Received: 21 August 2021 Accepted: 11 February 2022
DOI: 10.1111/syen.12541
OR I G I N A L A R T I C L E
Genomics-based higher classification of the species-rich
hairstreaks (Lepidoptera: Lycaenidae: Eumaeini)
Robert K. Robbins1 | Qian Cong2 | Jing Zhang3 | Jinhui Shen3 |
Robert C. Busby4 | Christophe Faynel5 | Marcelo Duarte6 |
Ananda R. P. Martins7 | Carlos Prieto8 | Gerardo Lamas9 | Nick V. Grishin3,10
1Department of Entomology, National Abstract
Museum of Natural History, Smithsonian
Institution, Washington, District of We propose a higher classification of the lycaenid hairstreak tribe Eumaeini – one of the
Columbia, USA youngest and most species-rich butterfly tribes – based on autosome, Lepidopteran Z sex
2McDermott Center for Human Growth and chromosome and mitochondrial protein-coding genes. The subtribe Neolycaenina Korb is a
Development, University of Texas
Southwestern Medical Center, Dallas, synonym of Callophryidina Tutt and subtribe Tmolusina Bálint is a synonym of Strephonotina
Texas, USA K. Johnson, Austin, Le Crom, & Salazar. Proposed names are Rhammina Prieto & Busby,
3Department of Biophysics and Biochemistry,
new subtribe; Timaetina Busby & Prieto, new subtribe; Atlidina Martins & Duarte, new
University of Texas Southwestern Medical
Center, Dallas, Texas, USA subtribe; Evenina Faynel & Grishin, new subtribe; Jantheclina Robbins & Faynel, new subtribe;
4Estero, Florida, USA Paiwarriina Lamas & Robbins, new subtribe; Cupatheclina Lamas & Grishin, new subtribe;
5Montaud, France
Parrhasiina Busby & Robbins, new subtribe; Ipideclina Martins & Grishin, new subtribe; and
6Museu de Zoologia, Universidade de S~ao
Paulo, S~ao Paulo, Brazil Trichonidina Duarte & Faynel, new subtribe. Phylogenetic results from the autosome and Z
7Redpath Museum, McGill University, sex chromosome analyses are similar. Future analyses of datasets with hundreds of terminal
Montreal, Canada taxa may be more practical time-wise by focussing on the smaller number of sex chromo-
8Departamento de Biología, Universidad del
some sequences (2.6% of nuclear protein-coding sequences). The phylogenetic classification
Atlántico, Barranquilla, Colombia and
Corporacion Universitaria Autonoma del and biological summaries for each subtribe suggest that a variety of factors affected
Cauca, Popayán, Colombia
Eumaeini diversification. About a dozen kinds of male secondary sexual organs with fre-
9Museo de Historia Natural, Universidad
Nacional Mayor de San Marcos, Lima, Peru quent evolutionary gains and losses occur in Atlidina, Evenina and Jantheclina (141 species
10Howard Hughes Medical Institute, combined). Females have been shown to use these organs to discriminate between conspe-
University of Texas Southwestern Medical
cific and nonconspecific males, facilitating sympatry among close relatives. Eumaeina,
Center, Dallas, Texas, USA
Rhammina and Timaetina (140 species combined) are overwhelmingly montane with some
Correspondence evidence for a higher incidence of sympatric diversification. Seven Neotropical lineages in
Robert K. Robbins, Department of Entomology,
National Museum of Natural History, PO Box five subtribes invaded the temperate parts of the Nearctic Region with a diversification
37012, NHB Stop 105, Smithsonian Institution, increase in the Callophryidina (262 species). North American Satyrium and Callophrys then
Washington, D.C., USA.
Email: robbinsr@si.edu invaded the Palearctic at least once each, with a major species-richness increase in Satyrium.
The evolution of litter-feeding detritivores within Calycopidina (172 species) resulted in an
Funding information
increase in diversification rate compared with its flower-feeding sister lineage. Atlidina,
Conselho Nacional de Desenvolvimento
Científico e Tecnologico, Grant/Award Strephonotina, Parrhasiina and Strymonina (562 species combined) each contain a mixture
Numbers: 305905/2012-0, 311083/2015-3,
~ of genera that specialize on one or two caterpillar food plant families and genera that are312190/2018-2; Coordenaçao de
Aperfeiçoamento de Pessoal de Nível Superior, polyphagous. These would be appropriate subtribes to assess how the breadth of caterpillar
Grant/Award Number: 440597/2015-3;
food plants and the frequency of host shifts affected diversification.
FAPESP, Grant/Award Numbers:
2002/13898-0, 2003/13985-3,
2016/50384-8; United State National
© 2022 Royal Entomological Society. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.
Syst Entomol. 2022;1–25. wileyonlinelibrary.com/journal/syen 1
2 ROBBINS ET AL.
Institutes of Health (NIH), Grant/Award
Number: GM127390; NMNH ADS core;
K E YWORD S
Welch Foundation, Grant/Award Number: I-
diversification, food plant specialization, male secondary sexual organs, polyphagy, Theclinae, Z sex
1505
chromosome
INTRODUCTION Robinson, 2008; Fiedler, 1991; Silva, Duarte, Diniz, & Morais, 2011).
Species in genera such as Rekoa Kaye, Strymon Hübner, Pseudolycaena
The tribe Eumaeini (Lepidoptera: Lycaenidae) is biologically notable Wallengren and Panthiades Hübner feed on plants in more than a
for being a rapidly diversifying clade. With an estimated age of dozen families (e.g., Austin, Miller, & Miller, 2007; Janzen &
approximately 30 million years, it is one of the youngest recognized Hallwachs, 2021; Monteiro, 1991; Robbins, 1991a; Robbins &
tribes of butterflies (Espeland et al., 2018; Valencia-Montoya Nicolay, 2002). The genus Callophrys is recorded from Gymnosperms,
et al., 2021). With about 1200 New World and 100+ Palearctic spe- Monocotyledons and Dicotyledons (Ehrlich & Raven, 1965).
cies (Robbins, 2004a; Weidenhoffer, Bozano, & Churkin, 2004), it is Calycopidina are leaf-litter detritivores (Duarte & Robbins, 2010;
also one of the most species-rich with 7%–8% of the world’s diurnal Robbins et al., 2010), including seeds and mushrooms (Basset
butterfly species (Lamas, 2008). et al., 2018; Gripenberg et al., 2019; Nishida & Robbins, 2020). In con-
The higher classification of Eumaeini has been unsettled. Many trast, the caterpillars of some genera specialize primarily on one plant
proposed lineages based on morphology were poorly characterized family. Examples include Eumaeus Hübner on Zamiaceae (Robbins
(Robbins, 2004a), and those that were readily recognized, such as the et al., 2021), Evenus Hübner on new leaves of Sapotaceae (Janzen &
Atlides Section and the Lamprospilus Section (later named Hallwachs, 2021; Robbins, 2004a), a lineage of Strymon Hübner on
Calycopidina), were based on homoplastic characters (Duarte & Bromeliaceae (Robbins, 2010a), Arawacus Kaye on Solanaceae
Robbins, 2010; Martins, Duarte, & Robbins, 2019a). A molecular anal- (Robbins, 2000) and Thereus Hübner on Loranthaceae (Heredia &
ysis based upon 187 eumaeines recognized eight lineages, but no tax- Robbins, 2016). The concept that caterpillar food plant associations,
onomic changes were proposed (Valencia-Montoya et al., 2021). such as specialization, generalization and the frequency of host shifts,
Congruence between morphological and molecular groupings was shape diversification has been widely supported (e.g., Braga,
limited. Guimara~es, Wheat, Nylin, & Janz, 2018; Fordyce, 2010; Hardy &
Eumaeines are evolutionarily notable for a variety of male sec- Otto, 2014; Janz, 2011; Janz, Nylin, & Wahlberg, 2006; St Laurent,
ondary sexual organs that have been associated with diversification Carvalho, Earl, & Kawahara, 2021). However, without a classification
(Valencia-Montoya et al., 2021). For example, males of the Neotropi- to use as a foundation, patterns of Eumaeini caterpillar food plant uti-
cal Arcas cypria (Geyer) have scent pads, scent patches, scent pouches lization have not been assessed.
and abdominal brush organs (Martins, Duarte, & Robbins, 2019b; Rob- Mutualistic interactions between caterpillars and ants occur
bins, Martins, Busby, & Duarte, 2012). Over 90% of eumaeine species widely among Lycaenidae and have been implicated in rapid diversifi-
possess at least one male secondary sexual organ (Valencia-Montoya cation in Old World lineages (i.e., Eastwood, Pierce, Kitching, &
et al., 2021). The seven male secondary sexual organs in one clade Hughes, 2006; Pierce, 1984). In Eumaeini, ants may provide enemy-
(Atlides Section) were gained or lost within the lineage, often multiple free space for caterpillars (Kaminski, Freitas, & Oliveira, 2010;
times (Martins et al., 2019b). Females in the Atlides Section use these Kaminski, Rodrigues, & Freitas, 2012), which may be facultatively myr-
organs to identify conspecific males (Martins et al., 2019b). In this mecophilous (DeVries, 1990, 1991), but obligate mutualism is
sense, the evolutionary gain of male secondary sexual organs might unrecorded. Whether myrmecophily is an important factor in the
have facilitated sympatric diversification. diversification of Eumaeini is largely unexplored.
Eumaeines occur in a wide range of habitats. They inhabit areas Sequencing complete genomes of Eumaeini butterflies using
with negligible rainfall, such as the Atacama Desert, to forests with museum specimens provides a prodigious quantity of phylogenetic
800 cm annual precipitation (Lamas, McInnis, Busby, & data (Cong et al., 2016, 2017; Robbins et al., 2021). In this paper, we
Robbins, 2021; Vargas & Duarte, 2016). They occur in areas up to infer phylogenetic relationships using three datasets of protein-coding
4000 m elevation (Bálint, Katona, & Kertész, 2019). They are wide- genes. The first dataset contains autosome sequences with ‘diploid’
spread in the Palearctic, whereas in the New World, they occur in inheritance. The second is the Lepidopteran Z sex chromosome genes
almost all areas with plants from the subarctic to the central valley of with ‘haplo-diploid’ inheritance and the third contains mitochondrial
Chile (Robbins, 2004a). One lineage of Eumaeini (Callophrys loci with ‘maternal’ inheritance.
Westwood and relatives) is distributed in the Palearctic, Nearctic and There are additional reasons for focussing on Z sex chromosome
Neotropical Regions. It was suggested that geography played a central loci. Sex chromosome loci outperform autosomal ones in resolving
role in the early divergence of Callophrys and relatives (Valencia- some species trees (Corl & Ellegren, 2013). Sex linked genes have
Montoya et al., 2021). been implicated in reproductive isolation because they have a dispro-
Eumaeines are ecologically notable for the diverse array of plants portionately large effect on hybrid sterility and viability (Coyne &
eaten by its caterpillars (e.g., Beccaloni, Viloria, Hall, & Orr, 1989; Payseur, Presgraves, & Filatov, 2018; Presgraves, 2018).
HAIRSTREAK SUBTRIBE CLASSIFICATION 3
There is an elevated differentiation of the sex chromosomes com- reads or failure in alignment due to low similarity in the DNA
pared to autosomes among closely related taxa, as shown in the Z sex sequences. Reference exons were filtered to remove short exons less
chromosome of Heliconius Kluk butterflies (e.g., Kronforst et al., 2013; than 12 amino acids or redundant exons with identity greater
Martin et al., 2013; Van Belleghem et al., 2018). These findings sug- than 95%.
gest that the sex chromosome might be useful for resolving phyloge- To remove adapter sequences and low-quality bases at the end
netic relations in a rapidly diverging clade such as Eumaeini. of reads, NGS reads were processed by Trimmomatic–0.39 (Bolger,
Here we propose a comprehensive subtribal classification based Lohse, & Usadel, 2014) with parameters seedmismatches 3, pal-
on a phylogenetic analysis of 202 eumaeine species representing indromeClipThreshold 25 and simpleClipThreshold 10. We searched
almost every available generic name. Second, we summarize conspicu- each reference exon against sequence reads of samples using DIA-
ous biological and biogeographic characteristics in each subtribe. Inte- MOND (Buchfink, Xie, & Huson, 2015) with parameters l 1, comp-
grating the proposed classification with the summary datasets, we based-stats 1, masking 0 and evalue 0.01. We computed the cumula-
then seek to identify taxa and characteristics that are associated with tive sequencing depth for each reference exon among all samples. We
fast or slow diversification. The goal is to advance the hypothesis that discarded the exons that have sequencing depth more than 2.5 times
a variety of factors are likely to have affected diversification of the median depth because they may arise from repeats in the
Eumaeini. genome.
From the DIAMOND results of exons in the Calycopis reference,
we kept the reads that could be unambiguously mapped to one locus
MATERIALS AND METHODS by both E-value (<1e5 
 E-value for other loci) and sequence iden-
tity (> identity for other loci +10). We further filtered the alignments
Terminal taxa by requiring at least 75% coverage over the reads of each reference
exon and sequence identity higher than that between the reference
The 266 Eumaeini generic names are listed in Data S1. Of these, two and the sample. Because we used a number of old, dry museum speci-
are suppressed by the International Commission on Zoological mens that can be contaminated by DNA from fungi, bacteria and sur-
Nomenclature (ICZN), 16 are homonyms, three are incorrect original rounding specimens, we applied the following two protocols to detect
spellings, 12 are subsequent misspellings and 12 are nomina nuda. and remove contaminants.
Another eight are objective synonyms (the same type species as First, we set the identity cutoff of mapped reads to each refer-
another generic name), and nine have a type species that is a subjec- ence exon using the reference genome of Calephelis nemesis (W.H.
tive synonym of a type species of another genus. We sequenced rep- Edwards) in Riodinidae (Cong et al., 2017). Lycaenidae samples are
resentatives of 202 of the remaining 204 generic names (Data S2). more closely related to C. cecrops than to Riodinidae (Espeland
For each generic name, we sequenced a specimen of the type species et al., 2018). We mapped Calycopis reference exons to the C. nemesis
or a closely related species based on morphological traits. We were genome and calculated the identity between them as the identity cut-
unable to sequence specimens representing Variegata K. Johnson and off for each exon. We kept sample reads with identity higher than the
Semonina Robbins. Representatives of four species in other tribes of cutoff for each exon.
Theclinae were used as outgroups. Taxonomic authors for generic Second, for each 30 bp sliding window applied to the alignment
names in the text below are given in Data S1. between the Calycopis reference and the sample reads, we clustered
the reads into groups of similar sequences using the following proce-
dure. We ranked reads by their sequence identity to the reference
Generation of sequence alignments exons from high to low. The first read initiated a cluster. Starting from
the second read, a new read was compared to the first sequence of
DNA was extracted and genomic libraries were prepared either from each cluster and assigned to the first cluster when the first sequence
freshly collected specimens stored in RNAlater or from abdomens/ had no more than one mismatch from the current sequence. If a new
legs of dry pinned specimens in museum collections according to read could not be assigned to existing clusters, a new cluster was initi-
established protocols (Li et al., 2019; Zhang, Cong, Shen, ated. For each cluster, we computed its size and the average number
Brockmann, & Grishin, 2019). All libraries were sequenced for 150 bp of mismatches to the reference exon, and we considered a cluster to
from both ends targeting with 
5 to 
10 coverage using Illumina be good if its size was at least half of the largest cluster size and the
HiSeq X10. number of mismatches was no larger than the minimal mismatches
We used the Calycopis cecrops (Fabricius) genome (Cong among all clusters. If the number of good clusters was no more than
et al., 2016) as a reference to assemble genomic sequences. We 2 (diploid genome), we discarded the reads not included in the good
focussed on coding sequences and aligned the sequencing reads of clusters. Alignments of mitochondrial genes were constructed using a
these samples with the amino acid sequences of exons in the similar pipeline, except that we allowed only 1 good cluster per win-
Calycopis reference. For more distantly related samples, alignment dow because the mitogenome is haploid.
based on protein sequences increases accuracy and sensitivity After this cleaning procedure, the dominant nucleotide
(Pearson, 2013), reducing the chance of aligning nonorthologous (frequency > 0.6) at each position in the sequence alignment was used
4 ROBBINS ET AL.
to generate the exon sequences of a sample. The exon sequences other possible subtribal circumscriptions and give the reasons for the
were further translated to amino acid sequences and sequences of options that we chose. In a few cases, subtribes were not monophy-
different exons of a protein were concatenated to obtain the protein letic in an analysis of mitochondrial sequences or in a previous molec-
sequence of a sample. Taxon information and sequencing data can be ular phylogeny (Valencia-Montoya et al., 2021). These results are
accessed through NCBI Bioproject ID PRJNA778531. summarized (Table 1), and we discuss possible reasons for them.
For each subtribe, we present a diagnosis that includes conspicu-
ous morphological traits that are characteristic of that subtribe. To
Construction of phylogenetic trees satisfy ICZN Code Articles 13.1 and 13.2, we differentiate each
subtribe with molecular sequence synapomorphies (Table 2). Nomen-
We constructed phylogenetic trees for autosomes, Z chromosome clatural issues are addressed in Data S1, where we modify those cur-
and mitogenome, respectively. Gene content of the Lepidoptera Z rently recognized genera that were not monophyletic in the
chromosome is highly conserved (Fraïsse, Picard, & Vicoso, 2017), phylogenetic results based on autosomal, Z sex chromosome and
therefore we aligned Calycopis cecrops exons using TBLASTN (evalue mitochondrial sequences (with one exception noted below). Provision-
0.001, seg no) (Altschul, Gish, Miller, Myers, & Lipman, 1990) with ref- ally included genera are also listed in each subtribe account to facili-
erence to the Heliconius melpomene genome where the Z chromosome tate communication.
sequence is known. We identified Calycopis cecrops exons as Z-linked The taxonomic composition of each subtribe is summarized. The
if their best TBLASTN hit was on the Heliconius Z chromosome. Genes number of included genera that we provisionally list in Data S1 is
with more than 80% exons mapped to the Z chromosome were con- given. The number of species is presented as a range. The lower
sidered Z-linked. bound is the number of species with scientific names. The upper
We removed positions from the alignments that are present in bound adds an estimate, based on our research, of the number of spe-
less than 40% of the samples, and the 13.87 million remaining posi- cies in museum collections that lack scientific names.
tions in the alignment were used for phylogenetic analysis. Analysing
almost 14 million bp poses practical difficulties, for which reason we
generated 50 samples consisting of 100,000 codons each using a Biological traits
“random.sample” function (//docs.python.org/3/library/random.html).
In this random sampling procedure, each autosomal position is We summarize the occurrence of male secondary sexual organs in
expected to be represented in one of the 50 trees each subtribe and note instances in which the morphology of these
(300,000 
 50/13,870,000 = 1.08). We used IQ-TREE (Nguyen, organs needs better documentation. Terminology for these organs fol-
Schmidt, von Haeseler, & Minh, 2015) (version 1.6.10) with the GTR lows Martins et al. (2019b).
+ GAMMA model to construct a maximum-likelihood phylogenetic All Eumaeini subtribes inhabit the American tropics, but we point
tree for each of the 50 sampled alignments. The resulting trees were out those subtribes that also occur elsewhere. We note subtribes that
used to generate a consensus tree with sumtrees.py in the dendropy are primarily restricted to montane or wet habitats. We present infor-
package (Sukumaran & Holder, 2010). A similar procedure was used mation, when available, on the incidence of sympatry among closely
to derive the Z chromosome-based phylogeny. Bootstrap support related species. For genera that have not been revised, distributional
values may not be appropriate for large datasets (Lemoine and habitat information is primarily taken from large faunal works
et al., 2018). The large number of base pairs in this study almost (e.g., Godman & Salvin, 1887; Draudt 1919–1920), augmented with
always resulted in 100% values. Instead, we calculated the proportion information from the museum collections with which we work. Biogeo-
of times that a node was present in each of the 50 trees. For graphic region for each generic type species is recorded in Data S4.
mitogenomes, a maximum-likelihood tree was constructed from the We summarize patterns of caterpillar food plant specificity and
13 mitochondrial protein-coding genes with IQ-TREE. The best model myrmecophily for each subtribe. Published caterpillar food plant
was automatically selected by a Bayesian information criterion records are yet scanty. Misidentifications and a lack of vouchers have
implemented in IQ-TREE. The ultrafast bootstrap (bb 1000) in IQ- been problematic (i.e., Cajé et al., 2021). For these reasons, we cite
TREE was used to estimate the confidence of the phylogenetic tree of original published records in which adults were illustrated or in which
mitogenomes. the identifications were verified by museum vouchers, such as the
food plant records reported in Guagliumi (1965) and Zikán and
Zikán (1968). We also add data from reared vouchers in public institu-
Taxonomy tions that we examined and identified. These food plant records are
intended to point out major patterns of specialization, generalization
We partitioned Eumaeini into subtribes based on monophyly in ana- and host shifts, but they are not exhaustive. Obligate myrmecophily is
