Abstract
A comprehensive tribal-level classification for the worldÕs subfamilies of Hesperiidae, the skipper butterflies, is proposed for the first time. Phylogenetic relationships between tribes and subfamilies are inferred using DNA sequence data from three gene regions (cytochrome oxidase subunit I-subunit II, elongation factor-1a and wingless

William Harry Evans' (1937 Evans' ( , 1949 Evans' ( , 1951 Evans' ( , 1952 Evans' ( , 1953 Evans' ( , 1955 monumental series of monographs represents the most recent revision of the worldÕs fauna of Hesperiidae, although he proposed somewhat independent classification schemes for each of the worldÕs regions. Evans arranged phenotypically similar genera into informal groups to aid in their identification, but rarely hypothesized relationships among groups in the same subfamily distributed in different parts of the world. Although there have been some modifications to EvansÕ classification (see Appendix 1), his taxonomic system remains largely intact in current treatments of the group. As a result, Hesperiidae is the only family of butterflies without a widely accepted tribal-level classification for all of the major subfamilies Lamas, 2004; . As noted by Voss (1952) , the skippersÕ ''remarkable uniformity of structure leaves us with so little upon which to base sound distinctions that we often are forced to consider significant any trivial character that appears to be a fairly consistent criterion to characterize a group''. Indeed, few morphological synapomorphies have been identified that can readily characterize any subfamily of skippers , and there is no general consensus on the composition of or relationships among the various subfamilies .
Considering the recent progress in understanding the higher-level systematics of other groups of Lepidoptera (e.g. Weller et al., 1994; Brower, 2000; Regier et al., 2000 Regier et al., , 2002 Wiegmann et al., 2000; Caterino et al., 2001; Bucheli and Wenzel, 2005; Wahlberg et al., 2005a,b; Braby et al., 2006; Brower et al., 2006; Pen˜a et al., 2006) , it is clear that molecular characters can be useful in delineating higher-level taxa and determining relationships. We agree with , who noted for the Hesperiidae, ''A molecular study to assist in the redefinition of subfamilies, tribes, and genera -and not least the relationships within the family worldwidewould be a worthwhile exercise''.
In the present study we test the monophyly of the most recent circumscriptions of the subfamilies of the Hesperiidae, as well as EvansÕ generic groupings within each subfamily. We endeavour to delineate tribes within the major subfamilies, and determine relationships among tribes and subfamilies of the Hesperiidae. We also aim to gain preliminary insights into certain historically controversial genus-level relationships. Hesperiidae is currently divided into seven subfamilies, namely Coeliadinae, Pyrrhopyginae, Pyrginae, Heteropterinae, Trapezitinae, Hesperiinae and Megathyminae (see Table 1 ), which include a total of 567 genera (see Appendix 1). Some recent authors (e.g. Atkins, 2005) recognize an eighth subfamily, Euschemoninae, while other authors treat the Heteropterinae (e.g. Bridges, 1993; Pyle, 2002) or Megathyminae (e.g. Ackery et al., 1999; Opler and Warren, 2002) as subordinate taxa of the Hesperiinae. Evans divided the Pyrrhopyginae, Pyrginae and Hesperiinae into a total of 28 generic groups, a few of which have since been modified and given formal recognition at the tribal or subfamily level (e.g. Higgins, 1976; ). Evans further divided nine of his generic groups into 38 subgroups, for a total of 58 suprageneric taxa. We have sampled one or more members from all but three of EvansÕ groups and subgroups, and two or more members from all but ten of these (excluding monotypic subgroups), allowing us to make a preliminary assessment of the monophyly of most of EvansÕ suprageneric hypotheses (see Appendix 1), and to evaluate the naturalness of these groups as a basis for a phylogenetic tribal classification. Our hypothesis of relationships is based upon DNA sequences from three gene regions: a contiguous region of mitochondrial cytochrome oxidase subunits I and II (COI-COII), and nuclear elongation factor-1a (EF-1a) and wingless.
Materials and methods

Taxon sampling
Adult butterflies were sampled with aerial nets in the field, by the authors and various colleagues. Specimens were preserved in 85-100% ethanol, with wings removed prior to submersion, or were preserved dry, in glassine envelopes. The species sampled and their collection localities are listed in Appendix 2. A total of 209 species in 198 genera are included in the combined analysis of three genes, discussed below, representing about 35% of the worldÕs skipper genera (sensu Ackery et al., 1999; see Table 1 , Appendix 1). Partial (two gene segments) or complete data were obtained for 22 additional genera and species (marked with an asterisk in Appendix 1), which were not included in the final combined analysis but were included in alternative analyses and are discussed below. Sequences for all taxa are new, except for outgroups and five skipper species, which were published in Wahlberg et al. (2005a) . Five outgroup species (see Appendix 2) were selected, one from each family of the Papilionoidea, the putative sister clade to the Hesperioidea (Wahlberg et al., 2005a) . Sequences for the outgroup species were obtained from GenBank.
Laboratory protocols
Total genomic DNA was extracted from individual butterflies, by using a standard phenol-chloroform extraction protocol (Brower, 1994 (Brower, , 2000 or QiagenÕs DNEasy extraction kits (Qiagen, Venlo, the Netherlands) according to the manufacturerÕs instructions. We extracted DNA from the thorax of specimens preserved in ethanol, or from two legs of dried butterflies. Vouchers consist of vials of DNA suspended in Table 1 Traditional and revised family-level classifications of Hesperiidae. Left column represents the classification proposed by Evans and subsequent authors, as detailed in Appendix 1. Right column represents the classification proposed in this paper. Dashes prior to names indicate the following taxonomic status: 1 ¼ family-level name; 2 =subfamily-level names; 3 ¼ tribal names; 4 ¼ subtribal names; 5 ¼ subjective junior synonyms; 6 ¼ unavailable names HPLC-grade water (final elution volume between 50 and 500 lL, depending on amount of starting tissues), frozen at )20°C, and corresponding wings and body parts (usually minus the thorax) stored in glassine envelopes. DNA and residual morphological materials will be permanently deposited in public institutions, as indicated in Appendix 2.
