Volume 8 Number

29 March 2019

W The Taxonomic Report

Expanded phenotypic diagnoses for 24 recently named

new taxa of Hesperiidae (Lepidoptera)

Nick V. Grishin 1 ’ 2

1 Howard Hughes Medical Institute and departments of Biophysics and Biochemistry, University of Texas Southwestern

Medical Center, 5323 Harry Hines Blvd., Dallas, TX, USA 75390-9050; grishin@chop.swmed.edu

ABSTRACT. Expanded diagnoses by phenotypic characters for the 24 new taxa named in the article "Genomes of

skipper butterflies reveal extensive convergence of wing patterns" by Li, W., Cong, Q., Shen, J., Zhang, J., Hallwachs, W.,

Janzen, D.H. and Grishin, N.V., 2019 and published in the Proceedings of the National Academy of Sciences of the United

States of America on March 15, 2019 are provided and illustrated. More detailed diagnoses will help identifying these

phylogenetic groups by their wing patterns and shapes and other morphological characters including the structures of antennae

and genitalia using this single publication, instead of obtaining the sequences or inspecting additional works referenced in the

original diagnoses for brevity.

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

ZooBank registration: http://zoobank.org/D0BA0046-6ACA-47B4-9E00-B0FD9QA99555

INTRODUCTION

Recently, we published genomics-based analysis of Hesperiidae that suggested 24 new taxa: 6

tribes, 6 subtribes, 9 genera, and 3 subgenera (Li et al ., 2019). We and others have argued that the best

way to define a higher level taxon (above species level) is by a clade in a phylogenetic tree that has strong

statistical support (Talavera et al ., 2012; Li et al ., 2019). While the exact criteria for matching the ranks

(e.g. genus vs. subgenus) to clades can be debated, if the taxa are desired to be monophyletic, the only

way to increase the probability of that would be by using reliable phylogenetic trees. While the definition

of a taxon by a branch in a tree is aimed at having it monophyletic to the best of our knowledge, such

definition does not agree with how the taxon has to be defined according to the ICZN Code (ICZN, 1999).

The Code requires defining "characters," i.e. some features of an organism that enable us to diagnose it as

belonging to a particular taxon. A list of characters should be provided as a "diagnosis" when a new name

is proposed. If an animal possesses these characters (individually or in combination), it belongs to the

taxon. While such definitions are reasonable and practically useful, they have obvious limitations. Most

importantly, it is challenging to extrapolate to yet undiscovered taxa. Thus some characters shared by the

currently know taxa may not hold for a new taxon to be discovered, which nevertheless belongs to the

group by the criterion of monophyly. The challenges are more severe in animals that experience high

frequency of phenotypic convergence, for instance Hesperiidae. To protect the name itself from the

problems with reporting characters that do not actually hold for the entire group, ICZN does not require

the characters to be meaningful, just "purported to differentiate" is sufficient. Thus, in principle, any

statement of any characters would be enough, even if they are wrong. While such an approach does not

sound appealing, it is not easy to suggest a better alternative.

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Nevertheless, researchers strive to find characters that are most meaningful given the current

knowledge. The best character would be a conserved synapomorphy, i.e. some feature that originated in

the common ancestor of the entire group before that group has diversified, and stayed in all member of the

group (i.e. "conserved" in the group). Thus, only the group that has this character and no animal outside

the group possesses it. An example would be osmeterium in Papilionidae (Cong et al ., 2015). However, it

is challenging to find such characters, and they may not even exist in some groups due to rapid evolution

reflected in character losses and gains. To address the problem of possible phenotypic convergence and

rapid divergence, and keeping in mind that the best way to define a higher-level taxon is from a

phylogenetic tree, we can look for possible synapomorphies in DNA that was used to construct the tree.

Such DNA-based characters are likely to be more reliable as diagnoses than phenotypic characters. An

approach to predict conserved synapomorphies in genomic sequences was described in the SI Appendix to

Li et al. (2019) and was used to find such characters for the new taxa proposed in that publication. These

potential synapomorphies were listed as a part of diagnoses in Tables 1 and 2 (Li et al ., 2019), and actual

sequences from the reference genome with positions highlighted (to prevent mishaps with position

numbers) were given in the SI Appendix.

While the DNA characters are expected to be more reliable in diagnosing the taxon, they are not

easy to use, because they require sequencing that is not available to everyone. Therefore, in addition to

DNA-based diagnoses, morphology-based diagnoses were also provided for each taxon in Tables 1 and 2

(Li et al ., 2019). Due to the need to fit the article into 6 pages, these diagnoses, while far from being

random, were brief and mostly referred to published statements in literature (Williams and Bell, 1934;

Evans, 1937; Evans, 1949; Evans, 1951; Evans, 1952; Evans, 1953; Evans, 1955; Bums, 1996). The

majority of diagnoses referenced the comprehensive Evans volumes. In Evans, diagnostic characters were

given as identification keys (he described dozens of new taxa by means of these keys), which may be a

better way of presentation, because a key allows comparisons of the alternatives. However, these keys

may not be straightforward to use, and Evans books are not readily accessible to everyone, similar to

sequencing. Here, I use the opportunity to elaborate on the diagnoses and rephrase the Evans keys. These

morphological characters (while not being original, but discovered by Evans and others) are put together

in this article that would be easier to use than going through the Evans volumes. In addition, many

essential characters are illustrated here, which was not possible in the original article due to space

constraints. The main value of this work is educational and I hope that this article, while not very original,

is nevertheless useful in the studies of Hesperiidae.

