharis C. Felder, 1862; Religuia Ackery, 1975 of Pontia

[Fabricius], 1807; Tatochila A. Butler, 1870, Piercolias Staudinger, 1894, Hypsochila Ureta, 1955, Theochila W. D. Field,

1958, Pierphulia W. D. Field, 1958, and Infraphulia W. D. Field, 1958 of Phulia Herrich-Schaffer, 1867; Mesapia Gray, 1856

of Aporia Hubner, [1819]; Catasticta Butler, 1870 of Archonias Hubner, 1827; Sandia Clench & P. Ehrlich, 1960 and Xamia

Clench, 1961 of Jncisalia Scudder, 1872; Hades Westwood, 1851 of Methone Doubleday, 1847; Semomesia Westwood, 1851,

Mesophthalma Westwood, 1851, Perophthalma Westwood, 1851 and Leucochimona Stichel, 1909 of Mesosemia Hubner,

[1819], Xynias Hewitson, 1874 of Mesenopsis Godman & Salvin, 1886; Stichelia J. Zikan, 1949 of Symmachia Hubner,

[1819]; Chimastrum Godman & Salvin, 1886 of Mesene Doubleday, 1847; Alethea Nielsen & Salazar, [2018] of Pirascca J.

Hall & Willmott, 1996; Panaropsis J. Hall, 2002 of Pterographium Stichel, 1910; Comphotis Stichel, 1910 of Phaenochitonia

Stichel, 1910; Colaciticus Stichel, 1910 of Baeotis Hubner, [1819]; Nahida Kirby, 1871 of /thomeis Bates, 1862; Machaya

Hall & Willmott, 1995 of Pachythone Bates, 1868; Percnodaimon Butler, 1876 and Erebiola Fereday, 1879 of Argyrophenga

Doubleday, 1845; Hestinalis Bryk, 1938 of Mimathyma Moore, 1896; Catacore Dillon, 1948 of Diaethria Billberg, 1820;

Mesotaenia Kirby, 1871 and Orophila Staudinger, 1886 of Perisama Doubleday, 1849; Paulogramma Dillon, 1948 of

Catagramma Boisduval, 1836; Panacea Godman & Salvin, 1883 of Batesia C. Felder & R. Felder, 1862; Napeocles Bates,

1864 of Siproeta Hubner, [1823]; Texola Higgins, 1959 and Dymasia Higgins, 1960 of Microtia H. Bates, 1864; Tisona

Higgins, 1981 of Ortilia Higgins, 1981; Abananote Potts, 1943 and Altinote Potts, 1943 of Actinote Hubner, [1819]; Episcada

Godman & Salvin, 1879 of Ceratinia Hubner, 1816; and Appia Evans, 1955 of Pompeius Evans, 1955. The following genera

are placed in taxonomic hierarchy: Prestonia Schaus, 1920 belongs to Euremini Grote, 1898; Petrocerus Callaghan, 1979

belongs to Theopina Clench, 1955; and Paralasa Moore, 1893 belongs to Ypthimina Reuter, 1896. The following taxa are

distinct species rather than subspecies (of species shown in parenthesis): Pyrisitia westwoodii (Boisduval, 1836) (not Pyrisitia

dina (Poey, 1832)), Biblis aganisa Boisduval, 1836 (not Biblis hyperia (Cramer, 1779)), Phystis variegata (Rober, 1913) and

Phystis pratti (A. Hall, 1935) (not Phystis simois (Hewitson, 1864)), Phocides batabano (Lucas, 1857) and Phocides bicolora

(Boddaert, 1783) (not Phocides pigmalion (Cramer, 1779)), Lobotractus mysie (Dyar, 1904) (not Lobotractus valeriana (P16tz,

1881)). Nahida coenoides (Hewitson, 1870) is conspecific with /thomeis aurantiaca H. Bates, 1862. Additional new and

revised combinations are: Teriocolias deva (E. Doubleday, 1847), Teriocolias reticulata (A. Butler, 1871), Hesperocharis

leucothea (Molina, 1782), Methone euploea (Hewitson, [1855]), Methone eucerus (Hewitson, 1872), Methone hypophaea

(Godman & Salvin, 1878), Methone eubule (R. Felder, 1869), Methone onorata (Hewitson, 1869), Methone authe (Godman,

1903), Methone dolichos (Staudinger, [1887]), Methone baucis (Stichel, 1919), Methone eucrates (Hewitson, 1872), Napaea

danforthi A. Warren & Opler, 1999, Napaea dramba (J. Hall, Robbins & Harvey, 2004), Napaea sanarita (Schaus, 1902),

Napaea agroeca Stichel, 1910, Napaea tumbesia J. Hall & Lamas, 2001, Napaea umbra (Boisduval, 1870), Napaea phryxe (C.

& R. Felder, 1865), Napaea cebrenia (Hewitson, [1873]), Napaea loxicha (R.G. Maza & J. Maza, 2016), Napaea maya (J.

Maza & Lamas, 2016), Napaea necaxa (R.G. Maza & J. Maza, 2018), Napaea totonaca (R.G. Maza & J. Maza, 2016), Mesene

aeolia (Bates, 1868), Pterographium hypochloris (Bates, 1868), Phaenochitonia florus (Fabricius, 1793), Ourocnemis

carausius (Westwood, 1851), Ourocnemis principalis (Hopffer, 1874), Ourocnemis renaldus (Stoll, 1790), and Ourocnemis

aerosus (Stichel, 1924), Hallonympha maculosa (Bates, 1868), Exoplisia aphanis (Stichel, 1910), Phystis fontus (A. Hall,

1928), Phocides batabano okeechobee (Worthington, 1881), and Phocides batabano batabanoides (W. Holland, 1902).

Finally, we confirm the combination Zabuella castanea (Prittwitz, 1865) and find Pyrgus centaureae dzekh Gorbunov, 2007 as

a new subspecies for North America.

Key words: taxonomy, classification, genomics, phylogeny, biodiversity.

ZooBank registration: http://zoobank.org/5027A DA7-E67E-415E-AE9C-D8E282AF942D

INTRODUCTION: METHODS AND CONCEPTS

Genome-scale DNA analysis opens a new dimension in exploration of butterfly taxonomy and

offers a promise of more objective and internally consistent classification firmly grounded in evolutionary

considerations and reliable phylogenies (Allio et al. 2019; Li et al. 2019; Zhang et al. 2019a; Zhang et al.

2019b; Zhang et al. 2020). Many conclusions we arrive at challenge current taxonomy based largely on

phenotypes: phylogenomic analysis shows that some taxa are not monophyletic, while others are either

too broad or too narrow in terms of genetic diversification compared to taxa of the same rank. Here, the

resultant taxonomic hypotheses are formalized and corresponding name changes are proposed.

Classification concepts and methods employed in this work do not differ from those in our

previous studies, where they were explained in more detail (Zhang et al. 2019c; Zhang et al. 2020). Here,

they are simply applied to additional taxonomic groups of butterflies. In brief, we do not amplify any

specific gene markers, but instead sequence all DNA molecules in a specimen that pass the procedure of

genomic library construction (Li et al. 2019; Zhang et al. 2019a). Therefore, all genes of a butterfly are

sequenced together. On the one hand, this approach allows us to obtain complete genomes of butterflies.

On the other hand, it enables DNA sequencing of century-old specimens with degraded DNA that may be

too short for the amplification procedure of standard gene markers (Cong et al. 2021). For fresher

specimens, the method produces nearly 99% of all genes, but for older specimens it could be only 10%

due to DNA degradation. Nevertheless, even 10% of about 15,000 total genes that constitute the gene set

of most butterflies (Zhang et al. 2019d) provides more information than a study based on amplification of

several genes. As a result, genome-scale phylogenetic trees are not biased by gene marker selection, but

represent the entire organism, and therefore enable us to judge more accurately about its evolution.

For phylogenetic analysis, all sequences are aligned to the closest reference genome and from this

alignment, we select positions in exons with a gap fraction less than 0.4: 1.e., such position in the

alignment is not a gap in more than 40% of specimens. We then compute a reference tree using IQ- TREE

version 1.6.8 with model GIR+G (Minh et al. 2020) from an alignment with 300K such positions

randomly sampled. Then, 100 partitions consisting of 30K randomly sampled such positions are generated

to construct 100 trees for estimating the confidence of each node in the reference tree. The support values

are assigned to each node (and shown in the trees below) by mapping trees from 100 partitions to the

reference tree using sumtrees.py script from the DendroPy package (Sukumaran and Holder 2010). The

same procedure is repeated for the Z chromosome tree, which is only composed of Z-linked positions.

The Z-linked exons are inferred by aligning exons of the closest genome reference to known Heliconius

melpomene Z chromosome (Davey et al. 2016) using tRLASTn (Gertz et al. 2006). Mitogenome tree is

based on all protein-coding regions and inferred by IQ-TREE with automatic model estimation and 1000

ultrafast bootstrap replicates. COI dendrograms are constructed with BioNJ (Gascuel 1997) using

Phylogeny.fr website (Dereeper et al. 2008) with the default model and 200 bootstrap replicates. DNA

characters are obtained using the approach we developed to increase their robustness to missing species as

described in our previous work (Cong et al. 2019b; Zhang et al. 2019e). The character states are given as

abbreviations: e.g., cne703.2.8:A1414C means position 1414 in exon 8 of gene 2 from scaffold 703 of

Calephelis nemesis (cne) reference genome (Cong et al. 2017) is C, changed from A in the ancestor; or

cne5129.1.5:A833A (not G), which means that position 833 in exon 5 of gene 1 on scaffold 5129 is

occupied by the ancestral base pair A, which was changed to G in the sister clade (it is not G in the

diagnosed taxon). We also use Calycopis cecrops (cce) (Cong et al. 2016) and Heliconius melpomene

(hm) (Davey et al. 2016) genomes as references. For H. melpomene, the abbreviation is like hm2009277-

RA.13:T2076C, where hm2009277-RA is the protein ID and 13 is exon number.

The resulting phylogenetic trees are inspected visually and compared with the current butterfly

classification. Close attention is paid to family-group and genus-group taxa that are not monophyletic in

the trees, 1.e., are not the groups of species that consists of all descendants of their common ancestor.

First, identification is checked for specimens that disrupt monophyly of higher level taxa by inspection of

sequenced specimens, their photographs or genitalia when necessary. Then, the quality of sequence

datasets is analyzed in detail for the possibility of insufficient coverage and contamination, both from

other butterfly specimens or bacteria and fungi. When we are convinced about the identifications and

appropriate data quality, suggestions are made as how to restore monophyly of these taxa, either by

combining a number of taxa into one that is monophyletic, by transferring species between the taxa, or by

splitting the non-monophyletic taxon into several monophyletic taxa. The names of these taxa follow the

placement of their type genera or species: the clade with the type species carries its genus name. Clades

that do not contain type genera or type species for any of the available names are named as new taxa.

In decisions about taxa and their ranks we rely on their prominence in phylogenetic trees and

internal consistency of definition based on _ genetic Present Proposed

diversification. The discussion below uses genus rank as a fgenust_ Classification classification

an example, but similar logic applies to other ranks. Our | _cssifcation Genus 1 | | Genus 1

criteria for genera were explained previously (Li et al.

2019; Zhang et al. 2020). Traditionally, genera were

defined using arbitrary and subjective criteria based on

prominent phenotypic characters that make a certain

group of species stand out from other similar groups of ee

Species. We attempt to define genera more objectively —

based on the patterns of branch lengths in genome-scale y

phylogenetic trees, while keeping in mind agreement with promineAt branch ca

the current classification. Most prominent tree branches Fig. 1. Bes anne re eenee mheidiversity:

near the origin of tribes and subtribes are typically

defined as genera. By "prominent", we mean that the branch is comparatively longer among neighboring

branches (Fig. 1), and therefore is more likely to define a better supported and more reliable clade in the

tree. Also, longer branches (branch length is proportional to the number of accepted mutations along the

branch) are expected to contain more phenotypic mutations and correlate with larger phenotypic

differences, which indeed may be the case due to generally good agreement between current mostly

phenotypic classification and our tree-based definition of genera. We note that these prominent branches

frequently "line up" under each other forming a level of classification (Fig. 1 green highlight, Figs. 13—

18) and naturally define genus level clades. This level dates to about 15-20 Mya (Chazot et al. 2019).

We note that, similar to species, genera can vary in genetic diversification. E.g., Genus 1| (Fig. 1

magenta) is genetically compact, while Genus 2 (Fig. | cyan) is genetically diverse. But both are

supported by prominent branches. However, Genus 3 and Genus 4 (Fig. 1 blue and red) are both compact

and not separated from each other by prominent branches. From genetic perspective, their definition

appears arbitrary and inconsistent with how Genus 2 is defined: they do not form the same level in the

classification. The level that defines Genus 3 and 4 is closer to the leaves, and therefore should not have

the same rank and the level that defines Genus | and 2. Thus, to achieve better consistency of the

classification, we propose to combine Genus 3 and 4 into one, emphasizing evolutionary relationships and

not subjective preferences of the authors who defined Genus 3 and 4. The next prominent level is defined

as subgenus. For species, we largely rely on Fst (relative genetic diversification) and Gmin (measure of

gene exchange) as detailed in Cong et al. (2019a). As a rough guide, but not a decisive criterion (Trujano-

Ortega et al. 2020), we frequently provide percent difference in COI barcodes: 2% is usual for different

species (Hebert et al. 2003), 10% for different genera, and about 7-8% for different congeneric subgenera.

The taxonomic rearrangements presented below follow the standardized format. Taxonomic act is

the title of each section. For cited genera and subgenera, type species are given, and if the type species are

synonyms, valid names are provided. When the species are listed with their originally proposed genus

name, author names are given without parenthesis. For each species and subspecies with changed rank,

type locality is specified. Most sections are illustrated by a segment of a nuclear genomic tree (or Z

chromosome tree when specified) with species minimally necessary to support the conclusion. Presently

employed genus-species combinations (Callaghan and Lamas 2004; Lamas 2004; Mielke 2005; Pelham

2008; Hall 2018) are used in the figures, including recently proposed changes (Pelham 2019; Zhang et al.

2019c; Pelham 2020; Zhang et al. 2020). New combinations and taxonomic changes are given in the text.

Colors highlight phylogenetic groups and inconsistencies within the present classification that require

Genus 2 Genus 2

Genus 3

Genus 4 |

attention and changes proposed here. The section ends with a conclusion and, if necessary, with a list of

Species with revised genus-species names combinations. The sections are ordered by family and generally

in their taxonomic order deduced from genome-scale phylogeny complemented by phenotypic

considerations. Whole genome shotgun datasets we obtained and used in this work are available from the

NCBI database <https://www.ncbi.nlm.nih.gov/> as BioProject PRJNA731937, and BioSample entries of

the project contain the locality and collection data of the sequenced specimens shown in the trees. Exon

sequences with diagnostic characters highlighted are also available from <https://osf.10/kj4es/>.

Family Papilionidae Latreille, [1802]

Boreographium Grishin, new subgenus

http://zoobank.org/32D82A5D-2DEB-4685-8179-D5FF9ESF2225

1 if] 1 Re Eurytides phaon phaon|19012407|Mexico:Chiapas

Type species. Papilio marcellus Cramer, 1777. eurytides thymbraeus] 1901 ay (ylexice Oaxaca|1988

Eurytides ilus|190 cee ea ‘Chiapas|1988

Eurytides ariarathes|1911 Nil|venezuelal 1984

Definition. As_ revealed by genome-scale me, Eurytides marcelus|34gelUSAITX,Tyler Co,|2015

Eurytides marcellus

phylogenetic trees (Zhang et al. 2019d; Zhang _ |_| ma t<——" ies ALN cere aL son

. . urytides zonafa OmMInitan REePUBIIC

et al. 2019c), the type species of this new See eee ee aaetenene ee

subgenus is in the same clade with the subgenus euryedes dolicaon|17112c04)Peru| 1986

‘ 4 : te) Eurytides leéucaspis|17112C12|Peru:Junin|2003

Mimoides K. Brown, 1991 (type species Papilio Fig. 2. Genus Eurytides with subgenera Boreographium

£ ry & srap

ariarathes Esper, 1788) (Fig. 2). However, its subgen. n. (red), Mimoides (blue), Neographium (cyan),

wing patterns and shapes are more similar to ROPES GUSH DONDIS PAGE WYO RES DIAC):

subgenus Neographium Mohn, 2002 (type species Papilio philolaus Boisduval, 1836). This phenotypic

distinction and early divergence from the common ancestor with Mimoides suggest that the marcellus

clade should be defined as a subgenus of its own. This new subgenus is diagnosed by male genitalia: a

unique broadly rounded ear-shaped harpe with a dorsal narrow keel projecting ventrad as a tooth for half

of its length, and with two prominent narrow teeth: dorsal tooth directed anteriad, ventral tooth directed

caudad, as illustrated on plate 66 in Tyler et al (1994). In other subgenera, the harpe is either narrower, or

the keel is broader or not projecting beyond harpe, or the teeth are smaller or directed differently.

Etymology. The name is a masculine noun in the nominative singular, formed from Boreo|tis| (Latin for

northern) + Graphium in reference to the northernmost representatives of the "Graphium" complex of taxa

in America distributed up to northern Minnesota (Lotts and Naberhaus 2021).

Species included. Only the type species.

Parent taxon. Genus Eurytides Hiibner, [1821].

Esperourus Grishin, new subgenus

http://zoobank.org/DCO85A9A-4391-49F3-B3 AE-F2F57B0BB76C

Type species. Papilio esperanza Beutelspacher, 1975.

Definition. Confidently placed by Zakharov et pterourus ee Oaxacal1968

al. (2004) as sister to the nominal subgenus of Prague planedes 166 S3/USA:TX.A Hardin co [2016

oF pterourus mullcaudata pao IBS| Lea UT Gomeld co Co,|2016

Pterourus Scopoli, 1777 (type species Papilio , Pterourus rutulus|6374/USA:CO,Grand Co./2016

. . . Pterourus eae cul 6389|USA: co, ale ne 2016

troilus Linnaeus, 1758) this new subgenus i Pterourus glaucus| 1670|USA:Tx,Denton Co.|20

A i ¥ eT. anni Pterourus alexiares garcia/5522|US5 ATX, Bre Aes Co, Bao

shows prominent genetic differentiation from Btareunie Vicornus|t74geF1ieso:san tue Bo rene

F F ° Pterourus eeseBaccu ad Pi 1cO aq Hexic eerne a tee

the three Species of the nominal subgenus (Fig. Pterourus scamander|14081C05|Brazil:SC,Joinville| 1986

3 d h di d fi h 1 h aS Eee Tee | ae ee ae entinal 1267 ‘Tingo Maria

) an as iIverge rom t em prior to t c 7 Pterourus elwesi|16106402|China|1987

diversification of all other species of the genus Fee oe ee ae

Pp g Fig. 3. Genus Prerourus with subgenera Esperourus

Pterourus. Due to its ancient (comparatively to subgen. n. (red), Pterourus (blue), Jasoniades (cyan

g yan),

other Pterourus species) origin, the clade with Pyrrhosticta (purple), and Agehana (green).

Pterourus esperanza represents a valid subgenus. This new subgenus is diagnosed by a unique serrated

lobe near the distal end of harpe in male genitalia, as illustrated on plate 100 in Tyler et al. (1994); and

may be distinguished from other Pterourus species by its ventral forewing discal cell with 4 yellow

longitudinal rays in the basal half, and from the three species in the subgenus Prerourus by its basal edge

of the postdiscal band on the ventral hindwing that are closer to the outer margin than to the discal cell.

Etymology. The name is a masculine noun in the nominative singular, formed as a fusion of the type

Species name with its genus name: Esper|anza] + [Pter]ourus.

Species included. Only the type species.

Parent taxon. Genus Pterourus Scopoli, 1777.

Ayppasonia Grishin, new subgenus

http://zoobank.org/B37D0541-CD40-4824-AE2 1-BD6E37D37CB7

Type species. Papilio hyppason Cramer, 1775. Seon pea vee 1908

. - Braciiges pelaus amalca

Definition. Previously, we noted this unnamed “Heracles eee

subgenus in the COI barcode dendrogram | 3 « ca Harselides andFogeus|1<08i¢0, |Nemce Siialoa2003

: . . oye . +_ Heraclides pallas|17116F08|Us ‘TX, Hidalgo Gic(oe 75

(Shiraiwa et al. 2014). Here, its definition is for- | 4 Heradlides aristodem rus] 2882 [Dominican Republic|2 8 BOF acieeniaet

e 6 o

malized. While the COI barcodes demonstrate arya ies Heracles hy {apatcok [Pens tinge plata 1985

3 di . . f h b f elds pbc taedae Fg ey satel Dallas Co./201

its istinction trom other subgenera oO Hee ees eee eo 7 Hart AS aria|1996

Heraclides andraemon 10301|Jamaic a|[20

Heraclides Hiibner, [1819] (type species Papilio 7 Heradides machaonides|14084¢03 Dominican Republic| 1982

thoas Linnaeus, 1771), its unexpected sister Fig. 4. Genus Heraclides with subgenera Hyppasonia subgen. n.

i ; ; ; d), Heraclides (blue), Calaid d Priamid le).

relationship with the nominal subgenus Co ERGO S AINE AGS Cyan) aes eitanntae SKOUrR Ie)

Heraclides—instead of with Priamides Hubner, [1819] (type species Priamides hipponous Hibner,

[1819], which is a junior objective synonym of Heraclides anchisiades (Esper, 1788)) as hinted by

similarities in wing shape and patterns—was discovered by Lewis et. al. (2015). Our genomic tree

confirms this placement, and at the same time reveals prominent genetic differentiation from the nominal

subgenus (Fig. 4). Curiously, the long branch in the tree that defines this subgenus suggests accelerated

evolution that may explain its phenotypic differences from the nominal subgenus and likely mimetic wing

patterns. This new subgenus is distinguished from others by a bilobed, crab-claw harpe with strongly

unequal lobes: one lobe broad, rounded and serrated distad, and the other lobe shorter, narrow, tooth-like

(for illustration see plate 83 in Tyler et al. (1994)). In other species with a bilobed harpe, the two lobes are

nearly equal or at least the larger lobe is narrower and prominently constricted before the serrated end.

Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.

Species included. Only the type species.

Parent taxon. Genus Heraclides Hiibner, [1819].

Family Pieridae Swainson, 1820

Prestonia Schaus, 1920 belongs to Euremini Grote, 1898

The monotypic genus Prestonia Schaus, 1920 (type and the only species Prestonia clarki Schaus, 1920)

has been tentatively placed near Phoebis Hiibner, [1819] (type species Phoebis cypris Hiibner, [1819], a

junior subjective synonym of Papilio argante Fabricius, 1775) due to phenotypic similarity and no DNA

sequences available for it. The genomic tree reveals that subfamily Coliadinae Swainson, 1821 splits into

two clades that we treat as tribes: Coliadini Swainson, 1821 (includes Phoebis) and Euremini Grote, 1898

(Fig. 5). Genomic-scale phylogeny confidently places Prestonia as sister to Kricogonia Reakirt, 1863

(type species Colias lyside Godart, 1819) and therefore Prestonia belongs to Euremini Grote, 1898 (the

clade that does not include Phoebis) and not to Coliadini.

Tertocolias deva (E. Doubleday, 1847) and Teriocolias reticulata (A. Butler, 1871)

new combinations

Currently placed in Eurema Hibner, [1819] (type species Papilio delia Cramer, 1780, a junior homonym:

valid name for this species is Pieris i Eurema hecabe|20039401|Thailand,Chai Prakan District

Ve laine a nea Ts , | [Eurema hapale|20078Guz|Etngpia Oromia 1957.

aira odart, ), WO Species . ee hiale|19078FO2|Brazil:Federal ae evi

; a urema daira daira onroe Co

Terias deva Doubleday, 1847 and = Pyrisitia lisa|3545|USA:TX,Hidalgo Co.|2015

. . P ith te 5167 |USA:TX,St Co.]2015

Terias reticulata Butler, 1871 are not abaeis albulal itoeeoiMewicn: Taraulpastio7

. : : Abae 4057 (USA;TX,D Co,{2015

monophyletic with E. daira, and my Peucidia brephos|190¢acOol French Ftianal1999

r F , Leucidia elvina] L9067HO5|Brazil:Rio de Janeiro|1996

instead are in the same clade with Teriocolias zelia|19067HO08|/Argentina:Tucuman|2013_

a ate” 1 Eurema deval|19078E08/Brazil:Minas Gerais|1994

Teriocolias zelia (Lucas, 1852), | Pum Purema (oN athals iole|3572/USAs1X Cameron Co.|2015

’ > £ Nathalis iole ameron Co,

; : : . NEC K lyside|3423|USA:TX,C Co.|2015

which is a valid name of Terias |-2 a8 SE eee SSAA REL Canoe A ae Taeo

arnae Hewitson 1874 the t e 4 Gonepteryx dala rae ge re ea Al ‘

0. De = =[|19067F01|M ‘J 1991

h f Teri : li Rob no aa eaniacs harnal e704 Myanmar Rakhine|2003

— | I Ay tat tat 16107012 |US5A4:TX Nueces Co./2013

species of Teriocolias Rober, cota — febarata TAT ies Gantana oe)

1 1 Catopsilia pyranthe|19067E09|Sri Lankal1976

(Fig. 5). Therefore, these two species — Anteos maerulal 10934|F|USATX.Camaran Co. Harlingen|2017

: aie Zerene cesonia allas Co

do not belong to Eurema and instead i Colas currtheme |OS30|USA:CO Bark Co.|3016

can be placed in_ Teriocolias Fig. 5. Coliadinae and its two tribes: Coliadini and Euremini.

implying Teriocolias deva_ (E. Eurema (blue), Terias (red), and Teriocolias (green and magenta).

Doubleday, 1847) comb. n. and Teriocolias reticulata (A. Butler, 1871) comb. n.

Terias Swainson, 1821 is a valid genus

Our genomic tree reveals that Ewrema Hiibner, [1819] (type species Papilio delia Cramer, 1780, a junior

homonym: valid name is Pieris daira Godart, 1819) is paraphyletic with respect to Pyrisitia Butler, 1870

(type species Papilio proterpia Fabricius, 1775) (Fig. 5). To restore monophyly, we choose to keep

Pyrisitia as a genus and therefore treat the Old World clade currently placed in Eurema as a distinct valid

genus. Terias Swainson, 1821 (type species Papilio hecabe Linnaeus, 1758) is its oldest available name.

Pyrisitia westwoodii (Boisduval, 1836) is a species

distinct from Pyrisitia dina (Poey, 1832)

Currently considered a subspecies of Pyrisitia dina (Poey, 1832) (type locality Cuba), Terias westwoodii

Boisduval, 1836 (type locality Mexico) IS prominently Pyrisitia dina westwoodii|17116H10|Mexico;QRoo|1979

: : ° Pyrisitia dina Bae Let ae neat

separated genetically from the insular taxa (Fig. 6). Pyrisitia dina helios|5351/USA:FL,Miami-Dade

oe : : *Bynstia gina helios)sis7 (USA iiam-Dade Co ors.

Compared to Pyrisitia dina helios (M. Bates, 1934) Brrsts poem 358 | jamateaS817 e Co,|

(type locality Bahamas) the Fst/Gmin statistics are Bytaiia parvumbra|.oeeallamaies|soi7

0.46/0.03 and their COI barcodes are 2.7% (18 bp) Pvrisitte leucsl oO 7004 erssil ondamil1996

different. Therefore, we propose that Pyrisitia Fig. 6. Pyrisitia westwoodii (red), and dina (blue)

westwoodii (Boisduval, 1836), reinstated status is a jae eal oa ee

species-level taxon and employ Pyrisitia westwoodii gabriela Le Crom & Llorente, 2004 comb. nov. We

also confirm Pyrisitia parvumbra (Kaye, 1925) as a species (Fig. 6; 2.6%, 17 bp barcode difference).