For each specimen, we amplified and sequenced a 943-bp fragment spanning the 3¢ end of COI, the tRNAleu and the 5¢ end of COII, 739 bp of EF-1a and 403 bp of the wingless gene (although in a few cases sequences for different genes were obtained from two specimens, as indicated in Appendix 2). Skipper-specific primers for COI-COII were developed (Gary and Susan, see Table 2 ), after obtaining preliminary sequences from primers listed in Brower and Jeansonne (2004) and Brower et al. (2006) . Primers for EF-1a were taken from Cho et al. (1995) and Monteiro and Pierce (2001) , and for wingless from Brower and DeSalle (1998) ; all primers used in this study are listed in Table 2 . PCR amplifications were performed in a 50-or 100-ll reaction volume, on a Peltier thermal cycler (PTC-100, MJ Research, c ⁄ o Biorad, Hercules, CA, USA). Amplifications conducted in a 50-lL reaction volume included 3 lL of template, 5 lL of 10 · buffer (0.1 m Tris-HCl, 0.1 m KCl, 1% Triton X-100, pH 8.3), 5 lL of 25 lm MgCl 2 , 1 lL of 10 lm dNTPs, 2 lL of each primer (10 lm), 0.3 lL Taq polymerase, and 31.7 lL distilled water. Amplifications conducted in a 100-lL reaction volume included 1 lL of template, 10 lL of 10 · buffer, 15 lL of 25 lm MgCl 2 , 2 lL of 10 lm dNTPs, 2 lL of each primer (10 lm), 0.2 lL Taq polymerase, and 69 lL distilled water. The cycling profile for COI-COII and wingless was 4 min at 92°C, and 40 cycles of 1 min at 94°C, 0.5 or 1 min at ------l Erynninae Swinhoe, 1913 *This synonymy is based on the arrangement detailed in Appendix 1. l = unavailable name. There are at least four family group names formed from the genus Erynnis, but only one of these is based on the genus properly identified. As dictated by Code articles 41 and 65.2.1 (ICZN, 1999) , the case should be referred to the Commission for a ruling. In the meantime, we treat these names as if the Commission has ruled to suppress all but the one properly proposed name (Erynninae Brues and Carpenter, 1932) . (Rijsewijk et al., 1987) ; of the COI-COII primers in the Drosophila yakuba mitochondrial genome sequence (Clary and Wolstenholme, 1985) , and the Ef-1a primers in the Drosophila melanogaster sequence as reported by Cho et al. (1995) . 46°C, and 2 min at 72°C, and that for EF-1a was 2 min at 94°C, and 32 cycles of 1 min at 94°C, 1 min at 60°C, and 1.5 min at 72°C, followed by 10 min at 72°C.
Amplified DNA fragments were cleaned with silica beads (Bio 101, Qbiogene, Irvine, CA, USA), or with Qiaquick PCR purification kits (Qiagen). Cleaned PCR products were cycle sequenced using ABI Prism or Big Dye kits (Applied Biosystems, Foster City, CA, USA), in a PTC-100, with the same primers as used for PCR. Recommended reaction conditions were used, along with the profile of 60 cycles of 0.5 min at 96°C, 0.25 min at 50°C, and 4 min at 60°C. Single-stranded products were cleaned using ethanol and sodium acetate precipitation, and run on an ABI 373A or 377 automated sequencer or outsourced to Macrogen (Seoul, South Korea). All sequences were generated in both directions. Automated sequence outputs were edited manually and aligned by eye. Other than some minor length heterogeneity at the beginning and end of the tRNA and a single one-codon deletion in wingless (present in two taxa), there was no ambiguity in the alignment. Heterozygous positions in the nuclear genes (where simultaneous chromatogram peaks for two nucleotides appeared almost or exactly equal) were coded according to the IUPAC ambiguity codes. The aligned data matrix is available on the web at http:// www.treebase.org. Individual sequences have been submitted to GenBank (accession codes given in Appendix 2).
Phylogenetic analysis
Data were concatenated and analysed as a single matrix under the parsimony criterion. Gaps were scored as missing; all characters and transformations were weighted equally. We searched for the most parsimonious cladograms from the unordered and equally weighted data matrix consisting of 215 taxa. Trees were rooted with Papilio, and other non-hesperiid taxa were included in the ingroup to test the monophyly of Hesperiidae. The parsimony analyses were performed in PAUP* 4.0b 10 (Swofford, 2002) using the parsimony ratchet (Nixon, 1999) as implemented in PAUP* by PAUPRat (Sikes and Lewis, 2001) . The general ratchet analysis conditions were as follows: seed = 0, nreps = 200, wtmode = uniform. The percentage of characters perturbed during each iteration (pct) varied between 5, 10 and 15%. The search was repeated five times for each level of character perturbation, yielding a total of 15 independent ratchet searches. The maximumparsiomiony (MP) tree length was corroborated in NONA 2.0 (Goloboff, 1999) using similar parameters as the PAUP* tree searches. In addition, we explored the structure of the data with separate analyses of each gene region, using heuristic searches with 1000 random addition replicates using tree bisection-reconnection (TBR) branch swapping with a single tree held during each step.
In the combined analysis, we evaluated character support and congruence among partitions for the clades in the strict consensus of the MP trees using branch support (BS : Bremer, 1988 : Bremer, , 1994 , partitioned branch support (PBS: Baker and DeSalle, 1997; Gatesy et al., 1999) and the partition congruence index (PCI: Brower, 2006b ; see also Brower et al., 2006) . Fractional PBS values were rounded to two decimal places. Due to the computationally intensive structure of the data set, BS values were calculated in PAUP* using PAUPRatgenerated batch files that were modified to search anticonstraint trees generated from the MP tree set using TreeRot v.2. (Sorenson, 1999) . Although tedious to set up by hand, the use of the parsimony ratchet to search for anti-constraint tree lengths consistently found shorter trees (resulting in lower BS values) than searches using standard PAUP* heuristic strategies. As in other recent studies (e.g. Wahlberg et al., , 2005b , we refer to the support values as giving weak, moderate, good or strong support when discussing our results. We define Ôweak supportÕ as BS values between 1 and 2, Ômoderate supportÕ as BS values between 3 and 5, Ôgood supportÕ as values between 6 and 10, and Ôstrong supportÕ as values of 11 and greater. We endorse BS values over bootstrap values as they are a parameter of the data, rather than an estimate of tree stability based on pseudoreplicated subsamples of the data, and because they have no upper bound (Brower, 2006b) .
Results and discussion

Characteristics of the data set
The total combined data consist of 2086 bp, 913 of which are invariant and 890 of which are parsimonyinformative. Combining the three data sets in simultaneous parsimony analysis yields 90 trees of 19,123 steps (CI = 0.091, RI = 0.422), the strict consensus of which is shown in Figs 1 and 2. Up to 35 positions were coded as gaps in some taxa, including one gap in the wingless data set and three gaps in the COI-COII data set; all of these were easily detected when aligning by eye, as flanking regions were conserved. A few sequences are incomplete and 11 taxa are missing wingless sequences (see Appendix 2). Basic statistics for the three gene regions are shown in Tables 3 and 4. In order to investigate incongruence (Mickevich and Farris, 1981; Farris et al., 1994) , we conducted separate analyses of the three gene regions. Overall, the phylogenetic signal of wingless strongly conflicts with the other two gene regions ( The classification of Hesperiidae discussed below is based on the results of the combined cladistic analysis. The nomenclatorial philosophy we employ is that all named taxa should be monophyletic, and that taxa meeting this criterion should bear names and ranks associated with them in the historical literature to the greatest degree possible. BS values for individual clades are indicated below (also see Table 4 ). In the text below, numbers in parentheses after the names of taxa refer to the numbered clades in Figs 1 and 2.
Subfamily-level relationships
This is the first comprehensive phylogenetic analysis of relationships within the family Hesperiidae. Previous phylogenetic studies of the family have been limited by insufficient taxon sampling, either lacking sufficient taxa for adequate resolution (e.g. de Jong et al., 1996; , or including taxa sampled on a regional basis only (Chiba et al., 2001 ), and therefore lacking major sections of diversity present in the family. Other family-level studies of the Hesperiidae have not employed a cladistic methodology, and ⁄ or have scored and analysed characters in an ambiguous way (Voss, 1952; Scott, 1985; Scott and Wright, 1990; Atkins, 2005) . In our study, we have identified several clades that are strongly supported by three gene regions, as well as clades that are less robust and likely to change with the addition of more characters.
