Review Article |
Corresponding author: Vladimir Lukhtanov ( lukhtanov@mail.ru ) Academic editor: Nazar Shapoval
© 2014 Vladimir Lukhtanov.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lukhtanov V (2014) Chromosome number evolution in skippers (Lepidoptera, Hesperiidae). Comparative Cytogenetics 8(4): 1-19. https://doi.org/10.3897/CompCytogen.v8i4.8789
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Lepidoptera (butterflies and moths), as many other groups of animals and plants, simultaneously represent preservation of ancestral karyotype in the majority of families with a high degree of chromosome number instability in numerous independently evolved phylogenetic lineages. However, the pattern and trends of karyotype evolution in some Lepidoptera families are poorly studied. Here I provide a survey of chromosome numbers in skippers (family Hesperiidae) based on intensive search and analysis of published data. I demonstrate that the majority of skippers preserve the haploid chromosome number n=31 that seems to be an ancestral number for the Hesperiidae and the order Lepidoptera at whole. However, in the tribe Baorini the derived number n=16 is the most typical state which can be used as a (syn)apomorphic character in further phylogenetic investigations. Several groups of skippers display extreme chromosome number variations on within-species (e.g. the representatives of the genus Carcharodus Hübner, [1819]) and between-species (e.g. the genus Agathymus Freeman, 1959) levels. Thus, these groups can be used as model systems for future analysis of the phenomenon of chromosome instability. Interspecific chromosomal differences are also shown to be useful for discovering and describing new cryptic species of Hesperiidae representing in such a way a powerful tool in biodiversity research. Generally, the skipper butterflies promise to be an exciting group that will significantly contribute to the growing knowledge of patterns and processes of chromosome evolution.
Lepidoptera , Hesperiidae , karyotype evolution, chromosome number, cryptic species, phylogeny, chromosomal conservatism, chromosomal instability
The main karyotypic features of organisms, particularly the number of chromosomes, tend to be stable within species (
In contrast to chromosomal conservatism, chromosomal instability characterizes situations where multiple closely related taxa (populations, subspecies and/or species) belonging to a single phylogenetic lineage differ drastically from each other by major chromosomal rearrangements, sometimes resulting in high variability in chromosome number.
Both phenomena - chromosomal conservatism and chromosomal instability - are clearly expressed in insects of the order Lepidoptera (butterflies and moths). The modal haploid number of chromosomes (n) of n = 31 or n = 30 (
Skippers (the family Hesperiidae) are studied to a lesser extent with the respect of karyotype evolution than the other butterfly families mentioned above (but see:
Here I provide a first world-wide survey of chromosome numbers in skippers based on intensive search and analysis of published data.
The results of literature search are presented in the Table below. It includes all the discovered chromosome counts except n=13 for Ochlodes venatus (Bremer et Grey, 1853), noted by
The classification of skippers accepted in this paper follows Warren and colleagues (
The table gives the chromosome numbers of 205 species of skippers, i.e. about 5% of the species of the world fauna. This number is not enough to infer any final statements about peculiarities of chromosome numbers distribution within the Hesperiidae. However, several tentative conclusions can be made. The haploid chromosome number n=31 was found in 50 studied species of skippers and, thus, it is a clear modal number for the family at whole. Interestingly, n=31 was found in representatives of all investigated subfamilies, except for Heteropterinae. However, in the last subfamily only one species was karyologically studied until now, and discovery of n=31 in Heteropterinae is not excluded in future. The next most common numbers are n=29 (43 species), n=30 (33 species) and n=28 (13 species).
Subfamilies Coeliadinae and Eudaminae have a sharp peak at n=31. In the subfamily Trapezitinae n=31 was also found (only one species studied).
Within the subfamily Pyrginae, the modal number n=31 is found in the tribe Erynnini. The tribe Pyrrhopygini is characterized by the most common n=28. The modal number in the tribe Tagiadini is n=30. The tribe Carcharodini has peaks at n=30 and n=31. In the tribe Pyrgini, n=29, n=30 and n=31 were found as the most common numbers.
In the family Heteropterinae n=29 was found (only one species studied).