Zegris Boisduval, 1836 is a subgenus

of Anthocharis Boisduval, Rambur, [Duménil] & Graslin, [1833]

Zegris Boisduval, 1836 (type species Papilio eupheme Esper, [1804]) originates within Anthocharis

Boisduval, Rambur, [Duménil] & Graslin, [1833] (type species Papilio cardamines Linnaeus, 1758),

rendering it paraphyletic, and is confidently placed as a sister to subgenus Paramidea Kuznetsov, 1929

(type species: Anthocharis scolymus Butler, 1866) (Fig. 7). Subgenera of Anthocharis are genetically

close to each other and even to the sister genus Euchloe (type species Euchloe ausonia var. esperi W. F.

Kirby, 1871, which is Euchloe crameri Butler

Anthocharis scolymus|19067G09|South Korea,Naksansal19388

1 Fegris eupheme|19068B02|Azerbaijan,vic. Agstafa|1972

(1869)) (Fig. 7). To restore the monophyly, ed die AAR Se cease eli : |

1 1 ; Anthocharis cardamines|17115H10|Spain:Cataloniai|/2005

instead of elevating subgenera of Anthocharis to Anthoacharis midea annickae|6134|US4:Wiv Pendleton Co,|2016

genus status we propose to treat Zegris Anthocharis cethura cethural6667|(US4:C4,San Diego Co,/2004

4 Euchloe charlonia]19067612|Tunisia]1981

Boisduval, 1836 as a subgenus of Anthocharis ca Euchloe cramer] 19068401|Portugal:Algarve|1978

B : d l R b D i 7 & G lj Euchloe tagis|19068402|M|Portugallold

oisduval, Rambur, [Dumenil] rash, | Rig. 7. Anthocharis (blue and red) and Euchloe (magenta).

[1833], along with its other two subgenera

Tetracharis Grote, 1898 (type species Anthocharis cethura C. & R. Felder, 1865) and Paramidea.

Cunizza Grote, 1900 is a junior subjective synonym of Hesperocharis C. Felder, 1862

Despite differing wing patterns, a monotypic Hesperocharis erota|19068408|Brazil:Rio de Janeiro|1995

; an Hesperocharis nereina (=chloris}|18097H12|LT|Bolivialold

genus Cunizza Grote, 1900 (type and the only = Cunizza hirlanda|19068411|Venezuela|1985

1 fy ; Mathania leucotheal15117G01|[Chile: Coquimbo|1991

i tars Papilio hirlanda Stoll, 1790) ap Hesperocharis emeris (=catasticta)|18097H10|LT|Brazil:SPlold

originates within Hesperocharis Felder 1862 Hesperocharis crocea|19068A406|Venezuelal1964

; z Hesperocharis nera aida (=boliviana)}|18097HO9|LT|Bolivial|old

(type species Pieris erota Lucas 1852) — Hesperocharis paranensis|10216/Argentina: Buenos Aires|2006

. : . : i Hesperocharis infrasignata|10218|4rgentina:Cordoba|2008

rendering It paraphyletic (Fig. 8). To restore _Mathania aureomaculata esther|19068A09|Peru:Huanuco|1973

. * Mathania aureomaculata esther|19068410|Peru:Hduanuco|1974

monophyly, we propose that Cunizza Grote, Eroessa chiliensis|19067G10|Chile:Palena|1987

1900 is a junior subjective synonym of Fig. 8. espero ie red and magenta)

Hesperocharis C. Felder, 1862. and Mathania (green).

Hesperocharis leucothea (Molina, 1782) new combination

Papilio leucothea Molina, 1782 (type locality Chile), currently placed in the genus Mathania Oberthiir,

1890, originates within Hesperocharis Felder, 1862 (type species Pieris erota Lucas, 1852) according to

the genome-scale tree (Fig. 8), which implies Hesperocharis leucothea (Molina, 1782) comb. n.

Reliquia Ackery, 1975 is a junior subjective synonym of Pontia [Fabricius], 1807

A monotypic genus Reliquia Ackery, 1975 (type and the only species Reliquia santamarta Ackery, 1975)

originates within Pontia [Fabricius], saw Pieris brassicae|15112G01|Germany,Egestorf| 1986

. oe — Pieris rapae|3537|(USA:TX,Dallas Co,Dallas|2015

1807 (type species Papilio daplidice 7 o.gpleris oleracea oleracea|16105B04|USA4:MI,St. Joseph Co,|1984

: : ; ; _ Pieris marginalis guppyi|16107B09|Canada:Yukon Territory|2016

Linnaeus, 1758), rendering Pontia Pieris napi napi|16105D07|Norway|1987

h 1 ‘i (Fi 9) Th t h Sisymbria z Bene Best ty LNasaltah wi Geeeee Bite

r 1 19, : r ontia sisymbrii elivata|8922|USA anta Fe Ca,|21

pe : 7 ‘ 8 : fi SSS 0.654 onbewchlora—, Pontia beckerti|15112F11|USA:NV Douglas Co,|1993

that Reliquia is a close sister to the ie Pontia chloridice|7722|Kazakhstan|1966

, ; Baltia shawii karakuli[15117G03|Tajikistan|2004

clade of three closely related species: | Balig Baltia rut lata soncomarcalis078t — evade de Santa al

. oer = aliquia santamarta|19078C06 mb a:Sierra Nevada de Santa Marta|1977

Pontia callidice Hiibner, [1800] (the [Rel quia santamarta|19078C07|Colombia:Sierra Nevada de Santa Martal19

, in Pontia occidentalis |9338|USA: WY, Park Co, Yellowstone NPI2017

type species of Synchloe Hibner, Bh se Pontia protodice|8433|USA:TX,Wise Ca.,LB] National Grassland|2017

: 7 i 7 Pontia callidice callidice|15117G05|Spain: Gironal1988

1818, currently a junior subjective Pontia daplidice|15117F12|Spain:Gironal 1989

. : Lex Phulia nymphula|19077411|Peru:duamangal2003

synonym of Pontia), Pontia callidice +, (Infraphulia ilyodes|15117G12|Chile,Tarapaca|1991

i D. ‘Infraphulia illirmani]19069603|Bolivia:La Paz|1984

(Boisduval & Le Conte, [1830]) and , Pierphulia nysias|19078B06|Bolivialold

: 1 : 4 i * Pierphulia rosea/19078B09|Chile:Antofagasta|1961

Pontia occidentalis (Reakirt, 1866). af ,Plercolias forsteri|19078B02|F|Bolivia,lllimani|1950

’ Piercolias huanaco|19078B03|Bolivialold

E.g%, COI barcodes of R. sSantamarta Theochila maenacte|/19078D03|Argentina: Buenos Aires|2008

ae 3 jHypsochila penai]/19069F12|Chile:Antofagasta|1961

and P. callidice differ by 4.7% (31 ss Hypsochila wagenknechti|19069G602|Chile:Coquimbo|1966

: ; : : : Tatochila autodice|19078C11|Bolivia: Cochabamba,Cereado| 1976

bp). Therefore, Reliquia Sa junior Ascia monuste monuste|4148|USA:Tx Jefferson Co,|2015

7 , , Sanyra josephina josepha|17116612|Mexico:Tamaulipas| 1974

subjective synonym of Synchloe. (L* Ganyra howarthi|15117H10|Mexico:Sonora|2003

0.98 Ganyra phaloe|17065401|Bolivia:Beni,Trinidad|1989

Because we do hot have sufficient Fig. 9. Genera Phulia (green) and Pontia, colored non-green, with

evidence to raise Synchloe from | subgenera Pontia (blue), Baltia (red), Pontieuchloia (cyan), and Sisymbria

synonymy with Pontia, we propose subgen. n. (magenta). Former genus Reliquia is shown in pink.

that Reliquia Ackery, 1975 is also a junior subjective synonym of Pontia [Fabricius], 1807.

Baltia Moore, 1878 and Pontieuchloia Verity, 1929

are subgenera of Pontia [Fabricius], 1807

A small genus Baltia Moore, 1878 (type species Mesapia shawii Bates, 1873) originates within Pontia

[Fabricius], 1807 (type species Papilio daplidice Linnaeus, 1758), rendering Pontia paraphyletic (Fig. 9).

Baltia is a confident but comparatively distant sister to the clade of two closely related species Pontia

chloridice Hiibner, [1813] (the type species of Pontieuchloia Verity, 1929, currently a junior subjective

synonym of Pontia) and Pontia beckerii (W. H. Edwards, 1871). E.g., COI barcodes of B. shawii and P.

chloridice differ by 8.5% (51 bp). To restore monophyly of Pontia and, at the same time, keep the relative

distinction of Baltia, we propose that Baltia Moore, 1878 is a subgenus of Pontia [Fabricius], 1807. If

Baltia is a subgenus and not a synonym, clades comparable to it in prominence should be defined as

subgenera. In addition to the nominal subgenus (Fig. 9 blue and pink), of which Synchloe Hiibner, 1818

(type species Papilio callidice Hiibner, [1800]) is kept as a junior subjective synonym, we propose that

Pontieuchloia Verity, 1929 is a valid subgenus (Fig. 9 cyan). For these subgenera to be monophyletic, a

new subgenus is proposed next for the clade with Pontia sisymbrii (Boisduval, 1852) (Fig. 9 magenta).

Sisymbria Grishin, new subgenus

http://zoobank.org/28C486B5-3F65-4CDD-AC44-3FE386B58D0B

Type species. Pieris sisymbrii Boisduval, 1852.

Definition. This new subgenus differs from other subgenera of Pontia by the following combination of

characters: forewing vein R3 longer than in other subgenera, about half of vein R4a+s length; androconia

present in the forewing discal cell spot in males, this spot is narrower than in other subgenera, with a

notch on the outer edge (smoothly curved of straight in Baltia) and without a line of white scales along

the discal cross-vein that is curved less strongly than in most other subgenera towards the wing base;

dorsal hindwing without prominent bar at the end of discal cell; aedeagus shorter and relatively broader

than in other subgenera, prominently curved at phallobase; hindwing below with gray or brown (not green

or yellow) scaling along yellowish veins; full-grown caterpillar with orange-yellow framed with black

rings on grayish segments; univoltine in spring. See Chang (1963) for elaboration on and illustrations of

some of these characters as they are given for P. sisymbrii.

Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.

Species included. Only the type species.

Parent taxon. Genus Pontia [Fabricius], 1807.

Comments. The genomic tree reveals markedly uneven rates of evolution within Pontia (Fig. 9): the

nominotypical subgenus evolves about 2 times faster than other subgenera. This observation combined

with rather substantial genetic differentiation among Pontia, including the COI barcode, which in P.

(Sisymbria) sisymbrii and P. (Pontia) daplidice differs by 8.7% (57 bp), and close similarity in

phenotypes of Pontia species creates a unique situation. On the one hand, Pontia (including Baltia) is a

morphologically compact genus. On the other hand, strong genetic diversification behind this apparent

phenotypic similarity may suggest elevating subgenera of Pontia to genera (which will return Baltia to the

genus status), a step that we refrain from.

Tatochila A. Butler, 1870, Piercolias Staudinger, 1894, Hypsochila Ureta, 1955,

Theochila W. D. Field, 1958, Pierphulia W. D. Field, 1958, and Infraphulia W. D.

Field, 1958 are junior subjective synonyms of Phulia Herrich-Schaffer, 1867

Phulia Herrich-Schaffer, 1867 (type species Pieris nvmphula Blanchard, 1852), Tatochila A. Butler, 1870

(type species Synchloe autodice Hiibner, [1818]), Piercolias Staudinger, 1894 (type species Trifurcula

huanaco Staudinger, 1894), Hypsochila Ureta, 1955 (type species Tatochila microdice f. wagenknechti

Ureta, 1938), Theochila W. D. Field, 1958 (type species Pieris maenacte Boisduval, 1836), Pierphulia W.

D. Field, 1958 (type species Phulia nysias Weymer, 1890), and /nfraphulia W. D. Field, 1958 (type

species Phulia nymphula var. illimani Weymer, 1890) cluster closely in the genomic tree without obvious

separation into groups (Fig. 9 green). E.g., COI barcodes of P. nymphula and T. autodice differ by 4.0%

(26 bp). However, being combined into one, all these genera together represent a prominently distinct

genetic group that is sister to Ascia Scopoli, 1777 (type species Papilio monuste Linnaeus, 1764) and

more distantly related to Ganyra Billberg, 1820 (type species Papilio amaryllis Fabricius, 1793, a junior

homonym: valid name for this species is Pieris josephina Godart, 1819). With genetic similarity between

these taxa being at the level of a species group, it may not be meaningful to consider these names as

denoting valid subgenera, and therefore we propose that Tatochila A. Butler, 1870, Piercolias Staudinger,

1894, Hypsochila Ureta, 1955, Theochila W. D. Field, 1958, Pierphulia W. D. Field, 1958, and

Infraphulia W. D. Field, 1958 are junior subjective synonyms of Phulia Herrich-Schaffer, 1867.

Calopierini Grishin, new tribe

http://zoobank.org/B7717ECE-C015-48AA-A57D-B7A6A3CCE4F6

Type genus. Calopieris Aurivillius, 1898.

Definition. In the genomic tree, this taxon is confidently placed as sister to the tribe Leptosiaini Braby,

2014 (Fig. 10, a monotypic tribe consisting of Leptosia Hiibner, 1818) and shares veins Mi and M2 being

connate with the hindwing discal cell, but genetically and phenotypically distant from it otherwise. Most

notably, the wings are not rounded as in Leptosia and venation differs: forewing veins R3 and Ra+s stalked

for less than half of their lengths and M: stalked with their stalk, resembling Coliadinae Swainson, 1821

(e.g., Phoebis Htibner, [1819]). In general appearance reminds more of Colotis, where it was formerly

included as a subgenus (Klots 1933), but is distinguished from it by very short and slender palpi that do

not protrude beyond the front of the head and are not visible from above. Antennae are short (about half

of forewing discal cell length) with large and flattened clubs. The combination of the abovementioned

characters of palpi, antennae and wing venation uniquely defines this new tribe.

Genera included. Only the type genus.

Parent Taxon. Subfamily Pierinae Swainson, 1820.

Comments. Although grammatically correct formation of this tribe name calls for insertion of "id" before

ini", these letters have been elided from the stem to agree with Pierini, under Art. 29.3.1.1 UCZN 1999).

Phrissura Butler, 1870 is a subgenus of Appias Hiibner, [1819]

Frequently treated as a separate and monotypic genus, Phrissura Butler, 1870 (type pia Pieris illana

C. & R. Felder, 1862, currently a subspecies of “=e Coleus tausta

Pieris aegis C. & R. Felder, 1861), is rather a2 ee ise au 2003370 ni

closely allied to Appias Hitibner, [1819] (type

species Papilio zelmira Stoll, 1780, currently a

subspecies of Papilio libythea Fabricius, 1775) ° rete ity 7ave|binparnes batsan(ald

(Fig. 10 magenta and orange). E.g., COI bar- aoa affnis|120¢8D07|Indonesia Sulawesilold

codes of P. illana and Appias olferna Swinhoe, a A ! Elk 8312 fae Guinea a

1890 differ by 9.9% (65 bp). To emphasize the i Abbie pandione|19066¢09| Myanmar § Shan State|2003

relationship between Appias and monotypic Fig. 10. Ca/opierini trib. n. (green), Appias (magenta and orange)

Phrissura, we propose treating the latter as a with its subgenus Phrissura (orange) and Catophaga (blue and

a i

subgenus of the former, as already adopted in red) with its subgenus Safetara (red)

some publications (Wahlberg et al. 2014).

Saletara Distant, 1885 is a subgenus of Catophaga Hiibner, 1819

Frequently treated as a subgenus of Appias Hiibner, [1819] (type species Papilio zelmira Stoll, 1780,

currently a subspecies of Papilio libythea Fabricius, 1775), Catophaga Hiibner, [1819] (type species

Papilio paulina Cramer, [1777]) is not monophyletic with it and instead, together with Sa/etara Distant,

1885 (type species Pieris nathalia C. & R. Felder, 1862, currently a subspecies of Papilio liberia Cramer,

[1779]), is sister to doa de Nicéville, 1898 (type and the only species Pieris affinis Vollenhoven, 1865)

(Fig. 10). Therefore, in agreement with Wahlberg et al. (2014), we treat Catophaga as a valid genus distinct

from Appias. Our genomic tree reveals close clustering of Saletara (Fig. 10 red) with Catophaga (Fig. 10

blue) at the level consistent with congeners, hence we propose that Sa/etara is a subgenus of Catophaga.

Mesapia Gray, 1856 is a junior subjective synonym of Aporia Hiibner, [1819]

A monotypic genus Mesapia Gray, 1856 (type and the only species Pieris peloria Hewitson, 1853)

originates within Aporia Hitibner, [1819] (type species oo agathon|19068D09|India:Uttar Pradesh|1969

Aporia agathon maltrechti|19068D 10|Taiwan:Nantou|1980

Aporia leucodice|19068E10|Tajikistan|1978

Aporia crataegi|19068E09|Albania:4pollonia/1991

Mesapia peloria|19069H10|India:Tamil Nadul|old

Papilio crataegi Linnaeus, 1758) rendering it paraphyletic

(Fig. 11). Due to the close relationship of these taxa, it is

not desirable to restore monophyly by breaking Aporia into Fig. 11. Aporia (blue and red).

at least 3 genera. Therefore, agreeing with previous studies (Ding and Zhang 2016; Todisco et al. 2020),

genomic-scale phylogeny provides the ultimate evidence that Mesapia Gray, 1856 is a junior subjective

synonym of Aporia Hiibner, [1819].

Leodonta Butler, 1870 is a subgenus of Pereute Herrich-Schaffer, 1867

Despite its nearly extreme difference in appearance: Leodonta Butler, 1870 (type species Euterpe dysoni

Doubleday, 1847) and Pereute Herrich- ~ LiMylothris sagala dentatus|20034E03|Malawi,Misuku Hills|1974

Mylothris narcissus mahale|20034E02|Tanzania:Mpanda|1974

Schaffer, 1867 (type Species Euterpe Mylothris poppea|19069HO7|Cameroon:Mount Kala|1987

ie n Leodonta dysoni|19077403|Peru:Cuzco,Cosnipata Valley|2015

callinice C. & R. Felder, 1861) are 4 Pereute callinice|19077406|Columbia:Cundinamarcal 1984

3 3 . Melete lycimnia lycimnia] 19069HO3|Guyanal1999

genetically close sisters (Fig. 12). * Melete lycimnia flippantha|19069H04|Brazil:Rio de Janeiro|1964

J A cl i i Archonias brassolis tereas|19068G02|Brazil:Rio de Janeiro|1994

Genetic diversification between them 1S 0. “archonias brassolis brassolis]19068G601|Guyana|2000

I 1, i : | Catasticta nimbice nimbice|17116G01|Mexico:Nueva Lean|1979

or Charonias eurytele|19068HO7 |Ecuador:Napo|1989

SMC E t is t at MAS ated genus Neophasia menapia magnamenapia|P40257|U54:C0,Larimer Co,.|2016

Mylothris Hiibner [1819] (Fig 12) Eucheira socialis|19069F10|Mexico:Michoacan,Uruapan|1980

A < .

Fig. 12. Pereute (purple and green) with its subgenus Leodonta (green)

and Archonias (blue and red) with its synonym Catasticta (red).

Disparate phenotypes are frequently

placed in the same genus, e.g.

Pterourus euterpinus (Salvin & Godman, 1868) and Pterourus eurymedon (Lucas, 1852) to some extent

resemble Pereute and Leodonta, respectively. Therefore, we propose adopting a more internally

consistent classification when groups of similar divergence are assigned similar taxonomic rank, and treat

Leodonta Butler, 1870 as a subgenus of Pereute Herrich-Schaffer, 1867 despite their wing pattern

differences.

Catasticta Butler, 1870 is a Junior subjective synonym of Archonias Hiibner, 1827

Catasticta Butler, 1870 (type species Euterpe nimbice Boisduval, 1836) clusters closely with Archonias

Hiibner, [1829] (type species Archonias marcias Hubner, 1825 which is currently treated as a subspecies

of Papilio brassolis Fabricius, 1777 from Brazil: Bahia) in the genomic tree (Fig. 12). Genetic

diversification between these genera is within the norm for most Pieridae genera, e.g., Mylothris shown in

the same tree. Moreover, previously published phylogenetic trees reveal that Catasticta is not

monophyletic (Padron 2014; Wahlberg et al. 2014), although sometimes with low support. Furthermore,

Padron (2014) discussed this issue at length and suggested synonymy between Catasticta and Archonias.

We agree with these conclusions, also supported by our genome-scale results. Although further studies are

required to develop a meaningful classification within this very large genus, three independent studies

provide phylogenetic data suggesting that Catasticta Butler, 1870 is a junior subjective synonym of

Archonias Hiibner, 1827. This name change may not be welcomed by those used to the name Catasticta,

but it will be necessary to accept at one point. While some may consider breaking this large genus into

several genera, the genomic tree indicates that other relatives of Catasticta, such as Charonias Rober,

1908 (type species Euterpe eurytele Hewitson, 1853), Neophasia Behr, 1869 (type species Pieris menapia

C. & R. Felder, 1859), and Eucheira Westwood, 1834 (type and the only species Eucheira socialis

Westwood, 1834) are all closely allied to each other and may be treated as subgenera of Archonias, along

with some others currently placed in Catasticta. However, details of such classification will be revealed

after genomic-scale sequencing of all major species groups in this complex is completed.

Family Lycaenidae [Leach], [1815]

Sandia Clench & P. Ehrlich, 1960 and Xamia Clench, 1961 are

junior subjective synonyms of /ncisalia Scudder, 1872

Appearance could be misleading, especially if unusual. Despite unique wing patterns, we find from the

genomic analysis that Sandia Clench & P. Ehrlich, 1960 (type and the only species Callophrys (Sandia)

mcfarlandi P. Ehrlich & Clench, 1960) originates within Deciduphagus K. Johnson, 1992 (type species

Thecla augustinus Westwood, 1852), and is sister to Callophrys fotis (Strecker, [1878]) (Fig. 13). Because

Deciduphagus is currently a junior subjective synonym of /ncisalia Scudder, 1872 (type species Lycus

niphon Hubner, [1819]) (Pelham 2008; Pelham 2020) due to evolutionary closeness, and because Sandia

renders both Deciduphagus and Incisalia paraphyletic, we propose that Sandia is a junior subjective

synonym of /ncisalia. Furthermore, uniquely patterned Xamia Clench, 1961 (type species Thecla xami

Reakirt, [1867]) also originates Subgeneta >>, Callophrys dumetorum|17107F08|USA:WA,Mason Co,|1989

. - . . ° -s- . ' ek Callopnrys viridis|965/US4:CA Monterey Co,j2012

within Deciduphagus and is sister Genie [eC Siaphrys sheridan PAOZ9SIUSA:CO. ee [2017

he Deciduph cae Paldahrys nishon|L6107A0s|0SA: oe sees Co.|2016

7 allophrys niphon ake Co,

to the €Cl up agus core group a ily ealep phrys lancraleensis| 17114E07 IME, saa pees

1 1 ss allophrys eryphon fusca a Fe Co,|2

that contains the type Species. 7 Callophrys henrici|[5807|USA:TX,Travis Co,|2016

. : : ,— Callophrys irus|S862|USA:TX,Freestone Co.|2016

Although Xamia is more distant Callophrys polios|17108A01/USA:WA,Kittitas Co. [1991

i Callophrys 55 Callophrys xami|[5428[USA4:TX, Cameron Co ), [2015

from other congeners than Sandia cE Callophrys fotis|17114E06|USA:NM,San Juan Co,|1980

at thi ; ca Gehrels easel isa ess cede oa

it still falls within the prominent Callophrys Seidman uae Eee Co.|2017

Incisalia clade. Thus leaving it as me

: 8 - a ab Callaphrys rubi]20037HO6|Spain| aoe

a valid subgenus would require a i= Callophrys avis|20037H11|Spain| 197

; llaphrys hassel i8GL4C1 1 USA: Nc, Gates Co,|1983

new name for at least Callophrys Callophrys gryneus|3521|USA:TX,San Jacinto Co, dare

: Cyanophrys Sig Celis eanaror col 1965

polios (Cook & F. Watson, 1907), : Cyanophrys herodotus|17117408|Mexico: SLP|197

which is. sister to the clade Fig. 13. Callophrys subgenera: Greenie subgen. n. (red), Cisincisalia (pink),

consisting of Xamia and the core Callophrys (blue), Mitoura (olive), and Incisalia (green) with its synonyms

Deciduphagus species, but is not Deciduphagus (cyan), Xamia (orange), and Sandia (magenta).

prominently distinct from its relatives. Therefore, we propose to treat Xamia as a junior subjective

synonym of /ncisalia. Finally, we note that Deciduphagus as originally defined (Fig. 13 cyan) (Johnson

1992) is also paraphyletic with respect to /ncisalia (Fig. 13 green), and Callophrys henrici (Grote &

Robinson, 1867) with Callophrys irus (Godart, [1824]) should be attributed to /ncisalia sensu stricto

instead of to Deciduphagus. As a result of this analysis (Fig. 13), we conclude that the genus Callophrys

diversified into 5 prominent clades that correspond to the level of subgenus: Callophrys, Incisalia,

Cisincisalia K. Johnson, 1992 (type species Cisincisalia moecki K. Johnson, 1992, a junior subjective

synonym of Callophrys guatemalena Clench, 1981), Mitoura Scudder, 1872 (type species Thecla smilacis

Boisduval & Le Conte, [1835]) and the fifth subgenus that includes Nearctic species superficially similar

to Palearctic Callophrys, but instead is a confidently supported sister to /ncisalia (Fig. 13 red) (ten Hagen

and Miller 2010). This subgenus does not have a name available for it and therefore is new.

Greenie Grishin, new subgenus

http://zoobank.org/73368 1 D0-34AF-4884-9396-BE415705E783

Type species. 7hecla sheridonii |sic] Edwards, 1877, presently Callophrys sheridanii (Edwards, 1877).

Definition. Previously placed in the subgenus Callophrys Billberg, 1820 (type species Papilio rubi

Linnaeus, 1758) but is not monophyletic with it, forming a prominent lineage of its own within the genus

Callophrys (Fig. 13). In appearance, remarkably similar to some species (including the type) from the

subgenus Callophrys in: wings rounded without tails, slightly lobed hindwings at anal angle, and green

color of ventral surface with postdiscal variously complete to absent row of white markings.

Distinguished from Callophrys by hindwing tornal area: less developed lobe and less crenulate outer

margin, and generally smaller and rounder forewing androconial patch. Diagnosed among the genus

Callophrys by a combination of the following characters in the nuclear genome: cce2400.8.3:T366C,

cce8426.17.3:C4290T, ccel10587.6.6:G181C, cce925.9.4:C351T, and cce2041.25.2:T1572A; and the COI

barcode: 479C (not T) and 610C (not T), differing from the subgenus Callophrys in 202T (not A), 512G

(not T), and 556T (not A). See <https://osf.10/kj4es/> for the sequences with these characters.

Etymology. The name is a feminine noun in the nominative singular, Latinized verbatim from the

affectionate English name of this butterfly group.

Species included. The type species, Thecla viridis W. H. Edwards, 1862, Thecla dumetorum Boisduval,

1852, and Thecla affinis W. H. Edwards, 1862.

Parent taxon. Genus Callophrys Billberg, 1820.

Family Riodinidae Grote, 1895

Takashia M. Okano & T. Okano, 1985 is a subgenus of Polycaena Staudinger, 1886

Monotypic genus Takashia M. Okano & T. Okano, 1985 (type and the only species Timelaea nana Leech,

1892) is a close sister to Polycaena Staudinger, 1886 (type species Polycaena tamerlana Staudinger,

1886) (Fig. 14 magenta and blue). To avoid yet another not truly distinct monotypic genus, we propose to

place Takashia M. Okano & T Okano, 1985 as a subgenus of Polycaena Staudinger, 1886.

