Melitaea Fabricius, 1807

Kolesnichenko, Kirill A. & Kotlobay, Anatoly A., 2022, Review of the fritillary species systematically close to Melitaea lutko Evans, 1932 (Lepidoptera: Nymphalidae) with analysis of their geographic distribution and interrelations with host plants, European Journal of Taxonomy 830 (1), pp. 1-60 : 5

publication ID	https://doi.org/ 10.5852/ejt.2022.830.1865
publication LSID	lsid:zoobank.org:pub:C1F47AD9-ECF6-4F0F-9928-55A45332FF4B
DOI	https://doi.org/10.5281/zenodo.6840162
persistent identifier	https://treatment.plazi.org/id/EB42F330-FFD3-B327-9147-F896FBF1F8B2
treatment provided by	Felipe
scientific name	Melitaea Fabricius, 1807
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Genus Melitaea Fabricius, 1807 View in CoL

Fig. 1–30 View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig View Fig , Tables 1–6 View Table 1 View Table 2 View Table 3 View Table 4 View Table 5

Our study indicates the existence of four independent species within the complex under consideration: M. lutko , M. mimetica , M. shahvarica sp. nov., and M. timandra . The latter species is well differentiated into two subspecies that are isolated both geographically and ecologically.

Figures

Fig. 1. Type specimens of M. lutko Evans, 1932, M. mimetica Higgins, 1940, and M. timandra Coutsis & van Oorschot, 2014. A, C, E. UPS. B, D, F. UNS. A–B. M. lutko, holotype, ♂ (BMNH). C– D. M. mimetica, paratype, ♀ (BMNH). E–F. M. timandra, holotype, ♂ (ZMA). A–D photos by A. Devyatkin; E–F, by Eulalia Gasso Miracle.

Fig. 2. Adults of Melitaea lutko Evans, 1932 and M. timandra binaluduca subsp. nov. A–H. UPS. I–P. UNS. A, I. M. lutko, ♂, Chitral Gol National Park, Chitral, Pakistan, alt. 2700 m. B, J. M. timandra binaluduca subsp. nov., ♂, Iran, S Khorasan Prov., 35 km N Birjant t. C, K. M. timandra binaluduca subsp. nov., ♂, Central Afghanistan, Bamian Prov., Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m. D, L. M. timandra binaluduca subsp. nov., ♂, Afghanistan, Bamian Prov., Band-e-Amir, alt. 3200 m. E, M. M. lutko, ♀, Chitral Gol National Park, Chitral, Pakistan, alt. 2700 m. F, N. M. timandra binaluduca subsp. nov., ♂, Iran, Mazandaran Prov., S macroslopes of Albors Mts, 80 km SE of Sari, 5 km NE Foulad Mahhaleh v., E slopes of Sultan Kuh Mt., alt. 2000 m. G, O. M. timandra binaluduca subsp. nov., ♀, Afghanistan, Bamian Prov., Band-e-Amir, alt. 3200 m. H, P. M. timandra binaluduca subsp. nov., ♀, Central Afghanistan, Bamian Prov., Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m.

Fig. 3. Male genitalia (a = valva; b = aedeagus). A–G. Melitaea mimetica Higgins, 1940. H–I. M. lutko Evans, 1932. A. Pakistan, Balochistan, Quetta, Urak, alt. 2400–2700 m. B. Pakistan, Balochistan, Quetta, Urak, alt. 2400–2700 m. C. Central Afghanistan, Ghor Prov., 17 km E Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m. D. Central Afghanistan, Ghor Prov., 17 km E of Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m. E. Afghanistan, Bamian Prov., 10 km S of Bamian t., Hushkak v. vicinity, alt. 2700–2800 m. F. Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. G. Central Afghanistan, Ghor Prov., 17 km E of Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m. H. Chitral Gol National Park, Chitral, Pakistan, alt. 2700 m. I. Chitral Gol National Park, Chitral, Pakistan, alt. 2700 m.

