Coleoptera: Carabidae) from the Eastern Rhodope Mts

FORESTRY IDEAS, 2020, vol. 26, No 1 (59): 77–96

ECOLOGICAL, FAUNISTIC AND ZOOGEOGRAPHICAL NOTES ON THE GROUND BEETLES (COLEOPTERA: CARABIDAE) FROM THE EASTERN RHODOPE MTS. OF BULGARIA

Teodora Marius Teofilova1* and Nikolay Dyankov Kodzhabashev2 1Institute of Biodiversity and Ecosystem Research (IBER), Bulgarian Academy of Sciences (BAS), 1, Tsar Osvoboditel Blvd., 1000 Sofia, Bulgaria. *Е-mail: [email protected] 2Department of Hunting and Game Management, Faculty of Forestry, University of Forestry, 10, Kliment Ohridski Blvd., 1797 Sofia, Bulgaria. Е-mail: [email protected]

Received: 20 January 2020 Accepted: 14 March 2020

Abstract The present study adds new data to the list of Carabidae species from the Eastern Rhodope Mts. of Bulgaria, and reveals some patterns of the species composition from zoogeographic, ecological and conservation point of view. A total of 5915 specimens of 138 species belonging to 50 genera and 19 tribes are found, including 13 endemic species and subspecies, one relict and some rare and stenotopic species. This represents, respectively, 18.5 % of all established for the Bulgarian carabid fauna species and 40 % of the genera. Eleven genera (Acupalpus La- treille, 1829, Anchomenus Bonelli, 1810, Anisodactylus Dejean, 1829, Carterus Dejean & Boisdu- val, 1829, Demetrias Bonelli, 1810, Elaphrus Fabricius, 1775, Gynandromorphus Dejean, 1829, Olisthopus Dejean, 1828, Parophonus Ganglbauer, 1891, Tachys Dejean, 1821 and Tachyura Motschulsky, 1862) are new for the Eastern Rhodope Mts. Two of the genera (Carterus and Gy- nandromorphus) are also new for the whole Rhodope Mts. The subgenus Philochthus Stephens, 1828 is new for the Eastern Rhodopes and subgenus Phyla Motschulsky, 1844 is new for the whole Rhodope Mts. Among the species, 60 species are new for the Eastern Rhodopes, and 24 species of them are new for the whole Rhodope Mts. The richest tribes are Harpalini (46 species), Amarini (14 species), Lebiini (11 species) and Carabini and Sphodrini (10 species each). Pteros- tichini and Bembidiini are represented with nine species each. Zoogeographical analysis shows that the Mediterranean faunal type prevailed. In relation to their life forms, zoophagous carabids predominate, presented by 78 species (56.5 %). Mixophyitophagous are 60 species (43.5 %). Most of the ground beetles are mesoxerophilous (51 species, 37 %). Key words: carabids, endemism, life forms, zoogeography, new data.

Introduction the territory of the mountain (Guéorguiev 2004), 146 species (20 % of all Bulgarian From the beginning of the 20th century until Carabidae species) occur in the Bulgarian now, about 40 studies concerning the ca- part of the Eastern Rhodopes. So far, 156 rabid fauna of the Eastern Rhodope Moun- species (21 % of all Bulgarian Carabidae tains have been published. According to species) are surely known, 54 species the latest survey of the carabid fauna on are likely to be found there and another 78 T. Teofilova and N. Kodzhabashev seven species are probably doubtfully the valley of the Krumovitsa River as an established for that region (Teofilova and Important Bird Area. Guéorguiev unpublished results). Consid- The aim of the present study is to add ering the degree of research, dimensions new data to the list of Carabidae species of the region and the habitat diversity, from the Eastern Rhodope Mts. of Bul- which it suggests, it can be argued that the garia, and to reveal some patterns of the expected real number of carabid species species composition from zoogeographic, is about 250, or one-third of those known ecological and conservation point of view. from Bulgaria (Guéorguiev 2004, Teofilo- va and Guéorguiev unpublished results). The Eastern Rhodopes occupy the Material and Methods lower part of the Rhodope Mts. A variety of habitats occur on their territory: oak for- Field work was carried out in two periods: ests, beech forests (limited to the south- 2005–2006 and 2019, in parallel with the ernmost parts of the mountain), conifer- conduction of monitoring researches in ous and broadleaved plantations, bushes, some target areas. Ground beetles were riparian woods and bushes, grasslands, collected by hand picking, and terrestrial pastures, rocky habitats, inland standing ‘pitfall’ traps made of plastic bottles, bur- and running surface waters, as well as ied at the level of the ground surface. Ta- many artificial landscapes – villages, cha- ble 1 contains detailed information about lets, huts, etc. This fact is a prerequisite the individual localities, periods and col- for the occurrence of species from differ- lectors of the material. ent faunistic complexes, which are more Collected material was deposited in or less attached to a particular setting of the Carabidae collection of the Institute natural conditions. of Biodiversity and Ecosystem Research The Eastern Rhodope Mts. are among (Bulgarian Academy of Sciences, Sofia). the sites of highest biodiversity in Bulgar- According to their zoogeographical ia. Within its territory fall several protected belonging, the ground beetles were sep- areas with different designation – ‘Valchi arated in zoogeographical categories and Dol’ Nature Reserve, four Managed Re- complexes according to Kryzhanovskij serves (‘Zhenda’, ‘Chamlaka’, ‘Boraka’, (1965, 1983, 2002), Vigna Taglianti et al. ‘Borovets’), 20 protected sites and sev- (1999) and Kodzhabashev and Penev eral natural monuments. The mountain (2006), with some changes and adjust- overlaps with some Natura 2000 zones: ments, made by the authors of the pres- seven SPA (BG0002012, BG0002013, ent study. BG0002014, BG0002019, BG0002071, According to their ecological require- BG0002073, BG0002106), included in ments in terms of humidity, established EU Directive 79/409, and three SAC carabid species were divided into six cat- (BG0000372, BG0001031, BG0001032) egories: hygrophilous, mesohygrophilous, of Directive 92/43. In 1998, the territory mesophilous, mesoxerophilous, xerophil- of the Arda Valley was determined as a ous and eurybionts. CORINE site, due to its European signif- Categorization of the species in re- icance in protection of rare and endan- spect of their life forms follows the classi- gered habitats, plants and animals. In fication of Sharova (1981). The following 1997, BirdLife International pronounced codes were used: Life form class 1. Zoo- Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 79 NK Hand picking 6.VI.2005 20. V.2006 І .IX–20.X.2005; 20 .IX.2006 18 .IX, 20 VIII– 20. .VIII– 3 [6, formaldehyde, NK] [6, formaldehyde, NK] [6, formaldehyde, NK] [10 tree traps, DC&ML] [12, formaldehyde, NK] [12, formaldehyde, NK] VIII, 15.IV–18.VI, 18.VI–VIII.2006 15.IV–18.VI, 18.VI–VIII.2006 VIII.2006 [6, formaldehyde, NK] .IV–6.VI, 6.VI–7.VII, 7.VII–12.VIII.2005; 17.VI.2006 [5, formaldehyde, DC&ML] .IV–6.VI, 1 15.IV–17.VI.2006 [5 tree traps DC&ML] 17.VI.2006 [10, formaldehyde, DC&ML] 20.X.2005; 15.IV–18.VI, 18.VI–VIII.2006 23.IV–5.VI, 14.VIII–6.IX.2005; 16. 25 .IV–6.VI, 6.VI–7.VII, 7.VII–12.VIII, 12.VIII–18. .IV–6.VI, 6.VI–7.VII, 7.VII–12.VIII, 12.VIII–21. .IV–6.VI, 6.VI–7.VII, 7.VII–13.VIII, 13.VIII–21. 25 20.X.2005–15.IV.2006; 15.IV–18.VI, 18.VI – 20.X.2005–15.IV.2006; IX, 18.IX–20.X.2005; 20.X.2005–15.IV.2006; IX, 18.IX–20.X.2005; 20.X.2005–15.IV.2006; IX, 21.IX–20.X.2005; 20.X.2005–15.IV.2006; IX, 21.IX–20.X.2005; 20.X.2005–15.IV.2006; 13.VIII–6.IX.2005 [5, formaldehyde, DC&ML] 5.VI–7.VII, 7.VII–12.VIII, 12.VIII–18.IX, 18.IX– 23.IV–5.VI, 4.VI–3.VII, 6.XI–21.X.2005; 16.IV– 20.X.2005–15.IV.2006; 15.IV–18.VI, 18. VI– 20.X.2005–15.IV.2006; 6.IX–21.X.2005; 21.X.2005–14.IV.2006; 15.IV– 6.IX–21.X.2005; 21.X.2005–14.IV.2006; 25 25 25 Traps [number of traps, fixative, collector(s)] Traps 15.IV–18.VI, 18. VI – 20.VIII, 20.VIII–18.XII.2006 - L.) L.) plan sp.) forest J.F.Arnold) J.F.Arnold) 2005–2006 Quercus tation Habitat Dere Site plantation Pinus nigra Robinia pseudoacacia Pinus sylvestris grazed pasture black pine forest Hill near Pobeda vill. Xerothermic oak forest border of the Valchi Dol Reserve border of the Valchi vill. in the vicinity of Kayadzhik Black pine ( Xerothermic oak forest and actively Scots pine ( Xerothermic oak ( Actively grazed pasture near Pobeda Xerothermic oak forest at the eastern Kayadzhik Dere river, riverbank forest Kayadzhik Dere river, plantation, probably mixed with natural Black locust ( ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ’’ ’’ ’’ ’’ ’’ ’’ ’’ - 9 5‘‘ 1 4 7 9 0 18 1 13 15 41 46 14 ‘2 ‘ ‘ ‘ ‘ ‘ ’13’’ ‘2 ‘ 6‘14‘‘ 5‘5 9 9 9 9 26 2 2 26 26 26 38 dinates GPS coor E 25°3 E 25°3 E 25°3 E 25°3 E 25°38‘3 E 25°38‘ E 25° E 25°38’32’’ E 25°38‘3 N 41°35’52’’ N 41° N 41° N 41° N 41° N 41° N 41° N 41°26‘2 N 41°35‘46‘‘

