that a genus-group taxon should be monophyletic, i.e., a group of species that consists of all descendants

of their common ancestor. Thus, if a strongly supported by statistics phylogenetic tree reveals that a genus

is not monophyletic (i.e., its member species are distributed across several clades in the tree intermixed

with species from other genera, or the clade with this genus includes some species that are not currently

placed in this genus), some action needs to be taken to restore monophyly of the genus. The genus could

either be split into several genera, or species that are currently not included into it, but are in the same

clade with it, could be transferred into this genus. An important consideration is to follow the type species

of the genus, because it defines the genus. Only the clade with the type species could carry this genus

name. In addition to monophyly, attention should be paid to the prominence of the genus and consistency

with how other genera in this group of organisms are defined, see Taxonomic Appendix to Li et al. (2019)

for discussion. In brief, the genus should be a major phylogenetic group, i.e., it is best if the phylogenetic

tree branch leading to the last common ancestor of the genus is longer compared to other nearby branches,

and the genetic diversity within different genera should be comparable, so that genera define more or less

equivalent groups.

We obtained and analyzed whole genome shotgun sequences of all butterfly species recorded from

Canada and the United States (Pelham 2008; Pelham 2019). The manuscript describing this work is

available from bioRxiv (Zhang et al 2019c). Focusing on general evolutionary principles we can learn

from butterflies, the manuscript does not go into taxonomic details. However, accurate phylogenetic trees

we obtained from the genomic data reveal that some genera are not monophyletic, and some are either too

broad or too narrow in terms of genetic divergence. Here, we propose taxonomic rearrangements that are

supported by these phylogenetic trees. The following sections are in the standardized format. Taxonomic

act is the title of each section. Relevant genera, subgenera and their type species, are listed giving valid

names if the type species are synonyms. When the species are listed with their original genus name,

author names are given without parenthesis. Currently used genus name for these species is clear from the

context. Each section is illustrated by a small segment of a nuclear genomic tree with species that are

needed to support the conclusion. The trees showing all US and Canada species are available from the

bioRxiv preprint (Zhang et al. 2019c). Previous genus-species combinations (per Pelham 2019, version

revised 7 October 2019) are used in the figures. New combinations are given in the text. Species of major

focus are shown in red, other species in the genus of interest are shown in blue. Trees on gray background

include species not recorded from the US. The section ends with a conclusion and in many cases with a

list of species with revised genus-species names combinations. The sections are ordered by family and in

their taxonomic order.

Family Papilionidae Latreille, [1802]

Mimoides Brown, 1991 is a subgenus of Eurytides Hubner, [1821]

Previously placed in the genus Mimoides Brown, 1991 (type species Papilio ariarathes , Esper, 1788),

Papilio phaon Boisduval, 1836 is sister to

Eurytides marcellus (Cramer, 1777) among

butterflies of Canada and the US, and is

phylogenetically close to Eurytides philolaus

(Boisduval, 1836), suggesting that it should be

placed in the genus Eurytides Hubner, [1821]

(type species Eurytides iphitas Hubner,

[1821]) (Fig. 1). Additionally taking into

account that Mimoides species are close to

each other as evidenced by their morphology

(Tyler et al. 1994) and COI barcodes

(Ratnasingham & Hebert 2007), we propose to

treat Mimoides as a subgenus of Eurytides among its other subgenera: Protesilaus Swainson, [1832] (type

species Papilio protesilaus Linnaeus, 1758) and Neographium Mohn, 2002 (type species Papilio

philolaus Boisduval, 1836). Curiously, Papilio marcellus Cramer, 1777 is in the same clade with

Mimoides , but is in a different clade from Neographium , and therefore should be included in the subgenus

Mimoides despite the similarity in wing patterns to Eurytides ( Neographium) philolaus. Finally,

application of the genus Protographium Munroe (1961) (type species Papilio leosthenes E. Doubleday,

1846) to the New World is unwarranted, because genomic data show that the Australian endemic P.

leosthenes is sister to another Old World genus Graphium Scopoli, 1777 (type species Papilio sarpedon

Linnaeus, 1758) and is in a different clade from Eurytides (including Mimoides and Neographium) (Fig.