Fig. 4. Female genitalia (without the anal papillae, copulatory bursa, and anterior and posterior apophyses). A–D. M. shahvarica sp. nov., Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. E. M. lutko Evans, 1932, Chitral Gol National Park, Chitral, Pakistan, alt. 2700 m. F–I. M. mimetica Higgins, 1940. F. Pakistan, Balochistan, Quetta, Urak, alt. 2400– 2700 m. G–H. Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. I. Central Afghanistan, Ghor Prov., 17 km E of Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m.

Fig. 5. Distribution of M. lutko Evans, 1932. For a description of the dots with letters see Fig. 30.

Fig. 6. Adults of M. mimetica Higgins, 1940. A–H. UPS. I–P. UNS. A–B, I–J. ♂, Pakistan, Balochistan, Quetta, Urak, alt. 2400–2700 m. C, K. ♂, Afghanistan, Ghowr Prov., Bayan Range, 15 km S of Chagcharan, Kindival vall. D, L. ♂, Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. E–F, M–N. ♀, Pakistan, Balochistan, Quetta, Urak, alt. 2400–2700 m. G, O. ♀, Central Afghanistan, Bamian Prov., 8 km S of Bamian, Koh-e-Baba Mts, Dara-e-Khushkak, alt. 2930 m. H, P. ♀, Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. B, J photos by Dr Eckweiler).

Fig. 7. Eggs of Melitaea mimetica Higgins, 1940, Afghanistan, Ghor Prov., 16 km E of Changcharan, Bandi-Ali v. vicinity, alt. 2400 m. A–C. Lateral view. D–F. View from above. G–I. Micropile area.

Fig. 8. Distribution of Melitaea mimetica Higgins, 1940. For a description of the symbols with letters, see Fig. 30. = M. mimetica mimetica Higgins, 1940; = M. mimetica delerei Heidemann, 1954.

Fig. 9. Adults of Melitaea timandra timandra Coutsis & van Oorschot, 2014. A–H. UPS. I–P. UNS. A–D. ♂, Turkmenistan, Sary-Yazy, alt. 300 m. E–H. ♀, Turkmenistan, Sary-Yazy, alt. 300 m. I–L. ♂, Turkmenistan, Sary-Yazy, alt. 300 m. M–P. ♀, Turkmenistan, Sary-Yazy, alt. 300 m.

Fig. 10. Male genitalia (a = valva; b = aedeagus). A–F. Melitaea timandra timandra Coutsis & van Oorschot, 2014. G–O. M. timandra binaludica subsp. nov. A–C. S Turkmenistan, Sary-Yazy, alt. 300 m. D. Paratype, Turkmenistan, Badhyz Reserve, Kepeli. E–F. S Turkmenistan, Sary-Yazy, alt. 300 m. G–L. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m. M. Central Afghanistan, Bamian Prov., Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m. N. Bande-Amir, Hazarajat, Afghanistan. O. Afghanistan, Bamian Prov., Band-e-Amir, alt. 3200 m.

Fig. 11. Female genitalia (without the anal papillae, copulatory bursa, and anterior and posterior apophyses). A–C. Melitaea timandra timandra Coutsis & van Oorschot, 2014, S Turkmenistan, SaryYazy, alt. 300 m. D–I. M. timandra binaludica subsp. nov. D–F. Iran, Rezavi Khorassan Prov., Kuh-eBinalud Mts, Dorrud v. vicinity, alt. 2430 m. G–H. Iran, Mazandaran Prov., S macroslopes of Albors Mts, 80 km SE of Sari, 5 km NE of Foulad Mahhaleh v., E slopes of Sultan Kuh Mt., alt. 2000 m. I. Afghanistan, Bamian Prov., Band-e-Amir, alt. 3200 m.

Fig. 12. Eggs of Melitaea timandra timandra Coutsis & van Oorschot, 2014, S. Turkmenistan, SaryYazy. A–C. Lateral view. D–F. View from above. G–I. Micropile area.

Fig. 13. Biotopes and biology of Melitaea timandra timandraCoutsis & van Oorschot, 2014.A. Bumpycellular overgrown sands on the left bank of the Murgab River, alt. 300 m, April 1993, the habitat of M. timandra timandra. B. Phlomoides regeliana (Aitch. & Hemsl.) Adylov, Kamelin & Makhm. – host plant for M. timandra timandra. The female laying eggs is visible under the leaves of the plant. C. The female of M. timandra timandra is ovipositing on the soil under the host plant leaf. D. Egg clutch of M. timandra timandra on the basal leaf rosette of Ph. regeliana.

