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Historia naturalis bulgarica, 23: 109-118, 2016

Endemics and relicts in the high- fauna of

Petar Beron

Abstract: The orophyte zone in Bulgarian is about 1.37% of Bulgarian territory in eight mountains higher than 2000 m). and are the only mountains higher than 2400 m and most of the relicts and endemics are centered in these mountains.

Key words: Endemics, relicts, high-mountain fauna, Bulgaria

The orophyte zone in the Bulgarian mountains uted greatly to the knowledge of these (several starts usually at 1900 – 2200 m a.s.l. This is the up- groups). Most animals living on the top of Bulgarian per limit of the closed high (the mountain fur (and other) mountains could be considered as neoen- excluded). This high mountain (orophytic) zone demics. of Bulgaria consists of 1.37% of Bulgarian territo- ry (above 2000 m are 1.05% of the total surface of Development of the high mountain environment Bulgaria, the land above 2500 m – 0.18%.) in eight in Bulgaria mountains (Rila – 2925 m a.s.l., Pirin – 2914 m, Stara The development of the fauna is closely related planina – 2386 m, – 2290 m, Ossogovska with the development of the plant communities and planina – 2261 m, – 2212 m, Rhodopes – is indeed determined by it. It is justified to consider 2191 m, Belassitsa – 2029 m). Particularly important here only the changes of the vegetation in the last are the mountains exceeding 2400 m a.s.l. (Rila and 15 000 years (the Glacial and Postglacial time, or Pirin). Here are represented the ‘’true” high-moun- the end of the and the entire Holocene). tain dwellers, excluding the inhabitants of the eco- According to Palamarev (1982), “The - tone and the carried – away specimens. ous become dominant formation in Rila, If we consider as high-mountain fauna the Pirin and Rhodopes between 1000 and 2000 m by animals living higher than 1900 m a.s.l., in the eight the end of the Subboreal and the beginning of the Bulgarian mountains exceeding 2000 m at least 2300 Subatlantic phase. In the same time above the timber species of terrestrial Metazoa have been recorded line there have been developed groups of mountain above 1900 m (Beron, 1999, with supplements). pines and alpine herbaceous communities”. The ma- The highest mountains (Rila and Pirin) have been jority of our knowledge on the history of the high- glaciated, but at present there are no . These mountain flora in the Rila and Pirin Mountains we mountains contain the highest number of glacial rel- owe to Dr E. Bojilova (Božilova) and her associates. icts, but such relicts are found also on the territory of According her (Božilova, 1995), our information Vitosha, Central Balkan (Stara planina), Ossogovska about the changes of the upper forest line in the planina and Slavyanka Mountains. Rila Mountains goes to the end of the last glaciation The glacial relicts in Bulgaria were firstly sum- (10 500 years ago). In the arid climatic conditions marised first by Buresch & Arndt (1926), than during the Glacial periods forest vegetation has not Beron (1969) collated a list of “boreoalpine” species been developed in the mountains of South . in Bulgaria. Recently several detailed studies of the The present-day upper forest line (formed by Pinus fauna of Rila, Pirin and Central Balkan (Sakalian V. , sylvestris, P. peuce and Picea abies) has taken shape Ed., 1997; Sakalian M., Ed., 2000a, 2000b) contrib- only in the Subatlantic time (less than 7000 years 110 Petar Beron ago). With the beginning of the Atlantic phase (8000 Fam. Scutacaridae: Scutacarus pirinicus Dobrev years BP) domination of and P. peuce – Pirin, Stara planina Mountains starts. By contrast with Stara planina, in the Rila Fam. Trombiculidae: Neotrombicula monticola Mountains the beech (Fagus) has never reached the Kolebinova – Pirin upper forest limit. About 4000 years ago the upper Fam. Erythraeidae: Erythraeus rilensis Beron, E. forest line has been fixed already at the present day bulgaromontanus Beron – Rila level of 2000 m a.s.l. Oribatida Fam. Carabodidae: Carabodes pirinensis Kunst Endemism. Here as endemics are considered – Pirin taxa which are found only on the Balkan Peninsula Fam. Eremaeidae: Eremaeus valkanovi Kunst or have a more restricted areal (usually recorded Pauropoda only on one or several of its mountains). Fam. Pauropodidae: Allopauropus doryphorus Endemism above 1900 m a.s.l. Remy – Rila Oligochaeta. From the nine species of Chilopoda Oligochaeta known from above 1900 m three are Lithobiomorpha endemics: Dendrobaena balcanica (Černosvitov), Fam. Lithobiidae: Lithobius (L.) borisi Verhoeff D. hrabei (Černosvitov), and D. rhodopensis – Pirin (Černosvitov). L. (L.) electron Verhoeff – W. Rhodopes Gastropoda. From the 31 species, found above L.(L.) glaciei Verhoeff – Ossogovo 1900 m, endemic for the higher parts of the Pirin Fam. Geophilomorpha Mountains are two species of Wladislawia: W. polin- Fam. Geophilidae skii (Wagner) and W. sztolcmani (Wagner). Geophilus rhodopensis Kaczmarek – W. Arachnida. Endemics or relicts that have Rhodopes, Vitosha been recorded above 1900 m a.s.l. are found among Diplopoda Araneae and Acariformes. Iulida Araneae. Endemism is high among the high- Fam. Iulidae: altitude . From 192 species living above 1900 Leptoiulus borisi Verhoeff – Rila, Vitosha, Pirin m, 12 (6,2 %) are Bulgarian and local endemics: Megaphyllum glossulifer (Schubart) – Rila Fam. Zodariidae: M. rhodopinus (Verhoeff) – Rhodopes, Zodarion pirini Drenski – Pirin, , Rila Slavyanka, Pirin and Vitosha Mountains. Chordeumatida Fam. Linyphiidae (incl. Erigonidae) Fam. Anthroleucosomatidae: Bulgarosoma su- Araeoncus clivifrons Deltshev – Rila and Pirin, perficiei Strasser – Vitosha not found below the upper border of the forest Insecta s.l. (Deltshev). Collembola. In the pot holes of the higher parts Erigone pirini Deltchev – Pirin, Rila, Stara of Pirin we have collected some species, described planina. as new and so far considered endemic (Pomorski, Diplocephalus altimontanus Deltshev – Pirin. 2006). Metopobactrus orbelicus Deltshev – Pirin and Rila. Fam. Onychiuridae Antrohyphantes rhodopensis (Drenski) – Pirin, Rila. Onychiuroides bojani Pomorski, O. peteri Drepanotylus pirinicus Deltchev – Pirin. Pomorski, O. pirinicus Pomorski – Pirin Mountains Cеntromerus paucidentatus Deltchev – Pirin Orthoptera. The endemism in the mountain and Rila. species of Orthoptera from Bulgaria has been out- Mughyphantes lithoclasicolus (Deltchev) – Pirin lined by Peschev (1987). Most of the species, enu- and Rila. merated as “paleoendemics” or “neoendemics”, are Tenuiphantes drenskii (Helsdingen) – Rila. not distributed above 1900 m. All the “paleoen- Mansuphantes rectilamellus (Deltshev) – Pirin. demic” species and the high-mountain “neoendem- Fam. Lycosidae: Pardosa drenskii Buchar – ics” are Poecilimon spp. from Pirin. According to Vitosha, Pirin, Rila, Stara planina Mountains. Popov (1997b), the species of Orthoptera from the Acari could be considered as endemics Acariformes. (neoendemics). However, only five of them are re- Prostigmata. Currently some terrestrial corded above 1900 m a.s.l.: Poecilimon harzi Peshev Prostigmata are considered endemic: (1500-2400 m, endemic for Pirin), P. m. mistshenkoi Endemics and relicts in the high-mountain fauna of Bulgaria 111

