Fifty new genera of Hesperiidae (Lepidoptera)

Qian Cong; Jing Zhang; Jinhui Shen; Nick V Grishin

doi:10.5281/zenodo.3677235

. Author manuscript; available in PMC: 2022 Jan 26.

Published in final edited form as: Insecta mundi. 2019 Oct 11;2019:0731. doi: 10.5281/zenodo.3677235

Fifty new genera of Hesperiidae (Lepidoptera)

Qian Cong ^1,^#, Jing Zhang ^2,^#, Jinhui Shen ³, Nick V Grishin ⁴

PMCID: PMC8791444 NIHMSID: NIHMS1724225 PMID: 35087260

Abstract

Genomic sequencing and analysis of worldwide skipper butterfly (Lepidoptera: Hesperiidae) fauna points to imperfections in their current classification. Some tribes, subtribes and genera as they are circumscribed today are not monophyletic. Rationalizing genomic results from the perspective of phenotypic characters suggests two new tribes, two new subtribes and 50 new genera that are named here: Ceratrichiini Grishin, trib. n., Gretnini Grishin, trib. n., Falgina Grishin, subtr. n., Apaustina Grishin, subtr. n., Flattoides Grishin, gen. n., Aurivittia Grishin, gen. n., Viuria Grishin, gen. n., Clytius Grishin, gen. n., Incisus Grishin, gen. n., Perus Grishin, gen. n., Livida Grishin, gen. n., Festivia Grishin, gen. n., Hoodus Grishin, gen. n., Anaxas Grishin, gen. n., Chiothion Grishin, gen. n., Crenda Grishin, gen. n., Santa Grishin, gen. n., Canesia Grishin, gen. n., Bralus Grishin, gen. n., Ladda Grishin, gen. n., Willema Grishin, gen. n., Argemma Grishin, gen. n., Nervia Grishin, gen. n., Dotta Grishin, gen. n., Lissia Grishin, gen. n., Xanthonymus Grishin, gen. n., Cerba Grishin, gen. n., Avestia Grishin, gen. n., Zetka Grishin, gen. n., Turmosa Grishin, gen. n., Mielkeus Grishin, gen. n., Coolus Grishin, gen. n., Daron Grishin, gen. n., Barrolla Grishin, gen. n., Brownus Grishin, gen. n., Tava Grishin, gen. n., Rigga Grishin, gen. n., Haza Grishin, gen. n., Dubia Grishin, gen. n., Pares Grishin, gen. n., Chitta Grishin, gen. n., Artonia Grishin, gen. n., Lurida Grishin, gen. n., Corra Grishin, gen. n., Fidius Grishin, gen. n., Veadda Grishin, gen. n., Tricrista Grishin, gen. n., Viridina Grishin, gen. n., Alychna Grishin, gen. n., Ralis Grishin, gen. n., Testia Grishin, gen. n., Buzella Grishin, gen. n., Vernia Grishin, gen. n., and Lon Grishin, gen. n. In addition, the following taxonomic changes are suggested. Prada Evans is transferred from Hesperiinae to Trapezitinae. Echelatus Godman and Salvin, Systaspes Weeks, and Oenides Mabille are removed from synonymy and are treated as valid genera. The following genera are new junior subjective synonyms: Tosta Evans of Eantis Boisduval; Turmada Evans of Neoxeniades Hayward, Arita Evans of Tigasis Godman, and Alera Mabille of Perichares Scudder. Eantis pallida (R. Felder) (not Achlyodes Hübner), Gindanes kelso (Evans) (not Onenses Godman and Salvin), Isoteinon abjecta (Snellen) (not Astictopterus C. and R. Felder), Neoxeniades ethoda (Hewitson) (not Xeniades Godman), Moeris anna (Mabille) (not Vidius Evans), and Molo pelta Evans (not Lychnuchus Hübner) are new genus-species combinations. The following are species-level taxa: Livida assecla (Mabille) (not a subspecies of Livida grandis (Mabille), formerly Pythonides Hübner) and Alychna zenus (E. Bell) (not a junior subjective synonym of Alychna exclamationis (Mabille), formerly Psoralis Mabille); and Barrolla molla E. Bell (formerly Vacerra Godman) is a junior subjective synonym of Barrolla barroni Evans (formerly Paratrytone Godman). All these changes to taxonomic status of names are propagated to all names currently treated as subspecies (for species), subgenera (for genera) and synonyms of these taxa. Finally, taxa not mentioned in this work are considered to remain at the ranks and in taxonomic groups they have been previously assigned to.

Keywords: Genomics, higher classification, taxonomy, biodiversity, phylogeny

Introduction

Hesperiidae, commonly known as skippers, are a charismatic group of butterflies that frequently look moth-like due to their stout bodies (Watson 1893), mostly brown and gray colors and fast wing beats. Some are crepuscular and even come to light (Austin 2008). However, recent DNA-based studies argue that they may have originated deep within butterfly radiation, after the swallowtails (Papilionidae) have split from the common ancestor (Wahlberg et al. 2005; Kawahara and Breinholt 2014; Espeland et al. 2018).

Having worldwide distribution, Hesperiidae are highly speciose with more than 3500 species described. However, they have received less attention than other butterfly families. The major milestone works remain those of Evans, who offered a comprehensive taxonomic treatment of the group in six volumes (Evans 1937, 1949, 1951, 1952, 1953, 1955). Refinement of this classification was catalyzed by new methodologies. The groundbreaking work of Warren et al. (2008, 2009) re-shaped the higher classification based on combination of DNA sequences of several genes with morphological characters. Several follow up studies employed a larger set of genes, up to several hundred (Sahoo et al. 2016, 2017; Toussaint et al. 2018). Then, a genome-scale revisionary work has followed (Li et al. 2019; Zhang et al. 2019b, c).

Genomic analyses revealed many nuances not readily apparent from a morphological standpoint. However, retrospective comparison of phenotypic characters is consistent with the picture emerging from the comparison of phylogenetic trees based on nuclear and mitochondrial genomes (Li et al. 2019; Zhang et al. 2019c). We carried out genomic sequencing of representative Hesperiidae species from all known genera. As a result, we found inconsistencies in assignment of species to genera, and many genera and some tribes and subtribes were not monophyletic as currently defined. While the details of the phylogenetic analysis and their implications for the higher classification of Hesperiidae will be presented elsewhere, some of these inconsistencies are corrected here by proposing new names for two tribes, two subtribes and 50 genera detected in phylogenetic trees. Here, we show only a subset of data necessary to justify our conclusions.

Materials and Methods

This study is based on whole genomic shotgun DNA sequences that were mostly obtained from pinned and dried specimens in collections. Many of these specimens were collected more than a century ago and a number of them were primary type specimens. See Table 1 for a brief data summary of the specimens used in this work, and Table S1 in the Supplemental file deposited at https://osf.io/5cfht/?view_only=21eb53b6f8f344afaee3de2be90bf5d2 for details. We use either an abdomen, pieces of muscle tissue taken from the thorax through the abdomen attachment site (for previously dissected specimens), or a leg for DNA extraction. The abdomen is used when genitalic dissection is needed. Muscle tissue is a viable alternative to the leg when leg material is not sufficient or using a leg is not possible. Legs were the most convenient choice because they were easier to sample and often yielded better-quality DNA. Therefore, most specimens were sequenced from legs. The details of protocols for DNA extraction, genomic library preparation, sequencing and analysis are given in our previous publications (Cong et al. 2015a, b, 2016a, b, 2017a, b, 2018; Shen et al. 2015, 2016a, b, 2017; Zhang et al. 2017a, b; Li et al. 2019; Zhang et al. 2019a, b). In the methods employed, this study is identical to that of Li et al. (2019), and only difference is that a larger number of species and specimens were used.

Table 1.

Data for 620 sequenced Hesperiidae specimens. See Table S1 in the Supplemental file deposited at https://osf.io/5cfht/?view_only=21eb53b6f8f344afaee3de2be90bf5d2 for detailed information about these specimens and Materials and Methods section for collection abbreviations. Type status abbreviations are: AT, allotype; HT, holotype; LT, lectotype; PT, paratype, ST, syntype; T, type.