Within the subfamily Hesperiinae, the tribes Taractrocerini, Thymelicini, Calpodini, Moncini and Hesperiini are characterized by the most common n=29. Very variable chromosome numbers (from n=5 to n=50) were found in the tribe Aeromachini. It is difficult to infer the modal number for the last tribe. However, it should be noted that one species, Thoressa varia, has n=31 as the majority of other skippers. The tribe Baorini (subfamily Hesperiinae) has a clear peak at n=16, so it is exceptional with respect to the modal number of chromosomes.
The overall evidence indicates that chromosome numbers of Coeliadinae, Eudaminae, Trapezitinae, Pyrginae and Hesperiinae conform to the lepidopteran modal of n=31 (
Several groups of skippers display extreme chromosome number variations at the within-species level (Table). The most extreme variations in number of chromosome elements were observed in first meiotic metaphase of Carcharodus boeticus, C. dravira and C. flocciferus (Table,
Second, this kind of variation can be caused by violations in meiotic chromosome pairing resulting in appearance of univalents (instead of bivalents) in meiotic prophase (
Between-species variation exists in numerous genera of skippers (Table 1) and is especially expressed in the Nearctic genus Agathymus Freeman, 1959, in which the range of haploid numbers was discovered from n =5 in A. aryxna to n=38 in A. alliae (
Chromosome number of skippers (Lepidoptera, Hesperiidae) of the world fauna (Us are univalents; 2n is diploid chromosome number). <br/> Years of the species descriptions are given square brackets in cases where they were not stated in the original sources but were inferred from reliable external evidence.
# | Species | Haploid chromosome number | Country | Reference |
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Subfamily Coeliadinae | ||||
1 | Bibasis aquilina (Speyer, 1879) | 29 | Japan |
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B. a. chrysaeglia (Butler, 1881) | 31 (2n=62) | Japan |
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2 | B. jaina formosana Fruhstorfer, 1911 | 31 | Taiwan |
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3 | Choaspes benjaminii (Guérin-Méneville, 1843) | 31 | Japan |
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Ch. b. japonica (Murray, 1875) | 31 | Japan |
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4 | Coeliades anchises jucunda (Butler, 1881) | 30 | Oman |
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5 | C. ernesti (Grandidier, 1867) | 31 | Madagascar |
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6 | C. fervida (Butler, 1880) | 23 | Madagascar |
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7 | C. forestan arbogastes (Guenee, 1863) | 31 | Madagascar |
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8 | C. ramanatek (Boisduval, 1833) | 31 | Madagascar |
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Subfamily Euschemoninae no chromosomal data available | ||||
Subfamily Eudaminae | ||||
9 | Achalarus casica (Herrich-Schäffer, 1869) | 29 | USA (Texas) |
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10 | A. lyciades (Geyer, 1832) | 31 | USA (Connecticut) |
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11 | A. toxeus (Plötz, 1882) | 16 | Mexico |
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12 | Astraptes anaphus (Godman et Salvin, 1896) | 31 | Bolivia |
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13 | A. fulgerator (Walch, 1775) | 31 | Peru |
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14 | A. naxos (Hewitson, 1867) | 31 | Brazil |
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15 | A. phalaecus (Godman et Salvin, 1893) | 25 | Guatemala |
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16 | A. longipennis (Plötz, 1882) | 31 | Costa Rica |
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31 | Peru |
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31 | Brazil |
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17 | Autochton sp. | 20, 21 | Brazil |
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18 | Chioides albofasciatus (Hewitson, 1867) | 31 | Mexico |
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Ch. albofasciatus (Hewitson, 1867) (as Ch. catillus) | 31 | Mexico |
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Ch. albofasciatus (Hewitson, 1867) | 31 | USA (Texas) |
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19 | Entheus priassus pralina Evans, 1952 | 22 | Brazil |
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20 | Epargyreus barisses (Hewitson, 1874) | 31 | Argentina |
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21 | E. clarus (Cramer, 1775) | 31 | USA (Florida) |
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22 | E. clavicornis tenda Evans, 1955 | ca 29–30 | Guatemala |
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23 | Oechydrus chersis (Herrich-Schäffer, 1869) | 31 | Bolivia |
