Кавказский энтомол. бюллетень 12(1): 145–156 © CAUCASIAN ENTOMOLOGICAL BULL. 2016

A molecular phylogeny of the subfamily Polyommatinae (Lepidoptera: Lycaenidae)

Молекулярная филогения подсемейства Polyommatinae (Lepidoptera: Lycaenidae)

B.V. Stradomsky Б.В. Страдомский

Institute of Arid Zones SSC RAS, Chekhov str., 41, Rostov-on-Don 344006 Russia. E-mail: bvstr@yandex.ru Институт аридных зон ЮНЦ РАН, пр. Чехова, 41, Ростов-на-Дону 344006 Россия

Key words: Lepidoptera, Lycaenidae, Polyommatinae, phylogeny, molecular genetics, morphological research. Ключевые слова: Lepidoptera, Lycaenidae, Polyommatinae, филогения, молекулярно-биологические и морфологические исследования.

Abstract. Molecular and morphological study of the subfamily Polyommatinae allows to make the following conclusions: the tribe Candalidini and the genus Cupidopsis should be excluded from the subfamily. The status of the tribe Niphandini should be reduced to a subtribe level. Thus, subfamily Polyommatinae consists of two tribes: Lycaenesthini and Polyommatini. Elimination of the non-taxonomic rank “section” and the combined morphological and genetic analysis make it possible to distinguish 22 subtribes within the tribe Polyommatini, which meet the requirements of monophyly: Brephidiina, Pithecopina, Niphandina, Danina, Azanina, Theclinesthina, Lycaenopsina, Jamidina, Cacyreina, Actizerina, Uranothaumatina, Lampidina, Zizulina, Catochrysopsina, Scolitantidina, Castaliina, Oboroniina, Leptotina, Zizeeriina, Fameganina, Everina, and Polyommatina.

Резюме. Молекулярно-генетическое и

морфологическое изучение голубянок подсемейства Polyommatinae позволяет прийти к следующим выводам: триба Candalidini и род Cupidopsis должны быть выведены из состава подсемейства, а триба Niphandini понижена в статусе до уровня подтрибы. Таким образом, в подсемействе Polyommatinae остаются две трибы: Lycaenesthini и Polyommatini. Упразднение внетаксономического ранга «секция» и комплексный морфолого-генетический анализ позволяют выделить в составе трибы Polyommatini следующие 22 подтрибы, отвечающие требованию монофилии: Brephidiina, Pithecopina, Niphandina, Danina, Azanina, Theclinesthina, Lycaenopsina, Jamidina, Cacyreina, Actizerina, Uranothaumatina, Lampidina, Zizulina, Catochrysopsina, Scolitantidina, Castaliina, Oboroniina, Leptotina, Zizeeriina, Fameganina, Everina и Polyommatina.

Currently the most commonly used and adequate system of Lycaenidae is the concept of Eliot [1973]. It is based solely on an analysis of morphological characters. However, the author uses unreasonably excessive fragmentation of many genera, as well as higher-level taxa.

The author also uses such non-taxonomic concept as a section (Section sensu Eliot).

The leading role in the study of taxonomy and systematics of organisms is currently allocated to molecular genetic research. This rule undoubtedly applies to Blue butterflies, and the publication of many articles in the last decade only emphasizes that [Wiemers, 2003; Kandul et al., 2004; Lukhtanov et al., 2005; Vodolazhsky et al., 2009; Wiemers et al., 2010; Ugelvig et al., 2011; Vila et al., 2011; Talavera et al., 2013, 2015; Stradomsky, 2014]. Traditional morphological methods can not ensure the construction of a natural system of Lycaenidae. High variability of the wing pattern, structure of genitalia and other features cannot create an adequate system of Blue butterflies. However, the analysis of some morphological characters can be used as an additional criterion for the construction of the system. First of all, this holds true for the study of genitalia.

The goal of this study was an attempt to build a natural phylogeny of one of the largest subfamilies of Blue butterflies, the Polyommatinae. This subfamily contains species with the extremely pronounced heterogeneity of external morphological characteristics and variability of the genital apparatus. The specimens of the subfamily Polyommatinae were studied with the use of molecular genetic methods. Specifically, we examined the following genetic markers: sections of the mitochondrial gene Cytochrome Oxidase subunit I, the nuclear Elongation Factor 1-alpha (the nuclear coding sequence) and the nuclear noncoding sequence internal transcribed spacer 2 (the nuclear noncoding sequence). At the same time the genital structures of Polyommatinae representatives have been investigated.