DNA voucher	Taxon name	Type	Brief data	Collection
NVG-18054G10	Acada biseriata		Zimbabwe, 1993	ZMHB
NVG-18081B11	Acerbas anthea		Malaysia, 1917, NHMUK_010430824, 0247275554	BMNH
NVG-17091E09	Acerbas duris duris		Philippines, old (around 1900)	USNM
NVG-18074B03	Acerbas sarala sarala		India, 1890	ZMHB
NVG-18011F11	Achlyodes busirus busirus		French Guiana, 1988	USNM
NVG-18081B04	Acleros leucopyga		Madagascar, 1921, NHMUK_010430828, 0247274717	BMNH
NVG-18081D02	Actinor radians		India, 1885, NHMUK_010430809, 0247275546	BMNH
NVG-18015A06	Adlerodea petrovna	T	Brazil: Rio de Janeiro, old (around 1900)	USNM
NVG-7988	Adopaeoides prittwitzi		USA: AZ, Santa Cruz Co., 1999	USNM
NVG-7915	Aeromachus stigmata shanda		Myanmar, 2002	USNM
NVG-18038C05	Aethilla chiapa		Honduras	RGallardo
NVG-18013G10	Aethilla echina		Costa Rica, 2005, 05-SRNP-20	USNM
NVG-18011F09	Aethilla eleusinia		Peru, 2012	USNM
NVG-18056D08	Aethilla gigas		Bolivia, old (around 1900)	ZfBS
NVG-7891	Aethilla lavochreaDHJ02		Costa Rica, 2013, 13-SRNP-22231	USNM
NVG-14064D01	Aethilla memmius		Venezuela, 1985	USNM
NVG-18012E06	Aides duma argyrina		Bolivia?, no date?1970	USNM
NVG-18054H04	Alenia namaqua		South Africa, 2002	ZMHB
NVG-15035E02	Alychna degener	ST	no data	ZMHB
NVG-18042G08	Alychna exclamationis	ST	Bolivia, old (around 1900)	ZMHB
NVG-8007	Alychna victa		Ecuador, 2004	USNM
NVG-18027A06	Alychna zenus	HT	Ecuador, 1939	AMNH
NVG-6470	Amblyscirtes vialis		USA: CO, Grand Co., 2016	UTSW
NVG-7288	Ampittia dioscorides singa		Sri Lanka, 1971	USNM
NVG-18012A02	Anastrus meliboea		Costa Rica, 1977	USNM
NVG-18013F12	Anastrus neaeris		Costa Rica, 2008, 08-SRNP-2106	USNM
NVG-18011H11	Anastrus obscurus		Brazil: Rondonia, 1991	USNM
NVG-18067H10	Anastrus ulpianus		Brazil: Santa Catarina, 2000	EBrockmann
NVG-18056D06	Anaxas obliqua		Bolivia, old (around 1900)	ZfBS
NVG-4461	Ancyloxypha numitor		USA: TX, Dallas Co., 2015	UTSW
NVG-18091C07	Anisochoria pedaliodina pedaliodina		Ecuador, 2012	EBrockmann
NVG-7909	Anisochoria pedaliodina polysticta		Costa Rica, 2004, 04-SRNP-15751	USNM
NVG-18011G07	Anisochoria quadrifenestrata		Colombia, old (around 1900)	USNM
NVG-18019B02	Anisynta cynone		Australia, 1961	AMNH
NVG-17069F11	Ankola fan		Kenya, 1955	USNM
NVG-17111E11	Anthoptus epictetus		Brazil: Santa Catarina, 1999	LACM
NVG-10646	Antigonus emorsa		Mexico: Michoacan, 1994	TAMU
NVG-7907	Antigonus erosus		Costa Rica, 2013, 13-SRNP-56479	USNM
NVG-18014A03	Antigonus nearchus		Costa Rica, 2006, 06-SRNP-32799	USNM
NVG-18082B12	Antipodia chaostola		Australia, 1934, NHMUK_012824109, 0247278477	BMNH
NVG-18079E09	Apallaga fulgens fulgens		Cameroon, 1987, EL63116	MNHP
NVG-18011B09	Apallaga mokeezi		South Africa, 1978	USNM
NVG-17108C09	Apallaga opalinus-cf		Kenya, 2001	LACM
NVG-18033B11	Apallaga opalinus-cf		Uganda, 2008	MWalker
NVG-18055G03	Apallaga pooanus obscuripennis	ST	Equatorial Guinea, 1906	ZMHB
NVG-18079F03	Apallaga rutilans rutilans		Cameroon, 1991, EL63122	MNHP
NVG-18012F08	Apaustus menes		Peru, 2011	USNM
NVG-18064D06	Argemma argyrosticta		Cameroon, old (around 1900)	USNM
NVG-7921	Argon lota		Costa Rica, 2007, 07-SRNP-55877	USNM
NVG-17069C01	Argopteron aureipennis		Chile, 1982	USNM
NVG-18019B09	Arnetta atkinsoni		India, 1927	AMNH
NVG-5065	Aroma aroma		Costa Rica, 04-SRNP-1707	USNM
NVG-5485	Arteurotia tractipennis tractipennis		USA: TX, Hidalgo Co., 1972	TAMU
NVG-18012H06	Artines aquilina		Brazil: Santa Catarina, 1999	USNM
NVG-18067F11	Artitropa erinnys		Comoros, 1987	EBrockmann
NVG-19022F06	Artonia artona		Brazil: Parana, 1995	USNM
NVG-4881	Asbolis capucinus		USA: FL, Monroe Co., 2015	UTSW
NVG-7394	Astictopterus jama jama		Myanmar, 2001	USNM
NVG-3718	Atalopedes campestris huron		USA: TX, Dallas Co., 2015	UTSW
NVG-18113D01	Atalopedes flaveola (=clarkei)	HT	Venezuela, 1985	USNM
NVG-17092D07	Atalopedes lina		Colombia, 1991	USNM
NVG-5313	Atalopedes mesogramma apa		Dominican Republic, 2015	UTSW
NVG-7903	Atarnes sallei		Costa Rica, 2007, 07-SRNP-59529	USNM
NVG-17119H02	Aurivittia aurivittata		Myanmar, 2002	USNM
NVG-18033B05	Aurivittia cameroni		Malaysia, 2003	MWalker
NVG-14101B01	Austinus heroica		Colombia, 1946	AMNH
NVG-18067C09	Austinus heros		Brazil: Para, 2009	EBrockmann
NVG-18014E05	Avestia avesta		Malaysia, old (around 1900)	USNM
NVG-17091G11	Baoris oceia		Phillipines, 1914	USNM
NVG-18089F07	Barca bicolor		China: Shaanxi, 2009	EBrockmann
NVG-18026C12	Barrolla molla	HT	Ecuador	AMNH
NVG-7246	Bolla brennus brennus		Panama, 1981	USNM
NVG-17108F05	Bolla cylindus		Mexico: Hidalgo, 1982	LACM
NVG-18071A09	Bolla evippe		Costa Rica, 2009, 09-SRNP-57008	USNM
NVG-18011H02	Bolla imbras		Panama, 1984	USNM
NVG-18083B12	Bolla saletas	T	Guatemala, BMNH(E)_1669828, 0247277236	BMNH
NVG-18067H11	Bralus albida albida		Peru, 1994	EBrockmann
NVG-18025F01	Brownus browni	HT	Costa Rica, 1946	AMNH
NVG-17095E11	Brownus browni		Costa Rica, 2006	USNM
NVG-17093D04	Burca concolor concolor		Cuba, 2010	USNM
NVG-7552	Burnsius communis		USA: TX, Bexar Co., 1977	TAMU
NVG-16108G04	Butleria flavomaculatus valdivianus		Chile, 1979, USNMENT 00894446	USNM
NVG-8052	Buzella mellanaformis		Panama, 1982	USNM
NVG-18013C12	Buzyges idothea		Costa Rica, 1980	USNM
NVG-5721	Cabirus procas		Ecuador, 2004	USNM
NVG-7754	Caenides dacela		Liberia, 1988	USNM
NVG-17092C10	Callimormus juventus		Costa Rica, 2012, 12-SRNP-20224	USNM
NVG-4591	Calpodes ethlius		USA: TX, Cameron Co., 2015	UTSW
NVG-18013G01	Camptopleura theramenes		Costa Rica, 2015, 15-SRNP-45798	USNM
NVG-7905	Canesia canescens		Costa Rica, 2014, 14-SRNP-1649	USNM
NVG-18013H12	Canesia meridensis		Costa Rica, 1997, 97-SRNP-1522	USNM
NVG-18012F06	Cantha calva		Peru, 1986	USNM
NVG-18033B07	Capila phanaeus		Cambodia, 2006	MWalker
NVG-7763	Carcharodus alceae		Greece, 1990	USNM
NVG-18014A02	Carrhenes fuscescens		Costa Rica, 1995, 95-SRNP-6819	USNM
NVG-18089C06	Carterocephalus alcina		China: Sichuan, 2006	EBrockmann
NVG-18021G04	Carterocephalus avanti flavostigma		Tibet, 1944	AMNH
NVG-17119H05	Carterocephalus christophi		China, old (around 1900)	USNM
NVG-18089B05	Carterocephalus gemmatus		China: Sichuan, 2002	EBrockmann
NVG-18089B08	Carterocephalus houangty houangty		China: Sichuan, 2007	EBrockmann
PAO-69	Carterocephalus mandan		USA: CA, Sierra Co., 2016	UTSW
NVG-18089C04	Carterocephalus micio		China: Yunnan, 2006	EBrockmann
NVG-18038G04	Carterocephalus silvicola		Russia: Buryatia, 2016, 5195	UTSW
NVG-18012D09	Carystina lysiteles		Peru, 1999	USNM
NVG-18012D05	Carystus jolus		Colombia, 1971	USNM
NVG-18011B06	Celaenorrhinus aegiochus		Panama, 1981	USNM
NVG-18064A03	Celaenorrhinus bettoni		Uganda, 1952	USNM
NVG-18013G07	Celaenorrhinus eligius		Costa Rica, 2010, 10-SRNP-20588	USNM
NVG-17093D03	Celaenorrhinus ficulnea ficulnea		Malaysia, old (around 1900)	USNM
NVG-18011B04	Celaenorrhinus monartus		Panama, 1981	USNM
NVG-18073G09	Celaenorrhinus rosetta		Gabon, old (around 1900)	ZMHB
NVG-18011B08	Celaenorrhinus spilothyrus		Sri Lanka, 1976	USNM
NVG-18019A07	Celaenorrhinus sumitra		Nepal, 1937	AMNH
NVG-7993	Celaenorrhinus syllius		Ecuador, 2002, USNMENT 01321833	USNM
11-BOA-13383BrockB12	Celotes limpia		USA: TX, Brewster Co.	JPBrock
NVG-3956	Celotes nessus		USA: TX, Hidalgo Co., 2015	UTSW
NVG-17067E03	Celotes spurcus	PT	Mexico: Queretaro, 2007, CSU_ENT1024631	CSUC
NVG-5684	Cephise aelius		Costa Rica, 2014, 14-SRNP-70854	USNM
NVG-7398	Ceratrichia flava		Cameroon, 1989	USNM
NVG-18053E03	Ceratrichia nothus		no data	ZMHB
NVG-7401	Ceratrichia wollastoni		Cameroon, 1989	USNM
NVG-18087B09	Ceratricula semilutea		Sierra Leone, 1906, EL62955	MNHP
NVG-18089G07	Cerba martini		Malaysia, 1993	EBrockmann
NVG-18039B11	Chaetocneme beata		Australia, old (around 1900)	FMNH
NVG-17108C12	Chaetocneme helirius		British New Guinea, 1962	LACM
NVG-17069A11	Chamunda chamunda		India, old (around 1900)	USNM
NVG-7981	Charidia lucaria		Peru, 2008	USNM
NVG-18057A01	Chiomara gundlachi		Cuba, 2014	ZSMC
NVG-17109F12	Chiomara mithrax		USA: AZ, Santa Cruz Co., 1992	LACM
NVG-14103A02	Chiothion asychis asychis		Suriname	USNM
NVG-17109F10	Chiothion basigutta		Brazil: Goias, 1956	LACM
NVG-5100	Chiothion georgina		USA: TX, Starr Co., 2015	UTSW
NVG-18081D11	Chiothion khalili		Brazil: AM, 1929, NHMUK_010430875, 0247277240	BMNH
NVG-14102F02	Chirgus limbata		Chile, 1952	FMNH
NVG-18025D04	Chitta chittara (=alis)	HT	Brazil: Santa Catarina, old (around 1900)	AMNH
NVG-5271	Choaspes hemixanthus furcata		China: Sichuan Prov., 2015	UTSW
NVG-17069G10	Chondrolepis niveicornis		Kenya, 1951	USNM
NVG-18117E12	Choranthus radians		Cuba, 2010	USNM
NVG-14107C04	Clito aberrans-cf		Peru, 1982	USNM
NVG-15117B05	Clytius clytius		Mexico: Sinaloa, 2003, CSU_ENT1039479	CSUC
NVG-17108F02	Clytius clytius		USA: AZ, Santa Cruz Co., 1991	LACM
NVG-18013A06	Cobalopsis autumna		Costa Rica, old (around 1900)	USNM
NVG-7927	Cobalus virbius		Costa Rica, 2012, 12-SRNP-22162	USNM
NVG-3354	Cogia calchas		USA: TX, Hidalgo Co., 2015	UTSW
NVG-7963	Conga chydaea		Costa Rica, 2009, 09-SRNP-68418	USNM
NVG-14107B12	Conognathus platon		Ecuador, 1989	USNM
NVG-15104C09	Coolus bushi	HT	Dominican Republic	AMNH
NVG-8381	Copaeodes aurantiaca		USA: TX, Blanco Co., 2017	UTSW
NVG-18063D07	Cornuphallus onoribo		French Guiana, 2004, H21114	BHermier
NVG-18071C04	Corra coryna conka		Costa Rica, 2007, 07-SRNP-35032	USNM
NVG-17092F04	Corticea corticea		Costa Rica, 2012, 12-SRNP-70085	USNM
NVG-14064H09	Crenda crenda		Brazil: Parana, old (around 1900)	USNM
NVG-18081D10	Crenda crenda	HT	Brazil: Parana, NHMUK_010430876, 0247278431	BMNH
NVG-17092G08	Cumbre cumbre		Brazil: Rio de Janeiro, 1994	USNM
NVG-7381	Cupitha purreea		Myanmar, 2001	USNM
NVG-15117B07	Cycloglypha thrasibulus thrasibulus		Mexico: Sinaloa, 2003, CSU_ENT1039523	CSUC
NVG-17105F01	Cyclosemia herennius		Guyana, 2000	USNM
NVG-4842	Cymaenes tripunctus tripunctus		USA: FL, Collier Co., 2015	UTSW
NVG-7960	Cynea cynea		Costa Rica, 2010, 10-SRNP-35740	USNM
NVG-18054C11	Dalla agathocles agathocles		Colombia, 1921	ZMHB
NVG-18017B10	Dalla agathocles lanna		Ecuador, 1993	USNM
NVG-18014F05	Dalla caicus inca		Peru, 2011	USNM
NVG-18014E11	Dalla costala		Peru, 2011	USNM
NVG-18014F06	Dalla cyprius quinka		Peru, 2013	USNM
NVG-18014E12	Dalla dimidiatus lilla		Peru, 2011	USNM
NVG-18017B08	Dalla dimidiatus pucer		Peru, 1992	USNM
NVG-18073F06	Dalla eryonas		Panama, old (around 1900)	ZMHB
NVG-18014F03	Dalla frater		Peru, 2013	USNM
NVG-18017B04	Dalla frontinia-cf		Ecuador, 1986	USNM
11-BOA-13383BrockB09	Dalla nona		Peru	JPBrock
NVG-18017B05	Dalla pincha		Ecuador, 1982	USNM
NVG-18017B12	Dalla semiargentea		Colombia, 1965	USNM
NVG-17111B10	Damas clavus		Brazil: Rondonia, 1993	LACM
NVG-7929	Damas immacula		Costa Rica, 2010, 10-SRNP-67003	USNM
NVG-7349	Dardarina dardaris		Costa Rica, 2004, 04-SRNP-13073	USNM
NVG-18114D07	Daron seron sexton		Ecuador, 2001	USNM
NVG-7330	Darpa striata striata		Malaysia, old (around 1900)	USNM
NVG-17095C10	Decinea decinea decinea		Brazil: Parana, 1995	USNM
11-BOA-13385E10	Diaeus lacaena		Brazil: Rio de Janeiro, 1996	USNM
NVG-7893	Doberes anticus		Costa Rica, 2004, 03-SRNP-23600	USNM
NVG-18054F12	Dotta callicles		Namibia, 1992	ZMHB
NVG-17093B03	Dotta stellata stellata		Kenya, 1957	USNM
NVG-8038	Dubia dubia		Guyana, 1999	USNM
NVG-18012D08	Dubiella dubius		Peru, 1989	USNM
NVG-17098F01	Duroca duroca duroca		Brazil: Rio de Janeiro, 1996, USNMENT 00913432	USNM
NVG-17108B11	Eagris tigris		Kenya, 1985	LACM
NVG-15033A04	Eantis minna		Brazil: Amazonas, 1886	ZMHB
NVG-18025A03	Eantis minor	HT	Dominica, 1934	AMNH
NVG-10487	Eantis mithridates		Jamaica, 2017	UTSW
NVG-18025A06	Eantis munroei	HT	Cuba	AMNH
NVG-1931	Eantis pallida		Mexico: Tamaulipas, 1974	TAMU
NVG-3758	Eantis tamenund		USA: TX, Hidalgo Co., 2015	UTSW
NVG-1900	Eantis thraso		Brazil: Rondonia	TAMU
NVG-16108E06	Eantis tosta		Peru, 1999	USNM
NVG-18012A05	Ebrietas osyris		Mexico: Oaxaca, 1992	USNM
NVG-7977	Eburuncus unifasciata		Panama, 1984	USNM
NVG-7920	Ebusus ebusus		Costa Rica, 2013, 13-SRNP-30669	USNM
NVG-7884	Echelatus sempiternus		Costa Rica, 2007, 07-SRNP-12147	USNM
NVG-18013B09	Enosis dognini		Peru, 1998	USNM
NVG-5730	Entheus Burns01		Costa Rica, 2008, 08-SRNP-35619	USNM
NVG-18019B11	Eogenes alcides		Turkey, old (around 1900)	AMNH
NVG-17093E07	Ephyriades arcas		St. Croix, 1996	USNM
NVG-7897	Eracon sarahburnsae		Costa Rica, 2010, 10-SRNP-67957	USNM
NVG-7910	Erionota thrax		USA: HI, Molokai, 2005	USNM
NVG-18068D04	Erynnis tages		Russia: Siberia, 1999	EBrockmann
NVG-15103B05	Euschemon rafflesia rafflesia		Australia	USNM
NVG-17092F09	Eutocus facilisDHJ01		Costa Rica, 2006, 06-SRNP-47351	USNM
NVG-18082B04	Exometoeca nycteris		Australia, NHMUK_010430873, 0247277190	BMNH
NVG-18012E10	Falga jeconia jeconia		Venezuela, 1985	USNM
NVG-18031H12	Festivia adamantinus		Ecuador, 2002	USNM
NVG-2073	Festivia cronion		Brazil: Parana, 2011	MEM
NVG-18031H06	Festivia festiva		Ecuador, 1990	USNM
NVG-18032A03	Festivia grippa		Ecuador, 2002	USNM
NVG-19017F02	Fidius fido		Brazil: Parana, old (around 1900)	USNM
NVG-19017E10	Flaccilla aecas		Guyana, 2000	USNM
NVG-14101G07	Flattoides amazonensis amazonensis	HT	Colombia, 1946	AMNH
NVG-17092A07	Fresna netopha		Uganda, 1960	USNM
NVG-7762	Fulda coroller		Madagascar, 1991	USNM
NVG-7778	Fulda rhadama		Madagascar, 1990	USNM
NVG-7808	Galerga hyposticta		Madagascar, 1990	USNM
NVG-8020	Gallio gallio		Peru, 1986	USNM
NVG-16108E10	Gamia shelleyi		Uganda, 1953	USNM
NVG-16108F09	Gangara thyrsis		Philippines, 1987	USNM
NVG-18057H08	Ge geta		no data, 1894	ZSMC
NVG-7336	Gerosis bhagava		Myanmar, 2003	USNM
NVG-7570	Gesta gesta		Dominican Republic, 1981	TAMU
NVG-18011E10	Gindanes brebisson		Peru, 2016	USNM
NVG-7901	Gindanes brontinus		Costa Rica, 2008, 08-SRNP-57426	USNM
NVG-18067H12	Gindanes kelso		Peru, 2000	EBrockmann
NVG-18026G08	Ginungagapus schmithi	HT	Brazil: Santa Catarina	AMNH
NVG-18087G04	Gomalia elma		Botswana, 1997	EBrockmann
NVG-18062A03	Gorgopas chlorocephala chlorocephala		Peru, 2016	USNM
NVG-18019E12	Gorgyra aburae		Uganda, 1954	AMNH
NVG-17067D06	Gorgythion begga pyralina		Mexico: Sinaloa, 2003, CSU_ENT1039357	CSUC
NVG-18091C03	Gorgythion beggina escalophoides		Ecuador, 2012	EBrockmann
NVG-15101B07	Gorgythion plautia (=beggoides)	ST	Trinidad, Type No. 5981 U.S.N.M.	USNM
NVG-7880	Grais stigmaticus		Costa Rica, 2014, 14-SRNP-30242	USNM
NVG-18019F12	Gretna cylinda		Uganda, 1953	AMNH
NVG-18053D11	Gyrogra subnotata		Gabun, 1892	ZMHB
NVG-7979	Haemactis sanguinalis		Ecuador, 1991	USNM
NVG-18013A12	Halotus angellus		Panama, 1976	USNM
NVG-17119G01	Halpe porus		Myanmar, 2001	USNM
NVG-18052D09	Haza hazarma	LT	no data	ZMHB
NVG-15039D09	Helias phalaenoides phalaenoides		Brazil: Rondonia, 1993	FMNH
NVG-7558	Heliopetes arsalte		Mexico: San Luis Potosi, 1980	TAMU
NVG-18064E01	Herila herilus		Tanzania, 1951	USNM
NVG-18068A06	Hesperia comma		Russia: Yakutia, 1990	EBrockmann
NVG-4767	Hesperia meskei straton		USA: FL, Levy Co., 2015	UTSW
NVG-18026D09	Hesperia nabokovi	AT	Haiti, 1922	AMNH
NVG-8392	Hesperia viridis		USA: TX, Blanco Co., 2017	UTSW
NVG-17069D12	Hesperilla ornata		Australia, old (around 1900)	USNM
NVG-17067A10	Hesperopsis alpheus alpheus		USA: CA, Kern Co., 2012, CSU_ENT1039256	CSUC
NVG-17067B02	Hesperopsis gracielae		USA: CA, Riverside Co., 1997, CSU_ENT1039348	CSUC
NVG-17067A09	Hesperopsis libya libya		USA: CA, Inyo Co., 2009, CSU_ENT1039161	CSUC
NVG-17069E08	Heteropterus morpheus		France, 1966, USNMENT 00894386	USNM
NVG-17108D07	Hewitsoniella migonitis		Papua New Guinea, 1996	LACM
NVG-7823	Hidari irava		Singapore, 1989	USNM
NVG-18072H08	Hollandus xanthopeplus		Cameroon, 1895	ZMHB
NVG-18072H07	Hollandus xanthopeplus		Equatorial Guinea, 1906	ZMHB
NVG-7882	Hoodus pelopidas		Costa Rica, 2008, 08-SRNP-55556	USNM
NVG-7816	Hovala dispar-cf		Madagascar, 1988	USNM
NVG-7767	Hovala pardalina		Madagascar, 1988	USNM
NVG-7766	Hovala saclavus		Madagascar, 1988	USNM
NVG-3607	Hylephila phyleus		USA: TX, Starr Co., 2015	UTSW
NVG-18081B02	Hylephila venustus		Chile, 1960, NHMUK_010430840, 0247274697	BMNH
NVG-17091H02	Hypoleucis tripunctata draga		Uganda, 1960	USNM
NVG-14103B04	Iliana romulus		Peru	USNM
NVG-17091E06	Ilma irvina		Indonesia, old (around 1900)	USNM
NVG-18059B11	Incisus incisus		Brazil: Rio de Janeiro, 1995	USNM
NVG-18025A09	Isoteinon abjecta (=niangarensis)	HT	Niangara, 1910	AMNH
NVG-17091A07	Isoteinon lamprospilus formosanus		Taiwan, 1980	USNM
NVG-14107C02	Jera tricuspidata		Ecuador, 1984	USNM
NVG-7953	Joanna joanna		Costa Rica, 2004, 04-SRNP-14377	USNM
NVG-18013B06	Justinia justinianus		Guyana, 2000	USNM
NVG-18053B08	Katreus johnstonii apicalis		Sierra Leone, 1887	ZMHB
NVG-17108F10	Kedestes lepenula		South Africa, 1943	LACM
NVG-17069F06	Kobelana kobela		South Africa, 1978	USNM
NVG-18093B06	Ladda connexa	LT	Colombia	SMF
NVG-18014F02	Ladda eburones eburones		Peru, 2008	USNM
NVG-18014F01	Ladda monospila		Peru, 2010	USNM
NVG-18093B08	Ladda ochrolimbata	HT	Peru, old (around 1900)	SMF
NVG-18014F04	Ladda plancus		Peru, 2013	USNM
NVG-18017B09	Ladda puracensis cotopa		Ecuador, 1993	USNM
NVG-17111D10	Ladda pura-cf		Ecuador, 1992	LACM
NVG-18014E10	Ladda quadristriga		Peru, 2013	USNM
NVG-18017B07	Ladda seirocastnia		Ecuador, 1986	USNM
8041	Lamponia lamponia		Brazil: Parana, 1995	USNM
NVG-18012F02	Lento lento		Ecuador, 1998	USNM
NVG-18019G02	Leona leonora dux		Malawi, 1938	AMNH
NVG-17069C05	Leptalina unicolor		Japan, 1951	USNM
NVG-1769	Lerema accius		USA: TX, Dallas Co., 2013	USNM
NVG-7738	Lerema lineosa		Brazil: Mato Grosso, 1991	USNM
NVG-4062	Lerodea eufala		USA: TX, Dallas Co., 2015	UTSW
NVG-18054G05	Leucochitonea levubu		Namibia, 2002	ZMHB
NVG-17092D04	Librita librita		Mexico: Oaxaca, 1992	USNM
NVG-18067H07	Lindra simulius		Ecuador, 2012	EBrockmann
NVG-16108E11	Lissia lissa lima		Kenya, 1956	USNM
NVG-18056B11	Livida assecla		Brazil: Goias, 1929	ZfBS
NVG-3288	Lobocla liliana liliana		China: Yunnan, 2009	UTSW
NVG-18054C08	Lon azin		Colombia, 1920	ZMHB
NVG-6175	Lon hobomok hobomok		USA: VA, Augusta Co., 2016	UTSW
NVG-18115E11	Lon inimica		Panama, 2007	USNM
NVG-18113H06	Lon macneilli	HT	Colombia, 1975	USNM
NVG-17111H01	Lon melane melane		USA: CA, San Luis Obispo Co., 1994	LACM
NVG-18089H01	Lon monticola		Mexico: Queretaro, 2004	EBrockmann
NVG-18115G04	Lon niveolimbus		Mexico: Chiapas, 1992	USNM
NVG-9774	Lon taxiles		USA: AZ, Cochise Co., 2017	UTSW
NVG-9209	Lon zabulon		USA: AR, Montgomery Co., 2017	UTSW
NVG-7373	Lotongus calathus balta		Myanmar, 2003	USNM
8026	Lucida lucia		Brazil: Minas Gerais, 1994	USNM