Peshev (1250-2000 m, endemic for Pirin), P. orbelicus tles (Carabidae) in Bulgaria has been studied by Panč. (900-2400 m, Balkan endemic), Metrioptera Guéorguiev in Sakalian & Guéorguiev (1997). arnoldi Ramme (1200-2200 m, Balkan endemic), According to these authors among the 756 species and Pholidoptera aptera karnyi (Ebner) (950-2400 and subspecies of Carabidae in Bulgaria 125 species m, Balkan endemic). and subspecies, belonging to 34 genera, are Balkan In the six species are con- endemics (some of them Bulgarian or local endem- sidered as Balkan endemics by Popov (in Hubenov ics). “The lowest level of endemism diversity has et al., 2000a) as Balkan endemics, four of them liv- been observed in the orophyte zone: 32 (34.04%). ing above 1900 m. Three of them coincide with Although as few as eight endemic carabid taxa occur the endemics recorded in the higher parts of Pirin in Bulgaria only within this zone, as many as seven Mountains: Poecilimon orbelicus Panč., Metrioptera of them, or 19,44% of the total local endemics, are arnoldi Ramme and Pholidoptera aptera karnyi confined to the orophyte belt, this share being the (Ebner). The fourth is Psorodonotus fieberi Friv. (re- highest among Bulgaria’s local epigean endemics – corded from the Rila, Rhodope, Belassitsa, and Stara (p.33-34). planina Mountains). In Central Balkan National Park Our analysis has shown that above 1900 m in five Balkan endemics were recorded so far (Popov, Bulgaria 30 species and 17 subspecies of endemic in Hubenov et al., 2000b), including four living ground have been found. They belong to 15 above 1900 m. Again three species are found also in genera‚ as follows: the high parts of Rila: Metrioptera arnoldi Ramme, Duvalius (Paraduvalius) klimai Janák et Moravec Pholidoptera aptera karnyi (Ebner) and Psorodonotus – Bulgarian endemic fieberi Friv. The fourth species is Isophya obtusa D. (Paraduvalius) kuboni Janák et Moravec – Br.-W. As local endemics is described the subspecies Bulgarian endemic Isophia pravdini bazyluki Peshev. Carabus violaceus azurescens Dejean – Balkan Altogether, from 44 species of Orthoptera, found endemic in Bulgaria higher than 1900 m, the following eight C. cavernosus cavernosus Friv. – Balkan endemic (18,2 %) could be considered endemics (all of them Cychrus semigranosus balcanicus Hopffg. – members of Tettigoniidae): Isophia obtusa Br.-W., Balkan endemic I. rhodopensis Ramme, I. pravdini bazyluki Peshev, Nebria eugeniae Daniel – Endemic to Rila Poecilimon harzi Peshev, P. m. mistshenkoi Peshev, P. N. hybrida hybrida Rottenberg – Rila orbelicus Panc. (Balkan endemic), Pholidoptera ap- N. hybrida pirinensis Horv. – Endemic to Pirin tera karnyi (Ebner)(Balkan endemic), Metrioptera N. hybrida rhodopensis Horv. – Rhodope arnoldi Ramme. N. rhilensis Friv. – Endemic to Rila and Pirin Plecoptera. From the 101 species of this order Trechus bohemorum Pawlowski – Endemic to in Bulgaria 16 live above 1900 m, including ten en- Rila and Pirin demic species (ca. 10%): T. cardioderus balcanicus Jeannel – Balkan en- Fam. Taeniopterygidae: demic Brachyptera bulgarica Raušer (Pirin) T. demircapicus Moravec – Endemic to Pirin Fam. Nemouridae: T. gulickai Löbl – Endemic to Pirin Nemoura bulgarica Raušer (Stara planina, Rila, T. matrismeae Pawlowski – Rhodope Pirin, Rhodope) T. merkli Pawl. – Western Stara planina (en- N. pirinensis Raušer (Stara planina, Rila, Pirin, demic to Stara planina) Rhodope) T. orphaeus Pawlovski – Endemic to Rila Protonemoura tarda Braasch (Rila and Pirin) T. pirinicus Pawlowski – Endemic to Pirin P. mattheyi (Aubert)(Balkan endemic) T. priapus Daniel – Balkan endemic Fam. Leuctridae: T. rambouseki Breit – Endemic to Rila Leuctra joosti Braasch (Vitoscha, Pirin) T. rhilensis Kaufmann – Endemic to Vitosha L. kumanskii Braasch et Joost (Pirin) and Rila Fam. Perlodidae: T. rhodopeius Jeannel – Endemic to Bulgaria Isoperla buresi Raušer (Rila, Pirin, Rhodope) T. szujeckii Pawlowski – Rhodope Chloroperla russevi Braasch (Balkan endemic) Asaphidion caraboides balcanicus Netolitzky – Ch. kozarovi Braasch (Balkan endemic) Balkan endemic Coleoptera Bembidion rhodopense Apf. – Balkan endemic Fam. Carabidae. Endemic ground bee- Xenion ignitum (Kraatz) – Balkan endemic 112 Petar Beron

