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Volume 11 Number 2 18 May 2023
The Taxonomic Report
OF THE INTERNATIONAL LEPIDOPTERA SURVEY
ISSN 2643-4776 (print) / ISSN 2643-4806 (online)
A taxonomic list of the Old World genera in the subfamily
Hesperiinae (Hesperiidae) arranged into tribes
Jing Zhang'’°, Qian Cong!”, Jinhui Shen'’, Leina Song!’, and Nick V. Grishin!"
Departments of 'Biophysics, 7Biochemistry, and >Eugene McDermott Center For Human Growth & Development,
University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX 75390, USA;
“Corresponding author: grishin@chop.swmed.edu
ABSTRACT. A taxonomic list of valid genera in the subfamily Hesperiinae from the Old World assigned to tribes is
provided. This list is based on phylogenetic analysis of genomic sequence data complemented by morphological
considerations. As a result, there are no incertae sedis non-fossil genera in the family Hesperiidae.
Keywords: taxonomy, classification, genomics, phylogeny, biodiversity.
ZooBank registration: http://zoobank.org/CCF59E03-048E-440A-B310-4830DD2BB5C5
Here, we summarize tribal assignment for the Old World Hesperiinae Latreille, 1809 genera currently
regarded as valid (Lees et al. 2003; Warren et al. 2008; Warren et al. 2009; Fan et al. 2016; Toussaint et
al. 2018; Cong et al. 2019; Huang et al. 2019; Jiang et al. 2019; Li et al. 2019b; Zhang et al. 2022), with
comments about several others. No changes to nomenclature or synonymy are proposed. These results are
not new but are mostly scattered throughout our previous publications (Cong et al. 2019; Li et al. 2019a;
Zhang et al. 2019; Zhang et al. 2022) and are catalogued here for convenience. The following treatment is
deduced from the genomic tree shown in Fig. 1 based on our previously published datasets (Zhang et al.
2022), complemented with morphological considerations. The methods used are detailed in Li (2019a) for
experimental work and data processing and in Zhang et al. (2022) for phylogenetic tree construction.
Twelve tribes are ordered phylogenetically. Genera are listed alphabetically within each tribe to facilitate
visual search. We conclude that presently there are no incertae sedis non-fossil genera in the entire family
Hesperiidae because all genera have been assigned to tribes. All tribes are monophyletic in our analyses.
Family Hesperiidae Latreille, 1809
Subfamily Hesperiinae Latreille, 1809
Tribe Aeromachini Tutt, 1906
Aeromachus Nicéville, 1890
Ampittia Moore, 1881
Arnetta Watson, 1893
Baracus Moore, 1881
Creteus Nicéville, 1895
Halpe Moore, 1878
Halpemorpha Huang, Fan & Chiba, 2019
Lepella Evans, 1937
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Onryza Watson, 1893
Pedesta Hemming, 1934
Pithauria Moore, 1878
Praethoressa Huang, Chiba & Fan, 2019
Prosopalpus Holland, 1896
Sebastonyma Watson, 1893
(includes Parasovia Devyatkin, 1996)
Sovia Evans, 1949
Thoressa Swinhoe, 1913
Tribe Ceratrichiini Grishin, 2019
Argemma Grishin, 2019
Ceratrichia Butler, 1870
Herila Larsen & Collins, 2012
Meza Hemming, 1939
Pardaleodes Butler, 1870
(includes Ankola Evans, 1937)
Tribe Astictopterini Swinhoe, 1912