lyses of both autosome and Z sex chromosome protein-coding DNA unreported in Eumaeini, but some species are facultatively myrme-
base pair sequences (Table 1). Among monophyletic lineages, we cophilous (DeVries, 1990, 1991). Myrmecophily is not noted in most
chose those that we considered to be most useful for communicating publications on food plants. When noted, the remarks are usually
biological and biogeographic information. In the discussion, we note anecdotal.
HAIRSTREAK SUBTRIBE CLASSIFICATION 5
T AB L E 1 Monophyly of subtribes from analyses of autosomes, Z sex chromosomes, mitochondrial DNA and a variety of DNA sequences
(Valencia-Montoya et al., 2021)
Subtribe Dataset autosomes Dataset Z sex chromosome Dataset mitochondria Valencia-Montoya et al. (2021)
Eumaeina monophyletic monophyletic monophyletic Thestius does not cluster
Rhammina monophyletic monophyletic Balintus does not cluster Balintus does not cluster
Timaetina monophyletic monophyletic Busbiina does not cluster monophyletic
Atlidina monophyletic monophyletic monophyletic monophyletic
Evenina monophyletic monophyletic monophyletic monophyletic
Jantheclina monophyletic monophyletic Paraphyletic with respect to Allosmaitia and Aveexcrenota do not
Evenina cluster
Paiwarriina monophyletic monophyletic monophyletic paraphyletic with Thestius
Cupatheclina monophyletic monophyletic monophyletic monophyletic
Parrhasiina monophyletic monophyletic monophyletic monophyletic
Ipideclina monophyletic monophyletic monophyletic not sequenced
Calycopidina monophyletic monophyletic monophyletic monophyletic
Strymonina monophyletic monophyletic monophyletic monophyletic
Strephonotina monophyletic monophyletic monophyletic monophyletic
Trichonidina monophyletic monophyletic Megathecla does not cluster monophyletic
Callophryidina monophyletic monophyletic monophyletic monophyletic
Note: The placements of six of 202 generic names – Thestius, Busbiina, Balintus, Allosmaitia, Aveexcrenota and Megathecla – are variable and account for
virtually all the nonmonophyletic results at the subtribal level.
T AB L E 2 Differentiating gene sequences for Eumaeini subtribes
Subtribe Distinguishing genetic sequences
Eumaeina cce.2894.13.5:G2312A, cce.117602.1.3:T503A, cce.117602.1.3:C514G,
cce.1920.4.2:A1922G
Rhammina cce.3413.13.1:T337A, cce.3413.13.1:G338T, cce.2784.3.6:A1508G, cce.3516.7.1:G692A
Timaetina cce.4657.1.3:C1418A, cce.6475.2.1:A4115T, cce.6475.2.1:A4115T, cce.2595.1.2:G1441A
Atlidina cce.6475.2.1:T3586A, cce4260.2.3:T859A, cce.694.7.7:G2008A, cce6475.2.1:G2620A
Evenina cce.179145.1.1:A511C, cce.5263.2.1:T2410A, cce.302667.2.2:T2894C, cce.72976.7.4:A4403T
Jantheclina cce.165326.13.1:C221A, cce957.9.4:A9254C, cce.621.3.3:T164C, cce.621.3.3:C142T
Paiwarriina cce.13174.19.5:T139A, cce.3074.1.4:G226C, cce.13686.1.3:T231C, cce.483.7.2:A10C
Cupatheclina cce.1467.11.2:A1126C, cce.2790.1.3:A67G, cce.5392.4.1:T200C, cce.3034.5.1:G97C
Parrhasiina cce.1806.8.2:A344G, cce.993.29.4:A43C, cce.1546.4.5:T827A, cce.3911.8.16:C119T
Ipideclina cce.557.5.1:A1330T, cce.1546.6.3:C158G, cce.557.5.1:A602C, cce2207.3.1:A712G and not cce.4319.10.3:943C,
not cce.4260.2.3:2090G, not cce.7187.5.1:G862A, not cce4319.10.3:294A
Calycopidina cce.312.2.3:A598G, cce.8343.11.4:C4475A, cce.2805.8.6:T109G, cce.2805.8.6:C110A
Strymonina cce.1367.1.1:A437T, cce.2423.1.2:T4300A, cce.9657.10.14:C25A, cce.2070.8.17:G251C
Strephonotina cce.663.6.2:A610G, cce.419.10.2:G433A, cce.2423.1.2:A2467C, cce.2041.8.30:A584G
Trichonidina cce.1162.12.1:A3868C, cce.3869.2.3:A106C, cce.3869.2.3:A110G, cce.1162.12.1:G6708C
Callophryidina cce.7057.16.1:A193C, cce.303173.8.11:T185C, cce.7057.16.1:A310G, cce.6582.6.9:C2185A
Note: Character states are given as abbreviations, such as cce.2894.13.5:G2312A. The “cce” refers to the Calycopis cecrops reference genome (Cong
et al., 2016). To satisfy ICZN code articles 13.1 and 13.2, the character “cce.2894.13.5:G2312A” in words is “position 2312 of gene 13 and exon 5 in
scaffold 2894 in the annotated Calycopis cecrops genome has nucleotide G in the coding strand (5’ to 3’ direction), which is differentiated from other
lineages, which have nucleotide A in the coding strand”.
We discuss diversification in each subtribe by briefly with the purpose of doing a quantitative analysis of diversifica-
assessing the evolution of morphological, biogeographic and eco- tion. Rather, as noted, the intention is to point those subtribes
logical traits using the phylogenetic results as a framework. The where different factors were likely to have influenced
dataset in this paper was selected for taxonomic reasons, not diversification.
6 ROBBINS ET AL.
Museum acronyms Entomology Research Museum); USNM (USA, Washington D.C.,
Smithsonian Institution, National Museum of Natural History); and
CPAC (Brazil, Distrito Federal, Planaltina, EMBRAPA, Centro de UWIZM (Trinidad and Tobago, St. Augustine, University of the West
Pesquisas Agropecuárias do Cerrado); DZUP (Brazil, Paraná, Curitiba, Indies Zoological Museum).
Universidade Federal do Paraná, Coleç~ao de Entomologia Pe. Jesus
Santiago Moure); FIOC (Brazil, Rio de Janeiro, Rio de Janeiro, Fun-
daca~o Instituto Oswaldo Cruz); MCZ (USA, Massachusetts, Harvard RESULTS
University, Museum of Comparative Zoology); MGCL (USA, Florida,
Gainesville, University of Florida, Florida Museum of Natural History, Phylogenetic analyses
McGuire Center for Lepidoptera and Biodiversity); MIZA (Venezuela,
Maracay, Museo del Instituto de Zoología Agrícola); MNCR We illustrate maximum likelihood phylogenetic trees for 202 genera
(Costa Rica, San Jose, Museo Nacional de Costa Rica); TAMU (USA, (sequenced specimens listed in Data S2) based on analyses of
Texas, College Station, Texas A & M University); UCRC (USA, Califor- autosomal (Figure 1), Z sex chromosome (Figure 2) and mitochondrial
nia, Riverside, University of California, Department of Entomology, (Figure 3) protein-coding sequences, with relative node support values
F I GU R E 1 Maximum likelihood phylogenetic relationships of Eumaeini based on 13.5 million autosome protein-coding base pairs. Ipideclina
contains only the genus Ipidecla (red)
HAIRSTREAK SUBTRIBE CLASSIFICATION 7
in a more traditional format in Data S3. Sequencing resulted in 13.87 mil- not cluster (Table 1). Topology of the autosome and Z sex chromosome
lion autosomal bp and 368 thousand Z sex chromosome bp (2.6% of the trees (Figures 1 and 2) is highly congruent from Callophryidina (pur-
sequenced nuclear genome). The 13 mitochondrial protein-coding genes ple upper right) to Calycopidina (green lower left). Relationships
were composed of 11,130 bp (less than 0.1% of the nuclear genome). among Eumaeina + Timaetina + Rhammina + Atlidina + Evenina +
The fifteen subtribes that we recognize are monophyletic in ana- Jantheclina + Paiwarriina + Cupatheclina + Parrhasiina differ, and
lyses of both the autosome and Z sex chromosome sequences. Eleven some lineages have short branch lengths (little genomic change).
subtribes are monophyletic in the analysis of mitochondrial sequences, Although topology of the mitochondria tree (Figure 3) is generally
and the other four subtribes are monophyletic except for one genus in similar to those of the autosome and Z sex chromosome trees, there
each that does not cluster or one case of paraphyly (Table 1). Ten sub- are many differences. Biogeographic region for each of the
tribes are monophyletic in the illustrated tree in Valencia-Montoya 202 generic type species is recorded in Data S4. In sum, 169 generic
et al. (2021), one subtribe was not sequenced and the other four sub- type species are Neotropical, 18 are primarily Nearctic and 15 are
tribes are monophyletic except for one or two genera in each that do Palearctic.
F I GU R E 2 Maximum likelihood phylogenetic relationships of Eumaeini based on 368 thousand Z sex chromosome protein-coding base pairs.
Ipideclina contains only the Ipidecla (red)
8 ROBBINS ET AL.
F I GU R E 3 Maximum likelihood phylogenetic relationships of Eumaeini based on 11,130 mitochondrial protein-coding base pairs. Ipideclina
contains only the genus Ipidecla (red). The subtribes Timaetina, Rhammina, Jantheclina, and Trichonidina are not monophyletic
Currently recognized genera that were not monophyletic in the
Systematics and biology
autosome, Z sex chromosome and mitochondria trees (with the
exception of Enos, discussed below) were modified. For example, the
former characterization of Thepytus (Robbins, Busby, & Duarte, 2010) EUMAEINA DOUBLEDAY
was not monophyletic in the autosome and Z chromosome analyses,
thus we provisionally split it into Thepytus and Beatheclus. As another Included Genera. Eumaeus Hübner, Theorema Hewitson, Mithras
example, Timaeta was paraphyletic in terms of Temecla (Robbins & Hübner, Micandra Staudinger, Brevianta K. Johnson, Kruse &
Busby, 2008) in all analyses, so we combined them. We tried to mini- Kroenlein, Thestius Hübner.
mize the number of changes, but the large genera Calycopis
(Calycopidina) and Nicolaea (Strephonotina) as previously recognized
were not monophyletic, resulting in the provisional listing of more Diagnosis
genera. Phylogenetic analyses of species within each subtribe, espe-
cially Calycopidina and Strephonotina, will be needed to propose a Eumaeina here is equivalent to the Eumaeus Section of Robbins (2004-
more stable generic classification. b), which had been characterized primarily by brush organ traits, plus
HAIRSTREAK SUBTRIBE CLASSIFICATION 9
F I GU R E 4 Legend on next page.
10 ROBBINS ET AL.
Micandra, Brevianta and Thestius sensu stricto, but without Paiwarria and in the aposematically coloured Eumaeus (Figure 4a) increased diversifi-
Paraspiculatus. A possible morphological synapomorphy for Eumaeina is cation rate, subsequent amplified rates of gene changes were arguably
the poor development of forewing veins mdc and ldc, as noted for more biologically significant (Robbins et al., 2021).
Eumaeus by Bates (1861). Subsequently, this character has been used
rarely, probably because development of these forewing veins is vari-
able and difficult to score. Eumaeina is monophyletic in the analyses of RHAMMINA PRIETO & BUSBY, NEW
autosomal, Z sex chromosome and mitochondrial DNA sequences, but SUBTRIBE
not in Valencia-Montoya et al. (2021) as Thestius s.s. was embedded in
the sister lineage to Eumaeina (Table 1). Eumaeina is distinguished from http://zoobank.org/ urn:lsid:zoobank.org:act:C9735B7D-DEF7-480C-
other subtribes based on DNA sequence data only (Table 2). 8D10-46B0F53D2532.
Type Genus. Rhamma K. Johnson.
Other Included Genera. Balintus D’Abrera, Johnsonita Salazar &
Male secondary sexual organs Constantino, Lathecla Robbins, Podanotum Torres & K. Johnson, Sal-
azaria D’Abrera & Bálint.
Semi-hemispherical abdominal brush organs are unique to species of
Eumaeus, Theorema and Mithras (Robbins et al., 2021). Piliform scales on
the dorsal hindwing of male Eumaeus childrenae (Grey) are absent in Diagnosis
females and may be androconia (Robbins et al., 2021). A dorsal forewing
scent pad occurs in all species of Mithras, Micandra and Brevianta, but in Rhammina represents part of theMicandra Section of Robbins (2004b) with
some species of the latter two, there are additional complex androconial the addition of Lathecla (Robbins & Busby, 2015). A “fan-shaped” signa of
structures that have not been detailed morphologically. A dorsal hindwing the female genitalia occurs in all genera other than Salazaria, but also occurs
scent patch occurs in Thestius, but is often absent in small individuals. in some species of Micandra (Eumaeina), in some genera of Timaetina, and
in most genera of Calycopidina, where this structure was detailed morpho-
logically (Duarte & Robbins, 2010). Rhammina is monophyletic in the ana-
Biogeography, habitat and larval food plants lyses of autosome and Z sex chromosome sequences. Except for the
monotypic Balintus, it is also monophyletic in the mitochondrial tree and in
The genera of Eumaeina occur in most of the forested Neotropics the results in Valencia-Montoya et al. (2021) (Table 1). Rhammina is distin-
(Data S4), including the northern Antilles. Some are primarily lowland guished from other subtribes based on DNA sequence data (Table 2).
(Mithras, Theorema, Thestius) whereas others are primarily montane
(Micandra, Brevianta). Eumaeus is a caterpillar food plant specialist on
Zamiaceae (Robbins et al., 2021), but the other genera are recorded Male secondary sexual organs
from Fabaceae and a variety of other Angiosperm families (Robbins
et al., 2021). Caterpillars of Micandra are myrmecophilous whereas Rhammina contains a variety of male secondary sexual organs on the
those of Eumaeus are not (DeVries, 1991). wings. Unique organs occur on the wings of Lathecla and Johnsonita
(Bálint et al., 2021; Robbins & Busby, 2015). Podanotum lacks male
secondary sexual organs (Busby, Faynel, Moser, & Robbins, 2017).
Diversification Scent pads are universal in Rhamma, the most species-rich genus,
except for species such as R. anosma (Draudt). Abdominal brush
Subtribe Eumaeina consists of six genera containing 34–38 species. No organs are unrecorded in Rhammina.
more than two species of the primarily lowland genera Theorema,
Mithras, Eumaeus and Thestius s.s. occur at a locality (13 species com-
bined). In contrast, the incidence of sympatry is higher in the primarily Biogeography, habitat and larval food plants
montane Micandra and Brevianta (25 species combined). For example,
five Micandra species are sympatric at 1600–1800 m elevation in east- The genera of Rhammina are Andean endemics (Figure 4b) except for
ern Ecuador, and five Brevianta species are sympatric in the mountains Lathecla, which occurs from Mexico to southern Brazil in lowland and
of Panama (unpublished data). Although the switch to eating cycads montane habitats (Robbins & Busby, 2015). Johnsonita was reported from
F I GU R E 4 (a) warningly coloured adult of Eumaeus atala (Poey) (Eumaeina); (b),(c) Andean Rhamma (Rhammina) (top) and Penaincisalia
(Timaetina), representative montane species; (d),(e) Arcas cypria (Geyer) (Atlidina) dorsal forewing (left) with a discal cell brown scent pad and a
grey scent patch, and ventral hindwing with scent pouch opening; (f),(g) Evenus regalis (Cramer) (left) and E. temathea (Hewitson) (Evenina). The
latter resembles the wing pattern of satyrine Nymphalidae; (h),(i) Annamaria rhaptissima (K. Johnson) (Paiwarriina) with male (left) and female
sexually dimorphic ventral wing patterns; (j) Dabreras teucria with a scent patch on the dorsal hindwing costa (Cupatheclina); (k) Oenomaus
ortygnus (Cramer) (Parrhasiina), a commercial crop pest of Annonaceae. Scale refers to set specimens
HAIRSTREAK SUBTRIBE CLASSIFICATION 11
Panama (Draudt, 1919–1920), but there are no vouchers. A photograph occur in habitats at 4000 m elevation (Bálint et al., 2019). Timaeta was
taken in Ecuador was incorrectly attributed to Panama (Bálint et al., 2021). reared from Melastomataceae (Badenes-Pérez, Alfaro-Alpízar, &
Efforts to sample high elevation habitats more broadly have improved the Johnson, 2010) under the generic name Temecla, but there are no
ability to assess intraspecific geographic variation in Rhamma (Prieto, records for myrmecophily.
Núñez, & Hausmann, 2018; Prieto & Vargas, 2016). Rhamma has been
reared from Fabaceae and Melastomataceae (Arregui & Onore, 1989;
Callaghan, 2008). There are no records of myrmecophily. Diversification
The subtribe Timaetina consists of seven genera containing 102–128
Diversification species, with the vast majority of the species with a montane distribu-
tion. Phylogenetic revisions of Timaeta and Paraspiculatus showed a
The subtribe Rhammina consists of 6 genera containing 58–74 species. high incidence of sympatric diversification (Busby et al., 2017;
With the exception of Lathecla, this subtribe is endemic to the Andes. Robbins & Busby, 2008).
TIMAETINA BUSBY & PRIETO, NEW ATLIDINA MARTINS & DUARTE, NEW
SUBTRIBE SUBTRIBE
http://zoobank.org/ urn:lsid:zoobank.org:act:C5D9818A-ABA9-4891- http://zoobank.org/ lsid:zoobank.org:act:D67CA146-F37A-46FB-
926A-1ADD3D304ACE 8DDB-2E754E0F57EF
Type Genus. Timaeta K. Johnson, Kruse & Kroenlein. Type Genus. Atlides Hübner.
Other Included Genera. Penaincisalia K. Johnson, Busbiina Robbins, Other Included Genera. Brangas Hübner, Denivia K. Johnson, Arcas
Phothecla Robbins, Marachina Robbins, Nesiostrymon Clench, Para- Swainson, Theritas Hübner, Pseudolycaena Wallengren.
spiculatus K. Johnson & Constantino.
Diagnosis
Diagnosis
Atlidina represents the Atlides Section of Martins et al. (2019a,
Timaetina represents part of theMicandra Section of Robbins (2004b) with 2019b) with Dabreras transferred to Cupatheclina. A phylogeny
the addition of Marachina and Paraspiculatus. “Fan-shaped” signa (see based on morphology (Martins et al., 2019a, 2019b) differed from
Rhammina) are present in all genera except Paraspiculatus and some Pen- the phylogeny in this paper primarily by the location of the root.
aincisalia (Busby et al., 2017; Prieto, Bálint, Boyer, & Mico, 2008; Prieto, Notable morphological features of Atlidina are homoplastic. A cleft
Grishin, Hausmann, & Lorenc-Brudecka, 2016). Timaetina is monophyletic hindwing anal lobe (Godman & Salvin, 1887; illustrated in Martins
in the analyses of autosome and Z sex chromosome sequences and in the et al., 2019a) occurs in all Atlidina (Martins et al., 2019a, 2019b),
results of Valencia-Montoya et al. (2021). It was monophyletic in the mito- but also in Dabreras (Cupatheclina) and some species of Panthiades
chondria tree except for Busbiina (Table 1). Timaetina is distinguished from (Strymonina). A process of the male genitalia vinculum lying under
other subtribes based on DNA sequence data only (Table 2). the brush organs occurs in all Atlidina that possess brush organs,
but also in Evenus (Evenina), Aveexcrenota (Jantheclina) and
Dabreras (Cupatheclina). Atlidina is monophyletic in analyses of
Male secondary sexual organs autosome, Z sex chromosome and mitochondrial sequences and is
congruent with results presented by Valencia-Montoya et al. (2021).
A variety of male secondary sexual organs on the wings occur in this Atlidina is distinguished from other subtribes based on DNA
subtribe. Although scent pads are a characteristic of most species of sequence data only (Table 2).
Penaincisalia, unique organs occur in Timaeta and Phothecla
(Robbins & Busby, 2008; Robbins & Duarte, 2004). The male second-
ary sexual organs on the forewings of Marachina need better docu- Male secondary sexual organs
mentation. Paraspiculatus lacks male secondary sexual organs (Busby
et al., 2017). Abdominal brush organs are not reported in Timaetina. Martins et al. (2019a, 2019b) documented seven different kinds of
male secondary sexual organs in Atlidina (Figure 4d,e). They found
that evolutionary gains of these organs occurred primarily when spe-
Biogeography, habitat and larval food plants cies were sympatric with their sister lineage and their loss when spe-
cies were not sympatric with their sister lineage. Many males in this
The genera of Timaetina are primarily montane. The species-rich Pen- subtribe produce scents that are perceptible to people (Robbins
aincisalia (Figure 4c) is endemic to the Andes, where species may et al., 2012).
12 ROBBINS ET AL.
Biogeography, habitat and larval food plants Valencia-Montoya et al. (2021). Evenina is distinguished from other
subtribes based on DNA sequence data only (Table 2).
Atlidina is a widespread Neotropical subtribe with one species in temper-
ate areas of North America (Atlides halesus [Cramer], Data S4) and one in
the subtropical areas in South America (Atlides thargelia [Burmeister]). Male secondary sexual organs
Although primarily denizens of wet forest, a few species, such as Atlides
gaumeri (Godman) and Pseudolycaena dorcas (H. H. Druce), occur As noted, upturned scales on an enlarged frons of the male head occur
most frequently in drier, sparsely-forested habitats. Caterpillars of Denivia in some species of Evenus as well as Janthecla and Laothus (Jantheclina)
are generalists on Bombacaceae, Chrysobalanaceae, Euphorbiaceae, and are usually associated with short “longitudinal” scales along the
Fabaceae, Lecythidaceae, Meliaceae and Sterculiaceae (Müller, 1878; forewing costa. Some Evenus, such as E. coronata (Hewitson), also have
Guppy, 1904; Hoffmann, 1930, 1933; Monte, 1934; Kirkpatrick, 1954; androconia on the costa of the dorsal hindwing, on the inner margin of
Zikán, 1956; Guagliumi, 1967; Muyshondt, 1973; Robbins & Aiello, 1982; the ventral forewing, and at the base of the cubital ventral forewing
reared adults in MIZA and USNM). Similarly, the larvae of Pseudolycaena vein. Evenus regalis (Cramer) lacks androconia on the wings. Paired dor-
are generalists on Anacardiaceae, Celastraceae, Combretaceae, sal brush organs are associated with a process of the vinculum, much as
Myrtaceae, Rosaceae, Ulmaceae, Urticaceae, Annonaceae, Fabaceae, they are in the Atlidina and in Aveexcrenota (Jantheclina).
Malpighiaceae, Meliaceae, Euphorbiaceae and Sapotaceae (summarized in
Austin et al., 2007). In contrast, the caterpillars of Atlides and Brangas spe-
cialize on mistletoe plant families Loranthaceae and Viscaceae Biogeography, habitat, and larval food plants
(Sepp, 1829–1852; Hoffmann, 1937; Zikán, 1956; Whittaker, 1984;
Janzen & Hallwachs, 2021; reared vouchers in USNM). Myrmecophily Evenus is Neotropical with one species in the subtropical areas in South
does not occur in Denivia and Pseudolycaena (DeVries, 1990, 1991). America (Evenus latreillii (Hewitson)). Most species occur in lowland wet
forest, but the E. coronata clade (three species) is strictly montane. Cat-
erpillars of Evenus eat the new growth of Sapotaceae (summarized in
Diversification Robbins, 2004a; Sermeño, Robbins, Lamas, & Gámez, 2013). Flowering
and new growth for some Sapotaceae trees is seasonal, so that there
This subtribe consists of six genera containing 74–81 species. Atlidina appears to be one adult brood a year (Janzen & Hallwachs, 2021;
is likely an appropriate subtribe for investigating the evolution of male Schultze-Rhonhof, 1938; Terra-Araújo, Faria, Ribeiro, &
secondary sexual organs because of the variety of these organs and Swenson, 2012). There are no records of myrmecophily in the Evenina.