270 390 480 480 280 290 280 395 410 a.s.l. de, m Altitu- - Table 1. List, average altitude and description of the sampling sites, methods dates visiting. Table Peak Locality Chal vill. Ada Tepe Tepe Ada Kechi Kaya near the top Ada Tepe Hill Tepe Ada Hill Tepe Ada Hill Tepe Ada Hill Tepe Ada Hill Tepe Ada Ada Tepe Hill, Tepe Ada W Potocharka vill., near Stari nitsa vill., near above Potoch S Krumovgrad, S Krumovgrad, S Krumovgrad, S Krumovgrad, S Krumovgrad, S Krumovgrad, P_O AT_A Code AT_O P_O1 AT_M KD_R AT_Ps AT_Pn AT_O1 80 T. Teofilova and N. Kodzhabashev NK Hand picking 6.VI.2005 18. –18. –20. VI .2005; VIII ] X . VIII– .2005–15. NK X , 18. 20 , 20. , ] –20. VI .2005 IX VIII NK , .2006 –18. , 18. –20. XII IV .2005; 20. IX VI X –18. –18. formaldehyde , 18. –20. VIII VI VIII IX .2006; 15. formaldehyde , 20. IV –18. , 12. , 21. [6, formaldehyde, NK] [12, [12, formaldehyde, NK] IV formaldehyde, DC&ML] VIII .IV–6.VI, 7.VII–12.VIII IX [5, formaldehyde, DC&ML] .2006 [12, VIII 22.IV–6.VI, 6.IX–21.X.2005 [11, formaldehyde, DC&ML] [11, 22.X.2005; 14.IV–19.VI.2006 25 XII.2006 [6, formaldehyde, NK] XII –21. IX.2005 [5, formaldehyde, DC&ML] –12. .2005–15. 19.VI.2006 [5, formaldehyde, DC&ML] X VIII VII 22.IV–5.VI, 5.VI–3.VII, 3.VII–15.VIII, 7.IX– .IV–6.VI, 6.VI–7.VII, 7.VII–12.VIII, 12.VIII–18. .IV–6.VI, 6.VI–7.VII, 7.VII–12.VIII, 12.VIII–21. 6.IX–21.X.2005 [5, formaldehyde, DC&ML] .2006; 15. 6.IX.2005–16.ІV.2006 [5 tree traps DC&ML] 6.IX.2005–16.ІV.2006 6.IX.2005–16.ІV.2006 [5 tree traps, DC&ML] 6.IX.2005–16.ІV.2006 IX, 21.IX–20.X.2005, 20.X.2005–15.IV.2006; IX, 21.IX–20.X.2005, 20.X.2005–15.IV.2006; 7. IX, 18.IX–20.X.2005; 15.IV–18.VI, 23.IV–5.VI, 5.VI–3.VII, 3.VII–14.VIII, 14.VIII–6. 5.VI–4.VII, 4.VII–15.VIII, 15.VIII–7.IX.2005 [10, 20. 13. IV 15.IV–18.VI, 18. VI–20.VIII, 20.VIII–18.XII.2006 25 25 Traps [number of traps, fixative, collector(s)] Traps 4.VI–3.VII, 3.VII–14.VIII, 6.IX–21.X.2005; 16.IV– -

- - on Tilia spp.), L.) plan brook

near dry pas epiphytes

Populus ture Habitat sp.) and sp.) forest Pinus sylvestris Salix the Arda River coast the terfall (Kodzha dere) Xerothermic oak forest Xerothermic oak forest Xerothermic oak forest mixed with sprout linden ( near never failing vegetation on sandy silt

Riverbank forest near Arda river Riverbank forest near Arda River coast, riverbank forest Very old mixed mesophylous forest Very willows ( Grouping of poplars ( tation Scots pine ( Xerothermic oak forest Arda River coast with natural riverine Xerothermic scrub near gully with wa ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ’’ ’’ ’’ ’’ ’’ ’’ - 8 2 5 6 3 8 2’’ 6’’ 0’’ 2’’ 35 34 33 01 34 18 19 2 2 2 2 40 51 ’31’’ ‘2 ‘ ‘2 ‘09‘‘ ‘ ‘ ‘ ‘ ‘5 ‘ ‘ ‘ ‘ ‘ ‘ ‘ ‘ 6 6 6 8 9 8 8 9 8 6 51 52 52 52 52 52 52 52 dinates GPS coor E 25°38’15’’ E 25°38‘3 E 25°38‘3 E 25° E 25° E 25° E 25° E 25° E 25° E 25° E 25° N 41°3 N 41°3 N 41°3 N 41°3 N 41°3 N 41°3 N 41°3 N 41°3 N 41°3 N 41°3 N 41°35‘5

5 195 165 162 13 135 200 220 145 190 210 420 a.s.l. de, m Altitu- - - vill. vill. vill. vill. vo vill. nets vill. Locality 2 km NE 2 km NE 2 km NE NE Studen near Studen Kladenets vill. Kladenets vill. NW Madzharo SE Madzharovo SE Madzharovo NE Madzharovo NE Madzharovo Madzharovo vill. Madzharovo vill. Madzharovo vill. E Studen Klade M_R M_O M_M Code M_Ps P_SK SK_R M_R1 M_O1 M_O2 M_O3 SK_R1 Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 81 Hand picking 17.VI., TT 17.VI., TT 17.VI., TT 15.VI., TT 18.VI., 17.VI–29.VII.2019 17.VI–29.VII.2019 15.VI–31.VII.2019 16.VI–30.VII.2019 18.VI–30.VII.2019 [5 tree traps DC&ML] [12, formaldehyde, NK] [12, formaldehyde, NK] ІV.2006, 14.ІV–19.VI.2006 ІV.2006, [5, formaldehyde, DC&ML] 5.VII–15.VIII, 15.VIII–7.IX, 7.IX–22.X.2005 [5, salt-vinegar saturated solution, TT&KD] [5, salt-vinegar saturated solution, TT&KD] [5, salt-vinegar saturated solution, TT&KD] [8, salt-vinegar saturated solution, .IV–5.VI, 5.VI–7.VII, 7.VII–13.VIII, 13.VIII–21. [10, salt-vinegar saturated solution, TT&KD] [10, salt-vinegar saturated solution, TT&KD] [12, salt-vinegar saturated solution, IX, 21.IX–20.X.2005; 20.X.2005–15.IV.2006; IX, 21.IX–20.X.2005; 20.X.2005–15.IV.2006; 5.VII–15.VIII, 7.IX–22.X.2005; 22.X.2005–14. 25.IV–6.VI, 6.VI–3.VII, 3.VII–14.VIII, 14.VIII–7. 15.IV–18.VI, 18.VI–20.VIII, 20.VIII–18.XII.2006 IX., 7.IX–22.X.2005 [5, formaldehyde, DC&ML] 15.IV–18.VI, 18. VI–20.VIII, 20.VIII–18.XII.2006 25 Traps [number of traps, fixative, collector(s)] Traps

- -

L.) Mill.) on Crataegus Quercus 2019 Rosa canina Ten.), black pine Ten.), Cotinus coggygria forest Scop.) glades Habitat slag-depot pine forests Willd.) forest with grazed Xerothermic woods Paliurus spina-christi well drained eroded terrain Lawn at the place of a recycled Sparse pubescent oak ( joining well preserved oak forests Quercus frainetto Forest habitat with Hungarian oak and autumn, near xerophilous oak groups of oak trees and bushes and smoke tree ( ( Abandoned pasture with Jerusalem thorn ( river with rich vegetation) and Yuren Yuren river with rich vegetation) and spp.) and dog rose ( pubescens Along the Arpa Dere River (full-water Along the Vast meadow with bushes and single Vast dles and rich vegetation), the ad Dere River (almost dry gully with pud Actively grazed pasture with scattered trees and adjoining pond, oak Jerusalem thorn, hawthorn ( Pasture with shrubs, grazed in summer ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ‘‘ ’’ ’’ ’’ ’’ ’’ ’’ ’’ ’’ ’’ - 7 5‘‘ 4 3 8 38 4 17 07 04 43 13 18 36 43 12 02 04 ‘ ‘2 ‘ ‘ ‘ ‘ ‘ ‘ ‘ ‘2 ‘3 ‘2 ‘ ‘ ‘ ‘ ‘ 7 9 0 0 0 40 24 26 52 55 49 49 28 28 29 31 55 ° ° ° ° ° 5 5 5 5 5 dinates GPS coor E 25° E 25° E 25° E 25° E 2 E 2 E 2 E 2 E 2 N 41°3 N 41°3 N 41° N 41°40‘43‘‘ N 41°3 N 41°3 N 41°3 N 41° N 41°