EXPANDED DIAGNOSES OF THE 24 TAXA

Here are the 24 standardized sections, each giving the name of the taxon; its type genus or species,

ZooBank registration, diagnosis that explains how the taxon was defined in the original diagnosis and

expands it to rephrase characters given in the referenced publication, all members placed in the taxon in

the original description (genera or species), and the parent taxon of the next rank. Species names are given

with their original genus in its original spelling ([sic] indicates spelling errors). Collating all this

information from the main text, tables and SI Appendix of the original publication (Li et al ., 2019) and

rephrasing characters from the publications referenced in the original diagnoses (Williams and Bell, 1934;

Evans, 1937; Evans, 1949; Evans, 1951; Evans, 1952; Evans, 1953; Evans, 1955; Burns, 1996) makes this

information more accessible and usable.

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Tribe Entheini Grishin, 2019

Type genus: Entheus Hubner, [1819].

ZooBank registration: 303C1FD0-07CB-4919-900E-EA3D6347E5DD

Diagnosis: The tribe was defined as a clade from about 40-45 Mya

that contains the genus Entheus , and putatively synapomorphic

DNA characters for this clade were given (Li et al., 2019).

Phenotypically, species in this tribe would key to B.3a in Evans

(1952), excluding B.9. Diagnosed by unusual palpi (Fig. 1): 3rd

segment divergent, stout, spatulate and set on the outer edge of the

2nd segment; and regular-shaped, not produced at vein M3

hindwing. The hindwing character needs to be added to avoid

inclusion of Phareas Westwood, 1852 (hindwing produced at vein

M3), which apparently converged to this unusual shape of palpi,

but is not monophyletic with Entheini as revealed by genomic trees

(Li et al., 2019).

Genera included: Drephalys E. Watson, 1893, Udranomia A. Butler, 1870, Phanus Hubner, [1819],

Hyalothyrus Mabille, 1878, Entheus Hubner, [1819], Augiades Hubner, [1819], and Tarsoctenus E.

Watson, 1893.

Parent Taxon: Subfamily Eudaminae Mabille, 1877.

Subtribe Loboclina Grishin, 2019

Type genus: Lobocla Moore, 1884.

ZooBank registration: C606FC35-323D-4E55-AF5A-A86C6366BAFA

Diagnosis: The subtribe was defined as a clade from about 30 Mya

that contains the genus Lobocla , and putatively synapomorphic

DNA characters for this clade were given (Li et al., 2019).

Phenotypically, species in this subtribe would key to B.4 in Evans

(1949) or C.5, C.lOa, C.15.2 or C.18 in Evans (1953). Genera in

this tribe are unified by the long forewing cell, at least 2/3 of costa,

lobed or tailed hindwing, short, non-divergent palpi with the 2nd

segment touching the face and the 3rd not protruding beyond the

2nd (except Zestusa). Most genera could be diagnosed by single

character and possible synapomorphy: broadly arcuate antennal

clubs (Fig. 2). Additionally, hyaline spot in forewing cell R2-R3

present even in species with hooked antennae. In species with hooked antennae that lack the spot,

genitalic valvae asymmetrical and very broad. Typically (except Aguna ), valvae rather broad, somewhat

rectangular, with harpe narrow, long upturned, hook-like, leaving a deep space between harpe and

ampulla, uncus divided.

Genera included: Aguna R. Williams, 1927, Zeutus Grishin, 2019, Lobocla Moore, 1884, Lobotractus

Grishin, 2019, Codatractus Lindsey, 1921, Zestusa Lindsey, 1925, Ridens Evans, 1952, and Venada

Evans, 1952.

Fig. 1. Entheini. Head of Entheus aureolus

Austin, O. Mielke & Steinhauser, 1997

Brazil: AM, with divergent 3 rd segment of palpi.

Parent Taxon: Tribe Eudamini Mabille, 1877.

Subtribe Cephisina Grishin, 2019

Type genus: Cephise Evans, 1952.

Diagnosis: The subtribe was defined as a clade from about 30 Mya

that contains the genus Cephise , and putatively synapomorphic

DNA characters for this clade were given (Li et al., 2019).

Phenotypically, species in this subtribe would be diagnosed by

"genitalia and palpi as described by Burns (1996: 182-183) for

Cephise " (Li et al., 2019). Note that this statement refers to the

characters of genitalia and palpi (they are "as described ... for

Cephise"), not to the genus Cephise. Thus, the referenced

publication could have been describing the characters of any other

genus. Not the genus is relevant here, but the characters

themselves, some of which are the following: uniquely short,

truncate uncus with a couple of teeth at each distal comer, harpe extending distad, separated from the

valva by a U-shaped notch; the first segment of palpi below the eyes with a shelf-like projection of scales

that looks like a triangle below the eyes (Fig. 3), more developed in males than in females.

Genera included: Cephise Evans, 1952.

Parent Taxon: Tribe Eudamini Mabille, 1877.

Subtribe Telemiadina Grishin,

Type genus: Telemiades Hubner, [1819].

ZooBank registration: 4AE0E59C-8B92-4C84-8651-E7A1C45C93C1

Diagnosis: The subtribe was defined as a clade from about 30 Mya

that contains the genus Telemiades , and putatively synapomorphic

DNA characters for this clade were given (Li et al., 2019).

