Butterfly diversity of Lička Plješevica Mountain chain, Croatia (Lepidoptera: Papilionoidea)

Diversidad de las mariposas de la cadena montañosa de Lička Plješevica, Croacia (Lepidoptera: Papilionoidea)

Introduction

Lička Plješevica massif is an elongated mountain chain that extends more than 100 kilometres in the northwest-southeast direction from Medved¯ak and Velika Plješevica in the north-eastern part of Croatian Lika region, through Mala Plješevica, Ozeblin peak, and complex of several lower mountains like Kremen, Mazinske planine and Urljaj in the south-eastern part of the chain. Isolated peaks Lisac east of Gračac and Mt. Poštak northwest of Knin are also considered as part of the massif (HORVAT, 1925; POLJAK, 1974). The whole range runs in parallel but further inland to Velebit Mts. while the southern part including Mt. Poštak is inserted between Velebit and Dinara Mountain chains. The northwestern border, right side of Upper Korana Gorge, Medved¯ak Hill, Donja Jezera, right bank of Kozjak Lake and Prijeboj are situated in National Park Plitvice Lakes. The western slopes of Lička Plješevica massif rise from the several karst poljes in Lika region: Koreničko polje, Bjelopolje, Krbavsko polje and Gračačko polje, while its eastern slopes descend into the gorges of upper Una River, Srebrenica and Butiznica which represent the natural boundary between Croatia and Bosnia and Herzegovina. Within Lička Plješivica range there are several small valleys and two karst poljes, the Lapačko polje and Gubavčevo polje. The highest peaks of the mountain chain are located in the northern part of the massif, Ozeblin (1657 m a.s.l.) and Gola Plješivica (1647 m a.s.l.) where the subalpine altitudinal belt with dwarf pine (Pinus mugo Turro) and alpine meadows is developed near the peaks above 1600 m a.s.l. (HORVAT, 1925). About 85 % of northern part of Mt. Lička Plješevica is covered with dense forests mostly of Euro-Siberian forest belt, with only small remnants of alpine and montane grasslands and pastures, usually in the upper parts of the mountains (POLJAK, 1998). Open dry calcareous grasslands and pastures in different stages of abandonment characterised by the presence of some typical sub-Mediterranean plants are still abundant in the central and southern part of the massif at lower elevations (TRINAJSTIĆ & ŠUGAR, 1968; ŠUGAR & PLAZIBAT, 1988).

The entire region has a very complex geological structure composed of diverse rocks, mostly of carbonate and dolomite originating from the Mesozoic period (the main ridge of GolaPlješevica is from Cretaceous carbonate and dolomite), but with incorporated mosaic of other types of rocks like Triassic siltstones and micaceous sandstones between Glogovo and Lisac hill (ANONYMUS, 1978: Geological map SFRJ 1:500000). The Lička Plješevica massif is located in continental region with strong influence of Mediterranean climate which penetrates deep into continent along Butižnica Valley and Una River Gorge, and along karst poljes west of the mountain chain (TRINAJSTIĆ & ŠUGAR, 1968). The average temperature in January is -4,5ºC and in July between 12ºC and 14ºC (BOGNAR et al., 1975).

The historical data about the butterfly fauna of Lička Plješevica chain are relatively few and scattered across a wide time frame. First record was published by Gjuro Koča (KOČA, 1901) who reported Erebia ligea (Linnaeus, 1758) in 1891, and the German malacologist Rudolf Sturany who collected Spialia sertorius (Hoffmansegg, 1804) (possibly S. orbifer (Hübner, [1823])), Vanessa atalanta (Linnaeus, 1758), Euphydras aurinia (Rottemburg, 1775), Melitaea aurelia Nickerl, 1850, M. didyma (Esper, 1778), Boloria euphrosyne (Linnaeus, 1758), and Cupido minimus (Fuessly, 1775) in the foothills of Velika Lička Plješevica near Plitvice Lakes (REBEL, 1895). Additionally, Hochetlinger noted Parnasius mnemosyne (Linnaeus, 1758) as “common species around touristic house below Pljesevica peak” (Gola Plješevica), together with records of three rare species in the region Apatura ilia ([Denis & Schiffermüller], 1775), Limenitis populi (Linnaeus, 1758), and probably regionally extinct Nymphalis vaualbum ([Denis & Schiffermüller], 1775) which he also observed near Plitvice Lakes (FRANIĆ, 1910).

In last week of July to 2^nd August 1922, students from Zagreb University Zdravko Lorkovič and Zvonimir Badovinac collected butterflies from Zrmanja River spring (LORKOVIĆ, 1974), near Dabašnica (LORKOVIĆ, 1989), on Ozeblin peak, in vicinity of forest hut near Karlović Korita, from alpine meadow Ruda Poljana on Mala Plješivica Mountain (MLADINOV & LORKOVIĆ 1979), and near Plitvice Lakes. Based on this collection STEINER (1938) published unexpected finding of a new butterfly species for the fauna of Croatia Boloria titania (Esper, 1793) from “Lička Plješivica”. The record was later confirmed by MLADINOV & LORKOVIĆ (1985) based on observations of Badovinac from Ruda Poljana, 1616 m a.s.l. Another 43 species observed in the area were noted in Lorković’s manuscript from 1954, which was published only recently after he passed away (LORKOVIĆ, 2009). Lorković also added his observation of Hipparchia statilinus (Hufnagel, 1766) from Zrmanja River spring (LORKOVIĆ, 1974), first record of Pieris balcana Lorković, 1969 near Dabašnica and after chromosome analysis also confirmed the contact zone between P. napi (Linnaeus, 1758) and P. balcana Lorković, 1969 at Plitvice Lakes (LORKOVIĆ, 1970, 1989). MLADINOV (1973) added records for eight additional species originating from the area, stored in the collection of Natural History Museum in Zagreb. MLADINOV & LORKOVIĆ (1979) additionally included a montane species Erebia oeme (Hübner, [1804]).