Corrachia Schaus, 1913 is a subgenus of Styx Staudinger, 1876

Two monotypic genera Styx Staudinger, 1876 (type and the only species Styx infernalis Staudinger, 1875)

and Corrachia Schaus, 1913 (type and the only ip | reaireeats Hamearis el el ar til

. . Styx infernalis|7280|Peru|2011

species Corrachia leucoplaga Schaus, 1913) are area Corrachia leucopiagal7281|Costa Rical2004

. . ‘s Olycaena tamernana Inala

close sisters (Fig. 14 red and green). Hindered by | | ra eure eee ao: SOP Ghipalgls

. . . A mene olycaena lama 2/Chinalo

substantial difference in appearance, their close Teenie er abe ala

inshi i 0.9 Dodona deodatal 1807 1F04|Chinalold

kinship has been revealed by DNA sequencing and |. Barone aed ta eee wale,

- Dodona durgal18071b07|Chinalold

USiboges (Espeland et al) abisara fyllal1807 1E02|Myanmar|2002

0.32 Stiboges nymphidia|18071005|Thailand|old

Zemeros Femeros flagyas|18071C11|Myanmar|2001

Zemeros emesoides|18071C12|Malayal1990

discussed in detail (Espeland et al. 2015). Their COI

barcodes differ by only 7.4% (49 bp). Here, we take

h lim} h

: See sti a pe ; 7 a voce, ceca Fig. 14. Nemeobiini genera: Hamearis (black), Styx

y proposing that 1s BOC OAS Oe au sw (red and green), Polycaena (blue and magenta),

subgenus of Styx Staudinger, 1876. Considering these Dodona (olive), new genus (Espeland et al. 2015),

two close relatives to be congeneric is more revealing Stiboges (cyan) and Zemeros (purple).

about their evolutionary relationship than keeping them in two not prominently distinct monotypic genera.

Elo

Hades Westwood, 1851 is a junior subjective synonym of Methone Doubleday, 1847

Despite their difference in appearance, monotypic genus Methone Doubleday, 1847 (type and the only

species Papilio cecilia Cramer, 1777) is genetically close to Hades Westwood, 1851 (type species Hades

noctula Westwood, 1851) (Fig. 15 brown and gray). Therefore, we propose that Hades is a junior

subjective synonym of Methone and deduce that the phenotypic difference between Hades and Methone

was caused by rapid evolution possibly driven by selection for mimetic appearance of their type species,

each in a different mimetic complex.

Methone Doubleday, 1847 receives 9 species from Euselasia Hiibner, [1819]

We find that Euselasia Hiibner, [1819] (type species Euselasia gelaena Hiibner, [1819], which is Papilio

gelon Stoll, 1787) as currently defined is paraphyletic with respect to Methone Doubleday, 1847 (type

species Papilio cecilia Cramer, 1777), and a number of Euselasia species belong to the clade with

Methone cecilia (Fig. 15 red and brown). The type species of Euselasia and Methone are genetically

distant from each other, e.g., their COI barcodes differ by 11.5% (76 bp), which is typical for species in

different genera. Therefore, to restore the monophyly, instead of placing Methone in Euselasia, we

transfer Methone-clade species (Fig. 15 red) and their phenotypically close relatives from Euselasia to

Methone to form the following new combinations: Methone euploea (Hewitson, [1855]), Methone

eucerus (Hewitson, 1872), Methone hypophaea (Godman & Salvin, 1878), Methone eubule (R. Felder,

1869), Methone onorata (Hewitson, 1869), Methone authe (Godman, 1903), Methone dolichos

(Staudinger, [1887]), Methone baucis (Stichel, 1919), and Methone eucrates (Hewitson, 1872).

Erythia Hiibner, [1819] and Marmessus Hiibner, [1819] are valid genera

Considered junior subjective synonyms of Euselasia Htibner, [1819] (type species Euselasia gelaena

Hiibner, [1819], which is Papilio gelon Stoll, 1787), Erythia Htibner, [1819] (type species Papilio

labdacus Stoll, 1780) (Fig. 15 olive) and = ass Pendle ra anus aoe eo Tracer) eaat

Hamearis lucinal]?282|Austria]19

Marmessus Hiibner, [1819] (type species r eee aes eee

Papilio lisias Cramer, 1777) (Fig. 15 cyan) a Euselasia hrysippe|727elcosta Rcal2021

. . : . Pelolasia| |. Euselasia eumedia a Te eee aad

are genetically distant from Euselasia (Fig. 15 Euselasia argenteal19036C08|Costa Rical201

i i ee amphidedta| 90360 :0|costa rica{2021

blue) and form two prominent clades in the Se SERS TTR IEEE ENS eee

tree that should be assigned a genus rank (Fig. wetonel! °

15 Th f ete pape d sche intial Ure acretsecle pth rls ELML Ae 2

USe@lasla a aea anama

). ererore, Me reinstate ryt la all wt Euselasia Sen anace icone Rica|2006

Marmessus as valid genera. As a result, we : = Euselasia Dulatal 19036A09|brazl Paral 1978

. ° , e . . Euse 19036412|G 2006

split Euselasia as it is currently circumscribed aba — Euselasia mystical is036409| costa Rical 2008

2 ’ 7 0.7: Euselasia Ee eneecer Seema

into 4 genera. Indeed, Euselasia sensu lato is a pars Euselasia proculal 19036810|Costa Rical2004 _

0.68 SS RR ertecr tects Jam|t97¢

tribe rank taxon by its genetic divergence Euselasia pusilla| 19036CO8|El Salvador|1033

comparable to the divergence between Taxila Pete

Doubleday, 1847 (in the subtribe Abisarina a Euselasia labdacus|19036C07|Bolivia|2003

e| huecydide 6 razil:

Lee Se edecese mined | Heenan cet oes

Stichel, 1928) and Hamearis Hitibner, 1819 (in ervthia (| Euselasia Fee eer tela Mller —

. oe . Tytha UsSélasla moda ne O54 Rica

the subtribe Nemeobiina Bates, 1868) (Fig. 15 * Euselasia cabsleuce 19036008) French Guiana]1988

; . . ® c ee leet cea eae beet

US@lasla4 opalescens cUadear

black). However, after this split, species s Euselasa dagait903¢ez0[8ra2i'es12969

2, 7 bs . is USe@lasSla @xtensa 2|Peru

remaining in Euselasia are not monophyletic | Euselasia oaxacensis|19036G07|PT|Costa Rical 1965

“aE 3 3 Euselasia lisias|19036G05|Guyana|2001

(all named Euselasia in Fig. 15) and there are |“ Euselasia euphaes|19036401|Peru|1998

. . . Maculasia nate

other prominent clades in the tree. The level in | = elms <hails]2303504)Peru| 1995 ae

: : : : 0.8 Euselasia gelon|19035606|Frenc Ginanal 1988

Euselasia corduena|19035G603|Peru:Cusco|2016

the tree with these clades is visually obvious, | duena| | |

4 ; , Euselasia q Euselasia phedica|19035E11|Guyana|2000

a Euselasia perisama|19035F02|PT|Peru|1994

because they diversified at about the same eee

time (Fig. 15 different colors, with names ee Bc pat oma he ea eer Mee

: p) Euselasia brevicauda|19036 5FO9 (Braz IRO[1991

along branches). These nine clades supported | Fig. 15. Euselasiinae genera: Pe/olasia gen. n. (magenta), Methone

by prominent branches of about the same (brown, gray, and red), Myselasia gen. n. (green), Eurylasia gen. n.

| i d Th ; (pink), Erythia (olive), Marmessus (cyan), Maculasia gen. n.

ength correspond to genera. Ihe type species | (orange), Euselasia (blue), Eugelasia gen. n. (purple). Compare

of Psalidopteris Hitibner, 1823, P. nycha Euselasiinae divergence to that of Taxila and Hamearis (black).

Hiibner, 1823, which is a junior subjective synonym of Hesperia thucydides Fabricius, 1793, is in the

Erythia clade, making Psalidopteris a junior subjective synonym of Erythia. Thus, four of these major

clades have names and five do not, corresponding to five new genera that are proposed next.

Pelolasia Grishin, new genus

http://zoobank.org/AYAC96C6-B5D0-4 16D-9CS5E-0853B7BD8339

Type species. Eurygona pelor Hewitson, [1853].

Definition. Currently within Euselasia Hubner, [1819] (type species Euselasia gelaena Hibner, [1819],

which is Papilio gelon Stoll, 1787) but is not monophyletic with it (Fig. 15 magenta). Instead, an

independent prominent lineage originating in the early radiation of Euselasiini Kirby, 1871 and a poorly

supported sister to the clade that includes Methone Doubleday, 1847 (type species Papilio cecilia Cramer,

1777) (Fig. 15 brown, gray and red) and Erythia Hiibner, [1819] (type species Papilio labdacus Stoll,

1780) (Fig. 15 olive). Therefore, it constitutes a new genus. Union of Stichel (1928) groups Peloriformes,

Eusepiformes, Melaphaeoformes, Argenteoformes and Eubuliformes (in part). Characterized by wings

rounder than in relatives, hindwing with undulate outer margin and checkered fringes, ventrally with

postdiscal reddish line and a row of marginal black dots framed with white, reddish, or both; or nearly

immaculate wings below, silvery to golden, may be with marginal black dots on hindwing and brown

postdiscal line; or with 3—5 brown narrow bands over broader than bands pale background, lacking

isolated large eyespots or dotted discal pattern, or with a row of small eyespots along hindwing margin.

The following combination of nuclear genome characters is diagnostic: cne2559.1.3:T87C, cne3355.

8.1:T362C, cne1314.4.1:A351G, cne2022.5.2:C448A, and cne599.10.1:T5841A.

Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type

Species name and its former genus name: Pelo|r| + [Euse]/asia to keep the word at 9 letters, which is the

number of letters in Euselasia and the number of genera Euselasia is being split into.

Species included. The type species, Eurygona amphidecta Godman & Salvin, 1878, Eurygona argentea

Hewitson, 1871, Eurygona artos Herrich-Schaffer, [1853], Eurygona aurantia Butler & Druce, 1872,

Eurygona bettina Hewitson, 1869, Eurygona candaria Druce, 1904, Eurygona cataleuca R. Felder, 1869,

Eurygona chrysippe Bates, 1866, Eurygona euboea Hewitson, [1853], Eurygona eumedia Hewitson,

1853, Eurygona eumenes Hewitson, 1853, Euselasia eupatra Seitz, 1916, Eurygona eusepus Hewitson,

1853, Eurygona fervida Butler, 1874, Euselasia hahneli Staudinger, [1887], Euselasia ignitus Stichel,

1924, Eurygona mazaca Hewitson, 1860, Erythia melaphaea Hibner, 1823, Eurygona mirania Bates,

1868, Euselasia misteriosa Salazar & J. Vargas, 2019, Euselasia nytua J. Hall & Willmott, 2009,

Euselasia pellonia Stichel, 1919, Euselasia rubrocilia Lathy, 1926, and Euselasia seitzi Lathy, 1926.

Parent taxon. Subfamily Euselasiinae Kirby, 1871.

Myselasia Grishin, new genus

http://zoobank.org/61CDF024-ADCB-49D0-82FA-9956EA9E5553

Type species. Eurygona mys Herrich-Schaffer, [1853].

Definition. Currently within Euselasia Hubner, [1819] (type species Euselasia gelaena Hubner, [1819],

which is Papilio gelon Stoll, 1787) but is not monophyletic with it (Fig. 15 green). Instead, an

independent prominent lineage in the same clade with and of the same rank as Methone Doubleday, 1847

(type species Papilio cecilia Cramer, 1777) (Fig. 15 brown, gray and red), thus is a new genus. Largely,

the Hygeniiformes group of Stichel (1928), sharing its diagnostic characters. Hindwing rounded, below

with a kinked-L-shaped central reddish band and an eyespot in the middle by the margin, but without

well-developed marginal longitudinal dashes; forewing without eyespots and with 1-2 narrow reddish

bands. The following combination of nuclear genome characters is diagnostic: cne23605.2.5:C468T,

cne13338.5.3: A334C, cne123.2.3:C1009T, cne18035.2.1:C172A, and cne81.14.5:A756G.

Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type

Species name and its former genus name: Mys + [Eus]le/asia to keep the word at 9 letters, which is the

number of letters in Euselasia and the number of genera Euselasia is being split into.

Species included. The type species, Eurygona alcmena Druce, 1878, Eurygona athena Hewitson, 1869,

Eurygona cafusa Bates, 1868, Euselasia crinon Stichel, 1919, Euselasia cucuta (Schaus, 1902), Euselasia

cyanofusa J. Hall & Willmott, 1998, Euselasia eberti Callaghan, 1999, Euselasia ella Seitz, 1916,

Eurygona eulione Hewitson, 1856, Euselasia eustola Stichel, 1919, Euselasia gradata Stichel, 1927,

Eurygona hieronymi Salvin & Godman, 1868, Papilio hygenius Stoll, 1787, Euselasia illarina J. Hall,

Willmott & R. Busby, 1998, Eurygona inconspicua Godman & Salvin, 1878, Euselasia janigena Stichel,

1919, Euselasia jigginsi J. Hall & Willmott, 1998, Eurygona leucon Schaus, 1913, Euselasia mapatayna

J. Hall & Willmott, 1998, Euselasia marica Stichel, 1919, Eurygona mystica Schaus, 1913, Euselasia

nauca J. Hall & Willmott, 1998, Euselasia pance Callaghan, 1999, Eurygona procula Godman & Salvin,

1885, Euselasia pseudomys Callaghan, 1999, Euselasia pullata Stichel, 1927, Eurygona pusilla R. Felder,

1869, Euselasia rhodon Seitz, 1913, and Eurygona sergia Godman & Salvin, 1885.

Parent taxon. Subfamily Euselasiinae Kirby, 1871.

Eurylasia Grishin, new genus

http://zoobank.org/44399705-A 1CF-43 AF-B35E-F510E964BDE0

Type species. Eurygona euryone Hewitson, 1856.

Definition. Currently within Euselasia Hubner, [1819] (type species Euselasia gelaena Hibner, [1819],

which is Papilio gelon Stoll, 1787) but is not monophyletic with it (Fig. 15 pink). Instead, an independent

prominent lineage in the same clade with and of the same rank as Methone Doubleday, 1847 (type species

Papilio cecilia Cramer, 1777) (Fig. 15 brown, gray and red), hence is a new genus. It is diagnosed by its

distal “4 of hindwing dusted pale-yellow below, large round marginal eyespot in the middle of it, and

either large eyespot in the middle by forewing margin below or diagonal orange patch on forewing above.

The hindwing tornus is pale above in some species. The following combination of nuclear genomic

characters is diagnostic: cne9878.8.1:C146A, cnel78.3.20:C595T, cne178.3.20:TS596C, cne7676.26.2:

T31C, and cne593 1.2.1:A478G.

Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type

Species name and its former genus name: Eury[one] + [Euse]/asia to keep the word at 9 letters, which is

the number of letters in Ewse/asia and the number of genera Euse/asia is being split into.

Species included. The type species, Eurygona effima Hewitson, 1869, Euselasia thusnelda Moschler,

1883, and Eurygona eunaeus Hewitson, 1855.

Parent taxon. Subfamily Euselasiinae Kirby, 1871.

Maculasia Grishin, new genus

http://zoobank.org/20121C52-93F8-4540-9070-FF2971B2D960

Type species. Euselasia albomaculiga Callaghan, 1999.

Definition. The genus is sister to Euselasia Hiibner, [1819] (Euselasia gelaena Hiibner, [1819], which is

Papilio gelon Stoll, 1787) and is prominently distinct from it genetically (Fig. 15 orange). Therefore, it

constitutes a new genus. Distinguished from its relatives by the characters given on pages 1047-1048 and

illustrated in Figs. 3-6, 46-47 for Euselasia albomaculiga by Callaghan (1999). In brief, its wings are

rounded, without blue scaling, its forewings are with a large pale spot, and its ventral hindwing

submarginal black spots are surrounded by grayish-white (not at the end of orange rays, except the central

large one that is framed by yellow basad), and have yellow streaks along the veins not between them;

valvae narrowing to a point, not bilobed, as long as tegumen with uncus, slightly wider than aedeagus,

vinculum angled in lateral view, with spurs in the middle directed caudad. Additionally, the following

combination of nuclear genomic characters is diagnostic: cne2298.2.2:A2104A (not C), cne5129.1.5:

A833A (not G), cne2685.14.3:A934A (not T), cnel1095.7.22:G1463G (not A), cne4870.1.18:T552T (not

A), cne2885.9.10:A4270C, cnel547.14.4:A2756G, cnel696.1.1:A2529T, cne7231.10.9:T567C, and

cne1036.6.11:G3200C.

Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type

Species name and its former genus name: [albo|Macu|liga] + [Euse]/asia to keep the word at 9 letters,

which is the number of letters in Euselasia and the number of genera Euselasia is being split into.

Species included. Only the type species.

Parent taxon. Subfamily Euselasiinae Kirby, 1871.

Eugelasia Grishin, new genus

http://zoobank.org/36F889D 1 -2E27-40BB-95B9-761903 LA223E

Type species. Eurygona eugeon Hewitson, 1856.

Definition. This taxon is sister to the clade formed by Euselasia Hiibner, [1819] (Euselasia gelaena

Hiibner, [1819], which is Papilio gelon Stoll, 1787) and Maculasia gen. n. and is prominently distinct

from it genetically (Fig. 15 purple); hence it is a new genus. It is distinguished from its relatives by its

inverted-drop shaped hindwing with extended anal lobe, as well as by its solid-brown color above and

lack of eyespots below, fringes dark, legs yellow; and the following combination of nuclear genomic

characters: cne1999.2.1:C203A, cnel015.3.2:T555C, cne2803.19.1:G88A, cne5471.1.1:T261C, and

cne703.2.8:AI1414C.

Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type

Species name and its former genus name: Euge[on] + [Euse]/asia to keep the word at 9 letters, which is

the number of letters in Ewse/asia and the number of genera Euse/asia is being split into.

Species included. The type species and Euselasia brevicauda Lathy, 1926.

Parent taxon. Subfamily Euselasiinae Kirby, 1871.

Eunogyrina Grishin, new subtribe

http://zoobank.org/CB55411C-D989-4F77-87C6-9E2E2023C608

Type genus. Eunogyra Westwood, 1851.

Definition. Placed in Mesosemiina Bates, 1859 on the basis of phenotypic assessment (Hall 2003), the

two genera Eunogyra Westwood, 1851 (type species Eunogyra satyrus Westwood, 1851) and

Teratophthalma Stichel, 1909 (type species Mesosemia phelina C. & R. Felder, 1862) form a clade that is

sister to the clade formed by Mesosemiina and Napaeina Hall, 2003 (Figs. 16, 27), and therefore

Eunogyra taken together with Teratophthalma constitute a substribe. The description and diagnostic

characters of this new subtribe are as those given for Eunogyra on page 463 by Westwood (1851) and

page 90 (illustrated in Fig. 16) by Stichel (1910), and for Teratophthalma on pages 76—77 (illustrated in

Fig. 11) by Stichel (1910). In brief, the subtribe belongs to Mesosemiini (see Hall (2003) for genera

Eunogyra and Teratophthalma), and is diagnosed by the following combination of characters: wings

without multiple narrow bands, eyespots either at the end of forewing discal cell or along wing margins;

genitalic valvae short (as long as tegumen) and triangular, simple with rounded or pointed apex, but

without two separated posterior projections with narrow intervening section, pedicel unsclerotized in the

middle or ventrally split (or both).

Genera included. The type genus and Teratophthalma Stichel, 1909.

Parent Taxon. Tribe Mesosemiini Bates, 1859.

Semomesia Westwood, 1851, Mesophthalma Westwood, 1851,

Perophthalma Westwood, 1851 and Leucochimona Stichel, 1909

are junior subjective synonyms of Mesosemia Hiibner, [1819]

The following four genera render Mesosemia Hiibner, [1819] (type species Mesosemia phicoclessa

Hiibner, [1819], which is Papilio philocles Linnaeus, 1758) paraphyletic, in agreement with Seraphim

(2018): Semomesia Westwood, 1851 (type species Papilio croesus Fabricius, 1777), Mesophthalma

Westwood, 1851 (type species Mesophthalma idotea Westwood, 1851), Perophthalma Westwood, 1851

(type species Mesosemia tenera Westwood, 1851 which is Papilio tullius Fabricius, 1787) and

Leucochimona Stichel, 1909 (type species Papilio philemon Cramer, 1775, homonym, current name

Mesosemia icare Hubner, [1819]) (Fig. 16). Our genomic phylogeny indicates that all these genera (Fig.

b f other li five lee Fore oithalma culliizigos|Peruigols

a num er Or olMer pee? curren y: ‘Teratophthalma Bc iRowiedene eedcderiiaeat

; Teratophthalma phelina|18121606|Ecuador|1969

placed in Mesosemia (Fig. 16 blue) Ectosemia|— esosemia stall 19037411 [Peru2015 vise

esosemla eumene Uy¥ana

diversified rapidly around the same Mesosemia Sun Wert ade Seo a ee csue aeee

. . . - M 19037E11|B RI/1995

time, leading to the comb-like structure i esosemia ibycus| 19037807 [Peru|2016

2) iC ’ : 7 Perophthalma lasus|18072606|Costa Rica|2006

of the tree rather than a well-resolved | = genni Perophthalma tullius|18122410|Guyana| 1999

nF a i | Thi . . = Mesosemia harvey!|19037HO2|Costa Rica|2005

i Mesophthalma idoteal18122B801|Brazil:RO|1996

; IC otomous Pp y ogeny. i situation O i Seiomesis marisal 1819iHi0|Srazh ro 1994

D Semome esus|19038404|G 2000

IS commonly encountered In genomic = : Mesosemia albipuncta|18121G09|Costa Rica|2009

. e i : : ‘F J

analyses of butterflies (Li et al. 2019; Mesosemiina zs MOS GRETA SUBeranvaLienavANid|PaMamalIgEe

0. M - 19037B09|Peru]1995

Zhang et al. 2019a; Zhang et al. 2019d) Zen Mesosemia philoces|19037D02[auyanal2001

° ‘ Mesosemia melpia|19037CO07|Guyana|2000

and Suggests existence of levels in re Leucochimona anophthalma|18122409|Ecuador|1999

di 7. hs h 5 a f, Leucochimona Hie ae seas eedt pice ae

- Mesose 18072E09|Costa Rica|2012

cL ication t at can - nes or = Me cee Tiree a ueeereo Ie aeeeidese

classification. This radiation of CE ettame ethene pel tela

. Mesosemia ulrica|L9037HO6|Guyana|2000

Mesosemia and related genera Was eae, ; Mesosemia macella|19037H10|Brazil:RO|1991

dated to about 15 Mya (Seraphim et al. x

2018) and therefore corresponds to the Fig. 16. Mesosemiini subtribes: Eunogyrina subtrib. n. (black),

2 : . Napaeina (gray) and Mesosemiina with genera: Ectosemia gen. n.

diversification of a genus. All these P Saas ate: o S

pony ; (magenta), Endosemia gen. n. (red), and Mesosemia (blue and other

Mesosemia-like genera are close to colors) with its synonyms Semomesia (green), Mesophthalma

each other both genetically (Fig. 16) (purple), Perophthalma (orange) and Leucochimona (olive).

and phenotypically, and we propose to treat Semomesia Westwood, 1851, Mesophthalma Westwood,

1851, Perophthalma Westwood, 1851 and Leucochimona Stichel, 1909 as junior subjective synonyms of

Mesosemia Hibner, [1819].

Inspection of the branching pattern in the tree (Fig. 16) reveals the next level: 1.e., diversification

of the subtribe Mesosemiina Bates, 1859 into 3 lineages. Their exact bifurcation order is not resolved

(support 0.4) due to closeness of these events in time (=rapid radiation) and possible incomplete lineage

sorting or introgression early in their origins. These 3 lineages correspond to genera, although currently

these species are included in the paraphyletic genus Mesosemia. One of these lineages is the genus

Mesosemia (sensu lato) that includes all available names in the group as synonyms (as detailed above),

together with Diophtalma Boisduval, 1836 (type species Diophtalma telegone Boisduval, 1836), which is

already considered to be its synonym. Therefore, the other two lineages do not have names and represent

new genera that are defined below. Furthermore, a possibility of additional new genera in Mesosemiina

has been suggested, although not yet formalized (Seraphim et al. 2018).

Ectosemia Grishin, new genus

http://zoobank.org/8AB8B269-CAE8-417D-BFOD-E46043B20CBE

Type species. Papilio eumene Cramer, 1776.

Definition. Species in this clade are currently in Mesosemia Hiibner, [1819] (type species Mesosemia

Phicoclessa Hiibner, [1819], which is Papilio philocles Linnaeus, 1758), but are quite distant from it

falling much outside of the Mesosemia radiation and originating during the earlier round of radiation (Fig.

16). Therefore this clade is a taxon of the same rank, a genus. This new genus differs from Mesosemia by

the following combination of characters: hindwing not lobed in the middle, each wing with 2 broad and

straight parallel bands: discal and postdiscal, bands do not encircle forewing eyespot as in most

Mesosemia, but discal forewing band bends distad at costa, no striations, discal cell eyespots developed

ventrally on both wings, elongated along the cell, with 3 white spots inside (sometimes 2 on hindwing).

Furthermore, the following combination of nuclear genomic characters is diagnostic: cne7048.1.3:

T2251C, cne3658.2.1:A638T, cne2957.11.4:A952G, cne3658.2.1:A608C, and cne3461.2.10:C1462A.

Etymology. The name is a feminine noun in the nominative singular formed by replacing prefix "Meso-"

with "Ecto-" in the name of the former genus for these species.

Species included. The type species, Mesosemia decolorata Lathy, 1932, Mesosemia erinnya Stichel,

1910, and Mesosemia steli Hewitson, 1858.

Parent taxon. Subtribe Mesosemiina Bates, 1859.

Endosemia Grishin, new genus

http://zoobank.org/AA 1AC39C-2696-46A8-9927-2A8CDBB14D28

Type species. Papilio ulrica Cramer, 1777.

Definition. Species in this clade are currently in Mesosemia Hiibner, [1819] (type species Mesosemia

Phicoclessa Hiibner, [1819], which is Papilio philocles Linnaeus, 1758), but are quite distant from it

falling much outside of the Mesosemia radiation and originating during the earlier round of radiation (Fig.

16). Therefore this clade is a taxon of the same rank, a genus. This new genus differs from both

Mesosemia and Ectosemia gen. n. by the following combination of characters: hindwing not lobed in the

middle, forewing eyespot not elongated along discal cell, discal band (if developed) narrow, bands do not

encircle forewing eyespot, or bands diffuse and poorly formed. Furthermore, the following combination

of nuclear genomic characters is diagnostic: cne703.2.8:T903C, cne1411.6.4:T1025A, cne2651.14.5:

A4602G, cne12205.6.2: G835A, and cne8028.2.1:T1787A.

Etymology. The name is a feminine noun in the nominative singular formed by replacing prefix "Meso-"

with "Endo-" in the name of the former genus for these species.

Species included. The type species and Mesosemia macella Hewitson, 1859.

Parent taxon. Subtribe Mesosemiina Bates, 1859.

Eucorna Strand, 1932 is a valid genus

Presently, Eucorna Strand, 1932 (type species Voltinia sanarita (Schaus, 1902)) is a junior subjective

synonym of Voltinia Stichel, 1910 (type species Esthemopsis (?) radiata Godman & Salvin, 1886), but is

not monophyletic with it, and is sister to all other Napaeina Hall, 2003 except Hyphilaria Hiibner, [1819]

(type species Hyphilaria nicia Hiibner, [1819]) (Fig. 17). Hence, Eucorna is a valid genus.