210 180 320 350 612 570 600 430 286 a.s.l. de, m Altitu- - vill. vill. vill. vill. vanak Locality 1 km S Dolni W Ribino vill. NW Obichnik NW Obichnik Glavanak vill. SE Dolni Gla SW Borislavtsi SE Rozino vill. N Obichnik vill. SE Madzharovo B_S O_P M_D O_O O_m Code R_AY Ru_R B_DG B_DG1 82 T. Teofilova and N. Kodzhabashev

phagous. Life form subclass: 1.1 – Phy- tobios; 1.2 – Epigeobios; 1.3 – Strato- TT Hand 31.VII.,

picking bios; 1.4 – Geobios. Life form groups: 19.VI., TT 19.VI., TT 18.VI., 1.1.2 – stem-dwelling hortobionts; 1.1.3 – leaf-dwelling dendrohortobionts; 1.2.2 – large walking epigeobionts; 1.2.2(1) – large walking dendroepigeobionts; 1.2.3 – running epigeobionts; 1.2.4 – flying epigeobionts; 1.3(1) – series crevice-dwelling stratobionts; 1.3(1).1 – surface & litter-dwelling; 1.3(1).2 – litter-dwelling; 1.3(1).3 – litter & crevice-dwelling; 1.3(1).4 – endogeobionts; solution, TT&KD] solution, TT&KD] solution, 1.3(1).5 – litter & bark-dwelling; 1.3(1).6 – bothrobionts; 1.3(2) – series digging stratobionts; 1.3(2).1 – litter & soil-dwelling; 1.3(2).2 – litter & crevice-dwelling; 19.VI–30.VII.2019 [6, salt-vinegar saturated 19.VI–31.VII.2019 [6, salt-vinegar saturated

Traps [number of traps, fixative, collector(s)] Traps 1.4.2(1) – small digging geobionts. Life form class 2. Mixophytophagous. Life -

form subclass: 2.1 – Stratobios; 2.2 – Stratohortobios; 2.3 – Geohortobios. Pistacia Tamarix Life form groups: 2.1.1 – crevice-dwelling stratobionts; 2.2.1 – stratohortobi- spp.),

L., etc. onts; 2.3.1 – harpaloid geohortobionts; 2.3.1(1) – crevice-dwelling harpaloid stands Habitat movitsa City park Fraxinus geohortobionts; 2.3.2 – zabroid geohortobionts; 2.3.3 – dytomeoid geohortobi- terebinthus onts. The first figure in the index shows the class of life form, the second – the and pseudomaquis with Jerusalem sp.) and surrounding poplar–willow House yard near densely vegetated River branch with tamarisk ( Dry stony riverbed of the Byala Reka thorn, ashes ( shore of a tributary the river Kru Rocks near mesoxerophilic oak forest subclass, the third – the life form group. In brackets after the subclass the series ‘‘ ’’ - 34 02 ‘ ‘ is shown, when it exists. 25 54

° The systematic list follows Kry- 5 41°28’15’’ dinates zhanovskij et al. (1995). GPS coor E 25°53’33” N 41°26’00” E 2 E 25°39’41’’ E 25°24’16’’ N N 41°31’50’’ N 41°

365 197 265 263 a.s.l. de, m Altitu- Results and Discussion

During the field work were captured 5915 specimens of 138 species belong- vill. ing to 50 genera and 19 tribes of ground Locality Krumovgrad Momchilgrad beetles. This represents, respectively, SW Rozino vill. . NW Byalgradets 18.5 % of all established for Bulgarian

Note: Collectors: DC – Dragan Chobanov, ML – Mario Langourov, NK – Nikolay Kodzabashev, TT – Teodora Teofilova, KD – Krasimir – KD Teofilova, Teodora – TT Kodzabashev, Nikolay – NK Langourov, Mario – ML Chobanov, Dragan – DC Collectors: Note: carabid fauna species and 40 % of the Kr Mo Code R_RF B_BR genera (Teofilova and Guéorguiev un- Donchev Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 83 published results). During the first study part of the material was collected by hand (2005–2006) were captured 120 species and with tree traps, since most of these (5688 specimens), and another 18 spe- beetles are rarely found in the pitfall traps. cies (227 specimens) were added with the Genera most rich in species were Har- recent samplings (2019). The complete palus Latreille, 1802 (25 species), Amara check-list of the established species with Bonelli, 1810 (11 species) and Carabus zoogeographic category, life form and Linnaeus, 1758 (eight species). Genera ecological group in relation of the humidity Ophonus Dejean, 1821, Calathus Bonelli, is given in the Appendix. 1810 and Bembidion were represented by Our study presents eleven new genera six species each. for the Eastern Rhodopes (Acupalpus, New highest altitudes were found in Anchomenus, Anisodactylus, Carterus, the distribution of five species: Amara Demetrias, Elaphrus, Gynandromorphus, parvicollis Gebler, 1833 – so far known Olisthopus, Parophonus, Tachys and only from the Black Sea coast and East- Tachyura) Two of the genera (Carterus ern Danube Plain under 50 m a.s.l., now and Gynandromorphus) are also new for found in localities M_R and M_R1 (for the whole Rhodope Mts. The subgenera locality codes see Table 1), at 135 m; Philochthus and Phyla (both from the ge- Anisodactylus intermedius Dejean, 1829 nus Bembidion Latreille, 1802) are new – so far known only under 300 m a.s.l., for the Eastern Rhodopes, and Phyla is now found in locality P_O1, at 410 m; also new to the whole Rhodope Mts. Six- Brachinus brevicollis Motschulsky, 1844 – ty species are new for the Eastern Rho- so far known only under 300 m a.s.l., now dopes; 24 species of them are new for the found in locality B_DG, at 320 m; Calathus whole Rhodope Mts. The record of Clivina cinctus Motschulsky, 1850 – so far known fossor (Linnaeus, 1758) gives confirma- only under 300 m a.s.l., now found in lo- tion for the presence of this species in this cality AT_O, at 390 m; Poecilus anatolicus part of the mountain. All new species are (Chaudoir, 1850) – so far known only marked in the Appendix. under 400 m a.s.l., now found in locality The richest tribes were Harpalini (46 Ru_R, at 430 m. It is hard to claim if these species), Amarini (14 species), Lebiini (11 new findings are evidences for the real species) and Carabini and Sphodrini (10 movement of the species in height. They species each). Pterostichini and Bembidi- are probably resulting of the less knowl- ini tribes were represented with nine spe- edge about the ecology of these species cies each. Harpalini and Amarini tribes in- and their distribution in Bulgaria. clude mostly ecologically plastic carabids. We found some species with conser- Carabini and Pterostichini are typical for- vation value. Our study reports one Ter- est dwellers; most of them are stenotop- tiary relict (Myas chalybaeus (Palliardi, ic and any impact on the forest habitats 1825)) and 14 endemic species and sub- where they occur, also affects the struc- species, of which Balkan endemics are 10 ture of their communities. Sphodrini tribe taxa (two species and eight subspecies), includes both forest dwellers and open Bulgarian endemics are two taxa (one living species. The increased richness of species and one subspecies), and one the tribes Lebiini and Bembidiini could be species and one subspecies are subend- resulting from the fact that a significant emic (Table 2). 84 T. Teofilova and N. Kodzhabashev

