Introduction

Tribe Tomarini is a monotypic genus represented by genus Tomares Rambur, 1840, which involves approximately 10 species and is distributed from Europe and North Africa to Central Asia. Taxonomy of the genus is controversial since the samples lack diagnostic characteristics by which they can be explicitly assigned to species. Generally, most of the Tomares species show individual and local variability in the size of adult individuals with the base colour intensity and the shade and size of the orange spots, which can sometimes completely disappear, on their wings ( Nazari et al. 2020). What they all have on their forewings are a red-orange spot with broad black-brown edges and a red-orange spot on the anal area of the black-brown hindwings. Caterpillars live as endophag on the flower clusters of specific Astragalus species; or they live as ektophag on the flowers of specific Onobyrchis species. Caterpillars live as endophag on the flower clusters of specific Astragalus species; or they live as ektophag on the flowers of specific Onobyrchis species ( Hesselbarth et al. 1995).

Tragacanth gum ( Astragalus spp.) is an important plant which is distributed considerably in Turkey. Its most important industrial area of usage is the production of tragacanth resin. They are destroyed by being disrooted to obtain tragacanth gum in some areas and to be used as fuel and provender in other areas in Anatolia. Some of its species are important food plants for larvae, especially for the Lycaenidae. There are 459 taxa known belonging to 60 sections of genus Astragalus L., which is widely distributed in Turkey with an endemism rate of %51 ( Ekici et al. 2015; Aytaç et al. 2020; Tunçkol et al. 2020; Duman et al. 2020). Larvae of Tomares prefer Astragalus, Astracantha, and Onocrychis from leguminosae ( Leesmants et al. 1986; Koçak, 1983, 1987; Weidenhoffer & Vanek, 1977; Hesselbarth et al. 1995; Tuzov et al. 2000; Rákosy & Craioveanu, 2015; Bury & Savchuk, 2015; Seven, 2014; Nazari & Ten Hagen, 2020).

The taxonomic status of some species remains controversial, due to distinct morphological characters ( Weidenhoffer & Bozano, 2007; Nazari & Ten Hagen, 2020) and differences in ecology and distribution ( Hesselbarth et al. 1995; Van Oorschot & Wagener, 2000). In our study, distribution maps of the Tomares Rbr. and larval food plant Astragalus L. species, which are distributed in Turkey, were prepared in order to contribute to the solution of these problems. The relationship between the distribution of adult Tomares and the larval food plant is insufficient for explaining the distribution model of Tomares in Turkey.

Material and Methods

In the study, distribution maps of Tomares and larval food plants were determined, and distribution maps were prepared. The necessary food plant data for distribution maps were obtained from the literature. Related literatures are given in the food plant section. Hesselbarth et al. (1995) and Koçak & Kemal (2018) are used for the names and distributions of the species without considering the most recent studies on their systematic status. Chamberlain et al. (1970) used for the distribution of Astragalus species. The distribution of species belonging to the genus Astragalus is shown in blue on the map of Turkey. Species are marked with symbols of different colors in blue. The study is related to the second author’s master’s thesis.

Results

The larvae of the genera Tomares prefer 14 species from the genus of Astragalus L. as food plants ( Astragalus macropterus DC , A. ponticus Pall. , A. dasyanthus Pall. , A. schahrudensis Bunge , A. utriger Pall. , A. suprapilosus Gontsch. , A. macrocephalus Willd., A. micropterus Fisch , A. ornithopodioides Lam. , A. densifolius Lam. , A. physodes L. , A. macrocarpus DC , A. leptostachys Pall. , A. vulpinus Willd.). Of these, A. macropterus DC, A. schahrudensis Bunge , A. dasyanthus Pall. , A. suprapilosus Gontsch. , A. macrocarpus DC, A. leptostachys Pall. and A. vulpinus Will. do not show distribution in Turkey.

The genera Tomares which show distribution in Turkey and their food plants.