NVG-7940	Ludens ludens		Costa Rica, 2012, 11-SRNP-33493	USNM
NVG-18015A10	Lurida lurida (=carinna)	T	Brazil: Parana, old (around 1900)	USNM
NVG-7987	Lycas argentea		Argentina, 1998	USNM
NVG-18111A07	Lycas godart boisduvalii		French Guiana, 1993	USNM
NVG-8009	Lychnuchoides ozias		Brazil: Rio de Janeiro, 1995	USNM
NVG-2076	Lychnuchus celsus		Brazil: Parana, 2011	MEM
NVG-5688	Marela tamba		Ecuador, 1991	USNM
NVG-7787	Matapa aria		Philippines, 1986	USNM
NVG-18014E07	Matapa cresta		Myanmar, 2001	USNM
NVG-1185	Megathymus yuccae yuccae		USA: SC, Aiken Co., 2013	UTSW
NVG-18081D08	Melphina melphis		Nigeria, 1958, NHMUK_010430880, 0247278457	BMNH
NVG-18074H11	Melphinyet tarace	T	Sierra Leone, 1889	ZMHB
NVG-17108A04	Methion melas		Guatemala, 1963	BMUW
NVG-18064B11	Methionopsis ina		Costa Rica, 2010, 10-SRNP-43176	USNM
NVG-18054G04	Metisella medea		Malawi, 1996	ZMHB
NVG-17093A02	Metisella metis paris		Uganda, 1958	USNM
NVG-18054G03	Metisella orientalis		Malawi, 1996	ZMHB
NVG-17091H09	Meza meza		Cameroon, old (around 1900)	USNM
NVG-7874	Mictris crispus		Costa Rica, 2009, 09-SRNP-71399	USNM
NVG-18093C03	Mielkeus diana diana (=seitzi)	LT	Brazil: Rio Grande do Sul, old (around 1900)	SMF
NVG-19022G01	Mielkeus klugi		Guyana, 1999	USNM
NVG-19022G03	Mielkeus lucretius		Brazil: Rio de Janeiro, 1995	USNM
NVG-18071B01	Mielkeus tertianus		Costa Rica, 2010, 10-SRNP-72549	USNM
NVG-7904	Milanion marciana		Costa Rica, 2005, 05-SRNP-41228	USNM
NVG-8043	Miltomiges cinnamomea		Brazil: Rio de Janeiro, 1995	USNM
NVG-7814	Miraja varians		Madagascar, 1988	USNM
NVG-17069F02	Misius misius		Guyana, 2001	USNM
NVG-17111C02	Mnasicles geta		Mexico: San Luis Potosi, 1981	LACM
NVG-7968	Mnasilus allubita		Costa Rica, 2002, 02-SRNP-13739	USNM
NVG-8030	Mnasinous patage		Panama, 1996	USNM
NVG-8027	Mnestheus ittona		Peru, 2012	USNM
NVG-8029	Moeris anna		Brazil: Rio de Janeiro, 1994	USNM
NVG-18012H11	Moeris striga		French Guiana, 1993	USNM
NVG-7941	Moeris stroma		Costa Rica, 2013, 13-SRNP-56538	USNM
NVG-15036C07	Moeros moeros	T	Suriname, 1874	ZMHB
NVG-18039F01	Molo mango		Guyana, 2003	FMNH
NVG-18012D03	Molo pelta		Peru, 2012	USNM
NVG-18019F10	Moltena fiara		Natal, 1924	AMNH
NVG-17092G12	Monca crispinus		Costa Rica, 2006, 06-SRNP-55847	USNM
NVG-17069H12	Monza cretacea		Nigeria, 1951	USNM
NVG-17069H09	Mopala orma		Cameroon, old (around 1900)	USNM
NVG-18019H04	Morvina morvus morvus		Colombia, 1945	AMNH
NVG-18013A08	Morys valerius		French Guiana, 1993	USNM
NVG-8048	Mucia zygia		Colombia, 1992	USNM
NVG-18041E02	Muschampia proto		France, 2012	EBrockmann
NVG-18015C07	Mylon ander ander		Guyana, 2000	USNM
NVG-18015D02	Mylon cajus cajus		Peru, 2016	USNM
NVG-18015C03	Mylon illineatus illineatus		Peru, 2016	USNM
NVG-7881	Mylon lassia		Costa Rica, 2009, 09-SRNP-36601	USNM
NVG-18013F07	Mylon maimon		Costa Rica, 2015, 15-SRNP-21203	USNM
NVG-18015C05	Mylon mestor		Colombia, 1992	USNM
NVG-18015C04	Mylon orsa		Panama, 1975	USNM
NVG-18013F09	Mylon salvia		Costa Rica, 2006, 06-SRNP-1769	USNM
NVG-18015C01	Mylon zephus albodiscus		Peru, 2008	USNM
11-BOA-13385F09	Myrinia myris		Brazil: Rondonia, 1989	USNM
NVG-7950	Naevolus orius		Costa Rica, 2010, 10-SRNP-72281	USNM
NVG-8137	Nastra lherminier		USA: FL, Liberty Co., 2017	UTSW
NVG-18053B01	Neohesperilla croceus		Australia, 1892	ZMHB
NVG-17095F04	Neoxeniades ethoda		Brazil: Santa Catarina, 1991	USNM
NVG-18069F03	Neoxeniades luda		Costa Rica, 2012, 12-SRNP-1402	USNM
NVG-18082E06	Neoxeniades musarion		Brazil: RJ, NHMUK_012824133, 0247279800	BMNH
NVG-7936	Neoxeniades pluviasilva		Costa Rica, 2012, 12-SRNP-30105	USNM
NVG-17112B12	Neoxeniades turmada		Peru, 1986	LACM
NVG-18113B11	Neposa heras		Mexico, old (around 1900)	BMNH
NVG-14063B05	Nerula fibrena		Venezuela, 1985	USNM
NVG-17091B06	Nervia nancy		Kenya, 1960	USNM
NVG-16106A03	Netrocoryne repanda		Australia, 1963	LACM
NVG-18053D01	Netrocoryne thaddeus		Indonesia, 1894	ZMHB
NVG-18013B02	Niconiades xanthaphes		Guyana, 2000	USNM
NVG-18011G09	Nisoniades mimas		Peru, 1982	USNM
NVG-18015E03	Noctuana lactifera lactifera		Costa Rica, 2003, 03-SRNP-22972	USNM
NVG-18086D12	Noctulana noctula		Ivory Coast, EL63203	MNHP
NVG-3845	Nyctelius nyctelius		USA: TX, Cameron Co., 2015	UTSW
NVG-15036C01	Nyctus crinitus	ST	Peru, old (around 1900)	ZMHB
NVG-17068C03	Oarisma poweshiek		USA: MN, Pipestone Co., 1986, CSU_ENT1025108	CSUC
NVG-14112G12	Ocella albata		Peru, no date	TLS
PAO-23	Ochlodes agricola		USA: CA, Sierra Co., 2016	UTSW
NVG-18039E08	Ochlodes subhyalina		Korea, 1956	FMNH
PAO-263	Ochlodes sylvanoides		USA: CO, Larimer Co., 2016	UTSW
NVG-7746	Ochlodes venata venata		Japan, 1933	USNM
NVG-17114B06	Ochlodes yuma		USA: CO, Mesa Co., 2001, LepNet 1024498	CSUC
NVG-5686	Oechydrus chersis chersis		Brazil: Rio de Janeiro, 1996, 14063A12	USNM
NVG-18014F09	Oenides vulpina		Peru, 2016	USNM
NVG-18021D06	Oeonus pyste		Mexico: San Luis Potosi, 1967	AMNH
NVG-14063C11	Oileides fenestratus		French Guiana, 1993, Hermier No 4771	USNM
NVG-18067C02	Oileides fenestratus		French Guiana, 2009	EBrockmann
NVG-18098F04	Oileides guyanensis		French Guiana, 2002, H19994	BHermier
NVG-18057C09	Oileides vulpinus		Brazil: Espirito Santo, old (around 1900)	ZSMC
NVG-5727	Oileides vulpinus		Brazil: Espirito Santo, 1969	USNM
NVG-15117B01	Onenses hyalophora		Mexico: Tamaulipas, 2003, CSU_ENT1039476	CSUC
NVG-17092D06	Onespa nubis		Mexico: Oaxaca, 1990	USNM
NVG-8040	Onophas columbaria		Ecuador, 2002	USNM
NVG-17109G08	Orphe gerasa		Venezuela, 1993	LACM
NVG-2761	Orphe vatinius		Colombia, 1976	JAScott
NVG-7956	Orses cynisca		Costa Rica, 2008, 08-SRNP-40358	USNM
NVG-18111A01	Orses itea		Brazil: Rio de Janeiro, 1995	USNM
NVG-8061	Orthos orthos		Brazil: Parana, 1991	USNM
NVG-18026A06	Orthos orthos hyalinus	HT	Brazil: Santa Catarina, old (around 1900)	AMNH
NVG-17119C02	Osmodes laronia		Uganda, 1961	USNM
NVG-18073E04	Osphantes ogowena	ST	Gabon, 1888	ZMHB
NVG-18011G05	Ouleus fridericus		Guyana, 2000	USNM
NVG-7894	Ouleus salvinaDHJ01		Costa Rica, 2011, 11-SRNP-57460	USNM
NVG-14113A02	Oxynetra aureopecta	HT	Mexico: Hidalgo, 1987	LACM
7962	Oxynthes coruscaDHJ02		Costa Rica, 2011, 11-SRNP-23410	USNM
NVG-7899	Paches loxus		Costa Rica, 2003, 03-SRNP-30995	USNM
NVG-18025B01	Pachyneuria lineatopunctata (=phintias)	HT	Peru, 1931	AMNH
NVG-15033D07	Pachyneuria obscura	T	Peru, old (prior to 1888)	ZMHB
NVG-4155	Panoquina panoquin		USA: TX, Jefferson Co., 2015	UTSW
NVG-17111G07	Papias subcostulata		Mexico: Hidalgo, 1981	LACM
NVG-8037	Paracarystus hypargyra		Peru, 2013	USNM
NVG-18063A09	Parachoranthus magdalia		Cuba, 2009	EBrockmann
NVG-17092H10	Paracleros biguttulus		Uganda, 1956	USNM
NVG-18027E08	Paramimus scurra scurra		Guyana, 2001	USNM
NVG-18015E01	Paratrytone rhexenor		Mexico: Veracruz, old (around 1900)	USNM
NVG-17108G05	Pardaleodes edipus		Zaire, 1981	LACM
NVG-18026E08	Pares pares	HT	Paraguay	AMNH
NVG-7290	Parnara guttatus		Myanmar, 2002	USNM
NVG-17108G12	Paronymus ligora		Sierra Lione, 1974	LACM
NVG-18013A05	Parphorus storax		Guyana, 2000	USNM
NVG-17094H03	Passova gellias		Costa Rica, 2013, 13-SRNP-6248	USNM
NVG-18082E01	Pastria pastria		Papua NG, 1982, NHMUK_010430882, 0247277187	BMNH
NVG-17116A09	Pellicia dimidiata dimidiata		Mexico: Tamaulipas, 1974	TAMU
NVG-18013C07	Penicula bryanti		Ecuador, 2001	USNM
NVG-18011F04	Penicula subviridis		Brazil: Santa Catarina, 1990	USNM
NVG-7955	Perichares adela		Costa Rica, 2013, 13-SRNP-65013	USNM
NVG-18065F12	Perichares butus		Peru, 2015	EBrockmann
NVG-18105G06	Perichares deceptus fulvimargo		Peru, 2013	USNM
NVG-18014F08	Perichares furcata		Ecuador, 1976	USNM
NVG-7986	Perichares haworthiana		Brazil: Rondonia, 1992	USNM
NVG-18105F10	Perichares lotus		Trinidad, 2000	USNM
NVG-18027A09	Perichares metallica (=zikani)	HT	Brazil: Rio de Janeiro	AMNH
NVG-10247	Perichares philetes		Jamaica, 2017	UTSW
NVG-18058H07	Perus coecatus (=tadus)	ST	Brazil: Rio de Janeiro	USNM
NVG-7826	Perus cordillerae		Peru, 1999	USNM
NVG-19055F06	Perus menuda	ST	Bolivia, 1899	MCZ
NVG-18059C09	Perus minor		Ecuador, 1988	USNM
NVG-17092F05	Phanes aletesDHJ02		Costa Rica, 2015, 15-SRNP-71060	USNM
NVG-18012F07	Pheraeus odilia epidius		Panama, 1982	USNM
NVG-18013C03	Phlebodes pertinax		Brazil: Amazonas, 1993	USNM
NVG-5316	Phocides pigmalion okeechobee		USA: FL, Monroe Co., 2015	UTSW
NVG-3990	Pholisora catullus		USA: TX, Starr Co., 2015	UTSW
NVG-7976	Pholisora mejicanus		USA: NM, Colfax Co., 1989	USNM
NVG-17068A11	Piruna aea mexicana		USA: AZ, Santa Cruz Co., 2016, CSU_ENT1033276	CSUC
NVG-6454	Piruna pirus		USA: CO, Grand Co., 2016	UTSW
NVG-17108G10	Platylesches picanini		South Africa, 1944	LACM
NVG-7982	Plumbago plumbago		Brazil: Rondonia, 1989	USNM
NVG-939	Poanes aaroni bordeloni		USA: LA, 2011	UTSW
NVG-4704	Poanes aaroni howardi		USA: FL, Levy Co., 2015	UTSW
NVG-17114B07	Poanes massasoit		USA: MD, Dorchester Co., 1976, LepNet 1031028	CSUC
NVG-6711	Poanes viator zizaniae		USA: TX, Dallas Co., 2016	UTSW
NVG-7046	Poanes yehl		USA: TX, Hopkins Co., 2016	UTSW
NVG-8170	Polites baracoa baracoa		USA: FL, Miami-Dade Co., 2017	UTSW
NVG-5968	Polites carus		USA: AZ, Santa Cruz Co., 2016	UTSW
NVG-4276	Polites peckius peckius		USA: IN, Montgomery Co., 2015	UTSW
NVG-4255	Polites themistocles themistocles		USA: IN, Montgomery Co., 2015	UTSW
NVG-7875	Polyctor polyctor		Costa Rica, 2012, 12-SRNP-4870	USNM
NVG-18115C01	Pompeius amblyspila		Brazil: Amazonas, 1993	USNM
NVG-17106B02	Pompeius pompeius		Costa Rica, 2010, 10-SRNP-103576	USNM
NVG-18038D12	Potamanaxas flavofasciata flavofasciata		Ecuador, 2008	EBrockmann
NVG-18082C08	Prada rothschildi	PT	Papua NG, 1899, NHMUK_012824115, 0247281590	BMNH
NVG-18082C09	Prada rothschildi	PT	Papua NG, 1899, NHMUK_010430826, 0247278442	BMNH
NVG-18114H04	Propapias sipariana		French Guiana, 1993	USNM
NVG-7331	Pseudocoladenia dan fabia		Myanmar, 2001	USNM
NVG-8050	Pseudocopaeodes eunus		USA: CA, Inyo Co., 1950	USNM
11-BOA-13382A03	Pseudodrephalys hypargus		Ecuador	USNM
NVG-18113B05	Pseudorphe pyrex		Peru, 1992	USNM
NVG-17112C01	Pseudosarbia phoenicicola		Uruguay, 1967	LACM
NVG-18021B02	Psoralis idee		Bolivia, old (around 1900)	AMNH
NVG-7757	Pteroteinon laufella		Liberia, 1988	USNM
NVG-7771	Pyrgus malvae		Greece, 1992	USNM
NVG-8060	Pyrrhocalles antiqua		Dominican Republic, 1994	USNM
NVG-17094C09	Pyrrhopyge hadassa pseudohadassa		Peru, 2013, USNMENT 00894886	USNM
NVG-7984	Pyrrhopygopsis socrates		Brazil: Mato Grosso, 1991	USNM
NVG-7896	Pythonides amaryllis		Costa Rica, 2006, 06-SRNP-7674	USNM
11-BOA-13382B01	Pythonides jovianus jovianus		Guyana, 2000	USNM
NVG-14102D02	Pythonides lerina		Guyana, 2003	FMNH
NVG-7902	Quadrus cerialis		Costa Rica, 2008, 08-SRNP-1186	USNM
NVG-18013H08	Quadrus lugubris		Costa Rica, 2008, 08-SRNP-57856	USNM
NVG-18018A03	Quadrus truncata		Ecuador, 1988	USNM
NVG-18015B10	Quasimellana mexicana		Mexico: Oaxaca, 1989	USNM
NVG-7959	Quinta cannae		Costa Rica, 2012, 12-SRNP-75508	USNM
NVG-18057H02	Rachelia extrusus		Papua New Guinea, old (around 1900)	ZSMC
NVG-18012A12	Racta plasma		Peru, 2011	USNM
NVG-17092G01	Radiatus bradus		Guyana, 1999	USNM
NVG-18091D12	Ralis concolor		Ecuador, 2012	EBrockmann
NVG-19021E08	Ralis coyana		Brazil: Rio de Janeiro, 1996	USNM
NVG-17092C08	Remella remus		Costa Rica, 2006, 06-SRNP-6640	USNM
NVG-18081A02	Repens repens	PT	Paraguay, 1904, NHMUK_010430831, 0247279233	BMNH
NVG-17069G04	Rhabdomantis galatia		Uganda, 1958	USNM
NVG-17116C08	Rhinthon osca		USA: TX, Hidalgo Co., 1974	TAMU
NVG-18093C02	Rigga auristriga	HT	Bolivia, old (around 1900)	SMF
NVG-19019G04	Rigga hesia		Ecuador, 2002	USNM
NVG-18093C05	Sacrator sacrator (=stupenda)	HT	Colombia, old (around 1900)	SMF
NVG-18012E03	Saliana salius		Guyana, 2000	USNM
NVG-8024	Saniba sabina		Brazil: Rio de Janeiro, 1996	USNM
NVG-18025B08	Santa santes	HT	Peru, 1924	AMNH
11-BOA-13382F10	Santa trifasciatus		Guyana	USNM
NVG-7345	Sarangesa dasahara		Myanmar, 2001	USNM
NVG-18037G08	Satarupa nymphalis		China: Sichuan, 2010	UTSW
NVG-18013C01	Saturnus saturnus		Guyana, 2000	USNM
NVG-7803	Semalea pulvina		Cameroon, 1989	USNM
NVG-7760	Signeta flammeata		Australia, 1985	USNM
NVG-7970	Sodalia sodalis		Costa Rica, 2011, 11-SRNP-75366	USNM
NVG-14107C10	Sophista aristoteles aristoteles		Peru, 2013	USNM
NVG-18011H10	Sostrata bifasciata		Brazil: Rio de Janeiro, 1995	USNM
NVG-7247	Sostrata nordica		Mexico: Veracruz, 1906	USNM
NVG-18032A04	Sostrata pusilla pulsa		Ecuador, 1976	USNM
NVG-18032A10	Sostrata pusilla pusilla		Peru, 1989	USNM
NVG-17108E02	Spialia galba galba		India, 1961	LACM
NVG-18013H11	Spioniades artemides		Costa Rica, 2010, 10-SRNP-67995	USNM
NVG-18071A04	Staphylus ascalaphus		Costa Rica, 2016, 16-SRNP-56091	USNM
NVG-18071A05	Staphylus caribbea		Costa Rica, 2009, 09-SRNP-42728	USNM
NVG-9694	Staphylus ceos		USA: AZ, Santa Cruz Co., 2017	UTSW
NVG-18025B10	Staphylus musculus (=similis)	HT	Brazil: Sta Catarina, old (around 1900)	AMNH
NVG-18025B03	Staphylus putumayo	HT	Peru, 1931	AMNH
NVG-18011H08	Staphylus vincula		Mexico: Oaxaca, 1988	USNM
NVG-18071A12	Staphylus vulgata		Costa Rica, 2010, 10-SRNP-55088	USNM
NVG-6016	Stinga morrisoni		USA: TX, Jeff Davis Co., 2016	UTSW
NVG-8044	Styriodes lyco		Guyana, 2000	USNM
NVG-18012D04	Synale hylaspes		Argentina, 1998	USNM
NVG-7937	Synapte salenus salenus		Costa Rica, 2007, 07-SRNP-21744	USNM
NVG-3621	Systasea pulverulenta		USA: TX, Duval Co., 2015	UTSW
NVG-6008	Systasea zampa		USA: TX, El Paso Co., 2016	UTSW
NVG-18011E02	Systaspes corrosus		Mexico: Oaxaca, 1992	USNM
NVG-7333	Tagiades litigiosus litigiosus		Myanmar, 2001	USNM
NVG-18012D01	Talides sinois		Peru, 2015	USNM
NVG-7375	Taractrocera maevius sagara		Myanmar, 2003	USNM
NVG-18116E01	Tava tavola	ST	Trinidad, old (around 1900)	USNM
NVG-18026A03	Tava tavola (=hoffmanni)	HT	Brazil: Santa Catarina, 1913	AMNH
NVG-5726	Telemiades fides		Costa Rica, 2011, 11-SRNP-20768	USNM
NVG-8008	Tellona variegata		Brazil: Rondonia, 1993	USNM
NVG-17119H08	Teniorhinus watsoni watsoni		Uganda, 1961	USNM
NVG-18026F01	Testia potesta	HT	Peru, 1931	AMNH
NVG-18098F08	Thargella caura		French Guiana, 2005, H22106	BHermier
11-BOA-13386C12	Theagenes aegides		Costa Rica, 1980	USNM
11-BOA-13386C11	Theagenes albiplaga		Peru, 2008	USNM
NVG-18013B04	Thoon modius		Ecuador, 1992	USNM
NVG-7383	Thoressa masoni		Myanmar, 2001	USNM
NVG-17112B06	Thracides phidon		Venezuela, 1993	LACM
NVG-18022H12	Thymelicus acteon acteon		Spain, 1953	AMNH
NVG-7888	Tiana niger		Costa Rica, 2011, 11-SRNP-35371	USNM
NVG-7944	Tigasis arita		Costa Rica, 2011, 11-SRNP-32281	USNM
NVG-18013A11	Tigasis zalates		Ecuador, 1977	USNM
NVG-7883	Timochares trifasciata		Costa Rica, 2005, 05-SRNP-12097	USNM
NVG-7908	Timochreon satyrus		Costa Rica, 2007, 07-SRNP-58884	USNM
NVG-18118A08	Tirynthia conflua		Brazil: Rio de Janeiro, 1995	USNM
NVG-18081D03	Toxidia thyrrhus		Australia, 1911, NHMUK_010430805, 0247281664	BMNH
NVG-16106A10	Trapezites symmomus		Australia, 1963	LACM
NVG-19023C11	Tricrista advena		Guyana, 2000	USNM
8049	Tricrista crista		Guyana, 2000	USNM
NVG-18025G05	Tricrista cristatus	HT	Brazil: Santa Catarina, old (around 1900)	AMNH
NVG-7983	Trina geometrina		French Guiana, 1993	USNM
NVG-18012D02	Tromba tromba		Peru, 2014	USNM
NVG-18019E10	Tsitana tsita		South Africa, 1924	AMNH
NVG-18111G06	Turesis complanula		Guyana, 2000	USNM
NVG-18082D06	Turmosa camposa		Brazil: RJ, 1883, NHMUK_012824124, 0247279797	BMNH
NVG-18056F01	Turmosa camposa	LT	no data	ZSMC
NVG-5699	Typhedanus ampyx		Mexico: Oaxaca, 1992, 14104H07	USNM
NVG-17108H12	Unkana ambasa		no data, no date	LACM
NVG-18081C02	Unkana mytheca		Indonesia, 1914, NHMUK_010430823, 0247278996	BMNH
NVG-4894	Urbanus proteus proteus		USA: FL, Miami-Dade Co., 2015	UTSW
NVG-17095C06	Vacerra litana		Venezuela, 1975	USNM
NVG-17098F07	Veadda veadeira		Brazil: Mato Grosso, 1991	USNM
NVG-17111G04	Vehilius stictomenes illudens		Mexico: San Luis Potosi, 1980	LACM
NVG-8022	Venas evans		Guyana, 2000	USNM
NVG-18021D01	Vernia dares		Mexico: San Luis Potosi, 1966	AMNH
NVG-18014H01	Vernia verna		USA: OH, Summit Co., 2012	USNM
NVG-18013B11	Vertica verticalis		Peru, 1983	USNM
NVG-18014G01	Vettius phyllus phyllus		Guyana, 2001	USNM
NVG-18071C05	Vettius picaDHJ01		Costa Rica, 2015, 15-SRNP-71571	USNM
NVG-18012H07	Vidius vidius		Paraguay, old (around 1900)	USNM
NVG-15097D10	Vinpeius tinga (=freemani)	HT	Mexico: Veracruz	CMNH
NVG-18011G11	Viola alicus	ST	Brazil: Rio de Janeiro, old (around 1900)	USNM
NVG-18061D04	Viola minor		Brazil: Rio de Janeiro, 1995	USNM
NVG-7973	Viola violella		Brazil: Mato Grosso, 1991	USNM
NVG-18012F11	Virga virginius		Paraguay, 1986	USNM
NVG-18026H05	Viridina subviridis	HT	Ecuador, 1938	AMNH
NVG-19055F04	Viridina viridenex	HT	Bolivia, 1899	MCZ
NVG-15104C11	Viridina viridis	HT	Ecuador, 1938	AMNH
NVG-18028C11	Viuria licisca		Costa Rica, 2003, 03-SRNP-27671	USNM
NVG-7972	Viuria lista		French Guiana, 1988	USNM
NVG-17107B06	Wahydra kenava		Venezuela, 1978	USNM
NVG-7421	Wallengrenia drury		Dominican Republic, 1981	USNM
NVG-17098E07	Wallengrenia ophites		French Antilles, 1989	USNM
NVG-17098E12	Wallengrenia premnas		Brazil: Mato Grosso, 1990	USNM
NVG-10332	Wallengrenia vesuria		Jamaica, 2017	UTSW
NVG-18079C02	Willema tsadicus (=birbiranus)	T	Ethiopia, 1925, EL63085	MNHP
NVG-18063B11	Willema willemi		Tanzania, old (around 1900)	ZSMC
NVG-17069E09	Willema willemi		Zimbabwe, 1961	USNM
NVG-15102D03	Windia windi		Mexico: Sonora, 1984	USNM
NVG-7765	Xanthodisca vibius		Cameroon, 1987	USNM
NVG-17091D05	Xanthoneura corissa corissa		Malaysia, old (around 1900)	USNM
NVG-18064D07	Xanthonymus xanthioides		Cameroon, old (around 1900)	USNM
NVG-17112A11	Xeniades orchamus		Brazil: Santa Catarina, 1999	LACM
NVG-7906	Xenophanes tryxus		Costa Rica, 2010, 10-SRNP-103428	USNM
NVG-15033D08	Xispia quadrata	T	Brazil: Amazonas, prior to 1889	ZMHB
NVG-17105B08	Zalomes biforis		Colombia, 1965	USNM
NVG-18074B01	Zela excellens	T	Palawan, 1888	ZMHB
NVG-18098G05	Zela smaragdinus		Malaysia, 2016	UTSW
NVG-18126A12	Zela zenon		Sabah, 1984	KMaruyama
NVG-18126A05	Zela zeus optima		Langkawi, 2018	UTSW
NVG-17112A04	Zenis jebus hemizona		Venezuela, 1993	LACM
NVG-18038B10	Zera hosta		Honduras	RGallardo
NVG-18091C05	Zera phila		Ecuador, 2012	EBrockmann
NVG-18011E12	Zera zera		Peru, 2014	USNM
NVG-18027A07	Zetka zeteki	HT	Panama, 1928	AMNH
NVG-15104C01	Zobera albopunctata	HT	Mexico: Colima	AMNH
NVG-17091C04	Zographetus satwa		no data	USNM
NVG-17069H08	Zophopetes dysmephila		South Africa, 1953	USNM
NVG-6998	Zopyrion sandace		Mexico: Oaxaca, 1981	TAMU
NVG-1670	Pterourus glaucus glaucus		USA: TX, Denton Co., 2013	USNM