the frequency with which they were gained and lost (Martins
et al., 2019a, 2019b).
Diversification
EVENINA FAYNEL & GRISHIN, NEW This subtribe currently consists of one genus and 16–18 species. The mon-
SUBTRIBE tane E. coronata and relatives appear to be elevationally parapatric with
the lowland E. regalis. The former has at least four kinds of male secondary
http://zoobank.org/ urn:lsid:zoobank.org:act:276DA02D-7331-4EC1- sexual organs on the head and wings whereas the latter lacks these organs.
BE6A-7F488329DA2F For this reason, Godman and Salvin (1887) did not consider them to be
Type Genus. Evenus Hübner. closely related. Evenus adults are large, conspicuous blue and green butter-
flies (Figure 4f). In three species in one lineage of Evenus, however, the
females resemble brown satyrine butterflies (Nymphalidae) (Figure 4g). The
Diagnosis only possible difference other than wing pattern in this “satyrine resem-
bling” lineage is that adults do not seem to be seasonally single-brooded.
Evenina contains the relatively autapomorphic genus Evenus.
Upturned scales on the enlarged frons of males (Godman &
Salvin, 1887; Robbins & Busby, 2009) occur in most species, but also JANTHECLINA ROBBINS & FAYNEL, NEW
in species of Janthecla and Laothus (Jantheclina). “Longitudinal” SUBTRIBE
androconia on the dorsal forewing costa (Godman & Salvin, 1887;
Neild & Bálint, 2014) occur in most species of Evenus, but also in some http://zoobank.org/ urn:lsid:zoobank.org:act:CC454EE6-F45B-4C6B-
Laothus (Jantheclina). Evenina is monophyletic and strongly supported A708-7B1B66C3EEF8.
in all molecular analyses (Table 1), but its phylogenetic placement is Type Genus. Janthecla Robbins & Venables.
inconsistent. It is the phylogenetic sister of Atlidina in the autosomal Other Included Genera. Aveexcrenota Salazar & K. Johnson; Con-
tree, of Jantheclina in the Z sex chromosome tree, of Allosmaitia trafacia K. Johnson; Allosmaitia Clench; Enos K. Johnson, Kruse &
(Jantheclina) in the mitochondria tree, and of part of the Jantheclina in Kroenlein; Laothus K. Johnson, Kruse & Kroenlein.
HAIRSTREAK SUBTRIBE CLASSIFICATION 13
Diagnosis Diversification
Jantheclina is equivalent to the Allosmaitia Section of Robbins (2004b), Jantheclina consists of six genera containing 38–39 species. The phy-
which was characterized by overall genitalic similarity, with the addi- logenetically inconsistent occurrence of male secondary sexual
tion of Contrafacia, Enos and Aveexcrenota. The homoplastic occur- organs, as noted, suggests a high incidence of evolutionary gains and
rence of male secondary sexual organs is noted in the accounts of losses.
Atlidina and Evenina. Jantheclina is monophyletic in analyses of auto-
some and Z sex chromosome sequences, but paraphyletic with
respect to Evenina in the mitochondria tree. It is monophyletic in PAIWARRIINA LAMAS & ROBBINS, NEW
Valencia-Montoya et al. (2021) except for Allosmaitia and SUBTRIBE
Aveexcrenota. Jantheclina are distinguished from other subtribes
based on DNA sequence data only (Table 2). http://zoobank.org/ urn:lsid:zoobank.org:act:02D4B672-34AA-
4DC2-A3CC-8471458B1A1D
Type Genus. Paiwarria Kaye.
Male secondary sexual organs Other Included Genera. Annamaria D’Abrera & Bálint, Fasslantonius
Bálint & Salazar, Kolana Robbins.
Male secondary sexual organs in Jantheclina may be located on the
head, legs, forewings, hindwings or abdomen. According to these phylo-
genetic results, each of these organs was gained or lost at least once in Diagnosis
the subtribe. The following is a brief overview of some of the diversity
of male secondary sexual organs in Jantheclina. Upturned scales on an Paiwarriina consists of the Kolana and Paiwarria clades. Some species
enlarged frons of the male head occur in some species of Janthecla and in each lineage have a conspicuous oval scent pad at the distal end of
Laothus (as well as in the Evenina) and are often associated with short the forewing discal cell, which may be a morphological synapomorphy
“longitudinal” scales along the forewing costa, as already noted. Laothus for the subtribe, albeit superficially similar scent pads occur else-
gibberosa (Hewitson) has a unique forewing costal “hump” associated where, such as Arcas splendor (H. H. Druce) (Atlidina) (Robbins
with these scales (Draudt, 1919–1920). A “scale brush” is located on et al., 2012). Paiwarriina is monophyletic in analyses of autosome, Z
the enlarged foreleg femur of one species of Janthecla (Robbins & sex chromosome and mitochondrial sequences (Table 1), but is
Venables, 1991). Forewing scent pads occur in Allosmaitia and many paraphyletic in Valencia-Montoya et al. (2021) in terms of Thestius
species of Janthecla. An additional dorsal forewing scent patch also (Eumaeina). Paiwarriina is distinguished from other subtribes based on
occurs in Allosmaitia and Enos myrtusa (Hewitson) in which the DNA sequence data only (Table 2).
androconia are interspersed with regular wing scales (Robbins, 1987).
Varied androconial structures occur on the costa of the dorsal hindwing
in some species Janthecla, Laothus and Allosmaitia. In some Enos, the Male secondary sexual organs
hindwing costal area lacks scales. The inner margin of the ventral fore-
wings may have a scent patch, sometimes accompanied by an enlarged All described species have a dorsal forewing scent pad. Males of
hindwing costa, in Laothus, Enos and Janthecla, which is superficially Annamaria have up to four male secondary sexual organs on the wings
similar to structures in Strephonota (Strephonotina). Brush organs in (Robbins & Lamas, 2008, as Lamasina). All have a dorsal forewing
Aveexcrenota resemble those of Atlidina and Evenina. Brush organs in scent pad and a scent patch, which is often covered by regular wing
Contrafacia resemble those in Kolana (Paiwarriina) and some genera of scales. Some males have a patch of presumed androconia at the
Strymonina (Robbins, 1991a). Some species such as Janthecla leea tornus of the dorsal forewing and/or a black androconial patch at the
Venables & Robbins and Laothus viridicans (C. Felder & R. Felder) lack base of the ventral forewing. Paired dorsal brush organs abut a pro-
male secondary sexual organs. cess of the male genitalia dorsal vinculum in all genera except
Annamaria. In Paiwarria and Fasslantonius, these structures are similar
to those in Radissima (Callophryidina). In Kolana, they are similar to
Biogeography, habitat and larval food plants those in Contrafacia (Jantheclina), Thereus, Rekoa, Heterosmaitia and
Arawacus (Strymonina).
The genera of Jantheclina are strictly Neotropical (Data S4). Two genera
appear to be food plant specialists as larvae: Laothus specialize on
Asteraceae (Hoffmann, 1935, 1937, Zikán, 1956; vouchers in USNM), Biogeography, habitat and larval food plants
whereas Allosmaitia feed on Malpighiaceae (Armas, 2004; Dewitz, 1879;
Gundlach, 1881; Kaminski & Freitas, 2010; Silva et al., 2011; Silva, The genera of Paiwarriina are Neotropical with most species in wet
Duarte, Araújo, & Morais, 2014). Larvae of Laothus and Allosmaitia are forest. A few, such as K. buccina (H. H. Druce) and P. aphaca
not myrmecophilous (DeVries, 1990, 1991; Kaminski & Freitas, 2010). (Hewitson), may occur in drier forested habitats. Caterpillar food plant
14 ROBBINS ET AL.
records for Kolana are Araliaceae, Connaraceae, Erythroxylaceae, Cupathecla are Flacourtiaceae and Meliaceae (vouchers in MIZA
Lythraceae, Malpighiaceae, Melastomataceae, Ochnaceae and and USNM).
Vochysiaceae (Silva et al., 2011, 2014; voucher in MNCR). Paiwarria
may specialize on Celastraceae (Diniz, Morais, & Camargo, 2001;
Silva et al., 2014), but data are scant. There are no records of Diversification
myrmecophily.
Cupatheclina consists of two genera containing three species. No
known aspects of morphology or biology support a relationship
Diversification between these two genera.
Paiwarriina consists of four genera containing 15–16 species. The
ventral wing patterns of most species in the Paiwarria clade are con- PARRHASIINA BUSBY & ROBBINS, NEW
spicuously sexually dimorphic (Figure 4h,i) whereas those of the SUBTRIBE
Kolana lineage are not. The former clade has almost twice as many
species as the latter. http://zoobank.org/ urn:lsid:zoobank.org:act:4B69E55B-49BC-493A-
A0F4-145C27B3BE61
Type Genus. Parrhasius Hübner.
CUPATHECLINA LAMAS & GRISHIN, NEW Other Included Genera. Ignata K. Johnson, Michaelus Nicolay,
SUBTRIBE Thepytus Robbins, Olynthus Hübner, Beatheclus Bálint & Dahners,
Oenomaus Hübner, Apuecla Robbins, Dicya K. Johnson, Caerofethra
http://zoobank.org/ urn:lsid:zoobank.org:act:ABE3BF4E-8E83-4C14-8CA4- K. Johnson, Symbiopsis Nicolay.
705EEFCA8D50
Type Genus. Cupathecla Bálint.
Other Included Genus. Dabreras Bálint. Diagnosis
Nicolay (1976, 1979, 1982) suggested a relationship among many of the
Diagnosis included genera on account of their robust male genitalia capsules, but
this trait also occurs in Panthiades (Strymonina). Although the male geni-
The original descriptions of Cupathecla and Dabreras did not note a talia capsules of Apuecla, Dicya and Symbiopsis are somewhat robust, the
relationship between them (Bálint, 2005; Bálint & Faynel, 2008) and relationship with the other genera is novel. Parrhasiina are monophyletic
the latter genus was not sequenced in Valencia-Montoya et al. (2021). in all molecular analyses (Table 1). Parrhasiina is distinguished from other
Cupatheclina is distinguished from other subtribes based on DNA subtribes based on DNA sequence data only (Table 2).
sequence data only (Table 2).
Male secondary sexual organs
Male secondary sexual organs
Dorsal forewing scent pads are universal except for Symbiopsis and
Cupathecla has a dorsal forewing scent pad in the discal cell extending Caerofethra. A dorsal forewing scent patch additionally occurs in some
across the disco-cellular veins, a characteristic that otherwise occurs species of Parrhasius and Michaelus (Nicolay, 1979). Paired abdominal
in Megathecla (Trichonidina). Dabreras has a dorsal scent patch on the brush organs occur in Caerofethra, Thepytus thyrea (Hewitson) and
hindwing costa (Figure 4j) and a ventral scent patch on the forewing T. epytus (Godman & Salvin) (Robbins, Busby, & Duarte, 2010). A unique
inner margin, analogous to those found in some Thereus (Robbins, single median brush organ occurs in Symbiopsis (Robbins, 2004a).
Heredia, & Busby, 2015). Dabreras has abdominal brush organs. A pro-
cess of the vinculum/tegumen abuts the ventral and inner surface of
the brush organs, a structure that otherwise occurs in Brangas Biogeography, habitat and larval food plants
(Atlidina).
This subtribe occurs widely throughout the forested Neotropics with one
species in temperate North America (Parrhasius m-album [Boisduval & Le
Biogeography, habitat and larval food plants Conte], Data S4). As far as is known most genera feed on only one or two
plant families. Olynthus is recorded from Lecythidaceae and Caryocaraceae
Cupathecla occurs in most of the forested Neotropics, including (Nicolay, 1982; Silva et al., 2011; reared adults in USNM), Oenomaus
montane habitats in the eastern Andes, whereas Dabreras is (Figure 4k, including Porthecla) from Annonaceae (Fennah, 1937;
restricted to the Amazonian Region. Larval food plant records for Ballou, 1945; Guagliumi, 1965, 1967; Silva et al., 1967–1968;
HAIRSTREAK SUBTRIBE CLASSIFICATION 15
Kendall, 1975; Kaminski et al., 2012; reared adults in CPAC, MIZA, and Diversification
USNM), Thepytus from Vochysiaceae (Silva et al., 2011), Beatheclus
from Loranthaceae (Janzen & Hallwachs, 2021; Silva et al., 2011; A single genus containing three species. Adults superficially resemble
Uchôa, Caires, Nicácio, & Duarte, 2012), Michaelus from Bignoniaceae adults of Riodinidae (Figure 5a), but there are no other evident differ-
and Fabaceae (summarized in Robbins, 2010b; Kaminski et al., 2010; ences from other subtribes.
Silva et al., 2011) and Symbiopsis from Fabaceae (Janzen &
Hallwachs, 2021; reared vouchers in USNM). In contrast, Parrhasius is
polyphagous on Araliaceae, Asteraceae, Bignoniaceae, Euphorbiaceae, CALYCOPIDINA DUARTE & ROBBINS
Fabaceae, Fagaceae, Malpighiaceae and Malvaceae (Zikán, 1956;
Clench, 1961b; Zikán & Zikán, 1968; Maes, Hellebuyck, & Included Genera. Calycopis Scudder, Serratofalca K. Johnson,
Gantier, 1999; Rodrigues, Kaminski, Freitas, & Oliveira, 2010; Pendantus K. Johnson & Kroenlein, Camissecla Robbins & Duarte,
Janzen & Hallwachs, 2021; vouchers in CPAC, FIOC). Caterpillars of Gigantorubra K. Johnson, Electrostrymon Clench, Rubroserrata
Parrhasius and Olynthus are myrmecophilous, but those of Symbiopsis K. Johnson & Kroenlein, Ziegleria K. Johnson, Arzecla Duarte &
are not (DeVries, 1990, 1991; Rodrigues et al., 2010). Robbins, Badecla Duarte & Robbins, Kisutam K. Johnson &
Kroenlein, Lamprospilus Geyer, Argentostriatus K. Johnson,
Mercedes K. Johnson.
Diversification
Parrhasiina currently consists of 11 genera containing 102–112 species. Diagnosis
Apparent caterpillar food plant specialization in most genera is contrasted
with polyphagy in Parrhasius highlights the need for more rearing data. Perhaps the most conspicuous morphological synapomorphy for
Calycopidina is a thickened lateral edge of the female 8th abdomi-
nal tergum, but this trait is homoplastic in its occurrence (Duarte &
IPIDECLINA MARTINS & GRISHIN, NEW Robbins, 2010) and is absent in Pendantus and Ziegleria. Fan-shaped
SUBTRIBE signa of the female genitalia is another widespread character in
Calycopidina (Duarte & Robbins, 2010), but also occurs in some
http://zoobank.org/ urn:lsid:zoobank.org:act:C0DFF57C-5974-478B- Rhammina, Timaetina and Eumaeina. A lack of male secondary sex-
9758-A6924D3124D7 ual organs on the wings is another widespread trait of Calycopidina
Type Genus. Ipidecla Dyar. (Duarte & Robbins, 2010), but also occurs in many other subtribes.
Duarte and Robbins (2010) inferred phylogenetic relationships
among the genera based on morphology. The primary difference in
Diagnosis the molecular phylogeny herein is that the subtribe is rooted within
Calycopis, rendering the previous concept of Calycopis not mono-
Ipidecla was proposed in Riodinidae even though some included spe- phyletic. Calycopidina is monophyletic in all molecular analyses
cies had previously been treated as lycaenids (Druce, 1909; (Table 1) and is distinguished from other subtribes based on DNA
Godman & Salvin, 1887). The genitalia are phenotypically similar to sequence data only (Table 2).
those of Penaincisalia (Robbins, 2004a), but this similarity is homoplas-
tic according to the phylogeny. Ipideclina is distinguished from other
subtribes based on DNA sequence data only (Table 2). Male secondary sexual organs
No male secondary sexual organs occur on the wings of Calycopidina.
Male secondary sexual organs However, most species possess abdominal paired brush organs
(Duarte & Robbins, 2010).
All species have a complex androconial cluster on the dorsal forewings that
has not been characterized morphologically, but appears to be a scent pad.
Biogeography, habitat and larval food plants
Biogeography, habitat and larval food plants This subtribe occurs in virtually all habitats below 3000 m elevation
from the temperate United States (C. cecrops) to subtropical parts
Ipidecla is Neotropical from Mexico to southern Brazil, most of southern South America (e.g., Calycopis caulonia (Hewitson),
commonly in deciduous forest. Larval food plant records are Badecla clarissa (Draudt)). Caterpillars of Calycopidina are leaf-litter
Anacardiaceae, Combretaceae and Fabaceae (Kaye, 1940; vouchers in detritivores (Duarte & Robbins, 2010; Robbins, Aiello, et al., 2010),
UCRC, MIZA). Myrmecophily is unknown. including seeds and mushrooms (Gripenberg et al., 2019; Nishida &
16 ROBBINS ET AL.
F I GU R E 5 Legend on next page.
HAIRSTREAK SUBTRIBE CLASSIFICATION 17
Robbins, 2020). Females lay eggs in the leaf litter where caterpillars scales, in much the same way that it is in Tmolus, Ministrymon and
feed (Duarte & Robbins, 2010). The only exception is Mercedes some Nicolaea (Strephonotina) (Robbins & Nicolay, 2002). In
(previously a part of Calycopis), in which caterpillars eat flowers on Panthiades, the scent pad is surrounded by a circle of scales that are
the plant (Silva et al., 2011; vouchers in USNM, UWIZM). Mercedes firmly embedded in the wing membrane (Robbins, 2005). The dorsal
is sister to the remainder of the subtribe in the phylogenetic forewing androconial cluster in two Thereus species was originally
results. The caterpillars of Camissecla are not myrmecophilous described as an “oblique ovular sac-like brand occupying most of the
(DeVries, 1990, 1991). cell of the forewing, but not distinctly apparent, with a longitudinal
opening and enclosing large whitish scales” (Druce, 1907: page 591).
It is a distinctive structure that has not been documented further.
Diversification Other kinds of scent patches in the Strymonina are not restricted
to the forewing discal cell. In Thereus, some species have a scent patch
Calycopidina currently consists of 14 genera containing 131–172 spe- on the costa of the dorsal hindwing, sometimes with associated hair-
cies. An evolutionary switch from phytophagy to detritivory usually like androconia, and a second one on the ventral forewing (Robbins
inhibits diversification (Mitter, Farrell, & Wiegmann, 1988), but et al., 2015). In two species, there are erect hair-like androconia on
Calycopidina appears to be an exception. Some species appear to ovi- the inner margin of the ventral forewing (Robbins et al., 2015). In
posit only on fallen flowers/seeds and others on fallen leaves. The Arawacus, a dorsal forewing scent patch occurs either in the discal cell
number of species without scientific names in this subfamily is large, or distal of it. In some species, a second scent patch is located on the
and the 172 species upper limit is conservative. The relatively nonde- dorsal surface of the hindwing (Robbins, 2010b). In Panthiades, a dor-
script wing patterns and lack of male secondary sexual organs on the sal forewing scent patch is located distal of the discal cell
wings make it difficult to recognize specific differences. Wing pattern (Robbins, 2005).
sexual dimorphism in some genera, such as Lamprospilus (Figure 5b), Paired abdominal brush organs are associated with dorsal pro-
also contributes to difficulties with species recognition. cesses of the vinculum in Thereus and Rekoa and in some species of
Arawacus (Robbins, 1991a, 2000; Robbins et al., 2015).
Robbins (1991a) detailed their structure, which is indistinguishable
STRYMONINA TUTT from those in Contrafacia (Jantheclina) and Kolana (Paiwarriina).
Paired dorsal and ventral brush organs occur in Thereus (Robbins
Included Genera. Thereus Hübner, Rekoa Kaye, Heterosmaitia Clench, et al., 2015), which otherwise only occur in some Chalybs
Arawacus Kaye, Strymon Hübner, Hypostrymon Clench, Panthiades (Callophryidina) (Faynel, 2019). Most Strymon have brush organs
Hübner. (Robbins & Nicolay, 2002), but the vinculum is unmodified. There
are no brush organs in Panthiades and Hypostrymon (Clench, 1975;
Nicolay, 1976).
Diagnosis
This subtribe was not predicted in classifications based on morphol- Biogeography, habitat and larval food plants
ogy. Despite the lack of supporting morphological evidence,
Strymonina is monophyletic in all analyses (Table 1) and is distin- Strymonina has the broadest geographic range of the subtribes in the
guished from other subtribes based on DNA sequence data only New World, extending from Canada to the central valley of Chile in
(Table 2). virtually all vegetated habitats. Strymonina contains many widespread
and common species, especially members of Heterosmaitia, Strymon
and Panthiades, noted for larval feeding on at least a dozen plant fami-
Male secondary sexual organs lies (Janzen & Hallwachs, 2021; Monteiro, 1991; Robbins, 1991a;
Robbins & Nicolay, 2002). Alternatively, some lineages primarily spe-
The typical organs in this subtribe are a scent patch in the dorsal fore- cialize on Loranthaceae (about 35 Thereus species), Solanaceae (about
wing discal cell and a scent pad on the radial/disco-cellular veins, as 16 Arawacus species) or Bromeliaceae (about 16 Strymon species)
documented in Rekoa (Robbins, 1991a). One or both may be absent. (Heredia & Robbins, 2016; Robbins, 1991a, 2000, 2010a). There are
In Strymon, the discal cell scent patch is covered with regular wing numerous records of myrmecophilous behaviour (i.e., DeVries, 1990,
F I GU R E 5 (a), Ipidecla schausi (Godman & Salvin) (Ipideclina) superficially resembles some Riodinidae; (b) Lamprospilus genius Geyer
(Calycopidina) male (left) and female with wing pattern sexual dimorphism; (c),(d) ventral “false head” wing patterns in Panthiades phaleros
(Linnaeus) (left) and Arawacus aetolus (Sulzer) (Strymonina); (e) Ostrinotes tarena (Hewitson) with a wing pattern representative of many
Strephonotina; (f),(g) male Trichonis hyacinthus (Cramer) (left) and male Bistonina bactriana (Hewitson) (Trichonidina) with little similarity in wing
pattern or male secondary sexual organs; (h),(i),(j), Chalybs janias (Cramer) (left), Erora badeta (Hewitson) (right, top) and Chlorostrymon simaethis
(Drury) (Callophryidina) with ventral green wings. Scale refers to set specimens
18 ROBBINS ET AL.
1991; Robbins, 1991b). Conspicuous false head wing patterns (charac- Biogeography, habitat and larval food plants
terized in Robbins, 1981) occur in some species of Thereus, Panthiades
and Arawacus (Figure 5c,d). Strephonotina is Neotropical except for the subtropical part of the distri-
bution of Ministrymon leda (W.H. Edwards) in North America and the sub-
tropical parts of the distributions of M. gamma (H.H. Druce) and
Diversification M. sanguinalis (Burmeister) in South America. Caterpillar food plant usage
in Strephonotina is variable. For example, Tmolus has been reared from
Strymonina currently consists of seven genera containing 123–138 Acanthaceae, Anacardiaceae, Boraginaceae, Campanulaceae, Celastraceae,
species. Repeated caterpillar food plant specialization and/or generali- Combretaceae, Connaraceae, Fabaceae, Gesneriaceae, Lecythidaceae,