Recently, KOREN (2010) published first findings of Polyommatus ripartii (Freyer, 1830), and KOREN et al. (2010) the observations of Proterebia phegea (Borkhausen, 1788) both from the Zrmanja spring area. KOREN et al. (2011) and KOREN et al. (2015) reviewed rich butterfly fauna of Zrmanja River source and Mt. Poštak (92 and 108 recorded species respectively). KOREN & ŠTIH (2013) noted Melitaea ornata Christoph, 1893 from Zrmanja spring area, and TVRTKOVIĆ et al. (2015) added some interesting species records from Lisac peak, Kremen Mt. and Una Gorge including Euphydrias maturna (Linnaeus, 1758). Taxonomical status and occurrence of species of the Polyommatus subgenus Agrodiaetus in this region was also recently discussed (KOREN & LAUŠ, 2015; LOVRENČIĆ et al., 2016).

The goal of this contribution is to present the overview of the butterfly diversity of the Lička Plješevica massif based on the recent faunistic surveys of the authors (2007 - 2019), with explanatory notes for some rare and interesting species recorded in the area. A comparison with neighbouring Velebit and Dinara Mountain chains is discussed in terms of diversity and zoogeography. We also discuss the main threats that could on a long run cause the decline of the high butterfly diversity in the region.

Materials and methods

Field surveys specifically addressing the butterfly fauna of the region were carried out mostly during the last five years, from 2014 onwards. Topography and habitat diversity were used to select the best suitable localities in the region. A total of 71 localities were visited during the surveys, some of them with butterfly rich habitats were visited on several occasions (Figure 1). The spatial processing and visualisation of data was made in the program ARC GIS desktop. Butterflies were identified using standard field guides (LAFRANCHIS, 2004; TOLMAN & LEWINGTON, 2008). Additionally, specimens of the genera Leptidea, Melitaea and Plebejus were collected and their genitalia were examined for correct identification. Revision of collected specimens from Pieris napi / balcana aggregate were examined with help of wing-markings (LORKOVIĆ, 1970). The nomenclature follows WIEMERS et al. (2018). The biogeographical affiliation adheres to KUDRNA et al. (2016).

List of localities

The list of localities contains the relevant toponyms, a short description of the habitat, altitude, coordinates, dates of the visits and observers. Localities are mostly arranged in geographical order from northwest towards southeast (Fig. 1).