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Due to this very large number of specimens we have sequenced, the phylogenetic trees were built for smaller phylogenetic groups (subfamilies, tribes and subtribes) and analyzed together with the “backbone” tree for the entire family constructed with selected reference species and given in Li et al. (2019). Type species (or their close relatives when the type is not available) of available genus-level names (including the names treated as synonyms) were marked on the trees to ensure that any available name for each clade is used. Manual analysis of the trees involved searching for non-monophyletic genera and checking clades that are prominently separated from others. Both nuclear genome and mitogenome trees were checked side-by-side to evaluate the consistency between them. Statistical support values (bootstrap) were taken into consideration to judge the validity of each observed clade. When a clade without an available name was found, identification of specimens in the clade was confirmed by the analysis of their wing patterns and genitalia. These new clades and the genus of former placement of each species proposed as a type species of a new genus name were rationalized in terms of genitalic morphology and wing patterns to search for diagnostic characters in phenotype. Finally, genitalia and wing patterns were used to decide the generic placement of species for which DNA data were not available.

Diagnostic DNA characters were identified in nuclear genomic sequences using our recently published procedure (see SI Appendix to Li et al. 2019). The positions in exons were found that are most likely synapomorphic to the clade defined as a genus. For the clades where we had several species sequenced, positions that are invariant in all species from this clade and have a base pair different from a (mostly invariant) base pair in the outgroups were found and those with the smallest number of species with missing data were selected. If a genus has only one species sequenced, it is difficult to distinguish between characters of the genus and characters of the species. Therefore, we frequently resorted to a different method of defining DNA characters of a genus that would increase the robustness of these characters. First, we looked for characters to define the sister clade of the genus. Sister clades usually included more than one species, and thus characters for the sister clade were better defined. We find the characters for the sister clade and take their states that differ from those in the sister clade as diagnostic for the genus in question. Second, we found synapomorphic characters for the clade that leads to the common ancestor of the genus and its sister clade. We used the combination of these latter characters with the diagnostic ones to define the genus. Such a treatment increases the chances that the character found is not a random non-conserved change or a sequencing error. Number of sequence reads covering this position was taken into account in choosing the characters, and those positions with better coverage were given priority. The character states are given in diagnoses below as abbreviations. For example, aly728.44.1:G672C means position 672 in exon 1 of gene 44 from scaffold 728 of the Cecropterus [formerly Achalarus] lyciades (aly) reference genome (Shen et al. 2017) is C, changed from G in the ancestor. When characters were found for the sister clade of the diagnosed taxon, the following statement was used: aly5294.20.2:A548A (not C), which means that position 547 in exon 2 of gene 20 on scaffold 5294 is occupied by the ancestral base pair A, which was changed to C in the sister clade (so it is not C in the diagnosed taxon). The sequences of exons from the reference genome with the positions used as character states highlighted in green are given in the supplemental file deposited at https://osf.io/5cfht/?view_only=21eb53b6f8f344afaee3de2be90bf5d2. Distribution of these sequences together with this publication ensures that the numbers given in the diagnoses can be easily associated with actual sequences. This publication has been registered with ZooBank as http://zoobank.org/BA35690A-FC73-4E5A-A805-FE9550275FEC and individual ZooBank registration numbers for each new name are given below.