zation, as noted above, as well as the variety of male secondary sexual Malpighiaceae, Melastomataceae, Ochnaceae, Simaroubaceae, Solanaceae,
organs suggest that each of these factors may have affected Verbenaceae and Vochysiaceae (Perkins & Swezey, 1924; Lima, 1936;
diversification. Zimmerman, 1958; Robbins & Aiello, 1982; Silva et al., 2011; vouchers in
CPAC, MIZA and USNM). In contrast, Ministrymon appears to specialize
on Fabaceae (Comstock & Dammers, 1935; Ballmer & Pratt, 1989; Silva
STREPHONOTINA K. JOHNSON, AUSTIN, LE et al., 2011; Janzen & Hallwachs, 2021; vouchers in DZUP, MIZA, TAMU,
CROM & SALAZAR UCRC, USNM and UWIZM). Myrmecophilous behaviour is recorded in
Tmolus and Gargina (DeVries, 1990, 1991).
Included Genera. Theclopsis Godman & Salvin, Ministrymon Clench,
Iaspis Kaye, Crimsinota K. Johnson, Aubergina K. Johnson, Tmolus
Hübner, Celmia K. Johnson, Rindgea K. Johnson, Nicolaea K. Johnson, Diversification
Terenthina Robbins, Siderus Kaye, Gossenia Bálint, Gargina Robbins,
Decussata K. Johnson, Austin, Le Crom & Salazar, Ostrinotes Strephonotina currently consists of 16 genera containing 148–228
K. Johnson, Austin, Le Crom & Salazar, Strephonota K. Johnson, Aus- species. Perhaps the most notable trait of Strephonotina is that a
tin, Le Crom & Salazar. majority of the species have a relatively nondescript grey ventral wing
pattern with a postmedian line (Figure 5e). Compared with Atlidina
and Evenina (Figure 4d–g), for example, the adults are less colourful
Diagnosis and have fewer types of male secondary sexual organs. As with
Calycopidina, there are many undescribed species.
Strephonotina represents the Tmolus Section of Robbins (2004b) with
four genera added from the Hypostrymon Section. Many species in
Strephonotina have the female genitalia corpus bursae constricted TRICHONIDINA DUARTE & FAYNEL, NEW
medially (Faynel & Robbins, 2014; Robbins & Duarte, 2004). Many SUBTRIBE
also have sexually dimorphic forewing venation in which vein M2
arises closer to vein M1 in males than it does in females (Robbins & http://zoobank.org/ urn:lsid:zoobank.org:act:36D5508F-9355-418C-
Duarte, 2004). Strephonotina was found monophyletic in all molecular 92B5-E5AE45951D36
analyses (Table 1) and is distinguished from other subtribes based on Type Genus. Trichonis Hewitson.
DNA sequence data only (Table 2). Other Included Genera. Bistonina Robbins, Manticia Bálint, Mega-
thecla Robbins.
Male secondary sexual organs
Diagnosis
Dorsal forewing scent pads and patches occur widely in the
Strephonotina, but not all. In Tmolus, Ministrymon and some This subtribe includes genera with strikingly different wing patterns,
Nicolaea, a discal cell dorsal forewing scent patch is covered by genitalia and androconia (Figure 5f,g). Trichonis contains two sexually
regular wing scales (Robbins & Glassberg, 2013), as in the genus dimorphic species that are conspicuously different from all other
Strymon (Strymonina). A conspicuous ventral forewing scent patch eumaeines (Robbins, 1987). Eliot (1973) placed Trichonis and Micandra
occurs in Strephonota strephon (Fabricius) and relatives accompa- (Eumaeina) in the Trichonis section, but the morphological basis for
nied by an enlarged hindwing costa. Scattered androconia on the this action was disputed (Robbins, 1987). Trichonidina was recognized
ventral surface of the hindwing occur in S. jactator (H.H. Druce) in Valencia-Montoya et al. (2021) and is monophyletic in all analyses
and relatives (unpublished data). Abdominal brush organs are based on molecular sequences except Megathecla in the mitochondrial
absent except for Siderus nouraguensis Faynel & Robbins tree (Table 1); it is distinguished from other subtribes based on DNA
(Faynel & Robbins, 2014). sequence data only (Table 2).
HAIRSTREAK SUBTRIBE CLASSIFICATION 19
Male secondary sexual organs synapomorphic. As noted in Robbins and Duarte (2004), the posteriorly
developed male genitalia labides and the female genitalia ductus seminalis
All species of Trichonidina have dorsal forewing scent pads except for arising from the middle of the ductus bursae in Erora (Field, 1941) superfi-
the two species of Trichonis. Megathecla has a scent pad that extends cially resemble the genitalia of the Old World Hypochrysops C. & R. Felder
from the discal cell to forewing cells M1-M3, which also occurs in (Sands, 1986) of the tribe Luciini (Australian Region). Callophryidina is
Cupathecla (Cupatheclina). Some Bistonina have dorsal forewing scent monophyletic in Valencia-Montoya et al. (2021) and in all analyses herein
patches. The overlapping dorsal hindwing and ventral forewing scent (Table 1); it is distinguished from other subtribes based on DNA sequence
patches in Trichonis are distinctive (Robbins, 1987). Abdominal brush data only (Table 2).
organs are not recorded in Trichonidina.
Male secondary sexual organs
Biogeography, habitat and larval food plants
Dorsal forewing scent pads occur widely in Callophryidina. The scent pad
Most species of Trichonidina are restricted to the in some species of Ocaria is a complex structure in need of further study.
Amazonian Region. No members of this subtribe appear to have A dorsal hindwing scent patch occurs in several species of Erora. Paired
been reared. abdominal dorsal brush organs occur widely in the Callophryidina. In Erora
and Semonina, some species have paired dorsal brush organs that do not
touch the vinculum (Robbins & Duarte, 2004). Callophrys henrici (Grote &
Diversification Robinson) lacks male secondary sexual organs.
Trichonidina consists of four genera containing 10–14 species. As
with Cupatheclina, no known aspects of morphology or biology sup- Biogeography, habitat and larval food plants
port a relationship among these genera.
Callophryidina occurs in the Palearctic, Nearctic and Neotropical Regions
(Data S4). All Palearctic Eumaeini belong to Callophrys or Satyrium, as del-
CALLOPHRYIDINA TUTT imited for over half a century (Ziegler, 1960). Classifications, such as
those in Clench (1978) and Weidenhoffer et al. (2004), in which Satyrium
Included Genera. Callophrys Billberg, Cyanophrys Clench, Chalybs is partitioned into smaller genera, are not supported by the results.
Hübner, Ocaria Clench, Magnastigma Nicolay, Satyrium Scudder, With few exceptions, the eumaeine species that inhabit areas
Thaeides K. Johnson, Kruse & Kroenlein, Radissima K. Johnson, Erora with freezing temperatures belong to this subtribe. As noted, caterpil-
Scudder, Semonina Robbins, Chlorostrymon Clench. lars of Callophrys eat Gymnosperms, Monocotyledons and Dicotyle-
dons (Ehrlich & Raven, 1965). More generally, a vast array of larval
food plants, as well as myrmecophilous behaviour, have been
Diagnosis recorded for Callophryidina in multiple regions. The species of Holarc-
tic Satyrium appear to be obligately univoltine but many of Holarctic
Widespread conspicuous morphological traits in Callophryidina are homo- Callophrys are multivoltine.
plastic in their occurrence. Chalybs, Callophrys, Cyanophrys, Erora, Sem-
onina and Chlorostrymon have species with a green ventral ground colour
(Figure 5h–j), but based on the autosome and Z sex chromosome results, Diversification
the occurrence of this trait is homoplastic. A serrate keel on the ventral
tip of the penis (Clench, 1961a) occurs in genera Phaeostrymon, Mag- Callophryidina consists of 11 genera containing 112–131 Neotropical
nastigma, Chlorostrymon, Ocaria and most species of Satyrium. It is homo- and Nearctic species and a similar number of Palearctic species. It is
plastic in our phylogeny, and also occurs in Podanotum in Rhammina the only subtribe with Palearctic species. Valencia-Montoya
(Robbins, 2004a). The structure of the male abdominal brush organs and et al. (2021) concluded that geography played a central role in the
the associated modification of the vinculum in Radissima is almost identi- early divergence of Callophryidina.
cal to that in Paiwarria (Paiwarriina). The female genitalia ductus seminalis
arises at the dorsal anterior edge of the ductus bursae with a posterior
lobe of the corpus bursae dorsal of it (Nicolay, 1980) in Magnastigma, DISCUSSION
Chlorostrymon, Ocaria and Satyrium alcestis (W.H. Edwards). This trait does
not occur elsewhere in Eumaeini, but its occurrence is homoplastic. The Subtribal classification
species in this subtribe tend to have more beta-trichoid sensilla (see
Chun & Schoonhoven, 1973) at the distal end of the mid- and hind tibia Eumaeini is divided into 15 subtribes, each of which is monophyletic
than other Eumaeini, but variability makes this trait unlikely in the tree based on autosome protein-coding genes (Figure 1) and in
20 ROBBINS ET AL.
the tree based on Z sex chromosome protein-coding genes (Figure 2). Lepidopteran Z sex chromosome
Subtribal nomenclature is addressed in Data S1. The classification of
102 generic names in Data S1 is provisional until most species in each The maximum likelihood tree based on Z sex chromosome loci is simi-
subtribe can be sequenced and analysed. Eumaeini currently contains lar to that based on autosome sequences (Figures 1 and 2). The topol-
1074–1323 species. ogy among subtribes is the same with respect to Callophryidina to
Using non-eumaeine fossils, the ancestor where Callophryidina Strymonina. The similarities allow us to propose a classification in
split from the remainder of Eumaeini was estimated at 13.8–18.8 Ma which subtribes are monophyletic in both trees (Figures 1 and 2).
(Chazot et al., 2019) and at 23.8–33.5 Ma (Espeland et al., 2018; When analysing hundreds of taxa, the analysis of the small number of
Valencia-Montoya et al., 2021). Given these nonoverlapping ranges Z sex chromosome sequences (2.6% of nuclear base pairs) may be
and the lack of fossils of eumaeines or closely related tribes (Valencia- more practical as far as time and expense. Moreover, sex chromo-
Montoya et al., 2021), dating divergence times of eumaeine lineages somes disproportionately affect hybrid sterility/nonviability and show
is premature. elevated differences among related taxa (e.g., Kronforst et al., 2013;
The differences between the results presented here and those of Martin et al., 2013; Payseur et al., 2018), suggesting some utility for
Valencia-Montoya et al. (2021) (Table 1) may be due, in part, to their their use in phylogenetic analyses. On the other hand, divergence pat-
smaller dataset. The analysis in Valencia-Montoya et al. (2021) was terns in sex chromosomes are influenced by changes in population
based upon 187 eumaeines with between five and 374 sequenced size (Pool & Nielsen, 2007; Van Belleghem et al., 2018), indicating that
loci. The species that varied in placement – Thestius meridionalis caution be exercised when interpreting phylogenetic signal in sex
(Draudt), Balintus tityrus (C. Felder & R. Felder), Allosmaitia sp. and chromosome sequences.
Aveexcrenota anna (H. H. Druce) – were among the taxa with five
sequenced loci.
Relationships in the clade comprising subtribes Eumaeina + Diversification and morphological homoplasy
Rhammina + Timaetina + Atlidina + Evenina + Jantheclina +
Paiwarriina + Cupatheclina + Parrhasiina were not strongly resolved Although rapidly diversifying clades may be accompanied by rela-
in our results, nor in Valencia-Montoya et al. (2021). We considered tively little morphological evolution (Adams, Berns, Kozak, &
lumping these clades into one large subtribe of about 500 species Wiens, 2009), our phylogenetic results show that Eumaeini suggest
even though it was not monophyletic in the mitochondrial results many instances of conspicuous morphological homoplasy. Male sec-
(e.g., Busbiina). An advantage of this action would be a simpler classifi- ondary sexual organs represent many proposed synapomorphies of
cation. A disadvantage is that it would be a classificatory taxon that a previous classification (Robbins, 2004a, 2004b). The homoplastic
conveyed little information because of morphological, biogeographical occurrence of these traits in the Lepidoptera has long been known,
and ecological heterogeneity. For that reason, we instead chose a based on phylogenetic incongruence with other morphological
classification that was intended to better communicate biological structures (summarized in Robbins et al., 2012). The molecular phy-
information. For example, Rhammina is primarily Andean endemics, logenetic results confirm this pattern; Atlidina, Jantheclina and
Atlidina is renowned for its variety of male secondary sexual organs Evenina provide many examples. Primary sexual structures in
and Paiwarriina has a stark contrast among wing pattern sexually Eumaeini are also homoplastic. For example, a dorsal process of the
monomorphic and dimorphic sister clades. fused male genitalia vinculum/tegumen in which the vinculum
Callophryidina is the most species-rich subtribe and is sister to groove is flush with the posterior margin of the process (see
the remainder of the Eumaeini. We considered splitting this clade in Robbins, 1991a) is present in Atlidina, Jantheclina and Paiwarriina.
two, but there was no partition in which each clade was monophyletic As another example, a “fan-shaped” signa (sensu Duarte &
in all the molecular analyses. Robbins, 2010) occur in many genera of Eumaeina, Rhammina,
The classification of Eumaeini presented herein contains 102 gen- Timaetina and Calycopidina, and based on these molecular results,
era. However, Enos + Chopinia + Falerinota is not monophyletic in they have been gained or lost in each subtribe.
analysis of mitochondrial sequences (Data S3). We recognize the Homoplasy of conspicuous morphological traits in Eumaeini is not
monophyly of Enos based upon monophyly in the autosome and Z sex restricted to sexual organs. For example, our results indicate that a
chromosome analyses herein, and based on results by Valencia- “cleft” anal lobe (Martins et al., 2019a) likely evolved three times;
Montoya et al. (2021) (see Robbins, 1987). once in the ancestor of Atlidina, within the genus Panthiades
The two proposed generic names that we did not sequence are (Strymonina) and in the ancestor of Dabreras (Cupatheclina). As
placed in Callophryidina. The type species of Variegatta has a complex another example, a thickened ridge on the lateral edge of the female
cornutus in the male genitalia penis that is unique among eumaeines to 8th abdominal tergum (see Duarte & Robbins, 2010) is absent in
Ocaria, for which reason it was treated as a junior synonym of Ocaria Pendantus and Ziegleria. Similarly, males and females in some genera,
(Robbins, 2004a, 2004b). The type species of Semonina and some species such as Lamprospilus (Calycopidina) and Annamaria (Paiwarriina), have
of Erora have a pair of male abdominal brush organs that do not touch sexually dimorphic ventral wing patterns (Figures 4h,i and 5b); related
the vinculum – a unique structure in Eumaeini – for which reason they genera, such as Arzecla (Calycopidina) and Kolana (Paiwarriina) respec-
were treated as close relatives (Robbins & Duarte, 2004). tively, have monomorphic ventral wing patterns.
HAIRSTREAK SUBTRIBE CLASSIFICATION 21
Diversification and biogeography Matthew Cock, Phil DeVries, Winnie Hallwachs, Dan Janzen, the late
Roy Kendall, Olaf Mielke, Deb Murray and many others for sharing
About 85% of the species in subtribes Eumaeina, Rhammina and vouchers of reared species and related information. Thanks to Niklas
Timaetina (240 species combined) occur in the Andes (Figure 4b,c). Wahlberg and Naomi Pierce for checking an identification in the
Although stratification in narrow elevational zones has been implicated in Chazot publication. We thank William Beck, David Geale and Andrew
the diversification of some Andean butterflies (e.g., Adams, 1985; Neild for the images in Figure 4A, 4H, 4I, 10A, 10B, respectively. We are
Pyrcz & Wojtusiak, 2002), such patterns do not occur in Eumaeini (Lamas especially grateful to the curators of the museum collections noted for
et al., 2021). On the contrary, the incidence of sympatry between sister allowing us access to their collections. Robert K. Robbins acknowledges
lineages in Timaetina may be high (Busby et al., 2017; Robbins & support from an ADS core proposal grant from the National Museum of
Busby, 2008). The subtribes Eumaeina, Rhammina and Timaetina may be Natural History. Nick V. Grishin acknowledges support from the NIH
appropriate taxa to study the apparent higher incidence of sympatry in (GM127390) and the Welch Foundation (I-1505). Marcelo Duarte
montane habitats on diversification. acknowledges support from FAPESP (grants 2002/13898-0,
The Neotropical Eumaeini likely invaded Nearctic areas with 2003/13985-3 and 2016/50384-8), Coordenaç~ao de Aperfeiçoamento
freezing weather seven times, according to our results (Data S4). de Pessoal de Nível Superior (PROTAX II – grant 440597/2015-3),
Atlidina (e.g., Atlides halesus), Parrhasiina (e.g., Parrhasius m-album), Conselho Nacional de Desenvolvimento Científico e Tecnologico (grants
Calycopidina (e.g., Calycopis cecrops) and Strymonina (e.g., Strymon 305905/2012-0, 311083/2015-3 and 312190/2018-2) and
melinus [Hübner]) each appear to have colonized the Nearctic once. In Universidade de S~ao Paulo (Projeto 1, Pro-Reitoria de Pesquisa).
contrast, the Neotropical Callophryidina invaded the temperate parts
of the Nearctic, including the mountains of Mexico and Guatemala, CONFLICT OF INTEREST
three times (Callophrys, Satyrium and within Erora). The Nearctic Cal- The authors declare that there is no conflict of interest.
lophrys and Satyrium each invaded the Palearctic at least once.
Although more sequenced taxa are needed, it appears that coloniza- DATA AVAILABILITY STATEMENT
tion of the Palearctic by Satyrium led to a vastly increased rate of Data that support the findings of this study are available in the
diversification. supporting information and in NCBI Bioproject ID PRJNA778531.
Diversification and larval food plants ORCID
Robert K. Robbins https://orcid.org/0000-0003-4137-786X
Christophe Faynel https://orcid.org/0000-0001-9330-4633
Two relationships between diversification and caterpillar food plants
Gerardo Lamas https://orcid.org/0000-0002-3664-6730
are evident despite the caveat that caterpillar food plant records are
Nick V. Grishin https://orcid.org/0000-0003-4108-1153
still not well documented. Calycopidina (172 species) (Figure 5b) con-
sists of leaf-litter detritus feeding caterpillars except for Mercedes
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Nicolay, S.S. (1982) Studies in the genera of American hairstreaks. 6. A Robbins, R.K. (2004b) Lycaenidae. Theclinae. Tribe Eumaeini. In: Lamas, G.
review of the Hübnerian genus Olynthus (Lycaenidae: Eumaeini). Bul- (Ed.) Checklist: part 4A. Hesperioidea – Papilionoidea. In: Heppner, J. B.
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Payseur, B.A., Presgraves, D.C. & Filatov, D.A. (2018) Introduction: sex Robbins, R.K. (2010a) The “upside down” systematics of hairstreak butter-
chromosomes and speciation. Molecular Ecology, 27(19), 3745–3748. flies (Lycaenidae) that eat pineapple and other Bromeliaceae. Studies
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Perkins, R.C.L. & Swezey, O.H. (1924) The introduction into Hawaii of Robbins, R.K. & Aiello, A. (1982) Foodplant and oviposition records for
insects that attack lantana. Bulletin of the Experiment Station of the Panamanian Lycaenidae and Riodinidae. Journal of the Lepidopterists’
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Pierce, N.E. (1984) Amplified species diversity: a case study of an Robbins, R.K., Aiello, A., Feinstein, J., Berkov, A., Caldas, A., Busby, R.C.
Australian lycaenid butterfly and its attendant ants. In: Vane- et al. (2010) A tale of two species: detritivory, parapatry, and
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Presgraves, D.C. (2018) Evaluating genomic signatures of “the large X– est endemic. Tijdschrift voor Entomologie, 151(2), 205–233.
effect” during complex speciation. Molecular Ecology, 27(19), 3822– Robbins, R.K. & Busby, R.C. (2009) Updated phylogeny, taxonomy, and
3830. https://doi.org/10.1111/mec.14777 diversification of Janthecla Robbins & Venables (Lycaenidae:
Prieto, C., Bálint, Z., Boyer, P. & Mico, E. (2008) A review of the “browni Theclinae: Eumaeini). Journal of Research on the Lepidoptera, 41,
group” of Penaincisalia with notes on their distribution and variability 5–13.
(Lepidoptera: Lycaenidae). Zootaxa, 1941, 1–24. Robbins, R.K. & Busby, R.C. (2015) Evolutionary gain of male secondary
Prieto, C., Grishin, N., Hausmann, A. & Lorenc-Brudecka, J. (2016) The Pen- sexual structures in the widespread Neotropical montane genus
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Colombia, insights from mtDNA barcodes and the description of a new Evolution, 46(1), 47–78.
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Prieto, C., Núñez, R. & Hausmann, A. (2018) Molecular species delimitation of Thepytus (Lycaenidae, Theclinae, Eumaeini). Arthropod Systemat-
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3897/zookeys.301.5081ZooKeys hyperdiverse tribe Eumaeini (Lepidoptera: Lycaenidae). Proceedings
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Robbins, R.K. & Nicolay, S.S. (2002) An overview of Strymon Hübner Weidenhoffer, Z., Bozano, G.C. & Churkin, S. (2004) Guide to the
(Lycaenidae: Theclinae: Eumaeini). Journal of the Lepidopterists’ Soci- butterflies of the Palearctic region. Omnes Artes, Milano: Lycaenidae
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Papillons de Surinam. Amsterdam, J. C. Sepp en Zoon. 3 Deel. pp. viii, III. Lepidoptera. Pesquisa agropecuária Brasileira (Agronomia), 3,
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Sermeño, J.M., Robbins, R.K., Lamas, G. & Gámez, J.A. (2013) Cría en Zimmerman, E.C. (1958) Insects of Hawaii, Vol. 7. Honolulu: Macrolepidop-
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Systematic Botany, 25(5), 295–303. 1–25. Available from: https://doi.org/10.1111/syen.12541
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silba species (Diptera: Lonchaeidae) and Thepytus echelta
Subtribe and Provisional Generic Classification of the Eumaeini 
New World citations listed in Lamas (2022) – 
http://www.butterfliesofamerica.com/L/ref_library.htm 
 