Table 2. List of the endemic ground beetles found in the Eastern Rhodope Mts. in 2005– 2006 and 2019. Species/subspecies Level Tapinopterus balcanicus balcanicus Ganglbauer, 1891 Bulgarian Zabrus balcanicus Heyden, 1883 Bulgarian Carabus scabriusculus bulgarus Lapouge, 1908 Balkan Carabus montivagus montivagus Palliardi, 1825 Balkan Carabus wiedemanni wiedemanni Ménétriés, 1836 Balkan Carabus scabrosus scabrosus Olivier, 1790 Balkan Cychrus semigranosus balcanicus Hopffgarten, 1881 Balkan Trechus subnotatus subnotatus Dejean, 1831 Balkan Molops dilatatus Chaudoir, 1868 Balkan Laemostenus cimmerius weiratheri J. Müller, 1932 Balkan Zabrus graecus subtilis Schaum, 1862 Balkan Licinus graecus Apfelbeck, 1901 Balkan Myas chalybaeus (Palliardi, 1825) Balkan subendemic Platynus scrobiculatus purkynei Obenberger, 1917 Balkan subendemic Carabus intricatus Linnaeus, 1760 is the Mediterranean (sensu lato) complex included in the IUCN Red List as ‘Near prevailed (47 species, 34 %). It was fol- Threatened‘. Along with Calosoma syco- lowed by the European-Asiatic (35 spe- phanta (Linnaeus, 1758), it is included in cies, 25 %) and Northern Holarctic and the Annexes of CORINE and ESC Red European-Siberian complexes (31 spe- List. Carabus scabrosus Olivier 1790, is cies, 23 %). European complex consist- enlisted in the Red Data Book of Bulgaria ed of 17 species (12 %) and Endemic as ‘Vulnerable‘ (Golemanski et al. 2015). complex – eight species (6 %) (Fig. 1). Some rare and stenotopic species oc- Mediterranean species are distributed in cur in the studied region: Amara saphy- the so-called region of the ‘Ancient Med- rea Dejean, 1828, Calathus micropterus iterraneum‘ (Popov 1927, Kryzhanovskij (Duftschmid, 1812), Calosoma sycophan- 1965, 1983, 2002). European-Asiatic spe- ta, Lebia marginata (Geoffroy, 1785), cies ranges lie between the Eurosiberian Pedius inquinatus (Sturm, 1824), Platy- and Mediterranean zones. Northern Hol- nus scrobiculatus (Fabricius, 1801), Za- arctic and European-Siberian complex brus graecus Dejean, 1828, most of the includes species distributed mainly in the endemites. Some of the species (e.g. Car- northern regions of the Holarctic, mostly abus intricatus and Carabus scabrosus) in Europe and Siberia. European complex have become rare under the influence of includes mostly forest dwelling species anthropogenic pressures and changes in connected to the middle and southern part their primary habitats. Calosoma inquis- of Europe. itor (Linnaeus, 1758), Calosoma syco- Greatest number of species had the phanta and some of the Carabus species European-Central Asian, European-Near- are usually highly sensitive to chemical eastern, Palaearctic, Balkan-Neareastern agents, which affects their ranges and and Northmediterranean zoogeographic numbers (Huusela-Veistola 2000). categories (Table 3). The predominance Zoogeographical analysis showed that of the Mediterranean species is not typi- Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 85

Endemic Northern 6% Holarctic and Mediterranean European- 34% Siberian 23%

European 12%

Euroasiatic 25%

Fig. 1. Distribution of the carabid species among the zoogeographical complexes. cal for the other mountain regions in Bul- The specific geographic position and cli- garia (e.g. Kostova 2009, Teofilova 2016, matic conditions are a precondition for the Teofilova 2019b), but it is common for the development of respective vegetation and habitats near the Black Sea coast (Popov habitat types, determining the composi- and Krusteva 1999, Teofilova et al. 2015). tion of the carabid coenoses.

Table 3. Zoogeographical categories of the ground beetles in the Eastern Rhodope Mts. (on species level). Zoogeographical Species, Species, Code Zoogeographical element complex No % OLA Holarctic 5 4 Northern Holarctic PAL Palaearctic 14 10 and European- WPAL Western Palaearctic 6 4 Siberian E-SI European-Siberian 5 4 E-WSI European and West Siberian 1 1 EUR European 5 4 CE-PAS Central European and Neareastern 6 4 European Central and Eastern European and CEE-PA 3 2 Neareastern CE-E Central and Eastern European 3 2 Euroasiatic steppe and forest-steppe E-AS 2 1 complex Euroasiatic E-CAS European and Central Asian 16 12 E-PAS European-Neareastern 15 11 B-CAS Balkan and Central Asian 2 1 86 T. Teofilova and N. Kodzhabashev

Zoogeographical Species, Species, Code Zoogeographical element complex No % E-CA-M European-Central Asian-Mediterranean 9 7 E-PA-M European-Neareastern-Mediterranean 9 7 MED-PA Mediterranean-Neareastern 2 1 MED Mediterranean 3 2 Mediterranean EMED Eastmediterranean 1 1 NM-CAS Northmediterranean-Central Asian 1 1 NMED Northmediterranean 10 7 P-SMED Pontic-Submediterranean 1 1 B-PAS Balkan-Neareastern 11 8 Balkan subendemic (+ Balkan- BAL-K 3 2 Carpathian) Endemic BAL Balkan endemic 4 3 BGE Bulgarian endemic 1 1

The ground beetles established dur- phytophagous harpaloid geohortobionts ing the study related to two classes of life from Class Mixophytophagous (32 spe- forms proposed by Sharova (1981), with cies, 23 %), followed by the crevice-dwell- a predominance of class Zoophagous, ing surface & litter stratobionts (20 spe- presented by 78 species (56.5 %). Mixo- cies, 14 %) from Class Zoophagous (figs phyitophagous were 60 species (43.5 %). 2 and 3). This ratio is most similar to that in the The harpaloid geohortobionts were meadow steppes from the Forest-steppe also dominating in the Western Rhodope zone of Eurasia (Sharova 1981), habitats Mts. (Teofilova 2018, Teofilova 2019a) near Cape Emine at the Black Sea coast and Vrachanska Planina Mts. (Teofilova (Teofilova et al. 2015) and unvegetated, 2019b). Surface & litter dwelling stratobi- sparsely vegetated, and bryophyte- or li- onts are most of the Bembidion and Ago- chen-vegetated cliffs, rock faces and rock num Bonelli, 1810 species, Notiophilus pavements in the inland areas in Bulgaria Duméril, 1806, Chlaenius Bonelli, 1810. (Teofilova 2019c), and differs from the es- They were also found as the most species tablished in other mountain regions in Bul- rich zoophagous category in the Western garia (e.g. Teofilova 2016, Teofilova 2018, Rhodope Mts. (Teofilova 2018, Teofilova Teofilova 2019b). 2019a) and Vrachanska Planina Mts. There were 21 life forms of ground bee- (Teofilova 2019b). tles – 15 zoophagous and six mixophy- In the studied part of the mountain, tophagous. Zoophagous life form groups according to their ecological preferenc- are normally more numerous, especially es, mesoxerophilous (51 species, 36 %) in stable ecosystems (Sharova 1981) and ground beetles prevailed (Fig. 4). Meso- in forest regions, as it was found in ‘Lesh- philous were 37 species (27 %), hygro- nitsa’ Reserve (Teofilova 2016), Vrachan- philous were 18 species (13 %), meso- ska Planina Mts. (Teofilova 2019b) and hygrophilous were 15 species (11 %), 12 in the Western Rhodope Mts. (Teofilova species were xerophilous (9 %), and only 2018, Teofilova 2019a). The largest share five species (4 %) were eurytopic. of the species belonged to the specialised These results showed the predom- Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 87

2.3.2. 2.3.3. 1.1.2. 1.1.3. 1.2.2. 1.2.2(1) 4% 1% 1% 1% 7% 1% 1.2.3. 2.3.1(1) 2% 1% 1.2.4. 2.3.1. 1% 23% 1.3(1).1 14%

1.3(1).2 2.2.1. 9% 9% 2.1.1. 4% 1.3(1).3 1.3(1).4 9% 1.4.2(1) 1.3(2).2 1.3(2).1 1.3(1).6 1.3(1).5 1% 1% 1% 6% 2% 1%

Fig. 2. Life forms of the ground beetles from the Eastern Rhodope Mts. Note: Explanations of the codes are given in the Material and Methods section.

Fig. 3. Proportions of the subclasses of life forms in the carabid complex. Note: I – class Zoophaga, II – class Mixophytophaga. 88 T. Teofilova and N. Kodzhabashev

E H X 4% 13% 9% MH 11%

MX 36% M 27%

Fig. 4. Humidity preferences of the carabids from the Eastern Rhodope Mts. (number of species). Note: H – hygrophilous, MH – mesohygrophilous; M – mesophilous, MX – mesoxerophilous, X – xerophilous, E – eurybiont. inantly mesoxerophilous nature of the accordance with the life forms structure, habitats, in contrast of the predominant- and the preference of the carabids for ly mesophilous conditions in the Western more mesoxerophilous habitats. part of the Rhodope Mts. (Teofilova 2018, The region of the Eastern Rhodope Teofilova 2019a) and Vrachanska Planina Mts. is treasuring specific assemblages Mts. (Teofilova 2019b), probably resulting and species with conservation signifi- from the typical forest-mountain condition cance, and the preserving of their char- in those territories. acteristic habitats is a keystone for their conservation, especially in that favoured mining prospecting region. Conclusions

The large number of the found new spe- Acknowledgments cies shows the great variety of the ground beetle fauna in the Eastern Rhodope Mts. Authors express their gratitude to Dr. As the studied area provides various and Dragan Chobanov, Dr. Mario Langourov diverse habitats, and considering the gen- and Krasimir Donchev for their help with eral richness of the carabids and the lack the collection of the material. We also of targeted methodical studies, it is quite thank to both reviewers for their fast re- possible that the carabid fauna is even views. more diverse. For general conclusions, however, more localities have to be ex- amined systematically. References Zoogeographical analysis shows a predominantly Mediterranean nature of Golemanski V., Beron P., Popov A., Popov V., the established carabid fauna, which is in Beshkov V., Zhivkov M., Spasov N., Michev Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 89