Pterostichus rh. rhilensis Rottb. – Rila, Rhodope, macrocephalus Zerche (Pirin), O. musalensis Zerche Stara planina (Rila), O. ilievi Zerche (Vitosha), O. behnei Zerche P. rhilensis kourili Mařan – Endemic subspecies (Vitosha, Rila), O. doeblerae Zerche (Rila), O. heide- in Pirin mariae Zerche (Pirin). Particularly interesting is the P. rhilensis vitosensis Mařan – Vitosha, Central Ophthalmoniphetodes, studied by L. Zerche. Balkan Fam. Curculionidae. Among the 63 weevil spe- P. macedonicus Apfelbeck – Belasitsa, Stara cies found above 1900 m the following eight (12,7%) planina (Balkan endemic) are endemic: Otiorrrhynchus biformatus Mazur Tapinopterus balcanicus balcanicus Ganglbauer (Pirin), O. rhilensis Stierlin (Rila, Rhodope, Central – Bulgaria Balkan, Pirin, Vitosha, 1700 – 2600 m), O. joakimo- T. balcanicus belasicensis Mařan – Belassitsa ffi Apfelbeck (Pirin, Rila), O. serdicanus Apfelbeck T. kaufmanni kalofirensis Mařan – Bulgarian (Pirin, Rila, Vitosha), Plinthus sturmi bulgaricus endemic Meregalli, Omias taygetanus Purkyne (Balkan en- T. kaufmanni winkleri Mandl – Stara planina, demic), Tachyphloeus bosnicus Apfelbeck (Balkan East (Balkan endemic) endemic), Alophus rhodopensis Reitter (Vitosha, Rila, T. kaufmanni kulti Mařan – Rila, Rhodope Pirin, Rhodope). Molops alpestris centralis Mlynař – endemic for Fam. Elateridae. From the nine species of click Bulgaria beetles living above 1900 m endemic for Bulgaria is M. alpestris kalofericus Mlynař – endemic for Ctenicera schneebergi Roubal. Bulgaria Fam. Chrysomelidae. Among the 43 leaf bee- M. alpestris rhilensis Apfelbeck – Endemic for tles that are found above 1900 m Balkan endemic is Bulgaria Oreina speciossisima drenskii (Gruev). Luperus rhilen- M. dilatatus dilatatus Chaudoir – Balkan endemic sis Weise is endemic for Rila Mountains. Among the M. dilatatus angulicollis G. Müller – endemic 289 species of leaf beetles in Pirin (Gruev, 2006) there for Bulgaria are six Balkan endemics and one (Longitarsus behnei M. piceus bulgaricus Mařan – , Gruev et Arnold) is local endemic. Bulgaria (Balkan endemic) Fam. Leiodidae. Catops pirinensis Zerche is en- M. rhodopensis rhodopensis Apfelbeck – demic for the Pirin Mountains (, 2100 -1500 m). Endemic for Bulgaria . Dimorphocoris fuscus Joakimoff Calathus ellipticus Reitter – Balkan endemic is endemic for Rila, Pirin and Vitosha. C. metallicus aeneus Putzeys – Balkan endemic Trichoptera. Among the 57 species of cad- Laemostenus plasoni (Reitter) – Endemic for dis flies known to occur in Pirin National Park (73 Bulgaria in the whole of Pirin), 13-14 species of the families Amara messae Baliani – Balkan endemic Rhyacophilidae, Glossosomatidae and Limnephilidae Zabrus rhodopensis Apfelbeck – Balkan endemic are considered by Kumanski (1997) to be endem- Fam. Dytiscidae. From 27 species living above ic. Some species or subspecies are Bulgarian en- 1900 m only Agabus balcanicus Hlisnikovsky (Pirin, demics (Rhyacophila pseudotristis Kumanski, Rh. Vitosha, Rila, Stara planina, Slavyanka Mountains), braaschi Malicky et Kumanski, Synagapetus mon- the subspecies Agabus solieri falcozi Guignot (Pirin) tanus Kumanski, Drusus romanicus meridiona- and Illybius fuliginosus pirinicus Guéorguiev (Pirin) lis Botosaneanu et Riedel, D. discophorus pallidus could be considered endemic. Kumanski, Chaetopteroides bulgaricus (Kumanski), Fam. Staphylinidae. From the 124 species of Psilopteryx schmidi Marinković, and Wormaldia bul- row beetles known from Bulgaria above 1900 m garica Novak). All endemics reach altitudes above there are 20 Bulgarian and local endemics (16.6%): 1900 m. Omalium bulgaricum Zerche, Deliphrosoma pirinense Kumanski (in Hubenov et al., 2000a) pro- Zerche (endemics for Pirin), Geodromicus schuberti vides a list of 11 Balkan, ten Bulgarian and one lo- Scheerpeltz, Gabrius beroni Raitschev (Ossogovska cal endemics, recorded in the Rila National Park. planina), Atheta scintillans Scheerpeltz, Leptusa rhilen- Among them the Balkan endemics Rhyacophila sis Pace, Ocalea bulgarica Scheerpeltz, Ocyusa ferdi- loxias Schmid, Wormaldia bulgarica Novak, Drusus nandicoburgi Rambousek, O. regisborisi Scheerpeltz, botosaneanui Kumanski, Rhadicoleptus alpestris Ophthalmoniphetodes longicornis Zerche (Pirin), O. macedonicus Botosaneanu et Riedel, Chaetopteryx maljovicensis Zerche (Rila), O. piger Zerche (Rila), O. stankovici Marinković, Annitella triloba Marinković- gracilis Zerche (Pirin), O. rhilensis Zerche (Rila), O. Gospodnetić, the Bulgarian endemics Rhyacophila Endemics and relicts in the high-mountain fauna of Bulgaria 113 pseudotristis Kumanski, Rh. obtusa Klapálek, For me this opinion is too generalising. Relics Synagapetus montanus Kumanski, Drusus romani- and even “living fossils” exist in many groups cus meridionalis Botosaneanu et Riedel, D. disco- (classic examples being animals like Latimeria, phorus pallidus Kumanski, Chaetopteroides bulgari- Sphenodon, and Okapia). For some groups such as cus (Kumanski), Psilopteryx schmidti Marinković, the Pseudoscorpions, the relics are indisputable. and the regional endemic Chionophylax monteryla Such are the Troglochthonius from ex-Yugoslavi- Botosaneanu, all being recorded above 1900 m. an and the species of the family Syarinidae Kumanski (in Hubenov et al., 2000b) reports (Chitrellinae). In the caves of Santorin and Iraklia we four caddis flies from the Central Balkan National had the chance to find the first Syarinidae in Balkan Park above 1900 m: one Balkan endemic and three Peninsula (Hadoblothrus aegaeus Beron). The spe- Bulgarian endemics in Central Balkan National Park cialists consider the Chitrellinae on the Europe as (Rhyacophila loxias Schmid, Rh. obtusa Klapálek, relicts. Beier (1969): “Syarinidae: Wohl bei keiner Rh. kownackiana Szczesny, and Drusus botosaneanui anderen Familie ist der Reliktcharakter so aus- Kumanski). geprägt”. Another good example is the Opilion Altogether, according to Kumanski (in Hubenov Paralola buresi (Laniatores, Phalangodidae), living in et al., 2000), out of 244 species of Trichoptera in four caves near Lakatnik (Stara planina). Here also Bulgaria, 17 species are endemic for Bulgaria (incl. there is another opinion of Martens (1972), disput- nine living above 1900 m) and 16 sp. are endemic ing the widely accepted relict nature of European for the Balkan Peninsula (incl. five living above 1900 Laniatores. He says that: “The European Laniatores m), or the endemic Trichoptera in the Bulgarian should no longer be regarded as tertiary relics for high-altitude zone are 14. Represented are the fami- they are nowadays widely distributed inhabitants of lies Rhyacophilidae (four sp.), Glossosomatidae litter in areas not covered by ice during the gla- (one species), Philopotamidae (one species), and cious periods”. Limnephilidae (eight species). According to Lopatin (1989): “As relicts are . In the Pirin National Park five spe- considered species (or genera), which have already cies have been found only in the alpine belt and are left behind their time of maximal development and considered “characteristic endemics”(Abadjiev, 1997): having areals either restricted or becoming restrict- Erebia rhodopensis