Our data imply that the family Hesperiidae (1), as currently circumscribed, is monophyletic with strong support (BS 13), in agreement with the results of Wahlberg et al. (2005a) . Six of seven currently recognized subfamilies of Hesperiidae are recovered as monophyletic clades (although not all represent subfamily-level taxa, see Figs 1 and 2), with the following relationships: (Coeliadinae + (''Pyrginae'' including Pyrrhopyginae + (Heteropterinae + (Trapezitinae + Hesperiinae including Megathyminae)))). Monophyly of Coeliadinae (2) receives strong support (BS 12), and its basal position sister to the rest of the Hesperiidae corroborates the results of de Jong et al. (1996) and Wahlberg et al. (2005a) . Although Pyrrhopyginae (45) (BS 5) . The two genera included in our analysis from Megathyminae (or ''giant skippers'') are sister taxa with strong support (129: BS 43), but this clade is placed deep within Hesperiinae, in a polytomy (110) with various Asian and African genera, also including the Neotropical genera Orses, Perichares (Carystus group) and Pyrrhopygopsis (Calpodes group). Additional taxa and characters will be needed to elucidate the phylogenetic position of the giant skippers.
In summary, our results imply that four subfamilies of Hesperiidae should be recognized: Coeliadinae, Heteropterinae, Trapezitinae and Hesperiinae. ''Pyrginae'' is a paraphyletic grade of seven clades, some of which should be recognized as tribal-level taxa. Further study is needed before a satisfactory classification of the ''Pyrginae'' will be possible, and additional characters and ⁄ or taxa are needed to elucidate the phylogenetic positions of Euschemon rafflesia and the giant skippers. 
Monophyly of EvansÕ generic groups and subgroups
As shown in Table 5 , only five of EvansÕ 28 generic groups within the various subfamilies of Hesperiidae were recovered as monophyletic clades, although we did not sample enough genera to assess the monophyly of the Isoteinon group or two groups of Pyrrhopyginae (Oxynetrini and the monotypic Zoniini). One of the monophyletic groups is Heteropterinae (89, equivalent to EvansÕ Carterocephalus group), which has since been widely regarded as a subfamily-level taxon (see . Two of EvansÕ monophyletic genus-groups are in Pyrrhopyginae (46, 50), and were subsequently modified and elevated to tribal-level taxa by . The other two monophyletic genus-groups are both within Hesperiinae: the Taractrocera group (132), which is strongly supported (BS 12; eight of 13 genera included), and the Gegenes group (151), which is also strongly supported (BS 30; three of 14 genera included) but is situated within a clade of hesperiines from several other groups (145, as discussed below). The remaining 23 generic groups defined by Evans are para-or polyphyletic, according to our results.
Only three of EvansÕ 38 generic subgroups were recovered as monophyletic clades (see Table 5 ), although we did not sample enough taxa to assess the monophyly of eight of these (excluding monotypic subgroups). The monophyletic groups are the Tagiades subgroup of the Tagiades group (54: BS 28; strongly supported but only two of ten genera included, which were once considered congeneric), the Paramimus subgroup of the Telemiades group (31: BS 55; very strongly supported, two of five genera included), and the Thymelicus subgroup of the Hesperia group (141: BS 4; moderate support, three of five genera included). The remaining subgroups are para-or polyphyletic. Based on these results, the use of EvansÕ generic groups and subgroups as a basis for a tribal classification cannot be considered satisfactory.
Paraphyly of Pyrginae
As noted above (Figs 1 and 2), our data suggest that Pyrginae of previous authors (e.g. Evans, 1937 Evans, , 1949 Evans, , 1952 Evans, , 1953 ) is a paraphyletic grade composed of several clades. This result is not surprising, as several recent workers have questioned the monophyly of the group. In the morphological analyses conducted by de Jong et al. (1996; see also Ackery et al., 1999) , Pyrginae was ''never'' recovered as a monophyletic group, although their study included just ten skipper taxa. went as far as to say that ''Pyrginae is certainly not monophyletic''. In the combined molecular (three genes) and total evidence (molecular plus morphological) analyses conducted by Wahlberg et al. (2005a) , only two species of pyrgines were included (Pyrgus and Urbanus), but these never formed a monophyletic group. Our data also failed to recover Pyrginae as a monophyletic group, although relationships implied ( Fig. 1 ) among the clades of ''Pyrginae' ' (8, 20, 33, 35, 42 : all BS 1) receive weak support with strong incongruence among data partitions, and the arrangement of these clades is not likely to be robust to the addition of more characters. However, two (9, 36) of the five ''major'' clades in this group are strongly supported, and appear to represent tribal-level taxa, while components of the remaining major clades (21, 43, 61) are also strongly supported and appear to represent tribal-level taxa, as discussed below. Until the clades of ''Pyrginae'' can be studied in more detail through the addition of more characters (e.g. morphology), and relationships between them can be better understood, we retain ''Pyrginae'' as an informal subfamily-level grouping composed of various tribes, but acknowledge its paraphyly by placing the name in quotation marks.
The sister taxon to other ''Pyrginae'' + Heteropterinae + Trapezetinae + Hesperiinae with weak support (8: BS 1) is the aptly named Clito aberrans. Throughout the course of this study, the position of C. aberrans has varied with the inclusion of additional taxa; various data sets have implied relationships with Quadrus + Pythonides, Milanion + Atarnes, and Eracon, among others. Evans (1953) placed Clito in his Antigonus subgroup of the Telemiades group, a placement retained by subsequent workers (e.g. Cock, 1998; , none of whom has questioned its genus-group placement or commented on unusual morphological features. Based on this, we consider the current basal position of Clito in Fig. 1 to be spurious, and do not believe it represents its actual relationship to other members of ''Pyrginae'' (trees just two steps longer place Clito between Quadrus + Pythonides and Milanion + Atarnes). Of the six Evans (1937 Evans ( , 1949 Celaenorrhinus group. This is the first study to challenge the subfamily-level status of Pyrrhopyginae ; but see Wahlberg et al., 2005a) ; however its phylogenetic position deeply nested within ''Pyrginae'' (clade 45) has been robust to the addition of taxa and characters over the course of this study (e.g. . When ''Pyrginae'' has appeared as a paraphyletic grade, as in the current study, Pyrrhopyginae never formed one of its ''major'' subdivisions. Given the topology of our cladogram, maintenance of the Pyrrhopyginae as a subfamily-level taxon would require recognition of at least seven additional subfamilies (clades 9, 22, 29, 36, 51, 55, 61) within what is currently circumscribed as ''Pyrginae''. In discussing the secondary sexual characters that partly serve to delineate groups within Pyrginae, Ackery et al. (1999) noted, ''At first sight there is no apparent reason why the Pyrrhopyginae could not be a subordinate taxon of Pyrginae''. Our results support that hypothesis.