Phenotypically, species in this subtribe would key to B.2, C.3, C.7a

(exclude C.7.6b), E.6a, or E.9 in Evans (1952, 1953). Includes a

phenotypically diverse array of Hesperiidae that were not brought

together before this study and do not possess an obvious

phenotypic synapomorphy. Many species included here have a

prominent tuft of long scales from a polished area near the base and

inner margin of forewing below, and correspondingly widened

costal area of hindwing to cover the tuft (Fig. 4). No such tufts on hindwing. If forewing tufts missing

then forewing apex not truncate, recurrent vein in the discal cell on forewing present and ends at the

origin of vein M3 (not before it), no spot in cell R2-R3, antennae angled (bent beyond ticker part of club),

discal cell about 2/3 of costa length. If discal cell longer, about 3/4 of costa, then antennae hooked, short,

not reaching the origin of vein CuAi, and no hyaline spots around mid costa or in cell CuA2-lA+2A.

Genera included: Ectomis Mabille, 1878, Telemiades Hubner, [1819], and Polygonus Hubner, [1825].

Parent Taxon: Tribe Eudamini Mabille, 1877.

2019

Fig. 4. Telemiadina. Wing bases of Ectomis

cythna cythna (Hewitson, 1878) S, French

Guiana, below, showing the tuft of scales and

expanded costal area of hindwing.

Fig. 3. Cephisina. Head of Cephise cephise

(Herrich-Schaffer, 1869) S, ventral view

showing lateral projections of scales from palpi

near eyes (by antennae).

Tribe Oileidini Grishin, 2019

Type genus: Oileides Hiibner, [1825],

ZooBank registration: CF9C3D29-523A-4D17-B140-9A69CFA98731

Diagnosis: The tribe was defined as a clade from about 40-45 Mya

that contains the genus Oileides , and putatively synapomorphic

DNA characters for this clade were given (Li et al., 2019).

Phenotypically, species in this tribe would key to C.6, D.9.2a,

D. 9.4, E.2, E.4, E.5b, or E.10 in Evans (1952, 1953). A possible

synapomorphy of the group: tufts of longer scales placed in a

groove at the base of hindwing near anal fold. The tufts present

either on dorsal or ventral side (Fig. 5), but not both. If tufts below,

then antennae bent not beyond the thickest part of the club, the

club more slender and uncus divided.

Genera included: Oileides Hiibner, [1825], Typhedanus A. Butler, 1870, Oechydrus E. Watson, 1893,

Cogia A. Butler, 1870, Nerula Mabille, 1888, and Marela Mabille, 1903.

Parent Taxon: Subfamily Eudaminae Mabille, 1877.

Subtribe Typhedanina Grishin, 2019

Type genus: Typhedanus Butler, 1870.

ZooBank registration: B4D56F93-67F9-476F-B69C-133D98BFBD58

Diagnosis: The subtribe was defined as a clade from about 30 Mya

that contains the genus Typhedanus , and putatively synapomorphic

DNA characters for this clade were given (Li et al ., 2019).

Phenotypically, species in this subtribe would key to C.6, E.2, E.4,

E. 5b, or E.10 in Evans (1952, 1953). Diagnosed by tufts of longer

scales placed in a groove at the base of hindwing near anal fold, on

dorsal side (Fig. 6), no tufts on ventral side.

Genera included: Typhedanus A. Butler, 1870, Oechydrus E.

Watson, 1893, Cogia A. Butler, 1870, Nerula Mabille, 1888, and

Marela Mabille, 1903.

Parent Taxon: Tribe Oileidini Grishin, 2019.

Tribe Netrocorynini Grishin, 2019

Type genus: Netrocoryne C. & R. Felder, [1867].

ZooBank registration: DE61F048-02CF-4F8E-9392-D18A4618BABD

Diagnosis: The tribe was defined as a clade from about 40-45 Mya that contains the genus Netrocoryne ,

and putatively synapomorphic DNA characters for this clade were given (Li et al ., 2019). Phenotypically,

species in this tribe would key to B.2, C.l, or C.15 in Evans (1949). Unexpected assemblage of genera

without obvious synapomorphies (Fig. 72). Previously placed in tribe Tagiadini Mabille, 1878, but can be

distinguished from them by the following somewhat convoluted combination of choices. If palpi erect,

then mid and hind tibiae without spines and antennae not less

than half of costa length in males, forewing vein M3 originates

midway between veins CuAi and Mi (not M2). If palpi porrect,

then antennal club not flattened or twisted, apiculus tapered to a

point (not blunt), palpi long, sometimes longer than head, and if

club angled or hooked towards apiculus, then fore wing cell not

sorter than outer margin, forewing produced and truncate at

apex, and hindwing produced at the end of vein 3; if the club

arcuate or obtuse from the thickest part, then apiculus short,

nudum with less than 17 segments, and hindwing costa

produced at apex and longer than outer margin.

Genera included: Netrocoryne C. Felder & R. Felder, [1867],

Chaetocneme Felder 1860, and Exometoeca Meyrick, 1888.

Parent Taxon: Subfamily Tagiadinae Mabille, 1878.

Tribe Jerini Grishin, 2019

Type genus: Jera Lindsey, 1925.