Nicholl (1900-2500 m, Balkan en- ed. The present day conditions of the environment demic), E. cassioides macedonica Buresch (2000-2600 do not correspond to their ecological requirements m, endemic for Rila and Pirin), cynthia le- – this is the main factor determining the relict forms. onhardi (Röber) (2000-2800 m, endemic for Rila and The flourishing of these species is now impossible, Pirin), Boloria pales rilaensis Varga (2600-2800 m, en- what is witnessed by the diminution of their number demic for Rila and Pirin) and coracina bureschi and the reducing of their areals. Relicts could be very Varga (2600-2900 m, local endemic). old (Hateria) or younger (the so-called glacial relicts, like Lepus timidus in Central Europe”. Relicts. The notion of “relict” is widely used by bioge- The notion of “relict”. ographers, despite of the warning of some of them to According to Wikipedia, “In biology a relict (or their colleagues to not allow themselves to be carried relic) is an organism that at an earlier time was abun- away too much (Еlenevskiy & Radigina, 2002). dant in a large area but now occurs at only one or a This notion was used widely by biospeleolo- few small areas”. gists, influenced by the book of Jeannel (1942) “Les This notion has been discussed still by fossils vivant des cavernes”. Some researchers oppose Wangerin (1912). However, some authors contest strongly this very notion, as does the prominent the very notion of “relict”. We may quote Brignoli Italian Arachnologist Brignoli (1979): “Le terme de (1979), discussing the cave spiders: “relicte” (ou même de “fossile vivant”) si souvent em- “pour connaître l’histoire de peuplement d’une ployé pour les troglobies, n’a pour moi aucun sense”. région, les troglobies n’ont aucune valeure spéciale. Checking in the dictionaries, we can find sev- Tous les animaux ont la même importance ...” eral interpretations of “relict”. “il n’est pas du tout vrai (ou, au moins, ce n’est Merriam-Webster Dictionary: “A surviving pas du tout sûr) que les troglobies sont anciens.” species of an otherwise extinct group of organisms; “La terme de “relicte” (ou même de “fossile also: a remnant of a formerly widespread species that vivant”) si souvent employé pour les troglobies, n’a persists in an isolated area”. pour moi aucun sense.” The Free Dictionary: “An organism or species 114 Petar Beron of an earlier time surviving in an environment that ous papers of Deltshev and other authors point at the has undergone considerable change”. following: The botanists have three concepts to classify Preglacial relicts relicts: geographic, taxonomic, and lineage relicts. Fam. Linyphiidae According Birstein (1947), confirmed by Antrohyphantes balcanicus (Drensky) Vandel (1964): “Les rélicts comme des types ani- Glacial relicts maux (ou végétaux) dont l’évolution est arrêtée ou du Fam. Linyphiidae moins fort ralentie, et qui ont conserve le facies de Diplocephalus foraminifer (O.P. Cambridge) leurs lointains ancêtres”. – Rila, Pirin, Stara planina (Botev Summit)(1895 – Darlington (1957, but translated from the 2925 m) Russian edition in 1966): “ or plant which: 1. Mecynargus paetulus (O.P. Cambridge) – Pirin, Keeps existing in given place after its extinction or the Stara planina (Summit Botev)(up to 2914 m) extinction of its related forms elsewhere (geographical Improphantes improbulus (Simon) – Pirin, Rila, relict) or 2: Exists after the extinction of most of the and Stara planina (up to 2925 m) group (evolutionary or phylogenetical relict).” Fam. Clubionidae In the recent paper by Grandcolas, Nattier Clubiona alpicola Kulczynski – Pirin, Rila, & Trewick (2014) relicts (geographical or phyloge- Rhodopes, and Vitosha (up to 2914 m) netic) are identified as „a species or a group of species Chilopoda. Following Stoev (2007), we should remaining from a large group that is mainly extinct”. take into account that “Several species have been re- They speak of “Relict species: a relict concept?”. corded from the subalpine and alpine zones of the A special type of relictual distribution is the old- Bulgarian mountains, but with almost no exception er notion of Boreo-Alpine distribution (now are used all were found in the lower, forest belts as well. Only mostly the terms of Arcto-Alpine and Boreomontane Lithobius electron, L. glaciei, and L. borisi have been distributions). These elements in Bulgarian fauna described from the altitudes above 2000 m a.s.l. in have been discussed by Beron (1969). the Rhodopes, , and Pirin Mts. respective- ly, and were not collected again As a result, no real Relicts in the high-mountain fauna of Bulgaria subalpine or alpine species have been found so far Glacial relicts are the most typical relicts. The among the Bulgarian myriapods. data about them, obtained by the beginning of XX Diplopoda. Stoev (in Deltchev et al., 1999a) century, have been summarised by Buresch & lists four species of Diplopoda as relicts for the Rila Arndt (1926), inspired by the work of K. Holdhaus. National Park, including two over 1900 m a.s.l.: These authors report 63 species from Bulgaria Leptoiulus borisi Verhoeff and Megaphyllum glossu- and Macedonia, considered relicts. Prevail the lifer Schubart. Lepidoptera, already well known in Bulgarian moun- Orthoptera. Popov (1997a) considers four tains. Other groups are Carabidae and Dytiscidae species of grasshoppers living in the Pirin National (Coleoptera). Many species are known from altitude Park as relicts – three glacial relicts: Melanoplus frig- much lower than 1900 m a.s.l. idus (Boh.), Gomphocerus sibiricus (L.), Aeropedellus Since then a lot of new data have been obtained variegatus F.-W., and one preglacial relict Anterastes by various researchers. The data about most relict serbicus Br.-W. All of them are typical for the highest from Pirin have been collated in the monograph, parts of the Bulgarian mountains. Anterastes serbicus edited by Sakalian (1997), but there are no such is considered (together with Gomphocerus sibiricus studies about other Bulgarian mountains. The eight and Aeropedellus variegatus) as glacial relict on Rila articles on Orthoptera, Plecoptera, Ephemeroptera, (Popov in Hubenov et al., 1999a) and (together Heteroptera, Neuroptera, Coleoptera, Trichoptera, with Gomphocerus sibiricus and Melanoplus frigidus) and Lepidoptera contain information about several as glacial relict for the Central species of , glacial relicts or tertiary relicts. (Popov in Hubenov et al., 1999b). Anterastes ser- Important information is contained in the volumes bicus lives also in the mountains Vitosha, Rhodope, on the Biodiversity of the National Parks Rila and Slavyanka, and Belassitsa, Melanoplus frigidus – also Central Balkan (Sakalian M. (Ed.), 1999a, 1999b). from Slavyanka Mountain, Gomphocerus sibiricus – Arachnida. also from Vitosha, Ossogovska planina, Belassitsa, Araneae. The orophyte zone of several Slavyanka, and . Detailed anal- Bulgarian mountains is inhabited by a number of ysis of the mountain Orthopterids in Bulgaria is spiders, – preglacial and glacial relicts. The numer- made by Popov (2007). According to this author, Endemics and relicts in the high-mountain fauna of Bulgaria 115