Fig. 14. Distribution of Melitaea timandra timandra Coutsis & van Oorschot, 2014, M. timandra binaludica subsp. nov. and M. shahvarica sp. nov. For a description of the points by letters, see Fig. 30. = Melitaea timandra timandra; = Melitaea timandra binaludica subsp. nov.; = Melitaea shahvarica sp. nov.;? = unconfirmed finds of Melitaea timandra; = Melitaea timandra with uncertain subspecies status.

Fig. 15. Adults of Melitaea timandra binaludica subsp. nov. A–H. UPS. I–P. UNS. A, I. Holotype, ♂ (SDM). B–H, J–P. Paratypes (EDMSU). A–D, I–L. ♂, Iran, Rezavi Khorassan Prov., Kuh-e Binalud Mts, Dorrud v. vicinity, alt. 2430 m. E–H, M–P. ♀, same data as for preceding.

Fig. 16. Male genitalia (without aedeagus) in dorsal projection. A–C. Melitaea shahvarica sp. nov. D–H. M. mimetica Higgins, 1940. I. M. lutko Evans, 1932. A–C. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. D. Pakistan, Balochistan, Quetta, Urak, alt. 2400– 2700 m. E. Afghanistan, Bamian Prov., 10 km S of Bamian t., Hushkak v. vicinity, alt, 2700– 2800 m. F. Central Afghanistan, Ghor Prov., 17 km E of Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m. G. Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. H. Afghanistan, Ghor Prov., 16 km E of Changcharan, Bandi-Ali v. vicinity, alt. 2400 m. I. Pakistan, Chitral, Chaghbini CGNP [Chitral Gol National Park], alt. 2700 m.

Fig. 17. Male genitalia (without aedeagus) in dorsal projection. A–C. Melitaea timandra timandra Coutsis & van Oorschot, 2014. D–I. M. timandra binaludica subsp. nov. A–C. S Turkmenistan, SaryYazy, alt. 300 m. D–F. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m. G. Iran, Horossan Prov., 35 km N of Birjant t. H. Afghanistan, Bamian Prov., Band-e-Amir, alt. 3200 m. I. Central Afghanistan, Bamian Prov., Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m.

Fig. 18. Male genitalia and harpe. A–C. Melitaea shahvarica sp. nov. D–E. M. lutko Evans, 1932. F–I. M. mimetica Higgins, 1940. A–C. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. D–E. Pakistan, Chitral, Chaghbini, CGNP [Chitral Gol National Park], alt. 2700 m. F–G. Afghanistan, Bamian Prov., Punjub Distr., 10 km. NE Varas v., alt. 2400 m. H–I. Afghanistan, Bamian Prov., Panjub Distr., 10 km. NE Varas vil., alt. 2400 m.

Fig. 19. Male genitalia and harpe. A–C. Melitaea timandra timandra Coutsis & van Oorschot, 2014. D–I. M. timandra binaludica subsp. nov. A–C. Turkmenistan, Sary-Yazy, alt. 300 m. D–F. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m. G. Afghanistan, Bamian Prov., Band-e-Amir, alt. 3200 m. H. Afghanistan, Bamian Prov., Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m. I. Afghanistan, Band-e-Amir, Hazarajat.

Fig. 20. Eggs of Melitaea timandra binaludica subsp. nov., Iran, Kuh-e-Binalud Mts. A–C. Lateral view. D–F. View from above. G–I. Micropile area.

Fig. 21. Adults of Melitaea shahvarica sp. nov. A–H. UPS. I–P. UNS. A, I. Holotype, ♂ (SDM). B–H, J–P. Paratypes (EDMSU). A–C. ♂. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mt., alt. 2200–2400 m. D. ♂. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mt., Tohar v. vicinity, alt. 2200 m. E–H. ♀. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mt., alt. 2200– 2400 m. I–K. ♂. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mt., 2200–2400 m. L. ♂. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, Tohar v. vicinity, alt. 2200 m. M–P. ♀. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mt., alt. 2200–2400 m.