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24 | Phocides polybius phanias (Burmeister, 1880) | 16 | Brazil |
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25 | Tarsoctenus praecia plutia (Hewitson, 1857) | 15 | Brazil |
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26 | Thorybes pylades pylades (Scudder, 1870) | 31 | USA (Connecticut) |
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27 | Udranomia spitzi (Hayward, 1942) | 29 | Brazil |
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28 | Urbanus dorantes dorantes (Stoll, 1790) | 31 | Mexico |
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29 | U. doryssus doryssus (Swainson, 1831) | 14 | Costa Rica |
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30 | Urbanus proteus (Linnaeus, 1758) | 31 | Bolivia |
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31 | Mexico |
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31 | USA (Florida) |
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31 | U. simplicius (Stoll, 1790) | 31 | Argentina |
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32 | U. teleus (Hübner, 1821) | 31 | Argentina |
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Subfamily Pyrginae | ||||
Tribe Pyrrhopygini | ||||
33 | Elbella lamprus (Hopffer, 1874) | 40 | Brazil |
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34 | (?) Jemadia sp. | 32(?) | Brazil |
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35 | Mimoniades montana J. Zikán, 1938 | 27 | Brazil |
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36 | M. nurscia (Swainson, 1821) | 28 | Ecuador |
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M. n. malis (Godman et Salvin, 1879) | 28 | Colombia |
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37 | Mimoniades sp. | 21 | Colombia |
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38 | Mimoniades sp. | 28 | Colombia |
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39 | M. versicolor (Latreille, [1824]) | 28 | Brazil |
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40 | Pyrrhopyge charybdis Westwood, 1852 | 14(?) | Brazil |
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41 | P. pelota Plötz, 1879 | 28 | Argentina |
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42 | Pyrrhopyge sp. | 15 | Brazil |
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43 | Sarbia sp. | 30 | Brazil |
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Tribe Tagiadini | ||||
44 | Daimio tethys (Ménétriés, 1857) | 30 | Japan |
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45 | D. t. moorei Mabille, 1876 | 30 | Taiwan |
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46 | Eagris lucetia (Hewitson, 1876) | 30 | Uganda |
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47 | E. sabadius astoria Holland, 1896 | 30 | Kenya |
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48 | Eretis lugens (Rogenhofer, 1891 | 28 | Kenya |
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Tribe Celaenorrhinini | ||||
49 | Sarangesa phidyle (Walker, 1870) | 29 | Senegal |
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Tribe Carcharodini | ||||
50 | Carcharodus alceae (Esper, [1780]) | 31 | Croatia |
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51 | C. boeticus Reverdin, 1913 | 43–47 | Spain |
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C. boeticus Reverdin, 1913 | 40–52 | France |
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C. boeticus Reverdin, 1913 | 38–46 | Italy |
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52 | C. dravira (Moore, 1874) | 37–48 (with Us) | Iran |
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53 | C. flocciferus (Zeller, 1847) | 32–41 (with Us) | France (Cauterets) |
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54 | C. flocciferus (Zeller, 1847) | 42–58 (with Us) | Italy |
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55 | C. lavatherae (Esper, [1783]) | 30 | France (Salau, Ariege) |
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56 | C. orientalis Reverdin, 1913 | 31–32 | Lebanon |
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30 | Turkey (Van) |
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30–37 (with Us) | Turkey (Amasya) |
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57 | C. stauderi ambiguus Verity, 1925 | 30 | Lebanon |
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30 | Turkey |
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58 | Hesperopsis alpheus (W. H. Edwards, 1876) (as Pholisora) | 34 | USA (Texas) |
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59 | Muschampia nomas (Lederer, 1855) | 30 | Lebanon |
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60 | M. proteides (Wagner, 1929) | 30 | Lebanon |
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61 | M. proto (Ochsenheimer, 1808) | 30 | Spain |
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30 | Lebanon |
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62 | Pholisora catullus (Fabricius, 1793) | 29 | ?USA | Lorković in |