In trees just two steps longer than the most parsimonious tree set, Pyrginae is recovered as a weakly supported monophyletic group, composed of two major clades. One of these clades includes members of clades 9, 21, 34, 36 and 43 (Fig. 1) , while the other major clade includes the same taxa as clade 61 (BS 5), including members of EvansÕ Augiades and Urbanus groups, also including a few species from the Celaenorrhinus and Telemiades groups (see below), as well as Euschemon rafflesia. The position of E. rafflesia at the base of this clade (62) is weakly supported (BS 1, with strong incongruence among partitions), and in trees just a few steps longer, E. rafflesia falls out of this clade into an unresolved polytomy including the rest of the Pyrginae. Thus, it would not be surprising if the phylogenetic position of E. rafflesia changes with the addition of further data. However, our results highlight the affinity of Euschemon with other Hesperiidae, and suggest that the species belongs in this family, despite its morphological peculiarities.
Circumscription of tribes
No tribal-level classification has been proposed for Coeliadinae (2) or Heteropterinae (89), both of which are ÔsmallÕ subfamilies with fewer than 15 genera. Morphology of the coeliadine genera is rather uniform , and it seems unlikely the subfamily will be further subdivided in the future. Morphology of Heteropterinae is also rather uniform , although its circumscription remains incomplete. Evans (1937) placed the genera Tsitana and Lepella in his African Astictopterus group (part of his Hesperiinae), together with Metisella and Hovala. Bridges (1993) retained all of these genera in the Astictopterus group, but included them all in Heteropterinae. When Tsitana is included in our combined analysis (data not shown), it groups with members of EvansÕ African Astictopterus and Ampittia groups (such as Astictopterus, Isoteinon and Kedestes, clade 121), well within Hesperiinae. We were unable to sample Hovala, but Evans (1937) believed it to be closely related to Metisella, and in our tree Metisella is sister to Carterocephalus (a genus undoubtedly related to Heteropterus), with strong support (91: BS 14). We were also unable to sample Lepella, and some other putative heteropterine genera, as indicated in Appendix 1. Therefore, we make no attempt to subdivide the Heteropterinae further, although further subdivision may be warranted with the addition of more taxa and characters. Voss (1952) divided Trapezitinae (95) into two tribes, ''Trapezitidi'' (explicitly including just Trapezites) and ''Hesperillidi''. He divided the latter into two unnamed groups based on the presence or absence of a stigma on the male forewing, and on the number of metatibial Table 5 Monophyly of EvansÕ subfamilies and generic groups (as modified by Ackery et al., 1999 and ), based on taxa sampled for this study. spurs. One group (with a stigma and two pairs of spurs) explicitly included Dispar, Hesperilla, Signeta, and Toxidia, while the other group (without a stigma and with one pair of spurs) explicitly included only Mesodina. However, Waterhouse (1932) and various subsequent authors have recognized three major groupings within the Trapezitinae (e.g. Atkins, 1973; Common and Waterhouse, 1981; Ackery et al., 1999 ) that do not directly overlap with VossÕ tribes, based on differences in larval foodplant families and characters of the larvae and pupae. These include (1) ÔtrapezitineÕ genera feeding primarily on Xanthorrhoeaceae and Poaceae (Trapezites, Anisynta, Pasma, Neohesperilla, Dispar, Toxidia, Signeta, and Croitana), (2) ÔhesperillineÕ genera feeding only on Cyperaceae (Oreisplanus, Hesperilla, and Motasingha), and (3) the ÔmesodineÕ genus feeding on Iridaceae (Mesodina). Larval foodplants of the New Guinean genera Hewitsoniella and Felicena remain unknown (Parsons, 1999) , and foodplants of the genus Rachelia have recently been found to be in the Flagellariaceae (Braby, 2004) . Recent research on relationships of trapezetine genera (e.g. Atkins, 1973 Atkins, , 1984 Atkins, , 1994 has not supported WaterhouseÕs groupings, and a separate informal grouping, the ÔProeidosa group,Õ has been proposed for Croitana and two recently described genera, Proeidosa and Antipodia (see Atkins, 1984 Atkins, , 1994 . Despite the informal groupings identified by various authors, no formal tribal-level classification for the subfamily Trapezitinae has been employed since VossÕ study (e.g. Bridges, 1993; Atkins and Edwards, 1996; Braby, 2000 Braby, , 2004 . Our results do not support the monophyly of VossÕ tribes, but do support the monophyly of WaterhouseÕs three ÔhesperillineÕ genera (100: BS 9). Our results also indicate that the ÔtrapezitineÕ genera are polyphyletic. When three other ÔtrapezitineÕ genera are added to our combined analysis (Anisynta, Neohesperilla and Pasma), for which data from only two genes is currently available, this arrangement does not change (data not shown). More genera are required to test the monophyly of the Proeidosa group, and its relationship to Mesodina. Until additional genera can be sampled and additional characters can be included (including those from immature stages), we feel it is premature to propose a tribal-level classification for the Trapezitinae. The Megathyminae (or ''giant skippers'', clade 129) have previously been divided into three tribes Turner, 1958, 1959) , an arrangement which has persisted among some authors (e.g. . However, many authors have treated the giant skippers as a family-level taxon within Hesperioidea (e.g. Comstock and Comstock, 1895; Barnes and McDunnough, 1912; Lindsey, 1921; Lindsey et al., 1931; McDunnough, 1938; Brown et al., 1956; dos Passos, 1964; Freeman, 1969b; Roever, 1975; Bridges, 1993) . Our results indicate that the giant skippers are apparently a highly derived group of hesperiines (see Table 1 ), corroborating the views of Scott and Wright (1990) and Ackery et al. (1999) . Furthermore, our results fail to support even tribal-level status for giant skippers, although such a status should not be ruled out until morphological characters are also considered.
As we have included only 35% of the worldÕs genera of Hesperiidae in our combined analysis, inclusion of all skipper genera into a tribal classification must await a comprehensive morphological study to put our results into a broader context (A. Warren, J.R. Ogawa and A.V.Z. Brower, unpublished data). However, we have been able to identify certain clades with good or strong support, which are likely to be robust to the addition of taxa and characters, and appear to represent tribal-level entities. Recent efforts to construct a tribal nomenclature for Pyrginae and Hesperiinae have been regional in nature and are largely based on EvansÕ regional generic groups (Chou, 1994 (Chou, , 1998 Koc¸ak and Seven, 1997) . Based on our results (Figs 1 and 2 ), we propose a cosmopolitan tribal classification for ''Pyrginae'' and Hesperiinae, using available family-level names, to complement our revised subfamily-level arrangement (see Table 1 ). As the tribes of ''Pyrginae'' are arranged in a weakly supported paraphyletic grade (Fig. 1) , the order in which they are discussed below does not imply any particular relationship among tribes, and mostly follows the order presented by Evans (1937 Evans ( , 1949 Evans ( , 1952 Evans ( , 1953 .
Eudamini, confirmed status (61). This clade includes members of EvansÕ Augiades and Urbanus groups, as well as some members of his Celaenorrhinus and Telemiades groups (see Appendix 1). Recently, has arranged genera in EvansÕ Augiades and Urbanus groups, and American representatives of the Celaenorrhinus group, under the tribe Eudamini. MielkeÕs Eudamini (62: BS 1) was recovered as a weakly supported monophyletic group, with the addition of Spathilepia, Cogia and Telemiades (from the Telemiades group), and the removal of Celaenorrhinus. The Asian genus Lobocla (75, from the Celaenorrhinus group) is also included in Eudamini. As discussed above, Euschemon rafflesia is situated at the base of this clade, in a sister relationship with MielkeÕs Eudamini. For now we include Euschemon within Eudamini, although it is stressed that this placement should be considered tentative, until morphological characters can also be evaluated together with our molecular data. For the most part, relationships within the Eudamini are poorly supported by our data, although the monophyly of the clade including Urbanus (which itself is paraphyletic, see below), Thorybes, Achalarus, and Autochton receives good support (85: BS 8), and the sister relationships between Phocides + Nascus (63: BS 18), Polygonus + Telemiades (69: BS 21), and Typhedanus + Codatractus (77: BS 27) are strongly supported. Eudamini was originally proposed by Mabille (1877) , and has been used at the tribal level by various authors (e.g. Mabille, 1878; Tutt, 1906 -in Tutt 1905 -1914 Clark, 1948; Mielke and Casagrande, 1998; Lamas, 2003; .