Reassessment of Voltinia Stichel, 1910 and Napaea Hiibner, [1819]

Our genome-level phylogeny combined with phenotypic assessment of species missing from the tree

reveals that Voltinia Stichel, 1910 (type Hyphilaria Hyphilaria anthias|18122804|/Guyana/1999

ies Esthemopsis (2) radiata Godman & SS eine ara TaS erste

ae s PUEOH ote, 7 radiata 0 a " eto Voltinia sanarita]18122C01|Brazil:RJ|1996

Salvin, 1886) consists of only two species: Cremna | r {Napaea heterova|16122608/[Guiyana[2c 50 Ds

< 5 ° apaea actoris|19038B0 raZzl va] nia a9

the type and Voltinia theata Stichel, 1910 r

1 1 Ith lat 180/78CO9|T|1896|2MHB

(Fig. 17 gray). Based on this phylogeny, we i Themicla corals] 184 2900 2jpanamalt977

transfer all other species presently in Voltinia sthomiotal lz Ithomiola Horsls feral) iBia3¢ G3|euyana{zo00

" ; : omiola orpheus 22 eru

to Napaea Hubner, [1819] (type Species 7 Teen ree a TE EEe Tien sentaleecader (i899

Cremna eucharila Bates, 1867) (Fig. 17 red 098 pew conus 1 (Seraphim etal) TETAS nlania thasus|1007E03| osta Rical2 2012

t bl ) Th f ll ¢ : d bi ti 7 ones ea See ae enelea eat ce

0.92 apaea calitra ies Cuador

0 uc). € 10 Owing revised COMOINAUONS ‘ volta ded ent tg cba rata ee eae :Sonora|2003

are proposed for them: Napaea danforthi A. eee ee esses alteatan 38 b3}ecuador| 1909.» mae

° Volt tot 19038C01|M ‘SLP|1982

Warren & Opler, 1999 (as_ originally 7 vottinia imbralie{22cOs| Costa Rica] 19:

: ee eae Peruyisve

he 22 eB

proposed!), Napaea dramba (J. Hall, Robbins ne ~ Nepaes fumbe isosesoe era Ral is 1979

° , apaeéda JOnvilea PazZil:

& Harvey, 2004), Napaea Sanarita (Schaus, 782 Napaea Zikani[19038804|Brazil:RI|/1959

1902). N, Stichel. 1910 oe Voltinis SoroscalABISeCOD|BrasLRy eos

), apaed = agroeca 1cnel, > Napaea aie igioon 1 tiGusanal.ono”

Napaea tumbesia J. Hall & Lamas, 2001 (as Pi Ngpaea sucharlial 18072FUi|costa Ries 2013

originally proposed!), Napaea umbra | Fig. 17. Napaeina genera: Hyphilaria (black), Eucorna (magenta),

: le, olive and gray), /thomiola (green and cyan), new

(Boisduval, 1870), Napaea phryxe (C. & R. | Cremna (purple gray

enus | (Seraphim et al. 2018) (orange) and NM blue and red).

Felder, 1865), Napaea cebrenia (Hewitson, See yore) x temic

[1873]), Napaea loxicha (R. G. Maza & J. Maza, 2016), Napaea maya (J. Maza & Lamas, 2016), Napaea

necaxa (R. G. Maza & J. Maza, 2018), Napaea totonaca (R. G. Maza & J. Maza, 2016).

Tonotus Hall, 2005 and Voltinia Stichel, 1910 are subgenera of

Cremna Doubleday, 1847, which is a valid genus

A genus comprised of two species, Voltinia Stichel, 1910 (type species Esthemopsis (?) radiata Godman

& Salvin, 1886) (Fig. 17 gray) is a close sister to Cremna Doubleday, 1847 (type species Papilio actoris

Cramer, 1776), which also consists of only two species: the type and Cremna heteroea Bates, 1867 (Fig.

17 purple); and Jonotus Hall, 2005 (type and the only species Hamanumida alector Geyer, 1837) (Fig. 17

olive) is sister to them combined. Hence, we reinstate Cremna as a valid genus (not a synonym of

Napaea), and due to genetic similarities place Jonotus Hall, 2005 and Voltinia Stichel, 1910 as its

subgenera.

Hermathena Hewitson, 1874 is a subgenus of Ithomiola C. & R. Felder, 1865

Despite its mostly white coloration, much different from its relatives, Hermathena Hewitson, 1874 (type

species Hermathena candidata Hewitson, 1874) (Fig. 17 cyan) clusters closely with Jthomiola C. & R.

Felder, 1865 (type species /thomiola floralis C. & R. Felder, 1865) (Fig. 17 green), which even now

includes species considerably different in appearance (Hall 2005). The male genitalia of these taxa are

rather similar as illustrated by Hall (2005). To achieve a more internally consistent classification, we

place Hermathena Hewitson, 1874 as a subgenus of /thomiola C. & R. Felder, 1865.

Lucillella Strand, 1932 is a subgenus of Esthemopsis C. & R. Felder, 1865

Lucillella Strand, 1932 (type species Lucilla camissa Hewitson, 1870) (Fig. 18 pale blue) and

Esthemopsis C. & R. Felder, 1865 (type species Esthemopsis clonia C. & R. Felder, 1865) (Fig. 18 gray)

are closely related sisters that form a clade prominently separated from others (Fig. 18) and sister to

Mesene Doubleday, 1847 (type species Papilio phareus Cramer, 1777) (Fig. 18 olive). Because Mesene is

already more diverse than Lucillella and Esthemopsis combined, to achieve better consistency of

classification, we propose treating Lucillella Strand, 1932 as a subgenus of Esthemopsis C. & R. Felder,

1865.

Xynias Hewitson, 1874 is a junior subjective synonym of Mesenopsis

Godman & Salvin, 1886, which along with Xenandra C. & R. Felder, 1865

are subgenera of Symmachia Hiibner, [1819], that contains

Stichelia J. Zikan, 1949 as its junior subjective synonym

First, Xynias Hewitson, 1874 (type species Xynias cynosema Hewitson, 1874, which is a subspecies of

Esthemopsis lithosina Bates, 1868) (Fig. 18 bright green) is in the same clade with Mesenopsis Godman

& Salvin, 1886 (type species —o

Limnas (?) bryaxis Hewitson,

1870) (Fig. 18 purple) and is

closely related to the type species

of Mesenopsis genetically and

possesses, similar to it, elongated

wing shape. Therefore, we place

Xynias as a junior subjective

synonym of Mesenopsis. Dis-

06|Peru;Ccusco|2009

Lucillalla pomposa ele eer oc] yictised

12 cliador|2

Lucillella camissal19

Mesene phareus|18072E07|Costa Rical2006

Mesene pyrinpe|19027F11|Brazil:RJ JAlold

Mesene veleda|1so0 yay ue Zil:

hHeasene margaretta|1902 aeeAlole i 2002

Esthemopsis aeolia|19028410|Suriname

adehimastrum argentea argenteal19028B i rete Ri

Chimastrum argentea carnutes| 19028B03|Panama

Mesene monostigmal19027G0

Mesen De ee FoI Es 27

Symmachia trica mel 18072F1

Mesenopsis bryaxis|1902

Masta ae elapse

Xynias lithosina oe

jesene A

Woe 5 ee

nel A04 cua a

O08

Mesenopsis jordanal18o78 eu 7[ald

Svinte probetor| 3.908803! Tnidadl1935

similar wing patterns in these

Species are caused by their

involvement in different mimicry

complexes.

Second, Stichelia J. Zikan,

1949 (type species Amarynthia

bocchoris Hewitson, 1876) (Fig.

Symmachia. technema walla

Syme miro aaa

sti relate oma 9

* Stic hella ag fu al

Symmac ae Lele

Symmachia elinas

symmachia phaedralts

7 Symmachia xype

Sy mmachi

colts Reslat 1995

arate Puede

cual virgatu

Asymma Symmachia satanal1

: * es 0.38 5 oun sunreenne 28D09)/Guyana

18 bright orange) originates — Comphotis sanes|19031403|Guyanal2000,

ithin Symmachia — Hiib Cs seperate ee:

within VYVNMNACHIA uoner, phaenochitonia cta [igo

[1819] (type species Symmachia

probetrix Hubner, [1819], which

is Papilio probetor Stoll, 1782)

sensu stricto (Fig. 18 blue). To

restore monophyly of Symmachia

sensu. stricto and_ considering

genetic closeness, we propose

treating Stichelia J. Zikan, 1949

as a junior subjective synonym of

Guyana]2006 |

ee Costa Rica|2012

= itsessines iWferiwisy| il es 78 ial ivevaih

sks

we saan a onla pyrsodes

aenoc tonte. cin tel Se00 aR ena nS

Pterographium sicoral19028F12|Brazil:ES|197

Panaropsis t ae ieee tlt Peru ee

Panaropsis elegans|19028F11|/Panamali984

Pirascca (pee oes |T 9028F02 Pane a

vas ee Mere loris|18072 orzoie

1 Alethea p dias| 9129E02/Ecua

Birger Styriotes [18072 O8 Cost a Rica re

Piranets sag BES eh ea oO ee nal200

Piraseca at Heals 9028E0 Guianal1992 Fa

Piraseca phoenicural19028F |Ecua or|200

Pirascca suapure vittata| 18078B805/T|Brazil:PAlold =

Argyrogrammana stilbe[190 agnor lr gt Tae iva dre de pigs |1990 és

Arayrogrammana occidentalis| 1902 éléu an nal 200 S

Arqyrogrammana banne[1s02 FO? sia al[19 a

Argqyragrammana praestigiosal1902 elt |Peru: RdD |1991 =5

Fig. 18. Symmachiini genera marked on the tree, including Tigria gen. n. (violet)

and Asymma gen. n. (magenta). Different colors denote different genera and

Sy mmachiini

Ptero-

gvaphivm :

Argyrogrammana

0.02 0.72

aeamesntontl

inconsistencies with the current classification.

Symmachia Hiibner, [1819].

Third, Xenandra C. & R. Felder, 1865 (type species Xenandra heliodes C. & R. Felder, 1865,

which is currently a junior subjective synonym of Limnas agria Hewitson, 1853) (Fig. 18 pale pink), falls

within a rapid radiation that dates past the diversification of most Symmachiini Reuter, 1896 genera. We

consider the radiation that led to the origin of Xenandra to represent the diversification within the genus

Symmachia, (Fig. 18 blue clade labeled with the name) and therefore propose treating Xenandra C. & R.

Felder, 1865 as a subgenus of Symmachia Hubner, [1819] because it forms a prominent clade within it.

Lastly, the clade with Mesenopsis (includes Xynias and Symmachia tricolor Hewitson, 1867)

originates early in the radiation of Symmachia and is not prominently distinct from it. Therefore, we treat

Mesenopsis Godman & Salvin, 1886 as a subgenus of Symmachia Hiibner, [1819]. As a result, genus

Symmachia (Fig. 18 blue, purple, bright green, pale pink, bright orange) consists of 3 subgenera:

Symmachia, Xenandra, and Mesenopsis. Finally, some species currently in Symmachia fall outside this

genus (e.g., Fig. 18 violet and magenta clades). Two of such major clades do not have names and are

proposed as new genera here.

Tigria Grishin, new genus

http://zoobank.org/1665C4A 1-7AF7-4426-855A-1799CC9FS9E4

Type species. Mesene xypete Hewitson, 1870.

Definition. A sister clade to other Symmachia Hiibner, [1819] (type species Symmachia probetrix

Hiibner, [1819], which is Papilio probetor Stoll, 1782), but prominently distinct from it, more so than

Symmachia sensu stricto species are from each other (Fig. 18 blue), and genetically distant from them at a

level where other Symmachiini genera are defined (Fig. 18), is therefore a genus. It is similar to

Symmachia and distinguished from it by the following combination of characters (at least in males):

forewing costa rather straight, not concave, apex produced, hindwing typically with angular tornus;

antennae long, about % of forewing length; eyes bare; wings red-orange, bordered and partly striped at

least along forewing costa with black or dark-brown, dark areas could take half of wings, no pale spot

mid-costa. The following combination of nuclear genome characters is diagnostic: cne1935.6.1:A2889G,

cne3461.1.14: G922C, cne3437.1.9:A1260G, cne3461.1.14:A846G, and cne2170.2.1:T2751C.

Etymology. The name is a feminine noun in the nominative singular given for the red-orange tiger-striped

appearance of these species.

Species included. The type species, Polystichtis rubrica Stichel, 1929, Cricosoma phaedra Bates, 1868

and Metacharis elinas Rebillard, 1958.

Parent taxon. Tribe Symmachiini Reuter, 1896.

Asymma Grishin, new genus

http://zoobank.org/901035D7-E6D1-4F16-A 1FD-BA85C48AAC50

Type species. Symmachia virgatula Stichel, 1910.

Definition. This group of species is currently placed in Symmachia Hiibner, [1819] (type species

Symmachia probetrix Hiibner, [1819], which is Papilio probetor Stoll, 1782) but is not monophyletic with

it (Fig. 18). It originates early in the radiation of the Symmachiini core group, a possible, but weakly

supported sister to Phaenochitonia Stichel, 1910 (type species Papilio cingulus Stoll, 1790) sensu lato

(see below), and therefore is a genus. Similar to Symmachia and Tigria gen. n. and is distinguished from

them by the following combination of characters (at least in males): forewing costa concave in the middle,

as in most Symmachia but different from Jigria gen. n., wings red-orange to yellow, bordered and partly

striped or spotted at least along forewing costa with black or dark-brown, without white spots by the

forewing apex. The following combination of nuclear genome characters is diagnostic: cne4291.7.6:

A1077G, cne4291.7.6:A1064G, cne4291.7.6:A1051G, cne3461.2.5:A1310G, and cne3461.1.15:A3342C.

Etymology. The name is a feminine noun in the nominative singular, formed from the beginning of the

former genus name of these species, prefixing it with "a" for "not", because these species cannot possibly

belong to Symmachia: A + symma|chia].

Species included. The type species, Synapta arion C. & R. Felder, 1865, Symmachia giffordi P. Jauftret

& J. Jauffret, 2010, Symmachia hippodice Godman, 1903, Symmachia pardalis Hewitson, 1867,

Symmachia satana J. Hall & Harvey, 2007, Symmachia stigmosissima Stichel, 1910, and Symmachia

virgaurea Stichel, 1910.

Parent taxon. Tribe Symmachiini Reuter, 1896.

Chimastrum Godman & Salvin, 1886 is a

junior subjective synonym of Mesene Doubleday, 1847

Chimastrum Godman & Salvin, 1886 (type species Mesene argentea Bates, 1866) (Fig. 18 red) originates

within Mesene Doubleday, 1847 (type species Papilio phareus Cramer, 1777) (Fig. 18 olive), thus

rendering it paraphyletic. To restore the monophyly, instead of breaking Mesene (that is currently already

assembled from close relatives) into several genus-group taxa, we treat Chimastrum as a junior subjective

synonym of Mesene.

Mesene aeolia (Bates, 1868), new combination

Kept in Esthemopsis C. & R. Felder, 1865 (type species Esthemopsis

clonia C. & R. Felder, 1865) since its description, E. aeolia Bates, 1868

(type locality Brazil: Para) is not monophyletic with it, and instead is

sister to Mesene argentea Bates, 1866, the type species of Chimastrum

Godman & Salvin, 1886 that originates within Mesene Doubleday, 1847

(type species Papilio phareus Cramer, 1777) (Fig. 18, sequenced :

specimen in Fig. 19), implying Mesene aeolia (Bates, 1868) comb. nov. Fig. 19. Esthemopsis aeolia

Alethea Nielsen & Salazar, [2018] is a junior subjective synonym of Pirascca J. Hall &

Willmott, 1996, which is a subgenus of Pterographium Stichel, 1910, that contains

Panaropsis J. Hall, 2002 as its junior subjective synonym

The monotypic Alethea Nielsen & Salazar, [2018] (type and the only species Siseme pedias Godman,

1903) (Fig. 18 dark blue), Panaropsis J. Hall, 2002 (type species Panara elegans Schaus, 1920) (Fig. 18

cyan-green), Pirascca J. Hall & Willmott, 1996 (type species Papilio sagaris Cramer, 1775) (Fig. 18

green) and monotypic Pterographium Stichel, 1910 (type and the only species Prerographium aphaniodes

Stichel, 1910, which is Panara sicora Hewitson, 1875) (Fig. 18 brown) taken together form a prominent

clade in the tree that originated early in the diversification of Symmachiini Reuter, 1896 into genera and

therefore corresponds to genus rank. This genus gets the name Pterographium as the oldest available for

its species. This species-rich Pterographium sensu lato splits into 2 well-defined and strongly supported

but not very prominent clades (100% support for each clade, Fig. 18) that we treat as subgenera. The

nominal subgenus (upper clade in Fig. 18) contains Panaropsis as sister to its type species, and therefore

Panaropsis becomes a junior subjective synonym of Prerographium, along with some other species such

as Phaenochitona |sic| interrupta Lathy, 1932 (currently in Pirascca). The second subgenus of

Pterographium (lower clade in Fig. 18) is Pirascca, a close sister to monotypic and unusually patterned

Alethea, which we place as junior subjective synonym of Pirascca due to genetic similarities. We reason

that considerable phenotypic differences of Pterographium (Pirascca) pedias (Godman, 1903) comb.

nov. from other Pirascca are caused by rapid evolution towards a different mimicry complex.

Pterographium hypochlioris (Bates, 1868), new combination

Currently in Symmachia Hibner, [1819] (type species Symmachia

probetrix Hiibner, [1819], which is Papilio probetor Stoll, 1782), Emesis

hypochloris Bates, 1868 (type locality Brazil: Amazonas) is not

monophyletic with it and instead originates within Prerographium Stichel,

1910 (type species Pterographium aphaniodes Stichel, 1910, which is

Panara sicora Hewitson, 1875) sensu lato as it is defined above (Fig. 18,

sequenced specimen shown in Fig. 20), hence Pterographium hypochloris

(Bates, 1868) comb. nov. Fig. 20. Symmachia hypochloris

Comphotis Stichel, 1910 is a junior subjective synonym

of Phaenochitonia Stichel, 1910

Comphotis Stichel, 1910 (type species Cricosoma irroratum Godman, 1903) (Fig. 18 pale orange) is

paraphyletic with respect to Phaenochitonia Stichel, 1910 (type species Papilio cingulus Stoll, 1790)

(Fig. 18 bright cyan), but taken together, they form a prominent lineage consistent with other

Symmachiini Reuter, 1896 genera. Therefore we place Comphotis Stichel, 1910-XH-31 as a junior

subjective synonym of Phaenochitonia Stichel, 1910-IX.

Phaenochitonia florus (Fabricius, 1793), new combination

Currently in Mesene Doubleday, 1847 (type species Papilio phareus Cramer,

1777), Hesperia florus Fabricius, 1793 (type locality "Indiis", likely SE Brazil)

is not monophyletic with it and originates within Phaenochitonia Stichel, 1910

(type species Papilio cingulus Stoll, 1790) (Fig. 18 yellow, highlighted in violet,

sequenced specimen shown in Fig. 21), implying Phaenochitonia florus

(Fabricius, 1793) comb. nov. Distinctness of this species from Mesene came as

a surprise considering how well it was fitting in that genus on the basis of wing

colors, patterns and shape, even reminding of Mesene type species.

Reassessment of Ourocnemis Bethune-Baker, 1887 and Anferos Hiibner, [1819]

The tree of Helicopini Stichel, 1928 revealed a distribution of species between the two genera

Ourocnemis Bethune-Baker, 1887 (type species

Anteros axiochus Hewitson, 1867) and Anteros

=- Ourocnemis archytas|19129D04|Ecuador|2018

Anteros principalis|19029C08|Bolivia|2003

Anteros renaldus|18072D01|Costa Rica|2015

Anteros aerosus|19029B12|Brazil:RO|1991

Hiibner, [1819] (type species Papilio formosus Cramer,

1777) not as presently attributed (Fig. 22). A number of

species placed in Anteros were in the same clade with

Ourocnemis, confirming the suspicion of Hall (1998)

Anteros allectus|19029B06|Peru:Cuseoa|2014

Anteros chrysoprastal19029B08|Costa Rica|2011

Anteros micon|19029B09|Costa Rica|2009

Anteras formosus|19029B10|Peru:Cusco|2016

Anteras cruentatus|19029C02|Panama|1983

Anteras kupris|19029C04|Ecuador|2008

Anteros bracteata|19029C06|Ecuador|2008

Helicopis gnidus|19028HO2|Venezuelal 19384

that Anteros may be paraphyletic. Using this

phylogenetic result and phenotypic similarities (cream = Sarota completal iS029805|Peru:Cuscol 2009

vs. dark background of wings below) for species with | Fig. 22. Helicopini genera with Ourocnemis (red and

missing DNA data, we transfer four species from ina cate ane A meron OMe):

Anteros to Ourocnemis to form the following new combinations: Ourocnemis carausius (Westwood,

1851), Ourocnemis principalis (Hopfter, 1874), Ourocnemis renaldus (Stoll, 1790), and Ourocnemis

aerosus (Stichel, 1924).

Helicopis cupido|19028HO3|Suriname|1982

Imelda Hewitson, 1870 is a subgenus of Echenais Hiibner, [1819]

Despite the differences in appearance, monotypic genus

Echenais Hiibner, [1819] (type species Lemonias alphaea

Hiibner, [1808], which is Papilio thelephus Cramer, 1775)

is genetically close to Imelda Hewitson, 1870 (type species

Imelda glaucosmia Hewitson, 1870, which is a subspecies .

of Nymphidium mycea Hewitson, 1865) (Fig. 23). E.g., COI Bigs23: ECHnaS CRIME) SUDBENUS UNEGLGE MCE),

barcodes of their type species differ by 8.4% (55 bp). Therefore we propose treating /me/da Hewitson,

1870 as a subgenus of Echenais Hiibner, [1819].

Echenais thelephus|19029H04|Peru|1993

Imelda mycea glaucosmia|19029HO1|Ecuador|2003

Imelda aenetus|19029HO2|Peru:Cusco|2013

Calydna calamisal19029004|French Guianal1992

Calydna sturnula|19029E05|Costa Rica|2011

Calydna thersander|19029D12|Brazil:RO|1993

Calydna caieta|19029D05|French Guianal198s8

Echydna chasebal|19029002|Brazil:sc|1991

Echydna punctatal19029003|Peru/2016

Colaciticus Stichel, 1910 is junior subjective synonym of Baeotis Hiibner, [1819]

Despite different coloration and wing shape, which are Baeotis capreolus|18076D10|Brazil:AM|old

os il . = Baeotis macularia|19027B802|Panamali977

probably due to mimicry, Colaciticus Stichel, 1910 284 paeotis kadenii| 19027B03|Venezuela| 1985

3 . . ,Colaciticus johnstani|19083HO5|old

(type species Monethe johnstoni Dannatt, 1904) “Colaciticus johnstoni|19083H06|old

tS Se < mu Colaciticus johnstoni|19129E05|Ecuador|2010

originates deep within Baeotis Hiibner, [1819] (type Baeotis euprepes|19027A06|Brazil:SP|1991

i . ‘ : ‘ Baeotis primal 19027408|Peru:Amazonas|1999

species Baeotis hisbaena Hubner, [1819] which is Baeotis nesaea|19027A04|Costa Rica] 1980

ui . : ae: Baeotis felix|19027409|Peru:Cusco|2016

Papilio hisbon Cramer, 1775) (Fig. 24). Baeotis is a ? Baeotis zonata| 10595|Mexico:SLP| 1980

ll-defined meat that docivot aeisturt ‘ ohare der tencietane aera el meteal

ae -Gerine prominen genus at we do not wis 0 om Baeotis alegantula|19026H12|Peru:Cuscol2012

Baeotis choroniensis|19027412|Venezuela|1985

split. Therefore, to restore the monophyly, WS Proreek Dachetola azora|18049401|Brazil:RJ|1998

that Colaciticus Stichel, 1910 is a junior subjective Deel etolavealigatal THES aAn2|Penama|la%3

Fig. 24. Baeotis (blue and red).

synonym of Baeotis Hubner, [1819].

Nahida coenoides (Hewitson, 1870) is conspecific with [thomets aurantiaca H. Bates,

1862 and Nahida Kirby, 1871 is a junior subjective synonym of /thomeis Bates, 1862

Monotypic genus Nahida Kirby, 1871 (type and the only species Threnodes coenoides Hewitson, 1870)

originates within Ithomeis aurantiaca H. ase Nahida coenoides coenoides (= eee lee eae

: P ; Nahida coenoides pipette eat aol eae Macas|old

Bates. 1862 (type locality Brazil: Amazonas) Ithomeis aurantiaca delectal18076F09|T|Brazil:SClo

4 ‘ . : ‘ 7 area Ste ace sa ace ee ‘Pebas|ald

amels aurantiaca mimica eruLoreta

which is the type species of /thomeis Bates, Romets aurartaca cabaites|is02eAt2|Gurenal2000

; : . ; ‘ ‘ ameis aurantiaca heliconina (=cumbasina Qo

1862 (Fig. 25) implying that Nahida IS a Junior ) Tahomeis aurantiaca Te iSaTieh lt seesgan Peru:Loreto|1995

. : : Ithomeis aurantiaca lauronial19026A10/Peru:Cusco|2016

subjective synonym of Ithomeis. Moreover, Ithomeis eulema serenal|18076F06|Colombiail1926

: : 3 1 ithomeis eulema serena|19026406|Ecuador|2011

Ithomeis coenoides (Hewitson, 1870) comb. Ithomeis eulema imitatrix|19026405|Panamal1982

nov. and various subspecies of I. aurantiaca Fig. 25. Nahida coenoides (red) is Ithomeis aurantiaca (blue).

are very close to each other genetically, e.g., their COI barcodes are mostly 0.15% (1 bp) different, at

maximum 0.9% (6 bp) with /thomeis aurantiaca satellites H. Bates, 1862 (type locality Brazil: Para).

Therefore, we consider /. coenoides to be conspecific with 1. aurantiaca, and propose the following new

Species-subspecies combinations: /thomeis aurantiaca coenoides (Hewitson, 1870), [thomeis aurantiaca

ecuadorica (Strand, 1911), /thomeis aurantiaca trochois (Hewitson, 1877).

Putridivora Grishin, new genus Parcella amarynthinal18049403|Ecuador|1999

- Pheles aca tale alana era

http://zoobank. org/AD6A 1874-503C-4A07-B079-79A 326060635 Lt casa neodes|laoszeoelerasiialiay7

* Lasaia cutisca|19027CO7|Ecuador|1999

Melanis pixe|5148|USA4:TX,Hidalgo Co,[2015

. 3 Lasala agesilas|19027B10|French Guianal1991

Type species. Charis argyrea Bates, 1868. , Lasaia meris|20036409|F|French Guiana|2014

* Lasaia sula|4026|USA4:TX,Cameron Co,.|2015

Detritivora argyrea|18048H04|Peru|2013

Definition. Currently within Detritivora Hall & Harvey, Detritivora smallil 18049A06|PT|Panamal 1975

; ; ; Detritivora nicolay!]18049405|PT|Panamal|1977

2002 (type species Charis matic Harvey & Hall, 2002), but Detivora maticii902sA10[HT|clombial 1968

is not monophyletic with it, and instead is sister to the clade phen dees eenuesimn

. te . Amphiselenis chama|19027604|Venezuelal1988

that includes Detritivora with a number of other genera such Calephelis mexicana|15109H03|Mexico:Ver|1961

: i ; : ee Calephelis virginiensis|4171|TX,Hardin Co./2015

as Charis Hubner, [1819] (type species Charis ania Hiibner, Fig. 26. Putridivora gen. n. (green), Detritivora

[1819], which is Papilio anius Cramer, 1776) and | (brown), Chadia (pink), Charis (olive), Inkana

Calephelis Grote & Robinson, 1869 (type species Erycina gen. n. (red), and Lasaia (blue).

virginiensis Guérin-Méneville, [1832]) (Fig. 26), hence a new genus. Similar to Detritivora as described

by Hall and Harvey (2002), and distinguished from it by 3 (not 4) dark marks in discal cell and signa

markedly elongate along corpus bursae wall, as described by Hall and Harvey (2001), see their Fig. 6a.