1. 1. Prijeboj, meadows toward Ličko Petrovo Selo, N: 44,846625, E: 15,682131, 661 m, 2-VIII-2015, BL, MZ
2. 2. Medvedak Hill, N of Prijeboj settlement, spring Dražica, forest edge, grasslands, N: 44,856174, E: 15,691113, 672 m, 28-VI-2015, TK
3. 3. Velika Plješevica, N of Korenica, meadows between Krgin vršak and Jukanova draga, bushy meadowsurrounded by forest, N: 44,801503, E: 15,694054, 773 m, 25-VII-2015, BL, MZ
4. 4. Velika Plješevica, N of Korenica, Stare paljevine, forest edge, N: 44,812725, E: 15,709914, 982 m, 25-VII-2015,BL, MZ
5. 5. Velika Plješevica, N of Korenica, between Jaraga and Torovi, forest edge, small patches of grasslands, N:44,815631, E: 15,722884, 1015 m, 25-VII-2015, BL, MZ, 21-V-2016, TK, MZ
6. 6. Udbina, Kurjak, small hill with a transmitter near the turn for the village, dry grasslands, abandoned quarry, N:44,486958, E: 15,732503, 775 m, 13-VIII-2016, RV, 6-VIII-2019, RV
7. 7. Velika Plješevica, Osinjak, toward the peak Gola Plješevica, forest edge, N: 44,799754, E: 15,738640, 1365 m,1-VIII-2007, NT, MV; 25-VII-2015, BL, MZ
8. 8. Velika Plješevica, Gola Plješevica, grasslands and forest edge near the peak, N: 44,791911, E: 15,745421, 1455m, 1-VIII-2007, NT, MV, 25-VII-2015, BL, MZ, 21-V-2016, TK, MZ
9. 9. Mala Plješevica, Ponor Korenički, banks of Matica River, grasslands, forest edge, N: 44,728688, E: 15,763533,712 m, 29-VII-2015, TK
10. 10. Ozeblin, Kozja Draga, slopes at the entrance to the valley NE of Danilovac hamlet, owergrown grasslands, forestedge, N: 44,546395, E: 15,810098, 818 m, 21-V-2016, TK, MZ
11. 11. Ozeblin, Frkasić, surroundings of pond Jezero, forest edge and grasslands, N: 44,676879, E: 15,815090, 840 m,29-VI-2015, TK
12. 12. Mt. Kremen: Ondić settlement, slopes above peak Rteševo, grasslands, forest edge, N: 44,483984, E: 15,821597,1359 m, 23-VII-2014, BL
13. 13. Lička Plješevica, area between Ondic and Rudopolje, overgrown dry grasslands, N: 44,437113, E: 15,825760, 735 m, 25-VI-2014, BL
14. 14. Mala Plješevica, E of Frkašič, Škorina poljana, dry grasslands, N: 44,701803, E: 15,826848, 1204 m, 24-VII-2015, BL, MZ
15. 15. Udbina, middle part of the Kozja draga Valley, rocky meadows along the road, N: 44,558682, E: 15,828372, 884 m, 2-VIII-2014, BL, 1-VI-2015, TK, 29-VII-2015, TK, 21-V-2016, TK, MZ, 13-VIII-2016, RV, 6-VIII-2019, RV
16. 16. Mala Plješevica, Frkašić, Ďevina uvala, meadows, N: 44,699963, E: 15,832812, 1168 m, 24-VII-2015, BL, MZ, 3-VII-2018, TK, BL, MZ
17. 17. Lička Plješevica, NE of Vugja poljana, grasslands at the forest edge, N: 44,585483, E: 15,839517, 1292 m, 2-VIII-2014, BL
18. 18. Mala Plješevica, road Korenica - Doni Lapac, turning toward Bijeli potoci village, Fagus forest, N: 44,670477,E: 15,842894, 1067 m, 24-VII-2015, BL, MZ
19. 19. Mala Plješevica, Bijeli potoci, Poljane meadows, forest edge, N: 44,679578, E: 15,845102, 1294 m, 3-VI-2015, NT, LL, 29-VI-2015, TK, 7-VII-2015, NT, 16-IX-2015, NT, 21-V-2016, TK, MZ, 2-VII-2018, TK, 12-VII-2018,TK
20. 20. Lička Plješevica, SE of Frkašić, meadows Unjka, dry grasslands, forest edge, N: 44,654388, E: 15,845329, 1129m, 23-VII-2015, BL, MZ
21. 21. Lička Plješevica, W of Ruda Polja, forest path with flowering patches, N: 44,691139, E: 15,848, 1294 m, 3-VII-2018, TK, BL, MZ
22. 22. Lička Plješevica, E of peak Trla, forest glade, N: 44,581353, E: 15,851600, 1291 m, 2-VIII-2014, BL
23. 23. Mala Plješevica, SE of Frkašić, old quarry NW of hill Šišinov tavan, forest clearing, N: 44,660843, E:15,855764, 1165 m, 23-VII-2015, BL, MZ
24. 24. Rudopolje Bruvanjsko, area of Gornja Klapavica, dry grassland, N: 44,439344, E: 15,858142, 746 m, 26-VII-2014, BL
25. 25. Lička Plješevica, Trlska draga, partially overgrown forest edge, N: 44,572187, E: 15,858182, 1206 m, 2-VIII-2014, NT
26. 26. Lička Plješevica, E of Frkašić, Ruda Polja meadows, owergrown grasslands, forest edge, N: 44,687917, E:15,862389, 1294 m, 3-VII-2018, TK, BL, MZ
27. 27. Lička Plješevica, Donji Lapac, hill Pogledalo, meadow Veliko Kamensko, grasslands, forest edge, N: 44,638324, E: 15,866548, 1051 m, 23-VII-2015, BL, MZ
28. 28. Lička Plješevica, Debeli Vrh, grasslands, N: 44,683239, E: 15,868475, 1294 m, 3-VII-2018, TK, BL, MZ
29. 29. Mt. Kremen, meadow Sarajevo, overgrown grasslands, N: 44,494742, E: 15,875780, 982 m, 22-VII-2014, BL
30. 30. Lika, Plješevica, Ozeblin, Trl peak, small forest glade, forest edge, N: 44,578888, E: 15,880833, 1536 m, 22-V-2016, TK
31. 31. Mt. Kremen, Meďugorje, Magarčev do, karstic grasslands, N: 44,513609, E: 15,888295, 1072 m, 4-VI-2015, NT, LL, 8-VII-2015, NT, LL, 23-IX-2015, NT, LL
32. 32. Mt. Kremen, large abandoned meadow S of the main ridge, overgrown grasslands, N: 44,459733, E: 15,888683, 1330 m, 1-VII-2015, RV
33. 33. Mt. Kremen, along the ridge above the tree line, montane grasslands, rocky slopes, N: 44,463333, E: 15,889583, 1500 m, 1-VII-2015, RV
34. 34. Lička Plješevica, Donji Lapac, meadow Malo Kamensko, grasslands, N: 44,617204, E: 15,896684, 1007 m, 1-VIII-2014, BL, 18-V-2017, MZ, 19-V-2017, MZ, 29-VI-2017, MZ
35. 35. Gračac, Mt. Kremen, near the road SE of the peak, dry grassland, forest edge, N: 44,460183, E: 15,8989, 1160 m, 1-VII-2015, RV, 14-VII-2015, NT
36. 36. Donji Lapac, Dnopolje, grasslands around a pond, N: 44,588682, E: 15,921794, 777 m, 29-VI-2015, TK