Melanoplus frigidus “is a young (glacial) relict with Fam. : Dichrooscytus valesianus Fieber, a typical Arctoalpine type of distribution”. A relict Dimorphocoris fuscus Joakimov Stenobothrus cotticus Kruseman et Jeekel surviv- Fam. – Carpocoris melanocerus ing in an interglacial refugium in Rila (2300-2650 (Mulsant et Ray) m) has been announced and analysed by Berger, Glacial relicts: Chobanov & Mayer (2010). Fam. Corixidae: Arctocorisa carinata (C. Coleoptera Sahlberg), A. germari (Fieber) Carabidae. From 108 species of ground beetles Fam. Miridae: Placochilus s. seladonicus in Bulgaria known to live above 1900 m a.s.l. the (Fallén), Lygus wagneri Remane, Monalocoris fili- following could be considered relicts (Sakalian in cis (L.), Orthops basalis (A. Costa), O. montanus Hubenov et al., 2000a, 2000b): (Schilling), Orthotylus virescens (Douglas et Scott), Amara erratica Duftschmid – G (Rila, Pirin, Phoenicocoris obscurellus (Fallén), Phytocoris pini Vitosha, Rhodope, Stara planina, and Ossogovska Kirschbaum, Pinalitus rubricatus (Fallén), Globiceps planina Mountains) dispar Boh., G. flavomaculatus (F.), Psallus haema- A. nigricornis Thomson – G (Rila, Stara planina) todes (Gmellin), Atractotomus magnicornis (Fallén), A. quenseli (Schönherr) – G (Rila, Pirin, Stara Bryocoris pteridis (Fallén), alpestris (Meyer planina, and Slavyanka Mountains) – Dür), C. sexguttatus (Fabricius) Bembidion bipunctatum nivale (Schönherr) – Fam. Anthocoridae: Anthocoris nemorum (L.), G (Rila, Pirin, Vitosha, Stara planina, and Rhodope Acompocoris alpinus Reuter Mountains) Fam. Reduviidae: Rhinocoris annulatus (L.) Nebria rufescens (Ström) (= gyllenhali Fam. Nabidae: Nabis brevis Scholtz, N. fla- Schönherr) – G (Rila, Pirin, Stara planina) vomarginatus Scholtz, N. limbatus Dahlbom, N. ru- Dytiscidae. In the Rila National Park six spe- gosus (L.) cies of this family are considered relicts (Sakalian Fam. Saldidae: Salda littoralis littoralis (L.), in Hubenov et al., 1999a), all of them living above Saldula orthochila (Fieber), S. c-album (Fieber), 1900 m: Oreodytes davisi Curtis, Hydroporus tartari- Macrosaldula scotica (Curtis) cus Le Conte, H. nivalis Heer, H. kraatzi Schaum, Fam. Aradidae: Aradus pallescens frigidus Potamonectes griseostriatus (De Geer) and Coelambus Kiritschenko novemlineatus Steph. Fam. Lygaeidae: Nithecus jacobaeae (Schilling), Agabus (Gaurodytes) solieri Aubé – Rila (2195- Nysius thymi (Wolff), Trapezonotus desertus 2460 m), Pirin (2190-2525 m), Vitosha (above 2000 Seidenstücker, and Megalonotus dilatatus (Herrich- m). West Palaearctic Boreo-Alpine species. Glacial Schäffer) relict. Fam. Stenocephalidae: Dicranocephalus medius The species Oreodytes davisi, Hydroporus tar- (Mulsant et Rey) taricus and H. nivalis have been recorded also from Fam. Coreidae: Ulmicola spinipes Fallén Vitosha and Pirin Mountains, Agabus solieri Aubé Fam. Rhopalidae: Stictopleurus crassicornis (L.) was found in Vitosha, endemic subspecies of it (A. s. Fam. Scutelleridae: Eurygaster dilaticollis Dohrn falcozi) – in Pirin. Fam. Pentatomidae: microphthalmus Flor, Hydrophilidae. The only endemic species is Aelia klugi Hahn, A. sibirica Reuter, Carpocoris pur- Helophorus glacialis Villa (1400-2600 m, Vitosha, pureipennis (De Geer), and Chlorochroa juniperina (L.) Rila, Stara planina, and Pirin Mountains). Fam. Cydnidae: Canthophorus impressus Curculionidae. As relicts is considered the spe- (Horváth) cies Otiorrhynchus dubius (Ström) (Rila Mts.) Neuroptera. According to Popov (1997b), Elateridae. Relict species are Ctenicera cuprea F. only Wesmaelius (Kimminsia) malladai (Navas) and Hypnoidus riparius (F.) (Hemerobiidae, 1600-2050 m) could be considered Chrysomelidae. Among the 289 species of leaf as a glacial relict in the Pirin Mts.(rarely found above beetles in Pirin (Gruev, 2006) there are also relict. the timberline). The same species is the only relict Heteroptera. Among the 101 species of Neuroptera in the Central Balkan Mts. (Popov in Heteroptera known to exist higher than 1900 m in Hubenov et al., 2000a). For the Rila National Park Bulgaria as much as 43 species could be considered Popov in Popov et al., 2000, mentions five relict relicts (Josifov in Hubenov et al., 2000a, 2000b; Neuroptera (all Hemerobiidae), but only two of the Josifov in Sakalyan (ed.), 1997: have been found higher than 1900 m: Wesmaelius Praeglacial relicts: malladai (Navas) and Hemerobius schedli Hölzel. 116 Petar Beron