Pyrrhopygini, reinstated status (45). Evans ' (1951) generic groups for Pyrrhopyginae were modified and given tribal-level status by . We were unable to sample representatives of two of these tribes, Oxynetrini and Zoniini (which is monotypic), but the monophyly of the two tribes we were able to sample, Pyrrhopygini (46: BS 24) and Passovini (50: BS 34), is strongly supported by our data. Inclusion of three additional genera for which we currently have only partial data (Yanguna, Jemadia and Mimoniades; data not shown) does not change the circumscription of MielkeÕs tribes. However, due to the position of Pyrrhopyginae within ''Pyrginae'' (Fig. 1) , we treat the former subfamily as a tribe of ''Pyrginae''. This action changes the status of the tribes described by , which can now be known as sub-tribes: Pyrrhopygina (46, new status), Zoniina (new status), Passovina (50, new status) and Oxynetrina (new status). Pyrrhopygini was originally proposed by Mabille (1877) , and was emended to Pyrrhopyginae by Watson (1893) , a spelling employed by all subsequent authors who recognized the group as a subfamily-level taxon .
Tagiadini, confirmed status (51). Monophyly of Tagiadini receives weak support (BS 2) from our data, although the sister relationship (44: BS 7) between the New World Pyrrhopygini (45) and the Old World Tagiadini (51) receives good support. Relationships within Tagiadini receive good (53: BS 6) and strong support (52: BS 11; 54: BS 28). Not all members of EvansÕ Tagiades group are included within Tagiadini as defined by our cladogram (Fig. 1) , which has the following topology: (Netrocoryne + (Darpa + (Eagris + (Daimio + Tagiades)))). We have incomplete data (two genes) for two additional genera, Gerosis and Odontoptilum, that when included in the combined analyses (data not shown) are also situated in this clade. Members of this tribe largely include those placed in the Tagiadini by Chou (1994 Chou ( , 1998 , with the exception of Sarangesa and Pseudocoladenia (see below). Tagiadini was first proposed by Mabille (1878) .
Celaenorrhinini, confirmed status (55). This clade (BS 13) is sister to Tagiadini + Pyrrhopygini, with good support (43: BS 7). According to our data, EvansÕ Celaenorrhinus group, given tribal status by Chou (1994 Chou ( , 1998 , is polyphyletic. As noted above, Lobocla is in the Eudamini, and Euschemon is also tentatively placed there. In addition, all members sampled from EvansÕ New World subgroups of the Celaenorrhinus group (including Bungalotis, Dyscophellus, Nascus, and Ocyba) are situated within Eudamini (63, 66, 71, 72) . However, Celaenorrhinus species, along with a few additional genera, do form a monophyletic clade with strong support (BS 13), which appears to represent a tribal entity (Fig. 1) . Genera in our study included within Celaenorrhinini are Celaenorrhinus, Pseudocoladenia, Sarangesa, Eretis, and Alenia. Eretis was formerly placed in EvansÕ Tagiades group, while Alenia was placed in EvansÕ Pyrgus group, based on similarities in wing pattern and antennal nudum number to the other Ôcheckered skippersÕ, such as Pyrgus and Spialia. Relationships within Celaenorrhinini receive moderate (56: BS 3; 57: BS 5) and good (58: BS 9; 59: BS 10) support. The name ''Celaenorrhinae'' was first proposed by Swinhoe (1912) , and was emended to Celaenorrhinini by Clark (1948) , who treated the group as a tribe of the Pyrginae, in which he included species from EvansÕ Telemiades, Erynnis, and Pyrgus groups.
Carcharodini, reinstated status (36). This clade is strongly supported (BS 14) by our data, and is composed of members of EvansÕ Telemiades (Pachyneuria, Viola, Cyclosemia, Staphylus) and Pyrgus (Spialia, Carcharodus, Pholisora) groups, with the following topology: (Cyclosemia + (Carcharodus + Spialia) + ((Pachyneuria + Viola) + (Staphylus + Pholisora))). Members of this clade occur widely in the Palaearctic (Carcharodus), African (Spialia), and Neotropical regions (remaining genera), extending to the Nearctic (Pholisora). Relationships within the Carcharodini receive good (37: BS 8; 38: BS 9) and strong support (40: BS 25; 41: BS 12), including the sister relationship between Staphylus and Pholisora (39: BS 40), corroborating LindseyÕs (1921; also see Lindsey et al., 1931 and Stanford, 1981) belief that these genera are closely related (contra Evans, 1953) . The name ''Carcharodidi'' was first proposed by Verity (1940) , was used as a tribal name by Picard (1947) , and was treated as a subtribe by Koc¸ak (1989) .
Achlyodidini, new status (29).
The union of Achlyodes + Aethilla (both from EvansÕ Erynnis group) is strongly supported by our data (32: BS 24), although the clade uniting these genera with Milanion + Atarnes (30: BS 1) is weakly supported, as is the union of Eracon (from EvansÕ Telemiades group) with the other four genera (29: BS 1). Although Achlyodes and Aethilla are fairly similar skippers on morphological grounds (e.g. , we see few characters that might suggest a close relationship between them, Atarnes + Milanion, and Eracon, and suggest that the clade (29) may not be robust to the addition of characters and taxa in future studies. However, the union of Atarnes + Milanion with Achlyodes + Aethilla has appeared in many anal-yses of these data over the course of this study, even though usually with weak support. In addition, Quadrus and Pythoniades, herein represented on their own clade within ''Pyrginae'' (Fig. 1) , have often grouped with Atarnes + Milanion in previous analyses. We therefore suspect that the position of Quadrus + Pythoniades is likely to change in future analyses employing additional characters and ⁄ or taxa. The name ''Achlyodidae'' was proposed by Burmeister (1878) and has not since been used at the family level.