37. 37. Mala Plješevica, Kalinovača, cca 7km N/NW of Donji Lapac, overgrown grassland, N: 44,612924, E:15,935532, 758 m, 30-VII-2014, BL
38. 38. Gubavčevo polje, NE part, dry grassland, N: 44,349671, E: 15,944296, 757 m, 27-VI-2014, BL
39. 39. Lička Plješevica, Glogovo, Cvijetkovici, overgrown grasslands, N: 44,310367, E: 15,944637, 845 m, 19-V-2015,NT
40. 40. Mt. Lisac, in the valley NE of the village towards Surle, partially overgrown meadows, N: 44,320513, E:15,967480, 870 m, 26-VI-2014, BL, 26-VI-2017, RV
41. 41. Mala Plješevica, meadow Lažipolje, cca 4km N/NE of Donji Lapac, dry grassland, forest edge, N: 44,588987, E:15,971300, 748 m, 31-VII-2014, BL
42. 42. Mt. Lisac, middle and lower part of Borovačka jaruga, dry grasslands, N: 44,318739, E: 15,973722, 920 m, 26-VII-2017, RV
43. 43. Mt. Lisac, ridge to the NW and above Borovačka jaruga, rocky slopes with sparse vegetation, N: 44,324497, E:15,981747, 1070 m, 18-IX-2015, NT, 26-VI-2017, RV
44. 44. Vršina peak between Mazin and Brezovac Dobroselski, grassland surrounded by forest, N: 44,447506, E:15,981851, 1068 m, 12-VII-2013, NT
45. 45. Mt. Lisac, road Surle - Strmica and the trail to Lisac between 860 - 1160 m, abandoned grasslands, mixed woods, N: 44,334173, E: 15,988206, 1019 m, 27-VI-2015, 17-IX-2015, NT
46. 46. Mt. Lisac, plateau below Lisac peak, western slopes between 1200 - 1340 m, dry grasslands, N: 44,330084, E:15,990722, 1206 m, 26-VI-2014, BL, 29-VII-2015, NT, LL, 28-VII-2017, BL, 29-VII-2017, BL
47. 47. Mt. Lisac, near the peak and southern slopes, dry rocky grasslands, tall grass meadow, N: 44,333575, E:15,994503, 1300 m, 26-VI-2017, RV, 28-30-VII-2017, BL
48. 48. Ozeblin, Gornji Lapac, settlement Gajine, dry grasslands with bushes and trees, N: 44,526731, E: 16,000374,597 m, 28-VII-2014, BL
49. 49. Mazin, SE of the village, Bukovačka Draga, forest path, N: 44,435060, E: 16,008641, 1146 m, 12-VII-2013, NT
50. 50. Mt. Kremen, Gornji Lapac, Poljana, meadow surrounded by forest, N: 44,482173, E: 16,011506, 1032 m, 29-VII-2014, BL
51. 51. Brezovac Dobroselski, SW of the village, toponym Klanci, forest edge and forest glades, N: 44,442276, E:16,028719, 996 m, 12-VII-2013, NT
52. 52. Gornja Suvaja, Čemernica meadows west of the settlement, N: 44,402098, E: 16,035524, 1094 m, 23-VI-2014,BL
53. 53. Veliko Popinsko polje, W of Lukici and toward Srpski Klanac, partially overgrown pastures, N: 44,282385, E:16,043992, 668 m, 20-VII-2016, 26-VII-2016, 7-VII-2017, NT
54. 54. Gračac, Zrmanja, at a large spring at the N edge of the village, forest edge, abandoned dry grassland, N:44,196406, E: 16,056494, 297 m, 8-VIII-2014, RV, 5-VIII-2015, RV, 13-VII-2018, RV, 6-VIII-2019, RV
55. 55. Gračac, Zrmanja, pastures on the E side of the river E of village Zrmanja Vrelo, N: 44,194686, E: 16,059772,380 m, 9-VII-2011, RV
56. 56. Gornji Srb, meadows Mala Čemernica, 5km NW of the settlement, forest edge, grasslands, N: 44,374213, E:16,064198, 1124 m, 24-VI-2014, BL
57. 57. Gornja Suvaja, surroundings of Čopići settlement, pastures, ruderal habitats, N: 44,406314, E: 16,071493, 571m, 23-VI-2014, BL
58. 58. Gračac, Zrmanja, valley at the spring of the river NE of village Zrmanja Vrelo, woods along the stream, rockyslopes with sparse vegetation, N: 44,203889, E: 16,074061, 303 m, 9-VII-2011, RV, 11-VIII-2011, RV, 8-VIII-2014, RV, 5-VIII-2015, RV, 20-VII-2016, NT, 26-VI-2017, RV
59. 59. Mt. Poštak, along the road NE fo Otrić, plateau at Kamenita glavica, abandoned dry grasslands, N: 44,261569,E: 16,089089, 980 m, 22-VI-2014, RV, 8-VIII-2014, RV, 1-VII-2015, RV, 5-VIII-2015, RV, 26-VI-2017, RV, 13-VII-2018, RV
60. 60. Kupirovo, between Srb and Srpski klanac, dry grasslands, N: 44,301546, E: 16,089298, 789 m, 6-VI-2015, NT
61. 61. Mt. Poštak, grasslands near small quarry at Ljubina Poljana, N: 44,261226, E: 16,102745, 1094 m, 7-VI-2014,NT, 22-VI-2014, RV, 1-VII-2015, RV, 5-VIII-2015, RV, 8-12-VII-2017, BL, 13-VII-2018, RV
62. 62. Mt. Postak, pastures near the top of the mountain and on SW slopes, N: 44,252803, E: 16,105508, 1250 - 1420m, 22-VI-2014, RV, 1-VII-2015, RV, 11-VII-2017, BL
63. 63. Mt. Poštak, meadow Ljubina poljana, northern slopes, dry grasslands, N: 44,262974, E: 16,106546, 1067 m, 21-VI-2014, BL
64. 64. Mt. Poštak, Med¯ugorje, N of Kirin vrh, near road to Jelovi Tavani, forest track, mixed woods, N: 44,287901, E:16,109056, 1066 m, 7-VI-2014, 23-IX-2014, NT
65. 65. Mt. Poštak, E slopes, dry rocky grasslands, N: 44,255135, E: 16,113831, 1374 m, 9-11-VII-2017, BL
66. 66. Srb, Donja Suvaja, along forest track E of Zalužje village, N: 44,417961, E: 16,121569, 390 m, 22-VI-2014, RV
67. 67. Kunovac Kupirovački, at Velika bara, wooded stream banks, dry grasslands, N: 44,345097, E: 16,135991, 446m, 22-VI-2014, BL
68. 68. Srb, Kunovac Kupirovački, along forest road in a valley SE of the village, N: 44,321044, E: 16,136575, 530 m,22-VI-2014, RV
69. 69. Srb, Donja Suvaja, along a track in the gorge of Una river E of Zalužje village, dry grasslands, rocky slopes, N:44,418592, E: 16,137892, 380 m, 22-VI-2014, RV
70. 70. Mt. Poštak, NE slopes of hill Visočica, meadows Palež, forest edge, N: 44,267572, E: 16,148236, 933 m, 21-VI-2014, BL, 31-VII-2014, BL
71. 71. Gornji Srb, Zavlaka village, meadows and forest edge, N: 44,315826, E: 16,176171, 749 m, 22-VI-2014, BL