Diptera. orientalis orientalis Elwes, E. pandrose ambicolorata Trichoptera. According to Kumanski (in Varga (= E. lappona), E. pronoe fruhstorferi Wrn., Hubenov et al., 2000a, 2000b) two members to the and E. cassioides macedonica Buresch family Limnephilidae are glacial relicts: Asynarchus Fam. Geometridae: Eupithecia fenestrata lapponicus (Zetterstedt) from Rila and Chionophylax Millière, limbaria rablensis Zeller, Gnophos mindszentyi bulgaricus Kumanski from the Central glaucinarius peruni Varga, G. obfuscatus (Denis et Balkan Mountains (endemic subspecies). Schiff.), Catascia dilucidaria (Denis et Schiff.) Lepidoptera. Among Lepidoptera of the Pirin Fam. Arctiidae: Arctia flavia (Fuessly) National Park 19 species are considered by Abadjiev Fam. : Parasemia plantaginis inter- (1997) as glacial relicts. They belong to the families rupta Draudt, Syngrapha interrogationis (L.), and S. Hesperiidae, Geometridae and Noctuidae, and are divergens rilaecacuminum Varga et Ronkay all inhabitants of open herbaceous formations: 11 Vertebrata have been recorded above 1900 m: Pyrgus cacaliae Amphibia. Two species are considered gla- (Rambur) (Hesperiidae, 2800 m), Entephria caesiata cial relicts: Rana temporaria L. and Ichthyosauria (Denis et Schiff.)(1500-2100 m), E. flavicinctaria [Triturus] alpestris (Laurenti) (Beshkov in Beron et Duponchel (1600-1950 m), Venusia cambrica Curtis al., 2000a, 2000b). (1950 m), Catascia dilucidaria (Denis et Schiff.) Reptilia. The lizard Zootoca vivipara (2236 m), Gnophos obfuscatus (Denis et Schiff.) (Lichtenstein) and the viper (L.) are the (1230-2000 m), Syngrapha interrogationis (L.)(1800- only relict reptiles in the high mountains of Bulgaria 2000 m), Discestra melanopa (Thunberg)(2700 m), (Beshkov in Beron et al., 2000a, 2000b). Lycophotia porphirea (Denis et Schiff.) (2000 m), Aves. Tengmalm’s owl (Aegolius funereus L.) grisescens (F.)(1950 m), and Xestia ashwor- is considered a glacial relict from Rila, Central thii candelarum (Staudinger)(1800-1950 m a.s.l.). All and Western Stara Planina, Pirin, and Rhodopes endemic Lepidoptera reach the highest areas above Mountains (Shurulinkov et al., 2003). 2500 m, but from the 19 relict species only two live in this zone. The others usually remain below 2100 m. Conclusions From 114 species of Lepidoptera in Bulgaria living In the eight mountains in Bulgaria higher than above 1900 m 26 are considered relicts (Abadjiev in 2000 m are distributed many species, considered Hubenov et al., 2000a, 2000b): local or Balkanic endemics, some of them relicts, Fam. : aethiopella (Duponchel), mostly preglacial and glacial. The groups with numer- Titanio [Metaxmeste] schrankiana (Hochenw.) ous endemics are Araneae, Orthoptera, Plecoptera, Fam. Hesperiidae: Pyrgos cacaliae (Rambur) Coleoptera, Trichoptera, and Lepidoptera. Relicts Fam. Pieridae: Colias caucasica balcanica Rebel are known among Araneae, Diplopoda, Orthoptera, Fam. : Boloria graeca balcanica Heteroptera, Coleoptera, Lepidoptera and even ver- (Rebel), B. pales rilaensis Varga, B. selene selene tebrates like Rana temporaria, Ichthyosauria alpestris (Denis et Schiff.), Euphydryas cynthia leonhardi (Amphibia), Zootoca vivipara, Vipera berus (Reptilia) (Röber), Erebia rhodopensis Nicholl, E. gorge pirini- and Aegolius funereus (Aves). Rila and Pirin are the ca Buresch, E. melas leonhardi Frühstorfer, E. oeme only mountains higher than 2400 m and most of the spodia Staudinger, E. ottomana balcanica Rebel, E. relicts and endemics are centered in these mountains. Endemics and relicts in the high-mountain fauna of Bulgaria 117