Eumaeina Doubleday, 1847  
Type Genus Eumaeus Hübner, [1819] 
= Eumenides Boisduval, 1836 [type genus Eumenia Godart, [1824]] 
Invalid under ICZN Article 55 because Eumenides is a junior homonym of the family-group name Eumenida Leach, 
1812 (Hymenoptera), based on the wasp genus Eumenes Latreille, 1802. 
 
Eumaeus Hübner, [1819], type species: Rusticus minyas Hübner, [1809] 
         Eumenia Godart, [1824], type species: Eumenia toxea Godart, [1824] 
         Eumaea Geyer, [1834], type species: Eumaea debora Geyer, [1834] 
         Epula Gistel, 1848, repl. name for Eumenia., ICZN art. 67.8, type species: Eumenia toxea Godart, [1824] 
Theorema Hewitson, 1865, type species: Theorema eumenia Hewitson, 1865 
Mithras Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio nautes Cramer, 1779 
Micandra Staudinger, 1888, cf. Prieto (2011) for authorship, type species designation, type species: Pseudolycaena 
platyptera C. Felder & R. Felder, 1865 
         Egides Salazar, 1995, not available, nom. nud.  
         Egides K. Johnson, Kruse & Kroenlein, 1997, type species: Pseudolycaena aegides C. Felder & R. Felder, 
1865 
Brevianta K. Johnson, Kruse & Kroenlein, 1997, type species: Thecla undulata Hewitson, 1867 
         Bussa K. Johnson, Kruse & Kroenlein, 1997, not available, preocc. (Ragonot 1888), type species: Thecla busa 
Godman & Salvin, 1887  
Thestius Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio pholeus Cramer, 1777 
 