ZooBank registration: AF3B5CEA-880A-4CB2-AF40-E6D87C39C040

Diagnosis: The tribe was defined as a clade from about 40-45 Mya

that contains the genus Jera , and putatively synapomorphic DNA

characters for this clade were given (Li et al. , 2019).

Phenotypically, species in this tribe would key to E.3 in Evans

(1953) and can be distinguished from all other Hesperiidae by

unique wing shape (Fig. 8): fore wing outer margin concave in the

cell M3-CUA1, hindwing produced with a short and stout bifurcated

tail extending cell CuAi-CuA 2 and a lobe at tornus, antennal club

blunt without apiculus, palpi long, forewing cell longer than 3/5 of

Fig. 8. Jerini. Unique wing shape of Jera

tricuspidata (Mabille, 1902) S, Ecuador.

Genera included: Jera Lindsey, 1925.

Parent Taxon: Subfamily Pyrrhopyginae Mabille, 1877.

Subtribe Pythonidina Grishin, 2019

Type genus: Pythonides Hubner, [1819].

ZooBank registration:

CB890271-5483-4B5A-A7BC-27DBC5E23DE5

Diagnosis: The subtribe was defined as a clade from about 30 Mya that contains the genus Pythonides ,

and putatively synapomorphic DNA characters for this clade were given (Li et al ., 2019). Phenotypically,

species in this subtribe would key to E.44a, E.49.1, or, if uncus undivided, then to E.37a or 40d in Evans

(1953). A heterogeneous assembly of mostly small and frequently brightly patterned Hesperiidae not

easily diagnosed morphologically. In all genera, palpi porrect, antennae not shorter than 1/2 costa,

forewing without recurrent vein and hindwing costal margin not shorter than anal margin. Includes two

subgroups by uncus morphology. If (1) uncus deeply divided (Fig. 9), then 3rd segment of palpi as long as

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2nd segment, antennal club nearly arcuate at its middle,

apiculus not shorter than unbent portion of the club. If uncus

undivided, and if (2) antennal apiculus shorter than unbent

region of the club, then apiculus obtuse, not hooked,

hindwing about triangular in shape, not quadrate, palpi not

longer than head, or if (3) antennal club nearly arcuate at its

middle and apiculus not shorter than unbent portion of the

club, then 3rd segment of palpi shorter than 2nd segment.

Genera included: Ouleus Lindsey, 1925, Zera Evans, 1953,

Quadrus Lindsey, 1925, Gindanes Godman & Salvin, 1895,

Pythonides Hubner, [1819], Haemactis Mabille, 1903,

Atarnes Godman & Salvin, 1897, Eburuncus Grishin, 2012,

Milanion Godman & Salvin, 1895, Paramimus Hubner,

[1819], and Charidia Mabille, 1903.

Parent Taxon: Tribe Achlyodini Burmeister, 1878.

Subtribe Clitina Grishin, 2019

Fig. 9. Pythonidina. Long uncus arms in Eburuncus

hierax (Hopffer, 1874), Peru (top left); palpi (long 3 rd

segment) of E. unifasciata (C. & R. Felder, 1867), Costa

Rica (top right); antennae, bottom left: Ouleus calavius

(Godman & Salvin, 1895), Panama (above), Pythonides

lerina (Hewitson, 1868), French Guiana (below); palpi

(short 3 rd segment) of Gindanes brebisson panaetius

Godman & Salvin, 1895, Panama (bottom right).

Type genus: Clito Evans, 1953.

Diagnosis: The subtribe was defined as a clade from about 30 Mya

that contains the genus Clito , and putatively synapomorphic DNA

characters for this clade were given (Li et al., 2019).

Phenotypically, species in this subtribe would key to E.52 or E.13.8

in Evans (1953). A possible synapomorphic character is a

conspicuously long sheath of aedeagus that distinguishes this

subtribe from its relatives. In addition, a combination of the

following characters is diagnostic: uncus undivided, tapered,

apiculus arcuate at its center, sharply pointed, nudum not equally

partitioned, 3/13: apiculus much longer than the club, mid tibiae

with spines, wings produced, fore wing inner margin straight,

hindwing anal margin longer than costal margin, outer margin irregular, hindwing without hyaline spots

but frequently with white areas (Fig. 10).

Genera included: Clito Evans, 1953.

Parent Taxon: Tribe Erynnini Brues & Carpenter, 1932.

Tribe Butleriini Grishin, 2019

Type genus: Butleria Kirby, 1871.

ZooBank registration: D621EF81-FA65-4858-9450-E0C041598D7A

Diagnosis: The tribe was defined as a clade from about 40-45 Mya

that contains the genus Butleria , and putatively synapomorphic

DNA characters for this clade were given (Li et al ., 2019).

Phenotypically, species in this tribe would key to H.4 and H.5 in

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Fig. 11. Butleriini. Apiculus of antenna (top

left), hind leg (bottom left), and characteristic

wing shape (right) of Argopteron aureipennis

(Blanchard, 1852), Chile

Fig. 10. Clitina. Characteristic wing shape and

pattern in Clito palotchka Grishin, 2014,

Ecuador (left) and Clito mnemon (Schaus,

1913), Panama (right).

Evans (1955). Belongs to the subfamily Heteropterinae and differs from the nominotypical tribe by the

blunt somewhat flattened apiculus, compressed at the blunt tip, antennae not shorter than 1/2 costa, and

hind tibiae with upper spurs (sometimes short) (Fig. 11).