Tomares romanovi (Christoph, 1882)

Distribution in Turkey: Ağrı, Bingöl, Bitlis, Diyarbakır, Elazığ, Hakkari, Kars, Mardin, Siirt, Tunceli, Van, Batman, Rırnak, Iğdır ( Map 1).

Host plants of larva: Astragalus finitimus, Onobrchis radiata, ( Weidenhoffer & Vanek, 1977; Hesselbarth et al. 1995; Tuzov et al. 2000); Astragalus schahrudensis (Christoph, 1882; Korb 1924; Zhdanko, 1997). ( Map 1)

Tomares callimachus (Eversmann, 1848)

Distribution in Turkey: Bitlis, Diyarbakır, Elazığ, Hakkari, Kars, Malatya, Kahramanmaraº, Mardin, Siirt, Ranlıurfa, Van, Rırnak, Iğdır ( Map 2)

Host plants of larva: Astragalus leptostachys, A. macropterus, A. physodes, A. vulpinus ( Nazari et al., 2020); Astragalus physodes ( Korb, 1924); Hedysarum candidum, Astragalus utriger, Astragalus suprapilosus ( Bury & Savchuk, 2015); Onobrychis radiata ( Weidenhoffer & Vanek, 1977) ( Map. 2)

Adult feeds on the nectar of Onobrychis cornuta and Aethionema speciosum and Geranium sp. ( Koçak & Kemal, 2011). The plants they visit sometimes, although rarely: Holosteum umbellatum, Geranium tuberosum, Erysimum cuspidatum, Iris pumila and Alyssum spp. ( Bury & Savchuk, 2015).

Tomares (nogelii) nesimachus (Oberthür, 1893)

Distribution in Turkey: Adana, Ankara, Antalya, Bilecik, Çankırı, Çorum, Elazığ, Erzincan, Gaziantep, Gümürhane, Hatay, Isparta, İçel, Kayseri, Konya, Malatya, Kahramanmarar, Mardin, Muğla, Niğde, Sivas, Tokat, Tunceli, Ranlıurfa, Yozgat, Bayburt, Karaman ( Map 3).

Host plants of larva: Astragalus densifolius (İçel) ( Leesmants et al. 1986); Astragalus macrocarpus in İsrail ( Larsen & Nakamura, 1983), Astracantha spp. ( Oorschot & Wagener 2000). Adult feeds on the nectar of Astragalus ponticus Çağlayan (Erzincan) ( Hesselbarth et al. 1995), A. macrocephalus finitimus, A. ornithopodioides ( Koçak, 1983).

Tomares (nogelii) nogelii (Herrich-Schäffer, [1851])

Distribution in Turkey: Adana, Afyon, Amasya, Ankara, Antalya, Bolu, Çankırı, Çorum, Diyarbakır, Elazığ, Eskirehir, Gaziantep, Isparta, İçel, Kayseri, Konya, Malatya, Kahramanmarar, Nevrehir, Niğde, Samsun, Sivas, Tokat, Tunceli, Yozgat, Kırıkkale ( Map 4).

Host plants of larva: Astragalus ponticus ( Hesselbarth et al. 1995); Astragalus micropterus ( Koçak, 1987). Adult feeds on the nectar of Astragalus microcephalus, A. ornithopodioides ( Koçak, 1983), Aethionema ( Koçak & Kemal, 2011), A. ponticus ( Hesselbarth et al. 1995) ( Map. 4).

Tomares (nogelii) dobrogensis Caradja, 2005

Distribution in Turkey: Adana, Ankara, Bolu, Nevrehir, Eskirehir ( Map 5).

Host plants of larva: Astragalus ponticus ( Rákosy & Craioveanu, 2015; Tuzov et al. 2000; Bury & Savchuk, 2015;), Astragalus macrocephalus ( Seven, 2014); Astragalus dasyanthus ( Bury & Savchuk, 2015) ( Map. 5).

Two subspecies are known to be from Turkey (ssp. monotana from Bolu, 1939, ssp. uighurica from Adana, Ankara, and Osmaniye).

Tomares nogelii cesaKoçak, Seven & Kemal, 2000

Distribution in Turkey: Elazığ (type locality) ( Map 6).