Acada Evans, 1937
Acleros Mabille, 1885
Actinor Watson, 1893
Andronymus Holland, 1896
Artitropa Holland, 1896
Astictopterus C. Felder & R. Felder, 1860
Caenides Holland, 1896
Ceratricula Larsen, 2013
Chondrolepis Mabille, 1904
Cupitha Moore, 1884
Dotta Grishin, 2019
Eogenes Mabille, 1909
Flandria Larsen, 2013
Fresna Evans, 1937
Fulda Evans, 1937
Galerga Mabille, 1898
Gamia Holland, 1896
Gorgyra Holland, 1896
Gyrogra Lindsey & Miller, 1965
Hidari Distant, 1886
Hollandus Larsen & Collins, 2015
Hypoleucis Mabille, 1891
Isoteinon C. Felder & R. Felder, 1862
Kedestes Watson, 1893
Lennia Grishin, 2022
Leona Evans, 1937
Lissia Grishin, 2019
Melphina Evans, 1937
Melphinyet Larsen, 2012
Moltena Evans, 1937
Monza Evans, 1937
Mopala Evans, 1937
Nervia Grishin, 2019
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Noctulana Larsen, 2012
Osmodes Holland, 1892
Osphantes Holland, 1896
Paracleros Berger, 1978
Paronymus Aurivillius, 1925
Parosmodes Holland, 1896
Perrotia Oberthiir, 1916
(includes Miraja Evans, 1937)
Platylesches Holland, 1896
Pteroteinon Watson, 1893
Rhabdomantis Holland, 1896
Semalea Holland, 1896
Teniorhinus Holland, 1892
Trida Grishin, 2022
Tsitana Evans, 1937
Xanthodisca Aurivillius, 1925
Xanthoneura Eliot, 1978
Xanthonymus Grishin, 2019
Zographetus Watson, 1893
Zophopetes Mabille, 1904
Tribe Gretnini Grishin, 2019
Gretna Evans, 1937
Tribe Taractrocerini Voss, 1952
Arrhenes Mabille, 1904
Banta Evans, 1949
Bibla Mabille, 1904
Cephrenes Waterhouse & Lyell, 1914
Kobrona Evans, 1935
Mimene Joicey & Talbot, 1917
Ocybadistes Heron, 1894
Oriens Evans, 1932
Pastria Evans, 1949
Potanthus Scudder, 1872
Sabera Swinhoe, 1908
Suniana Evans, 1934
Taractrocera Butler, 1870
Telicota Moore, 1881
Tiacellia Evans, 1949
Tribe Erionotini Distant, 1886
Acerbas Nicéville, 1895
Avestia Grishin, 2019
Cerba Grishin, 2019
Erionota Mabille, 1878
Gangara Moore, 1881
Ge Nicéville, 1895
Hyarotis Moore, 1881
Ilma Swinhoe, 1905
Lotongus Distant, 1886
Matapa Moore, 1881
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Oerane Elwes & Edwards, 1897
Pirdana Distant, 1886
Plastingia Butler, 1870
Ploetzia Saalmiiller, 1884
Praescobura Devyatkin, 2002
Pseudokerana Eliot, 1978
Pseudopirdana Chiba & Tsukiyama, 1993
Pudicitia Nicéville, 1895
Pyroneura Eliot, 1978
Quedara Swinhoe, 1919
Salanoemia Eliot, 1978
Scobura Elwes & Edwards, 1897
Suada Nicéville, 1895
Suastus Moore, 1881
Unkana Distant, 1886
Zela Nicéville, 1895
Tribe Notocryptini Swinhoe, 1913
Ancistroides Butler, 1874
(includes Notocrypta Nicéville, 1889 and Udaspes Moore, 1881)
Kerana Distant, 1886
Tamela Swinhoe, 1913
anbris Watson, 1893
Idmon Nicéville, 1895
Isma Distant, 1886
Tribe Eetionini Grishin, 2022
Eetion Nicéville, 1895
Tribe Psolosini Grishin, 2022
Koruthaialos Watson, 1893
(includes Stimula Nicéville, 1898)
Psolos Staudinger, 1889
Tribe Baorini Doherty, 1886
Afrogegenes de Jong & Coutsis, 2017
Baoris Moore, 1881
Borbo Evans, 1949
Brusa Evans, 1937
Caltoris Swinhoe, 1893
Gegenes Hibner, [1819]
Iton Nicéville, 1895
Parnara Moore, 1881
Pelopidas Walker, 1870
Polytremis Mabille, 1904
Prusiana Evans, 1937
Pseudoborbo Lee, 1966