Genera included: Butleria Kirby, 1871 and Argopteron E. Watson, 1893.

Parent Taxon: Subfamily Heteropterinae Aurivillius, 1925.

Tribe Pericharini Grishin, 2019

Type genus: Perichares Scudder, 1872.

ZooBank registration: 94B68BD2-7F83-4E58-80E1-7F5AC8C56511

Diagnosis: The tribe was defined as a clade from about 32 My a

that contains the genus Perichares , and putatively synapomorphic

DNA characters for this clade were given (Li et al., 2019).

Phenotypically, species in this tribe would key to K.27a in Evans

(1955). Belongs to the "K. Carystus group" (not a monophyletic

assemblage) of Evans, characterized by the broad "quadrantic"

palpi, inner side of 2nd segment longer than the side contacting

head in dorsal view and the 3rd segment short and stout, nipple¬

like, antennae not constricted before apiculus, nudum 10-16

segments, its portion on the club shorter than the long apiculus.

Distinguished from others within this group by forewing discal cell not shorter than inner margin, and

vein CuAi opposite to vein Ri (not R2 or R3) at their origins (Fig. 12).

Genera included: Perichares Scudder, 1872, Alera Mabille, 1891, Orses Godman, 1901, Lycas Godman,

1901, Lychnuchoides Godman, 1901, Pseudorphe A. Warren & Dolibaina, 2015, and Orphe Godman,

1901.

Parent Taxon: Subfamily Hesperiinae Latreille, 1809.

Genus Tekliades Grishin, 2019

Type species: Thymele ramanatek Boisduval, 1833.

ZooBank registration: 081564BA-DA0C-4C46-AEAB-6C00131AC8BD

Diagnosis: The genus was defined as a clade from about 15 Mya

that contains the species Thymele ramanatek Boisduval, 1833, and

putatively synapomorphic DNA characters for this clade were

given (Li et al ., 2019). Phenotypically, species in this genus would

key to 1.1.9 in Evans (1937). The distinction of this genus from

Coeliades Hubner, 1818 was not obvious before DNA studies due

to similarity in appearance and genitalia. Distinguished from other

genera of Coeliadinae by undivided uncus, bulkier gnathos and

aedeagus, notch at the distal end of valva, white hindwing fringes,

and a white postdiscal band on hindwing below (Fig. 13).

Species included: Thymele ramanatek Boisduval, 1833.

Parent Taxon: Subfamily Coeliadinae Evans, 1937.

Fig. 13. Tekliades. Uniquely characteristic

ventral wing pattern of Tekliades ramanatek.

Fig. 12. Pericharini. Characteristic palpi,

antennae, and long forewing cell in Perichares

aurina Evans, 1955 S, Brazil: Parana.

Genus Salantoia Grishin, 2019

Type species: Eudamus eriopis Hewitson, 1867.

ZooBank registration: 3F82E9DE-A5A2-44B3-A13D-53CF8A673FAE

Diagnosis: The genus was defined as a clade from about 15 Mya

that contains the species Eudamus eriopis Hewitson, 1867, and

putatively synapomorphic DNA characters for this clade were

given (Li et al ., 2019). Phenotypically, species in this genus would

key to D.3.2 or D.3.3 in Evans (1952). Previously placed in

Sarmientoia Berg, 1897, but distinguished from it by rectangular,

not hook-shaped harpe in male genitalia and a lack of white spot in

cell M2-M3 on forewing above. Either character is diagnostic (Fig.

14).

Species included: Sarmientoia dinka (Evans, 1952) and Eudamus

eriopis (Hewitson, 1867).

Parent Taxon: Tribe Phocidini Tutt, 1906.

Genus Spicauda Grishin, 2019

Type species: Goniurus procne , Plotz, 1881.

Diagnosis: The genus was defined as a clade from about 15 Mya

that contains the species Goniurus procne , Plotz, 1881, and

putatively synapomorphic DNA characters for this clade were

given (Li et al ., 2019). Phenotypically, species in this genus would

key to C. 13.13c in Evans (1952). Previously placed in Urbanus

Hlibner, [1807] owing to long hindwing tails, but distinguished

from it by unique shape of genitalic harpe that is upturned and with

a spike-like process (or two) at its dorsal margin (Fig. 15).

Externally, can be distinguished from other Eudaminae with long

tails by a combination of the following characters: wings and body

brown, without extensive green scales; fringes not checkered, brown or paler, but not white; if fore wing

cell M3-CUA1 with hyaline spot, it forms part of the discal band, not detached from it.

Species included: Goniurus teleus Hubner, 1821, Urbanus tanna Evans, 1952, Urbanus ambiguus de

Jong, 1983, Urbanus cindra Evans, 1952, Goniurus zagorus Plotz, 1881, Papilio simplicius Stoll, 1790,

and Goniurus procne Plotz, 1881.

Parent Taxon: Subtribe Eudamina Mabille, 1877.

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Subgenus Urbanoides Grishin, 2019

Type species: Goniurus esmeraldus Butler, 1877.