The specimens were examined and sampled for sequencing in the following collections (abbreviations in parenthesis, used in Table 1 and Table S1 in the Supplemental file deposited at https://osf.io/5cfht/?view_only=21eb53b6f8f344afaee3de2be90bf5d2): American Museum of Natural History, New York, NY, USA (AMNH), Natural History Museum, London, UK (BMNH), Burke Museum of Natural History and Culture, Seattle, WA, USA (BMUW), Carnegie Museum of Natural History, Pittsburgh, PA, USA (CMNH), Colorado State University Collection, Fort Collins, CO, USA (CSUC), The Field Museum of Natural History, Chicago, FL, USA (FMNH), Los Angeles County Museum of Natural History, Los Angeles, CA, USA (LACM), Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA (MCZ), Mississippi Entomological Museum, Starkville, MS, USA (MEM), Muséum National d’Histoire Naturelle, Paris, France (MNHP), Natural History Museum, Frankfurt, Germany (SMF), Texas A&M University Insect Collection, College Station, TX, USA (TAMU), National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (USNM), University of Texas Southwestern, freezers of the Grishin lab, Dallas, TX, USA (UTSW), Zentrum fur Biodokumentation des Saarlandes, Schiffweiler, Germany (ZfBS), Museum für Naturkunde, Berlin, Germany (ZMHB), Zoologische Staatssammlung München, Germany (ZSMC), and research collections of Jim P. Brock (JPBrock), Ernst Brockmann (EBrockmann), Robert Gallardo (RGallardo), Bernard Hermier (BHermier), Kiyoshi Maruyama (KMaruyama) James A. Scott (JAScott), Texas Lepidoptera Survey, Houston, TX, USA (TLS, since then donated to McGuire Center for Lepidoptera and Biodiversity, Gainesville, FL, USA, MGCL), and Mark Walker (MWalker).

Results and Discussion

Analysis of the trees revealed rampant inconsistencies in the current classification of Hesperiidae into genera. These inconsistencies were not evenly distributed among phylogenetic lineages of Hesperiidae. Fewer problems were observed among the Old World taxa due to better knowledge about them, and maybe due to their smaller number. Most problems involved non-monophyletic genera with one or more clades of the former genus not having an available name. In several instances, a genus remained monophyletic as currently circumscribed but was prominently divided into two clades. Genetic diversification within each clade was comparable to that in other genera. For instance, the COI barcode difference taken as a proxy for diversification was typically below 10%, and the distances in nuclear and mitochondrial genomic DNA trees were about the same as in other species-rich genera. However, the distance between the clades was prominently larger than within each clade, typically more than 10% identity in the barcode.

These clades are named here and the following are the standardized descriptions of new taxa found during this analysis. The names were chosen to be simple and short, and mostly either reflect names or properties of their type species to facilitate memorization, or are fusions of genus names, euphonized and shortened. For each genus, a ZooBank registration number is given. The type species name is listed in its original genus combination and spelling, followed by the author and year the name was made available (not a bibliographic reference, but part of the name). Definition section indicates closest genera, mostly sisters or a group of genera if there are no clear sister genera, states the generic placements prior to this study (type species are given where appropriate to help assign a clade to a genus), gives reference to diagnostic characters as they are given in previous publications mostly in the Evans volumes (Evans 1937, 1949, 1951, 1952, 1953, 1955). It was almost always possible to trace the genus observed in the phylogenetic trees to the morphological characters given in the Evans’ identification keys. We think that referencing the keys rather than comprehensively listing all the characters would facilitate identification. Nevertheless, a brief morphological diagnosis for each genus is given, summarizing the most prominent phenotypic traits. DNA characters found by our recently developed method (Zhang et al. 2019c) are given at the end of each definition. This method should increase the robustness of the generic-level characters allowing for addition of species to a genus once they are sequenced. Then the gender of the name and an explanation about its origin is provided, species placed in the genus are listed (in their original genus-species name combinations with authors and dates), a parent taxon (mostly a subtribe, tribe or subfamily per Li et al. (2019)) is given, sometimes followed by comments about species involved and proposed additional taxonomic changes.

All of the changes to taxonomic status of names are propagated to all names currently treated as subspecies (for species), subgenera (for genera) and synonyms of these taxa. For instance, we do not list subspecies and synonyms unless changes are made to their status, and we assume that these names go with the species they were previously placed with. Finally, taxa not mentioned in this work are considered to remain at the ranks and in taxonomic groups they have been previously assigned to (Evans 1937, 1949, 1951, 1952, 1953, 1955; Mielke 2005; Li et al. 2019; Zhang et al. 2019b).

Furthermore, several tribes and subtribes would no longer be monophyletic if the higher-level classification remained unchanged. To solve this problem, we establish two new tribes and two new subtribes, which are described here. The following sections follow the standardized format and are either new taxa descriptions or taxonomic changes to existing taxa, as stated in the titles of these sections. Sections dealing with family-group names are given first, so that the new names can be used in the following sections dealing with genus-group names. Then, the sections are arranged in phylogenetic order approximating how these taxa appear in the figured trees.

Ceratrichiini Grishin, new tribe

http://zoobank.org/FC72EB8A-495D-4439-AA53-C310CD0735DE

Type genus.

Ceratrichia Butler, [1870].

Definition.

Placed near the mostly African tribe Astictopterini Swinhoe, 1912, this tribe is a sister to all Hesperiinae except Aeromachini Tutt, 1906. Backed by the maximal statistical support in all trees (Fig. 1), this clade of closely related genera keys to VI.A.(b) or VII.B.(a)(a¹) in Evans (1937). Antennae long, longer than half of costa, 2nd segment of palpi directed up, forewing vein CuA₂ originates in the middle of discal cell, end of discal cell straight, hindwing vein M₂ prominent, originates closer to vein M₁ than M₃, vein CuA₂ originates before or opposite to RS, vein 3A long. Male genitalia simple in most species: uncus undivided (except Herila), terminally narrows to a point, narrow in lateral view, gnathos either short or lacking, valva 2–3 times longer than wide, without prominent elaborations, may have small teeth on harpe. In DNA, a combination of the following base pairs is diagnostic: aly1121.3.2:A429G, aly669.27.2:A50G, aly374.13.3:A242T, aly216.78.1:A568C, aly1155.14.6:T406G.

A sister genus to Oileides Hübner, [1825] (type species Oileides vulpinus Hübner, [1825]), where C. amazonensis was placed previously (Fig. 2). However large genetic distance between O. vulpinus and O. amazonensis (COI barcodes differ by 12.3%, the difference is typically smaller than 10% for species within a genus) argues for their distinction. Keys to D.9.4 in Evans (1952). Genitalia illustrated by Bell in his fig. 2 (Bell 1947). Distinguished from its relatives by the following combination of characters: shorter than tegumen and broad uncus with small (not longer than wide) arms; prominent gnathos not shorter than uncus; valva gradually curved dorsad, ending with two pointed teeth; tuft of apricot-colored modified scales in a groove along the vein near the base of ventral hindwing in males; forewing with a macular white band from mid-costa to tornus. In DNA, a combination of the following base pairs is diagnostic: aly8661.4.1:A916G, aly10226.44.1:C1975G, aly709.1.2:T194C, aly36556.1.1:T2416T (not A), aly274.33.1:A188A (not G), aly82.28.5:T398T (not G).

Figure 2. — Genomic trees of Eudaminae and representatives of other Hesperiidae subfamilies. The trees are built from protein-coding regions in different genomic partitions: a) Nuclear genome. b) Z chromosome. c) Mitochondrial genome; d) COI barcode. The trees are rooted with *Pterourus glaucus* (NVG-1670), not shown. See Table 1 and Table S1 in the Supplemental file deposited at https://osf.io/5cfht/?view_only=21eb53b6f8f344afaee3de2be90bf5d2 for additional data about these specimens. Where possible, taxa are ordered similarly to that in the nuclear genome tree. Names of species placed in new genera described in this work are highlighted in yellow and clades of these genera are colored in red. Clades for genera where these species were placed previously are colored in blue and green arrow points from the former genus to the new genus (only on nuclear genome tree). Names of genera resurrected from synonymy are highlighted in cyan. Names of genera that form new genus-species combinations proposed in this work are highlighted in magenta. Statistical support values are shown by nodes in all but the COI barcode trees. COI barcode NJ dendrogram is given for comparison and is not expected to reflect phylogeny. The same notations are used in Fig. 2–13.

Etymology.

The name is a masculine noun in the nominative singular, for the type species that resembles butterflies from the genus Celaenorrhinus known as “Flats” in some English-speaking countries.

Species included.

Only the type species.

Parent taxon.

Subtribe Oileidina Grishin, 2019.

Aurivittia Grishin, new genus

http://zoobank.org/C2C0299C-67C8-4F0E-ACB8-29A7292E10BD

Type species.

Plesioneura aurivittata Moore, 1878.

Definition.