Host plants of larva: Unknown.

Tomares desinens Nekrutenko & Effendi, 1980

Distribution in Turkey: Van (Koçak & Kemal, 2005), Iğdır ( Koçak & Kemal, 2012) ( Map 7).

The population in Van province is represented by the subspecies ssp.mebeb Koçak & Kemal, 2005. Koçak & Kemal (2018) have not included these species in their studies. The existence and distribution of the species in Turkey ought to be researched.

Host plants of larva: Unknown. Adults feed on the nectar of Erysimum sp. ( Koçak & Kemal, 2011).

Results and Discussion

Nazari & Ten Hagen (2020) examined various species of genus Tomares by using the barcoding and nuclear DNA methods. Molecular studies demonstrate that the genus is monophyletic and is divided into two in terms of biotope, being Africa and Asia ( Krupitsky et al. 2021). Genus Tomares came in sight between early Oligocene and early Miocene, probably in Southwestern Asia. The division of the youngest ancestor of Tomares occurred between mid to late Miocene and mid to late Pliocene, possibly as a result of increasing drought and habitat fragmentation. The differentiation of the Asian lineage appeared during Pliocene and Pleistocene in Southwestern Asia, and it overlapped with Astragalus (Fabaceae), the host plant of Tomares ( Krupitsky et al. 2021). There are approximately over 400 species of Astragalus known in Turkey ( Donner, 1990). Currently, there are 459 species reported, %51 of them being endemic ( Duman et al. 2020). According to the literature, larvae of Tomares prefer Astragalus plants the most among leguminosae. Species of Astragalus genus are common in Turkey; however only 7 species of it are known as food plants of Tomares larvae. This can be associated with the fact that lack of studies on the biology of the Tomares species. When looking at the distribution of Tomares species and food plants, it is seen that the distribution of food plants and Lepidoptera does not match in some places, and the are distributed outside the food plants ( Map 1-7).

Tomares nogelii is one of the most widely distributed species. Its larvae have been reported to feed on Ast. micropterus and Ast. ponticus ( Hesselbarth et al. 1995). Generally, the distribution of this species and the distribution of the food plants in Turkey draw a correspondence ( Map 4). Yet, in areas having records of some species, the relevant Astragalus species are absent. Conceivably, T. nogelii larvae may prefer other species of Astragalus. It has already been reported in some studies that they are fed on Astragantha ( Oorschot & Wagener, 2000).

Kovancı et al. (2009) was recorded of the T. nogelii from Bursa. In that study, the dominant plants in the habitat were Astragalus angustifolius Lam. and Astragalus sibthorpianus Boiss. however, no larvae were obtained from these plants. In the study, it was reported that these two plant species could be larval food plants of T. nogelii. However, this Astragalus species has not been recorded as a food plant for nogelii in any study to date.

Tomares nesimachus larvae feed on Astragalus densiflora ( Leesmants et al. 1986). The distribution of this species and the larvae food plant show correspondence ( Map 3). However, the relevant Astragalus species cannot be seen in places having records of some species. In these places, it might be possible that T. nesimachus larvae prefer distinct Astragalus species. Thus, Astragalus macrocarpus is given as a food plant in Israel ( Larsen & Nakamura, 1983).

Koçak (2000) were discussed the taxonomical status of Tomares nogelii. In that study, are offered superspecies categorie for nogelii. In the sudy, is classified nogelii, dobrogensis and nesimachus as distinct species in the superspecies of nogelii. Koçak et al. (2000) were definied ssp. uighurica under the species dobrogensis from Ankara and ssp. cesa were definied under the species nogelii from Elazığ. Nazari & Ten Hagen (2020) state that T. nogelii and T. nesimachus are different sister species. In the study by Krupitsky et al. (2021), the two species were declared to be the same species and offered as synonyms for each other. On the maps prepared with data analyses in our study, it is evident that there is a correlation between the geographical distribution of these two species. The large overlapping of the distribution range of the two species in the same geography raises doubts about them being separate subspecies. It is also noteworthy that the Astragalus species which are preferred by the larvae are different. These inconsistencies indicate that the discussions about these taxa will continue.