References [Elenovskiy A.G., Radigina V.I. ] Еленовский А.Ж., Радигина Abadjiev S. 1997. Lepidoptera. – In: Sakalian V. (ed.): Endemic В.И. 2002. О понятии “реликт” и реликтомании в and Relict Insects in the Pirin National Park‚ Bulgaria. географии растений [On the notion of “relict” and the rel- Pensoft Publ.‚ – Moskow‚ 69 -77. ictomania in the ]. – Buletin Moskovskogo Beier M. 1969. Reliktformen in der Pseudoscorpioniden-Fauna Obshtestva Ispitateley Prirodi, Otdel Biologii, 107 (3): 39 Europas. – Memorie Societa Entomologica Italiana, 48: -48. (In Russian). 317-323. Grandcolas Ph., Nattier R., Trewick S. 2014. Relict species: Berger D., Chobanov D., Mayer F. 2010. Interglacial refugia and a relict concept? – Trends in Ecology & Evolution, 29(12): range shifts of the alpine grasshopper Stenobothrus cotticus 655-663. (Orthoptera: Acrididae: Gomphoceratinae). – Organisms, Gruev B.A. 2004. The leaf beetles (Insecta: Coleoptera: Chry- Diversity & Evolution, 10 (2): 123-133. somelidae) of the Rila Mountain (Bulgaria). Fauna and Beron P. 1969. Sur les éléments boréo-alpins de la faune bulgare. Zoogeography. – Travaux scientifiques de l’Université de ‘Bulletin de l’Institut de Zoologie et Musée, Sofia, 30: , Animalia, 40(6): 97-114. 115-132. Gruev B. 2006. The leaf beetles (Coleoptera: Chrysomelidae) of Beron P. 1985. On the cave fauna of the Greek islands of Santorin the Pirin Mountain (Bulgaria). – Historia naturalis bulga- and Iraklia, with preliminary description of a new Pseu- rica, 17: 51-79. doscorpion. – Grottes bulgares, Sofia, 3: 64-71. Guéorguiev V. 1988. Coleopters (Coleoptera) in the orophyte zone Beron P. 1999. Biodiversity of the High Mountain Terrestrial Fau- of the Pirin Mountain. – Fauna of Southwestern Bulgaria, Ed. na in Bulgaria. – Historia naturalis bulgarica, 10: 13-33. BAN, Sofia, 2: 74-87 (Bulg., summ. Russ., Engl.). Beron P. 2008. High-mountain Isopoda Oniscidea, Arachnida Georgiev (= Guéorguiev) V. 1990. Coleoptera (Insecta, Co- and Myriapoda in the Old World. – Bureschiana, Sofia, 1: leoptera) from the orophyte zone of Vitoša. – Fauna of Pensoft & Nat. Mus. Natur. Hist. Sofia, 556 pp. Southwestern Bulgaria. Part 3: 134 – 145 (Bulg., summ. Beron P., Beshkov V., Popov V., Vassilev M., Pandurska R., Russ., Engl.). Ivanova T. 2000a. Biodiversity of Small Vertebrates (Pisces, Guéorguiev V.B., Sakalian V.P., Guéorguiev B.V. 1997. Amphibia, Reptilia, Mammalia – Insectivora, Chiroptera, Biogeography of the endemic Balkan Ground – beetles Lagomorpha and Rodentia) in the Rila National Park. – In: (Coleoptera: Carabidae) in Bulgaria. – Pensoft Series Faun. Sakalian M. (ed.): Biological Diversity of the Rila National No 6, Pensoft Publ., Sofia: V+73 p. Park, Sofia, 333-360. Hanák V., Horácek I. 1986. Zur Südgrenze des Areals von Ep- Beron P., Beshkov V., Popov V., Vassilev M., Pandurska R., tesicus nilssoni (Chiroptera: Vespertilionidae). – Annalen Ivanova T. 2000b. Biodiversity of Small Vertebrates (Pisces, Naturhistorischen Museum Wien, 88 – 89: 377 – 388. Amphibia, Reptilia, Mammalia – Insectivora, Chiroptera, Hubenov Z. 2015. Two-winged insects (Insecta: Diptera) of Pirin. Lagomorpha and Rodentia) in the Central Balkan National – Historia naturalis bulgarica, 21: 215-256. Park. – In: Sakalian M.(ed.): Biological Diversity of the Hubenov Z., Beshovski V., Josifov M., Popov A., Kumanski K., Central Balkan National Park, Sofia, 363-391. Sakalian V., Abadjiev S., Vidinova Y., Lyubomirov T. 2000a. Božilova E. 1995. The upper forest limit in the Rila Mts. in Entomofaunistic Diversity of the Rila National Park. – In: postglacial time – palaeoecological evidence from pollen Sakalian M. (Ed.) Biological Diversity of the Rila National analysis‚ macrofossil plant remains and C14 dating. – In: Park, Sofia: 285-331. Advances in holocene palaeoecology in Bulgaria‚ Pensoft Hubenov Z., Beshovski V., Josifov M., Popov A., Kumanski Publ.‚ Sofia – Moscow, 1-8. K., Sakalian V., Abadjiev S., Vidinova Y., Lyubomirov Brignoli P.M. 1980. La valeur biogéographique des Araignées T. 2000b. Entomofaunistic Diversity of the Central Balkan cavernicoles. – 8. Internat. Arachnologen – Kongr., Wien, National Park. – In: Sakalian M. (ed.): Biological Diversity 1980: 427-432. of the Central Balkan National Park, Sofia: 319-362. Buresch I., Arndt W. 1926. Die glazialrelicte stellenden Tierarten Josifov M. 1997. Heteroptera. – In: Sakalian V. (Ed.). Endemic and Bulgariens und Mazedoniens. – Zeitschrift für Morphologie Relict Insects in the Pirin National Park‚ Bulgaria. Pensoft und Ökologie der Tiere, 5 (3): 381-405. Publ.‚ Sofia – Moscow‚ 31-42. Darlington P.J. 1957. Zoogeography: The Geographical Distri- Kumanski K. 1997. Trichoptera. – In: Sakalian V. (Ed.). Endemic bution of Animals. Wiley, New York. and Relict Insects in the Pirin National Park‚ Bulgaria. Deltschev Ch. 1990. The high-altitude spiders (Araneae) in Pensoft Publ.‚ Sofia – Moscow‚ 59-68. the Pirin Mountains, Bulgaria. – Acta Zoologica Fennica, [Lopatin I.K.] Лопатин И.К. 1980. Основы зоогеографии 190: 111-115. [Fondaments of the Zoogeography]. Minsk, “Visshaya Deltschev Ch., Beron P., Blagoev G., Golemansky V., Najde- shkola”, 199 pp. nov V., Peneva V., Stoev P., Todorov M., Hubenov Z. Martens J. 1972. Ausobskya athos, der erste Krallenweberknechte 2000a. Faunistic Diversity of Invertebrates (non Insecta) of aus Griechenland (Opiliones: Phalangodidae). Mit Be- the Rila National Park. – In: Sakalian M. (ed.): Biological merkungen zum Familien-Gliederung der europäischen Diversity of the Rila National Park, Sofia, 249-284. Laniatores. – Senckenbergiana biologica, 53(5/6): 431-440. Deltschev Ch., Beron P., Blagoev G., Golemansky V., Na- Palamarev E. 1982. Principal stages in development of the flora jdenov V., Peneva V., Stoev P., Todorov M., Hubenov and vegetation. .Physical geography. Z. 2000b. Faunistic Diversity of Invertebrates (non Insecta) Publ. House of Bulg. Acad. of Sciences, Sofia., pp. 413-420. of the Central Balkan National Park. – In: Sakalian M. (In Bulgarian with English summary). (ed.): Biological Diversity of the Central Balkan National Peschev G.P. 1987. Endemism among the Orthopterans (Or- Park, Sofia, 289-317. thoptera) in the Bulgarian mountains. – In: Baccetti B. 118 Petar Beron