Rhammina Prieto & Busby, new subtribe 
Type Genus Rhamma K. Johnson, 1992   
 
Balintus D'Abrera, 2001, ICZN Opinion 2358 (2015), type species: Pseudolycaena tityrus C. Felder & R. Felder, 
1865  
Johnsonita Salazar & Constantino, 1995, type species: Johnsonita johnsoni Salazar & Constantino, 1995 
         Owda K. Johnson, Kruse & Kroenlein, 1997, type species: Thecla auda Hewitson, 1867  
Rhamma K. Johnson, 1992, type species: Thecla oxida Hewitson, 1870 
         Pontirama K. Johnson, 1992, (Robbins 2004, first reviser, ICZN art. 24.2), type species: Pontirama brunea K. 
Johnson, 1992 
         Shapiroana K. Johnson, 1992, (Robbins 2004, first reviser, ICZN art. 24.2), type species: Shapiroana shapiroi 
K. Johnson, 1992 
         Paralustrus K. Johnson, 1992, (Robbins 2004, first reviser, ICZN art. 24.2), type species: Thecla commodus C. 
Felder & R. Felder, 1865  
Lathecla Robbins, 2004, type species: Thecla latagus Godman & Salvin, 1887  
Podanotum Torres & K. Johnson, 1996, type species: Podanotum clarissimus Hall, Willmott & K. Johnson, 1996 
Salazaria D'Abrera & Bálint, 2001, ICZN Opinion 2358 (2015), type species: Thecla sala Hewitson, 1867 
 
Timaetina Busby & Prieto, new subtribe 
Type Genus Timaeta K. Johnson, Kruse & Kroenlein, 1997 
= Thecloxurina K. Johnson, 1992 [type genus Thecloxurina K. Johnson, 1992]. 
Unavailable under ICZN Articles 13.1 and 13.2 because Thecloxurina is a family-group name proposed after 1930 
without “a description or definition that states in words characters that are purported to differentiate the taxon.” 
 