Surprisingly, a possible sister genus to Alenia Evans, 1935 (type species: Pyrgus sandaster Trimen, 1868), and in the same clade with Apallaga Strand, 1911 (type species Apallaga separata Strand, 1911), but genetically far removed from these and all other taxa (Fig. 3). Species in this genus were formerly placed in Celaenorrhinus Hübner, [1819] (type species Papilio eligius Stoll, [1781];) and key to B.6.30 in Evans (1949), notably lacking hair pencil on hind tibiae, which other Celaenorrhinus-like taxa possess. Morphologically, distinguished from Celaenorrhinus (sensu stricto) by essentially undivided uncus, sometimes with small knobs (not processes) as arms and the lack of hair pencil on hind tibiae; and from Alenia (a genus with undivided uncus) by the shape of valva that is not terminally split like a crab claw, and the presence of yellow band across the forewing. In male genitalia most similar to Apallaga, a genus with many species characterized by undivided uncus, but differs in longer gnathos (not less than half of the uncus length), thicker penis, valva with more robust and broad harpe that is gradually curved dorsad (not ventrad), does not carry any processes and is not forked, but narrows to a single point, and ampulla with a long process (style) along the harpe. In wing patterns, hindwing is not prominently variegated dorsally and is mostly unmarked, forewing with a compact yellow discal band with regular edges that does not extend into the discal cell and frequently with a triplet of apical yellow spots. In DNA, a combination of the following base pairs is diagnostic: aly2532.2.1:T488A, aly997.12.1:C310A, aly527.19.4:T178A, aly822.15.1:C589A, aly235.8.18:G1178A.

Figure 3. — Genomic trees of Tagiadinae. See Fig. 2 legend for notations.

Etymology.

The name is a feminine noun in the nominative singular, formed to reflect the golden stripe on the forewing, similar to the name of the type species.

Species included.

The type species, Plesioneura cameroni Distant, 1882, and Celaenorrhinus viet-namicus Devyatkin, 1998.

Parent taxon.

Tribe Celaenorrhinini Swinhoe, 1912.

Comments.

These south Asian butterflies are not so prominently distinct from the striped species of Celaenorrhinus. Therefore, their uniqueness was not apparent prior to our genomic study, although genitalic features revealed a posteriori can diagnose the genus morphologically. Phylogenetic placement of this Asian genus in the African clade of Celaenorrhinini Swinhoe, 1912 is interesting.

Viuria Grishin, new genus

http://zoobank.org/1D91E55F-38B7-4E1E-997F-49FBFD10CE02

Type species.

Pellicia licisca Plötz, 1882.

Definition.

A sister genus to Viola Evans, 1953, distant from Pachyneuria Mabille, 1888 (type species Pachyneuria obscura Mabille, 1888), where species of this genus were formerly placed (Fig. 4). Keys to E.20.7 in Evans (1953). Differs from Pachyneuria by the presence of long tufts of scales near hindwing costa above in males and swollen hindwing vein Sc+R₁, similar to Viola and Nisoniades Hübner, [1819], among others. Differ from Viola and other genera in genitalia and diagnosed by asymmetric uncus with asymmetric processes, reduced harpe of the right valva compared to the expanded rounded ampulla. In DNA, a combination of the following base pairs is diagnostic: aly1222.46.1:A28C, aly207.4.6:C1025G, aly1019.7.7:T59A, aly2012.7.7:C98G, aly638.27.5:T694A.

The name is a masculine noun in the nominative singular, derived from Peru, the locality of the type species and the county where many other species of this genus occur.

Species included.

The type species, Staphylus mossi Evans, 1953, Staphylus minor Schaus, 1902, Antigonus coecatus Mabille, 1891, and Nisoniades menuda Weeks, 1902.

Parent taxon.

Tribe Carcharodini Verity, 1940.

Tosta Evans, 1953 is a synonym of Eantis Boisduval, 1836

Phylogenetic analysis confidently reveals (Fig. 5) that Tosta Evans, 1953 (type species Tosta tosta Evans, 1953) originates within Eantis Boisduval, [1836] (type species Urbanus thraso Hübner, [1807]) and therefore is best considered its junior subjective synonym.

Figure 5. — Genomic trees of Achlyodini. See Fig. 2 legend for notations.

Eantis pallida (R. Felder, 1869), new combination

In all trees, Helias pallida R. Felder, 1869 is not monophyletic with Papilio busirus Cramer, [1779], the type species of Achlyodes Hübner, [1819], but originates within Eantis Boisduval, [1836] (type species Urbanus thraso Hübner, [1807]) (Fig. 5). Therefore, H. pallida is placed in this genus to form a new combination: Eantis pallida.

Pythonides assecla Mabille, 1883, reinstated status

We consider Pythonides assecla Mabille, 1883 to be a species distinct from Pythonides grandis Mabille, 1878 due to difference in size and extent of the blue coloration of the hindwing, both in males and females, and differences in shade of blue, being paler in P. grandis.

Livida Grishin, new genus

http://zoobank.org/24179CB6-20EF-4DF2-B545-CFF25554444C

Type species.

Pythonides assecla Mabille, 1883.

Definition.

A genus clearly near Gindanes Godman and Salvin, 1895, Pythonides Hübner, [1819], Quadrus Lindsey, 1925, Zera Evans, 1953 and Ouleus Lindsey, 1925, but without closer affinity to any one of them (Fig. 5). The species included here in this genus were formerly placed in Pythonides, but are not monophyletic with the type species of this genus, Papilio jovianus Stoll, 1782. Keys to E.41.5 in Evans (1953). Morphologically, distinguished from related genera by the combination of the shape of uncus (flattened at the tip with a short and thin central projection) with the shape of valva (broad at the base with concave costa and diamond-shaped harpe longer than the rest of the valva, with serrated dorsoposterior edge), only one upper spur on hind tibiae, the lack of tuft in hind tibiae and thoracic pouch in males, ventral hindwing mostly pale blue, dorsal hindwing pale blue in males at least near tornus. In DNA, a combination of the following base pairs is diagnostic: aly1672.3.1:T128G, aly1672.3.1:C110G, aly2618.5.1:C3626G, aly140.1.2:T1822T (not A), aly1349.7.9:C1010C (not G), aly1877.13.1:A785A (not C), aly23605.1.46:T5641T (not A), aly23605.1.46:C5642C (not G), aly536.102.1:C56C (not A), aly1935.2.1:G242G (not C), aly345.3.1:C328C (not G), aly767.12.13:C118C (not A).

Etymology.

The name is a feminine noun in the nominative singular, from the Latin lividus (blue), to indicate the distinctive cornflower blue color of the hindwing in males of the type species.

Species included.

The type species and Pythonides grandis Mabille, 1878.

Parent taxon.

Subtribe Pythonidina Grishin, 2019.

Comments.

Another genus that would be very difficult to detect without genomic data, because by appearance the adults of this species do not look noticeably different from Pythonides. However, genomic analysis suggests that the two are distant.

Gindanes kelso (Evans, 1953), new combination

Phylogenetic analysis shows (Fig. 5) that Onenses kelso Evans, 1953 falls within Gindanes Godman and Salvin, [1895] (type species Gindanes panaetius Godman and Salvin, [1895], treated as a subspecies of Hesperia brebisson Latreille, [1824]) and in a different tribe (Achlyodini) from Onenses Godman and Salvin, [1895] (type species Leucochitonea hyalophora R. Felder, 1869, tribe Pyrgini). Therefore, we introduce a new combination: Gindanes kelso.

Festivia Grishin, new genus

http://zoobank.org/16737C47-F8DA-40C1-8D05-B670F1E6FDD0

Type species.

Syrichthus [sic] festiva Erichson, [1849].

Definition.

A sister genus of Sostrata Godman and Salvin, 1895 (Type species. Leucochitonea scintillans Mabille, 1876, which is treated as a junior subjective synonym of Sostrata bifasciata (Ménétriés, 1829)), where these species were placed before (Fig. 6). Keys to E.42.1a in Evans (1953). Distinguished from Sostrata by blue ventral hindwing, broader wings and in male genitalia by longer harpe, clearly separated from the rest of valva. In DNA, a combination of the following base pairs is diagnostic: aly276665.9.3:C95G, aly923.23.1:G274A, aly2790.3.6:A367C, aly923.23.1:T688G, aly2101.22.7:A31C.

http://zoobank.org/86DDFA39-B2D5-44DB-843C-CA314846FBF4

Type species.

Carrhenes santes Bell, 1940.

Definition.

A genus related to Plumbago Evans, 1953, Paches Godman and Salvin, 1895 and Carrhenes Godman and Salvin, 1895 (Fig. 7). Keys to E.43.3 or E.52.4 in Evans (1953), one species previously placed in Paches Godman and Salvin, [1895] (type species Pythonides loxus Westwood, [1852]) the other one in Carrhenes Godman and Salvin, 1895 (type species Leucochitonea fuscescens Mabille, 1891). Distinguished from others by brown wings with darker spots and stripes above and a black spot near the hindwing tornus below. Male genitalia with a process (style) from ampulla, uncus divided, arms about as long as wide, harpe longer than wide. In DNA, a combination of the following base pairs is diagnostic: aly5021.3.22:G847C, aly2103.4.2:G472A, aly528.2.2:G1190T, aly528.2.2:G1191T, aly839.26.5:A464T.

Yet another genus that would be difficult to detect without DNA analysis.

Ladda Grishin, new genus

http://zoobank.org/4BBF25C1-3C31-4B69-9499-516C8251D74E

Type species.

Cyclopides eburones Hewitson, 1877.

Definition.

A sister genus to Dalla Mabille, 1904 (type species Cyclopides eryonas Hewitson, 1877), where these species were placed previously (Fig. 8). Consists of the ibhara, part of quadristriga and part of caenides groups of Evans (1955) and keys to H.6.56b, H.6.44b, or H.6.53b. Distinguished by the combination of straight (non-concave) costa of forewing with unmarked, dark brown above hindwing. If hindwing is spotted above, then the spot is single, centered around the cell, ventral hindwing not plain yellow and without a pale ray from base to margin and if a pale spot is present in M₃-CuA₁ cell, then it is either joined with the spot in CuA₁-CuA₂ cell, or closer the discal cell spot than to margin and ventral hindwing, and the pale spot in CuA₁-CuA₂ cell on ventral hindwing is not connected with distal margin by a broadening pale area (but could be connected with the area of the same width as the spot or narrower). In DNA, a combination of the following base pairs is diagnostic: aly103.37.1:A134T, aly638.15.11:T5060C, aly23605.1.35:A566G, aly1139.25.1:G552T, aly23605.1.35:C580G.

Figure 8. — Genomic trees of Heteropterinae and Trapezitinae. See Fig. 2 legend for notations.

Etymology.

The name is a feminine noun in the nominative singular, a play of letters, from Dalla.

Species included.

The type species, Dalla tona Evans, 1955, Dalla decca Evans, 1955, Butleria morva Mabille, 1898, Thanaos ibhara Butler, 1870, Carterocephalus plancus Hopffer, 1874, Cyclopides crithote Hewitson, 1874, Dalla calima Steinhauser, 1991, Dalla rosea Evans, 1955, Butleria granites Mabille, 1898, Dalla parma Evans, 1955, Dalla mars Evans, 1955, Pamphila cuadrada Weeks, 1901, Butleria quadristriga Mabille, 1889, Dalla rubia Evans, 1955, Argopteron xicca Dyar, 1913, Dalla pedro Steinhauser, 2002, Dalla connexa Draudt, 1923, Dalla disconnexa Steinhauser, 2002, Dalla vista Steinhauser, 2002, Butleria ticidas Mabille, 1898, Cyclopides caenides Hewitson, 1868, Dalla bos Steinhauser, 1991, Dalla pura Steinhauser, 1991, Dalla mora Evans, 1955, Dalla carnis Evans, 1955, Butleria monospila Mabille, 1898, Dalla simplicis Steinhauser, 1991, Dalla seirocastnia Draudt, 1923, Dalla celsus Steinhauser, 2002, Dalla pantha Evans, 1955, Dalla puracensis Steinhauser, 1991, Dalla ochrolimbata Draudt, 1923.

Parent taxon.

Tribe Heteropterini Aurivillius, 1925.

Comments.

Although this genus does not disrupt the monophyly of Dalla when combined with it, Ladda is split from Dalla, due to the large genetic distance between the two genera, each forming a tight cluster of species well separated from the other. Genetic differentiation between these two genera is similar to that between traditional genera such as Heteropterus Duméril, 1806 and Leptalina Mabille, 1904, or Carterocephalus Lederer, 1852 and Metisella Hemming, 1934.

Willema Grishin, new genus

http://zoobank.org/D92BF2E1-F842-44C6-A924-023F495F9589

Type species.

Heteropterus willemi Wallengren, 1857.

Definition.

A sister genus to Hovala Evans, 1937 (type species Cyclopides pardalina Butler, 1879), more distant from Metisella Hemming, 1934 (type species Papilio metis Linnaeus, 1764), where it was placed previously (Fig. 8). Keys to IV.20.B in Evans (1937). Shares many features with Metisella, such as hindwing discal cell longer than half of the wing, forewing veins R₁ and R₂ very near each other or touching (not separated). Distinguished from its relatives by the pattern of hindwing underside, which is with dark web-like pattern along veins with some streaks across and orange, ochreous, yellow or white spots. Hindwing below pale-yellow with dark web-like pattern along veins with some streaks across, but without darker markings. Genitalia with narrow and long penis, uncus with tegumen bulbous in dorsal view, uncus narrows to a point (except in W. willemi where distal end is mushroom-shaped and base of uncus with side processes), gnathos better developed than in Metisella, ampulla separated from harpe, protruding beyond its caudal end (except in W. willemi where harpe upturned embracing ampulla). In DNA, a combination of the following base pairs is diagnostic: aly363.37.1:A929C, aly1450.10.1:A576G, aly283.4.1:T1129C, aly3555.2.2:T116C, aly451.23.2:C1177A.

Etymology.

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

Species included.

The type species, Cyclopides formosus var. tsadicus Aurivillius, 1905, Cyclopides angolana Karsch, 1896, Metisella kumbona Evans, 1937, Cyclopides carsoni Butler, 1898, Heteropterus formosus Butler, 1894, and Cyclopides perexcellens Butler, 1896.

Parent taxon.

Tribe Heteropterini Aurivillius, 1925.

Prada Evans, 1949 belongs to Trapezitinae Waterhouse and Lyell, 1914

Prada (type species Plastingia rothschildi Evans, 1928) is transferred to Trapezitinae because it is monophyletic with this subfamily, receiving the strongest statistical support in all trees (Fig. 8), and is not monophyletic with Hesperiinae where it was formerly placed (Warren et al. 2009).

Argemma Grishin, new genus

http://zoobank.org/3A82BAF3-A287-4EBD-8969-327471780BC1

Type species.

Apaustus argyrosticta Plötz, 1879.

Definition.

A sister genus of Ceratrichia Butler, [1870] (type species Papilio nothus Fabricius, 1787), where these species were formerly placed, but separated from it by a prominent genetic distance consistent with how other genera are defined (Fig. 9). Keys to VI.31.B in Evans (1937) and differs from its relatives by pale apical spot on ventral hindwing in cell R₅-M₁ is in line with the spots in cells next to it (not offset strongly basad); uncus bulbous in ventral view, which gives tegumen with uncus a bowling duckpin shape; harpe widening distad, separated from ampulla; ventral hindwing ochreous with many small silvery spots, dorsal forewing brown with large orange-yellow spots. In DNA, a combination of the following base pairs is diagnostic: aly669.9.1:A231G, aly214.13.5:A98G, aly4389.5.1:G869A, aly1329.4.13:T479C, aly214.13.5:A97C.

http://zoobank.org/199EACE1-E34F-4266-8CAA-46783F667A0A

Type species.

Zea martini Distant and Pryer, 1887.

Definition.