Etymology. The name is a feminine noun in the nominative singular formed by replacing prefix "Detriti-"

with "Putridi-" in the name of the former genus for these species, but keeping the meaning.

Species included. The type species and Charis smalli Hall & Harvey, 2001.

Parent taxon. Tribe Riodinini Grote, 1895.

Chadia Grishin, new genus

http://zoobank. org/10295164-45F5-4C4E-8323-FBCOD37E802D

Type species. Charis cadytis Hewitson, 1866.

Definition. Currently one of the two species of Charis Hiibner, [1819] (type species Charis ania Hiibner,

[1819], which is Papilio anius Cramer, 1776), but is not monophyletic with the second (and the type)

Species, and instead sister to the clade that includes a number of other genera such as Charis and

Calephelis Grote & Robinson, 1869 (type species Erycina virginiensis Guérin-Méneville, [1832]) (Fig.

26). Therefore, it constitutes a new genus. Distinguished from its relatives by atypical for the group more

elongated wings (without pointed apex), orange band at forewing apex, prominent silver wing margins in

males below; exceptionally curved, earlobe-shaped aedeagus and very broad valvae, as broad as long.

Etymology. The name is a feminine noun in the nominative singular formed from the beginning of the

type species name with insertion of h for Charis.

Species included. Only the type species.

Parent taxon. Tribe Riodinini Grote, 1895.

Inkana Grishin, new genus

http://zoobank.org/4FEC8FF0-3B35-4B6A-BEC4-5CF06240CA5C

Type species. Charis incoides Schaus, 1902.

Definition. Currently in the genus Lasaia Bates, 1868 (type species Papilio meris Stoll, 1781) but is not

monophyletic with it and is far removed from it in the tree, being sister to the clade with several genera

such as Pheles Herrich-Schaffer, [1858] (type species Pheles heliconides Herrich-Schaffer, [1853]) and

Parcella Stichel, 1910 (type and the only species Amblygonia amarynthina C. & R. Felder, 1865) (Fig.

26) and therefore is a genus of its own. Similar to Lasaia in wing shape and color and male genitalia.

Description and diagnostic characters for this new genus are as given for Lasaia cutisca Hall & Willmott,

1998 and Lasaia incoides on pages 23—24 and illustrated in Figs. 3-4, 9-10 by Hall & Willmott (1998). In

brief, brown above and lacks blue or green scaling of Lasaia males, wings below uniform in background

without paler patches and bands of most Lasaia species, hindwing outer margin lacks prominent

concavity of Lasaia at vein M2 (manifested also as a "tooth" at M1); eyes setose; genitalic valvae narrow,

about 3 times longer than broad, processus superior with rounded projection from ventral margin,

processus inferior \ong and narrow, with a pointed tip.

Etymology. The name is a feminine noun in the nominative singular formed from the type species name

with k for c to avoid a homonym.

Species included. The type species and Lasaia cutisca Hall & Willmott, 1998.

Parent taxon. Tribe Riodinini Grote, 1895.

Callistiumini Grishin, new tribe

http://zoobank.org/3 BFAAAD2-83E4-48EF-9952-CA85DA8DEB9E

Type genus. Callistium Stichel, 1911.

Definition. Presently, Callistium (type species Charis cleadas Hewitson, 1866) is kept in incertae sedis

(Seraphim et al. 2018). Genomic analysis reveals that the type species of Callistium may be sister to

Calydnini Seraphim, Freitas & Kaminski, 2018, albeit with very weak support (Fig. 27), while other

Species currently placed in Callistium belong to other genera (see below). Therefore, the Callistium

lineage is of ancient origin, not confidently associated with any Riodinidae tribes and thus is a distinct

tribe. Description and diagnostic characters of this new tribe are as those given for Callistium on page 258

and illustrated in Fig. 68 by Stichel (1911). Abisara bifasciata| 1807112 |Myanmar|2001

- Hamearis lucina]?282|A4ustria|1981

Briefly, thorax bulky, head large, eyes setose, Euselasia chrysippe|7279|Costa Rica|2011

2 : ; Eurybia lyeiseal 18072009 [Costa Rical2013

antennae longer than *% of forewing, venation elle ely ela eee CSE Lear ae

afher ° Teratophthalma axilla] 18121G04|Peru|2015

similar to Anteros, uncus. with tegumen large, Mesosemia grandis|18072E09|Costa Rica|2012

: ‘ : Napaea eucharilal18072F01|Costa Rica|2013

about the same length as vinculum height in lateral = ° Symmgchia inal 180r00|costaFica|2012

view, hood-shaped, well separated from each a0 r _Befrostia e éaia|{s052r02|sTleraci fala =

7 ook efrostia lalannel French Guianal2

other, uncus hooked at the tip, falces curved and a aL AS fea ecg ere ee

short, not reaching middle of uncus, vinculum bent i. Sertania guttatal 4s0s3Hos|stiargentinalald

d d J h iddl l : l if ag Hala Se eee Aenea

cauda Ay the middle, valvae twice as long as = : Melanis pice[>148|USA:TX,Hidalgo Co.|2015

broad, slightly longer than tegumen, rounded, Cllephelis perditais|sase Decameron co [2015

fused together, with small lobes on dorsal and =*Zabueiatenalus|iS0saE33Iar ntnafold -

; ymphidium onaeum i Zi 2|Costa ale 2

ventral margins, aedeagus long, slightly curved "had Hiiiabe Iyconaelisoy200o [Costs Riesleooe

. ; E Theope publius|18072G01|Costa Rica|2013

and terminally pointed. = Pachythone gigas|18072F03|Costa Rica|2014

: Stalachtis halloweeni| 15028802 |Venezuela|2011

Genera included. Only the type genus Fig. 27. Callistiumini trib. n. (red) among Riodinidae tribes

and subtribes, including Riodinini (blue), Nymphidiini (cyan)

Parent Taxon. Subfamily Riodininae Grote, 1895. and Eunogyrina subtrib. n. (magenta).

Comments. Callistus Bonelli, 1810 (Coleoptera: Carabidae) is the type genus of Callistini. According to

Art. 55.4. of the ICZN Code ICZN 1999), one letter difference, which in this case would be an extra "1"

from the suffix "-ini", avoids homonymy. However, it does not avoid confusion, including internet search

engines that correct possible spelling errors. Therefore, the choice was to form the name as Callistiumini

using the entire word as a stem (Art. 29.1.), instead of Callistiini.

7 7 Fabuella tenellus|19044G605|Argentina|1992

Zabuella castanea (Prittwitz, 1865), cSfistium castanea) 180436 1218 Brazil:SPold

, 2 eta Sale eg Paes ae ae Seer p

= e aculosa|LS048E02|Frenc anal1993

confirmed combination , =3istiym race ose 280 89502 French Guranal

Behemothia godmanii|18048E10|Mexico: Qaxaca|1990

: ee : . = i ea eogeme Decale ye ee

o. alicosama robbinsi 2 ama

Placed in Callistium (type species Charis cleadas arshaeanympha unc [19032601 |Panamal 1977

: 7 Theope nycteis|19032B01|Guyanal2000

Hewitson, 1866) by Hall (2018), Calydna ~ Theope orphana| 19032807 Breen Mal 1994

9032C04|Costa Rica|2011

‘ : ; ; ; wo, Theope devriesi

castanea Prittwitz, 1865 is not monophyletic with r ane excels

eecacae French Buena ane

Theope je comosa| taaaecue Peru|1993

9032E09]Peru:Loreto|1

its type species, but instead is sister to the type Theape foliorum|19032F 11 Perul1993

: p ‘i ; 7yPseudotinea hemis|18072HO1/HT|Brazil:SC|old

species of Zabuella Stichel, 1911, which is pee eat eI ered cored Ea

Lemonias tenellus Burmeister, 1878 (Fig. 28), 1 Petocarus aitens190ssHOeiBrzilold

and is genetically close to it. Therefore, we agree ae 28. aataae genera: me Sue Heer

. . magenta), and Teenie gen. n. (orange); an eopina

with Seraphim et al. (2018) who reached the genera: Behemothia (magenta with subgenus Calicosama

same conclusion, and confirm the combination in cyan), Archaeonympha (black), Theope (blue),

Zabuella castanea (Prittwitz, 1865). Pseudotinea (black), and Petrocerus (red).

Hallonympha Penz & DeVries, 2006, valid genus, and

Hallonympha maculosa (Bates, 1868), new combination

Although synonymized with Zabuella Stichel, 1911 (type species Lemonias tenellus Burmeister, 1878) by

Hall (2018), Hallonympha Penz & DeVries, 2006 (type species Apodemia paucipuncta Spitz, 1930) is

genetically distant from it (Fig. 28), in agreement with Seraphim et al. (2018). E.g., COI barcodes of the

type species of these genera differ by 9.1% (60 bp) and their divergence dates to about 20 Mya (Seraphim

et al. 2018). Therefore, we reinstate Hallonympha as a valid genus. Currently in Callistium (type species

Charis cleadas Hewitson, 1866), Calydna maculosa Bates, 1868 (type locality Brazil: Amazonas) is not

monophyletic with its type species and instead is sister to the type species of Hallonympha (Fig. 28).

Their COI barcodes are only 7.4% (49 bp) different. Hence, we propose Hallonympha maculosa (Bates,

1868) comb. nov.

Teenie Grishin, new genus

http://zoobank.org/BDF61309-B594-4355-B2ED-306025A2E8D9

Type species. Calydna tinea Bates, 1868.

Definition. Currently in the genus Zabuella Stichel, 1911 (type species Lemonias tenellus Burmeister,

1878) as suggested by Hall (2018), this group is not monophyletic with it and is sister to Hallonympha

Penz & DeVries, 2006 (type species Apodemia paucipuncta Spitz, 1930), but prominently differs from it

genetically and thus constitutes a genus (Fig. 28). Diagnostic characters for the new genus are as those

detailed for Z. tinea and Z. argiella in Hall (2018). Briefly, similar to Zabuella and Hallonympha in wing

patterns and genitalia, but aedeagus narrower, and signa highly asymmetrical both in position and size.

Etymology. The name is a feminine noun in the nominative singular inspired by the type species name.

Species included. The type species (including Lemonias eudocia Godman & Salvin, 1897 as it synonym)

and Calydna argiella Bates, 1868.

Parent taxon. Subtribe Zabuellina Seraphim, Freitas & Kaminski, 2018.

Petrocerus Callaghan, 1979 belongs to Theopina Clench, 1955

Currently in incertae sedis, Petrocerus Callaghan, 1979 (type species Calydna catiena Hewitson, 1875) is

a confident sister to Pseudotinea Hall & Callaghan, 2003 (type species Calydna volcanicus Callaghan &

Salazar, 1997) (Fig. 28). Together they are sister to the clade with Theope Doubleday, 1847 (type species

Polyommatus terambus Godart, [1824]), and therefore, Petrocerus belongs to Theopina Clench, 1955.

Calicosama J. Hall & Harvey, 2001 is a subgenus of Behemothia Hall, 2000

Monotypic genus Behemothia Hall, 2000 (type and the only species Pandemos godmanii Dewitz, 1877) is

closely related to nearly monotypic Calicosama J. Hall & Harvey, 2001 (type species Nymphidium lilina

Butler, 1870) (Fig. 28). Both genera are mostly Central American. To highlight their close kinship and

reduce the number of unnecessarily monotypic genera we propose that Calicosama J. Hall & Harvey,

2001 is a subgenus of Behemothia Hall, 2000.

Exoplisia aphanis (Stichel, 1910), new combination

Chamaelimnas tircis|18048G02|Brazil:RO|1991

Placed in Seco Hall & Harvey, 2002 (type species Charis Barbicornis basilis|19026E11|Brazil:RGS|1956

: F . ‘ seco calagutis|19044G07|Ecuador|1914

calagutis Hewitson, 1871), Charmona aphanis Stichel, 1910 Caria plutargus|19026G11|Argentina|2012

7 ic with j ee cchabiane Bee a oe

eco ace oO eo

is not monophyletic with its type species, but is sister to oer es ane eee orbs

Exoplisia Godman & Salvin, 1886 (type species Amarynthis : Seco aphansliads4bivlstine datalad

. . 9

hypochalybe C. & R. Felder, 1861) and is not prominently fan Wee ea cadmeisl807/2A0¢ Ecuador 1998

ene . . “, ° E rmis|18072408/B 2003

distinct from it (Fig. 29). Therefore, we place it in this genus pe ern is ase

a : Fig. 29. Seco (purple), Oco gen. n. (magenta),

to form Exoplisia aphanis (Stichel, 1910) comb. nov. Exoplisia (cyan and orange).

Oco Grishin, new genus

http://zoobank. org/1D2F2454-EFC9-473A-A91D-17097BB3DB03

Type species. Symmachia ocellata Hewitson, 1867.

Definition. Currently in the genus Seco Hall & Harvey, 2002 (type species Charis calagutis Hewitson,

1871), but is not monophyletic with its type species, and instead forms an independent lineage in early

radiation of the core Riodinini group (Fig. 29). Diagnostic characters for this new genus are as those given

for Seco (except the characters specific to either Seco calagutis (Hewitson, 1871) or Exoplisia aphanis

(Stichel, 1910) comb. nov. [then placed in Seco]) on pages 415-417 in Hall and Harvey (2002). Readily

distinguished from its relatives by a black yellow-bordered eyespot at forewing apex.

Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type

Species name with the name of the genus it was formerly placed in: O[cellata] + [Se]co for its eyespots.

Species included. Only the type species.

Parent taxon. Tribe Riodinini Grote, 1895.

Machaya Hall & Willmott, 1995 is

a junior subjective synonym of Pachythone Bates, 1868

1 1 1 Pachyth qi 18072F03|Costa Rica|2014

Despite the difference in appearance, Machaya Hall & rary tian fart] 19029409) French Culana|1988

: : F ar \ i] a

Willmott, 1995 (type species Machaya obstinata Hall & Machel abstinatali2i2eboalecuador|2010

i Fe 2 ° = t 19029H12|P 1991

Willmott, 1995) originates deep within Pachythone Bates, ®pachythone lateritial20036A06|French Guianal2016

‘ : Pachythone distigma|19031401|Peru|2014

1868 (type species Pachythone erebia Bates, 1868) at the Pachythone erebial19129C01|Ecuador|2015

1 f id di ificati f it Fi 30 SEE PR Toi alae gues an Gotan Wises eile laa

time of rapid diversification of its crown group (Fig. 30). a RR ere eee eee a

Therefore, to restore the monophyly, we propose that Pec One Soave [Aa0a1ADE Manus SUA Gee

Machaya is a junior subjective synonym of Pachythone. Fig. 30. Pachythone (blue and red).

Family Nymphalidae Rafinesque, 1815

Polygrapha Staudinger, 1887 and Fountainea Rydon, 1971

are subgenera of Anaea Hiibner, [1819]

Our genomic phylogeny of Anaeini Reuter, 1896 is fully consistent with that of Toussaint et al. (2019) in

identifying major clades in the tribe and re- Hypna clytemnestra|19095C08|Guyanal2000

: ‘ A faretis isidoral19095C04|Guyanal1999

aligning species of the former Polygrapha . Siderone nemesis|19095C06|Cubalold

i . . F * Siderone galanthis|19095C07|Brazil:Mato Grosso|1969

Staudinger, 1887 (type species Paphia cyanea Phantos callidryas|19095C05|Costa Rica|2010

. . Consul ena al aoes eaten Maret |

Anae 3982|USA:TX,M Co.|201

Salvin & Godman, 1868) (Fig. 31). We note - -fiiaea trosiodytal 17 Lior {0(UsarL Monroe Co,|1967

that relative branch lengths also agree between Lease Gyenum|t7 TBA hence: ‘Tam|1994

the two phylogenies. Assigning all species in See ERE Re eeecanieecaigeak

7 . Polygrapha cyaneal19095C10|Peru|/1981

one of the sister clades to the single genus Polygrapha suprema|19095D01|no datalold

Memphis Hiibner, [1819] (t ae ee MeMlemphie echamus|17119F12/Cuba|1962

emp LS U ner, f ; Oe Pee CS aie LO : Hes Ae aaciat eaeered ee ene or

oe Ls Me es|19095D03|Peru|2016

odilia Stoll, 1780, which IS a Jumor subjective i Sanania Mmoruus|1909SD04]Brazil:Rondonial1991

synonym of Papilio polycarmes Fabricius, F vamphis faleatal 12095E0Iecuaderiold

Rig ; j . M 19095D06|Costa Rical2003

1775), but dividing its sister clade of equal | Memphis wenocrates|19095C12|Perul 197 1

diversification into 3 genera: Polygrapha, | Fig. 31. Anaeini genera: Anaea (red, cyan and olive), Memphis

Fountainea Rydon, 1971 (type species Anaea (blue), Zaretis (green, purple and orange), Consul, and Hypna.

phidile Geyer, 1837, a subspecies of Papilio ryphea Cramer, 1775) and Anaea Hiibner, [1819] (type

species Papilio troglodyta Fabricius, 1775) is internally inconsistent. Not willing to break Memphis into

several genera, we propose treating Polygrapha and Fountainea as subgenera of Anaea. This generic

realignment results in two large and equally diversified sister genera: Anaea and Memphis, with Consul

Hiibner, [1807] (type species Papilio fabius Cramer, 1776) being their sister.

Siderone Hibner, [1823] and Phantos Dias, 2018

are subgenera of Zaretis Hiibner, [1819]

Genetic differentiation within the clade of three genera: Siderone Hibner, [1823] (type species Siderone

ide Hiibner, [1823], a junior subjective synonym of Papilio nemesis Illiger, 1801), Phantos Dias, 2018

(type species Nymphalis callidryas R. Felder, 1869) and Zaretis Hiibner, [1819] (type species Papilio

isidora Cramer, 1779) is less than that of Memphis and Anaea (sensu lato, as above) (Fig. 31), suggesting

that Siderone and Phantos are subgenera of Zaretis to restore internal consistency of the classification.

According to Fig. 3 in Toussaint et al. (2019), all non-monotypic genera of Anaeini as we define them

(Anaea, Memphis, Consul and Zaretis) diversified around the same time 17-20 Mya, indicating internal

consistency of our definition; and monotypic genera Coenophlebia C. & R. Felder, 1862 (type and the

only species Siderone archidona Hewitson, 1860, we have not sequenced this species yet) and Hypna

Hiibner, [1819] (type species Papilio clytemnestra Cramer, 1777) are more distant from others to maintain

their distinction.

Maniolina Grote, 1897 and Melanargiina Wheeler, 1903 are junior subjective

synonyms of Erebiina Tutt, 1896 and Satyrina Boisduval, 1833, respectively

The nuclear genomic tree of Satyrini Boisduval, 1833 reveals a confidently supported clade that contains

a number of subtribes as they are currently defined (Pefia et al. 2006; Wahlberg 2019a), including the

nominotypical subtribe Satyrina. We call this clade the Satyrini crown group (Fig. 32). The tree

confidently groups the monotypic subtribes Erebiina Tutt, 1896 and Melanargiina Wheeler, 1903

(Wahlberg 2019a) with Maniolina Grote, 1897 and Satyrina Boisduval, 1833, respectively (Fig. 32).

Subtribes in each pair are more closely related to each other compared to other pairs of subtribes in

Satyrini Boisduval, 1833, e.g., Euptychiina Reuter, 1896 and Pronophilina Reuter, 1896. Therefore, to

simplify subtribal classification and

avoid indistinct monotypic subtribes, we

propose that Maniolina Grote, 1897 is a

junior subjective synonym of Erebiina

Tutt, 1896 and Melanargiina Wheeler,

1903 is a junior subjective synonym of

Satyrina Boisduval, 1833. The Satyrini

crown group first splits into two most

distinct sister clades (Fig. 32). One is the

subtribe Satyrina. The other contains all

other subtribes and has undergone rapid

radiation that largely obscures the order

of bifurcations near its origin thus

creating obstacles for its classification. It

is possible to consider this entire group

as a single subtribe, because it is the

Euptychia mollina|19117F06|Guyanal|2000 Subtribes

Megisto cymela|3689|USA;TX,Harrison Co.J2015 Euptychiina

Cyllopsis gemma|9181|/USA:4R,Seott Co,|2017

Etcheverrius chillensis|19115610|4rgentina|1992

Steremnia umbracina|19117D010|Peru|2012 Pronophilina

Pherepedaliodes pheretiades|19117C12|Peru|2013

Gyrocheilus patrobas|10223/USA;AZ,Santa Cruz Co,|2017 G¥rochellina

Strabena smithii]19115C11|Madagascar|1988 _

Ypthima huebneri|19115E04|Myanmar|2002

Melampias huebneri]19115C12|South Africa] 1949

Neocoenyra duplex|19115006|Kenyal1960

Pseudonympha poetula|19115D12|South Africal1959

Cassionympha cassius|19115D04|South Africa] 1945

Paternympha naryecia]/19115011|South Africa? |1955

: an : Ypthimina

Stygionympha vigilans|19115E01/South Africa] 1945 includes

Paralasa herse|19115D07|Chinalold genus Paralasa

va nee jordanali9115D08|"Tura"|old

aralasa hades|19115D10|Russia:Alay|1971

Paralasa maracandica|19115009|Uzbekistan?|oald

Proterebla afra]19114D03|Russia:Altailold

Argestina waltoni]19115C09|Tibet|1980

Callerebia scandal19115D03|Indialald i

Loxerebla pratorum|19115D05|Chinal old en

Physcaeneura pandal19115D001|S Africa] 1968

Physcaeneura leda|19115D02|Kenyal1950

Erebia epipsodeal9331|USA:wY,Park Co.]2017 Erebiina

Maniola jurtina|20035B06|France|2018 =Maniolina

Cercyonis pegalal4506|USA4:TX, Wise Co,|2015

Melanargia galathea|19114B11|Albania|1991

Hipparchia statilinus|20035B05|France|2018 Satyrina

Satyrus actaeal19114407|Spain|1975 =Melanargiina

Oenels polixenes|16106G612|Canada:yT|2016

most prominent and well-supported large Fig. 32. Subtribes in the Satyrini crown group, each in its own color,

clade in the tree. However, we refrain except Ypthimina is blue + cyan. Subtribe names are listed on the right.

from this unification because of the number of species involved and the practicality of partitioning them

into phylogenetically meaningful smaller groups of ICZN-administered rank (ICZN 1999). Therefore we

keep the current classification of the group essentially as proposed by Pefia et al. (2006) with the two

exceptions mentioned above. However, as a consequence, several lineages borne out of that rapid

radiation, and therefore consistent with the Pefia et al. (2006) definition, do not have available names and

need to be named as subtribes, which is accomplished below. The following acts also eliminate "Subtribe

uncertain" group of available genus-group names as listed by Wahlberg (2019a).

Paralasa Moore, 1893 belongs to Ypthimina Reuter, 1896

Listed among the "Subtribe uncertain" genera by Wahlberg (2019a), Paralasa Moore, 1893 (type species

Erebia kalinda Moore, 1865) is a confident sister to Ypthimina Reuter, 1896 (Fig. 32). Therefore, instead

of proposing a monotypic subtribe for this genus, we include Paralasa in the subtribe Ypthimina.

Callerebiina Grishin, new subtribe

http://zoobank.org/46A68C99-BBC2-4F6C-BB06-4CA2AD2343B8

Type genus. Callerebia Butler, 1867.

Definition. Several genera of uncertain tribal placement (Wahlberg 2019a) confidently grouped with

others sometimes placed in Ypthimina Reuter, 1896 (Fig. 32 red). Close relationship of Callerebia,

Loxerebia and Argestina that are in the red clade has been reported before (Yang and Zhang 2015). This

prominent red clade is indeed a weakly supported sister to Ypthimina (Fig. 32 blue and cyan). However,

due to the weak statistical support (therefore possibly erroneous sister relationship with Ypthimina) and

the origin of both clades near the rapid radiation of many Satyrini subtribes, this clade is proposed as a

new subtribe. A phenotypically diverse assembly of species, generally characterized by Erebia-like

appearance, typically with a row of orange black-centered eyespots on each wing, but these could be

reduced to a couple of eyespots, similar to Ypthimina, or wings could be unspotted brown or even mostly

white above (with brown borders) in some species; gnathos developed (absent in many Ypthimina), but

arms shorter than uncus, saccus reduced, valva rather stout, flattened and rounded or excavated near the

apex, aedeagus rather short, shorter than valva, typically bent and twisted, boomerang-shaped

differentiating the subtribe from Erebia Dalman, 1816. Due to phenotypic diversity, best diagnosed by the

DNA characters in the nuclear genome: hm2009277-RA.13:T2076C, hm2009277-RA.13:A130C,

hm2015715-RA.2:C2137T, hm2009379-RA.4:T1I915C, ahm2002906-RA.2:A128G, where the part before

the first dot (e.g. hm2009277-RA) is the protein ID, next number (e.g. 13) is exon of the Heliconius

melpomene genome assembly (Davey et al. 2016), and combination like A130C means base pair C at

position 130, changed from A in the ancestor. See <https://osf.i0/kj4es/> for the sequences of these exons.

Genera included. The type genus, Proterebia Roos & Arnscheid, 1980, Argestina Riley, 1923, Loxerebia

Watkins, 1925, and Physcaeneura Wallengren, 1857.

Parent Taxon. Tribe Satyrini Boisduval, 1833.

Gyrocheilina Grishin, new subtribe

http://zoobank.org/EF7C543 A-88B9-4DCC-9847-139CC7763B84

Type genus. Gyrocheilus Butler, 1867.

Definition. Previously placed in Pronophilina Reuter, 1896 (Wahlberg 2019a), but is not monophyletic

with it. Instead, it originates in early radiation of the clade that is sister to Satyrina Boisduval, 1833 (Figs.

32, 33 orange), likely prior to divergence of subtribes Ypthimina Reuter, 1896 and Erebiina Tutt, 1896,

and therefore is a subtribe. Diagnosed by the combination of the following characters: most prominently,

female foreleg 1s much reduced, with 2 tarsal subsegment, not spined; then forewings apically rounded

with somewhat undulate outer margin, hindwing with strongly undulate, almost toothed outer margins;

forewing discal cell short and broad: slightly longer than half of wing, width ‘4—'4 of its length; forewing

vein Ri starts at distal '4 of discal cell, R2 very near discal cell apex (only slightly stalked), R3 and R4

stalked for 4 of their length, R4 and Rs stalked for about half of their length, recurrent vein in discal cell

from the middle of discocellular vein for '4 of discal cell length, discocellular vein straight between R2

and Mi and between M2 and M3 origins, but V-shaped between Mi and M2 origins; forewing with 4

(rarely 3) white-centered round eyespots, hindwing without such eyespots but frequently with several

cream-colored ovals or crescents; palpi long and porrect, scales on the second segment long, up to 5 times

the width of the segment, eyes bare, antennae shorter than half of forewing; pupa suspended, spindle-

shaped, smooth, head capsule apically extended similar to cremaster, forked at the tip.

Genera included. Only the type genus.

Parent Taxon. Tribe Satyrini Boisduval, 1833.

Calistina Grishin, new subtribe

http://zoobank.org/F 1 DIA7TDC-F549-4EC9-8C68-D8032DC4FB95

Type genus. Calisto Hiibner, [1823].

Definition. The phylogenetic position of Calisto Htibner, [1823] (type species Papilio zangis Fabricius,

1775) remains uncertain (Pefia et al.