ZooBank registration: 20FAC3B6-F038-40A0-B182-3C7F32A40702

Diagnosis: The subgenus was defined as a clade from about 12

Mya that contains the species Goniurus esmeraldus Butler, 1877,

and putatively synapomorphic DNA characters for this clade were

given (Li et al., 2019). Phenotypically, species in this subgenus

would key to C. 13.6a in Evans (1952). Member of this subgenus

are diagnosed by an apparent synapomorphy: dorsally directed

process on genitalic valvae, lacking in the nominotypical subgenus,

where valva may have a small "nose"-shaped projection (Fig. 16).

Species included: Goniurus esmeraldus A. Butler, 1877, Urbanus esma Evans, 1952, Urbanus prodicus

E. Bell, 1956, Urbanus elmina Evans, 1952, Urbanus evona Evans, 1952, Urbanus esta Evans, 1952,

Urbanus viridis H. Freeman, 1970.

Parent Taxon: Genus Urbanus Hubner, [1807].

Genus Zeutus Grishin, 2019

Type species: Cecropterus zeutus Moschler, 1879.

ZooBank registration: 75715B9C-46AB-40F5-B738-420DABD56B63

Diagnosis: The genus was defined as a clade from about 15 Mya

that contains the species Cecropterus zeutus Moschler, 1879, and

putatively synapomorphic DNA characters for this clade were

given (Li et al ., 2019). Phenotypically, species in this genus would

have genitalia as those described for zeutus by Williams & Bell

(1934: 27). More specifically, uncus arms shorter than in Calliades

Mabille & Boullet, 1912 where this species resided formerly;

valvae asymmetrical, broad, diamond-shaped; both harpes narrow,

deeply separated from the rest of valvae; aedeagus widens toward

its apex, with a rounded flange on its dorsal side (Fig. 17).

Species included: Cecropterus zeutus Moschler, 1879.

Parent Taxon: Subtribe Loboclina Grishin, 2019.

Genus Lobotractus Grishin, 2019

Type species: Eudamus valeriana Plotz, 1881.

ZooBank registration: C6E5B5DF-1C74-4DBD-85C3-7285209F6F03

Diagnosis: The genus was defined as a clade from about 15 Mya that contains the species Eudamus

valeriana Plotz, 1881, and putatively synapomorphic DNA characters for this clade were given (Li et al .,

2019). Phenotypically, diagnosis for this genus is the same as given for the "cyda group" by Bums (1996:

196). The following combination of characters unifies all known species in the genus and distinguishes

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them from all other genera: 3 rd segment of palpi long, longer than

in Codatractus (Fig. 18); costal fold absent; uncus undivided, in

dorsal view longer than wide and slightly concave terminally and

on the sides ("scalloped" per Evans (1952)), but not prominently

bilobed; tegumen humped in lateral view; shorter, fan-like comutus

with many sharp branches; valvae with a fang-like harpe bending

dorsad and enlarged, rounded ampulla protruding caudad to reach

the end of harpe; lamella antevaginalis expanded in two large

plates, each on the side of ostium bursae, plates reach the end of

lamella postvaginalis and protrude further narrowing towards the

middle into a sharp tooth, plates fully separated mid-ventrad

exposing ostium bursae ventrally. All these characters are illustrat

them and described in detail.

Species included: Eudamus valeriana Plotz, 1881, Thorybes uvydixa Dyar, 1914, and Heteropia cyda

Godman, 1901.

Parent Taxon: Subtribe Loboclina Grishin, 2019.

Subgenus Caudatractus Grishin, 2019

Type species: Eudamus alcaeus Hewitson, 1867.

ZooBank registration: DF0F3C91-F56E-4B65-B86C-385A36F9D7FD

Diagnosis: The subgenus was defined as a clade from about 7 Mya

that contains the species Eudamus alcaeus Hewitson, 1867, and

putatively synapomorphic DNA characters for this clade were

given (Li et al. , 2019). Phenotypically, species in this subgenus

would key to C. 11.1b in Evans (1952). These species share the

following characters of Codatractus : broadly arcuate antennae,

forewing with conjoined apical spots just beyond of discal cell in

cells R2-R3 and R3-R4, spot in fore wing cell M3-CUA1 midway

between the discal band and outer margin, ventral hindwing with 3 dark irregular bands. Species in this

subgenus are distinguished from all other Codatractus species by the tail near hindwing tornus (Fig. 19).

Species from the nominotypical subgenus lack tails, but their hind wing may be lobed at tornus.

Species included: Codatractus carlos Evans, 1952, Codatractus rowena Evans, 1952, Eudamus alcaeus

Hewitson, 1867, Codatractus apulia Evans, 1952, Codatractus yucatanus H. Freeman, 1977, and

Eudamus aminias Hewitson, 1867.

Parent Taxon: Genus Codatractus Lindsey, 1921.

Subgenus Asina Grishin, 2019

Type species: Eudamus asine Hewitson, 1867.

Diagnosis: The subgenus was defined as a clade from about 14 Mya that contains the species Eudamus

asine Hewitson, 1867, and putatively synapomorphic DNA characters for this clade were given (Li et al.,

2019). Phenotypically, species in this subgenus would key to C.7.2a in Evans (1952). The following

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Fig. 19. Caudatractus. Left: Codatractus

( Caudatractus) aminias, Brazil: MG, with tailed

hindwing vs. right: Codatractus ( Codatractus )

imalena (A. Butler, 1872), Brazil, AM.