(Ed.). Evolutionary biology of orthopteroid insects. Ellis Делчев Хр. 2005. Биоразнообразието на Национален Horwood Limited, Chichester: 414-417. парк Пирин. Българска фондация Биоразнообразие Pešev G. 1990. Biogeographical significance of the disjunctive [The Biodiversity of Pirin National Park. Bulgarian Founda- distribution on mountaneous Orthoptera in Bulgaria. – tion Bioraznoobrazie], Sofia, 95 p. (In Bulgarian). Acta zoologica bulgarica, 39: 16-24. Sakalian V. 1997. General Results and Discussion. – In: Sakalian Pomorski R.J. 2006. Revision of the genus Onychiuroides Bagnall, V. (ed.): Endemic and Relict Insects in the Pirin National 1948 (Collembola: Onychiuridae: Onychiurinae). – Park‚ Bulgaria. Pensoft Publ.‚ Sofia–Moscow‚ 78-91. Systematics and Evolution, 37 (1): 39-69. Sakalian V., Guéorguiev B. 1997. Coleoptera. – In: Sakalian Popov A. 1997a. Orthoptera. – In: Sakalian V. (Ed.). Endemic and V. (ed.): Endemic and Relict Insects in the Pirin National relict insects in the Pirin National Park, Bulgaria. – Pensoft Park, Bulgaria. Pesoft Publ., Sofia–Moscow, 45-58. Publ., Sofia – Moskow, 12-24. Shurulinkov P., Stoyanov G., Tzvetkov P., Vultchev K., Kol- Popov A. 1997b. Neuroptera. – ibidem, 43-44. chagov R., Ilieva M. 2003. Distribution and abundance of Popov A. 2007. Fauna and Zoogeography of the Orthopterid Tengmalm’s Owl Aegolius funereus on Mount Pirin, south- Insects (Embioptera, Dermaptera, Mantodea, Blattodea, west Bulgaria. – Sandgrouse, 25 (2): 103-117. Isoptera, and Orthoptera) in Bulgaria. – In: V. Fet & A. Stoev P. 2007. Fauna and Zoogeography of Myriapoda in Bulgaria. Popov (Eds.) Biogeography and Ecology in Bulgaria, – In: V. Fet & A.Popov (Eds.), Biogeography and Ecology Springer, Monographiae Biologicae, 82: 233-295. of Bulgaria, 379-404. Popov A., Deltshev Ch., Hubenov Z., Beshovski V., Dobrev Varga Z. 1995. Isolates of Arctic-alpine and Alpine Lepidoptera D., Gueorguiev B. 2000. Invertebrate fauna. – In: Popov in SE Europe. – Proceedings of the EIS Colloquium, Hel- A., T. Meshinev, (eds.): High mountain treeless zone of sinki, p. 140 -151. the Central Balkan National Park. Biological diversity and Wangerin W. 1912. Über den Reliktbegriff und die Konstanz der problems of conservation. Sofia, BSBCP, 339-416. Pflanzenstandorte. – Festschrift des Preußischen Botanis- [Popov V., Dimova D., Deltshev Hr.] Попов В., Димова Д., chen Vereins, 158-184.

Author’s address:

Petar Beron, National Museum of Natural History, 1 Tsar Osvoboditel Blvd, Sofia 1000, Bulgaria. E-mail: [email protected]

Реликти и ендемити във високопланинската фауна на България

Петър Берон

(Резюме) Разглежданата тук област (над 1900 м) обхваща около 1.37% от българската територия в осем планини, от които Рила и Пирин надхвърлят 2400 м. Там са съсредоточени и повечето ендемични и реликтни видове животни (изброени по-горе). Има както тесни локални ендемити, така и балкански такива. Осоено добре са застъпени групите Araneae, Acari, Orthoptera, Coleoptera, Heteroptera, Lepidoptera.