T., Delchev C. (Eds) 2015. Red Data Book animal assemblages from two landscapes of Republic of Bulgaria. Vol. 2. Animals. of the Bulgarian Black Sea Coast: relation- BAS & MOEW, Sofia. 383 p. ship to environmental gradients, assem- Guéorguiev B.V. 2004. Adephagous beetles blage structure and biodiversity. Part I: and some staphyliniform beetles (Insec- Ground beetles. Acta Zoologica Bulgarica ta: Coleoptera) in the Eastern Rhodopes 51(1): 81–114. (Bulgaria and Greece). In: Beron P., Popov Sharova I. 1981. Life forms of carabids. Nauka, A. (Eds). Biodiversity of Bulgaria. 2. Biodi- Moskow. 360 p. (in Russian). versity of Eastern Rhodopes (Bulgaria and Teofilova T. 2016. Initial study of the ground Greece). Pensoft & Nat. Mus. Natur. Hist., beetles (Coleoptera: Carabidae) and other Sofia: 379–411. Invertebrates from ‘Leshnitsa’ nature re- Huusela-Veistola E. 2000. Effects of pesticide serve (Central Stara Planina Mountains, use on non-target arthropods in a Finnish Bulgaria). Ecologia Balkanica 8(1): 79–87. cereal field. In: Cook S. K. et al. (Eds.). Teofilova T.M. 2018. A contribution to the study Farming systems for the new Millennium. of ground beetles (Coleoptera: Carabidae) Aspects of Applied Biology 62: 67–72. in the Western Rhodope Mts. (Bulgaria). Kodzhabashev N.D., Penev L.D. 2006. The Journal of BioScience and Biotechnology ground beetles (Coleoptera: Carabidae) of (2017) 6(3): 203–209. South Dobrudzha, Bulgaria. Acta Zoologi- Teofilova T.M. 2019a. Errata: A contribution to ca Bulgarica 58(2): 147–180. the study of ground beetles (Coleoptera: Kostova R. 2009. Ground beetles (Coleoptera: Carabidae) in the Western Rhodope Mts. Carabidae) in two Biosphere Reserves in (Bulgaria), published in J. BioSci. Biotech. the Rhodope Mountains, Bulgaria. Acta Zo- 2017, 6(3): 203–209. (Errata) Journal of ologica Bulgarica 61(2): 187–196. BioScience and Biotechnology 8(1): 39– Kryzhanovskij O.L. 1965. Composition and or- 44. igin of the terrestrial fauna of Middle Asia. Teofilova T.M. 2019b. Ground beetles Nauka, Moscow-Leningrad. 420 p. (in Rus- (Coleoptera: Carabidae) from the region sian). of Vrachanska Planina Mts. In: Bechev Kryzhanovskij O.L. 1983. Fauna of the USSR. D., Georgiev D. (Eds). Faunistic diversity Coleoptera, vol. 1, is. 2. The Ground-bee- of Vrachanski Balkan Nature Park. Part tles of suborder Adephaga: families Rhyso- 2. ZooNotes, Supplement 7, Plovdiv didae, Trachypachidae; family Carabidae University Press: 39–58. (introduction, overview of the fauna of the Teofilova T.M. 2019c. Ground beetles (Coleop- USSR). Nauka, Leningrad. 341 p. (in Rus- tera: Carabidae) from sparsely vegetated sian). lands in Bulgaria. North-Western Journal of Kryzhanovskij O.L. 2002. Composition and Zoology (2018) 15(1): 30–41. distribution of entomofaunas of the globe. Teofilova T., Markova E., Kodzhabashev N. KMK, Moscow. 237 p. (in Russian). 2015. Ground beetles (Coleoptera: Car- Kryzhanovskij O.L., Belousov I.A., Kabak I.I., abidae) from the region of Cape Emine Kataev B.M., Makarov K.V., Shilenkov V.G. (Central Bulgarian Black sea coast). Part 1995. A checklist of the ground-beetles I. Taxonomic and zoogeographic structure, of Russia and adjacent lands (Insecta, life forms, habitat and humidity preferenc- Coleoptera, Carabidae), Series Faunistica es. ZooNotes 68: 1–18. No 3. PENSOFT Publishers, Sofia-Mos- Vigna Taglianti A., Audisio A.P., Biondi M., Bolo- cow. 271 p. gna M.A., Carpaneto G.M., De Biase A., Fat- Popov M.G. 1927. The main features of the his- torini S., Piattella E., Sindaco R., Venchi torical development of the flora of Middle A., Zapparoli M. 1999. A proposal for choro- Asia. Bulletin of Middle Asian State Univer- type classification of the Near East fauna, sity 15: 239–292 (in Russian). in the framework of the Western Palaearc- Popov V.V., Krusteva I.A. 1999. Epigeobiont tic region. Biogeographia 20: 31–59. 90 T. Teofilova and N. Kodzhabashev

Appendix , , R _ _ , DG _ M O _ 1, B M_O, 1, SK _ B , 2♀, O , 9♀9♂; 1, 1♀; M _ , 1♀; O O O DG P _ _ O _ _ , 2♂; _ P P B , 1♀; 1, 3♀3♂; . P AT , 1♂; O , 1♀4♂; O _ M _ _ , 3♀1♂[h]; , 1♂; M , 1♂; DG M AT_Ps , 15♀6♂2ex.; O _ , 1♂; B_BR, 3♂; AT 1, 1♀; Pn _ , 143♀125♂16ex.; O SK B , 1♀4♂; _ P _ _ O 1, 1♂ RF , 1♂; _ Pn P _ M 1, 1♀; AT , 10♀17♂; DG _ , 2♀; R , 1♂ _ DG , 15♀15♂3ex.; R AT O , 3♀; , 1♀1♂; _ O _ B AT _ O Pn S _ O B _ M _ _ AT_M _ AT P M , 1♀1♂; 1, 1♀; B M AT R , 1♀1♂; Ps _ 1♂ _ , 1♀2♂; O

M , 1♀1♂; _ O , 2♂1ex.; NA 1♀5♂ , 2♂1ex.; NA , 2♀2♂; AT S , 1♂; _ , 2♀1♂; , 4♀7♂; NA 1, 1♀; AT _ R ; 1, 1♂[t]; O R , 6♀6♂; , 4♀11♂; , 4♀11♂; _ B R AT DG _ _ O , 2♀; A _ , 10♀3♂1ex.; _ 1♀ M Ps _ M _ 1♀ 1, 1♀3♂ B AT O _

M P SK AT_Pn _ 1, 1♀ M AT _ Locality code, material DG P , 41♀32♂10ex.; AT _ 1♂; DG , 3♀2♂; , 1♂; NA 7♀2♂ , 1♂; NA , 2♂; B , 1♀; Ps _ , 8♀6♂3ex.; 2♀1♂ R R M _ O , 3♀1♂6ex.; B_DG1, B , 3♀3♂; _ 1, 1♀1♂; _ _ M , 1♀; NA 1, 4♀5♂; _ A _ A O , 3♀2♂; M M AT P R _ _ _ KD 1♂; _ AT M SK AT AT AT_O1, AT_O1, M_M, , 1♀1♂; _ AT_O SK P DG , 1♂; 3♀2♂ , 1♀3♂; , 2♀; , 2♀4♂; , 3♂; , 1♀; _ , 12♀19♂5ex.; , 1♀ 1, 1♀; 1, 2♀3♂; ; B_DG, B Pn A M 1, 3♀5♂; Ps O O SK _ _ _ _ 1, 9♀4♂; M_D, 2♀1♂; _ _ _ R_RF, 1ex.[h] R_RF, AT_O 1ex. Ru_R, 2♀; B_BR, 1♀; AT 3♂2♀[h]; 167♀154♂5ex.; P_O O_m, 1♀; 1♂; R_AY, 2♀; O_O, 1ex.[h]; O_m, 1♂[h]; O_O, 1 ex.[h]; P P_SK, AT_O 3♀4♂; O_O, 1ex.[h]; Ru_R, 3♀1♂; R_AY, AT AT 12♀7♂; DG 12♀9♂3ex.; O_m, 1♀1♂; Ru_R, 1♀; R_AY, P M 1♀1♂ 18♀13♂; NA 1ex. 1ex.[h]; R_AY, O_O, 1ex.[h]; R_RF, P 1♀[h]; P_O1, 1♀2♂; O HP MX M MH M M M M MX X M MX MX MH MX LF 1.2.4 1.3(1).1 1.3(1).1 1.3(1).1 1.3(1).1 1.3(1).1 1.2.2(1) 1.2.2(1) 1.2.2 1.2.2 1.2.2 1.2.2 1.2.2 1.2.2 ZC CE-PAS EUR E-PAS WPAL E-PA-M E-PAS PAL PAL CE-PAS CE-E B-PAS E-PAS EUR B-PAS Olivier, Olivier,