1). Eurytides versus Protographium is yet another case of striking wing pattern convergence in butterflies.

Fig. 2. Eurema, Abaeis, Sphaenogona, and Lucidi

Family Pieridae Swainson, 1820

Sphaenogona Butler, 1870 and Lucidia Lacordaire, 1833 are subgenera of Abaeis

Hiibner, [1819] and not of Eurema Hiibner, [1819], new placement

Previously junior subjective synonyms of Eurema Hiibner, [1819] (type species Papilio delia Cramer,

[1780], a junior subjective synonym of Pieris daira Godart, 1819), Sphaenogona Butler, 1870 (type

species Terias bogotana C. & R. Felder, 1861, which is treated as a subspecies of Terias mexicana

Boisduval, 1836) and Lucidia Lacordaire, 1833 (type species Papilio albula Cramer, 1775) are not

monophyletic with Eurema , but are instead in the same clade with Abaeis Hiibner, [1819] (type species

Papilio nicippe Cramer, 1779) (Fig.

2). This genomic tree shows notable

genetic divergence among Abaeis ,

Sphaenogona and Lucidia that is

only slightly less than the divergence

between Eurema and Pyrisitia But¬

ler, 1870 (type species Papilio pro-

terpia Fabricius, 1775) suggesting

that Sphaenogona and Lucidia are

not synonyms, but can be treated as subgenera of Abaeis. As a result, we use the following new or revised

combination for the US species: Abaeis (Sphaenogona) boisduvaliana (C. Felder & R. Felder, 1865),

Abaeis (Sphaenogona) mexicana (Boisduval, 1836), Abaeis (Sphaenogona) salome (C. Felder & R.

Felder, 1861), and Abaeis (Lucidia) albula (Cramer, 1775). Consequently, only a single US species

remains in Eurema , the type species of the genus: Eurema daira. Our proposed changes keep the number

of genera in this group at 3 ( Eurema , Abaeis , and Pyrisitia ), and simply rearrange species between these

genera. This rearrangement agrees with wing pattern characters on the dorsal side, making identification

of the genus in the US easier. Both Eurema and Pyrisitia lack darker expanded area near fore wing tornus

and their males possess darker scaling along the outer margin of hindwing, at least by the veins. Eurema

males can be distinguished by a long dark bar near forewing inner margin, which Pyrisitia species lack.

Males of Abaeis species either have a dark forewing tornus (forewing mostly orange with a dark cell spot

in the nominal subgenus and yellower, without the cell spot in the subgenus Sphaenogona ), or lack dark

scaling by the hindwing outer margin (USA only) and wings are mostly white with variable extent of dark

margins (subgenus Lucidia). Although it is tempting to unite all these medium-sized white-yellow-orange

butterflies in a single genus Eurema , their genetic divergence is very large (Fig. 2), and the group is

divided into two prominent clades (Eurema + Pyrisitia and Abaeis ), one of which splits into two more

(Eurema and Pyrisitia). Therefore, we keep the three-genus arrangement of the group. Moreover, Abaeis

as defined here is a broad and diverse genus. Comprehensive sequencing of the worldwide fauna of the

group is likely to substantiate further splits rather than lumps.

Family Lycaenidae [Leach], [1815]

Philotiella Mattoni, [1978] is a subgenus of Euphilotes Mattoni, [1978]

1993 (Thecla syllis Godman & Salvin, 1887), we resurrect from synonymy the genus Pendantus K.

Johnson & Kroenlein, 1993 (type species Thecla plusios Godman & Salvin 1887, currently treated as a

junior subjective synonym of Tmolus denarius Butler & H. Druce, 1872) and form new or revised

combinations: Pendantus guzanta , Pendantus thurman (Thompson & Robbins, 2016), Pendantus

denarius (A. Butler & H. Druce, 1872), and Pendantus perisus (H. Druce, 1907).