Penaincisalia K. Johnson, 1990, type species: Thecla ? culminicola Staudinger, 1894 
         Thecloxurina K. Johnson, 1992, type species: Thecla loxurina C. Felder & R. Felder, 1865 
         Pons K. Johnson, 1992, type species: Pons magnifica K. Johnson, 1992 
         Abloxurina K. Johnson, 1992, subsequent designation, Prieto & Lorenc-Brudecka, 2014, type species: 
Penaincisalia ismaeli Busby & Hall, 2005 
         Candora K. Johnson, 1992, type species: Candora fallacandor K. Johnson, 1992 
         Ianusanta Bálint, 2011, type species: Ianusanta ianusi Bálint, 2011  
Busbiina Robbins, 2004, type species: Thecla bosora Hewitson, 1870  
Timaeta K. Johnson, Kruse & Kroenlein, 1997, type species: Pseudolycaena timaeus C. Felder & R. Felder, 1865 
         Trochusinus K. Johnson, Salazar & Vélez, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type species: 
Thecla trochus H.H. Druce, 1907 
         Jagiello Bálint & Wojtusiak, 2000, type species: Jagiello molinopampa Bálint & Wojtusiak, 2000 
         Temecla Robbins, 2004, type species: Thecla tema Hewitson, 1867  
Phothecla Robbins, 2004, type species: Thecla photismos H.H. Druce, 1907 
Marachina Robbins, 2004, type species: Thecla maraches H.H. Druce, 1912 
Nesiostrymon Clench, [1964], type species: Thecla celida shoumatoffi Comstock & Huntington, 1943 
         Terra K. Johnson & Matusik, 1988, type species: Thecla tera Hewitson, 1878 
         Sipaea K. Johnson, 1991, type species: Thecla hyccara Hewitson, 1868 
Paraspiculatus K. Johnson & Constantino, 1997, type species: Paraspiculatus colombiensis K. Johnson & 
Constantino, 1997 
 
Atlidina Martins & Duarte, new subtribe 
Type Genus Atlides Hübner, [1819] 
 
Atlides Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio halesus Cramer, 1777 
         Riojana D'Abrera & Bálint, 2001, ICZN Opinion 2358 (2015), type species: Thecla thargelia Burmeister, 
1878  
Brangas Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio caranus Stoll, 1780  
Denivia K. Johnson, 1992, type species: Thecla deniva Hewitson, 1874 
         Lucilda D'Abrera & Bálint, 2001, ICZN Opinion 2358 (2015), type species: Thecla crines H.H. Druce, 1907 
         Margaritheclus Bálint, 2002, type species: Pseudolycaena danaus C. Felder & R. Felder, 1865  
Arcas Swainson, 1832, type species: Papilio imperialis Cramer, 1775  
Theritas Hübner, 1818, subsequent designation, Scudder, 1875, type species: Theritas mavors Hübner, 1818  
Pseudolycaena Wallengren, 1858, type species: Papilio marsyas Linnaeus, 1758 
 
Evenina Faynel & Grishin, new subtribe 
Type Genus Evenus Hübner, [1819] 
= Macusiina K. Johnson, Kruse & Kroenlein, 1997 [type genus Macusia Kaye] 
Unavailable under ICZN Articles 13.1 and 13.2 because Macusiina is a family-group name proposed after 1930 
without “a description or definition that states in words characters that are purported to differentiate the taxon.” 
 
Evenus Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio regalis Cramer, 1775 
         Euenus Hübner, [1826], not available, missp.  
         Endymion Swainson, 1831, type species: Papilio regalis Cramer, 1775 
         Macusia Kaye, 1904, type species: Thecla satyroides Hewitson, 1865 
         Cryptaenota K. Johnson, 1992, type species: Thecla latreillii Hewitson, 1865 
         Ipocia Brévignon, 2000, type species: Papilio gabriela Cramer, 1775 
         Poetukulunma Brévignon, 2002, type species: Thecla sponsa Möschler, 1877 
         Poetuculunma Brévignon, 2002, not available, incorrect original spelling, type species: Thecla sponsa 
Möschler, 1877 
         Suneve Bálint, 2006, type species: Thecla coronata Hewitson, 1865 
 
Jantheclina Robbins & Faynel, new subtribe 
Type Genus Janthecla Robbins & Venables, 1991 
 
Aveexcrenota Salazar & K. Johnson, 1997, type species: Thecla anna H.H. Druce, 1907  
Contrafacia K. Johnson, 1989, type species: Contrafacia mexicana K. Johnson, 1989 
         Orcya K. Johnson, 1990, type species: Thecla orcynia Hewitson, 1868  
Allosmaitia Clench, [1964], type species: Thecla coelebs Herrich-Schäffer, 1862  
Enos K. Johnson, Kruse & Kroenlein, 1997, type species: Thecla thara Hewitson, 1867 
         Falerinota K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Thecla falerina Hewitson, 1867 
         Chopinia D'Abrera, 2001, ICZN Opinion 2358 (2015), type species: Thecla mazurka Hewitson, 1867 
         Ricuchallo Piñas, 2006, not available, nom. nud.  
Janthecla Robbins & Venables, 1991, type species: Thecla janthina Hewitson, 1867  
Laothus K. Johnson, Kruse & Kroenlein, 1997, type species: Thecla barajo Reakirt, [1867] 
         Gibbonota Salazar & López, 1996, not available, preocc. (Heinrich, 1937), type species: Thecla gibberosa 
Hewitson, 1867 
         Gibbossa Salazar & López, 2001, repl. name for Gibbonota, (ICZN art. 67.8), type species: Thecla gibberosa 
Hewitson, 1867 
         Runalatus Piñas, 2006, not available, nom. nud.  
 
Paiwarriina Lamas & Robbins, new subtribe 
Type Genus Paiwarria Kaye, 1904 
 
Paiwarria Kaye, 1904, type species: Papilio venulius Cramer, 1779  
Annamaria D'Abrera & Bálint, 2001, ICZN Opinion 2358 (2015), type species: Thecla draudti Lathy, 1926 
         Eucharia Boisduval, 1870, not available, subsequent designation, Kirby, 1871; preocc. (Hübner, [1820]), type 
species: Papilio ganimedes Cramer, 1775 
         Lamasina Robbins, 2002, repl. name for Eucharia, (ICZN art. 67.8), type species: Papilio ganimedes Cramer, 
1775 
         Airamanna Bálint, 2009, type species: Annamaria columbia Bálint, 2005  
Fasslantonius Bálint & Salazar, 2003, type species: Thecla episcopalis Fassl, 1912  
Kolana Robbins, 2004, type species: Thecla ligurina Hewitson, 1874 
 
Cupatheclina Lamas & Grishin, new subtribe 
Type Genus Cupathecla Bálint, 2005 
 
Cupathecla Bálint, 2005, type species: Papilio cupentus Stoll, 1781 
Dabreras Bálint, 2008, type species: Thecla teucria Hewitson, 1868 
 
Parrhasiina Busby & Robbins, new subtribe 
Type Genus Parrhasius Hübner, [1819] 
 
Parrhasius Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio polibetes Stoll, 1781 
         Eupsyche Scudder, 1876, type species: Thecla m-album Boisduval & Le Conte, 1833 
         Sumapanda Piñas, 2006, not available, nom. nud.  
Ignata K. Johnson, 1992, type species: Ignata ignobilis K. Johnson, 1992  
Michaelus Nicolay, 1979, type species: Thecla vibidia Hewitson, 1869  
Thepytus Robbins, 2004, type species: Thecla epytus Godman & Salvin, 1887  
Olynthus Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio narbal Stoll, 1790  
Beatheclus Bálint & Dahners, 2006, type species: Beatheclus beatrizae Bálint & Dahners, 2006  
Oenomaus Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio ortygnus Cramer, 1779 
         Draudtiana Kesselring & Ebert, [1982], not available, nom. nud.  
         Porthecla Robbins, 2004, type species: Thecla porthura H.H. Druce, 1907  
Apuecla Robbins, 2004, type species: Thecla upupa H.H. Druce, 1907  
Dicya K. Johnson, 1991, type species: Thecla dicaea Hewitson, 1874  
Caerofethra K. Johnson, 1991, type species: Thecla emendatus H.H. Druce, 1907  
Symbiopsis Nicolay, 1971, type species: Thecla strenua Hewitson, 1877 
 
Ipideclina Martins & Grishin, new subtribe 
Type Genus Ipidecla Dyar, 1916 
 
Ipidecla Dyar, 1916, type species: Ipidecla miadora Dyar, 1916 
 
Calycopidina Duarte & Robbins, 2010 
Type Genus Calycopis Scudder, 1876 
= Calycopina K. Johnson & Kroenlein, 1993 [type genus Calycopis Scudder, 1876] 
= Angulopina K. Johnson & Kroenlein, 1993 [type genus Angulopis K. Johnson, 1991] 
Unavailable under ICZN Articles 13.1 and 13.2 because Calycopina and Angulopina are family-group names 
proposed after 1930 without “a description or definition that states in words characters that are purported to differentiate 
the taxon.” 
 
Calycopis Scudder, 1876, type species: Rusticus poeas Hübner, [1811] 
         Calystryma Field, 1967, type species: Calystryma blora Field, 1967 
         Femniterga K. Johnson, 1988, type species: Femniterga notacastanea K. Johnson, 1988 
         Tergissima K. Johnson, 1988, type species: Tergissima mosconiensis K. Johnson, 1988 
         Morphissima K. Johnson, 1991, type species: Morphissima scalpera K. Johnson, 1991 
         Antrissima K. Johnson, 1991, type species: Antrissima varicolor K. Johnson, 1991 
         Fieldia K. Johnson, 1991, not available, preocc. (Walcott, 1912), type species: Fieldia yungas K. Johnson, 
1991 
         Cyanodivida K. Johnson, 1991, type species: Cyanodivida fornoi K. Johnson, 1991 
         Gigantofalca K. Johnson, 1991, type species: Gigantofalca stacya K. Johnson, 1991 
         Kroenleina K. Johnson, 1991, type species: Kroenleina panornata K. Johnson, 1991 
         Serratoterga K. Johnson, 1991, type species: Serratoterga larsoni K. Johnson, 1991 
         Klaufera K. Johnson, 1991, type species: Thecla pisis Godman & Salvin, 1887 
         Distissima K. Johnson, 1991, type species: Distissima spenceri K. Johnson, 1991 
         Terminospinissima K. Johnson, 1991, type species: Terminospinissima serratissima K. Johnson, 1991 
         Furcovalva K. Johnson, 1991, type species: Furcovalva extensa K. Johnson, 1991 
         Reversustus K. Johnson, 1991, type species: Thecla puppius Godman & Salvin, 1887 
         Profieldia K. Johnson, 1992, repl. name for Fieldia, (ICZN art. 67.8), type species: Fieldia yungas K. Johnson, 
1991  
Serratofalca K. Johnson, 1991, type species: Thecla cerata Hewitson, 1877  
Pendantus K. Johnson & Kroenlein, 1993, type species: Thecla plusios Godman & Salvin, 1887  
Camissecla Robbins & Duarte, 2004, type species: Thecla camissa Hewitson, 1870  
Gigantorubra K. Johnson, 1993, type species: Thecla collucia Hewitson, 1877 
         Arumecla Robbins & Duarte, 2004, type species: Thecla aruma Hewitson, 1877  
Electrostrymon Clench, 1961, type species: Papilio endymion Fabricius, 1781 
         Angulopis K. Johnson, 1991, type species: Thecla autoclea Hewitson, 1877  
Rubroserrata K. Johnson & Kroenlein, 1993, type species: Thecla mathewi Hewitson, 1874  
Ziegleria K. Johnson, 1993, type species: Ziegleria bernardi K. Johnson, 1993  
Arzecla Duarte & Robbins, 2010, type species: Thecla arza Hewitson, 1874  
Badecla Duarte & Robbins, 2010, type species: Thecla badaca Hewitson, 1868  
Kisutam K. Johnson & Kroenlein, 1993, type species: Thecla syllis Godman & Salvin, 1887  
Lamprospilus Geyer, 1832, type species: Lamprospilus genius Geyer, 1832  
Argentostriatus K. Johnson, 1991, type species: Thecla tamos Godman & Salvin, 1887  
Mercedes K. Johnson, 1991, type species: Thecla demonassa Hewitson, 1868 
 
Strymonina Tutt, 1907 
Type Genus Strymon Hübner, 1818 
Strymonini Tutt was placed on the Official List of Family-Group Names in Opinion 541 (ICZN, 1959). 
= Thereusina K. Johnson & Kroenlein, 1993 [type genus Thereus Hübner, [1819]].  
Unavailable under ICZN Articles 13.1 and 13.2 because Thereusina is a family-group name proposed after 1930 
without “a description or definition that states in words characters that are purported to differentiate the taxon.” 
 