Ménétriés, 1836 Linnaeus, 1760 Palliardi, 1825 (Linnaeus, 1758) Fabricius, 1775 Dejean, 1822 scabriusculus ) (Fabricius, 1792) rufomarginatus ) (Fabricius, 1779) Mannerheim, 1830 intricatus Species ) (Linnaeus, 1758) convexus Systematic list of the ground beetles found in Eastern Rhodopes 2005–2006 and 2019 montivagus wiedemanni ) sylvicola sycophanta ) ) morio G. R. Waterhouse, 1833 G. R. Waterhouse, ) brevicollis (s.str.) (s.str.) Trachycarabus Curtis, 1829 ( inquisitor Pogonophorus ( (s.str.) Table A1. Table Archicarabus Archicarabus Tomocarabus Chaetocarabus Pachystus substriatus (s.str.) ( ( ( ( ( . rufipes . . *Cicindela Leistus (Duftschmid, 1812) Nebria *Notiophilus biguttatus *N N Calosoma C Carabus 1795 C. C. C. C. C. 1. 2. 3. 4. 5. 6. 7. 8. 9. 11. No 10. 12. 13. 14. Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 91 , D 1, _ , R M M _ _ M AT , 1♀2♂; , 8♀7♂9ex.; 1, 1♀1♂; M M O _ _ _ , 1♀4♂ M M AT , 2♀1♂1ex.; , 30♀24♂6ex.; DG R A _ _ _ B , 67♀73♂9ex.; , 91♀69♂9ex., 1♂[h]; M AT , 1♀[t]; O , 1ex.; 1, 3♀1♂; NA 7♀6♂1ex. 1, 3♀1♂; NA Pn _ _ O SK P , 6♂; _ _ DG M AT _ P _ 3♀1♂ AT B , 4♀2♂; M , 79♀119♂14ex.; , 79♀119♂14ex.; SK , 1♂; _ 1, 1♂; , 2♀ Ps , 1♂; P O _ 1, 1♀1♂; SK ; M_R1, _ _ Ps O M 3♀3♂6ex. P _ P _ , 2♀4♂; SK_R, 3♀; Locality code, material M 1♂ , 35♀43♂17ex.; AT_Pn AT 1♀ 2♀2♂ SK , 2♀3♂ Ps , 22♀23♂5ex.; , 2♂; _ _ 3ex. ; M_M, O P , 2♂; , 1♂; DG _ DG AT _ O P _ B_DG, M_R, _ B B , 8♀5♂1ex.; ; M_R1, M AT_O O _ ; M_R1, 1♀ M 1♀1♂ 1♀ 5♀7♂9ex. 2♂ 1♀ 5♀2♂ 1♂ , 1♀1♂; , 19♀16♂6ex.; , 1♂; , 2♀3♂; O M S _ _ _ R_AY, 1♀[h]; R_AY, 3♀3♂; 1♀[h]; R_AY, R_AY, AT 32♀25♂17ex.; P_O1, 25♀17♂; 1♀1♂; B 1ex.[h]; Kr, M_Ps, M_R, B_BR, 2♀ M_R, O_O, 1♂; O_m, 1♀1♂; AT 4♀1♂, 1♀[t]; AT_A 1♂ R_AY, P_SK, SK_R, M_R, M_R, B_DG, B_BR, 6♀6♂ B_BR, 2♀1♂; B_BR, 2♀3♂; 1♀[h] R_AY, 1♂[h] R_AY, B_BR, 3♀1♂ HP E M M H MH MH MX M M H H MH MH M H H H H MH H LF 1.2.2 1.2.2 1.2.2 1.2.3 1.4.2(1) 1.4.2(1) 1.3(1).2 1.3(1).2 1.3(1).2 1.3(1).4 1.3(1).1 1.2.3 1.2.3 1.3(1).2 1.3(1).1 1.3(1).1 1.3(1).1 1.3(1).1 1.3(1).1 1.3(1).2 ZC E-PAS B-PAS BAL-K EUR PAL E-PAS E-CA-M B-PAS NMED E-PA-M E-PA-M WPAL B-CAS E-WSI MED PAL B-PAS E-PA-M EUR CEE-PA Peyron, 1858 (Stephens, P. W. J. Müller, Müller, J. W. P. (Duftschmid, (Duftschmid, (Schrank, 1781) (Motschulsky, (Motschulsky, Crotch, 1866 Audinet-Serville, Linnaeus, 1758 aureus Olivier, 1789 Olivier, (Fabricius, 1792) ) Palliardi, 1825 properans bistriatus ) Dejean, 1831 ) effluviorum (Linnaeus, 1758) ) (Linnaeus, 1760) Species Netolitzky, 1926 Netolitzky, (Herbst, 1784) Piochard de la Brûlerie, 1876 deletum stephensii quadrisignata guttula ) ) ) s.str. ( coriaceus quadristriatus subcostatum ) (Solsky, 1874) (Solsky, scabrosus ) fossor Metallina ) ( tethys Elaphroterus ) (s.str.) ( Paratachys collaris ( crucifer subnotatus (s.str.) (s.str.) Phyla ( Philochthus Peryphus Peryphanes Peryphanes flavicorne (s.str.) ( ( ( ( Procrustes Procerus .

(s.str.) (s.str.) ( ( .

. . r Asaphidion flavipes B. C. C. Cychrus semigranosus *Elaphrus 1821 Clivina *C Trechus Tr T **Tachys 1812) *Tachyura 1812) * **A *Bembidion 1828) **B. *B. * 1850) *B. 1821 *B. Sinechostictus . No 15. 16. 17. 18. 19. 20. 21. 22. 23. 24 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 92 T. Teofilova and N. Kodzhabashev , , S , Ps _ , O _ , 1♀ B _ R M , _ O DG AT M _ _ B M 4♀9♂ , 1♀, 1♂[t]; NA , 1♀, 1♂[t]; NA , 12♀8♂5ex.; , 1♀1♂[t]; , 1♀2♂2ex.; 1, 1♂; 1, 3♀2♂; R , 1♀; D , 11♀17♂5ex.; , 11♀17♂5ex.; R _ O Ps DG _ R _ _ M _ _ _ M _ ; M_M, KD B , 1♂2ex.; , 12♀13♂22ex.; P_ M , 4♀1♂4ex.; M SK AT M A O M 1♂ _ _ _ P M AT 1, 5♀; , 1ex.; , 1♀2♂; , 1♀2♂; M_O, , 1♀1♂; O DG S , 1♂; Ps _ R _ _ _ A P _ B B _ M , 8♀6♂12ex.; SK AT O _ , 2♀6♂; , 33♀50♂47ex.; , 4♀1♂1ex.; , 61♀66♂19ex.; , 1♀; M M Ps 1, 16♀6♂ Ps _ O _ _ _ P_O1 AT M M M DG , 1♂; Locality code, material _ , 6♀1♂8ex.; R B M _ 1, 4♀, 1♂[t]; 1♀1♂; M_D, 1♂ _ , 1♀; , 93♀85♂52ex.; M O 1, 3♀; M 1, 1♂; R _ 2♀[t] , 2♀1♂; _ O R P DG 1♀1♂ _ _

_ M Pn M M B _ P_SK, NA P_O1, 5♀1♂ ; 1♀ AT 1♂[h]; B_BR, 1♀1♂ 1♀1♂ 2♀ , 8♀14♂2ex.; 1♀ 1, 5♀2♀; NA 1♀ 1♀ 1♂ , 1♀1♂ , 1♂; , 9♀11♂, 2♀2♂[h]; , 9♀11♂, , 1♀9ex.; , 9♀1♂; DG O O DG 1, 1♀4♂, 1♀[t]; _ _ _ _ O_O, 1ex.[h]; 1♀; O_m, 1♂; 3♀3♂3ex.; B Ru_R, 2♀9♂ Ru_R, 1♂ B_BR, 3♀ M_R, SK_R, M_M, P_O M_M, 1♂; O_P, 1♀1♂; Ru_R, 1♀6♂; B_BR, 4♀3♂; O_P, 2♀1ex.; P 8♀3♂7ex.; 4♀1♂4ex.; 2♀2♂ M_D, 25♀10♂, M_R, AT_O AT_M B_DG1, P 1♀; Ru_R, 1♂; O_m, O_P, O 163♀74♂11ex.; B M_R, B_BR, 7♀9♂ HP M MX MX H MH H MH M M MX E MX M MX MX M M M M H LF 1.3(2).1 1.3(2).1 1.3(2).1 1.3(2).1 1.3(2).1 1.3(2).1 1.3(2).1 1.3(2).1 1.3(2).2 1.3(1).2 1.3(1).2 1.3(1).2 1.3(1).2 1.3(1).2 1.3(1).2 1.3(1).6 1.3(1).6 1.3(1).6 1.3(1).2 1.3(1).1 ZC BAL-K B-PAS B-PAS CE-E E-AS E-SI E-SI BAL BAL B-PAS PAL E-CAS PAL E-PAS E-SI NMED WPAL P-SMED E-SI E-SI (Linnaeus, J. Schmidt, (Dejean, 1828) Chaudoir, 1846 Chaudoir, Ganglbauer, Ganglbauer, (Duftschmid, Apfelbeck, 1904 (Chaudoir, 1850) (Chaudoir, (Paykull, 1790) (Fischer von (Paykull, 1790) Motschulsky, 1850 Motschulsky, (Herbst, 1784) (Sturm, 1824) (Palliardi, 1825) Chaudoir, 1868 Chaudoir, (Illiger, 1798) (Illiger, venustus leonisi ) duftschmidi nigrita (Schaller, 1783) (Schaller, balcanicus ) ) Species cinctus micropterus (Sturm, 1824) (Goeze, 1777) ) ) anatolicus ambiguus melanocephalus terricola cimmerius vivalis distinguendus ) ) ) ) (s.str.) (s.str.) dilatatus ovoideus Argutor niger ) ( (s.str.) ) chalybaeus Olisares (s.str.) ( (s.str.) (s.str.) fuscipes (s.str.) (s.str.) Phonias Neocalathus Neocalathus ( ( ( Platysma Pseudomaseus . Neocalathus Neocalathus ( ( t Pristonychus Pristonychus (s.str.) ( ( ( (