Family Nymphalidae Rafinesque, 1815

Neominois Scudder, 1875 is a subgenus of Oeneis Hiibner, [1819]

In agreement with the previous study (Kleckova et al. 2015), we find that Neominois Scudder, 1875 (type

species Satyrus ridingsii W. H. Edwards, 1865) originates within Oeneis Hlibner, [1819] (type species

Papilio norna Thunberg, 1791, represented by O. polixenes (Fabricius, 1775) in the US) as it is currently

defined, and is sister to the subgenus Oeneis.

COI barcode difference between ridingsii and

polixenes is only 6.2%. However, subgenus

Protoeneis Gorbunov, 2001 (type species

Chionobas nanna Menetries, 1858, represented

by Oeneis uhleri (Reakirt, 1866) in the US

fauna) that is placed in the genus Oeneis , is a

sister to the clade consisting of Neominois and subgenus Oeneis (Fig. 6). The entire group of 3 taxa is

compact (Fig. 6), prominent, and genetic divergence within it agrees with the expected divergence within

a genus. Therefore, we treat Neominois as a subgenus of Oeneis.

Agraulis Boisduval & Le Conte, [1835] is a subgenus of Dione Htibner, [1819]

Monotypic genus Agraulis Boisduval & Le Conte, [1835] (type species Papilio vanillae Linnaeus, 1758)

is a close sister to Dione Htibner, [1819] (type species Papilio juno Cramer, 1779) (Fig. 7). COI barcode

difference between A. vanillae and Dione

moneta Htibner, [1825] is 7.9%. Time-

calibrated nuclear genomic tree shows that

genetic divergence (Fig. 7) between Agraulis

and Dione is nearly the same as the divergence

among species of Boloria Moore, 1900 (type

species Papilio pales [Denis & Schiffermiiller],

1775) and smaller than the divergence between Heliconius Kluk, 1780 (type species Papilio charithonia

Linnaeus, 1767) and Eueides Hiibner, 1816 (type species Nereis dianasa Hiibner, [1806]) (Fig. 7).

Therefore, we treat Agraulis as a subgenus of Dione.

Family Hesperiidae Latreille, 1809

Urbanus alva Evans, 1952, new status

Described as a subspecies of Urbanus viterboana (Ehrmann, 1907) by Evans (1952), alva (the holotype,

male, from Mexico: Veracruz, Atoyac, examined by NVG) was placed in synonymy with Urbanus belli

(Hayward, 1935) (the holotype, female, from Argentina: Salta, photographs examined) by Steinhauser

(1981), who was unable to see the belli holotype. Our genomic analysis of U. belli from Argentina

indicates that it is not conspecific with belli-like specimens from Mexico or anywhere else in North

America due to genetic divergence between them. Evans noted shorter hindwing tails in alva compared to

belli, and this character holds true comparing the holotypes of these taxa and additional belli specimens

from Argentina we sequenced, including males. For these reasons we resurrect alva from synonymy with

Fig. 7. Agraulis and Dione.

belli and treat it as a distinct species Urbanus alva, new status. Consequently, we exclude Urbanus belli

(Hayward, 1935) from the North American fauna.

isisrssssissia,,,.

Fig. 8. Erynnis and Gesta.

Erynnides Burns, 1964 is a subgenus of Gesta Evans, 1953

and not of Erynnis Schrank, 1801, new placement

Gesta Evans, 1953 (type species Thanaos gesta Herrich-Schaffer, 1863) is a sister to subgenus Erynnides

Burns, 1964 (type species Nisoniades propertius Scudder & Burgess, 1870), with the exclusion of

subgenus Erynnis Schrank, 1801

(type species Papilio tages Linnaeus,

1758, represented by Erynnis brizo

(Boisduval & Le Conte, [1837]) and

Erynnis icelus (Scudder & Burgess,

1870) in the US), thus rendering

Erynnis paraphyletic (Fig. 8). There

are three possible solutions. First

(splitting), treat all three 0 Erynnis ,

Gesta , and Erynnides) as valid genera. However, genetic divergence between Gesta , and Erynnides is

moderate (about 8% in the COI barcode), and no prominent tree branches separate the two taxa. Their

genitalia are also quite similar. Therefore, these two taxa are best viewed as subgenera. Second (lumping),

treat all three as subgenera of Erynnis , thus eliminating genus-species combinations involving Gesta.