Erites medura|19099D02|Javalold Subtribes

2006; Wahlberg 2019a). Previously it radi

: ope a = Lethi

was placed in Pronophilina Reuter, Pararge Beyerials00oAgSIrrance|20i8 | angie

1896 (Miller 1968), but is not = Heteropsis drepana|19114H03 biMadagascar| 1988 Mycalesina

Seen ee ‘ Seer a ee BaP mcm

monophyletic with it (Fig. 33). Appa- Casto zangis|io245pamarajzoi7

. . bd s oreu nla arnaca OSTa Rica

rently, it is an ancient phylogenetic | , | relies Splendeuptychia ashna 1o11BA09|Perulz015

. . oe e e

lineage (Matos-Maravi et al. 2014) a. Hermeuipt this sosybius| 3310/1 Sabine Co|2015

a enone ee ae ane see aera | Comer tiem

ating to the rap fadiauion O € Inbio hilara]19118B02|Costa Rica|2012

. . . = Cyll 9181/USA:AR,Scott Co.|2017

clade sister to Satyrina Boisduval, a Caenopty chia eae 1885

1833 (Fig 33) restricted to Caribbean Etcheverrius chiliensis| 19115 iE iojArgentina|1902

revue peed ele ae a isi2largentina| 199: 93

Islands and not closely associated with steremnia umbracinal (0 Li7bid)peruiz0oi2 meine

et ge . rT LUun@a doraete eru

P hila thelebe/19117D04|Ve | 8

anya SrOUp OLE Satylitls WHGTENOLe 13 2 oe oa

a a e

subtibe. This new subtie e somn ra

tinguished from other Satyrini by Parsee hades|19115D10 Russia: ere oaeee:

tion: forewing vein R1 originates ee

vena : oO. subtrib. n.

t b d b t t b fe th d ee Martolaquttina S0035806 Seae cane perio Erebiina

at or beyond (but not before) the en a yegroyonis pegalal4206|USA.1x Wise Co|2015 ceili

. . e e

of discal cell, all other R veins stalked, eee eae galethes 30035B05|France| 2016 Satyrina

; eine : 1.08 Satyrus actaeal19114407|Spain|1975 =Melanargiina

discocellular vein irregular, veins Mi Oeneis polixenes|16106G12|Canada:yT/2016

Fig. 33. Satyrini subtribes marked with alternating shading, names listed on

the right. All Z chromosome protein-coding genes are used for this tree.

and M2 invade more into the discal

cell, bases of Sc and Cu, but not A, are

inflated; humeral vein weak, short, curved parallel to Sc. Further details about these diagnostic venation

characters for the subtribe are as given for Calisto on page 104 by Brown and Heineman (1972).

Genera included. Only the type genus.

Parent Taxon. Tribe Satyrini Boisduval, 1833.

Comments. The phylogenetic tree constructed from all protein-coding regions predicted to be on the Z

chromosome (Fig. 33) reveals some affinity of Calisto to Euptychiina Reuter, 1896. Although this

relationship seems possible, it is not obvious from morphology, and only 76% out of 100 selections of

positions from the genomic alignment support this placement (0.76 value at the node in Fig. 33). In our

experience, this value is too low for confident classification. Therefore, placing Calisto in Euptychiina

could be incorrect. Also, it is clearly incorrect to keep Calisto in Pronophilina, because as our tree shows,

support for the Pronophilina clade that includes a diverse sample of genera is very strong, at 100% (Fig.

33 green), and Calisto is placed outside of this clade. Therefore, the solution was to propose a new

substribe for the Calisto clade. Another curious observation is that the Euptychia Hitibner, 1818 (type

species Oreas mollina Hubner, [1813]) clade (1.e., Euptychia sensu lato) is quite removed from the rest of

Euptychiina, and their association is only weakly supported (0.56, Fig. 33). This sister to Euptychia sensu

lato clade is more prominent than Euptychiina as currently defined, and may deserve subtribal status: a

question that could be answered in future studies.

Harsiesis Fruhstorfer, 1911 is a subgenus of Platypthima Rothschild & Jordan, 1905

Harsiesis Fruhstorfer, 1911 (type species Hypocista [sic] hygea Hewitson, 1863) and Platypthima

Rothschild & Jordan, 1905 (type species Platypthima ornata Rothschild & Jordan, 1905) are

characterized by an elevated evolutionary rate as revealed by their longer branches (Fig. 34). Despite their

faster evolution, the two are more genetically similar to each other than other pairs of sister genera, e.g.,

COI barcodes between their type species (GenBank accession GQ357203 for P. ornata) differ by 9.4%

(62 bp). Therefore Harsiesis Fruhstorfer, 1911 is a subgenus of Platypthima Rothschild & Jordan, 1905.

Percnodaimon Butler, 1876 and Erebiola Fereday, 1879 are junior subjective

synonyms of Argyrophenga Doubleday, 1845

Two monotypic genera Erebiola Fereday, 1879 (type and the ony species Erebiola butleri Fereday,

1879) and Percnodaimon Butler, 1876 (type and the ‘ Harsiesis hygea|19099406|P.N.Guinea| 1972

Platypthima homocroal19099C02|P.N.Guineal1983

only species Percnodaimon pluto Butler, 1876, “Atgpa deol 2903900 Papua ew Gunea| 1903

homonym, valid name Erebia merula Hewitson, | | —acsuonestiisosoatl|new zesendio70

1875) cluster closely with Argyrophenga Doubleday, | |[de— ooaiuacimieniltSedontonen Seugraies

1845 (type species Argyrophenga antipodum a55 Paratisiphone lyrnessa|19099812 |New Caledonial1984

Doubleday, 1845) (Fig. 34), all being from New Fig, 34, Platypthima (magenta) and Argyrophenga (blue).

Zealand and phenotypically similar. Therefore, Percnodaimon Butler, 1876 and Erebiola Fereday, 1879

are junior subjective synonyms of Argyrophenga Doubleday, 1845.

Magda Grishin, new subgenus

http://zoobank.org/4AF74BDC-1FCD-42B6-8BFB-A3 16FOFO9D93

Type species. Erebia magdalena Strecker, 1880.

Definition. Forms a rather prominent clade within Erebia Dalman, 1816 (type species type species

Papilio ligea Linnaeus, 1758) sister to the clade with subgenera Atercoloratus Bang-Haas, 1938 (type

species Coenonympha alini Bang-Haas, 1937) and Boeberia Prout, 1901 (type species Papilio parmenio

Bober, 1809), thus is not monophyletic with the subgenus Erebia (Figs. 35, 36). A COI barcode

dendrogram constructed for the type species of all 11 available genus-group names associated with Erebia

reveals that this clade does not contain any of them, and therefore is a new taxon (Fig. 35). To maintain a

broadly defined genus Erebia, this taxon is assigned a subgenus rank. This new subgenus consists of the

magdalena and embla groups of Pefia et al. (2015). Phenotypically, a diverse assembly of species

previously placed in different species groups (Warren 1936; Pelham 2008) and strongly associated

together only from DNA sequence data be it gene markers (Pefia et al. 2015), COI barcodes (Fig. 35), or

all genomic protein-coding regions (Fig. 36). Morphologically, diagnosed either by a very short gnathos:

shorter than third of uncus in lateral

view (the magdalena group except

E. discoidalis (W. Kirby, 1837)); or

in species with longer gnathos (the

embla group and E, discoidalis) || | !— jo» _ {Shia taste maaians Meni e isco St

arms directed posterior-ventrad in a om oe

‘ ‘ pies

Erebia (Atercolorat | Atercolorat L€340573|China:Li

lateral view, rounded and inflated at a crebia(Boeberia) oarmenta(Bosbera/|12118Ci0/Russia ae

7 e Ere t T 19099C111/F e

the tips; or if upturned at the tips Brenig (Erebia} medusa = Madusia]]19099C06 [Albania

then only slightly and arms more _| efa08 Beebe tere Ole renga singe | 103ac Ul rence

rebia (Erebia) ligea [Erebia rance

: ° = Erebia (Erebia) aethiops [=Truncaefalcia]|19099C12|Austria

parallel to uncus in lateral view than ro. Erebia (Erebia) alberganus [=Gorgo]|19099C05|France

in other subgenera due to a stron 22 See er Brebla (Erebia) epistygne [~Phoreis][19099C0715

rebia (Erebia) epistygne orcis ain

kink hei 8 ies h bend 8 Maniola Erebia Merial aaniel " Simplospinosial|19029¢08] spain J

r hl rt hi 20035B06|F 2

ink at their oo: so t eye Ren to = aa aes “dercyonis (Cercyanis) pegala [Cercyonis]|4506|USA:TX,Wise Co.

position their axes parallel to the Cercyonis : Cercyonis (Hyponephele) lycaon [Hyponephele)|19113HO6|N Macedonia

Fig. 35. COI barcode dendrogram of Erebia (blue, purple, and magenta) with

its subgenera Magda subgen. n. (purple), Atercoloratus and Boeberia

(magenta). Non-type species of available genus-group names are in gray, the

distance between gnathos and lectotype of the type species of Magda is in red, genus-group names each

vinculum along tegumen_ shorter species is the type of are given in brackets (= for synonyms), Accession

uncus near their origin (instead of

being directed more ventrad),

than tegumen height in lateral view numbers are given for GenBank sequences, which are marked with an asterisk.

and valva simple, without heel-like expansion, tip rounded, valva either short, about 3 times as long as

broad with many small teeth in distal half (E. discoidalis), or long, more than 4 times longer than wide

and with a broad-W-shaped dorsal margin (£. rossii (J. Curtis, 1835)). The following combination of

characters in the COI barcode region is diagnostic: 271C (not T), 274C (not T), 421T or C (not A), 424T

(not A), 451T or C (not A), 484not T, and 622T (not A).

Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.

Species included. The type species, Hipparchia cyclopius Eversmann, 1844, Papilio disa Thunberg,

1791, Hipparchia discoidalis Kirby, 1837, Erebia edda Ménetriés, 1851, Papilio embla Thunberg, 1791,

Erebia erinnyn Warren, 1932, Erebia fasciata Butler, 1868, Erebia mackinleyensis Gunder, 1932, Erebia

mancinus Doubleday, 1849, Hipparchia rossii Curtis, 1835, Erebia magdalena sachaensis Dubatolov,

1992, Erebia tristis tristior Goltz, 1937, and Erebia wanga Bremer, 1864.

Parent taxon. Genus Erebia Dalman, 1816.

Additional genomic support for the broadly defined genera

Cercyonis Scudder, 1875, Maniola Schrank, 1801 and Erebia Dalman, 1816

Due to the pattern of diversification in the genomic tree, we proposed to place Hyponephele Muschamp,

1915 (type species Papilio lycaon Rottemburg, 1775) as a subgenus of Cercyonis Scudder, 1875 (type

species Papilio alope Fabricius, 1793, currently a subspecies of Papilio pegala Fabricius, 1775) and

Pyronia Hubner, [1819] (type species Papilio tithonus Linnaeus, 1771) with Aphantopus Wallengren,

1853 (type species Papilio hyperantus Linnaeus, 1758) as subgenera of Maniola Schrank, 1801 (type

species Maniola lemur Schrank, 1801, which is a junior subjective synonym of Papilio jurtina Linnaeus,

1758) (Zhang et al. 2020). Although the three genus-level names (Hyponephele, Pyronia, and

Aphantopus) have been in wide historical use, we have prioritized internal consistency of the

classification and selection of the most 7 Erebia 6 (ie Seema meats

; . : Mag disa|16106E

prominent clades in the genomic tree as Naas] map Mee Hele fies, Bee 2016

genera, as argued by Zhang et al. (2020) Wanage 198

: A : A ; ears ssia: SBerialold

in the Introduction and Discussion a |. cable rai its man ae quRl! i

: 5 k 6GO1[C ‘¥T/2016

sections. Internal consistency allows us rsp (Espa) young 19 etsleane NT20i6

to apply more objective and reproducible ee ee celal Ebtrss

criteria and relate ranks (genus or hus LSogSCOs|ersncel198

: 19099CO09/Spa 982

subgenus) to comparable evolutionary : ia Fance| 1380.

: 9CO4/F |/19%5

events. Here, we expand the genomic |§. 1 ae aus

tree with additional taxa (Fig. 36). The |i PK cércyons yes a se Lee

results strengthen our conclusion and Riacrentaks es cas yp ses le Salt as

agree with the recently published case an ee cae ae

h . hv] fE Cercyonis eremingp ele co eeers nis Ce remin a snpriLigi Ter Oa Rus la! al Alta 1909

comprehensive phylogeny of European (ek “Seine Cereaah Spare A, ee tee

. : ercyonis reyonis | meadil AB. os 6 ‘

butterflies (Wiemers et al. 2020). The - sonemmopitt Eerehonis ae oe USA:NY Elko Co: 208 co e017

three clades that we consider genera: 7{Pasiohana ante ; ee cals

: : Maniola [| ——— Bale

Erebia Dalman, 1816 (type species ee aa faniola) iu uiioe eo See iaetecentete

Maniola Idata) cecilia[19113HO8 Bain oan

Papilio ligea Linnaeus, 1758), Cercyonis

Fig. 36. Erebiina genera: Erebia (blue, purple, and magenta) with its

sensu lato and Maniola sensu lato that | subgenera Magda subgen. n. (purple) and Boeberia (magenta), Cercyonis

we unified into the subtribe Erebiina | (cyan) with its subgenera Hyponephele and Ereminephele, and Maniola

(red) with its subgenera Aphantopus, Pasiphana, Pyronia, and Idata.

Tutt, 1896, prominently stand out (e.,

tree branches leading to them are the longest internal branches in the tree) and diversified around the same

time (i.e. distances from the last common ancestors of these genera to the leaves are approximately the

same for all three). Moreover, more divergent taxa that have been at times treated as genera (listed as

subgenera here): Cercyonis (Ereminephele) huebneri Kocak, 1980 (formerly Hyponephele) and Erebia

(Boeberia) parmenio Bober, 1809 originate near the last common ancestors of their genera, further

supporting evolutionary significance of this time point. It does not seem accidental that 3 distinct but

related phylogenetic lineages diversified around the same time, and if we chose the names of genera to

reflect the pivotal common point in their evolution, the three-genus classification fits it best. As an

alternative, we would be left with many small lineages as genera, and for internal consistency (i.e.

taxonomic category represents a level in the classification), would need to split Erebia into at least 3

distinct genera, and Maniola into at least 5 (Fig. 36). We prefer to treat these smaller lineages as

subgenera, as they do not represent the most prominent level of diversification, but the next one to It.

Hestinalis Bryk, 1938 is a junior subjective synonym of Mimathyma Moore, 1896

In the genomic tree Hestinalis Bryk, 1938 (type species Hestina mimetica Butler, 1874) originates within

Mimathyma Moore, 1896 (type species Athyma chevana Apatura irls]19094C01|Slovalia:vihorlat/ 196

ae © (Fi : sibs lel Poss sl ena

Moore, 1866) rendering it paraphyletic (Fig. 37). To | 13 : bimathyma chavanali90 sac ale

restore monophyly and considering genetic closeness of Ta Mimathyma echrendhif9084004)Kore a}old :

A , i eng imathyma nycteis

these species, we propose that Hestinalis Bryk, 1938 isa |! * ——""Hestina taponical 19094007 Japani1934

junior subjective synonym of Mimathyma Moore, 1896. Big .3 7 ongioma sbluciand tee).

Curiously, Hestina Westwood, 1850 (type species Papilio assimilis Linnaeus, 1758) is in a different clade

and thus is not synonymous with Hestinalis.

Vila Kirby, 1871 is a subgenus of Biblis Fabricius, 1807

A monotypic genus Biblis Fabricius, 1807 (type species Papilio biblis Fabricius, 1775, a junior

homonym, valid name for this Species IS ‘Dynamine tithia|19095404|Brazil:Bahia|1991

Papilio hyperia Cramer, 1779) is a close Dynamine dyonis[as03|USA4Xcamaron Co [2015

. r . . : 7 Wvila seers rena etl tela

relative of Vila Kirby, 1871 (type species Olina * Vila eueidifarmis| 19095805 |Peru| 1962

i SIS Abana Hyberial 19094+05|BVIcGusna felana|ie4

azeca Doubleday, [1848]) (Fig. 38, compare Mestra amymone|3631/U5A4:TX,Cameron Co, pore

. . ‘ . . 5 be Archi tra teleb [19095B03/D¢ Republicl 1994

with diversification in Dynamine Hiibner, Roe rene ioney eo

[1819]). The COI barcodes of the type species

of these two genera differ by 7% (46 bp). Due to genetic similarities, we propose treating Vila as a

subgenus of Biblis. The wing pattern differences between all these species are likely caused by different

mimetic complexes they take part in.

Fig. 38. Vila (blue), Biblis (red) and Dynamine (purple).

Biblis aganisa Boisduval, 1836 is a species distinct from Biblis hyperia (Cramer, 1779)

Genetic diversification between the nominotypical Biblis (Biblis) hyperia (Cramer, 1779) (type locality

St. Thomas) and Biblis (Biblis) hyperia aganisa Boisduval, 1836 (type locality inferred as Mexico) is at

the level of that between two species from the subgenus Vila: Biblis (Vila) azeca (Doubleday, [1848]) and

Biblis (Vila) eueidiformis (Joicey & Talbot, 1918) (Fig. 38). The COI barcodes of the two taxa differ by

4.6% (30 bp). The nominotypical B. hyperia is characterized by more prominent dorsal forewing marginal

bands than B. b. aganisa. Taken together, these arguments suggest that Biblis aganisa Boisduval, 1836

reinstated status is a species distinct from Biblis hyperia (Cramer, 1779).

The following taxa are junior subjective synonyms: Catacore Dillon, 1948 of Diaethria

Billberg, 1820; Mesotaenia Kirby, 1871 with Orophila Staudinger, 1886 of Perisama

Doubleday, 1849; and Paulogramma Dillon, 1948 of Catagramma Boisduval, 1836

Genomic sequencing and analysis of the type species of available genus-group names of the subtribe

Callicorina Orfila, 1952 resulted in a fully resolved confident phylogeny (Fig. 39) that revealed close

relationships between some of them. For instance, even the most distinctive of the type species for

available genus-group names in the Diaethria Billberg, 1820 (type species Papilio clymena Cramer,

1775) clade sometimes separated in the monotypic genus Catacore Dillon, 1948 (type and the only

species Catagramma kolyma Hewitson, 1851) clusters closely with Diaethria. The COI barcodes of

Diaethria and Catacore type species are only 6.7% (44 bp) different. In agreements with Wahlberg

(2019a), we treat Catacore Dillon, 1948 as a junior subjective synonym of Diaethria Billberg, 1820.

Next, the three genera Perisama Doubleday, 1849 (type species Catagramma bomplandii Guérin-

Méneville, [1844]), Mesotaenia Kirby, 1871 (type species Callitaenia doris C. & R. Felder, 1861, a

Call tarte|19094F10|(P 2016

subspecies of Catagramma vaninka Hewitson, 1855) . Diacthvia anna[i7117612IMesico's Sree:

and Orophila Staudinger, 1886 (type species Cybelis Dissthris pandanma|19094G03| Costa Rica] 1973

Hewit 1869. a subspecies of Cvbdeli Grechie caaeae ion94603 eevaler|3001

COMI ASPE Wy) SON, 2 UDSp ay “ - Mesotaenia vanink ape Teeeeaey 2016

cardases Hewitson, 1869) cluster closely together (Fig. mma’ <r /so3 bomplandil| 19094G04|Colombial1983

. e ° il: d ig

39). Their COI barcodes also indicate a close Hasmnatera prrame| 12004612 Vanecuala moss

. : . ; 7 Antigonis p aaah apo saal eee Reon Pa

relationship, e.g., the type species of Mesotaenia and Lucite eedmme [1802 2A02 [Dominican Republic 199%

Perisama are only 6.2% (41 bp) different. Therefore, ee So Omit Bene Ta ueHia al):

. aematera (red: subgenus Antigonis, and cyan),

we agree with Wahlberg (2019a) and confirm that | Catagramma (olive & pink), and Callicore (purple &

Mesotaenia Kirby, 1871 and Orophila Staudinger, 1886 yellow: subgenus Diaethria, magenta & green:

are junior subjective synonyms of Perisama Doubleday, subgenus Perisama, and blue).

1849. Furthermore, the genomic tree confirms the close relationship between Paulogramma Dillon, 1948

(type species Nymphalis pyracmon Godart, [1824]) and Catagramma Boisduval, 1836 (type species

Catagramma hydaspes Boisduval, 1836, which is a junior subjective synonym of Nymphalis pygas

Godart, [1824]). The reasons why Freitas et al. (2014) resurrected Paulogramma in favor of the older

name Catagramma that they did not use are unclear. Their tree (Freitas et al. 2014: Fig. 3) shows

"Callicore pygas", the current name for the type species of Catagramma, inside Paulogramma as they

define it. Furthermore, Wahlberg (2019a) lists Paulogramma in synonymy with Catagramma. Therefore,

we confirm that Paulogramma Dillon, 1948 is a junior subjective synonym of Catagramma Boisduval,

1836. Finally, we confirm that Lucinia Hiibner, [1823] belongs to Callicorina (Fig. 39).

Diaethria Billberg, 1820 and Perisama Doubleday, 1849

are subgenera of Callicore Hiibner, [1819]

Inspection of genetic diversification and prominence of tree branches, we find that, contrary to their wing

pattern similarity, Catagramma Boisduval, 1836 (type species Catagramma hydaspes Boisduval, 1836,

which is a junior subjective synonym of Nymphalis pygas Godart, [1824]) is a prominent genus strongly

separated from others in the Callicore group, in agreement with Freitas et al. (2014) who called this genus

by its junior synonym Paulogramma Dillon, 1948 (type species Nymphalis pyracmon Godart, [1824])

(Fig. 39). Therefore, we retain Catagramma as a genus. However, the other three taxa: Callicore Hiibner,

[1819] (type species Papilio astarte Cramer, 1779), Diaethria Billberg, 1820 (type species Papilio

clymena Cramer, 1775) and Perisama Doubleday, 1849 (type species Catagramma bomplandii Guérin-

Méneville, [1844]) cluster more closely and are less prominently separated from each other in the tree

(Fig. 39), e.g., COI barcodes of the type species of differ by 8.8% (58 bp). Therefore, we propose that

Diaethria Billberg, 1820 and Perisama Doubleday, 1849 are subgenera of Callicore Hiibner, [1819]. This

adjustment of status (from genus to subgenus) makes their classification more internally consistent, and

this newly broader Callicore becomes similar in genetic divergence to its relatives Eunica Hubner,

[1819], Hamadryas Hiibner, [1806], and Dynamine Hiibner, [1819]. In wing patterns and shapes, the

currently proposed subgenera of Callicore are rather similar to each other, and a number of erroneous

attributions to former genera have been made in classifying these species (Freitas et al. 2014).

Antigonis C. Felder, 1861 is a subgenus of Haematera Doubleday, 1849

A monotypic genus Antigonis C. Felder, 1861 (type and the only species Cybdelis pharsalia Hewitson,

1852) is rather closely related to another monotypic genus Haematera Doubleday, 1849 (type species

Haematera thysbe Doubleday, 1849, which is a subspecies of Callidula pyrame Hiibner, [1819]) (Fig.

39), e.g., their COI barcodes differ by 9.3% (61 bp), suggesting that Antigonis C. Felder, 1861 is a

subgenus of Haematera Doubleday, 1849, thus eliminating two not truly distinctive monotypic genera.

Asterope Hiibner, [1819], Nica Hiibner, [1826], Peria Kirby, 1871,

and Callicorina Smart, 1976 are subgenera of Temenis Hiibner, [1819]

We obtained and analyzed genomic data for the type species of all 10 available genus-group names of the

subtribe Epiphilina Jenkins, 1987 that resulted in a Sm Aigienis laothioe|17 118803 |Mexica-Tamaulinas|1094

well-resolved phylogeny (Fig. 40). We find that ; gs ee PRE el eae arse Paral 1978

avilla] 19094HO

Temenis Hiibner, [1819] (type species Papilio merione Perry ams| 19084410 |Guyana|2000 :

Fabricius, 1781, which is Papilio laothoe Cramer, Epiphie adastal ty bau iMexico:Tamaulpaslives

1777) may not be monophyletic, with its type species | — eee neaerea| 9094H04/euyan|2000

being sister to Asterope Hiibner, [1819] (type species Fig. 40. Temenis (colored) with its subgenera Asterope

¥ ce ; (purple), Nica (green), Callicorina (red), Peria (cyan).

Oreas sapphira Hiibner, [1816]) with medium support,

and Temenis pulchra Hewitson, 1861, the type species of Callicorina Smart, 1976 possibly originating

among other related genera, two of which are monotypic: Nica Hibner, [1826] (type and the only species

Nymphalis flavilla Godart, [1824]) and Peria Kirby, 1871 (type and the only species Papilio lamis

Cramer, 1779). Species of this group are close to each other, e.g., COI barcodes of the type species of

Peria and Temenis differ by only 6.8% (45 bp) and taken together prominently separate from the rest of

the subtribe. Even if Temenis is monophyletic (1.e., if the topology in Fig. 40 is incorrect), it is not

prominently distinct from its relatives, and therefore we propose that they all are congeneric. Thus,

Asterope Hubner, [1819], Nica Htibner, [1826], Peria Kirby, 1871, and Callicorina Smart, 1976 are

subgenera of Temenis Hiibner, [1819]. Curiously, the subgenus Callicorina does not belong to the

subtribe Callicorina.

O5|Mexico:Tamaulipas|1

Panacea Godman & Salvin, 1883 is a junior subjective synonym

of Batesia C. Felder & R. Felder, 1862

A monotypic genus Batesia C. Felder & R. Felder, 1862 (type species Batesia hypochlora C. & R. Felder,

1862) is closely related to Panacea prola

Panacea Boles 9094E10|Colombia|1973

5 Bat hi Se P

(Doubleday, [1848]), the type species of ‘ Hamar ae Februalt7137 178 b Delton. Tamaipaslsen er

: : bez Hamadryas glauconome exico:Tamaulipas

Panacea Godman & Salvin, [1883] (Fig. 41), Hamadry as iphthime| 17 119009 | Mexico: vucatan|197

me sinat-th Rice pase a hasan SO eae

suggesting that the two genera are Su jective Hamadeae Stlantis| 17118 aan Me Qaxacal19

synonyms, thus eliminating the need for yet Pia i RS o aria Si are

another indistinct monotypic genus. The wing

shape and general patterns of these species are quite similar as well, Batesia differing in the absence of

black striations characteristic of Panacea, giving Batesia a more distinctive "cleaner" appearance

otherwise not much different from Panacea, but clearly distinct from their relatives: sister genera Ectima

Doubleday, [1848] and Hamadryas Hiibner, [1806].

Fig. 41. Panacea (red), Batesia (blue) and Hamadryas.

Introductory comments on Nymphalinae

Five genera from the subfamily Nymphalinae Rafinesque, 1815 were placed as incertae sedis by

Wahlberg (2019a): Pycina Doubleday 1849 (type and the only species Pycina zamba Doubleday, [1849]),

Rhinopalpa C. & R. Felder 1860 (type and the only species Rhinopalpa fulva C. & R. Felder, 1860, a

junior subjective synonym of Vanessa eudoxia Guérin-Méneville, 1840, with is a subspecies of Papilio

polynice Cramer, 1779)), Kallimoides Shirézu & Nakanishi 1984 (type and the only species Kallima

rumia Doubleday, 1849), Vanessula Dewitz 1887 (type and the only species Vanessula buchneri Dewitz,

1887, a junior subjective synonym of Liptena milca Hewitson, 1873), and Doleschallia C. & R. Felder

1860 (type species Papilio bisaltide Cramer, 1777). The reasons behind the incertae sedis placement,

such as ancient origin of these taxa forming long branches in the trees, were given by Wahlberg (2019b),

who concluded: "Clearly more data are needed for these ancient taxa." To shed light on their

classification, we obtained whole genome shotgun for the type species of these five genera and placed

them in the phylogenetic context of other Nymphalinae. The tree constructed from protein-coding genes

of the Z chromosome is illustrated (Fig. 42). In this mostly well-supported phylogeny, all currently

defined tribes receive 100% statistical support. The five taxa in question are found in deeper radiation

near the origins of Nymphalinae and are not closely associated with any of the currently defined tribes.