. . Laemostenus . t t Myas **Poecilus Pedius inquinatus *Pterostichus P P **P *Molops Tapinopterus 1891 *Calathus C C. C. 1758) **C. **C. 1812) ** L. L. 1823) Waldheim, Synuchus **Agonum 1994 No 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 93 1, 1, R _ O _ , 3♀2♂; P SK R _ 1, 1♂; NA 1, 1♂; NA SK DG _ B , 1♀; , 1♀1♂[h]; O R , 5♀1♂2ex.; _ _ O P _ , 4♀2♂ SK P , 17♀10♂; DG , 1♂; _ B 2♀1♂ DG Ps , 17♀27♂ _ _ 1♀1♂ B 1, 1♀2♂; AT DG , 3♀1♂[h]; _ O O B _ , 1♂[t]; _ ; B_DG, D P AT _ B_DG, M 1, 1♀1♂; , 3♀2♂; Locality code, material 33♀43♂ R 1♀ 2♀3♂ M , 2♀; , 1♂; _ _ 1♀2♂ 1♂ 1, 2♀2♂; O M , 2♀4♂; _ 1♂ R AT _ M AT M_R, _ AT_O M M_D, 1♀ ; B_DG, , 2♂; ; M_R1,

B_DG, AT B_DG, R _ , 11♀2♂1ex.; , 11♀2♂1ex.; ; M_O, 1♀ [t] R M 5♂[h] 1♀ 1♂ _ 4♀3♂ 1♀, 1♀[t] 1♀1♂ 1♂[t] 1♀ 2♂ , 1♀; , 6♀2♂; , 1♂; , 2♀ 3♂[h] 2♀[h]; 1♂[h] , 1♀; , 27♀26♂; , 2♀2♂; , 9♀8♂; , 1♀ , 3♀9♂; , 1ex. M A A A R _ _ _ _ M_M, M_R, P_O, P_O AT 1♀; AT M M_Ps, AT_M AT_A AT 2♀2♂; 1♀ AT_A P_O, M_D, M_R, M_D, M_R, P_O1 P_O1 P_O, SK_R, M_R, B_BR, 1♀ SK_R1, B_BR, 1♀ AT_M HP M E M M MX MX M MH M MX MH X MX M MX M M H MX MH H MH H H M LF 1.3(1).1 1.3(1).1 1.3(1).2 2.2.1 2.3.1 2.1.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1(1) 2.3.1(1) 2.3.2 2.3.2 2.3.2 2.3.1 2.2.1 2.1.1 2.1.1 2.1.1 2.1.1 2.1.1 2.2.1 ZC BAL-K PAL NMED E-CA-M OLA E-PAS OLA E-PA-M PAL CE-E E-CA-M B-PAS OLA E-CAS E-CAS BAL BGE MED-PA NMED ЕMED CE-PAS E-CAS E-PAS E-CAS E-PAS

(Duftschmid, intermedius ) (Fabricius, (Quensel en (Schrank, 1781) (Sturm, 1825) Dejean, 1831 (Goeze, 1777) Apfelbeck, 1904 Gebler, 1833 Gebler, (Germar, 1817) (Germar, (Pontoppidan, 1763) (Paykull, 1790) (Fischer von Waldheim, (Linnaeus, 1760) maculicornis Heyden, 1883 Species teutonus (Panzer, 1796) (Panzer, A. Villa et G. B. Villa, 1833 et G. B. Villa, A. Villa Dejean, 1829 scrobiculatus Dejean, 1828 ) flavicollis Dejean, 1828 (Gyllenhal, 1810) Pseudodichirius ( (Fabricius, 1792) (De Geer, 1774) (De Geer, tricuspidata (Duftschmid, 1812) ) (s.str.) serdicana tenebrioides ) parvicollis (s.str.) apricaria ) ) (s.str.) meridianus exiguus Batenus anthobia eurynota ovata discophorus graecus balcanicus ( aenea lucida saphyrea similata Zezea ) ) ( (s.str.) (s.str.) (s.str.) Paracelia

(s.str.) (s.str.) (s.str.) ( Bradytus Amathitis (s.str.) Pelor Pelor . .

(s.str.) (s.str.) (s.str.) (s.str.) ( ( ( ( . . . . A . . . . t A A A Platynus 1801) *Anchomenus dorsalis **Olisthopus glabricollis *Amara A * * A * A A *A. A. A. Zabrus Z. Z. **Anisodactylus Dejean, 1829 **Gynandromorphus etruscus 1806) Schönherr, Stenolophus S 1823) *Acupalpus ** **A **Parophonus 1812) No 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 94 T. Teofilova and N. Kodzhabashev , 1, R 1, 1, , ,

, R _ O O A _ R _ M , _ _ DG _ , 1♀; P P R _ SK R M AT _ B _ , 1♀2♂; M 2♀1♂1ex. SK , 1♀2♂; 1♂ , 1♀; O 1, 1♀; , 3♀2♂; _ P_O , 1♀1♂; , 3♀8♂[h]; R SK , 1♀; O M _ R _ , 18♀39♂; R _ 1, 1♀2♂ 1♀ _ _ P SK , 1♀2♂; M P ; B_DG, R SK _ , 1♂; _ O _ SK SK ; B_DG, P _ 2♀ M , 4♀1♂; , 22♀18♂8ex.; AT 1, 1♀; P DG B_S, 1♂ , 1♀; _ R , 1♂; _ B DG , 1♀; O Pn _ M , 32♀43♂ B_S, _ _ O AT_M B , 1♀1♂; P _ 1♂[h]; M_O, A AT DG P , 18♀39♂; _ 1, 4♀1♂; _ , 1♂; , 1♂; M B O , 1♀6♂; AT , 1♀; _ _ , 2♀; , 3♀6♂2ex.; , 2♀2♂; , 7♂; , 8♀19♂1ex.; M 1♀ , 1♂; SK_R, , 1♀5♂; , 1♂ _ Ps M R P_O AT R AT O M S _ _ _ _ _ M _ AT_O _ , 1♀; DG P_O M M , 2♂ B AT AT AT _ , 3♀9♂; AT , 1♀; , 3♂; 1♂ SK B B_DG, S 1♂

_ DG Locality code, material , 1♂; ; _ , 1♀; _ P , 1♂; B , 2♂; 1, 1♀; NA 1♂ 1, 1♀; NA B , 9♀17♂; Ps , 4♀8♂; 3♀2♂ 1♂ 1♀ M , 12♀18♂; _ M O A AT_M AT_M _ B_S, , 7♀3♂; , 1♀2♂; _ _ _ , 1♀1♂; O 1, 3♀3♂; M , 13♀12♂ A _ A M SK_R1, , 1♀2♂; M AT_M , 1♀; O AT _ AT AT _ _ AT_A S _ R DG AT , 1♂[t]; _ M _ P B_DG, _ AT B_DG, AT M_R1, B D B M , 1♂; , 1ex.; _ O O 1♀1♂[h]; M _ _ , 2♀11♂; , 1♂; SK_R, 1♂ , 2♂; , 1♀; , 1♂ , 3♀2♂; , 4♀10♂; , 1♀1♂1ex.; , 6♀9♂1ex.; , 1♂; 1, 2♀; 1, 1♂; M M A O O SK _ _ _ _ AT_A SK_R, AT_M AT 2♀1♂; AT_M M_D, 1♀ Mo, 1♀[h]; AT_A O_O, 1♀; O_m, 1♂; 1♀; 1ex.[h]; O_P, P_O O_m, 1♂[h]; Ru_R, 1♂; M 1♂; R_AY, 1♀; Ru_R, 1♀[h]; 21♀27♂; P 1♀3♂[h]; AT_M AT_A 1♀[h]; B_BR, 1♂; O_m, 2♀1♂; 1ex.[h]; O_P, Kr, 109♀316♂; 5♀5♂2ex.; M 1♂ 4♀2♂; NA AT_A M_D, 1♀1♂ B_BR, 5♀4♂; Ru_R, 1♀ HP M M MX E MX X MX MX M MX MX MX MX MX MX X MX X X MX MX LF 2.2.1 2.2.1 2.2.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 ZC NMED B-PAS PAL PAL PAL PAL E-CA-M E-PAS CEE-PA OLA E-PA-M E-CAS WPAL CE-PAS E-CAS E-CAS E-CAS E-PAS E-AS E-CAS NMED

(Duftschmid, Dejean, 1829 (Panzer, 1796) (Panzer, signaticornis (De Geer, 1774) (De Geer, ) Dejean, 1829 (Duftschmid, 1812) Germar, 1823 Germar, griseus calceatus (Dejean, 1829) ) (Duftschmid, 1812) Dejean, 1829 (Ménétriés, 1832) Stephens, 1828 ) Reitter, 1900 Reitter, Species rufipes tenebrosus Dejean, 1829 (Duftschmid, 1812) ) ) (Duftschmid, 1812) Sturm, 1818 (Quensel, 1806) Latreille, 1804 1796) (Panzer, Semiophonus ( planicollis laeviceps flavicornis albanicus smaragdinus cupreus honestus sulphuripes rubripes attenuatus atratus serripes pumilus subcylindricus tardus autumnalis Pseudoophonus Cryptophonus (s.str.) (s.str.) Pseudoophonus ( ( (s.str.) (s.str.) (s.str.) (s.str.) Pseudophonus

( . . ( (s.str.) (s.str.) (s.str.) (s.str.) (s.str.) (s.str.) (s.str.) (s.str.) (s.str.) (s.str.) . . . .