Torbenlarsenia Kemal & Kogak, 2020
(Larsenia Chiba, Fan & Safian, 2016 is preoccupied)
Tsukiyamaia Zhu, Chiba & Wu, 2016
Zenonia Evans, 1935
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Aeromachini
70
=
=
1
=
_
oO
he
o
1S)
1
Zenonoida Fan & Chiba, 2016
Zinaida Evans, 1937
Tribe Hesperiini Latreille, 1809
bys
40
Hesperia Fabricius, 1793
Ochlodes Scudder, 1872
Thymelicus Hubner, [1819]
Barca bicolor|20058H02|China: Yunnan ——Barcinae
Trapezites symmomus|16106A10|Australia—Trapezitinae
Creteus cyrina|22017G08|Borneo
Arnetta atkinsoni|18019B09|India:Sikkim
Sovia lucasii]16108H11|China:Sichuan
100
1
100
Aeromachus stigmata|7915|Myanmar Hesperiini
Prosopalpus debilis|17092H04|Uganda
Halpemorpha eminens|19081G11|China
Halpe porus|17119G01|Myanmar
Praethoressa varia|18099B11|Japan
Pithauria murdava|18074B07|Myanmar
Pedesta masuriensis|17091A04|Manchuria
Thoressa masoni|7383|Myanmar
Sebastonyma dolopia|17091A01|India:Khasi Hills
Ampittia maro|7288|Sri Lanka
Lepella lepeletier|17093A04|Uganda
Baracus vittatus|16108HO09|Sri Lanka
Argemma argyrosticta|18064D06|Cameroon
Ceratrichia nothus|18053E03|no data
Pardaleodes edipus|17108G05|Zaire
Herila herilus|18064E01|Tanzania
Meza meza|17091H09|Cameroon
Platylesches picanini|KSO7|Cameroon
Melphina melphis|20125A03|Gabon
Osphantes ogowena|18087A10|Guinea
Gorgyra aburae|18019E12|Uganda
Cupitha purreea|7381|Myanmar
Parosmodes morantii|17092B08|Africa
Melphinyet flavina|19043E06|Uganda
Gyrogra subnotata|18053D11|Gabun
Noctulana noctula|18086D12|Ivory Coast
Zographetus satwa|17091C04|no data
Osmodes laronia|17119C02|Uganda
Rhabdomantis galatia|17069G05|Congo
Acleros leucopyga|18019F01|Madagascar
Paracleros biguttulus|17092H10|Uganda
Ceratricula semilutea|19043B08|Uganda
Paronymus ligora|18073B10|Sierra Leone
Fresna netopha|17092A07|Uganda
Andronymus caesar|17092B02|Uganda
Teniorhinus watsoni|20126F12|Gabon
Acada biseriata|17069G02|Kenya
Caenides dacela|7754|Liberia
Hollandus xanthopeplus|18072HO07|Eq. Guinea
Xanthonymus xanthioides|18064D07|Cameroon
Semalea pulvina|7803|Cameroon
Xanthodisca vibius|7765|Cameroon 98 100
Hypoleucis tripunctata|17091HO2|Uganda
Moltena fiara|18019F10|Natal
Eogenes alcides|18019B11|Turkey
Kedestes lepenula|17108F10|South Africa
Trida barberae|20126F11|South Africa
Tsitana tsita|18096G05|South Africa
Nervia nancy|17091B06|Kenya
Astictopterus jama|7394|Myanmar
Xanthoneura corissa|17091D05|Borneo
Fulda coroller|7762|Madagascar
Galerga hyposticta|7817|Madagascar ”
Perrotia albiplaga|21101B06|Madagascar
Dotta stellata|17093B03|Kenya
lsoteinon lamprospilus|17091A07|Taiwan
Chondrolepis niveicornis|17069G10|Kenya
Lennia lena|18087A07|Ivory Coast
Leona leonora|18087A09|Gabon
Mopala orma|17069H09|Cameroon
Hidari irava|7823|Singapore
Pteroteinon laufella|7757|Liberia
Monza cretacea|17069H12|Nigeria
Zophopetes dysmephila|17069H08|South Africa
Flandria weberi|20125A10|Cameroon
Lissia lissa|16108E11|Kenya
Artitropa erinnys|17092B07|Zimbabwe
Baorini
70
1
ERGoMi
Erionotini
00
ah_100
Qo}
Taractrocerini
Gretnini
Hesperiinae
ae
Psolosini
100
10
0.06
Hesperia comma|16108F07|Russia:Tien-Shan
Ochlodes agricola|PAQ23|USA:CA,Sierra Co.