63 | Spialia orbifer (Hübner, [1823]) | 30 | Croatia |
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31 | Turkey |
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64 | S. phlomidis (Herrich-Schäffer, [1845]) | 31 | Turkey |
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65 | S. sertorius (Hoffmannsegg, 1804) | 31 | Slovenia |
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Tribe Erynnini | ||||
66 | Chiomara asychis georgina (Reakirt, 1868) | 31 | Mexico |
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Ch. asychis georgina (Reakirt, 1868) | 32 | USA (Texas) |
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67 | Chiomara sp. | 31 | Trinidad |
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68 | Ebrietas anacreon (Staudinger, 1876) | 31 | Argentina |
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69 | E. osyris (Staudinger, 1876) | 31 | Argentina |
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70 | Erynnis baptisiae (W. Forbes, 1936) | 31 | USA (Connecticut) |
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71 | E. funeralis (Scudder et Burgess, 1870) | 31 | Argentina |
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72 | E. horatius (Scudder et Burgess, 1870) | 31 | USA (Florida) |
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73 | E. icelus (Scudder et Burgess, 1870) | 30 | USA (Connecticut) |
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74 | E. juvenalis juvenalis (Fabricius, 1793) | 30 | USA (Connecticut) |
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75 | E. lucilius (Scudder et Burgess, 1870) | 31 | USA (Connecticut) |
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76 | E. marloyi (Boisduval, [1834]) | 31 | Lebanon |
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77 | E. montanus (Bremer, 1861) | 31 (2n=62) | Japan |
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E. montanus (Bremer, 1861) | 31 | Japan |
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78 | E. persius (Scudder, 1863) | 31 | USA (Connecticut) |
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79 | E. tages (Linnaeus, 1758) | 31 | Croatia |
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31 | France |
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31 | England |
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80 | E. tristis tatius (W. H. Edwards, 1883) | 31 | USA (Texas) |
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81 | Gesta gesta (Herrich-Schäffer, 1863) | 32 | Tobago |
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82 | Grais stigmaticus (Mabille, 1883) | 31 | Mexico |
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83 | Theagenes albiplaga (C. Felder et R. Felder, 1867) | 31 | Bolivia |
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Tribe Achlyodidini | ||||
84 | Achlyodes pallida (R. Felder, 1869) (as A. selva) | 15 | Bolivia |
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15 | Mexico |
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85 | Zera zera zera (Butler, 1870) | 34 | Brazil |
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Tribe Pyrgini | ||||
86 | Anisochoria sublimbata Mabille, 1883 | 31 | Argentina |
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87 | Antigonus erosus (Hübner, [1812]) | 31 | Mexico |
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88 | A. liborius Plötz, 1884 | 31 | Argentina |
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89 | Celotes nessus (W. H. Edwards, 1877) | 14, 13 | USA (Texas) |
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90 | Heliopetes arsalte (Linnaeus, 1758) | 30 | Bolivia |
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H. arsalte (Linnaeus, 1758) | 30 | Mexico |
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91 | H. laviana (Hewitson, 1868) | 29 | USA (Texas) |
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92 | H. macaira (Reakirt, [1867]) | 29 | USA (Texas) |
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93 | H. omrina (Butler, 1870) | 30 | Argentina |
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94 | Heliopyrgus americanus (Blanchard, 1852) | 30 | Chile |
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95 | Paches loxus (Westwood, [1852]) | 31 | Guatemala |
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96 | Pyrgus aladaghensis De Prins et van der Poorten, 1995 | ca 18–21 | Turkey | Lukhtanov and Kandul 1995 (in |
97 | P. albescens Plötz, 1884 | 30 (2n=60) | USA (Texas) |
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P. albescens Plötz, 1884 | 28 | USA (Texas) |
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98 | P. alveus (Hübner, [1803]) | 24 | Finland |
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24 | Croatia |
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24 | Turkey | Lukhtanov and Kandul 1995 (in |
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99 | P. bellieri (Oberthür, 1910) | 27 | France |