P P ...... ** ** **Harpalus (Duftschmid, 1812) H *H. *H. 1812) *H. H H H H H H *H H H H *H *H H *H No 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. Ecological, faunistic and zoogeographical notes on the ground beetles (Coleoptera: ... 95 1, , D , _ DG ; _ M M ; M_O, , , 2♀9♂; B _ 1, 1♀; O 1, 4♀3♂; AT _ DG O O 3ex. _ , 1♂; , 1♀1♂; NA , 1♀1♂; NA _ _ M B P AT SK DG _ _ P M_R, B , 1♀1♂; 1, 1♂; , 4♀3♂2ex. [t] , 1♀; R D Pn O _ , 1♀1♂; _ _ , 16♀34♂2ex.; _ S M M , 1ex.; 1, 1♂; _ AT AT DG B O , 1♀1♂; _ _ O B P , 1♂; P_O _ R M 1ex. _ , 3♀6♂1ex., 1♀[h]; M , 1♀; , 2♀1♂; O _ O O , 20♀19♂; 1, 1♀, 1♂[t]; , 1♀; , 1♀1♂; _ _ P A O O S P _ M _ _ _ ; KD_R, 1, 1♀; P , 4♀1♂6ex.; P B AT O 2♂ _ , 4♀3♂ Locality code, material , 1♂; P , 1ex.; 1♀ , 1♀3♂; 2♀1♂ R , 2♂; 4♂3ex. AT_M _ 1♀ DG 2♀[h] Ps Ps M 2♂[t] _ , 1♀4♂; _ M , 2♀2♂; _ _ , 1♂; B M SK_R1, M M AT O _ AT _ _ ; M_R1, ; B_DG, P AT AT SK_R, M_D, B_DG, , 1♀; ; B_DG, S 1♀ _ 1♂ 6♀3♂ 1♀[h] 2♂[h] B 1♀1♂ 1♂[t] , 1♀5♂[h]; , 1♀1♂ , 1♂; , 1♀ , 2♀2♂; , 1♂; , 1♀ , 1♂ 1♀7♂[h]; , 2♀; , 12♀3♂5ex.; 1, 2♀3♂; , 1♂ R M M A O _ _ _ _ _ O_O, 1♂; Ru_R, 6♀18♂2ex.; B_BR, 26♀83♂2ex.; SK 25♀43♂3ex. AT_M P_O1 P_O, SK_R, B_DG, Ru_R, 2♀1♂; O_m, 1♀; AT M 1♂ B_DG, AT_M AT 1♂[t]; M_R, AT 1♀ Ru_R, 1♂; SK_R1, M_D, AT_M AT_M AT_A 3♀1♂5ex. SK_R, B_BR, 1♀; Ru_R, 1♀; P_O HP MX MX X MX E MX MX M MX MX MX MX MX X MX X X H H M X LF 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.3.1 2.2.1 2.2.1 2.2.1 2.2.1 2.2.1 2.2.1 2.3.2 2.3.2 2.3.2 2.3.3 2.3.3 1.3(1).1 1.3(1).1 1.3(1).1 1.3(1).1 ZC E-PAS NMED CE-PAS E-CAS PAL CEE-PA E-CAS WPAL EUR E-CA-M E-PA-M E-CAS NMED CE-PAS NMED MED NM- CAS E-CAS WPAL B-PAS BAL Dejean, (Dejean, (P. Rossi, (P. Mannerheim, (Dejean, 1829) (Fabricius, 1775) (Fabricius, 1792) (Dejean, 1829) (Paykull, 1790) Dejean, 1829 (Panzer, 1796) (Panzer, laticollis ) (Olivier, 1795) (Olivier, (Duftschmid, 1812) (P. Rossi, 1792) (P. Apfelbeck, 1901 aeneocephalus azureus ) Dejean, 1829 subquadratus cribricollis (P. Rossi, 1790) (P. ) ) ) Species megacephalus vestitus ) Dejean, 1829 rufibarbis ) Sturm, 1818 dama ) parallelus festivus ) picipes (Schrank, 1781) (Dejean, 1825) graecus ammophilus Metophonus ) (s.str.) ( (s.str.) (s.str.) dimidiatus pygmaeus hospes affinis distinguendus saxicola (s.str.) Chlaeniellus Trichochlaenius Oedematicus ( ( Metophonus (

( Metophonus Hesperophonus Hesperophonus Hesperophonus Osimus . . ( ( ( ( (s.str.) (s.str.) (s.str.) (s.str.) (s.str.) (s.str.)

( A...... hl hl . H H H H H H *Ophonus 1825 *O. O. O. O. 1829) O. Acinopus A ** 1794) **Carterus Dixus obscurus Chlaenius C C 1826 Licinus No 111. 110. 112. 113. 114. 115. 116. 117. 118. 119. 101. 102. 103. 104. 105. 106. 107. 108. 109. 120. 121. 96 T. Teofilova and N. Kodzhabashev , 1, DG R _ _ B SK 1, 2♀; O , 1♀1♂1ex.; _ M 1, 1♀, 1♀[t]; P 1, 1♀; _ , 18♀12♂ O R M _ _ , 2♀1♂; P DG M R _ _ B , 1♀[h]; SK O , 1♂; , 1♀[h]; _ R R P _ _ , 4♀2♂[h], M SK O , 3♀3♂; _ M P , 2♀2♂6ex.; , 1♀2♂; _ S , 1♂; M 1♂ _ , 2♀3♂; O AT _ B _ M P AT _ , 5♀4♂; AT M_R, Locality code, material , 1♀; M , 6♀4♂ _ R , 1♀; , 3♀6♂; , 1♂; M_D, 1♂ 2♂ _ ; A A M AT DG , 1♀ , 1♀; M _ _ _ _ 1♀ B AT AT AT AT_M AT_M B_DG, B_S, , 1♂; 1ex. 2♂ 1♀1♂ R , 1♂ Table 1. Table , 1♀1♂6ex. , 82♀30♂ _ , 5♀4♂[h]; , 1ex. , 1♀; 1♂[h] , 4♀4♂; , 34♀, 32♂; R M A A _ DG DG _ _ _ _ AT_M SK_R1 B_DG, P_O, 1♂; Ru_R, 1♂; R_AY, 4♀3♂ Ru_R, 2♀1♂; 1♀; B_BR, 1♂; 1♂; O_P, Ru_R, 1♀1♂; B B_DG, B_BR, 4♀; 1♀ O_P, AT B B_DG, AT_M Ru_R, 1♀ AT SK HP MX X MH MH M MX M M MX MX H MX MX MX MX M MX LF – new species for the whole Rhodope Mts. ZC – zoogeographic category 1.3(1).1 1.1.3 1.1.2 1.3(1).5 1.3(1).3 1.3(1).3 1.3(1).3 1.3(1).3 1.3(1).3 1.3(1).3 1.3(1).1 1.3(1).3 1.3(1).3 1.3(1).3 1.3(1).3 1.3(1).3 1.3(1).3 ** ZC NMED E-PAS E-CA-M E-PAS E-PA-M E-CA-M MED E-CAS OLA MED-PA E-PAS E-PA-M PAL E-CA-M B-CAS E-CA-M E-CAS (Olivier, (Olivier, Apfelbeck, (Linnaeus, Motschulsky, Motschulsky, Duftschmid, 1812 (Duftschmid, (Duftschmid, 1812) (Linnaeus, 1758) (Fabricius, 1794) (Geoffroy, 1785) (Geoffroy, (Reitter, 1901) (Reitter, brevicollis bodemeyeri Species (P. Rossi, 1790) (P. Audinet-Serville, 1821 (Wollaston, 1854) (Wollaston, atricapillus ) ) explodens ) quadrillum crepitans axillaris (Dejean, 1825) (Goeze, 1777) marginata (s.str.) (Dejean, 1825) (s.str.) (s.str.) psophia (s.str.) silphoides (s.str.) Brachynidius Brachynidius ( ( Brachynidius impressus * – new species for the Eastern Rhodope Mts.; (s.str.) ( minutulus . . pallipes

. (s.str.) . :

. L *Lebia **Demetrias 1758) Paradromius (Manodromius) linearis 1759) Syntomus obscuroguttatus *S **S *Microlestes fissuralis *M Microlestes negrita Lionychus 1812) Cymindis Brachinus *Br **Br. 1844 **Br. 1904 Br. Note No 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. (explanations are given in Table 3 the Results section of the main Methods paper); section LF of – the life main form paper); (explanations HP are – given humidity preferencein (H the – mesoxerophilous, Material X – hygrophilous, and xerophilous, E MH – eurybiont); ♀ – – female, mesohygrophilous; ♂ – male, M ex. – – not determined; [h] mesophilous, – hand MX collection; [t] – – tree trap; NA – label destroyed, material collected in 2005–2006. Locality codes are given