Material and methods

All specimens examined in this study are archived at the museum of the Institute of Arid Zones SSC RAS (Rostov-on-Don, Russia) as voucher specimens. Features of studied specimens are presented in the Table 1. Eighty nine species were examined.

Table 1. List of material with voucher codes and GenBank accession numbers.

Таблица 1. Исследованный материал: музейные номера и присвоенные номера Генбанка.


Table 1 (continuation).

Таблица 1 (продолжение).


Parameters for methods of DNA extraction were described previously [Vodolazhsky, Stradomsky, 2008].

We amplified DNA 5’ section of the mitochondrial gene Cytochrome Oxidase subunit I (COI), the nuclear Elongation Factor 1-alpha (Ef-1a) and the nuclear noncoding sequence internal transcribed spacer 2 (ITS2) on the Mastercycler gradient (Eppendorf). The following cycling protocols were used: an initial 4 min denaturation at 95° C and 40 cycles of 30 s denaturation at 95° C, 30 s annealing at 53° C and 60 s extension at 72° C.

We used the following PCR primer pairs: forward,

5’-TAG CGA AAA TGA CTT TTT TCT A-3’ (reserve forward 5’-GGT CAA CAA ATC ATA AAG ATA TTG G-3’) with reverse, 5’-TTG CTC CAG CTA ATA

CAG GTA A-3’ (reserve reverse 5’-TAA ACT TCA GGG TGA CCA AAA AAT CA-3’) were used to amplify COI. Ef-1a was amplified with forward, 5’-TAC CAT CGA GAA GTT CGA GAA G-3’ (reserve forward 5’-TGA AGG CCG AAC GTG AAC GTG G -3’) and reverse, 5’-GCC ACC

CCT TGA ACC AGG GCA T-3’. ITS2 was amplified with forward, 5’-GGG CCG GCT GTA TAA AAT CAT A-3’

(reserve forward 5’-ACT CCT GTC TGA GGG CCG

GCT G-3’) and reverse, 5’-AAA AAT TGA GGC AGA

CGC GAT A-3’ (reserve reverse 5’-TGA GGC AGA CTC GAT ATC CGT C-3’) [Stradomsky, Fomina, 2013; Stradomsky, 2014].

Amplified fragments were separated using an automated sequencing machine (Applied Biosystems 3500). The analysis of primary nucleotide sequences was made with the help of the application BioEdit Sequence

Alignment Editor, version [Hall, 1999].

Summary COI-Ef-1a-ITS2 nucleotide sequences were treated quantitatively using MEGA5 [Tamura et al., 2011] methods Minimum-Evolution (ME) and Maximum Likelihood (ML) and were represented as ME- и ML- cladograms.

Results and discussion

This study based on a complex analysis of three genetic markers of blues butterflies from 4 tribes and 30 sections (sensu Eliot) of subfamily Polyommatinae, as well as an outgroup which includes some members of the subfamilies Lycaeninae, Theclinae and Miletinae. We used nucleotide sequences that are associated with various types of the evolutionary process as a marker: mitochondrial gene COI, the nuclear gene encoding a protein Ef-1a, as well as nuclear nucleotide sequence noncoding a protein ITS2, which is largely nondependent on external factors selection.

Obtained ME- and ML-cladograms (Figs 1, 2) have the maximum similarity, except for a insignificant displacement of some small clades. It should be noted that the obtained dendrograms have significant similarity with the ML-phylograms of tribe Polyommatini, which is based on the analysis of species and genes which are in many ways rather different [Vila et al., 2011: Fig. 2]. This coincidence suggests that the relationship reflected in the cladogram is very close to the natural relationship of taxa in the subfamily Polyommatinae.

In the first instance we consider an outgroup which includes representatives of subfamilies Lycaeninae, Theclinae and Miletinae. It should be noted that in this group there are some taxa previously attributed to the subfamily Polyommatinae, namely, representatives of the tribe Candalidini (C. helenita) and the section Cupidopsis sensu Eliot (C. cissus and C. iobates) of the tribe Polyommatini.

A comparison of the morphology of genitalia of

C. helenita, S. lemolea (Miletinae) and H. astyla (Theclinae) (Figs 3–5) demonstrates their undeniable similarities. In addition, the structure of male fore tarsus in Candalidini is different from that of Polyommatinae. Consequently, the molecular genetic and some morphological characteristics suggest the need for an exclusion of tribe Candalidini from the subfamily Polyommatinae.