Erynnini, confirmed status (9). This clade is strongly supported (BS 18) by our data, and has been surprisingly robust to the addition of taxa and characters over the course of this study. Erynnini is composed of most members of EvansÕ Erynnis group, excluding Achlyodes and Aethilla (see above), and including some members of EvansÕ Telemiades group (Gorgythion, Sostrata, Mylon). Relationships within Erynnini mostly receive good (10: BS 6; 11: BS 10; 12: BS 10) and strong support (14: BS 13; 15: BS 14; 17: BS 16; 18: BS 17; 19: BS 13). Recently, Chou (1994 Chou ( , 1998 ) resurrected use of the name Erynnini at the tribal level for the sole Chinese representative of this clade, Erynnis. There is some question as to the correct authorship of the name Erynnini. At least four family-group names have been formed from the genus Erynnis (see Table 1 ), but only one of these, Erynnini Brues and Carpenter, 1932 ; is based on the genus as properly identified. As dictated by Code article 65.2.1 (ICZN, 1999) , the case should be referred to the Commission for a ruling on each of these names. In the meantime, we treat these names as if the Commission has ruled to suppress all but the one properly proposed name (Table 1) . credited Barnes and Lindsey (1922) with the authorship of ''Erynninae''. However, Barnes and Lindsey merely mentioned Erynninae as a possible replacement name for the subfamily Hesperiinae (known in recent decades as Pyrginae), and explicitly chose ''Urbaninae'' as their replacement name. Thus, it is unclear if Erynninae Barnes and Lindsey, 1922 , can be considered to be validly proposed (ICZN, 1999 art. 12) . If so, it has precedence over Brues and CarpenterÕs (1932) authorship.
Pyrgini, confirmed status (22). Whereas Chou (1994 Chou ( , 1998 applied the name Pyrgini to Chinese members of EvansÕ Pyrgus group, recently applied Pyrgini in a much broader way, to all New World genera of Pyrginae that were not included in Eudamini (sensu . As currently composed, with good support (22: BS 7), Pyrgini ( Fig. 1) includes members of EvansÕ Pyrgus and Telemiades groups. Relationships within Pyrgini receive weak (23: BS 1), moderate (24: BS 4), good (25: BS 6; 26: BS 7; 28: BS 10), and strong support (27: BS 19). Xenophanes, for which we currently have only partial data (two genes), is also situated in this clade when it is included in our combined analyses (data not shown). One noteworthy aspect of our results is that the genera of Ôcheckered skippersÕ (Pyrgus, Spialia, Alenia), placed by Evans in his Pyrgus group, are undoubtedly polyphyletic, and are placed in three separate tribes (Pyrgini, Carcharodini and Celaenorrhinini, respectively).
Tribes of Hesperiinae
Aeromachini, new status (105). This clade is strongly supported (BS 12) by our data, and is sister to the rest of the Hesperiinae. Aeromachini includes some (but not all) members of EvansÕ Astictopterus group, including all three members of EvansÕ Halpe subgroup that were included in our analysis (Halpe, Thoressa, Sovia). The sister relationship between Halpe and Thoressa is strongly supported (107: BS 25), although our data provide only moderate support for the sister relationship between Ampittia and Sovia (106: BS 3). Although we were unable to sample the genus Aeromachus (the type genus of Aeromachini), its close relationship to Ampittia, Halpe, Thoressa, and Sovia is supported by the great similarity of male genital structures across these genera (as discussed and figured by Evans, 1937 , and Inoue´and Kawazoe´, 1966 , and we do not hesitate to associate Aeromachus at the tribal level with the four genera we studied. Aeromachini is apparently equivalent to the ''Halpe group'' proposed by Inoue´and Kawazoe´(1966) , probably excluding Arnetta (see Eliot, 1978) . Tutt (1906) originally proposed ''Aeromachinae'' as a subfamily and ''Aeromachidi'' as a tribe for Aeromachus, Ampittia, and Taractrocera, although Taractrocera belongs in a different tribe (see below).
Clade 110. This clade receives good support from our data (BS 6), but we consider its present composition to be tentative, as it contains a disparate mix of taxa that we feel are unlikely to be monophyletic, based on their morphology. In addition, the composition of this clade has varied widely over the course of this study (data not shown), and the large polytomy at clade 110 demonstrates the unresolved nature of relationships among taxa currently placed here. This clade mostly includes Old World genera from EvansÕ Astictopterus, Isoteinon, Ceratrichia, Acleros, Ploetzia, Ancistroides, and Plastingia groups, but also includes New World genera from EvansÕ Carystus and Calpodes groups, and Megathyminae (giant skippers). Despite the presence of a few strongly supported relationships (e.g. 121: BS 15), we feel that the composition of this clade is likely to change with the addition of more taxa and characters, and that its subdivision into more than one tribe in the future seems likely. However, the addition of Koruthaialos, Notocrypta, Pemara, Pyroneura, Gretna, and Pterotei-non in alternative analyses, for which we had only partial data (two genes), does not change the overall composition of this clade (data not shown).
As discussed above, the placement of giant skippers in this clade is not supported by any obvious morphological or biological evidence, but does suggest that they are Ôhighly derivedÕ hesperiines that do not represent a family-or subfamily-level taxon. Although we are confident that the giant skippers are a derived hesperiine clade, we stress that additional study is required to determine their phylogenetic position within the Hesperiinae.
The tropical American genera Perichares and Orses were included in EvansÕ Carystus group. Their sister relationship is strongly supported (127: BS 27) by our data, but their relationship to other taxa in this clade is unresolved. Like the giant skippers, their placement in this clade has been robust to the addition of taxa over time, although their position within the clade has varied (data not shown). The placement of Pyrrhopygopsis (from EvansÕ Calpodes group) in this clade has been less stable (e.g. .
Although the current composition of this clade is highly heterogenous, most of the Old World genera (at least) are likely to be closely related to each other, with respect to other tribes in the subfamily. Should clade 110 or groups therein prove to be robust to the addition of characters and taxa in future studies, several familygroup names are available for members of this clade (Table 1) , and we have sampled type genera of many of these (Appenedix 2). However, none of these names has been widely used in the literature (see . Because of the heterogeneous nature of this grouping, we apply no family-group name to Clade 110 at this time, until its monophyly can be corroborated in future studies; the tentative placement of all associated familygroup names is incertae sedis (Table 1) .
Taractrocerini, confirmed status (132). Voss (1952) was the first to treat this group as a tribal entity within Hesperiinae, but his concept of the group also included members of Thymelicini (from EvansÕ Thymelicus subgroup). Recently, Chou (1994 Chou ( , 1998 employed the tribe Taractrocerini for Chinese members of EvansÕ Taractrocera group. de Jong (1990 ) studied relationships of 13 genera in this group, sensu Evans (1949, minus Prusiana) , and noted that the group is apparently monophyletic. One of the few genus groups proposed by Evans that formed a monophyletic group in our study, the clade comprising Taractrocerini (132) is strongly supported by our data (BS 11), and is sister to the remaining tribes of the Hesperiinae, discussed below. Relationships between genera of Taractrocerini, as indicated by our data, do not entirely agree with those proposed by , suggesting that more taxa need to be sampled in order to better resolve relationships in this tribe. For the most part, relationships among genera in this clade receive moderate or good support from our data, although two primary clades are strongly supported (133: BS 18; 137: BS 12).