Previously placed in Acerbas de Nicéville, 1895 (type species Hesperia anthea Hewitson, 1868), this genus is not monophyletic with it and is sister to Zela de Nicéville, 1895 (type species Zela zeus de Nicéville, 1895) instead (Fig. 10). Keys to J.22.4 in Evans (1949). Diagnosed by the following combination of characters: mostly brown above wings with small forewing spots, milky areas by hindwing anal margin and broad central band on hindwing below; forewing vein CuA₂ originates in the middle between wing base and the origin of vein CuA₁; uncus broad, bilobed caudad, indistinctly separated from tegumen; tegumen with small side processes and a long central process; penis bulky and stout; valva twice as long as broad, with a prominent and rounded ampulla that is separated from a narrow but large harpe gradually upturned and rounded at the dorsal end protruding above ampulla. In DNA, a combination of the following base pairs is diagnostic: aly103.51.12:A1212T, aly2618.5.1:T1489A, aly420.23.4:A277C, aly276561.5.1:G1724G (not C), aly276561.5.1:A1721A (not G), aly1624.1.10:G534G (not A).

Figure 10. — Genomic trees of Erionotini, Carystina and relatives. See Fig. 2 legend for notations.

Etymology.

The name is a feminine noun in the nominative singular, formed by removing the first and last letters from the name of the genus that the type species was previously attributed to.

Species included.

Only the type species.

Parent taxon.

Tribe Erionotini Distant, 1886.

Avestia Grishin, new genus

http://zoobank.org/F54B9854-755F-4050-8D47-88EDB994DCED

Type species.

Hesperia avesta Hewitson, 1868.

Definition.

A genus sister to Acerbas de Nicéville, 1895 (type species Hesperia anthea Hewitson, 1868) with Zela de Nicéville, 1895 (type species Zela zeus de Nicéville, 1895), previously placed in Lotongus Distant, 1886 (type species Eudamus calathus Hewitson, 1876), but not monophyletic with it (Fig. 10). Keys to J.11.4 in Evans (1949). Morphologically, distinguished from its relatives by hourglass-shaped, terminally bulbous uncus in ventral view; harpe terminally rounded, not prominently separated from ampulla, with a single tooth directed caudad; ventral hindwing yellow band from costa to anal margin, closer to the base, veins brown below (not yellow). In DNA, a combination of the following base pairs is diagnostic: aly1838.39.2:A631G, aly2165.18.2:A22T, aly2096.50.1:A679T, aly1405.13.10:C136C (not G), aly537.26.8:A1279A (not G), aly1264.14.2:A37A (not T).

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.

Tribe Erionotini Distant, 1886.

Zetka Grishin, new genus

http://zoobank.org/C331EB66-E1BB-4C06-9A8B-22A293353C6D

Type species.

Mnasitheus zeteki Bell, 1931.

Definition.

Formerly placed in Styriodes Schaus, 1913 (type species Styriodes lyco Schaus, 1913, in Moncina, Fig. 10–12) this genus does not have close relatives, but belongs in the same clade as Neoxeniades Hayward, 1938 (type species Neoxeniades musarion Hayward, 1938). Keys to K.2.6. in Evans (1955). Can be told apart from its relatives by the following combination of characters: males with long brands over vein 1A+2A and under vein CuA₂ but not above vein CuA₂, antennae as long as forewing discal cell, palpi broad, 3rd segment short, stout, wings and body brown. Genitalia illustrated by Bell (1931: fig. 1), uncus shorter than tegumen, stout, with small rounded arms, gnathos as long as uncus, saccus vestigial, valva twice as long as wide, harpe angled and squared caudad, upturned dorsocephalad, serrated at the dorsal end that protrudes from valva (Bell 1931). In DNA, a combination of the following base pairs is diagnostic: aly1146.42.8:A1629G, aly1222.14.14:T7181A, aly1405.10.1:C373T, aly251.9.1:C1499T, aly103.32.1:A1000G.

Subtribe Falgina trib. n., described above.

Comments.

A genus with surprising phylogenetic placement that yet again indicates the value of DNA-based methods at the genomic scale, and instills confidence in the results due to the large datasets used for phylogeny reconstruction and highly confident nodes in phylogenetic trees.

Brownus Grishin, new genus

http://zoobank.org/99C1730A-E965-410D-A15A-D8754C5E3FF0

Type species.

Paratrytone browni Bell, 1959.

Definition.

Unexpectedly, a relative of Amblyscirtes Scudder, 1872 (type species Hesperia vialis Edwards, 1862) and is sister to Remella Hemming, 1939 (type species Hesperia remus Fabricius, 1798) with Mnasicles Godman, [1901] (type species Mnasicles geta Godman, 1901), which all are in a different subtribe (Moncina) from Paratrytone Godman, [1900] (type species Paratrytone rhexenor Godman, 1900) (in Hesperiina) where this lineage was placed previously (Fig. 11–12). Distinguished from its relatives by sacculus about as long as vinculum with tegumen in lateral view, valva simple, nearly diamond-shaped with vary narrow harpe ending in a point directed dorsoposteriad and not separated from ampulla, very long antennae reaching apical spots near costa, all (except the small cell spot) forewing spots forming a continuous line (separated by black veins only) and most spots in a line on ventral hindwing. In DNA, a combination of the following base pairs is diagnostic: aly28779.8.7:T461C, aly577.34.1:C115G, aly1370.21.5:A2861G, aly531.10.5:G517C, aly2124.3.99:A3193G.

A sister genus to Oxynthes Godman, [1900] (type species Goniloba corusca Herrich-Schäffer, 1869) combined with Oeonus Godman, [1900] (type species Oeonus pyste Godman, [1900]) and in the same clade with Lindra Evans, 1955 (type species Carystus simulius Druce, 1876) (Fig. 13). Not monophyletic with Orthos Evans, 1955 (type species Eutychide orthos Godman, [1900]), which is in a different subtribe (Carystina Mabille 1878). Keys to L.15.5 in Evans (1955). Distinguished from its relatives by a combination of the following characters (Bell 1941). Antennae about 2/3 of costa in length, male no secondary sexual characters, wings unmarked dark brown above with shiny green hair-like scales in tornal area of hindwing (and on body), with cream spots below: on hindwing near its middle plus diffuse cream area in distal half of CuA₂-1A+2A cell, small spots on hindwing in discal cell and as a postdiscal row in each cell between veins M₂ and 1A+2A. Male genitalia (see Bell 1941: fig. 3 for illustration) with uncus deeply divided, arms long (only slightly shorter than tegumen), parallel, near each other, penis widening terminally (nearly as wide as valva), bulky, with terminal spikes, valva nearly rectangular, sacculus without processes, harpe short (shorter than 1/3 of valva), angular, with a stout tooth pointed dorsad, separated from ampulla by a narrow notch. In DNA, a combination of the following base pairs is diagnostic: aly2874.22.9:G1030A, aly3512.12.2:T234C, aly2811.6.1:T166C, aly1139.93.1:C452C (not G), aly862.12.2:A1755A (not C), aly5021.5.1:G1325G (not C), aly537.7.1:A181A (not G), aly2012.14.2:T317T (not C), aly577.34.1:A485A (not T).

The type species, Hesperia hobomok T. Harris, 1862, Pamphila inimica Butler and Druce, 1872, Pamphila taxiles Edwards, 1881, Pamphila azin Godman, 1900, Poanes macneilli Burns, 1992, Hesperia ulphila Plötz, 1883, Atrytone monticola Godman, [1900], Pamphila niveolimbus Mabille, 1889, Hesperia melane Edwards, 1869.

Parent taxon.

Subtribe Hesperiina Latreille, 1809.

Alera Mabille, 1891 is a synonym of Perichares Scudder, 1872

We find (Fig. 13) that Alera furcata Mabille, 1891 and Perichares philetes (Gmelin, [1790]), the type species of both genera (technically, Papilio coridon Fabricius, 1775, a homonym, considered to refer to Perichares philetes, is the type species of Perichares) are genetically close (COI barcodes differ by about 6.5%). Moreover, several species of Alera that we sequenced are not monophyletic and are interspersed between species of Perichares. For these reasons, we consider Alera to be a subjective junior synonym of Perichares.

Oenides Mabille, 1904 is a valid genus

Hesperia vulpina C. and R. Felder, 1867 is not monophyletic with Alera furcata, the type species of Alera, where H. vulpina was previously placed, and since it is the type species of the available genus-group name Oenides Mabille, 1904, that was considered a subjective synonym of Alera, we resurrect this genus from synonymy and use it as a monotypic valid genus with the species Oenides vulpina (Fig. 13).

Supplementary Material

Supplemental file

NIHMS1724225-supplement-Supplemental_file.pdf^{(1.2MB, pdf)}

Acknowledgments

We are grateful to David Grimaldi and Courtney Richenbacher (American Museum of Natural History, New York, NY, USA), Blanca Huertas, David Lees and Geoff Martin (Natural History Museum, London, UK), Jonathan Pelham (Burke Museum of Natural History and Culture, Seattle, WA, USA), John Rawlins (Carnegie Museum of Natural History, Pittsburgh, PA, USA), Paul A. Opler and Boris Kondratieff (Colorado State University Collection, Fort Collins, CO, USA), Crystal Maier and Rebekah Baquiran (Field Museum of Natural History, Chicago, FL, USA), Weiping Xie (Los Angeles County Museum of Natural History, Los Angeles, CA, USA), Naomi Pierce, Philip Perkins and Rachel Hawkins (Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA), John R. MacDonald and Richard L. Brown (Mississippi Entomological Museum, Starkville, MS, USA), Rodolphe Rougerie (Muséum National d’Histoire Naturelle, Paris, France), Wolfgang A. Nässig (Natural History Museum, Frankfurt, Germany), Edward G. Riley, Karen Wright, and John Oswald (Texas A&M University Insect Collection, College Station, TX, USA), Robert K. Robbins, John M. Burns, and Brian Harris (National Museum of Natural History, Smithsonian Institution, Washington, DC, USA), Wolfram Mey and Viola Richter (ZMHB: Museum für Naturkunde, Berlin, Germany), Axel Hausmann and Ulf Buchsbaum (Zoologische Staatssammlung München, Germany) for granting access to the collections under their care and for stimulating discussions; 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 (Wildlife Biologist Raymond Skiles) for the Big Bend National Park research permit BIBE-2004-SCI-0011; to the National Environment & Planning Agency of Jamaica for the permission to collect specimens; to Paul A. Opler and Rongjiang Wang for specimens; to Jim P. Brock, Ernst Brockmann, Robert Gallardo, Bernard Hermier, the late Edward C. Knudson (specimens now at McGuire Center for Lepidoptera and Biodiversity, Gainesville, FL, USA), John MacDonald, Kiyoshi Maruyama, James A. Scott, and Mark Walker for leg samples of their specimens, to Ernst Brockmann for help with sampling specimens for DNA, to Gerardo Lamas, Bernard Hermier and Jonathan Pelham for fruitful discussions and to Bernard Hermier for critical reading of the manuscript at its early stages and copious helpful suggestions and corrections. We are indebted to Paul Opler and John Shuey for critical reviews of the manuscript. We acknowledge the Texas Advanced Computing Center (TACC) at The University of Texas at Austin for providing HPC resources. The study was supported in part by the grants (to NVG) from the National Institutes of Health GM094575 and GM127390 and the Welch Foundation I-1505.

Contributor Information

Qian Cong, Institute for Protein Design and Department of Biochemistry, University of Washington, 1959 NE Pacific Street, HSB J-405, Seattle, WA, 98195 USA.

Jing Zhang, Departments of Biophysics and Biochemistry, University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX, 75390-8816 USA.

Jinhui Shen, Departments of Biophysics and Biochemistry, University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX, 75390-8816 USA.

Nick V. Grishin, Howard Hughes Medical Institute and Departments of Biophysics and Biochemistry, University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX, 75390-9050 USA