Thymelicus acteon|18022H12|Spain
> Zenonia zeno|1/091B09|Kenya
Zenonoida eltola|7372|Myanmar
Zinaida nascens|18011A03|China
Caltoris kumara|17092A03|Borneo
Prusiana prusias|17119H01|Philippines
Baoris oceia|17091G11|Phillipines
Pelopidas thrax|7768|Greece
Pseudoborbo bevani|18011A04|Thailand
Torbenlarsenia holtzi|18014H02|Africa
Afrogegenes letterstedti|18064D03|South Africa
Gegenes pumilio|7764|Greece
Borbo borbonica|7818|Madagascar
Polytremis lubricans|7822|Singapore
Iton semamora|7393|Myanmar
Parnara guttatus|7290|Myanmar
Koruthaialos rubecula|18101C05|Myanmar
Psolos fuligo|7379|Myanmar
Eetion elia|7869|Malaysia
5 Isma protocliea|17091D04|Malaysia
Idmon distanti|7806|Malaysia
lambrix salsala|7376|Myanmar
Kerana armatus|18089E12|Malaysia
Tamela maura|18038H12|Malaysia
Ancistroides longicornis|18019C12|Sulawesi
Gangara thyrsis|16108F09|Philippines
Erionota thrax|7910|USA:HI,Molokai
Avestia avesta|18014E05|Borneo
Cerba martini|18089G07|Malaya
Zela zeus|18126A06|Langkawi
- Acerbas anthea|18095A06|Java
Matapa aria|7787|Philippines
IIma irvina|17091E06|Sulawesi
Unkana ambasa|18089G05|Philippines
Lotongus balta|7373|Myanmar
Pyroneura callineura|21105F10|Java
Pseudokerana fulgur|18057C01|Sumatra
Pseudopirdana albicornis|18053B07|Borneo
Ge geta|18057H08|M|no data
Oerane microthyrus|17091C01|N. Borneo
Suastus gremius|17108H04|India
Salanoemia sala|18073B06|Philippines
Plastingia flavescens|18057H05|Sulawesi
Ploetzia amygdalis|18053C07|Madagascar
Pirdana hyela|18089G06|Malaysia
Hyarotis adrastus|17091C12|Malaysia
Quedara monteithi|18014E01|Borneo
Scobura cephala|18019C05|India:Sikkim
Suada swerga|21105D10|N. India
Tiacellia tiacellia|22017G09|Indonesia: Papua Region
Pastria pastria|18082E01|NHMUK0247277187|PNG
Oriens gola|18103C03|Thailand
Taractrocera sagara|7375|Myanmar
Kobrona kobros|18063C05|Aru
Ocybadistes walkeri|19042B07|Australia:SA
Potanthus omaha|7388|Malaysia
Suniana lascivia|18075F11|Australia:NQ
Bibla papyria|18011A07|Australia:NSW
Telicota colon|7783|Philippines
Cephrenes augiades|7758|Australia:NSW
Sabera caesina|19067A02|Australia
Mimene miltias|18063C09|no data
Arrhenes marnas|17091G03|Papua New Guinea
Gretna cylinda|18019F12|Uganda
100
Gamia shelleyi|16108E10|Uganda
ee i |
Fig. 1. A phylogenetic tree of representative Hesperiinae (mostly type species of their genera) constructed from all protein-
coding autosomal genes in the nuclear genome. Specimen/sample codes and general localities are given. The numbers of
dataset partitions (Zhang et al. 2022) supporting the branches are shown by corresponding nodes. The tree is folded over to fit
on a page, and black arrows guide the direction of viewing. Detailed analysis of the results will be presented elsewhere.