Results and discussion

In total, 142 species were recorded during our surveys of Mt.Lička Plješevica range, including 12 new for this region (Heteropterus morpheus (Pallas, 1771), Colias caucasica Staudinger, 1871, Favonius quercus (Linnaeus, 1758), Leptotes pirithous (Linnaeus, 1767), Plebejus argyrognomon (Bergsträsser, 1779), Thecla betulae (Linnaeus, 1758), Araschnia levana (Linnaeus, 1758), Boloria selene ([Denis & Schiffermüller], 1775), Erebia aethiops (Esper, 1777), Melitaea britomartis Assmann, 1847, Melitaea ornata Christoph, 1893, and Minois dryas (Scopoli, 1763). In the previous surveys in the southern part of the massif, in particularly Mt.Poštak and the source of Zrmanja River (KOREN et al., 2015), three additional species not recorded during our survey were found: Polyommatus escheri (Hübner, [1823]), Proterebia phegea (Borkhausen, 1788), and Melanargia larissa (Geyer, [1828]). Additional species mentioned for Mt. Plješevica but not confirmed during recent surveys are Euphydryas maturna (Linnaeus, 1758) found in Una River canyon (TVRTKOVIĆ et al., 2015), Boloria titania (Esper, 1793) recorded almost 100 years ago (STEINER, 1938) and Nymphalis vaualbum ([Denis & Schiffermüller], 1775) recorded by Hochetlinger in 1909 (FRANIĆ, 1910) and Lorković in 1923 (LORKOVIĆ, 2009). With these records, the butterfly fauna of Lička Plješevica range comprises 152 species. This positions the Lička Plješevica range as the second butterfly richest mountain chain in Croatia, just after Velebit Mts., for which a total of 153 species were recorded (TVRTKOVIĆ et al., 2015).

The comparison of biogeographical affiliation between Mts. Lička Plješevica, Velebit and Dinara revealed a similar number of species per affiliation type (Table III). The most noticeable difference is in the smaller number of montane (MON) species on Lička Plješevica (5) in comparison with Velebit (10) and Dinara (7). This is likely a result of general lower altitude of Lička Plješivica and very small and isolated subalpine habitats limited to a few highest peaks in the range. Entire Lička Plješevica massif is also shorter than Velebit Mountain chain. Throughout its length, it stretches in the northwest-southeast direction - the position supporting microrefugia’s and north south migrations of more northern continental species during climatic fluctuations.

Among the species listed, several were recorded at 30 or more sites and could be considered common and widespread in the Lička Plješevica range: Ochlodes sylvanus (Esper, 1777), Papilio machaon Linnaeus, 1758, Colias crocea (Geoffroy, 1785), Gonepteryx rhamni (Linnaeus, 1758), Aporia crataegi (Linnaeus, 1758), Polyommatus icarus (Rottemburg, 1775), Vanessa cardui (Linnaeus, 1758), Maniola jurtina (Linnaeus, 1758), Melanargia galathea (Linnaeus, 1758), and Coenonympha arcania (Linnaeus, 1760).

On the other hand, several species were observed only at a single location and are possibly extremely rare in the region: Heteropterus morpheus (Pallas, 1771), Colias caucasica Staudinger, 1871, Favonius quercus (Linnaeus, 1758), Iolana iolas (Ochsenheimer, 1816), Boloria selene ([Denis & Schiffermüller], 1775), Aphantopus hyperantus (Linnaeus, 1758), Hyponephele lycaon (Kühn, 1774), Hyponephele lupinus (Costa, 1836), Limenitis populi (Linnaeus, 1758), Melitaea ornata Christoph, 1893, and Neptis rivularis (Scopoli, 1763).

According to the Red Data List of Croatian Butterflies (ŠAŠIĆ et al., 2015), four of the recorded species are considered vulnerable (VU): Phengaris alcon f. rebeli (Hirschke, 1904), Phengaris arion (Linnaeus, 1758), Polyommatus damon ([Denis & Schiffermüller], 1775), and Parnassius apollo (Linnaeus, 1758). Additionally, 11 species among those observed are considered near threatened (NT): Heteropterus morpheus (Pallas, 1771), Papilio machaon Linnaeus, 1758, Zerynthia polyxena ([Denis & Schiffermüller], 1775), Parnassius mnemosyne (Linnaeus, 1758), Glaucopsyche alexis (Poda, 1761), Polyommatus thersites (Cantener, 1835), Pseudophilotes vicrama (Moore, 1865), Scolitantides orion (Pallas, 1771), Apatura ilia ([Denis & Schiffermüller], 1775), A. iris (Linnaeus, 1758), Euphydryas maturna (Linnaeus, 1758), Euphydras aurinia (Rottemburg, 1775), and Erebia medusa ([Denis & Schiffermüller], 1775). Four species Thymelicus action (Rottemburg, 1775), Pieris brassicae (Linnaeus, 1758), Melitaea aurelia Nickerl, 1850, and M. britomartis Assmann, 1847 are considered as data deficient (DD).

The highest diversity with more than 50 species per locality was recorded in the southernmost part of the range near Zrmanja Vrelo (KOREN et al., 2011: 86 species), in the Ozeblin Mt. - Kozja Draga (76 species), Mala Plješevica Mt.: Bijeli potoci, Poljane (68), Mt. Poštak: plateau at Kamenita glavica (66) and grasslands at Ljubina Poljana (56).