Comparison of genitalia of C. cissus and H. astyla (Figs 5–8) shows the uniformity of their structure in a generalized plan of the lateral view (Figs 5, 7), as well as of separate structures, for example, uncus and gnathos in ventral view (Figs 6, 8). We can note a homogeneous structure of the lobes of uncus C. cissus and H. astyla, which form rounded diamond-shaped outgrowths with

torsion on the apex and long sickle-shaped branches of gnathos.

We should also note that the butterflies of the genus Cupidopsis Karsch, 1895 have 10 veins on the forewing, which is characteristic only for some genera of the subfamily Theclinae, including the genus Hypothecla Semper, 1890.

Thus, the results of molecular genetic analysis, as well as some significant morphological characteristics, indicate that the species of the genus Cupidopsis (and, consequently, Cupidopsis section sensu Eliot), most likely do not belong to the subfamily Polyommatinae.

All the other taxa of the subfamily Polyommatinae, except those mentioned above, form a monophyletic group. The first of the isolated clades of the subfamily includes genera Cupidesthes Aurivillius, 1895 and Anthene Doubleday, 1847, which constitute the tribe Lycaenesthini. Members of the tribe are still close to the species of the outgroup based on morphological features, which is especially characteristic for the genus Cupidesthes. The genitalia of representatives of the genera Cupidesthes (subfamily Polyommatinae) and Satyrium Scudder, 1876 (subfamily Theclinae) are very similar in the lateral and in the ventral view (Figs 9–12). They have an expressed saccus, wide domed dorsal structures, a long thin aedeagus with adjacent small elongated valvae, long widely rounded branches of gnathos, and small ventrally oriented lobes of uncus.

The next three sister clades are two sections and one tribe sensu Eliot: Brephidium section, Pithecus section and the tribe Niphandini sensu Eliot. Since all of these clades are at least equivalent, it is necessary to set a lower status for the tribe Niphandini and set it as a subtribe Niphandina, placed in the tribe Polyommatini. In addition, there is a need to clarify a taxonomic status of two sections: Brephidium section and Pithecops section sensu Eliot, thus setting subtribe Brephidiina and Pithecopina also within the tribe Polyommatini.

The analysis of genitalia of the genera representing these subtribes, namely Brephidium Scudder, 1876, Pithecops Horsfield, [1828] and Niphanda Moore, [1875] (Figs 13, 14, 16) indicates that all of them have an original, distinct structure. This fact also substantiates an independent status of the designated subtribes. Genitalia in the genus Brephidium are especially unique. They are characterized by very little vulva, thin vinculum and large tegument with thick spinous hairs and processes. On the contrary, the genitalia of the tribe Niphandini are very primitive. They have a dome-shaped dorsal structure with little lobes of uncus and crescent-shaped branches of gnathos similar to those of genera Cupidesthes and Satyrium (Figs 9–12).

Further, we distinguish two sister clades on both cladograms. The first clade includes Danis section and Nacaduba section sensu Eliot or Danis section sensu Eliot, Nacaduba section sensu Hirowatari [Hirowatari, 1992] and Prosotas section sensu Hirowatari. At the same time, this clade does not include the genus Psychonotis Toxopeus, 1930, which is traditionally included in Danis section. In connection with monophyletic type of this clade there is a need to designate subtribe Danina.

Analysis of the genital structure of the genera


Fig. 1. Lycaenidae: ME-cladogram based on the Minimum Evolution method of analysis of distances for COI, Ef-1a and ITS2 DNA sequences.

Рис. 1. Lycaenidae: ME-кладограмма, построенная методом Минимальной Эволюции на основе последовательностей ДНК COI, Ef-1a и ITS2.


Fig. 2. Lycaenidae: ML-cladogram based on the Maximum Likelihood method of analysis of distances for COI, Ef-1a and ITS2 DNA sequences.

Рис. 2. Lycaenidae: ML-кладограмма, построенная методом Максимального Правдоподобия на основе последовательностей ДНК COI, Ef-1a и ITS2.

Danis Fabricius, 1807, Nacaduba Moore, [1881], Ionolyce Toxopeus, 1929 and Prosotas Druce, 1891 shows that the male genitalia in these taxa have large wide aedeagus, very convex tegumen, well expressed branches of gnathos, moderately narrow valva (Figs 17–20). At the same time

genitalia of the genus Psychonotis are clearly different: tegumen is not convex, little expressed branches of gnathos, valva is very wide (Fig. 46). Thus, the members of the genus Psychonotis clearly stand out from the subtribe Danina by structure of the genitalia.