5
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ACKNOWLEDGMENTS
We acknowledge 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), Blanca Huertas, David Lees, and Geoff Martin (BMNH: Natural History Museum, London, UK),
Jim Fetzner, Bob Androw, Vanessa Verdecia, Cat Giles, and the late John Rawlins (CMNH: Carnegie
Museum of Natural History, Pittsburgh, PA, USA), Weiping Xie (LACM: Los Angeles County Museum
of Natural History, Los Angeles, CA, USA), Théo Léger, Wolfram Mey, and Viola Richter (MFNB:
Museum fiir Naturkunde, Berlin, Germany), Rodolphe Rougerie (MNHP: Muséum National d'Histoire
Naturelle, Paris, France), Matthias Nuss (MTD: Museum fiir Tierkunde, Dresden, Germany), 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), Axel Hausmann, Andreas Segerer, and Ulf Buchsbaum
(ZSMC: Zoologische Staatssammlung Miinchen, Germany), for granting access to the collections under
their care and stimulating discussions; Ernst Brockmann and Kojiro Shiraiwa for specimens and leg
samples, the late Paul A. Opler and Evi Buckner-Opler for collecting specimens under the California
Department of Fish and Game permit SC13645; Gerardo Lamas for critical review of the manuscript and
correcting several mistakes. We acknowledge the Texas Advanced Computing Center (TACC) at The
University of Texas at Austin for providing HPC resources. This study was supported in part by the
HHMI Investigator funds and by grants from the National Institutes of Health GM127390 and the Welch
Foundation I-1505.
LITERATURE CITED
Cong, Q., J. Zhang, J. Shen, and N. V. Grishin. 2019. Fifty new genera of Hesperiidae (Lepidoptera). Insecta Mundi
0731: 1-56.
Fan, X., H. Chiba, Z. Huang, W. Fei, M. Wang, and S. Safian. 2016. Clarification of the phylogenetic framework of
the tribe Baorini (Lepidoptera: Hesperiidae: Hesperiinae) inferred from multiple gene sequences. PLoS One
11(7): e0156861.
Huang, Z.-F., H. Chiba, J. Jin, A. G. Kizhakke, M. Wang, K. Kunte, and X.-L. Fan. 2019. A multilocus
phylogenetic framework of the tribe Aeromachini (Lepidoptera: Hesperiidae: Hesperiinae), with implications for
taxonomy and biogeography. Systematic Entomology 44(1): 163-178.
Jiang, W., H. He, Y. Li, Y. Wang, C. Ge, J. Zhu, and W. Yu. 2019. Molecular phylogeny of the butterfly tribe Baorini
Doherty, 1886 (Hesperiidae, Hesperiinae) in China. Zootaxa 4565(1): 108-116.
Lees, D. C., C. Kremen, and H. Raharitsimba. 2003. Classification, diversity and endemism of the butterflies
(Papilionoidea and Hesperioidea): a revised species checklist. Jn: M., G. S., and J. P. Benstead (Eds.). The
Natural History of Madagascar. University of Chicago Press; Chicago, pp. 762—793.
Li, W., Q. Cong, J. Shen, J. Zhang, W. Hallwachs, D. H. Janzen, and N. V. Grishin. 2019a. Genomes of skipper
butterflies reveal extensive convergence of wing patterns. Proceedings of the National Academy of Sciences of
the United States of America 116(13): 6232-6237.
Li, Y., J. Zhu, C. Ge, Y. Wang, Z. Zhao, S. Ma, A. A. Hoffmann, N. M. Endersby, Q. Liu, W. Yu, and W. Jiang.
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Hesperiidae) from China. Cells 8(4): 294.
Toussaint, E. F. A., J. W. Breinholt, C. Earl, A. D. Warren, A. V. Z. Brower, M. Yago, K. M. Dexter, M. Espeland,
N. E. Pierce, D. J. Lohman, and A. Y. Kawahara. 2018. Anchored phylogenomics illuminates the skipper
butterfly tree of life. BMC Evolutionary Biology 18(1): 101.
Warren, A. D., J. R. Ogawa, and A. V. Z. Brower. 2008. Phylogenetic relationships of subfamilies and
circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea). Cladistics 24(5): 642-676.
Warren, A. D., J. R. Ogawa, and A. V. Z. Brower. 2009. Revised classification of the family Hesperiidae
(Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomology
34(3): 467-523.
Thang, J., Q. Cong, J. Shen, E. Brockmann, and N. V. Grishin. 2019. Three new subfamilies of skipper butterflies
(Lepidoptera, Hesperiidae). Zookeys 861: 91-105.
Zhang, J., Q. Cong, J. Shen, and N. V. Grishin. 2022. Taxonomic changes suggested by the genomic analysis of
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