However, genetic divergence between Erynnis (sensu stricto) and Gesta + Erynnides is prominent (Fig.

8), comparable to that between other Erynnini Brues & F. Carpenter, 1932 genera. While the lumping

solution is more compatible with how these taxa were viewed historically, it is not consistent with how

other members of Erynnini are partitioned into genera. Third (middle ground), is a two-genus solution,

i.e., to transfer Erynnides from Erynnis to Gesta. Phylogenetic trees show the two prominent clades

corresponding to these two genera, and the clade leading to their common ancestor is shorter and thus less

prominent (Fig. 8). Genetic divergence between these two genera is the same magnitude as between other

sister genera of Erynnini. This divergence is equally profound in nuclear (autosomes and Z chromosome)

and mitochondrial genomes. Therefore, we prefer this two-genus solution. As a result, all species formerly

placed in the subgenus Erynnides of Erynnis would change their genus name to Gesta. This action results

in many name changes, but highlights deep genetic divergence between mostly Old Worth Erynnis and

exclusively New World Gesta and thus seems to be more biologically meaningful. Although the switch of

names is bothersome in short run, it may be beneficial long term.

Copaeodes Speyer, 1877 is a subgenus of Oarisma Scudder, 1872

Previously placed in the genus Oarisma Scudder, 1872 (type species Hesperia powesheik Parker, 1870),

Thymelicus edwardsii W. Barnes, 1897 is

not monophyletic with it, and is a close sister

to the two US species from the genus Copa¬

eodes Speyer, 1877, including its type spe¬

cies Heteropterus procris W. H. Edwards,

1871, a junior subjective synonym of Ancy-

loxipha [sic!] aurantiaca Hewitson, 1868

(Fig. 9). Investigation of other Oarisma and

Copaeodes species reveals that they are

close to each other and difficult to partition

between the two genera (Fig. 9). Only the

close cluster of 3 species O. poweshiek (Parker, 1870), O. garita (Reakirt, 1866), and O. era Dyar, 1927

constitute Oarisma sensu stricto. Others fall in a different and broad clade, which in addition to

Copaeodes includes other species previously placed in Oarisma , e.g., O. edwardsii (W. Barnes, 1897) and

O. nanus (Herrich-Schaffer, 1865), and we treat this clade as a subgenus Copaeodes within a broader-

defined Oarisma.

Saliana Evans, 1955 is a junior subjective synonym of Calpodes Hiibner, [1819]

Monotypic genus Calpodes Hiibner, [1819] (type species Papilio ethlius Stoll, 1782) is genetically close

to Saliana Evans, 1955 (type species Papilio salius Cramer, 1775) (Fig. 10) and their type species differ

by only about 6.5% in COI barcodes. Moreover, Calpodes

+ Saliana clade experienced rapid radiation over a short

period of time and, as a result, the tree looks more like a

comb than a bi-branching structure (Fig. 10). It is not clear

whether Saliana is monophyletic: 64% bootstrap in the

nuclear genome tree suggests that it is not, and that Saliana

fusta Evans, 1955 is sister to the rest of Saliana +

Calpodes. Due to rapid radiation, this genus is difficult to

partition into meaningful subgenera, because no clear clusters of species are apparent in the tree (Fig. 10).

For all these reasons, we propose that Saliana is a junior subjective synonym of Calpodes , new status.

Interestingly, C. ethlius diverged strongly in wing shapes and patterns from all other members of the

genus (although their male genitalia are similar) and this new synonymy is unexpected. Therefore, we

tested the results using multiple specimens of C. ethlius (7 shown in Fig. 10) and they all cluster together,

within former Saliana. The newly expanded Calpodes is a strongly supported (bootstrap 100%) and

prominent (long tree branch is leading to it) genus.

Nastra perigenes (Godman, 1900), new combination

Previously placed in the genus Vidius Evans,

1955 (type species Narga vidius Mabille, 1891),

Mastor perigenes Godman, 1900 is not mono¬

phyletic with the type species of Vidius , and

instead forms a clade with Nastra Evans, 1955

(type species Hesperia Iherminier Latreille,

[1824]) (Fig. 11) and therefore is transferred to

this genus to form a new combination Nastra perigenes.