Sister clade comprises the following sections sensu Eliot: Azanus, Petrelaea and Una. An obviously monophyletic type of clade allows us to combine these sections into subtribe Azanina. Male genitalia in genera Azanus Moore, [1881], Petraea Toxopeus, 1929 and Orthomiella de Nicéville in Marshall et de Nicéville, 1890 (Figs 21, 24, 27) are elongated, flattened, with a long thin aedeagus. The lobes of uncus in these genera in the ventral projection have a triangular shape (Figs 22, 25, 28). Valvae tend to expand from Azanus to Orthomiella (Figs 23, 26, 29). Genitalia of the species of the genera Petrelaea, Orthomiella and Una de Nicéville in Marshall et de Nicéville, 1890 have a small saccus.

Theclinesthes section sensu Eliot is a sister to the subtribe Danina and Azanina clade. The clade slightly changes its position depending of the type of cladograms (ME or ML), but it is always an independent clade. Accordingly, it is possible to establish the monophyletic subtribe Theclinesthina. Male genitalia of the genus Theclinesthes Röber, 1891 have a peculiar structure (Fig. 15): vinculum is very wide, lobes of uncus and branches of gnathos are directed ventrally.

The subtribe Lycaenopsina was designated previously [Stradomsky, 2014; Talavera et al., 2015]. The only question is the scope of taxa included in it. According to Talavera et al. [2015] as well as our ML-cladogram, monophyletic clade combines Lycaenopsis, Eicochrysops and Jamides sections sensu Eliot. However, ME-cladogram defines the Jamides section in the capacity of sister to subtribe Lycaenopsina, and in the case of their joining there will be a paraphyletic taxon. In addition, the analysis of genitalia reveals that male of genera Eicochrysops Bethune-Baker, 1924, Lycaenopsis

C. & R. Felder, [1865], Acytolepis Toxopeus, 1927 (Figs 30– 32), Celastrina Tutt, 1906 have a vinculum with wide wing-shaped expansion on the apex, the lobes of uncus are oriented caudally, and the branches of gnathos are mostly reduced to some extent. At the same time, genitalia of the members of Jamides section have a different shape (Fig. 33): vinculum has no widening, branches of gnathos are well developed, lobes of uncus are directed ventro- cranial. Therefore, there is a need to allocate Jamidina to a separate subtribe.

The following small related subtribes present exclusively African taxa. The members of subtribes Cacyreina (Cacyreus section sensu Eliot) and Actizerina (Actizera section sensu Eliot) have peculiar genitalia (Figs 34, 35). Each subtribe includes only one section sensu Eliot. The genitalia of the genus Actizera Chapman, 1910 have very long, apically pointed lobes of uncus and small branches of gnathos. The lobes of uncus in the genus Cacyreus Butler, 1897 are small, conical, have extra long styloid process, branches of gnathos are long. There is a need to include two sister clades into the subtribe Uranothaumatina: Uranothauma section sensu Eliot and Phlyaria section sensu Eliot. While the genitalia of genera Uranothauma Butler, 1895 and Phlyaria Karsch, 1895 in the lateral projections (Figs 38, 40) have some minor differences, the similarity of genitalia of species of these sections in the caudal projection (Figs 39, 41) is obvious.

Sister subtribes Lampidina (one species, one genus) and Zizulina (two species, one genus) related to African

subtribe are very small specieswise and each one includes only one section sensu Eliot. Male genitalia of the genus Lampides Hübner, [1819] (Fig. 36) are essentially similar to male genitalia of Glaucopsyche section (Figs 42–44), except short branches of gnathos. At the same time, genitalia in the genus Zizula Chapman, 1910 is very peculiar (Fig. 37): aedeagus has a deep cleavage laterally, valve has numerous bristles and processes of a specific shape.

Catochrysopsina stands as a very separate subtribe. It includes a single Catochrysops section sensu Eliot. Male genitalia of the genus Catochrysops Boisduval, 1832 are very primitive (Fig. 45) and similar to male genitalia of the genera Cupidesthes and Satyrium (Figs 9, 11).