T. dobrogensis, found in Eastern Romania, Ukraine, and Crimea in Europe, are reported from four provinces in Turkey ( Koçak & Kemal, 2018). Known in Europe, the larvae food plant Astragalus ponticus is extensively distributed in Turkey. However, the distribution of this plant species does not correspond to the records of Tomares nogelii dobrogensis in Turkey. Another food plant record known in Europe is A. dasyanthus ( Bury & Savchuk, 2015). This plant species is not distributed in Turkey. In the study of Seven (2014), it is noted that the lays eggs on Astragalus macracephalus and the photographs of the egg are given. In the distribution map prepared, it is seen that the records of the larval food plant and the distribution areas of the adults do not overlap with each other ( Map 5). Although molecular studies confirm that this species is a subspecies of nogelii, it is thought-provoking that both subspecies are distributed in the same region. This raises the following question; I wonder if the nominant species in Europe and the populations in Turkey are different from each other? Could the larvae of this taxon prefer only Astragalus macrocephalus plant in Turkey? In fact, Koçak et al. (2000) named the populations of sp. dobrogensis in Central Anatolia as ughurica. In order to answer these questions, more samples from different regions in Turkey and studies on their biology are needed. Insufficient published data for this species may elucidate the question of why species are more limited than their food crop distribution.

Adults of Tomares romanovi were reported in a hilly area covered with mixed forest ( Quercus, Crataegus, Prunus, Acer, Celtis), malacophil plants and grasses ( Koçak & Kemal, 2011). Known larvae food plant is Astragalus finitimus ( Weidenhoffer & Vanek, 1977Hesselbarth et al. 1995; Tuzov et al. 2000) and Astragalus schahrudensis (Christoph, 1882; Korb, 1924; Zhdanko, 1997). A. schahrudensis, do not show distribution in Turkey. Astragalus finitimus is considered as A. macrocephalus ssp. finitimus in Turkey. Upon examining the distribution, it can be seen that it is also distributed in such areas where the larvae food plant is not distributed at ( Map 1). Additionally, even though the larvae food plant is widely distributed, the distribution of sp. romanovi is limited to Eastern Anatolia. Thus, it bears the possibility of this species benefitting from Astragalus species different from the known ones.

T. callimachus feeds on Astragalus physodes ( Nazari & Ten Hagen, 2020; Korb, 1924). Ssp. acikirensis, the subspecies of this plant species is distributed in Turkey (Aytaç, 2020). The distribution of this plant species does not tally with the distribution range of sp. callimachus ( Map 2). The distribution range of utriger, another larvae food plant, and the distribution range of this species overlaps only in two places. This situation reinforces the possibility of sp. callimachus larvae feeding on other Astragalus species, just like sp. romanovi larvae. In other respects, it is reported by Koçak & Kemal (2011) that adults of this species feed on Onocrychis. This species, not being reported as a larvae food plant thus far, might be used as larvae food plant for the reason that the adults feed on them. In fact, it is stated in the study of Hesselbarth et al. (1995) that Tomares live as ectophag on the flowers of specific Onobyrchis species. Thus, it can explain why the species do not correspond to the distribution of Astragalus species.

T. desinens is represented by mebeb subspecies in Turkey ( Krupitsky et al. 2021). Faunistic records belong to only two places ( Map 7). Its habitat tragacanthic mountain steppe has been greatly degraded and overgrazed, on the roadside where Astragalus campylosema (?) and Hedysarum grow, Hilly place covered by mixed forest ( Quercus, Crataegus, Prunus, Acer, Celtis), with malacophyllous plants and grasses. T. desinens adults feed on the nectar of Erysimum sp. ( Koçak & Kemal, 2011). Each of which can be potential larvae food plants.

Current data on the larvae food plants of Tomares species is not enough to clarify the relation between the distributions of the species and food plants. More data on the biologly of Tomares species are needed.

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Author notes

*Autor para la correspondencia / Corresponding author