Amblyteria Grishin, new subgenus

http://zoobank.org/ClF98F8D-C366-4065-9317-EA039D15CB22

Type species. Goniloba exoteria Herrich-Schaffer, 1869.

Definition. A prominent clade within Amblyscirtes Scudder, 1872 (type species Hesperia vialis W. H.

Edwards, 1862) without a name (Fig. 11, 12). Keys to N.2.2, 6, 8, or 22 in Evans (1955). Phenotypically

diverse lineage of species that differs from its relatives by the following combination of characters: males

either with stigma long, narrow and rather straight, hindwing below with many small white spots in some

species, and size small (forewing length mostly < 15mm), or if larger, then forewing pale spot in cell Mi-

M 2 strongly offset towards outer margin; or with brands at the base of vein CuA 2 , and brands either very

conspicuous on bronze-colored wings, or wings black, head orange and fringes not orange but pale. Due

to this pronounced phenotypic variation, the subgenus is best defined by DNA characters. A combination

Fig. 11. Nastra and no Vidius.

. 1 1 . I+1 -’ 1

gJS/5S^ n 7 co|2Qis

aflf cat

lEJimisg:

Fig. 10. Calpodes and Saliana.

of the following base pairs in the standard DNA COI barcode region (658 bp) is diagnostic: T548C,

A550T (these two characters are synapomorphic), 343(not G as in Stomyles ), T346T(not C as in

Stomyles ), T553T(not A as in the nominal subgenus), and A637A(not T as in Stomyles).

Etymology. The name is a feminine noun in the nominative singular, a fusion of the type species name

epithets: Arafry[scirtes] + [exo ]teria.

Species included. Amblyscirtes elissa Godman, [1900], Pamphila oslari Skinner, 1899, Amblyscirtes

brocki Freeman, 1992, Goniloba exoteria Herrich-Schaffer, 1869, and Pamphila phylace Edwards, 1878.

Parent taxon. Genus Amblyscirtes Scudder, 1872.

Assignment of species to subgenera. With the

description of Amblyteria subgen. n., we consider

that the genus Amblyscirtes consists of 4 subgenera

(Fig. 12). Subgenus Stomyles Scudder, 1872

(Eastern US clade) consists of A. Carolina (Skinner,

1892), A. reversa F. Jones, 1926, A. aesculapius

(Fabricius, 1793), and A. hegon (Scudder, 1863).

Subgenus Mastor Godman, [1900] (Southern clade)

consists of A. fimbriata (Plotz, 1882) (the only USA

species), A. anubis (Godman, 1900), A. novimmaculatus A. Warren, 1998, A. raphaeli H. Freeman, 1973,

A. patriciae (E. Bell, 1959), and A. folia Godman, 1900. Names of type species are underlined. Other

Amblyscirtes species (Mielke 2005; Pelham 2008) not mentioned here belong to the nominal subgenus.

Comment. A surprising result is that two very similar-looking species A. (Amblyteria) phylace and A.

(Mastor) fimbriata are not each other's closest relatives and belong to different subgenera (Fig. 12).

Troyus fantasos (Cramer, 1780), Troyus onaca (Evans, 1955), Troyus aurelius (Plotz,

1882), Troyus marcus (Fabricius, 1787), Troyus diversa (Herrich-Schaffer, 1869), and

Troyus drova (Evans, 1955), new combinations

Previously placed in the genus Vettius Godman, 1901 (type species Papilio phyllus Cramer, 1777),

Papilio fantasos Cramer, 1780 is not monophy-

letic with Vettius type species, and is a close

relative of the monotypic genus Troyus A. Warren

& Turland, 2012 (type and the only included

species Troyus turneri A. Warren & Turland,

2012). Their sister genus is Monca Evans, 1955

(type species Cobalus telata Herrich-Schaffer,

1869) (Fig. 13). Therefore, we establish a new

combination Troyus fantasos. Due to genetic and

morphological similarities, we additionally propose the following new combinations: Troyus onaca

(Evans, 1955), Troyus aurelius (Plotz, 1882), Troyus marcus (Fabricius, 1787), Troyus diversa (Herrich-

Schaffer, 1869), and Troyus drova (Evans, 1955). All these species were in Vettius before.