Two large sister clades which comprise a significant number of genera are subtribes Scolitantidina and Castaliina. The subtribe Scolitantidina includes only one Glaucopsyche section sensu Eliot and was designated previously [Stradomsky, 2014]. It was explored in detail by molecular methods [Ugelvig et al., 2011]. It should be noted that the male genitalia in this subtribe are quite homogeneous (Figs 42–44): gnathos has long branches, apex is directed caudally, lobes of uncus are short, aedeagus is short, thick, with blunt apex.

Subtribe Castaliina contains a large number of

sections: Castalius section sensu Eliot, Zintha section sensu Eliot, Callictita section sensu Eliot, Upolampes section sensu Eliot, and the genus Psychonotis, previously included in the Danis section. Genitalia in the subtribe Castaliina look heterogeneous (Figs 46, 49, 52, 55, 58), but there is a need to note the following factors. Located basally in subtribe, the genus Psychonotis has small folds on valva (Fig. 47), which are greatly expanded in the genus Zintha Eliot, 1973 (Fig. 53) or even form a separate structure in genera Tarucus (Fig. 55), Castalius (Fig. 59) and Callictita Bethune-Baker, 1908. Furthermore, the valvae of the genera Psychonotis (Fig. 47), Caleta Hiwatari, 1992 (Fig. 50), Zintha (Fig. 53), Tarucus (Fig. 56) have long pointy processes. Also the juxtas of members of all these genera (Figs 48, 51, 54, 57, 60) have a tendency to expand from moderate (Psychonotis) to wide wing-shaped (Tarucus) branches. Thus, noteworthy is a complication of the structure of the ventral parts in male genitalia of all members of the subtribe Castaliina.

The subtribe Oboroniina comprises Euchrysops

section sensu Eliot. Male genitalia in the subtribe are characterized by high affinity (Figs 61–63). They have long narrow valva, very long branches of gnathos and very massive aedeagus.

The subtribe Leptotila contains only Leptotes section sensu Eliot. This subtribe was designated previously [Talavera et al., 2013]. The main feature of the genus Leptotes Scudder, 1876 is aedeagus apically splitted in the ventral view (Figs 66, 67).

The clade that is a sister to the subtribe Leptotina unites members of Zizeeria section sensu Eliot and can be designated as a subtribe Zizeeriina. Male genitalia in genera Zizeeria Chapman, 1910 and Zizina Chapman, 1910 (Figs 64, 65) are very peculiar: aedeagus is very swollen, valvae have numerous long setae, branches of juxta and gnathos are elongated and thin.

Three sister clades, apical in the subfamily Polyommatinae and monophyletic, represent three


Figs 3–12. Genitalia of Lycaenidae species (3–5, 7, 9, 11 – lateral view; 6, 8 – uncus and gnathos; 10, 12 – ventral view).

Рис. 3–12. Гениталии представителей Lycaenidae (3–5, 7, 9, 11 – боковая проекция; 6, 8 – ункус и гнатос; 10, 12 – вентральная проекция).

3 – Spalgis lemolea; 4 – Candalides helenita; 5–6 – Hypothecla astyla; 7–8 – Cupidopsis cissus; 9–10 – Satyrium pruni; 11–12 – Cupidesthes thyrsis.

subtribes: Fameganina and previously named Everina [Talavera et al., 2015] and Polyommatina [Talavera et al., 2013]. They include three sections sensu Eliot: Famegana section, Everes section and Polyommatus section respectively.

The close connection between these subtribes is confirmed by the details of the structure of the male genitalia. Genitalia of the three taxa look different in the lateral view (Figs 68, 71, 74). However, study of the dorsal structures (eg. in genera Famegana Eliot, 1973, Cupido Schrank, 1801 and Plebejus Click 1780) suggests that the structures of lobes of uncus are similar to the subtribes Fameganina (Fig. 69) and Polyommatina (Fig. 75). The uncuses are represented by binate elongated lobes. On the contrary, the unpaired uncus in the subtribe Everina (Fig. 72). At the same time, the branches of gnathos do not

reach uncus in ventral view in genitalia of Famegana and Cupido (Figs 69, 72). A specimen of the genus Plebejus has branches of gnathos located directly under the lobes of uncus (Fig. 75).

Valvae in these three subtribes also have certain similarities. The specimen of the genus Plebejus has a valve with shortened costal and caudal processes (Fig. 76). Specimens of the genus Famegana have greatly elongated processes (Fig. 70). Males of the genus Cupido have elongated, thin and crisscrossing each other processes (Fig. 73). Thus, we can see transitional forms between the subtribe in morphological features, confirming, together with the molecular genetic results, close relationship of Fameganina, Everina and Polyommatina.