Pycina zamba and Rhinopalpa polynice are confidently placed prior to divergence of other tribes such as

Victorinini Scudder, 1893 and Junoniini Reuter, 1896 (Fig. 42). Doleschallia bisaltide is a confident sister

to Melitaeini Herrich-Schaffer, 1843, but is prominently separated from them, originating well prior to

Coeini Baeotus beotus|19093F01|Costa Rical2009

ots giclee see ie aed cal 1984 Lise

inini istoris acherantal19093F05|Venezuelal1985

fous Pycina pera 095209 bane aliS 3

Smyrna blomfildia] 1¥117HO9|USA4:TR, Nueces Co,|1984

- Colobura dirce|19093F02(Guyanal|2000

at Tigridia acestal]19093F08|Guyanal1999

Nymphalini Hypanartia lethe|17118C08|Mexico: Oaxaca

1 = Antanartia delius|19093E01|Congo|1977

— Nymphalis polychloros|19093E04|France|1976

Vanessa atalanta|19124002|France|1983

Araschnia leyvana|19093E02|Czech Bemublie 1984

Symbrenthia hippoclus|19093 tree Ailippines

Rhinopalpini Mynes halli[19122G01|Indonesial2013

ws ; Rhinopalpa polynice 19092H11i|Malaysia]1974

Kallimini— Kallima paralekta|19092H06|Javalold

Kallimoidini Catacroptera Ca Oot oa 1964

Kallimoides rumial 19092HO?|Cameroon|1987

1 0.82 Siproeta Lata nse e eA CHEST ‘Santa Catarinal1984

ae Napeocles jucunda|19093D11|Brazil: Poa On aa eae

Victominini_— Metamorpha elissa|19093D10|French Guianal1993

Anartia jatrophae Bet JUSA:TX, sald Co,/2015

0.46 Salamis augustina|19092G08 Madagascarlold

Juno ini Yoma sabinal19092610|Myanmar|2001

ube SA, anacardii a 19092611|Ugandal1939

ne Junonia cone 8165/USA: nh man Dade Co,|2017

0.42 : 5 ae actavial {9 Sareea aaa sagas

- ypolimnas pandarus mbon

— Vanessula ee cet: Cameroon|~1980

Doleschallia bisaltide| eee eal

; nathotriche a ae Ge

Higginsius fasciata|19077G610|Peru:Cuzco|1959

Mazia amazonica|19077HO5|Peru|i990

i Ortilia Pee eee a agente cacue

Tisana ae 190784 alargentinatold

aa Tegosa lukal190 aCO3| Mog

Anthanassa texanal4378|USA4:TX, setae Co,J2015

Phyciodes tharos|3395|(USA:TX Hidalgo Co,|2015

Atlantea perezi| 19077407 |Cubal 1990

Antillea 5|19077HO9|Dom.Rep|i994

Melitaea cinxia|19078 clop: Albania|1991

Bikale ate aves MUM pn hears CA,Tulare Co,|[1974

i soil Aue Pours SA:TX, Starr Co. }2015

ra a eUnes a oee Se AZ, Santa Cruz Co,|2016

Euphydryas phae onl 1711 SB0R USE: H Portage Co,[1983

Fig. 42. Nymphalinae tribes. New tribes are colored in red. Specimens shown are actual specimens sequenced.

Doleschalliaini

radiation of Melitaeini into subtribes. Notably, Melitaeini are characterized by an elevated evolutionary

rate reflected in much longer branches within the tribe compared to those of most other Nymphalinae,

including Doleschallia. Finally, although Kallimoides rumia and Vanessula milca are confidently placed

in the clade consisting of Victorinini, Junoniini, Melitaeini and Doleschallia, their exact phylogenetic

position remains weakly supported because they originated during a rapid radiation event near the last

common ancestor of this clade. It is likely that K. rumia is a distant sister of Victorinini. However, the

association of V. milca with Junoniini is questionable. It is not likely that the lack of confidence in this

phylogenic placement is caused by the shortage of sequence data. It is plausible that these taxa are of

hybrid origin, or their evolution involved incomplete lineage sorting resulting in phylogenetic

incongruence among different genes. Regardless of their exact phylogenetic origins, and exactly due to

this phylogenetic uncertainty, these taxa do not belong to any existing tribes. For all the reasons stated

above, each of these five lineages represents a tribe of Nymphalinae, as named below.

Pycinini Grishin, new tribe

http://zoobank. org/DS5COAE41-2615-4852-9B59-8533C27E4798

Type genus. Pycina Doubleday 1849.

Definition. Formerly placed within the tribe Coeini Scudder, 1893, this tribe is not monophyletic with it

and instead is sister to all other Nymphalinae Rafinesque, 1815 excluding Coeini (Fig. 42), which justifies

its status as a new tribe. The tribe is diagnosed by the characters given for Pycina on pages 305-306 by

Westwood (1850) and on page 330 by Godman & Salvin (1884). In brief, diagnosed by densely hairy

eyes, relatively small and thin palpi not extending beyond middle of eyes in lateral view, long slender

antennae more than half of forewing length, elongated forewings, more so than in similarly patterned

Smyrna Hubner, [1823], and undulating costal margin of hindwing: 1.e., prominently concave near the

middle instead of evenly convex throughout as in its relatives.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815.

Rhinopalpini Grishin, new tribe

http://zoobank.org/4A3 ACC99-A761-49B9-BA00-00F83FF443 DE

Type genus. Rhinopalpa C. & R. Felder 1860.

Definition. Originates early in the Nymphalinae radiation and is sister to the clade consisting of several

tribes, e.g., Kallimini Doherty, 1886 and Melitaeini Herrich-Schaffer, 1843 among others (Fig. 42),

indicating that this new taxon is a tribe. Diagnosed by long and densely scaled stout palpi (as suggested

by the name, but not longer than the head as in Libytheinae Boisduval, 1833), discal cells short on both

wings, about 3 of wing length, all forewing R veins run very close together and R3, R4 and Rs stalked for

most of their length, forewings produced into apical and tornal lobes, hindwings with stubby tail in the

middle of outer margin; below with a row of 6 eyespots along the margin of both wings; caterpillar with

long scoli turned anteriad at tips past thoracic segments; pupa unusually shaped somewhat resembling a

dead leaf, with 3 pairs of horn-like protuberances, first pair very prominent, directed posteriad.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815.

Comments. The name Rhinopalpini as published by Teshirogi (2016) appears to be a nomen nudum:

Referred to Wahlberg et al. (2005), a work that does not mention the name, and the name is not "explicitly

indicated as intentionally new" in Teshirogi (2016), thus failing Art. 16.1. of the ICZN Code UCZN

1999).

Kallimoidini Grishin, new tribe

http://zoobank. org/22 C74 A4F-6B24-4502-99BB-B8E3C66AFABE

Type genus. Kallimoides Shiré6zu & Nakanishi 1984.

Definition. This tribe is a distant sister of Victorinini Scudder, 1893 with moderate support (Fig. 42).

Being an ancient lineage as indicated by its phylogenetic placement and only moderate statistical support

those given in detail for the genus Kallimoides on pages 107-108 and wing venation and genitalia are

illustrated in Figs. 5 and 40-45, respectively, by Shirdézu & Nakanishi (1984). Most notably, the new tribe

is diagnosed by S-shaped vein Ri on forewing, partially coalescing with Sc and then with R2; prominently

reduced tegumen and large S-shaped aedeagus terminally thinning into a hook in male genitalia; and the

absence of genital plate in females.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815.

Vanessulini Grishin, new tribe

http://zoobank. org/D584B74C-8DC8-479A-AD25-9197A03A843B

Type genus. Vanessula Dewitz 1887.

Definition. Belongs to the same clade with Victorinini, Junoniini, Melitaeinit and Doleschallia, but not

closely allied to any of them as judged by its ancient origin and poor statistical support for its placement

within this clade, justifying its status of a tribe (Fig. 42). The tribe is diagnosed by the characters given for

Vanessula on pages 145-146 by Dewitz (1887). In brief, forewing vein Ri near its origin partly coalescent

with Sc, vein R2 stalked with R3, R3 with R4 and R4 with Rs for at least half of their lengths each, discal

cell closed, short, about 1/3 of forewing; eyes bare, palpi long and thin, slightly longer than head,

antennae half of forewing length; wings moderately elongated, wing shape similar to Melitaeini.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815.

Doleschalliaini Grishin, new tribe

http://zoobank. org/88D32044-DB8E-486F-8C09-0D 188ES5CDC93

Type genus. Doleschallia C. & R. Felder 1860.

Definition. Formerly included in Kallimini Doherty, 1886 due to extensive superficial similarity. In the

genomic tree, confidently placed as sister to Melitaeini Newman, [1870], but more distant from them than

they are from each other, and is of an ancient origin estimated nearly 70 Mya (Su et al. 2017), which

justifies its status of a tribe (Fig. 42). Similar to Kallima Doubleday 1849 in wing shape and coloration,

apparently due to convergence. Diagnosed by its produced forewing apex and hindwing tornus extended

in a short tail, below colored as a dead brown leaf with mid-rib (as in Kallimini), with several eyespots

along the wing margins; discal cells open on both wings (closed in Kallimini), all forewing R veins close

together, R4 and Rs stalked for 2/3 of their length, hindwing humeral vein simple (branches into 2 in

Kallimini); eyes bare, pupa smooth and rounded, abdomen segments without protuberances that are

present in Kallimini.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815.

Comments. Doleschalla Walker 1861 (Diptera: Tachinidae) is the type genus of Doleschallini. According

to Art. 55.4. of the ICZN Code ICZN 1999), one letter difference, which in this case would be an extra

"i" from the suffix "-ina", avoids homonymy. However, it does not avoid confusion, including internet

search engines that correct possible spelling errors, especially in this case, because it is a duplication of a

letter in the suffix formation from the correctly determined stem. The Code gives a choice to alleviate the

problem: Art. 29.4. (original stem formation to be maintained) and Recommendation 29A are followed

here, and the entire name of the type genus is taken as a stem with hopes to diminish confusions. The

choice of being ridiculed for grammatically incorrect stem formation is a "lesser evil" compared to

increased confusion of those who use the name.

Additional genomic support to partition Nymphalis genus group into 4 genera:

Aypanartia Hiibner, [1821], Vanessa |Fabricius], 1807,

Antanartia Rothschild & Jordan, 1903, and Nymphalis Kluk, 1780

To achieve a more objective, internally consistent and eventually stable classification, we have placed

Aglais Dalman, 1816 (type species Papilio urticae Linnaeus, 1758) and Polygonia Hubner, [1819] (type

species Papilio c-aureum Linnaeus, 1758) as subgenera of Nymphalis Kluk, 1780 (type species Papilio

polychloros Linnaeus, 1758) on the basis of genomic evidence (Zhang et al. 2020). Here, we obtained and

H rtia lett 17118C08|Me a 1987

analyzed whole genome shotgun nearer ere ee ee eee

a4 : Hypanartia bella|19122E06|Brazil:Paranal1995

sequences of additional taxa from Hypanartia| C——_ypanartia paullus|19122€07 [Dominican Republic|1981

2 7 Hypanartia kefersteini|19122E08|Peru|2015

the Nymphalis group, defined as a : Hypanartia cinderella|19122E09|Peru|2015

3 , Hypanartia christophori|19122E11/Colombial1982

moderately prominent clade = fol UE ee elec ave Co,|2017

. es o Vanessa carye|19093E12|Bolivia|2009

formed by Hypanartia Hitibner, oh — Vanessa SEE aiBentaiNce Zealand|1968

1821 t : H : | Vanessa Vanessa Dat Ee ee reuleea ten nee isemetiont:

= 1 anessa carduli allas Co,

[ | (type species ypanartia os] : vanlente elt dl Be en Fealand|1968

. . . . . E = Vanessa myrinnal19121H10 cra face

1S a junior subjective synonym of = Vanessa terpsichore|19121H06|Chile|19

“ps yon vanessa abyssitical1g{22A00/ethopia| i963

a anessa dejeanii avalo

Papilio lethe Fabricius, 1793) and ma vanesee vuleaniali9 z2A0elno dstalold

7 : : 5 anessa indica anmar

its sister taxa that include Vanessa 7— Vanessa atalanta|S826|USA. Tx pallas C012017 |

fs - anessa tameamea awail

[Fabricius], 1807 (type species ee ee ae erie laae

: anessa dimorphica anzania

Papilio atalanta Linnaeus 1758) Antanartia 7 Antanartia schaeneial 19122611|Scuth Afr |

) ) i Antanartia schaeneia|19122B11|South Africal1978

2 . ieces Nymphalis Pee io |PAOE20|Germany|2017

Antanartia Rothschild & Jordan, ; Aa ll fAglais SET eat ag

iT; Si ue ele la acre ere MIELE

a 22 é

1903 (type SPecles Papilio delius Agiais| Nymphs Aglais miberti|s4o0/USA vi Fayico. yellowstone, NP|2017

° . a

Drury, 1782) and Nymphalis (Fig. Nymphalis| * 1 RPRASIASIStArTaliEn polychioros|19093E04|France|1976

A? Th Iti h | a Nymphhalk aoa fil lee cab Uae hep te pratt cae

ymphalis (Nymphalis} xanthomelas ina

1 1 1 — Nymphalis (Nymphalis) cyanomelas|15101E02|Mexico:Veracruz|o

agreement with previous studies Kaniska_ Nymhalls (rank) canacel 1909308) Myanmari20oi.

. . e a

(Wahlberg and Nylin 2003; s Nymphalls (Pelygoria) egeal19122305|Greece| 1983

W hlb d R bi ff 2011) Polygonia : 296 Nymphals (Pelygonial caibum|paceoe|Swizariand[2017 mete

a erg an ubino ymp alis (Polygonia) faunus arimer Co,

mphalis (Polygonia) undinal19122B07 [China| 1

Fi 43 N t bl 1 K, . k Bers at (Paty aa Dee Pate e race naerieea

te ae) ns NOD oniely aise sf wheat degdaeny aeraaseaeleasaL ss ramalas its

Moore, [1899] (type and the only i ane! sa ile g Se ones IER eee vagaries

1 ymphalis (Polygonia) commal alk Co

1 V7 1 Nymphalis (Polygonia) gracilis|P40246|US4:CO,Larimer Co,.|2016

species Papilio candace Linnaeus, - Nymphalis (Pal annals Harel yi 15B06|Mexico: Hidalgo|1981

: “1: é ; Nymahalis ace tile oreas|17115B05|US4:OR,Yamhill Co.]1983

1763) 1S placed within Nymphalis Nymphalis (Polygonia) progne|17115B04|USA:SD,Roberts Co,|1981

as a subgenus, sister to subgenus Fig. 43. Vanessa group genera: Hypanartia (purple), Vanessa (cyan),

Dal vara whe Odecates suBDOr: Antanartia (red), and Nymphailis (blue, subgenera labeled at their clades).

VE pport,

(2) Roddia Korshunov, 1995 (type and the only species Papilio l-album Esper, 1781) is situated at a

distance from and thus is a valid subgenus sister to subgenus Nymphalis; (3) Inachis Hubner, [1819] (type

and the only species Papilio io Linnaeus, 1758) is similarly distanced from Aglais as Roddia from

Nymphalis sensu stricto and therefore is a valid subgenus; (4) the genus Antanartia is sister to Nymphalis,

is prominently distinct from it, and forms a long branch in the tree indicating accelerated evolution, which

resulted in its relatively unusual wing shape and pattern, likely due to convergence more similar to the

two Vanessa species formerly placed in Antanartia (Vanessa hippomene (Hiibner, [1823] and Vanessa

dimorphica Howarth, 1966) rather than to Nymphalis. Overall, the Vanessa group has split over a

relatively narrow time period into 4 most prominent clades, which (except Antanartia) diversified around

the same time, indicating evolutionary significance of that time point. Therefore, it is meaningful to define

genera as these 4 clades, assigning more recent diversifications to subgenera. This study completes

genomic coverage of all distinct lineages in the Nymphalis group, and the results are unlikely to change

after inclusion of the remaining species.

Napeocles Bates, 1864 is a junior subjective synonym of Siproeta Hiibner, [1823]

We obtained genomic data for all 10 known species of Victorinini Scudder, 1893, and the resulting

phylogenetic tree revealed that the monotypic genus Siproeta epaphus|19093007 |Brazil:Santa Catarinal1984

Napeocles Bates, 1864 (type species Hamadryas Siprocta sislones|3969 [USA TX,start Co, [2015.

jucunda Hiibner, [1808]) clusters very closely with SR eC IGOra ne alicea GOGAT eran aceraiTeas

. A . : . , Anartia chrysopelea|17119E08|Cuba:Habana/1959

Siproeta Hubner, [1823] (type species Siproeta trayja * anartia iyereali9093006|Dominican Republic|1994

os ‘ . Anartia Jatrophae|3s615|USA:Tx,Starr Co,[2015

Htibner, [1823], currently a subspecies of Siproeta ; Anartia fatima|17117012/USA:TXHidalgo Co.|1972

* Anartia amatheal19093001|Surinamel1982

epaphus (Latreille, [1813])) (Fig. 44). The 4 species in

this clade clump together, suggesting that Napeocles

Bates, 1864 is a junior subjective synonym of Siproeta Hiibner, [1823] despite profound differences in

wing shape and patterns, a result of apparently convergent similarity with dead leaf mimics.

Fig. 44. Siproeta (blue and red) among Victorinini.

Tribe Melitaeini Herrich-Schaffer, 1843 consists of two subtribes

We sequenced and analyzed genomic data for all valid genera and nearly all available genus-group names

of the tribe Melitaeini Herrich-Schaffer, 1843. The resulting nuclear all-gene phylogeny of selected most

distinct taxa agrees with previous studies based on gene markers (Wahlberg et al. 2005; Wahlberg and

Freitas 2007; Long et al. 2014) (Fig. 45). The genome-based phylogeny reveals nearly extreme

evolutionary rate heterogeneity among different clades of Melitaeini, as much as 3-fold difference: with

Euphyadryas Scudder, 1872 (type species Papilio phaeton Drury, 1773) having the lowest rate (=shortest

distance in horizontal dimension from

the root on the left to the tree leaves

on the right) and insular taxa such as

F T

Euplaseying ydryas aurinia| 1907 ¥G04|5 eden/1974

Euphydryas phaeton|17115B80 Port

‘ sal i Maltaea gingia/ 130 bania

Melitaea > Mell fea athalia]190

Melittaea ornatal190

a Euphydryas gillettii|17115B07 |USA:MT Cascade Co.|1996

— uphyd ?

qe Co.J1983

991

con

Atlantea Higgins, 1959 (type species

Synchloe perezi Herrich-Schaffer,

18620) having the highest rate

(=longest distances: branches for these

taxa are sticking out to the right from

the rest). This heterogeneity is likely

connected to the high diversification

rate in the tribe (species richness) and

leads to the difficulties with its

classification. From its last common

Melitaeini

0.96

Poladryas

4 1

0.98

Chlosyne||— ik

Microtia —=

Gnathotriche 7

Aigginsius 7

i Antiliea

Atlantea

Phystis

Wazia

0.85

Chlosyne fulvia

Chigsyne nyctels | 44

Chia

Gnathotriche exclamatia

hla

syne ite

exo erse

faxola e Seirsde

Texola. coracaral1

Dymasia dymas|5

Melitaea didymali

Poladryas ar cee

Pola ryas minuta

— as

—s

fat

far

syne eArennerat

syne janals

y efinita are |

icrotia elval106

aha

_ Gnathotriche mundina

Higginsius fasci Plis07

HIQginsius mirilam|l9o7 "G1

Antillaa pel

Antilea croc

tlantea

_ Atlantea pa

“Atlantea, perez

Phystis simois/19

Ortilta

ancestor, the tribe divides into two

most prominent clades: Euphydryas

evolving with the speed typical for

Nymphalidae (Fig. 42) and its sister

that includes all other taxa that evolve

at progressively elevated _ rates.

Despite the elevated rates, the internal

branches in this sister to Euphydryas

clade are shorter than the branch

leading to it, suggesting that the taxa

within this clade are closely related to

each other and are not as prominently

distinct so to define as subtribes ae = Bagon eels gentina Tysuman:

compared to the whole clade itself. cee CCC cuyanal2900 BBtalp lea?

Therefore, instead of dividing this : seri: Cuzco |3¢

clade into additional subtribes, we

propose that the Melitaeini consists of

only two subtribes: Euphydryina

Higgins, 1976 and Melitaeina Herrich-

Schaffer, 1843, which is the major and

unquestionable division of the tribe. The most prominent (i.e. the longest compared to their surrounding)

internal branches within Melitaeina are defined here as genera (Fig. 45). Many of these genera correspond

to those in use today, 1.e., Melitaea Fabricius, 1807 (type species Papilio cinxia Linnaeus, 1758),

Poladryas Bauer, 1975 (type species Melitaea arachne W. H. Edwards, 1869), Chlosyne Butler, 1870

(type species Papilio janais Drury, 1782), Gnathotriche C. & R. Felder, 1862 (type species Euterpe

exclamationis Kollar, 1849), Higginsius Hemming, 1964 (type species Melitaea fasciata Hopffer, 1874),

Antillea Higgins, 1959 (type species Papilio pelops Drury, 1773), and Atlantea are traditionally used

genera that are prominent clades in the genomic tree (Fig. 45). Other cases suggest adjustment to ranks

and names that are detailed below.

0.6 .

Phyclodes mylitta]1

Pp an es bearas 33

0.54

v. hyciodes ao

FP ydodes obichelia|

Phyciodes graphica

i Tegosa lukal106

Tegosa nigrellal1

Tegosa etia/19

Tegosa flay

7 Ortilia

Ort

Phyciodes

sensu

o4s| —-Sticto

Subgenera

of Fresia

Notilia subgen. n.

Ortilig sejo

Ortilla dicomal19o7s

Eresia levina|19077H

Anthanassa ptolycall

Anthanassa drusilla]1

Levinata subgen. n.

Anthanassa

Casfilia

an usta

Castilia caspila 19

Telenassa abas|19

Telenassa delphial1 LI!

[>> lelenassa berenic 6[Peru:A

Telenagsa sts 08|Brazil: 996

Telenaséa

Ortila tthral 1907 zil:sao aie 1991

Dagon catylal19 rgentina:Tucuman|2005

Janatella

Fig. 45. Melitaeini subtribes Euphydryina and Melitaeina, and genera,

including Microtia (green), Ortilia (olive and pink), Phyciodes (purple),

Tegosa (cyan) and Eresia (red, orange, blue, and magenta).

Texola Higgins, 1959 and Dymasia Higgins, 1960

are junior subjective synonyms of Microtia H. Bates, 1864

Microtia H. Bates, 1864 (type and the only species Microtia elva H. Bates, 1864), Dymasia Higgins, 1960

(type and the only species Melitaea dymas W. H. Edwards, 1877) and Texola Higgins, 1959 (type species

Eresia elada Hewitson, 1868) cluster closely in the genomic tree (Fig. 45). Our result agrees with the

previous assessment based on gene markers (Wahlberg et al. 2005; Wahlberg and Freitas 2007; Long et

al. 2014) and morphological study by Kons (2000), who already synonymized Texola and Dymasia with

Microtia, a suggestion cited by Wahlberg et al. (2005) five years later but not followed either in that work

(Wahlberg et al. 2005) or in subsequent publications (Pelham 2008; Long et al. 2014) and many on-line

resources (Warren et al. 2016; North American Butterfly Association 2018; Wahlberg 2019a; Pelham

2020). Finally, more than two decades after these studies, we reach the same conclusion on a much larger

DNA dataset: Texola Higgins, 1959 and Dymasia Higgins, 1960 are junior subjective synonyms of

Microtia H. Bates, 1864. A curious observation is that Microtia |=Texola| coracara (Dyar, 1912) appears

to be more distant from other Texola than Dymasia from Microtia (Fig. 45), further supporting unification

of these species. The unification is a more meaningful solution, because this clade actually consists of

four semi-equal lineages, which are (1) M. elada (Hewitson, 1868) species group; (2) M. anomalus

(Godman & Salvin, 1897) species group that includes M. coracara; (3) M. dymas; and (4) M. elva; rather

than the traditional three (Texola, Dymasia and Microtia). An alternative could be to split the group into

these 4, rather than 3, evolutionary lineages, but each of these 4 groups is nearly monotypic (or monotypic

under some species concepts) and is a species group rather than a genus or even subgenus. Finally,

Microtia sensu lato is characterized by an elevated evolutionary rate compared to its sister Chlosyne

Butler, 1870 (type species Papilio janais Drury, 1782): in Fig. 45, branches of Microtia (green) are longer

(i.e., stick out to the right more) than branches of Chlosyne (black). This elevated rate may be behind

more pronounced phenotypic differences between Microtia species compared to Chlosyne that resulted in

the oversplit classification of Microtia into genera.

Tisona Higgins, 1981 is a junior subjective synonym of Ortilia Higgins, 1981

The monotypic genus Jisona Higgins, 1981 (type and the only species Phyciodes saladillensis

Giacomelli, 1911) was proposed on the basis of genitalic

differences, most significantly "penis apex with crossing

ostium-folds" (Higgins 1981), 1.e., with two twisted finger-like

processes at the tip. Higgins likened Tisona to Tegosa Higgins,

1981 (type species Acraea claudina Eschscholtz, 1821). To ae

better understand its phylogenetic affinities, we sequenced the Mr er

only available although century-old specimen of Tisona | Gonick. Grishin’ Eas

saladillensis in the National Museum of Natural History

collection (USNM) (Fig. 46). It lacks the right hindwing, and

one of its labels, probably written by William Schaus, reads ||

Argentina | Giacomelli ||, suggesting that this specimen may

have been part of the type series. In the genomic tree, it clusters

closely and confidently with Ortilia Higgins, 1981 (type species

Papilio liriope Cramer, 1775), away from Tegosa (Fig. 45).

COI barcodes of O. liriope and T. saladillensis differ by 5.5%

(36 bp). This small difference strongly suggests that the two |

species are congeneric, and we place Tisona Higgins, 1981 as a Fig. 46. Sequenced specimen of Ortilia

junior subjective synonym of Ortilia Higgins, 1981. Although saladillensis. Labels reduced 2 times

the unusual genitalia of this species revealed its uniqueness Commparsehtgie Specumen,

despite the common Phyciodes-like wing patterns, DNA analysis was necessary to understand its origins

and classification.

Anthanassa Scudder, 1875, Castilia Higgins, 1981, Telenassa Higgins, 1981, Dagon

Higgins, 1981, and Janatella Higgins, 1981 are subgenera of Eresia Boisduval, 1836

The most inclusive prominent clade that contains Phyciodes Hiibner, [1819] (type species Papilio cocyta

Cramer, 1777) and is consistent in genetic diversification (corrected for its higher evolutionary rate) with

how other Melitaeina genera are defined is labeled "Phyciodes sensu lato" in Fig. 45. It includes the

majority of Melitaeina genera and species as they are presently defined. This current classification that

remains largely unchanged since the Higgins revision (1981), represents the other extreme and appears to

be oversplit, because many genera defined by Higgins are too close genetically and do not stand out as

prominent clades in the tree (Fig. 45, first word in species names). Most notably, Anthanassa Scudder,

1875 (type species Melitaea texana Edwards, 1863), Castilia Higgins, 1981 (type species Eresia castilla

C. & R. Felder, 1862), Telenassa Higgins, 1981 (type species Argynnis teletusa Godart, [1824]), Dagon

Higgins, 1981 (type species Eresia catula Hopffer, 1874), Janatella Higgins, 1981 (type species Eresia

leucodesma C. & R. Felder, 1861), and Eresia Boisduval, 1836 (type species Nereis eunice Hiibner,

[1807]) cluster closely together, and the internal branches separating them are short and indistinct. While

most of Higgins genera are indeed monophyletic, with some exceptions noted in the tree by colors (Fig.