Hedone Scudder, 1872 is a valid genus

We find that Polites Scudder, 1872 (type species Hesperia peckius W. Kirby, 1837) is paraphyletic with

respect to Wallengrenia Berg, 1897 (type species Hesperia premnas Wallengren, 1860). Genetic

divergence within Polites that includes Wallengrenia is too large compared to how most Hesperiidae

genera are defined. Therefore, instead of including Wallengrenia into Polites , it makes sense to restore

‘J 35# ilia

llllifpSl:

life £ llf ~

Fig. 12. Amblyscirtes.

monophyly of these genera by splitting

Polites into genera with divergence consistent

with that of most other Hesperiidae genera.

Previously a junior subjective synonym of

Polites Scudder, 1872 (type species Hesperia

peckius W. Kirby, 1837), Hedone Scudder,

1872 (type species Hesperia brettus

Boisduval & Le Conte, [1837], a junior

subjective synonym of Thymelicus vibex

Geyer, 1832) forms a clade sister to the rest of Polites + Wallengrenia (Fig. 14). Therefore, Hedone is a

valid genus, and Hedone vibex (Geyer, 1832) is a revised combination. Due to morphological similarities,

we additionally propose the following new combinations: Hedone bittiae (Lindsey, 1925), Hedone

vibicoides (de Jong, 1983), and Hedone dictynna (Godman & Salvin, 1896).

Limochores Scudder, 1872 is a valid genus

Sister to the rest of Polites + Wallengrenia excluding Hedone , Limochores Scudder, 1872 (type species

Hesperia manataaqua Scudder, 1863, which is a junior subjective synonym of Hesperia origenes

Fabricius, 1793) was treated as a junior subjective synonym of Polites (Fig. 14). For the reasons given

above for Hedone , we propose the following new or revised combinations: Limochores origenes

(Fabricius, 1793), Limochores mystic (W. H. Edwards, 1863), Limochores sonora (Scudder, 1872),

Limochores puxillius (Mabille, 1891), and Limochores pupillus (Plotz, 1882). Interestingly, two difficult-

to-distinguish species that frequently fly together at the same location, L. origenes and Polites

themistocles (Latreille, [1824]), ended up in different genera. Even though their genitalia are similar, they

belong to distant from each other clades in the tree (Fig. 14): P. themistocles is closely related to P.

peckius , the type genus of Polites , while L. origenes is closer to L. mystic.

Coa Grishin, new subgenus

http://zoobank.org/CD8143FC-839D-408E-A114-3FFEEA7AE349

Type species. Hesperia baracoa Lucas, 1857.

Definition. A sister to subgenus Yvretta Hemming, 1935 (type species Pamphila citrus Mabille, 1889,

which is treated as a junior subjective synonym Hesperia subreticulata Plotz, 1883), Hesperia baracoa

Lucas, 1857 prominently stands out from other Polites (Fig. 14). Therefore, this lineage is given a

subgenus status and a name. This new subgenus keys to M.13.4 in Evans (1955). Distinguished from its

relatives within Polites by the combination of the following characters: presence of apiculus (longer than

1 segment); diagnostic shape of stigma: rather short, relatively straight and narrower than in other species

with defined apiculus; the lack of spot before the end of discal cell on plain gray-brown ventral hindwing

without dark spots (but sometimes with a row of pale discal spots) combined with orange area by the

fore wing costa below, stemming from the wing base and reaching apical spots.

Etymology. The name is a feminine noun in the nominative singular, the ending of the type species name.

Species included. Only the type species.

Parent taxon. Genus Polites Scudder, 1872.