Fig. 22. Male genitalia of Melitaea shahvarica sp. nov. (a = valva; b = aedeagus). Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m.

Fig. 23. Eggs of Melitaea shahvarica sp. nov., Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. A–C. Lateral view. D–F. View from above. G–I. Micropile area.

Fig. 24. First instar caterpillar of Melitaea shahvarica sp. nov. A. Head, bottom view. B. Head, front view. C–D. Head, lateral view. E–F. Caterpillar, lateral view.

Fig. 25. I–IV instar caterpillars of Melitaea shahvarica sp. nov. (a = view from above; b = lateral view). A. First instar caterpillar after hatching. B. First instar caterpillar before molting. C. Second instar caterpillar. D. Third instar caterpillar. E. Fourth instar caterpillar.

Fig. 26. V–VI instar caterpillars of Melitaea shahvarica sp. nov. (a = view from above; b = lateral view). A. Fifth instar caterpillar. B. Sixth instar caterpillar. C. Sixth instar caterpillar, head capsule, front view. D. Sixth instar caterpillar, head capsule, lateral view.

Fig. 27.Biotope, host plant, and adult of Melitaea shahvaricasp. nov. in nature. A. Shahvar Mt., E Elburs Ridge. B. Biotope with Phlomoides molucelloides (Bunge) Salmaki. C. Host plant Ph. molucelloides. D. Female on Ph. molucelloides.

Fig. 28.Eggs and caterpillars of Melitaea shahvarica sp. nov. in nature and in the laboratory.A–B. Freshly laid eggs under a leaf of a host plant, May 2018, Iran, Shahvar Mt., alt. 2200 m. C–D. IV–V instar caterpillars on the leaves of the host plant Phlomoides molucelloides (Bunge) Salmaki, July 2019, Iran, Shahvar Mt., alt. 2500 m. E. I instar caterpillars in the laboratory, Moscow, May 2018. F. VI instar caterpillars during diapause, Moscow, October 2018.

Fig.29.Differencesinthestructureofvalvaandaedeagusofthelutko speciesgroup.A.Melitaeashahvarica sp. nov. B, D, H. M. timandra binaludica subsp. nov. C–E. M. mimetica Higgins, 1940. F. M. lutko Evans, 1932. G. M. timandra timandra Coutsis &van Oorschot, 2014. A. Iran, Semnan Prov., Shahrud area, S macroslope of Shahvar Mts, alt. 2200–2400 m. B. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m. C. Afghanistan, Bamian Prov., Punjub Distr., 10 km NE of Varas v., alt. 2400 m. D. Afghanistan, Band-i-Amir, Hazarajat. E. Pakistan, Balochistan, Quetta, Urak, alt. 2400– 2700 m. F. Pakistan, Chitral, Gol National Park, alt. 2700 m. G. Turkmenistan, Sary-Yazy, alt. 700 m. H. Iran, Rezavi Khorassan Prov., Kuh-e-Binalud Mts, Dorrud v. vicinity, alt. 2430 m.