Faunistically all compared mountain ranges are similar in composition. For example, Mt. Dinara when compared to Lička Plješevica has only four additional species, all limited to highest elevations: Polyommatus eros (Ochsenheimer, 1808), Erebia euryale (Esper, 1805), Erebia ottomana Herrich-Schäffer, 1847, and Erebia triarius (Prunner, 1798) (TVRTKOVIĆ et al., 2012; KOREN & LAUŠ 2013). The Velebit Mts. however, host 18 additional species so far not recorded on Mt. Lička Plješevica (MIHOCI et al., 2007; TVRTKOVIĆ et al., 2015). Among these, Gegenes pumilio (Hoffmansegg, 1804), Cacyreus marshalli Butler, 1898, Lampides boeticus (Linnaeus, 1767), Tarucus balkanica (Freyer, 1844), Charaxes jasius (Linnaeus, 1767), Polygonia egea (Cramer, 1775), and Gonepteryx cleopatra (Linnaeus, 1767) are Mediterranean species, mostly confined to the coastal areas at lower elevations and the foothills of Velbit Mts., and are not to be expected so far inland on Lička Plješevica. Velebit Mts. also harbour some high alpine species not present on Lička Plješevica: Polyommatus eros (Ochsenheimer, 1808), Erebia epiphron (Knoch, 1783), Erebia euryale (Esper, 1805), Erebia gorge (Hübner, [1804]), Erebia ottomana Herrich-Schäffer, 1847, Erebia pronoe (Esper, 1780), and Erebia stiria (Godart, 1824). Additionally, there are single records form Velebit Mts. of species that could eventually be found also on Lička Plješevica such as Satyrium pruni (Linnaeus, 1758), Cupido alcetas (Hoffmansegg, 1804), Colias hyale (Linnaeus, 1758), and Neptis sappho (Pallas, 1771).

During this survey, several interesting or unexpected species were recorded. For each of them additional information on their ocurrence and distribution in Croatia is discussed.

Pyrgus serratulae (Rambur, 1839)

This species was considered very local and rare in Croatia, with only several existing records from Velebit Mts. (LORKOVIĆ, 2009). However, in the recent surveys it was recorded on several new mountain ranges including Mt. Poštak (KOREN et al., 2015a), Dinara Mts. (TVRTKOVIĆ et al., 2012, Tvrtković pers. obs. from 550 to 1450 m a.s.l.), Biokovo Mts. (KAČÍREK, 2017) and Mt. Kozjak (KOREN et al., 2019). We recorded it at seven localities: Frkašić - surroundings of pond Jezero, Kozja draga valley, Bijeli potoci - Poljane meadow, Meďugorje - Magarčev do, Mt. Kremen, and Mt. Lisac. The species is not rare on Mt. Plješevica and in some localities it was even abundant. The main reasons for the lack of records in Croatia are the similarity with the other larger Pyrgus species and the absence of targeted surveys.

Pyrgus sidae (Esper, 1784)

One of the most local and rare species of the genus Pyrgus in Croatia was not even included in the overview of the butterflies of Croatia written by Zdravko Lorković in the 1950’s (LORKOVIĆ, 2009), and neither was it represented in the collection of Croatian National Zoological Museum (today Croatian Natural History Museum) in Zagreb (MLADINOV, 1975). So far this species has been recorded from Josipdol, Gorski Kotar (MANN, 1867), Mt. Poštak (KOREN et al., 2015), Mt. Dinara, from 250-780 m a.s.l. (TVRTKOVIĆ et al., 2012; KOREN & LAUŠ, 2013) in NW Dalmatia (HABELER, 1976) and Mt. Biokovo at 650 and 700 m a.s.l. (MIHOCI et al., 2011). While most of the records originate from montane areas, the records from Tisno and Vodice, near the sea level in Dalmatia (HABELER, 1976) show that it also inhabits lowland calcareous areas. Beside the above mentioned record for Mt. Poštak, we recorded this species at two new localities, between Ondić and Rudopolje Bruvanjsko and on Mt. Lisac. The species was recently recorded also from further localities in central and Southern Dalmatia (KOREN et al., 2019).

Colias caucasica Staudinger, 1871

This species has been only recently added as a member of the fauna of Croatia, with several males collected and observed at Mt. Dinara but whithout finding of the larval host-plant (TVRTKOVIĆ et al., 2011). The observation from Mt. Poštak represents the second sighting for this species in Croatia, and it is at a distance of about 30 km north-west from Mt. Dinara. Only a single fresh male was observed so far, despite repeated efforts to make additional observations in the main flight period for the species. The potential host plant of the genus Chamaecytisus sp. is abundant on the southern slopes of Mt. Poštak, indicating a possible residence of the species on the mountain. It is also possible that it will be recorded in other high alititude grassland areas of southern Lička Plješevica range in the future.

Pieris napi / balcana

With the chromosome examination of Pieris napi and P. balcana, LORKOVIĆ (1989) found their contact zone and noted hybridisation in Plitvice area in the foothills of the Velika Plješevica. We confirmed another contact zone in Mala Plješevica. From the collected specimens, several were barcoded (Lovrenčić, personal information). Pieris napi was recorded in Bijeli potoci: Poljana (specimen with typical P. napi wing markings), and P. balcana from Poštak Mt. (890 m a.s.l,), Glogovo (830 m a.s.l.), Brezovac Dobroselski: Klanci (950 m a.s.l.) and Kremen: Kremenščak (110 m a.s.l.). Typical P. balcana wing characters were found in collected specimens from Ponor Korenički, Bijeli Potoci, Lisac, Surle, Glogovo, Kremen, Brezovac Dobroselski, Zrmanja Vrelo and Poštak (W­slope and Ljubina Poljana). Published records for P. napi based only on observaions (KOREN et al., 2011; KOREN et al., 2015) in the southern part of the range require confirmation.