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100 | P. bocchoris (Hewitson, 1874) | 30 | Argentina |
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101 | P. bolkariensis De Prins et van der Poorten, 1995 | 30 | Turkey | Lukhtanov and Kandul 1995 (in |
102 | P. cacaliae (Rambur, 1839) | 30 | Italy |
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103 | P. carlinae (Rambur, [1839]) | 30 | Italy |
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104 | P. carthami (Hübner, [1813]) | 29 | Italy |
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105 | P. cirsii (Rambur, [1839]) | 30 | France (Peyreleau, Aveyron) |
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106 | P. fides Hayward, 1940 | 30 | Chile |
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107 | P. maculates (Bremer et Grey, 1852) | 31 (2n=62) | Japan |
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108 | P. malvae (Linnaeus, 1758) | 31 | Finland |
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33 | England |
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109 | P. oileus (Linnaeus, 1767) | 30 (2n=60) | USA (Texas) |
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32 | USA (Texas) |
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110 | P. onopordi (Rambur, [1839]) | 30 | France |
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111 | P. serratulae (Rambur, [1839]) | 30 | France |
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112 | Trina geometrina geometrina (C. Felder et R. Felder, 1867) | 31 | Brazil |
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Subfamily Heteropterinae | ||||
113 | Butleria quilla Evans, 1939 | 29 | Chile |
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Subfamily Trapezitinae | ||||
114 | Trapezites eliena Hewitson, 1868 | 31 | Australia |
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Subfamily Hesperiinae | ||||
Tribe Aeromachini | ||||
115 | Aegiale hesperiaris (Walker, 1856) | 24 | Mexico |
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116 | Agathymus alliae (Stallings et Turner, 1957) | 38 | USA (Arizona) |
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117 | A. aryxna (Dyar, 1905) | 5 | Mexico |
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118 | A. baueri (Stallings et Turner, 1954) | 15 | USA (Arizona) |
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119 | A. chisosensis (Freeman, 1952) | 18 | USA (Texas) |
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120 | A. estelleae valverdiensis Freeman, 1966 | 9 | USA (Texas) |
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A. e. estelleae (Stallings et Turner, 1958) | 9 | Mexico |
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121 | A. freemani Stallings, Turner et Stallings, 1960 | 15 | USA (Arizona) |
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122 | A. gilberti Freeman, 1964 | 21 | USA (Texas) |
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123 | A. mariae chinatiensis Freeman, 1964 | 22 | USA (Texas) |
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A. mariae lajitaensis Freeman, 1964 | 22 | USA (Texas) |
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A. mariae mariae (Barnes et Benjamin, 1924) | 22 | USA or Mexico |
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A. mariae rindgei Freeman, 1964 | 22 | USA (Texas) |
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124 | A. micheneri Stallings, Turner et Stallings, 1961 | 20 | Mexico |
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125 | A. neumoegeni florenceae (Stallings et Turner, 1957) | 10 | USA (Texas) |
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A. neumoegeni macalpinei (Freeman, 1955) | 10 | USA (Texas) |
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126 | A. polingi (Skinner, 1905) | 10 | USA (Arizona) |
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127 | A. remingtoni (Stallings et Turner, 1958) | 9 | Mexico |
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128 | Alera vulpina (C. Felder et R. Felder, 1867) | ca27 | Ecuador |
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129 | Ankola fan (Holland, 1844) | 10 | Uganda |
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130 | Arotis derasa (Herrich-Schäffer, 1870) (as Euphyes) | 28 | Brazil |
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131 | Erionota thrax thrax (Linnaeus, 1767) | 29 | Malaysia |
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132 | Euphyes leptosema Mabille, 1891 | ca28 | Argentina |
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133 | Megathymus coloradensis coloradensis Riley, 1877 | 27 | USA |
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134 | M. coloradensis kendalli Freeman, 1965 | 27 | USA (South central Texas) |
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M. coloradensis louiseae Freeman, 1963 | 27 | USA (Western Texas) |
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M. coloradensis navajo Skinner, 1911 | 27 | USA |