Fig. 30 (continued on next page). Distribution of the lutko species group fritillaries in combination with the distribution of known and suspected species of host plants from the genus Phlomoides Blume. = Melitaea lutko Evans, 1932; = M. timandra timandra Coutsis & Oorschot, 2014; = M. timandra binaludica subsp. nov.; = M. shahvarica sp. nov.; = M. mimetica mimetica Higgins, 1940; = M. mimetica delerei Heidemann, 1954;? = unconfirmed findings of M. timandra; = M. timandra with an unclear subspecies status; = Phlomoides regeliana (Aitch. & Hemsl.) Adylov, Kamelin & Makhm.; = Phlomoides boissieriana (Regel) Adylov, Kamelin & Makhm.; = Phlomoides laciniata (L.) Kamelin & Makhm.; = Phlomoides labiosiformis (Popov) Adylov, Kamelin & Makhm.; = Phlomoides loasifolia (Benth.) Kamelin & Makhm.; = Phlomoides molucelloides (Bunge) Salmaki; = Phlomoides acaulis(Beck ex Rech.f.) Salmaki; = Phlomoides labiosa(Bunge) Adylov, Kamelin & Makhm. A. Pakistan, Chitral, Chaghbini CGNP, alt. 2700–3000 m. B. Pakistan, Khyber Pakhtunkhwa, Drosh. C. Pakistan, Khyber Pakhtunkhwa, Keon Nullah. D. Pakistan, Khyber Pakhtunkhwa, Malakand. E. Pakistan, Khyber Pakhtunkhwa, Birmoglasht. F. Turkmenistan, Badkhyz, Kepeli, alt. 700 m. G. Turkmenistan, Badkhyz, Kyzyl-Jar, alt. 700 m. H. Turkmenistan, Kushka, alt. 700 m. I. Turkmenistan, Murgab river, Sary-Yazy, alt. 300 m. J. Turkmenistan, 30 km E of BairamAli, Zahmet, alt. 240 m. K. Turkmenistan, Bairam-Ali, alt. 230 m. L. Turkmenistan, Kara-Kum desert, 30 km W of Mary, alt. 200 m.M. Turkmenistan, Dushak, alt. 250 m.N. Turkmenistan, Chaacha, alt. 400 m. O. Turkmenistan, Bakharden, alt. 200 m. P. Iran, Khorossan Razavi, Kuh-e-Binalud Mts, Qadamgah area, Gerina, alt. 2000 m. Q. Iran, Khorasan Razavi, Kuh-e-Binalud Mts, 15 km SW of Zoshk, alt. 2300– 2500 m. R. Iran, S Khorosan, 75 km N of Birjant, Sedeh, alt. 1500 m. S. Iran, S Khorosan, 35 km N of Birjant, alt. 1500 m. T. Afghanistan, Bamian, Band-e-Amir, Dzhudoi-Kvak Gorge, alt. 3200 m. U. Afghanistan, Bamian, Band-e-Amir, Hazarajat, alt. 3000–3200 m. V. Afghanistan, Bamian, Koh-iBaba Mts, Joshanak, alt. 2800 m. W. Afghanistan, Heart, Qala-i-Naw, Kashka pass. X. Iran, Semnan, Shahvar Mt., alt. 2200–2500 m. Y. Turkmenistan, Kara-Kala, Monjukly Ridge, 300–700 m. Z. Iran, Golestan, E Maraveh Tappeh, N Ghazan Ghayeh, Palizan Mts. A”. Pakistan, Balochistan, Quetta, Urak, alt. 2500 m. B”. Pakistan, Balochistan, Ziarat, alt. 2500 m. C”. Pakistan, Balochistan, Khojak, alt. 1700 m. D”. Pakistan, Balochistan, Zaghum, alt. 1600 m; E”. Pakistan, Punjab, Gawar, alt. 500 m. F”. Pakistan, Balochistan, Sheik Wazil, alt. 1600 m. G”. Afghanistan, Bamian, Hushkak, alt. 2700– 2800 m. H”. Afghanistan, Bamian, Punjub Distr., 10 km NE of Varas, alt. 2400 m. I”. Afghanistan, Ghor, 17 km E of Changcharan, 15 km S of Bandi-Ali, Gazak Mts, alt. 2400 m. J”. Afghanistan, Ghor, Bayan Range, 15 km S of Changcharan, Kindival valley, alt. 2700 m. K”. Afghanistan, Bamiyan, Kohi-Baba Mts, Panjao, alt. 3000 m. Afghanistan, Bamiyan, Koh-i-Baba Mts, Shah-tu-Kotal, alt. 4000 m. L”. Afghanistan, Kapisa, Pandshir valley, alt. 2200–2800 m. M”. Afghanistan, Kabul. N”. Iran, Tehran, Elburz Ridge, Demavend Mt., Ask, alt. 1800 m. O”. Iran, Semnan, Foulad Mohaleh, alt. 2200 m. P”. Pakistan, Punjab, Murree.

Kingdom	Animalia
Phylum	Arthropoda
Class	Insecta
Order	Lepidoptera
Family	Nymphalidae
SubFamily	Nymphalinae

Melitaea Fabricius, 1807

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