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Cong Q, Shen J, Borek D, Robbins RK, Opler PA, Otwinowski Z, and Grishin NV. 2017a. When COI barcodes deceive: complete genomes reveal introgression in hairstreaks. Proceedings of the Royal Society B: Biological Sciences 284(1848): 20161735. [DOI] [PMC free article] [PubMed] [Google Scholar]
Cong Q, Shen J, Borek D, Robbins RK, Otwinowski Z, and Grishin NV. 2016a. Complete genomes of hairstreak butterflies, their speciation, and nucleo-mitochondrial incongruence. Scientific Reports 6: 24863. [DOI] [PMC free article] [PubMed] [Google Scholar]
Cong Q, Shen J, Li W, Borek D, Otwinowski Z, and Grishin NV. 2017b. The first complete genomes of metalmarks and the classification of butterfly families. Genomics 109: 485–493. [DOI] [PMC free article] [PubMed] [Google Scholar]
Cong Q, Shen J, Warren AD, Borek D, Otwinowski Z, and Grishin NV. 2016b. Speciation in cloudless sulphurs gleaned from complete genomes. Genome Biology and Evolution 8(3): 915–931. [DOI] [PMC free article] [PubMed] [Google Scholar]
Espeland M, Breinholt J, Willmott KR, Warren AD, Vila R, Toussaint EFA, Maunsell SC, Aduse-Poku K, Talavera G, Eastwood R, Jarzyna MA, Guralnick R, Lohman DJ, Pierce NE, and Kawahara AY. 2018. A comprehensive and dated phylogenomic analysis of butterflies. Current Biology 28(5): 770–778. [DOI] [PubMed] [Google Scholar]
Evans WH 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum. British Museum (Natural History); London. xii + 212 p., 30 pl. [Google Scholar]
Evans WH 1949. A catalogue of the Hesperiidae from Europe, Asia, and Australia in the British Museum (Natural History). British Museum (Natural History); London. xix + 502 p., 53 pl. [Google Scholar]
Evans WH 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. x + 92 p., pl. 1–9 [Google Scholar]
Evans WH 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. v + 178 p., pl. 10–25 [Google Scholar]
Evans WH 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. v + 246 p., pl. 26–53 [Google Scholar]
Evans WH 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. v + 499 p., pl. 54–88 [Google Scholar]
Grishin NV 2012. A new Central American Anastrus with unexpectedly distinct genitalia (Lepidoptera, Hesperiidae, Pyrginae). Tropical Lepidoptera Research 22(1): 1–7. [Google Scholar]
Hancock DL, and Gardiner AJ. 1982. The Kedestes nerva group of species (Lepidoptera: Hesperiidae). Arnoldia Zimbabwe 9(8): 105–123. [Google Scholar]
Kawahara AY, and Breinholt JW. 2014. Phylogenomics provides strong evidence for relationships of butterflies and moths. Proceedings of the Royal Society B: Biological Sciences 281(1788): 20140970. [DOI] [PMC free article] [PubMed] [Google Scholar]
Li W, Cong Q, Shen J, Zhang J, Hallwachs W, Janzen DH, and Grishin NV. 2019. 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. [DOI] [PMC free article] [PubMed] [Google Scholar]
Mielke CGC 1995. Papilionoidea e Hesperioidea (Lepidoptera) de Curitiba e seus arrededores, Paraná, Brasil, com notas taxonômicas sobre Hesperiidae. Revista brasileira de Zoologia 11(4): 759–776. [Google Scholar]
Mielke OHH 1980. Contribuição ao estudo faunístico dos Hesperiidae americanos. VI. Nota suplementar às espécies de Hesperiinae do Rio Grande do Sul, Brasil (Lepidoptera). Acta Biologica Paranaense 8–9: 127–172. [Google Scholar]
Mielke OHH 2005. Catalogue of the American Hesperioidea: Hesperiidae (Lepidoptera). Sociedade Brasileira de Zoologia; Curitiba, Paraná, Brazil. xiii + 1536 p. [Google Scholar]
Nicolay SS 1980. Descriptions of new Hesperiidae from Panama and Ecuador (Pyrginae and Hesperiinae). Bulletin of the Allyn Museum 59: 1–17. [Google Scholar]
Sahoo RK, Warren AD, Collins SC, and Kodandaramaiah U. 2017. Hostplant change and paleoclimatic events explain diversification shifts in skipper butterflies (Family: Hesperiidae). BMC Evolutionary Biology 17(1): 174. [DOI] [PMC free article] [PubMed] [Google Scholar]
Sahoo RK, Warren AD, Wahlberg N, Brower AV, Lukhtanov VA, and Kodandaramaiah U. 2016. Ten genes and two topologies: an exploration of higher relationships in skipper butterflies (Hesperiidae). PeerJ 4: e2653. [DOI] [PMC free article] [PubMed] [Google Scholar]
Shen J, Cong Q, Borek D, Otwinowski Z, and Grishin NV. 2017. Complete genome of Achalarus lyciades, the first representative of the Eudaminae subfamily of skippers. Current Genomics 18(4): 366–374. [DOI] [PMC free article] [PubMed] [Google Scholar]
Shen J, Cong Q, and Grishin NV. 2015. The complete mitochondrial genome of Papilio glaucus and its phylogenetic implications. Meta Gene 5: 68–83. [DOI] [PMC free article] [PubMed] [Google Scholar]
Shen J, Cong Q, and Grishin NV. 2016a. The complete mitogenome of Achalarus lyciades (Lepidoptera: Hesperiidae). Mitochondrial DNA Part B: Resources 1(1): 581–583. [DOI] [PMC free article] [PubMed] [Google Scholar]
Shen J, Cong Q, Kinch LN, Borek D, Otwinowski Z, and Grishin NV. 2016b. Complete genome of Pieris rapae, a resilient alien, a cabbage pest, and a source of anti-cancer proteins. F1000Research 5: 2631. [DOI] [PMC free article] [PubMed] [Google Scholar]
Smith DS, Miller LD, and Miller JY. 1994. The butterflies of the West Indies and South Florida. Oxford Univ. Press; Oxford, New York, Tokyo. x + 264 p. [Google Scholar]
Steinhauser SR 1989. Taxonomic notes and descriptions of new taxa in the Neotropical Hesperiidae. Part I. Pyrginae. Bulletin of the Allyn Museum 127: 1–70. [Google Scholar]
Toussaint EFA, Breinholt JW, Earl C, Warren AD, Brower AVZ, Yago M, Dexter KM, Espeland M, Pierce NE, Lohman DJ, and Kawahara AY. 2018. Anchored phylogenomics illuminates the skipper butterfly tree of life. BMC Evolutionary Biology 18(1): 101. [DOI] [PMC free article] [PubMed] [Google Scholar]
Wahlberg N, Braby MF, Brower AV, de Jong R, Lee MM, Nylin S, Pierce NE, Sperling FA, Vila R, Warren AD, and Zakharov E. 2005. Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proceedings of the Royal Society B: Biological Sciences 272(1572): 1577–1586. [DOI] [PMC free article] [PubMed] [Google Scholar]
Warren AD, Ogawa JR, and Brower AVZ. 2008. Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea). Cladistics 24(5): 642–676. [Google Scholar]
Warren AD, Ogawa JR, and Brower AVZ. 2009. Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomology 34(3): 467–523. [Google Scholar]
Watson EY 1893. A proposed classification of the Hesperiidae, with a revision of the genera. Proceedings of the Zoological Society of London 1893(1): 3–132. [Google Scholar]
Zhang J, Cong Q, Rex EA, Hallwachs W, Janzen DH, Grishin NV, and Gammon DB. 2019a. Gypsy moth genome provides insights into flight capability and virus-host interactions. Proceedings of the National Academy of Sciences of the United States of America 116(5): 1669–1678. [DOI] [PMC free article] [PubMed] [Google Scholar]
Zhang J, Cong Q, Shen J, Brockmann E, and Grishin NV. 2019b. Genomes reveal drastic and recurrent phenotypic divergence in firetip skipper butterflies (Hesperiidae: Pyrrhopyginae). Proceedings of the Royal Society B: Biological Sciences 286(1903): 20190609. [DOI] [PMC free article] [PubMed] [Google Scholar]
Zhang J, Cong Q, Shen J, Brockmann E, and Grishin NV. 2019c. Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae). Zookeys 861: 91–105. [DOI] [PMC free article] [PubMed] [Google Scholar]
Zhang J, Cong Q, Shen J, Fan XL, Wang M, and Grishin NV. 2017a. The complete mitogenome of Euschemon rafflesia (Lepidoptera: Hesperiidae). Mitochondrial DNA Part B: Resources 2(1): 136–138. [DOI] [PMC free article] [PubMed] [Google Scholar]
Zhang J, Cong Q, Shen J, Wang R, and Grishin NV. 2017b. The complete mitochondrial genome of a skipper Burara striata (Lepidoptera: Hesperiidae). Mitochondrial DNA Part B: Resources 2(1): 145–147. [DOI] [PMC free article] [PubMed] [Google Scholar]

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Supplemental file

NIHMS1724225-supplement-Supplemental_file.pdf^{(1.2MB, pdf)}

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[R16] Cong Q, Li W, Borek D, Otwinowski Z, and Grishin NV. 2018. The bear giant-skipper genome suggests genetic adaptations to living inside yucca roots. Molecular Genetics and Genomics 294(1): 211–226. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R17] Cong Q, Shen J, Borek D, Robbins RK, Opler PA, Otwinowski Z, and Grishin NV. 2017a. When COI barcodes deceive: complete genomes reveal introgression in hairstreaks. Proceedings of the Royal Society B: Biological Sciences 284(1848): 20161735. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R18] Cong Q, Shen J, Borek D, Robbins RK, Otwinowski Z, and Grishin NV. 2016a. Complete genomes of hairstreak butterflies, their speciation, and nucleo-mitochondrial incongruence. Scientific Reports 6: 24863. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R19] Cong Q, Shen J, Li W, Borek D, Otwinowski Z, and Grishin NV. 2017b. The first complete genomes of metalmarks and the classification of butterfly families. Genomics 109: 485–493. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R20] Cong Q, Shen J, Warren AD, Borek D, Otwinowski Z, and Grishin NV. 2016b. Speciation in cloudless sulphurs gleaned from complete genomes. Genome Biology and Evolution 8(3): 915–931. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R21] Espeland M, Breinholt J, Willmott KR, Warren AD, Vila R, Toussaint EFA, Maunsell SC, Aduse-Poku K, Talavera G, Eastwood R, Jarzyna MA, Guralnick R, Lohman DJ, Pierce NE, and Kawahara AY. 2018. A comprehensive and dated phylogenomic analysis of butterflies. Current Biology 28(5): 770–778. [DOI] [PubMed] [Google Scholar]

[R22] Evans WH 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum. British Museum (Natural History); London. xii + 212 p., 30 pl. [Google Scholar]

[R23] Evans WH 1949. A catalogue of the Hesperiidae from Europe, Asia, and Australia in the British Museum (Natural History). British Museum (Natural History); London. xix + 502 p., 53 pl. [Google Scholar]

[R24] Evans WH 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. x + 92 p., pl. 1–9 [Google Scholar]

[R25] Evans WH 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. v + 178 p., pl. 10–25 [Google Scholar]

[R26] Evans WH 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. v + 246 p., pl. 26–53 [Google Scholar]

[R27] Evans WH 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. v + 499 p., pl. 54–88 [Google Scholar]

[R28] Grishin NV 2012. A new Central American Anastrus with unexpectedly distinct genitalia (Lepidoptera, Hesperiidae, Pyrginae). Tropical Lepidoptera Research 22(1): 1–7. [Google Scholar]

[R29] Hancock DL, and Gardiner AJ. 1982. The Kedestes nerva group of species (Lepidoptera: Hesperiidae). Arnoldia Zimbabwe 9(8): 105–123. [Google Scholar]

[R30] Kawahara AY, and Breinholt JW. 2014. Phylogenomics provides strong evidence for relationships of butterflies and moths. Proceedings of the Royal Society B: Biological Sciences 281(1788): 20140970. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R31] Li W, Cong Q, Shen J, Zhang J, Hallwachs W, Janzen DH, and Grishin NV. 2019. 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. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R32] Mielke CGC 1995. Papilionoidea e Hesperioidea (Lepidoptera) de Curitiba e seus arrededores, Paraná, Brasil, com notas taxonômicas sobre Hesperiidae. Revista brasileira de Zoologia 11(4): 759–776. [Google Scholar]

[R33] Mielke OHH 1980. Contribuição ao estudo faunístico dos Hesperiidae americanos. VI. Nota suplementar às espécies de Hesperiinae do Rio Grande do Sul, Brasil (Lepidoptera). Acta Biologica Paranaense 8–9: 127–172. [Google Scholar]

[R34] Mielke OHH 2005. Catalogue of the American Hesperioidea: Hesperiidae (Lepidoptera). Sociedade Brasileira de Zoologia; Curitiba, Paraná, Brazil. xiii + 1536 p. [Google Scholar]

[R35] Nicolay SS 1980. Descriptions of new Hesperiidae from Panama and Ecuador (Pyrginae and Hesperiinae). Bulletin of the Allyn Museum 59: 1–17. [Google Scholar]

[R36] Sahoo RK, Warren AD, Collins SC, and Kodandaramaiah U. 2017. Hostplant change and paleoclimatic events explain diversification shifts in skipper butterflies (Family: Hesperiidae). BMC Evolutionary Biology 17(1): 174. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R37] Sahoo RK, Warren AD, Wahlberg N, Brower AV, Lukhtanov VA, and Kodandaramaiah U. 2016. Ten genes and two topologies: an exploration of higher relationships in skipper butterflies (Hesperiidae). PeerJ 4: e2653. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R38] Shen J, Cong Q, Borek D, Otwinowski Z, and Grishin NV. 2017. Complete genome of Achalarus lyciades, the first representative of the Eudaminae subfamily of skippers. Current Genomics 18(4): 366–374. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R39] Shen J, Cong Q, and Grishin NV. 2015. The complete mitochondrial genome of Papilio glaucus and its phylogenetic implications. Meta Gene 5: 68–83. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R40] Shen J, Cong Q, and Grishin NV. 2016a. The complete mitogenome of Achalarus lyciades (Lepidoptera: Hesperiidae). Mitochondrial DNA Part B: Resources 1(1): 581–583. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R41] Shen J, Cong Q, Kinch LN, Borek D, Otwinowski Z, and Grishin NV. 2016b. Complete genome of Pieris rapae, a resilient alien, a cabbage pest, and a source of anti-cancer proteins. F1000Research 5: 2631. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R42] Smith DS, Miller LD, and Miller JY. 1994. The butterflies of the West Indies and South Florida. Oxford Univ. Press; Oxford, New York, Tokyo. x + 264 p. [Google Scholar]

[R43] Steinhauser SR 1989. Taxonomic notes and descriptions of new taxa in the Neotropical Hesperiidae. Part I. Pyrginae. Bulletin of the Allyn Museum 127: 1–70. [Google Scholar]

[R44] Toussaint EFA, Breinholt JW, Earl C, Warren AD, Brower AVZ, Yago M, Dexter KM, Espeland M, Pierce NE, Lohman DJ, and Kawahara AY. 2018. Anchored phylogenomics illuminates the skipper butterfly tree of life. BMC Evolutionary Biology 18(1): 101. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R45] Wahlberg N, Braby MF, Brower AV, de Jong R, Lee MM, Nylin S, Pierce NE, Sperling FA, Vila R, Warren AD, and Zakharov E. 2005. Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proceedings of the Royal Society B: Biological Sciences 272(1572): 1577–1586. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R46] Warren AD, Ogawa JR, and Brower AVZ. 2008. Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea). Cladistics 24(5): 642–676. [Google Scholar]

[R47] Warren AD, Ogawa JR, and Brower AVZ. 2009. Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomology 34(3): 467–523. [Google Scholar]

[R48] Watson EY 1893. A proposed classification of the Hesperiidae, with a revision of the genera. Proceedings of the Zoological Society of London 1893(1): 3–132. [Google Scholar]

[R49] Zhang J, Cong Q, Rex EA, Hallwachs W, Janzen DH, Grishin NV, and Gammon DB. 2019a. Gypsy moth genome provides insights into flight capability and virus-host interactions. Proceedings of the National Academy of Sciences of the United States of America 116(5): 1669–1678. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R50] Zhang J, Cong Q, Shen J, Brockmann E, and Grishin NV. 2019b. Genomes reveal drastic and recurrent phenotypic divergence in firetip skipper butterflies (Hesperiidae: Pyrrhopyginae). Proceedings of the Royal Society B: Biological Sciences 286(1903): 20190609. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R51] Zhang J, Cong Q, Shen J, Brockmann E, and Grishin NV. 2019c. Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae). Zookeys 861: 91–105. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R52] Zhang J, Cong Q, Shen J, Fan XL, Wang M, and Grishin NV. 2017a. The complete mitogenome of Euschemon rafflesia (Lepidoptera: Hesperiidae). Mitochondrial DNA Part B: Resources 2(1): 136–138. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R53] Zhang J, Cong Q, Shen J, Wang R, and Grishin NV. 2017b. The complete mitochondrial genome of a skipper Burara striata (Lepidoptera: Hesperiidae). Mitochondrial DNA Part B: Resources 2(1): 145–147. [DOI] [PMC free article] [PubMed] [Google Scholar]

PERMALINK

Fifty new genera of Hesperiidae (Lepidoptera)

Qian Cong

Jing Zhang

Jinhui Shen

Nick V Grishin

Abstract

Introduction

Materials and Methods

Table 1.

Results and Discussion

Ceratrichiini Grishin, new tribe

Type genus.

Definition.

Figure 1.

Genera included.

Parent taxon.

Comments.

Gretnini Grishin, new tribe

Type genus.

Definition.

Genera included.

Parent taxon.

Comments.

Falgina Grishin, new subtribe

Type genus.

Definition.

Genera included.

Parent taxon.

Comments.

Apaustina Grishin, new subtribe

Type genus.

Definition.

Genera included.

Parent taxon.

Comments.

Composition of the subtribes Calpodina Clark, 1948 and Thymelicina Tutt, 1905

Flattoides Grishin, new genus

Type species.

Definition.

Figure 2.

Etymology.

Species included.

Parent taxon.

Aurivittia Grishin, new genus

Type species.

Definition.

Figure 3.

Etymology.

Species included.

Parent taxon.

Comments.

Viuria Grishin, new genus

Type species.

Definition.

Figure 4.

Etymology.

Species included.

Parent taxon.

Clytius Grishin, new genus

Type species.

Definition.

Etymology.

Species included.

Parent taxon.

Incisus Grishin, new genus

Type species.

Definition.

Etymology.

Species included.

Parent taxon.

Comments.

Perus Grishin, new genus

Type species.

Definition.

Etymology.

Species included.

Parent taxon.

Tosta Evans, 1953 is a synonym of Eantis Boisduval, 1836

Figure 5.