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M. coloradensis reinthali Freeman, 1963 | 27 | USA (Texas) |
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M. coloradensis reubeni Stallings, Turner et Stallings, 1963 | 27 | USA (Texas) |
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M. coloradensis stallingsi Freeman, 1943 | 27 | USA |
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M. coloradensis wilsonorum Stallings et Turner, 1958 | 27 | ?Mexico |
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135 | M. violae Stallings et Turner, 1956 | 27 | USA |
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136 | M. yuccae buchholzi Freeman, 1952 | 26 | USA (Florida) |
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137 | Pardaleodes incerta (Snellen, 1872) | 17 | Uganda |
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138 | Stallingsia maculosus (Freeman, 1955) | 50 | USA (Texas) |
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139 | Suastus gremius (Fabricius, 1798) | 23 | Taiwan |
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140 | Thoressa varia (Murray, 1875) | 31 (2n=62) | Japan |
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141 | T. varia (Murray, 1875) | 31 | Japan |
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Tribe Baorini | ||||
142 | Gegenes gambica (Mabille, 1878) | 41 | Yemen |
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41 | Turkey |
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41 | Lebanon |
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143 | Gegenes nostrodamus (Fabricius, 1793) | 15 | Egypt |
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15 | Israel |
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144 | Gegenes pumilio (Hoffmansegg, 1804) | 24 | France |
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24 | Alger |
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145 | Parnara guttata (Bremer et Grey, 1852) | 16 | Japan |
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16 | China |
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146 | Pelopidas conjucta conjucta (Herrich-Schäffer, 1869) | 16 | Hong Kong |
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147 | P. jansonis (Butler, 1878) | 16 (2n=32) | Japan |
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148 | P. mathias (Fabricius, 1798) | 16 | Japan |
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149 | P. thrax (Hübner, [1821]) | 16 | Lebanon |
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150 | Polytremis lubricans (Herrich-Schäffer, 1869) | 16 | Taiwan |
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151 | P. pellucida (Murray, 1875) | 16, 17, 18 (2n=32, 33) | Japan |
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16 | Japan |
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152 | Zenonia zeno (Trimen, 1864) | 16 | Uganda |
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Tribe Taractrocerini | ||||
153 | Ocybadistes walkeri sothis Waterhouse, 1933 | 28 | Australia |
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154 | Potanthus flavus (Murray, 1875) | 29 (2n=58) | Japan |
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155 | Telicota ancilla horisha Evans, 1934 | 29 | Taiwan |
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156 | Telicota colon stinga Evans, 1949 | 29 | Japan (Okinava) |
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157 | T. ohara formosana Fruhstorfer, 1911 | 29 (2n=58) | Taiwan |
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Tribe Thymelicini | ||||
158 | Copaeodes minima (W.H. Edwards, 1870) | 29 | USA (Florida) |
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159 | Thymelicus sylvestris (Poda, 1761) | 27 | England |
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160 | Th. sylvaticus (Bremer, 1861) | 10 (2n=20) | Japan |
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161 | Th. acteon (Rottemburg, 1775) | 28 | Spain |
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162 | Th. hyrax (Lederer, 1861) | 29 | Lebanon |
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163 | Th. leoninus (Butler, 1878) | 9 (2n=18) | Japan |
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164 | Th. lineola (Ochsenheimer, 1808) | 29 | Finland |
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29 | Lebanon |
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Tribe Calpodini | ||||
165 | Ebusus ebusus (Cramer, [1780]) | 29 | Mexico |
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166 | Lychnuchus celsus (Fabricius, 1793) | 30 | Brazil |
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167 | Panoquina hecebolus (Scudder, 1872) | 29 | USA (Texas) |
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168 | Panoquina ocola (W. H. Edwards, 1863) | 29 | USA (Texas) |
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169 | P. panoquin (Scudder, 1863) | 29 | USA (Florida) |