45) and discussed below, the lack of their distinctiveness and low divergence is more consistent with that

of subgenera. E.g., COI barcodes of the type species of Eresia and Anthanassa differ by only 6.5% (43

bp), without any correction for the generally elevated evolutionary rate of these lineages. Moreover, even

Phyciodes is not particularly removed from this cluster of closely related "genera": COI barcodes of

Phyciodes tharos (Drury, 1773) and Anthanassa texana (W. H. Edwards, 1863) differ by only 7.3% (48

bp). Therefore, we would generally favor broader defined genera and could propose "Phyciodes sensu

lato" (Fig. 45) as a genus to combine most of these Higgins genera. However, looking for a compromise

to meaningfully classify this species-rich lineage, we opt for a centrist solution. We note that Phyciodes

sensu stricto and Tegosa Higgins, 1981 (type species Acraea claudina Eschscholtz, 1821) do form

somewhat prominent clades in the tree, and we keep them as genera. As a result, clades split prior to them

also keep their genus rank: Phystis Higgins, 1981 (type and the only species Eresia simois Hewitson,

1864), Mazia Higgins, 1981 (type and the only species Melitaea amazonica Bates, 1864) and Ortilia

Higgins, 1981 (type species Papilio liriope Cramer, 1775). All others: Anthanassa Scudder, 1875,

Castilia Higgins, 1981, Telenassa Higgins, 1981, Dagon Higgins, 1981, and Janatella Higgins, 1981 are

placed as subgenera of Eresia Boisduval, 1836. Finally, we note several clades in the tree that disrupt

monophyly of the genera as defined by Higgins (Fig. 45: red and magenta "Ortilia" and orange "Eresia").

These clades also appear in the phylogenetic studies based on gene markers, although their exact positions

relative to the other clades varied (Wahlberg and Freitas 2007; Long et al. 2014). To remove ambiguity

about their status, these clades are named as subgenera of Eresia below.

Notilia Grishin, new subgenus

http://zoobank.org/3 A8FA 139-BE8C-444A-A212-6108DA6D3EAB

Type species. Eresia orthia Hewitson, 1864.

Definition. Previously placed in Ortilia Higgins, 1981 (type species Papilio liriope Cramer, 1775), this

group is not monophyletic with it. Instead it is sister to all other Eresia sensu lato, and is close to them

(Fig. 45). Previously discovered and defined as the Brazilian "Ortilia" clade by Wahlberg and Freitas

(2007), this phylogenetic group was also confirmed and discussed in a more recent study suggesting "that

this clade requires a new name" (Long et al. 2014). Due to its genetic closeness to Eresia sensu lato, this

distinctive lineage is named as a subgenus of Eresia rather than a separate genus pending further analysis.

It keys out to Ortilia in Higgins (1981) sharing the following diagnostic combination of characters with it:

antennal club pyriform, aedeagus end without a pair of twisted processes, tegumen reduced, scaphial

extensions small, without hooks and spines at angles, saccus single, narrow, finger-like, not expanding

terminally, without a cleft. Differs from Ortilia in shorter and straighter harpe projecting directly

caudodorsad (not arched at its origin changing direction from cephalodorsad) and less extensive or absent

fulvous markings on wings above.

Etymology. The name is a feminine noun in the nominative singular, formed as a fusion Not + [Ort]ilia to

indicate distinction from Ortilia.

Species included. The type species, Phyciodes orticas Schaus, 1902, Phyciodes sejona Schaus, 1902,

Eresia velica Hewitson, 1864, Eresia dicoma Hewitson, 1864, and Phyciodes polinella Hall, 1928.

Parent taxon. Genus Eresia Boisduval, 1836.

Levinata Grishin, new subgenus

http://zoobank.org/S5C69C6A2-7ADD-4E42-8818-23CCB9IA042CE

Type species. Eresia levina Hewitson, 1872.

Definition. Previously placed in Eresia Boisduval, 1836 (type species Nereis eunice Hiibner, [1807]), this

group is not monophyletic with it. Instead it is sister to all other Eresia sensu lato but Notilia subgen. n.

(Fig. 45). It is a phenotypically distinctive subgenus diagnosed by wings rounder than Eresia with broad

discal band across forewing, blue in the type species. Its male genitalia are distinctive, with diagnostic

characters for this new subgenus as described for Eresia levina on page 150 and illustrated in Figs. 430—

431 in Higgins (1981): 1.e., genital capsule larger than in other Eresia, tegumen sclerotized, shoulders

well-developed, scaphial extension expanded, sclerotized and terminally with many small teeth, valvae

with fine teeth on inner sides near apex, aedeagus terminally rounded, without ostium keel.

Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.

Species included. Only the type species.

Parent taxon. Genus Eresia Boisduval, 1836.

Ithra Grishin, new subgenus

http://zoobank. org/BA DF7F37-1CFB-4846-A C56-6B 125E8B6708

Type species. Phyciodes ithra Kirby, 1900.

Definition. Previously placed in Ortilia Higgins, 1981 (type species Papilio liriope Cramer, 1775), this

group is not monophyletic with it. Instead it 1s sister to Dagon Higgins, 1981 (type species Eresia catula

Hopffer, 1874), but not with decisive statistical support (Fig. 45), and therefore is a distinctive lineage of

the same rank, 1.e., subgenus. Diagnosed by male genitalia and the characters for this subgenus are as

those given for Ortilia ithra on page 120 and illustrated in Figs. 318-320 in Higgins (1981). Keys out to

Ortilia in Higgins (1981) due to the following combination of characters this new subgenus shares with

Ortilia and Notilia subgen. n.: pear-shaped (not extended) antennal club, no elongated processes at

aedeagus end, reduced tegumen with small scaphial extensions not armed with hooks or spines, saccus

single, terminally narrower, without a cleft. Differs from Ortilia and Notilia subgen. n. in having larger

genital capsule, longer and bulkier saccus, prominent but short in dorsal view tegumen with very short

scaphial extensions, short apical sections of valvae with thicker and rather straight harpes directed

caudodorsad, aedeagus narrowing from phallobase caudad in basal half.

Etymology. The name is a feminine noun in the nominative singular, tautonymous with the type species

name.

Species included. Only the type species.

Parent taxon. Genus Eresia Boisduval, 1836.

Phystis fontus (A. Hall, 1928), new combination

To our initial surprise, the distinctive species Telenassa fontus (A. Hall, 1928) (type locality Guyana, Fig.

47 middle) was placed among subspecies of Phystis simois (Hewitson, 1864) (type locality Brazil, Fig. 47

left and right) in the genomic tree, rendering P. simois paraphyletic (Fig. 48). Suspecting contamination

from the specimens of P.

simois that were sampled next

to it, we initially removed this

otherwise excellent sample

(NVG-19077H04) from the

analysis pending DNA re-

extraction and re-sequencing.

However, 7. fontus and P.

simois were found to be

sisters based of gene markers

(Wahlberg and Freitas 2007;

Long et al. 2014) supporting

the validity of our genomic

results. Due to this genetic similarity, we place 7. fontus in Phystis to form Phystis fontus (A. Hall, 1928),

comb. n. and marvel about evolutionary plasticity leading to distinctly different wing shapes in this small

genus.

Phystis pratti stat. nov. Phystis fontus fontus comb. nov. Phystis variegata stat. nov.

Fig. 47. Phenotypic diversity in Phystis. Sequenced specimens are shown.

Phystis variegata (Roéber, 1913) and Phystis pratti (A. Hall, 1935) are species distinct

from Phystis simois (Hewitson, 1864)

Confident paraphyly of the species Phystis simois (Hewitson, 1864) (type locality Brazil, Fig. 47 left and

right) with respect to Phystis fontus (A. Hall, 1928), Phystis simois pratti|19077HO3|Peru:Piura/2000

A . 3 Telenassa Ul rt oes GORA ANSI ls oe

comb. n. was one of the more unsettling results in this Phystis simois wr ale 77HO6|Paraguay1986

; 7 Mazia amazonical19077HO5|Peru:Loreto|1990

project (Fig. 48). However, COI barcodes of the two a ,Ortiia Iriope|19077H11|Guyanal2000

sgh faisusten oopna ding eee ee ee

* orti la gentina razil:Rondonia

subspecies Phystis Aaa pratt (A. Hall, 1935) (type : Grtlie pentinal19078A11 [Bolivia ‘La Paz|2003

locality North Peru, Fig. 47 left) and Phystis simois piace sal eneie danas Praenena le

Fig. 48. Phystis (blue, red) and Ortilia (purple, cyan).

variegata (R6ber, 1913) (type locality Argentina, Fig.

A7 right) differ by 6.4% (42 bp), which is more than the difference between some of the Higgins genera

(Higgins 1981), see above. Inspection of specimens (Fig. 47 left and right) reveals marked difference in

wing shapes of the two subspecies, not commonly found within species. Therefore, these two subspecies

are species that are also distinct from the nominotypical Phystis simois (Hewitson, 1864) (type locality

Brazil) due to phenotypic differences and COI barcode differences (7.1% & 8.1%) between our specimens

and P. simois available from GenBank (accession EF493956) (Wahlberg and Freitas 2007). Hence, the

names for these species are Phystis variegata (Réber, 1913) stat. nov. and Phystis pratti (A. Hall, 1935)

stat. nov. We have not studied Phyciodes chinchipensis Hayward, 1964 (type locality Peru: Rio

Chinchipe) currently treated as a subspecies of P. simois. Nevertheless, examination of the holotype

photographs (Warren et al. 2016) suggests that it is not P. simois. Due to wing pattern resemblance and

locality (both are from North Peru), we tentatively place it as a subspecies of P. pratti instead, keeping in

mind that it may be a distinct species and not a mere color variation.

Abananote Potts, 1943 and Altinote Potts, 1943 are

junior subjective synonyms of Actinote Hiibner, [1819]

In agreement with previous studies (Silva-Brandao et al. 2008; Carvalho et al. 2020), we find that genera

Actinote Hiibner, [1819] (type species Papilio thalia Linnaeus, 1758), Abananote Potts, 1943 (type

species Acraea abana Hewitson, 1868) and A/tinote Potts, 1943 (type species Heliconius neleus Latreille,

[1813]) are closely related and some of them are not monophyletic as currently circumscribed (Fig. 49).

The type species of Abananote and Altinote are quite gaa fendva(ena)isa7e.0alMadabedca a9!

closely related: e.g. their COI barcodes differ by 6.4% pote an orisit | S083 91 saLstralsi 2013

(42 bp), the same divergence as between the two former FF eRe ESTE anee ae TEE

e e e . 1 ~ 1

subspecies of Phystis simois (see above), and therefore Dae tacae ineraaoatioi signs Lecnaiola

are congeneric. The Actinote clade is separated from Fe eR ett oerGr] ISGSSHeINETaI tas

them by a prominent gap (Carvalho et al. 2020), and the | | °/4-e=4 TEHInS Sefsns|{edseeorlMadausecsrlteco

barcodes of A. thalia and A. neleus differ by 8.5% (56 = Felehinid camivittaa| 1900404 | Camarcon|1972

: . . . : Talchinia ancedon|19086403|Madagascar|1985

bp), which would suggest their attribution to different Telchinia lycoa|19086406|Cameroon| 1987

‘ 3 F Actinote thalia]19076H12|Suriname|old

subgenera. However, species phylogeny is at odds with —“Altinote ozomene|15098D09]ST orizava|Mexicolold

: ae : . peg. Abananote abanal19086A401|Ecuador|2002

intuitive phenotypic assessment (Lamas 2004): there is “‘Altinote neleus|19086A02|Peru:San Martin|1998

an additional third clade unexpected from phenotypes Fig. 49. Actinote (blue, magenta and orange).

(Silva-Brandao et al. 2008; Carvalho et al. 2020). Therefore, until these inconsistencies between

phenotypic and genetic classifications of this group are sorted out, we propose placing Abananote Potts,

1943 and Altinote Potts, 1943 as junior subjective synonyms of Actinote Hubner, [1819]. This treatment is

consistent with the unified, and more genetically diverse, Old World genus Te/chinia Hiibner, [1819]

(type species Papilio serena Fabricius, 1775) that is sister to Actinote sensu lato (Fig. 49).

Episcada Godman & Salvin, 1879 is a junior subjective

synonym of Ceratinia Hiibner, 1816

Genome-level phylogeny confirms problems with the current classification of Episcada Godman &

Salvin, 1879 (type species /thomia salvinia Bates, 1864) Callithomia lenéal19097F01|Panamal1976_

d Ceratinia Hiibner, 1816 (t ies Neréi Haenschia dersmallg0S7Fo9|perujs0L1

Hiibner, [1806]) (Willmott and Freitas 2006): the two { Pteronyinva lett a7Ft2| Janezuel os

. a. a. c=] 2 ce ‘

genera are closely related and Episcada renders Ceratinia Ceratinia neso]19097F02|Brazil:Paral1978

hvletic (Fig. 50). i t with other studi as — Foiscada dote|19007F09 [Braci:Paralold

parap y CIC ( 1g. ), In agreemen wil O er StudIes - ESS REE salvinia| 19097F06|Mexico:Veracruz| 1984

(Chazot et al. 2020). COI barcodes of the Episcada and Sitgpaent si sella mareauuc ne ge Calas

Ceratinia type species differ by only 4.4% (29 bp), less Fig. 50. Ceratinia (blue and red).

than between the two former subspecies of Phystis simois (6.4%, see above). Therefore, we propose

treating Episcada Godman & Salvin, 1879 as a junior subjective synonym of Ceratinia Hitibner, 1816.

Family Hesperiidae Latreille, 1809

Phocides batabano (Lucas, 1857) and Phocides bicolora (Boddaert, 1783) are species

distinct from Phocides pigmalion (Cramer, 1779)

In his key, Evans (1952) stated that uncus flanges in genitalia of continental subspecies of Phocides

pigmalion (Cramer, 1779) (type locality Suriname) are as long as the uncus, but are shorter in the island

subspecies, being similar to other Phocides Hiibner, [1819] (type species Phocides cruentus Hiibner,

[1819], which is Hesperia polybius Fabricius, 1793). Consistently with this notable genitalic difference,

the genomic tree partitions ig pigmalion into several > moocides pigmalion okeechobee|5316|USA:FL,Monroe Co.|2015

. . . . . — Phocides pigmalion batabanocides|17099412|Bahamas|1981

groups (Fig. 51) rendering it paraphyletic with Phocides pigmalion batabano|14103F05|Cubal2010

’ Z Phocides pigmalion bicolora]17099410|Dominican Republic|1973

respect to Phocides belus Godman & Salvin, 1893 Phocides pigmalion bicolora]17099411|Dominican Republic] 1948

locality Mexi d Phocides li Herrich F- Phoddes eigmaionl170oSDO4|Perul1977

= oe ocides pigmalion eru

(type OCallly ex1CO) an . ocides incea ( CIT1e sex Phocides belus|17113G07|Mexico:Sinaloa| 1996

y Phocides lincea]15029C07 |old

Schiiffer, 1869) (type locality not stated, probably meal eae eet ae

the Guianas), and suggesting that P. pigmalion is a PDUs Bg stall ea UB sMalpe aaa

Fig. 51. Phocides batabano (red), bicolora (magenta),

pigmalion (blue), lincea (green) and perkinsi (cyan).

complex of several species. While the inclusion of

P. belus in the pigmalion group is expected because

Evans (1952) treated it as a subspecies of P. pigmalion, P. lincea was a surprise. Linked by Evans (1952)

with Phocides perkinsi (Kaye, 1931) (type locality Jamaica) as its subspecies that was elevated to species

only recently (Turner and Turland 2017), and placed at the end of Evans' key, P. lincea has not been

associated with the pigmalion group before. Our genomic results definitively confirm P. perkinsi (Fig. 51

cyan) as a species-level taxon, because it is far removed from P. lincea (Fig. 51 green). Furthermore, due

to genetic and genitalic differences, we reinstate Phocides batabano (Lucas, 1857) (type locality Cuba)

and Phocides bicolora (Boddaert, 1783) (type locality not stated, likely Haiti) as species. The COI

barcodes of P. batabano and P. bicolora differ by 2% (13 bp), and P. pigmalion (from Ecuador) and P.

batabano by 3.3% (22 bp). Wing patterns agree with this partitioning of P. pigmalion as it was defined by

Mielke (2005) previously, into at least three species: forewing hyaline spots absent (P. batabano), present

but narrow and wings green-striped (P. bicolora), and present and broader, wings blue-striped (P.

pigmalion). To accommodate this treatment, we revise species-subspecies combinations as: Phocides

batabano okeechobee (Worthington, 1881) and Phocides batabano batabanoides (W. Holland, 1902).

Lobotractus mysie (Dyar, 1904) is a species

distinct from Lobotractus valeriana (Plétz, 1881)

Correctly associating then "Codatractus" mysie (Dyar, 1904) with "Codatractus" valeriana (P\6tz, 1881)

as close relatives by visual inspection of photographs say Lobotractus valeriana (=mysie)|15101C05|LT|AZ,Sta Cruz Co.

. . . . ef obotractus Valeriana|15102C12|USA:42,Santa Cruz Co,|1983

of their primary type specimens, Mielke and Warren pelea valeriana|15105B05|USA:Az,Santa Cruz Co.|2005

(2004) " i 3 mi Loboatractus valeriana|5675|USA‘AZ,Santa Cruz Co.)1986

concluded that "valeriana ... is clearly Lobotractus valeriana|5674|USA;AzZ,Santa Cruz Co,|1991

conspecific with T m se " We sequenced the oats EAE at ani teres anaeenie | Wesieselie Take

eee Ee ohotractule valarara|is1g2cd4|Mextco-Higaldalie60.

lectotypes of L. valeriana and L. mystle and found a — Lobotractus valerianal15102C06|Mexico:DE|1917

that their COI barcodes differ by 2.1% (14 bp). |. - tobotracus valerian Le095e08 Mexco-cuarero(l912.

Furthermore, the genomic tree of L. valeriana Fig. 52. eremaatts valeriana (blue), aa mysie (purple).

Specimens from across the range revealed their

partitioning into two clades, each clade with its type specimen: mysie from the US (Fig. 52 purple), and

valeriana from various localities in Mexico (Fig. 52 blue). Fst/Gmin statistics for these clades were

0.41/0.04, suggesting that they represent distinct species (Cong et al. 2019a; Zhang et al. 2020).

Therefore, we reinstate Lobotractus mysie (Dyar, 1904) as a species different from Lobotractus valeriana

(Plétz, 1881), and consequently exclude L. valeriana from the US fauna. Curiously, Burns (1996) and

(2001) reached similar conclusion as far as the US fauna is concerned, but likely for incorrect reasons.

Pyrgus centaureae dzekh Gorbunov, 2007 is a new subspecies for North America

The genomic tree of Pyrgus centaureae (Rambur, 1839) (type locality Sweden) taxa reveals that a

specimen collected by J. L. Harry west of Galbraith = ,Pyrgus centaureae|20081A01|Sweden|1979

Lake around Dalton Hwy mi. 274 in Alaska is not Toe Py rgus centaureae|20081A03|F |Russia:altal Kuray 1998

; : . : Pyrgus centaureae dzekh|17069B07|(USA:AK|1991

monophyletic with North American populations, but Be oa arty arts Pee USER Wat Ee ae

instead belongs to the Old World lineage (Fig. 53). [J] te eee ilboeocosjusa coat cs ons

Therefore, it is not Pyrgus centaureae freija (B. pyrgus andromedas|20081E04|Russia:Kolal 1993

Warren, 1924) (type locality Labrador), but Pyrgus Fig. 53. Pyrgus centaureae: Palearctic lineage (blue)

centaureae dzekh Gorbunov, 2007 (type locality and Nearctic lineage (magenta).

Russia: Chukotka). Hence, we add this subspecies to North American fauna. The tree reveals partitioning

of P. centaureae into two clades: mostly Palearctic lineage (Fig. 53 blue) and Nearctic lineage (Fig. 53

magenta). However, COI barcodes of specimens from different lineages (e.g. Sweden vs. USA: WV)

differ by only 0.15% (1 bp) and Fst/Gmin statistics of the two lineages are 0.21/0.08, suggesting rather

limited genetic differentiation and continuing gene exchange between them. On the basis of statistics from

this small sample of specimens we sequenced, we cannot yet support the two lineages as distinct species,

and the Nearctic assemblage of subspecies may be conceptualized as a semi-species instead.

Appia Evans, 1955 is a junior subjective synonym of Pompeius Evans, 1955

The monotypic genus Appia Evans, 1955 (type and the only species Appia appia Evans, 1955) originates

within Pompeius Evans, 1955 (type species Hesperia pompeius Latreille, [1824]) being sister to Pompeius

amblyspila (Mabille, 1897) and rendering Pompeius paraphyletic (Fig. 54). All of these species are

genetically close, e.g., COI barcodes of the type species of Appia and Pompeius differ by only 5% (33

bp). Therefore, to restore the monophyly, we treat Appia Evans, 1955 as a junior subjective synonym of

Pompeius Evans, 1955. Both taxa were proposed in the same work issued on the same date (Evans 1955),

and using ICZN Code Art. 24, we give priority to Pompeius, because this name was used more frequently

in the literature and the genus is not monotypic as Appia.

Wallengrenia Berg, 1897 is a subgenus of Polites Scudder, 1872

We sequenced all major phenotypically distinct taxa from the Hylephila Billberg, 1820 (type species

Papilio phyleus Drury, 1773) group of | Pompeius pompaius| 17196802] Costa Hiosleare

. . j ; Appia appia|18021B11|Brazil:RGS|1961

genera (Fig. 54, rooted with Hesperia Pompeius ST Gl ree aE Brien

ae : Hedone vibex praeceps|4942|US5A: tal Ze o,[20

Fabricius, 1793). Confirming our weaeae Sse 2002

previous assessment (Zhang et al.

2019c), we see that Polites Scudder,

1872 (type species Hesperia peckius :

W. Kirby, 1837) i tically close t tes peckius|4276|USA'IN Montgomery Co.[2

° If y> ) 1S gene 1Ca ye CLOSE lO ; ‘ i ites 5 aCe sh: en Warmer Corl 1996

. e

Wallengrenia Berg, 1897 (type - Polites baracoa|8170|USA, FLMiami-Dade’ Co,|2017

. : Wallengrenia ophites|17098E07|Martinique| 1989

species Hesperia premnas Wallengrenia vesuria]/10259|Jamaical2017

: uidenbe aurea SUS a ane COL2U1S, oi

Wallengren, 1860) and is _ not : "Hvlgphila phyleus|>174/USA'TX Hidalgo co.[2045

separated from it by a long internal Hesperia comms| 16108209 |Russia‘Tien-Shen|1991

branch (Fig. 54). Both branches that | Fig. 54. Genera: Pompeius (green and orange), Hedone (cyan), Limochores

are labeled "Polites" in Fig. 54 are (magenta) and Polites (blue and red) with subgenus Wallengrenia (red).

longer than the branch between them and therefore assigning a taxonomic rank to them seems more

appropriate than to the shorter branch between them. Moreover, COI barcodes of W. otho and P. peckius

differ by 5.2% (34bp) further confirming their close relationship. Therefore, we propose treating

Wallengrenia Berg, 1897 as a subgenus of Polites Scudder, 1872. The resulting classification of the

Hylephila Billberg, 1820 (type species Papilio phyleus Drury, 1773) group is marked on the tree rooted

with Hesperia comma (Linnaeus, 1758) (Fig. 54). Hylephila is sister to all others in the group. Then, there

are two major levels in the tree. First, the genus Polites that includes Wallengrenia originates at about the

same level as other three genera in the group: Pompeius Evans, 1955 (type species Hesperia pompeius

Latreille, [1824]), Hedone Scudder, 1872 (type species Hesperia brettus Boisduval & Le Conte, [1837], a

junior subjective synonym of Thymelicus vibex Geyer, 1832), and Limochores Scudder, 1872 (type

species Hesperia manataaqua Scudder, 1863, which is a junior subjective synonym of Hesperia origenes

Fabricius, 1793). This diversification event dates to about 15 Mya (Zhang et al. 2019d) and therefore

corresponds to genera. Second, Polites diversifies into 4 prominent lineages: Polites, Yvretta Hemming,

1935 Coa Grishin, 2019 and Wallengrenia Berg, 1897. This more recent diversification (~10 Mya)

corresponds to subgenera.

ACKNOWLEDGMENTS

We acknowledge Leina Song, Ping Chen and Ming Tang for excellent technical assistance. We are

grateful to David Grimaldi and Courtney Richenbacher (AMNH: American Museum of Natural History,

New York, NY, USA), Jonathan P. Pelham (BMUW: Burke Museum of Natural History and Culture,

Seattle, WA, USA), Vince Lee and the late Norm Penny (CAS: California Academy of Sciences, San

Francisco, CA, USA), Boris Kondratieff and Chuck Harp (CSUC: Colorado State University Collection,

Fort Collins, CO, USA), Crystal Maier and Rebekah Baquiran (FMNH: Field Museum of Natural History,

Chicago, FL, USA), Michael S. Engel and Jennifer Thomas (KUE: KU Biodiversity Institute & Natural

History Museum, Lawrence, KS, USA), Weiping Xie (LACM: Los Angeles County Museum of Natural

History, Los Angeles, CA, USA), Andrew D. Warren, Debbie Matthews-Lott and Keith R. Willmott

(MGCL: McGuire Center for Lepidoptera and Biodiversity, Gainesville, FL, USA), Matthias Nuss (MTD:

Museum fir Tierkunde, Dresden, Germany), Larry F. Gall (PMNH: Peabody Museum of Natural History,

Yale University, New Haven, CT, USA), Edward G. Riley, Karen Wright, and John Oswald (TAMU:

Texas A&M University Insect Collection, College Station, TX, USA), Alex Wild (TMMC: University of

Texas Biodiversity Center, Austin, TX, USA), Jeff Smith and Lynn Kimsey (UCDC: Bohart Museum of

Entomology, University of California, Davis, CA, USA), Robert K. Robbins, John M. Burns, and Brian

Harris (USNM: National Museum of Natural History, Smithsonian Institution, Washington, DC, USA)

and Wolfram Mey and Viola Richter (ZMHB: Museum fiir Naturkunde, Berlin, Germany) for granting

access to the collections under their care, sampling specimens, and stimulating discussions; to David H.

Ahrenholz, Rich Bailowitz, Maurizio Bollino, Ernst Brockmann, Christian Brévignon, Jim P. Brock, Bill

R. Dempwolf, Jason P. W. Hall, Jeremy J. Kuhn, James McDermott, Andrew F. E. Neild, James A. Scott,

and Mark Walker for specimens and leg samples, to Gerardo Lamas, Vladimir Lukhtanov and Kojiro

Shiraiwa for discussions, advice and helpful suggestions, to Jonathan P. Pelham and David M. Wright for

insightful discussions and critical review of the manuscript. Evi Buckner-Opler assisted by providing

emotional and logistic support and helped to collect specimens. We are indebted to California Department

of Fish and Game for collecting permit SC13645, Texas Parks and Wildlife Department (Natural

Resources Program Director David H. Riskind) for the research permit 08-02Rev, to U. S. National Park

Service for the research permits: Big Bend (Raymond Skiles) for BIBE-2004-SCI-0011 and Yellowstone

(Erik Oberg and Annie Carlson) for YELL-2017-SCI-7076 and to the National Environment & Planning

Agency of Jamaica for the permission to collect specimens. We acknowledge the Texas Advanced

Computing Center (TACC) at The University of Texas at Austin for providing HPC resources. The study

has been supported in part by grants from the National Institutes of Health GM127390 and the Welch

Foundation I-1505.

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