Thymelicini, confirmed status (141). Tutt (1905 , in Tutt 1905 -1914 proposed the subfamily ''Thymelicinae'' and tribe ''Thymelicidi'' for members of the genus Thymelicus (an arrangement followed by Tutt, 1906; and 1906 and in Tutt 1905 and -1914 , but Evans (1949) included Thymelicus in his Hesperia group, an action followed by Voss (1952) , who placed the genus in his tribe ''Hesperiidi''. Subsequently, Evans (1955) created the Thymelicus subgroup of his Hesperia group, in which he included Thymelicus, Adopaeoides, Ancyloxypha, Oarisma, and Copaeodes (see Appendix 1), the last three genera included by Voss in his tribe ''Taractroceridi''. Recently, Chou (1994 Chou ( , 1998 employed the name Thymelicini at the tribal level, in which he included just Thymelicus, the sole Chinese representative of this group. Although Thymelicus was not included in our study, as noted by Evans (1949 Evans ( , 1955 , Thymelicus species share morphological features of the antennae, palpi, and male genitalia with the other four genera in his Thymelicus subgroup. de Jong (1984) and de Prins et al. (1992) figured the female genitalia of several Thymelicus species. Ha¨user (1993) commented on the peculiar structure of the corpus bursae in female Thymelicus, with a sclerotized ductus bursae interrupted by a membranous region where the ductus seminalis originates, and suggested this condition may represent a synapomorphy for the Thymelicus subgroup. Examination of the female genitalia of Ancyloxypha, Oarisma and Copaeodes (A.D.W. pers. obs.) has shown that the structure of the ductus bursae in these three species is similar to that found in Thymelicus, adding further evidence of a close relationship between them. Based on these morphological similarities, we apply the name Thymelicini to our clade (141) containing Ancyloxypha, Oarisma, and Copaeodes. However, monophyly of this clade receives only moderate support (BS 4) by our data, and the addition of Thymelicus and Adopaeoides in future studies is needed to test the stability of Thymelicini. Nevertheless, the sister relationship between Oarisma and Copaeodes, as implied by our data, is strongly supported (142: BS 14).
Baorini, new status (151). As noted above, our data strongly support the monophyly of EvansÕ Old World Gegenes group (151: BS 24), although only three genera from the group were included in our final analysis. We were unable to sample Baoris, but it shares many pupal and genitalic characters with Pelopidas, Polytremis, and Iton, as shown by Evans (1937 Evans ( , 1949 and especially by Bascombe et al. (1999) , and we do not hesitate to associate Baoris with our three sampled genera, at the tribal level. The genus Caltoris, for which we currently have only partial data (two genes), is also situated in Baorini (151) when included in alternative analyses (data not shown). ''Baorinae'' was proposed by Doherty (1886) , and was subsequently used at the subfamily level by Bell (1920 Bell ( , 1921 Bell ( , 1926 , who included Baoris, Caltoris, Chapra (a junior subjective synonym of Pelopidas), Parnara, Gegenes, and Iton in the group (all of which were subsequently placed in Evans' 1949 Gegenes group, and in ChouÕs 1994 .
However, our final analysis placed Talides, a New World genus from EvansÕ Carystus group, as sister to Baorini, with moderate support (150: BS 5). In addition, a moderately supported clade (146: BS 5) including members of EvansÕ Vinius (Synapte), Carystus (Dubiella) and Calpodes (Calpodes, Saliana, Thracides) groups is sister to the clade including Talides + Baorini, with moderate support (145: BS 3). While Dubiella, Calpodes, Saliana, and Thracides share various morphological characters (e.g. Evans, 1955) , the inclusion of Synapte in this clade defies any obvious explanation, as it is a much smaller skipper and is morphologically more similar to some other genera in the Vinius group. These genera (excluding Synapte but possibly including Talides), along with related taxa (various genera from the Carystus and Calpodes groups), may eventually warrant tribal status, but for now we do not associate any family-group name with this clade (146). Should these genera occupy a tribal-level position in future studies, two names are potentially applicable, Carystini Mabille, 1878; and Calpodini Clark, 1948 (see Table 1 ).
Clade 144. A strongly supported clade (144: BS 18), including the New World genera Anthoptus and Corticea (from EvansÕ Vinius group), is part of a polytomy including Baorini and associated clades, and the following two tribes (Fig. 2) . The position of this clade basal to the following two tribes, or in a polytomy with them, has been consistent over the course of this study, as characters and taxa have been added (data not shown). Although it is possible that this clade represents a triballevel entity, we feel that the addition of more characters or taxa is needed to corroborate our results, and for now do not associate any family-group name with this clade. We also note that no family-group name is currently available for this clade (Table 1) .
Moncini A. Warren, new tribe (154). Type Genus: Monca Evans, 1955; -This clade receives moderate support (154: BS 5), and includes genera from EvansÕ Vinius (Lento, Vinius), Apaustus (Callimormus, Virga, Mnasicles, Sodalia, Lucida, Vidius, Monca, Cymaenes, Vehilius, Mnasilus, Remella, Papias, Morys, Cumbre, Vettius, Eutychide), Phlebodes (Saturnus, Penicula), Lerodea (Amblyscirtes exoteria -see below) and Calpodes (Panoquina, Niconiades) groups. Additional genera, for which we have incomplete data (two of three genes), are situated in this tribe in alternative analyses (data not shown), including Lerodea (from the Lerodea group), Parphorus (from the Apaustus group), Mucia (from the Phlebodes group), and Halotus (from the Calpodes group). We note that Halotus is sister to Niconiades, as predicted by Burns (1992a) based on morphological similarities. Although many relationships among genera in the Moncini receive good or strong support, the large polytomy at clade 160 probably reflects the need to sample additional taxa. We were surprised to find that, despite the abundance of familygroup names that have been proposed for Old World groups of Hesperiinae, no name is available to apply to clade 154.
Morphology of genera in this clade is rather diverse, and despite molecular characters that differentiate (ICZN 1999 Art. 13.1.1) Moncini from other tribes in our analysis, no putative morphological synapomorphies have yet been identified to diagnose the tribe. However, all genera we include in Moncini have forewing vein M2 originating much nearer to M3 than M1, and most species are ''little brown skippers'' (although some have yellow, tawny, or other colourful markings). Adults of some genera (e.g. Callimormus, Virga) have a long, slender, pointed third segment of the labial palpi (like that found in Thymelicini and some Taractrocerini). Secondary sexual characters of males include the variable presence of forewing stigmata, and in some genera (e.g. Vinius), a tuft of hair-like scales on the dorsal hindwing.
Hesperiini, confirmed status (170). Clark (1948) first recognized the tribe Hesperiini, in which he included various members of EvansÕ Apaustus, Hesperia, and Lerodea groups. Voss (1952) recognized the tribe ''Hesperiidi'', which included some members of EvansÕ Taractrocera, Hesperia, and Lerodea groups. Recently, Chou (1994 Chou ( , 1998 employed the name Hesperiini at the tribal level to represent Hesperia and Ochlodes, the sole Chinese genera in this group. In our study, this clade (170) is composed of members of EvansÕ Phlebodes, Hesperia, Lerodea, and Calpodes groups, and receives good support from our data (BS 8). Other than genera now placed in Thymelicini (141, see above), all genera in EvansÕ Hesperia group appear to be members of Hesperiini (except Halotus, see above). Some genera from EvansÕ Oeonus subgroup of the Phlebodes group are situated in this clade (Decinea, Caligulana, Conga), as are some members of EvansÕ Calpodes group (Thespieus, Nyctelius, Lindra) and one species from EvansÕ Lerodea group (Notamblyscirtes simius-see below). In addition, Xeniades (from EvansÕ Calpodes group), for which we currently have only partial data, is situated in this clade when included in our analyses, as sister to Thespieus (data not shown). For the most part, relationships between genera of Hesperiini receive moderate or good support by our data. Two clades receive strong support, including 178 (BS 11), and Appia + Pompeius (184: BS 17).