Thus molecular genetic and morphological study of the subfamily Polyommatinae allows to make the


Figs 13–29. Genitalia of Lycaenidae species (13–21, 24, 27 – lateral view; 22, 25, 28 – right lobe of uncus; 23, 26, 29 – ventral view).

Рис. 13–29. Гениталии представителей Lycaenidae (13–21, 24, 27 – боковая проекция; 22, 25, 28 – правая лопасть ункуса; 23, 26, 29 – вентральная проекция).

13 – Brephidium exilis; 14 – Pithecops dionisius; 15 – Theclinesthes miskini; 16 – Niphanda fusca; 17 – Danis danis; 18 – Nacaduba kurava; 19 –

Ionolyce helicon; 20 – Prosotas dubiosa; 21–23 – Azanus jesous; 24–26 – Petrelaea dana; 27–29 – Orthomiella pontis.

following conclusions: the tribe Candalidini and the genus Cupidopsis should be excluded from the subfamily. The status of the tribe Niphandini should be reduced to the subtribe level. Therefore, the subfamily Polyommatinae consists of two tribes: Lycaenesthini and Polyommatini. Elimination of a non-taxonomic rank “section” and the combined morphological and genetic analysis make possible to discriminate subtribes from the tribe Polyommatini, which meet the requirements of monophyly: Brephidiina, Pithecopina, Niphandina, Danina, Azanina, Theclinesthina, Lycaenopsina, Jamidina, Cacyreina, Actizerina, Uranothaumatina, Lampidina,

Zizulina, Catochrysopsina, Scolitantidina, Castaliina, Oboroniina, Leptotina, Zizeeriina, Fameganina, Everina and Polyommatina.


The author is grateful to S. Schröder (Köln, Germany), R. Vila (Barcelona, Spain), V. Dinca (Ontario, Canada), Z.F. Fric (České Budějovice, Czech Republic),

G.V. Kuznetsov (Volgograd, Russia), A.M. Dumchus (Novocherkassk, Russia), E.S. Koshkin (Khabarovsk, Russia) for the provided material.


Figs 30–45. Genitalia of Lycaenidae species (30–38, 40, 42–45 – lateral view; 39, 41 – caudal view).

Рис. 30–45. Гениталии представителей Lycaenidae (30–38, 40, 42–45 – боковая проекция; 39, 41 – каудальная проекция).

30 – Eicochrysops hippocrates; 31 – Acytolepis puspa; 32 – Lycaenopsis haraldus; 33 – Jamides celeno; 34 – Actizera stellata; 35 – Cacyreus marshalli; 36 – Lampides boeticus; 37 – Zizula hylax; 38–39 – Uranothauma delatorum; 40–41 – Phlyaria cyara; 42 – Scolitantides orion; 43 – Glaucopsyche alexis; 44 – Phengaris arion; 45 – Catochrysops panormus.


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Figs 46–60. Genitalia of Lycaenidae species (46, 49, 52, 55, 58 – lateral view; 47, 50, 53, 56, 59 – left valve, inner side; 48, 51, 54, 57, 60 – juxta).

Рис. 46–60. Гениталии представителей Lycaenidae (46, 49, 52, 55, 58 – боковая проекция; 47, 50, 53, 56, 59 – левая вальва, вид изнутри; 48, 51,

54, 57, 60 – юкста).

46–48 – Psychonotis caelius; 49–51 – Caleta roxus; 52–54 – Zintha hintza; 55–57 – Tarucus balkanicus; 58–60 – Castalius rosimon.


Figs 61–76. Genitalia of Lycaenidae species (61–66, 68, 71, 74 – lateral view; 67 – aedeagus; 69, 72, 75 – uncus and gnathos; 70, 73, 76 – left valve, inner side).

Рис. 61–76. Гениталии представителей Lycaenidae (61–66, 68, 71, 74 – боковая проекция; 67 – эдеагус; 69, 72, 75 – ункус и гнатос; 70, 73, 76 – левая вальва, вид изнутри).

61 – Oboronia ornata; 62 – Thermoniphas alberici; 63 – Lepidochrysops intermedia; 64 – Zizina otis; 65 – Zizeeria knysna; 66–67 – Leptotes pirithous; 68–70 – Famegana alsulus; 71–73 – Cupido minimus; 74–76 – Plebeius argus.