Eumedonia eumedon (Esper, 1780)

Most records of E. eumedon in Croatia originate from the mountains located along the coastline of Croatia, from Mt. Učka (WITHRINGTON, 1984; MLADINOV & LORKOVIĆ, 1985), to Gorski Kotar (MLADINOV & LORKOVIĆ, 1985; KOREN, 2012), and across Velebit Mts. (GRUND, 1916; MLADINOV, 1973; LORKOVIĆ, 2009). During our surveys it was recorded on Mala Plješevica (Bijeli potoci), Mt. Kremen, Mt. Lisac, Gornja Suvaja, and confirmed for Mt. Poštak. These records, along with a recent record from Markov grob, Dinara (Verovnik, pers. obs) represent the southernmost records in Croatia, and fill the distribution gap towards known range in the neighbouring Bosnia and Herzegovina (LELO, 2008).

Cupido osiris (Meigen, 1829)

This species is very locally distributed in Croatia (LORKOVIĆ, 2009), with only a few historical records. MANN (1869) reports it from Central Dalmatia, HABELER (1976) from Podgora at the foothills of Mt. Biokovo, HAFNER (1994) from surroundings of Knin and BURGERMEISTER (1964) from Ombla. Recently it was found on Dinara Mt. (TVRTKOVIĆ et al., 2012; Verovnik pers. obs.) and Mt. Poštak (KOREN et al., 2015). During this survey, we recorded it on two localities, Bijeli potoci (Mala Plješevica Mt.) and Zrmanja spring. Only individual specimens were observed on both localities. Despite its limited distribution and specific habitat requirements (VEROVNIK, 2011a) it is not listed in the Red book of butterflies of Croatia (ŠAŠIĆ et al., 2015), therefore its status in Croatia should be rewised.

Phengaris arion (Linnaeus, 1758)

This is a rather localised but widely distributed species in montaneous parts of Croatia (ŠAŠIĆ et al., 2015). It inhabits the continental area of Croatia, where it is very local and always connected to thermophilous slopes with larval host plants (Thymmus sp., Origanum sp.). It is more widespread in the mountainous areas of Gorski Kotar and Velebit Mts. In the surveyed area, we recorded it at 12 localities, with several observed specimens. Highest densities were noted on Mt. Lisac and Mt. Poštak, while in other localities only single specimens were observed.

Plebejus argyrognom / idas

The Reverdin’s Blue is distributed in Croatia only in northern lowland part of the country (LORKOVIĆ, 2009), with the southernmost locality near Bosiljevo, in the northern foothills of the Mt. Velika Kapela. (Gorski Kotar) (MLADINOV, 1973). New records from the localities in Lička Plješevica range (Zrmanja source, Donja Suvaja near Srb, Kupirevo) are important because they represent isolated populations at the southern border of species distribution in Croatia. Two previusly published records of P. argyrognomon from Velebit Mts. and Dinara Mt. (MIHOCI et al., 2007; TVRTKOVIĆ et al., 2012), are possibly not valid. MIHOCI et al. (2007) repeated the quotation of KUČINIĆ et al. (1995) who erroneously listed the species after GRUND (1916). In Grund’s time (see GRUND, 1913) P. idas was considered as a subspecies of P. argyrognomon. The findings from Velebit’s localities are correctly interpreted by MLADINOV (1973) as P. idas.TVRTKOVIĆ et al. (2012) wrongly noted P. argyrognomon for Dinara Mt. based on barcoded female with identical barcode as several P. argyrognomon specimens from Europe (GenBank, accessed on 3-XII-2012). Later all collected males from the same locality on Dinara Mt. showed genitalia typical for P. idas. Specimens of P. idas from the area have very variable wing-markings, sometimes similar to those of P. argyrognomon.

Polyommatus thersites (Cantener, 1835)

As it is the case with many species in Croatia, our knowledge about the occurrence and distribution of the species has considerably improved in the past several years. Only a decade ago, less than ten localities were known for this species in Croatia (HABELER, 2003; MIHOCI & ŠAŠIĆ, 2006). It has been recently recorded also from Istria Peninsula (KOREN & LADAVAC, 2010), northern Dalmatia hinterland (VEROVNIK et al., 2015), in different localities in southern part of Lička Plješivica range (KOREN et al., 2011; KOREN et al., 2015), south-eastern Velebit Mts. (Javornik, leg. NT, July 2016), and Dinara Mt. (TVRTKOVIĆ et al., 2012; KOREN & LAUŠ, 2013). During this survey it was recorded at four new localities: Udbina near Kurjak, Kozja draga valley, Mt Lisac Surle -Strmica, and Veliko Popinsko polje. Additionally, it was confirmed at Zrmanja vrelo and Mt. Poštak. It was usually found in open dry grasslands or road verges with abundance of its larval hostplant, Onobrychis arenaria (Kit.) DC and/or O. vicifollia Scop. The species was moderately abundant at most localities with suitable habitat. Additional efforts are needed in order to remove potential gaps in the knowledge of its distribution in Croatia.

Thecla betulae (Linnaeus, 1758)

The Brown Hairstreak is one of the last species of butterflies to emerge in late summer, and due to that and its secretive lifestyle it may be easily overlooked during the butterfly surveys. Adults are generally scarce and local, usually flying in the treetops of ash trees and rarely visiting flowers, like Eupatorium cannabinum L. In Croatia, this is a rather local species, distributed in the continental biogeographic region, while in the coastal region it is extremely local, so far recorded only from Plitvice spring, Drežnik and Ogulin (LORKOVIĆ, 2009). Recently it was for the first time recorded on Velebit Mts. in Paklenica Gorge (TVRTKOVIĆ et al., 2015). Our records from Mt. Velika Plješevica and Ličko Petrovo selo represent important new records and fill a distribution gap towards the known range in Bosnia and Herzegovina (LELO, 2004).