Genus-level relationships
More than one species from certain genera were included in our analysis. In both cases where two individuals of the same species were included, they emerged as sister taxa (116 Ancistroides nigrita, 136 Suniana sunias). However, in cases where two or more species from a genus were included, some congeners emerged as sister taxa (13 Ebrietas infanda + E. anacreon; 19 Erynnis afranius + E. horatius; 27 Pyrgus scriptura + P. ruralis; 153 Pelopidas mathias + P. thrax; 157 Euytchide olympia + E. paria; 158 Panoquina ocola + P. hecebolus; 164 Morys micythus + M. valda), while congeners discussed below did not.
We sampled two species of Urbanus (sensu Evans, 1952) , U. dorantes and U. simplicius, members of Eudamini (''Pyrginae''). These did not emerge as sister taxa in our analysis, supporting SteinhauserÕs (1987) conclusion that the genus Urbanus is polyphyletic. Urbanus dorantes emerged in a sister relationship with Thorybes pylades (88: BS 7), and U. simplicius is sister to (Autochton + (Achalarus + (U. dorantes + T. pylades))), with good support (85: BS 8).
As noted by various authors (e.g. Lindsey and Miller, 1965; de Jong and Treadaway, 1993; Austin and Steinhauser, 1996; , the pyrgine genus Celaenorrhinus is the only pan-tropical skipper genus. As discussed by de Jong (1982), this genus displays considerable morphological diversity, both in wing pattern and in the distribution of secondary sexual characters. We sampled one New World (C. eligius) and one Old World (C. leona) species of Celaenorrhinus, which did not appear as sister taxa in our analysis. Celaenorrhinus eligius emerged as sister to the remaining genera of Celaenorrhinini (55: BS 13), but C. leona is sister to Alenia, with moderate support (57: BS 5). Despite this, as we sampled just two of over 90 currently recognized species of Celaenorrhinus (Vane-Wright and de Jong, 2003), we feel it is premature to challenge the monophyly of the genus, as defined by de Jong (1982) .
As currently circumscribed, the genus Pyrgus has an unusual Holarctic and Neotropical distribution (Warren, 1926; de Jong, 1972) . We sampled three New World species of Pyrgus: P. ruralis, P. scriptura and P. communis. Two of these, P. ruralis and P. scriptura, emerged as sister taxa, with strong support (27: BS 18), while P. communis emerged as sister to Heliopetes, with good support (28: BS 7). This suggests that the genus Pyrgus may be paraphyletic with respect to Heliopetes and Heliopyrgus (see Austin and Warren, 2001) . Until additional species of Pyrgus, Heliopyrgus, and Heliopetes can be sampled, we retain P. communis and its New World relatives (e.g. P. c. chloe, P. albescens, P. adepta, P. orcynoides, P. oileus, P. orcus, P. brenda, P. philetas, P. veturius; see Austin and Warren, 2001 ) in the genus Pyrgus, but stress that this arrangement requires further study, and note that a new genus is perhaps needed at least for the primarily Neotropical P. communis group.
Within Trapezitinae, we sampled two species of Toxidia and two species of Hesperilla, but neither genus emerged as a monophyletic clade. Toxidia peron emerged as sister to Signeta flammeata (104: BS 6), and Toxidia doubledayi emerged as sister to T. peron + S. flammeata (103: BS 9). As noted by Atkins et al. (1991) based on the morphology of immatures and adults, Signeta is very closely related to Toxidia, and the two genera are separated primarily on the basis of differences in the size and shape of the male forewing stigma. Our results suggest that Toxidia may be paraphyletic with respect to Signeta, but we feel that the other species of Signeta (S. tymbophora), and additional species of Toxidia should be sampled and analysed before formally changing the composition or synonymy of these genera. The genus Hesperilla is morphologically diverse, with multiple species groups (Atkins, 1978) . The two Hesperilla species we sampled are H. ornata and H. donnysa. Hesperilla ornata emerged as sister to Oreisplanus perornata, with strong support (102: BS 19), while H. donnysa is sister to H. ornata + O. perornata (101: BS 5). These results suggest that Hesperilla may be paraphyletic with respect to Oreisplanus, and that Oreisplanus might best be considered a Ôspecies groupÕ of Hesperilla. However, until the remaining species of Oreisplanus (O. munionga) and the 12 remaining species of Hesperilla can be sampled, we hesitate to disrupt the current generic arrangements (e.g. Atkins and Edwards, 1996; Braby, 2000 Braby, , 2004 . Burns (1990) commented on the hesperiine genus Amblyscirtes, placed by Evans (1955) in his Lerodea group. He suggested that Amblyscirtes is not related to other members of the Lerodea group, and that it was closely related to genera in EvansÕ Apaustus group, such as Mnasicles and Remella. He also noted that one species, simius, did not belong in Amblyscirtes, based on male genitalia that ''differ radically'' from other species in the genus. However, over concern that simius may be related to a Neotropical genus unfamiliar to him, Burns treated simius as incertae sedis, and did not suggest to which of EvansÕ groups of hesperiine genera it may belong. Scott (2006) subsequently proposed the generic name Notamblyscirtes for simius. In addition to N. simius, we sampled one Amblyscirtes species, A. exoteria, whose presence in Amblyscirtes has not been disputed (e.g. Burns, 1990) . According to our results, the two species are situated in separate tribes. Notamblyscirtes simius is in Hesperiini, in a sister relationship with Euphyes (177: BS 5). Amblyscirtes exoteria, presumably along with other Amblyscirtes species, is situated in Moncini, in a sister relationship with Mnasicles + Remella (167: BS 8), corroborating Burns' (1990) conclusion.
Conclusion
Here we have proposed a new family-level synonymy for the Hesperiidae, and have made a preliminary effort to establish a tribal nomenclature for the family (Table 1) . We have identified several strongly supported monophyletic taxa, such as Pyrrhopygini, Erynnini, Trapezitinae, Aeromachini, and Taractrocerini, and have demonstrated strong support for the monophyly of the family. We have defined several unresolved issues that require further study, such as the paraphyly of ''Pyrginae'' and the phylogenetic position of ''Megathyminae'', a group we tentatively consider to be infra-tribal. We feel that the addition of more taxa and characters will be required to strengthen hypotheses of relationships presented here, but that our current arrangement represents a more natural classification than that proposed by Evans and modified by subsequent authors. We plan a second publication that will combine these data with morphological characters, and will use comparative morphology to integrate all genera of the Hesperiidae into a tribal classification (A. Warren, J.R. Ogawa and A.V.Z. Brower, unpublished data). 
EU364300
Full data are preserved with each voucher specimen and are available upon request. Name combinations used in this list mostly follow Bridges (1993) and . Species-level names applied in this table that do not follow Bridges or Mielke are not indended to represent new taxonomic acts.
As shown by Hebert et al. (2004 , also see Brower, 2006a , Astraptes ''fulgerator'' in Guanacaste, Costa Rica, apparently represents multiple species. ADW = Collection of Andrew D. Warren, Castle Rock, Colorado, USA; vouchers will eventually be placed in a public institution. AVZB = Collection of Andrew V. Z. Brower, Murfreesboro, Tennessee, USA; vouchers will eventually be placed in the AMNH, NY, USA. MCZ = Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA.

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