Assignment of species to subgenera. With the description of Coa subgen. n., we consider that Polites

consists of 3 subgenera (Fig. 14). Subgenus Yvretta consists of P. subreticulata (Plotz, 1883), P. cams

(W. H. Edwards, 1883), and P. rhesus (W. H. Edwards, 1878). Other Polites species (Mielke 2005;

Pelham 2008) not mentioned in this work belong to the nominal subgenus. Interestingly, Wallengrenia

Berg, 1897 (type species Hesperia premnas Wallengren, 1860) is not prominently distinct genetically

from Polites (green branch in Fig. 14) and may therefore be included in Polites as the 4th subgenus,

depending on researcher's taste.

Problema Skinner & R. Williams, 1924 is a subgenus of Atrytone Scudder, 1872

Currently monotypic genus Atrytone Scudder, 1872 (type species Hesperia iowa Scudder, 1868) is sister

to Problema Skinner & R. Williams, 1924

(type genus Pamphila byssus W. H.

Edwards, 1880), a genus of two species,

including also Hesperia bulenta Boisduval &

Le Conte, [1837]. All three species are close in their genomic sequences (Fig. 15). Their genetic

divergence is more consistent with them being congeners. Therefore, we treat Problema as a subgenus of

Atrytone to form new or revised combinations Atrytone byssus and Atrytone bulenta.

Oligoria percosius (Godman, 1900), Oligoria rindgei (H. Freeman, 1969), Oligoria

lucifer (Hiibner, [1831]), and Oligoria mustea (H. Freeman, 1979), new combinations

Previously placed in the genus Decinea Evans, 1955 (type species Hesperia decinea Hewitson, 1876),

Cobalus percosius Godman, 1900 is not monophyletic with Decinea type species, which is in the same

clade with Buzyges Godman, 1900. Instead,

percosius is a very close sister of the mono¬

typic genus Oligoria Scudder, 1872 (type

species Hesperia maculata W. H. Edwards,

1865) (Fig. 16), which is in the same clade

with Xeniades Godman, 1900. COI barcode

difference between maculata and percosius

is only 4%. Therefore, we establish a new

combination: Oligoria percosius. Due to

morphological and genetic DNA similarities,

we additionally propose the following new

combinations: Oligoria rindgei (H. Freeman, 1969), Oligoria lucifer (Hiibner, [1831]), and Oligoria

mustea (H. Freeman, 1979) (Fig. 16). All these species were previously placed in Decinea.

ACKNOWLEDGMENTS

We acknowledge Leina Song and Ping Chen for excellent technical assistance. We are grateful to David

Grimaldi and Courtney Richenbacher (AMNH: American Museum of Natural History, New York, NY,

USA), Jason Weintraub (ANSP: Academy of Natural Sciences of Drexel University, Philadelphia, PA,

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 (CSUC: Colorado State University Collection, Fort Collins, CO, USA), Crystal Maier

and Rebekah Baquiran (FMNH: Field Museum of Natural History, Chicago, FL, USA), Weiping Xie

(LACM: Los Angeles County Museum of Natural History, Los Angeles, CA, USA), Andrew D. Warren

and Debbie Matthews-Lott (MGCL: McGuire Center for Lepidoptera and Biodiversity, Gainesville, FL,

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) for granting access to the

collections under their care and for stimulating discussions; to Jim P. Brock, Jack S Carter, Bill R.

Dempwolf, James McDermott, the late Edward C. Knudson (TLS: Texas Lepidoptera Survey, specimens

now at MGCL), Harry Pavulaan, James A. Scott, John A. Shuey, and Mark Walker for specimens and leg

samples, to Jonathan P. Pelham for insightful discussions and critical review of the manuscript. Greg

Kareofelas (Davis, CA) and Matthew Garhart (Grand Junction, CO) collected selected species and placed

them in RNAlater for molecular analysis. Boris Kondratieff, Chuck Harp, and James Scott curated the

butterfly collection at the C. P. Gillette Museum of Arthropod Diversity, Colorado State University,

which facilitated the accurate sampling of remaining species needed to complete the analysis. Evi

Buckner-Opler assisted by providing emotional and logistic support and helped to collect specimens. We

are indebted to 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 1-1505.

LITERATURE CITED

Steinhauser, S. R. 1981. Revision of the proteus group of the genus Urbanus Hubner (Le

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