Fig. 18. Lobotractus. 3rd segment of palpi

longer in Lobotractus (left) than in Codatractus

(Godman & Salvin, 1893), Mexico: Gro.

in Burns (1996), who discovered

characters distinguish them from other Hesperiidae except some

Ectomis : antennal club angled, hindwing elongated with a long tail

by the tomus, apical spots on fore wing in a straight line, spot in

forewing cell M3-CUA1 near discal band and farther from outer

margin, males with costal fold. The following characters

differentiate species in this subgenus from the nominal Ectomis :

head a thorax above brown (without extensive green scales),

forewing above with a dark spot about 1/3 from the base of cell

CuA 2-1A+2A, no spot in this cell past the discal band, but 2 small

spots (one frequently hyaline) under the pale spot from the discal

ventrally, aedeagus with a terminal spine projected to the left.

Species included: Polythrix gyges Evans, 1952, Goniuris [sic] hirtius A. Butler, 1870, Polythrix roma

Evans, 1952, Eudamus asine Hewitson, 1867, and Polythrix mexicanus H. Freeman, 1969.

Parent Taxon: Genus Ectomis Mabille, 1878.

Genus Tiana Grishin, 2019

Type species: Ebrietas niger Williams & Bell, 1940.

ZooBank registration: B9382699-24FB-4466-B39B-94E6B544C425

Diagnosis: The genus was defined as a clade from about 10 My a

that contains the species Ebrietas niger Williams & Bell, 1940, and

putatively synapomorphic DNA characters for this clade were

given (Li et al., 2019). Phenotypically, species in this genus would

key to F.7.3 or F.7.4 in Evans (1953). Combination of the

following characters is diagnostic of the genus (Fig. 21): forewing

discal cell very short, outer margin of both wings evenly convex,

palpi short, antennal shaft plain and club slender, nudum of 21-24

segments, males with costal fold on forewing, no tibial tufts, both

wings dark, almost unmarked or with two paler, ochreous brown bands above cut by dark veins without

violet sheen, below distal half of hindwing paler with darker veins. Differing from Tosta Evans, 1953 and

Iliana E. Bell, 1937 by uncus without projections, either undivided or with very short knob-like arms,

harpe simple, without processes, almost rectangular, unturned with serrated dorsal margin, not

prominently separated from the ampulla.

Species included: Ebrietas niger Williams & Bell, 1940 and Anastrus platypterus Mabille, 1895.

Parent Taxon: Tribe Carcharodini Verity, 1940.

Genus Chirgus Grishin, 2019

Type species: Hesperia limbata Erschoff, 1876.

ZooBank registration: 7B1905F1-9471-4BBF-90BF-32360783AB1E

Diagnosis: The genus was defined as a clade from about 10 Mya that contains the species Hesperia

limbata Erschoff, 1876, and putatively synapomorphic DNA characters for this clade were given (Li et

al ., 2019). Phenotypically, species in this genus would key to G.1.2e or G.1.9 in Evans (1953). Similar to

Pyrgus and Burnsius in checkered appearance, produced wings: fore wing costa longer than outer margin,

12

hindwing costa about the same length as outer margin, short,

extensively scaled palpi with inconspicuous 3rd segment, but differ

in the following characters (Fig. 22): tibial tuft present, no costal

fold in males, fore wing with a white spot at the base of cell CuAi-

CuA 2 , hyaline spots in cells R3-R4, R4-R5, and R5-M1 not produced

into cell R2-R3, and (1) if cell R1-R2 without a spot between the

apical spots and discal cell spot, then forewing lacks hyaline dot at

base of cell R3-R4 between apical spots and discal cell spot,

hindwing apex somewhat produced and outer margin slightly

concave in cell CuA 2-1A+2A, no submarginal white spots on

wings; or (2) if cell R1-R2 with a hyaline spot by the costa midway between the cell spot and apical spots,

then no streaks marginally from the forewing discal cell spot and no discal spots in cells Mi-M2 and M2-

M 3 .

Species included: Hesperia (Syrichthus [sic]) limbata Erschoff, 1876, Syrichthus [sic] nigella Weeks,

1902, Pyrgus barrosi Ureta, 1956, Pyrgus fides Hayward, 1940, Syrichtus bocchoris Hewitson, 1874, and

Pyrgus (Scelothrix [sic]) veturius Plotz, 1884.

Parent Taxon: Tribe Pyrgini Burmeister, 1878.

Genus Burnsius Grishin, 2019

Type species: Syricthus [sic] communis Grote, 1872.

ZooBank registration: 48996B74-3AB1-4DEA-9A64-B8F112E62343

Diagnosis: The genus was defined as a clade from about 10 Mya

that contains the species Syricthus [sic] communis Grote, 1872, and

putatively synapomorphic DNA characters for this clade were

given (Li et al. , 2019). Phenotypically, species in this genus would

key to G.1.5, G1.8, or G. 1.10a in Evans (1953). Similar to Pyrgus

and Chirgus in checkered appearance, produced wings: forewing

costa longer than outer margin, hindwing costa about the same

length as outer margin, short extensively scaled palpi with

inconspicuous 3rd segment, but differ in the following characters

(Fig. 23): forewing with a white spot at the base of cell CuAi-CuA 2 , hyaline spots in cells R5-M1, R4-R5,

R3-R4 not produced into cell R2-R3, and (1) if cell R1-R2 without spot between the apical spots and discal

cell spot, then forewing with hyaline dot at base of cell R3-R4 between apical spots and discal cell spot,

but (2) if no such dot, then hindwing outer margin evenly convex, not produced apically and white bar at

the end of discal cell absent both above and below; or (3) if cell R1-R2 with hyaline spot by the costa

midway between the cell spot and apical spots, then forewing with pale streaks marginally from the

fore wing discal cell spot and discal spots in cells Mi-M2 and M2-M3.