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170 | P. panoquinoides (Skinner, 1891) | 29 | USA (Texas) |
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Tribe Anthoptini no chromosomal data available | ||||
Tribe Moncini | ||||
171 | Amblyscirtes aenus W.H. Edwards, 1878 | 28, 29 | USA (Texas) |
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172 | A. cassus W. H. Edwards, 1883 | 29 | USA (Texas) |
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173 | A. celia (Skinner, 1895) | 29 | USA (Texas) |
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174 | A. phylace W.H. Edwards, 1878 | 29 | USA (Texas) |
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175 | A. texanae Bell, 1927 | 29 | USA (Texas) |
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176 | A. vialis (W. H. Edwards, 1862) | 29 | USA (Connecticut) |
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177 | Cymaenes sp. | 31 | Tobago |
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178 | Enosis immaculata immaculata (Hewitson, 1868) | 29 | Ecuador |
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179 | Lerema accius (Smith, 1797) | 29 (2n=58) | USA (Texas) |
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29 | USA (Texas) |
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180 | Moeris vopiscus (Herrich-Schäffer, 1869) | 27 | Peru |
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181 | Nastra lherminier (Latreille, [1824]) | 30 | USA (Connecticut) |
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182 |
Thargella caura (Plötz, 1882) |
25 | Brazil |
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183 | Vettius coryna (Hewitson, [1866]) | 31, ca32 | Ecuador |
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184 | V. phyllus prona Evans, 1955 | 26 | Brazil |
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185 | V. triangularis (Hübner, [1831]) | 26 | Brazil |
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Tribe Hesperiini | ||||
186 | Asbolis capucinus (Lucas, 1857) | 48 | USA (Florida) |
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187 | Cynea iquita (Bell, 1941) | 29 | Argentina |
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188 | Hesperia comma (Linnaeus, 1758) | 28 | Italy |
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28 | Lebanon |
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189 | H. florinda Butler, 1878 | 28 (2n=56) | Japan |
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190 | Hylephila fasciolata (Blanchard, 1852) | 29 | Argentina |
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191 | H. phyleus (Drury, 1773) | 29 | Argentia |
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29 | USA (Florida) |
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192 | H. signata (Blanchard, 1852) | 29 | Chile |
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193 | Ochlodes ochraceus (Bremer, 1861) | 29 (2n=58) | Japan |
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24 | Japan |
|
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194 | O. sylvanoides (Boisduval, 1852) | 29 | USA |
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195 | O. sylvanus (Esper, 1777) | 29 | Finland |
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29 | Croatia |
|
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196 | O. venatus (Bremer et Grey, 1853) (as sylvanus Esper, 1777) | 29 (2n=58) | Japan |
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197 | Oligoria maculata (W. H. Edwards, 1865) | 29 | USA (Florida) |
|
198 | Poanes hobomok hobomok (Harris, 1862) | 29 | ?USA | Lorković in |
199 | P. taxiles (W. H. Edwards, 1881) | 29 | USA |
|
200 | P. zabulon (Boisduval et Le Conte, [1837]) (as Polites zabulon) | 29 | USA (Connecticut) |
|
201 | Polites themistocles (Latreille, [1824]) | 29 | USA (Florida) |
|
202 | P. vibex catilina (Plötz, 1886) | 29 | Argentina |
|
P. vibex praeceps (Scudder, 1872) | 27 | USA (Texas) |
|
|
P. vibex vibex (Geyer, 1832) | 29 | USA (Florida) |
|
|
203 | Wallengrenia egeremet (Scudder, 1863) | 28 | USA (Texas) |
|
204 | W. otho curassavica (Snellen, 1887) | 28–30 | USA (Texas) |
|
205 | W. premnas (Wallengren, 1860) | 27 | Argentina |
|
Recent years karyological data have been widely used in studies of butterfly taxonomy and in biodiversity research as main or additional chracters for detecting cryptic species (e.g.
In the genus Pyrgus Hübner, [1819], our unpublished chromosome data (see Table) were used to recognize and then to describe two morphologically similar species, P. bolkariensis and P. aladaghensis (
Thus, interspecific chromosomal differences are useful for discovering and describing new cryptic species of Hesperiidae representing in such a way a powerful tool in biodiversity research.
I thank A. Warren (University of Florida) for help and consultations in taxonomy and nomenclature of skippers. The study was supported by the Russian Foundation for Basic Research: mainly by grant RFBR 13-04-92716-IND-a and partially by grant RFBR 14-04-01051-a.