Brenthis ino (Rottemburg, 1775)

The species was considered to be one of the rarest nymphalids in Croatia (MIHOCI & ŠAŠIĆ, 2005), however, it has been recorded in several new regions during the last years, although not from Mt. Poštak (KOREN et al., 2015a) or Dinara Mt. (KOREN & LAUS, 2013). The closest records to the study area are from northern Velebit Mts., where it was recorded only a decade ago (MIHOCI et al., 2007). Its distribution in Croatia includes northern Istria, and stretches across the montaneous parts of Croatia to the north as far as Hrvatsko Zagorje (KOREN et al., 2017), and parts of Slavonia (KOREN & LETIĆ, 2014). During our survey, we recorded it at nine localities at Velika Plješevica (meadows between Krgin vršak and Jukanova draga), Mala Plješevica (Bijeli potoci, Ruda Polja), Mt Kremen, Mt. Lisac and Mt. Poštak. (Tab. 1). It is possible that it will be recorded in other humid grassland areas in the region.

Limenitis populi (Linnaeus, 1758)

This species has very limited records in Croatia, and is mostly confined to the woodland areas of the north-western part of the country, with only a few scattered records from Gorski Kotar (Fužine) region (LORKOVIĆ, 2009). A single specimen was observed at the locality Bukovačka Draga at 1138 m a.s.l.. This is only the second record for Mt. Plješevica (FRANIČ, 1910), with the closest records originating from Velebit Mts. (BERRA, 1982; MIHOCI et al., 2007). The species is listed as near threatened in Croatia (ŠAŠIĆ et al., 2015), therefore further studies in suitable forest dominated areas are required to assess its distribution and endangerment.

Neptis rivularis (Scopoli, 1763)

In the view of the recent observations from Dinara Mts. (TVRTKOVIĆ et al., 2012) and Mt. Kamesnica (KOREN & LAUŠ, 2013), the record from Lička Plješevica fills the distribution gap of this species in Croatia, and indicates it has a wider distribution in the region. Several individuals were observed gliding near the forest edge at the locality Kozja Draga. The species was recently observed also in other lower parts of the Lika region outside studied area (personal observations of the authors).

Melitaea britomartis Assmann, 1847

The distribution of this species in Croatia has been poorly known until recently (KOREN & JUGOVIC, 2012). In general, this species is distributed in north-western part of Croatia, with a single record from the Lika region, and one from the Mt. Troglav in Bosnia and Herzegovina (KOREN & JUGOVIC, 2012). During this survey, it was recorded on Mt. Plješevica at the locality Bijeli Potoci. This is the second record of this species for Lika region. It indicates that further records of this species may be expected, so sampling of small Melitaea in the region is mandatory.

Araschnia levana (Linnaeus, 1758)

According to Lorkovic manuscript from 1954 (LORKOVIĆ, 2009) this species is widespread in most of the northern continental part of Croatia, especially near rivers. This species is notably absent from Istria, and across most of the Adriatic coastline. The only notable exceptions are the records from Split (STAUDER, 1922), Mljet Island (KUČINIĆ et al., 2011) and Vis Island (WITHRINGTON & VEROVNIK, 2008), but these findings were not confirmed by recent surveys (TVRTKOVIĆ et al., 2010; TVRTKOVIĆ et al., 2015; KOREN et al., 2015). During our surveys, we recorded the species at four localities in the Ligka Pljesivica range at Prijeboj, meadows between Krgin vrsak and Jukanova draga, Bijeli potoci at Poljane, and along forest track E of Zaluzje village near Donja Suvaja. These are the first records of this species for the Lika region.

Conclusions

During the last decade, a significant increase in the number of publications dealing with the butterfly fauna of previously understudied areas of Croatia is evident. This includes the studies of the butterfly diversity of the Adriatic islands (WITHRINGTON & VEROVNIK, 2008; KOREN et al., 2014; KUČINIĆ et al., 2011; VEROVNIK, 2011b), coastal areas of Dalmatia (KUČINIĆ et al., 2015; VEROVNIK et al., 2015) as well as the montaneous areas in southern Croatia (MIHOCI et al., 2011; KOREN & LAUŠ, 2013; KOREN et al., 2015; KOREN et al., 2019). Still, many areas remain poorly surveyed, and their fauna almost completely unknown. One of such areas was Lička Plješevica range which has proved to host one of the most diverse butterfly faunas in Croatia.

Such high species diversity could be explained by high habitat heterogenity, especially with pastures and grasslands in different stages of abandonment. For many centuries, herdsmen from the Lika and Velebit Mts. region led their livestock to summer pastures at higher elevations throughout most of the Lička Plješevica range (HORVAT, 1925; MARKOVIC, 2003). These practices are nowadays almost completely abandoned, and almost no pasturing was observed in the region during our visits. The only exception is Mt. Lisac, where small number of cows and sheep were observed grazing at lower altitudes. Although mowing is still practiced in the region, it is only evident near the settlements where people still persist. One of the highest species diversity was observed at Kozja Draga where some of the grasslands are still mowed. Such open-grassland localities are in danger of disappearing in the near future as grazing and mowing in the region of Lička Plješevica are almost entirely abandoned. Even within the span of our surveys in the last decade the succession is already visible, with small patches of previously open meadows becoming almost entirely covered with bushes. While it is difficult, or almost impossible to return the farming practices common only few decades ago, given the current demographic trends in the region, it is important to survey all remaining butterfly suitable areas in the region and maintain at least some sort of management in the areas with high diversity or presence of threatened butterfly species.

Agradecimientos

We are grateful to Mladen Zadravec (MZ), Leona Lovrenagic (LL), Marijana Vukovic (MV) and Ivona Buric for the help during the field trips.

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Author notes

*Autor para la correspondencia / Corresponding author Rudi.Verovnik@bf.uni-lj.si