Species included: Syrichtus notatus Blanchard, 1852, Pyrgus crisia Herrich-Schaffer, 1865, Syricthus

[sic] communis Grote, 1872, Pyrgus albescens Plotz, 1884, Pyrgus adepta Plotz, 1884, Hydraenomia

orcynoides Giacomelli, 1928, Pyrgus chloe Evans, 1942, Hesperia titicaca Reverdin, 1921, Pyrgus

philetas W. H. Edwards, 1881, Papilio oileus Linnaeus, 1767, Papilio orcus Stoll, 1780, and Pyrgus

brenda Evans, 1942.

Fig. 22. Chirgus. Variation in the genus: C.

limbata, Chile, C. bocchoris trisignatus, Peru,

and C. veturius, Brazil: BA (left to right), all

Parent Taxon: Tribe Pyrgini Burmeister, 1878.

Genus Duroca Grishin, 2019

Type species: Hesperia duroca Plotz, 1882.

Diagnosis: The genus was defined as a clade from about 7 My a

that contains the species Hesperia duroca Plotz, 1882, and

putatively synapomorphic DNA characters for this clade were

given (Li et al. , 2019). Phenotypically, species in this genus would

key to J.39.5a in Evans (1955) and a combination of the following

characters is diagnostic: antennae about half of costa length, nudum

2/9 to 4/9, palpi narrow, 3rd segment short, conically shaped, mid

tibiae with spines, males with broad tripartite stigma from base of

vein CuAi to vein 1A+2A, origin on vein CuA2 on fore wing is

closer to the base than to vein CuAi. Stigma longer than in Lerema Scudder, 1872 and reaches vein

1A+2A closer to the base of wing (Fig. 24). Uncus and gnathos deeply divided, and this genus best

distinguished from other genera by broad valva with large harpe, upturned and shaped like a hook, being

more similar to some Phlebodes Hubner, [1819] and Saturnus Evans, 1955 than to Lerema , but uncus and

gnathos divided less deeply in these genera with hook-shaped harpe.

Species included: Hesperia duroca Plotz, 1882.

Parent Taxon: Subtribe Moncina A. Warren, 2008.

ACKNOWLEDGMENTS

The author is grateful to Jonathan P. Pelham for the encouragement and rich exchange of ideas that

formed the premise for and the basis of this paper, introduction in particular, and to Bernard Hermier for

helpful discussions. The study has been supported by grants from the National Institutes of Health

GM127390 and the Welch Foundation 1-1505.

LITERATURE CITED

Burns, J.M., 1996. Genitalia and the proper genus: Codatractus gets mysie and uvydixa- in a compact

cyda group-as well as a hysterectomy, while Cephise gets part of Polythrix (Hesperiidae:

Pyrginae). J. Lep. Soc. 50, 173-216.

Cong, Q., Borek, D., Otwinowski, Z. and Grishin, N.V., 2015. Tiger Swallowtail Genome Reveals

Mechanisms for Speciation and Caterpillar Chemical Defense. Cell Rep 10, 910-919.

Evans, W.H., 1937. A Catalogue of the African Hesperiidae indicating the classification and

nomenclature adopted in the British Museum, British Museum (Natural History), London.

Evans, W.H., 1949. A Catalogue of the Hesperiidae from Europe, Asia, and Australia in the British

Museum (Natural History), British Museum (Natural History), London.

Evans, W.H., 1951. A catalogue of the American Hesperiidae indicating the classification and

nomenclature adopted in the British Museum (Natural History). Part I. Introduction and Group A

Pyrrhopyginae., British Museum (Natural History), London.

14

Evans, W.H., 1952. A catalogue of the American Hesperiidae indicating the classification and

nomenclature adopted in the British Museum (Natural History). Part II. Pyrginae. Section I.,

British Museum (Natural History). London.

Evans, W.H., 1953. A catalogue of the American Hesperiidae indicating the classification and

nomenclature adopted in the British Museum (Natural History). Part III. Pyrginae. Section 2.,

British Museum (Natural History), London.

Evans, W.H., 1955. A catalogue of the American Hesperiidae indicating the classification and

nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae and

Megathyminae., British Museum (Natural History), London.

ICZN, 1999. International Code of Zoological Nomenclature. Fourth edition. The International Trust for

Zoological Nomenclature, London, UK.

Li, W., Cong, Q., Shen, J., Zhang, J., Hallwachs, W., Janzen, D.H. and Grishin, N.V., 2019. Genomes of

skipper butterflies reveal extensive convergence of wing patterns. Proc. Natl. Acad. Sci. U.S.A.

116(3): 6232-6237.

Talavera, G., Lukhtanov, V.A., Pierce, N.E. and Vila, R., 2012. Establishing criteria for higher-level

classification using molecular data: the systematics of Polyommatus blue butterflies (Lepidoptera,

Lycaenidae). Cladistics 29, 166-192.

Williams, R.C. and Bell, E.L., 1934. Studies in the American Hesperioidea. Paper II (Lepidoptera). Trans

Am Entomol Soc 60, 17-30.

15

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