Thereus Hübner, [1819], type species: Papilio lausus Cramer, 1779 
         Molus Hübner, [1819], subsequent designation, Scudder, 1875 (Robbins 2004, first reviser, ICZN art. 24.2), 
type species: Papilio phalanthus Stoll, 1780 
         Noreena K. Johnson, MacPherson & Ingraham, 1986, type species: Noreena maria K. Johnson, MacPherson 
& Ingraham, 1986 
         Solanorum K. Johnson, 1992, type species: Solanorum gentilii K. Johnson, 1992 
         Timokla K. Johnson, Kruse & Kroenlein, 1997, type species: Papilio erix Cramer, 1775 
         Pedusa D'Abrera, 2001, ICZN Opinion 2358 (2015), type species: Thecla pedusa Hewitson, 1867  
Rekoa Kaye, 1904, type species: Papilio meton Cramer, 1779  
Heterosmaitia Clench, [1964], type species: Thecla bourkei Kaye, 1925  
Arawacus Kaye, 1904, type species: Papilio aetolus Sulzer, 1776 
         Polyniphes Kaye, 1904, (Robbins 1991, first reviser, ICZN art. 24.2), type species: Polyommatus dumenilii 
Godart, [1824] 
         Dolymorpha Holland, 1931, type species: Thecla jada Hewitson, 1867 
         Tigrinota K. Johnson, 1992, type species: Thecla ellida Hewitson, 1867  
Strymon Hübner, 1818, subsequent designation, Riley, 1922, type species: Strymon melinus Hübner, 1818 
         Callipareus Scudder, 1872, type species: Strymon melinus Hübner, 1818 
         Calliparus Hemming, 1967, not available, missp.  
         Callicista Grote, 1873, type species: Callicista ocellifera Grote, 1873 
         Uranotes Scudder, 1876, repl. name for Callipareus, (ICZN art. 67.8), type species: Strymon melinus Hübner, 
1818 
         Eiseliana Ajmat, 1978, type species: Eiseliana koehleri Ajmat, 1978 
         Heoda K. Johnson, Miller & Herrera, 1992, type species: Thecla heodes H.H. Druce, 1909  
Hypostrymon Clench, 1961, type species: Thecla critola Hewitson, 1874  
Panthiades Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio pelion Cramer, 1775 
         Cycnus Hübner, [1819], subsequent designation, Scudder, 1875 (Robbins 2004, first reviser, ICZN art. 24.2), 
type species: Papilio phaleros Linnaeus, 1767 
         Laotisama Piñas, 2006, not available, nom. nud.  
 
Strephonotina K. Johnson, Austin, Le Crom & Salazar, 1997 
type genus Strephonota K. Johnson, Austin, Le Crom & Salazar, 1997 
Emended under ICZN Articles 19.2, 29, and 32.5.3 from Strephonina K. Johnson, Austin, Le Crom & Salazar, 1997 
= Tmolusina Bálint, 2014 [type genus Tmolus Hübner, [1819]] new synonym 
 
Theclopsis Godman & Salvin, 1887, type species: Thecla lebena Hewitson, 1868 
         Asymbiopsis K. Johnson & Le Crom, 1997, type species: Asymbiopsis designarus K. Johnson & Le Crom, 
1997  
Ministrymon Clench, 1961, type species: Thecla leda W.H. Edwards, 1882  
Iaspis Kaye, 1904, subsequent designation, ICZN, 1967, type species: Thecla temesa Hewitson, 1868  
Crimsinota K. Johnson, 1993, type species: Thecla socia Hewitson, 1868  
Aubergina K. Johnson, 1991, type species: Thecla alda Hewitson, 1868  
Tmolus Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio echion Linnaeus, 1767 
         Cyclotrichia K. Johnson, Austin, Le Crom & Salazar, 1997, type species: Thecla wickhami Riley, 1919 
         Jahuaima Piñas, 2006, not available, nom. nud.  
Celmia K. Johnson, 1991, type species: Papilio celmus Cramer, 1775 
         Uzzia K. Johnson, 1991, (Robbins 2004, first reviser, ICZN art. 24.2), type species: Thecla uzza Hewitson, 
1873  
Rindgea K. Johnson, 1993, senior homonym of Rindgea Ferguson 2008 in Geometridae, type species: Rindgea 
umuarama K. Johnson, 1993  
Nicolaea K. Johnson, 1993, type species: Thecla cauter H.H. Druce, 1907 
         Nicolea D'Abrera, 2001, not available, missp.  
Terenthina Robbins, 2004, type species: Thecla terentia Hewitson, 1868  
Siderus Kaye, 1904, type species: Siderus parvinotus Kaye, 1904 
         Bithys Hübner, 1818, not available, subsequent designation, Riley, 1922; suppr. (ICZN, Opinion 541), type 
species: Bithys leucophaeus Hübner, 1818 
         Bythis Hübner, [1831], not available, missp.  
Gossenia Bálint, [2019], type species: Papilio lycabas, 1777 Cramer  
Gargina Robbins, 2004, type species: Thecla gargophia Hewitson, 1877  
Decussata K. Johnson, Austin, Le Crom & Salazar, 1997, type species: Decussata colombiana K. Johnson, Austin, 
Le Crom & Salazar, 1997  
Ostrinotes K. Johnson, Austin, Le Crom & Salazar, 1997, type species: Thecla ostrinus H.H. Druce, 1907  
Strephonota K. Johnson, Austin, Le Crom & Salazar, 1997, type species: Thecla strephon occidentalis Lathy, 1926 
         Strephonota Salazar, 1995, not available, nom. nud.  
         Zigirina K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Thecla zigira Hewitson, 1869 
         Syedranota K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Thecla syedra Hewitson, 1867 
         Treboniana K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Thecla trebonia Hewitson, 1870 
         Serratonotes K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Thecla porphyritis H.H. Druce, 1907 
         Letizia K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Thecla phoster H.H. Druce, 1907 
         Robustana K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Robustina prima K. Johnson, Austin, Le Crom & Salazar, 1997 
         Diminutina K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), type 
species: Thecla tyriam H.H. Druce, 1907 
         Dindyminotes K. Johnson, Austin, Le Crom & Salazar, 1997, (Robbins 2004, first reviser, ICZN art. 24.2), 
type species: Papilio dindymus Cramer, 1775 
         Exorbaetta K. Johnson, Austin, Le Crom & Salazar, 1997, type species: Thecla metanira Hewitson, 1867 
         Meridaria Bálint, 2014, type species: Meridaria hightoni Bálint, 2014 
 
Trichonidina Duarte & Faynel, new subtribe 
Type Genus Trichonis Hewitson, 1865 
 
Bistonina Robbins, 2004, type species: Thecla biston Möschler, 1877  
Trichonis Hewitson, 1865, type species: Papilio theanus Cramer, 1777  
Manticia Bálint, 2019, type species: Thecla mantica H.H. Druce, 1907  
Megathecla Robbins, 2002, repl. name for Gullivera, (ICZN art. 67.8), type species: Thecla gigantea Hewitson, 
1867 
         Gulliveria D'Abrera & Bálint, 2001, not available, preocc. (Castelnau, 1878)., type species: Thecla gigantea 
Hewitson, 1867 
         Gullivera D'Abrera, 2001, not available, incorrect original spelling, type species: Thecla gigantea Hewitson, 
1867 
         Gullicaena Bálint, 2002, repl. name for Gulliveria. (ICZN art. 67.8), type species: Thecla gigantea Hewitson, 
1867 
 
Callophryidina Tutt, 1907 
Type Genus Callophrys Billberg, 1820 
Emended under Articles 19.2, 29, and 32.5.3 from Callophryidi Tutt, 1907 
= Neolycaenina Korb, 1997 [Type Genus Neolycaena de Nicéville, 1890], new synonym 
= Satyriumina Bálint & Costa, 2012 [Type Genus Satyrium Scudder, 1876]  
Unavailable under ICZN Article 16.1 because Satyriumina was introduced after 1999 without being “explicitly 
indicated as intentionally new.” 
 
Callophrys Billberg, 1820, subsequent designation, Scudder, 1875, type species: Papilio rubi Linnaeus, 1758  
         Lycus Hübner, [1819], not available, preocc. (Fabricius, 1787), type species: Papilio rubi Linnaeus, 1758 
         Licus Hübner, 1823, not available, missp. 
         Mitoura Scudder, 1872, type species: Thecla smilacis Boisduval & Le Conte, [1835] 
         Mitouri Scudder, 1872, not available, incorrect original spelling, type species: Thecla smilacis Boisduval & Le 
Conte, [1835] 
         Mitura Butler, 1875, not available, missp. 
         Incisalia Scudder, 1872, type species: Lycus niphon Hübner, [1819] 
         Satsuma Murray, 1874, not available, preocc. (Adams, 1868), type species: Lycaena ferrea Butler, 1866 
         Ahlbergia Bryk, 1946, repl. name for Satsuma. (ICZN art. 67.8), type species: Lycaena ferrea Butler, 1866 
         Ginzia Okano, 1947, repl. name for Satsuma. (ICZN art. 67.8); Hemming (1967) gave the year as 1941, type 
species: Lycaena ferrea Butler, 1866 
         Calophrys Barragué, 1954, not available, missp.  
         Sandia Clench & Ehrlich, 1960, type species: Callophrys (Sandia) mcfarlandi Ehrlich & Clench, 1960 
         Xamia Clench, 1961, type species: Thecla xami Reakirt, [1867] 
         Cisincisalia K. Johnson, 1992, type species: Cisincisalia moecki K. Johnson, 1992 
         Novosatsuma K. Johnson, 1992, type species: Novosatsuma monstrabila K. Johnson, 1992 
         Deciduphagus K. Johnson, 1992, type species: Thecla augustinus Westwood, 1852 
         Cissatsuma K. Johnson, 1992, type species: Satsuma albilinea Riley, 1939 
         Loranthomitoura Ballmer & Pratt, 1992, type species: Thecla spinetorum Hewitson, 1867  
Cyanophrys Clench, 1961, type species: Strymon agricolor Butler & H. Druce, 1872 
         Plesiocyanophrys K. Johnson, Eisele & MacPherson, 1993, type species: Thecla goodsoni Clench, 1946 
         Antephrys K. Johnson, Eisele & MacPherson, 1993, type species: Antephrys prestoni K. Johnson, Eisele & 
MacPherson, 1993 
         Mesocyanophrys D'Abrera, 1995, not available, nom. nud.  
         Paracyanophrys D'Abrera, 1995, not available, nom. nud.  
         Mesacyanophrys D'Abrera, 1995, not available, nom. nud.  
         Mesocyanophrys K. Johnson, 1997, type species: Thecla lycimna Hewitson, 1868 
         Apophrys K. Johnson & Le Crom, 1997, type species: Hesperia herodotus Fabricius, 1793  
Chalybs Hübner, [1819], subsequent designation, Scudder, 1875, type species: Papilio janias Cramer, 1779 
         Chalyps Hübner, [1826], not available, missp.  
Ocaria Clench, 1970, type species: Thecla ocrisia Hewitson, 1868 
         Arases K. Johnson, 1992, type species: Arases clenchi K. Johnson, 1992 
         Galba K. Johnson, 1992, not available, preocc. (Schrank 1803)., type species: Galba elvira K. Johnson, 1992 
         Lamasa K. Johnson, 1992, type species: Thecla calesia Hewitson, 1870 
         Variegatta K. Johnson, 1992, type species: Thecla elongata Hewitson, 1870 
         Ocacuni Piñas, 2006, not available, nom. nud.  
         Kurtaria Bálint, 2019, repl. name for Galba. (ICZN art. 67.8), type species: Galba elvira K. Johnson, 1992  
Magnastigma Nicolay, 1977, type species: Thecla tegula Hewitson, 1868 
Satyrium Scudder, 1876, type species: Lycaena fuliginosa W.H. Edwards, 1861 
         Chrysophanus Hübner, 1818, not available, subsequent designation, Riley, 1922; suppr. (ICZN, Opinion 541), 
type species: Chrysophanus mopsus Hübner, 1818 
         Chrysophanes Weidemeyer, 1864, not available, missp.  
         Argus Gerhard, 1850, not available, preocc. (Bohadsch, 1761), type species: Lycaena ledereri Boisduval, 1848 
         Callipsyche Scudder, 1876, (Ziegler 1960, first reviser, ICZN art. 24.2), type species: Thecla behrii W.H. 
Edwards, 1870 
         Neolycaena Nicéville, 1890, type species: Lycaena sinensis Alphéraky, 1881 
         Nordmannia Tutt, 1907, type species: Lycaena myrtale Klug, 1834 
         Edwardsia Tutt, 1907, not available, preocc. (Costa, 1838), type species: Papilio w-album Knoch, 1782 
         Klugia Tutt, 1907, not available, preocc. (Robineau-Desvoidy, 1863), type species: Papilio spini Fabricius, 
1775 
         Kollaria Tutt, 1907, not available, preocc. (Pictet, 1841), type species: Thecla sassanides Kollar, 1849 
         Felderia Tutt, 1907, not available, preocc. (Walsingham, 1887), type species: Thecla w-album var. eximia 
Fixsen, 1887 
         Erschoffia Tutt, 1907, not available, preocc. (Swinhoe, 1900), type species: Thecla lunulata Erschoff, 1874 
         Fixsenia Tutt, 1907, type species: Thecla herzi Fixsen, 1887 
         Leechia Tutt, 1907, not available, preocc. (South, 1901), type species: Thecla thalia Leech, 1893 
         Bakeria Tutt, 1907, not available, preocc. (Kieffer, 1905), type species: Lycaena ledereri Boisduval, 1848 
         Chattendenia Tutt, 1908, repl. name for Edwardsia. (ICZN art. 67.8), type species: Papilio w-album Knoch, 
1782 
         Strymonidia Tutt, 1908, repl. name for Leechia (ICZN art. 67.8), precedence over Chattendenia, type species: 
Thecla thalia Leech, 1893 
         Strymonidea Dunk, 1952, not available, missp. 
         Pseudothecla Strand, 1910, repl. name for Erschoffia. (ICZN art. 67.8), type species: Thecla lunulata Erschoff, 
1874 
         Superflua Strand, 1910, repl. name for Kollaria. (ICZN art. 67.8), type species: Thecla sassanides Kollar, 1849 
         Thecliolia Strand, 1910, repl. name for Felderia. (ICZN art. 67.8), type species: Thecla w-album var. eximia 
Fixsen, 1887 
         Thecliola Waterhouse, 1912, not available, missp. 
         Tuttiolia Strand, 1910, repl. name for Klugia. (ICZN art. 67.8), type species: Papilio spini Fabricius, 1775 
         Tuttiola Hemming, 1967, not available, missp. 
         Necovatia Verity, 1951, type species: Papilio acaciae Fabricius, 1787 
         Euristrymon Clench, 1961, type species: Papilio favonius J.E. Smith, 1797 
         Phaeostrymon Clench, 1961, type species: Thecla alcestis W.H. Edwards, 1871 
         Harkenclenus dos Passos, 1970, repl. name for Chrysophanus. (ICZN art. 67.8), type species: Chrysophanus 
mopsus Hübner, 1818 
         Rhymnaria Zhdanko, 1983, type species: Lycaena rhymnus Eversmann, [1832] 
         Armenia Dubatolov & Korshunov, 1984, repl. name for Argus and Bakeria (ICZN art. 67.8), type species: 
Lycaena ledereri Boisduval, 1848 
         Neosatyrium Fisher, 2009, type species: Rusticus calanus Hübner, [1809]  
Thaeides K. Johnson, Kruse & Kroenlein, 1997, type species: Thecla theia Hewitson, 1870  
Radissima K. Johnson, 1992, type species: Sithon umbratus Geyer, 1837  
Erora Scudder, 1872, type species: Thecla laeta W.H. Edwards, 1862 
         Androcona K. Johnson, Eisele & MacPherson, 1993, type species: Thecla muridosca Dyar, 1918 
         Sarracenota K. Johnson, Eisele & MacPherson, 1993, type species: Thecla opisena H.H. Druce, 1912 
         Necmitoura K. Johnson, Eisele & MacPherson, 1993, type species: Necmitoura marcusa K. Johnson, Eisele & 
MacPherson, 1993  
Semonina Robbins, 2004, type species: Thecla semones Godman & Salvin, 1887  
Chlorostrymon Clench, 1961, type species: Thecla telea Hewitson, 1868