Arch. Biol. Sci., Belgrade, 65 (4), 1675-1686, 2013 DOI:10.2298/ABS1304675C

REMARKS ON SOME TRECHINE GROUND TAXA FROM THE BALKAN PENINSULA (COLEOPTERA: CARABIDAE: TRECHINI)

S. B. ĆURČIĆ, B. P. M. ĆURČIĆ and MAJA VRBICA

Institute of Zoology, University of Belgrade - Faculty of Biology, 11000 Belgrade,

Abstract – The purpose of this note is to offer some corrections and supplements to the paper of Lohaj et al. (2013), to prove the validity of recently described trechine genera in Serbia, and to comment on other erroneous facts presented in the analyzed paper. In particular, we will show that the taxonomical remarks on the genus Duvalius Delarouzée, 1859, are wrong. The following genera are clearly delimited and represent valid taxa: Serboduvalius Ćurčić, Pavićević & Ćurčić, 2001, comb. n., Rascioduvalius Ćurčić, Brajković, Mitić & Ćurčić, 2003, comb. n., Curcicia Ćurčić & Brajković, 2003, comb. n., and Javorella Ćurčić, Brajković & Ćurčić, 2003, comb. n. The following new combinations are established for the fol- lowing Duvalius species: Serboduvalius gejzadunayi (Lohaj, Čeplík & Lakota, 2013), comb. n., S. starivlahi (Guéorguiev, Ćurčić & Ćurčić, 2000), comb. n., S. dragacevensis Ćurčić, Pavićević & Ćurčić, 2001, comb. n., S. droveniki (Magrini, 1998), comb. n., Rascioduvalius cvijici (Jeannel, 1923), comb. n., R. stopicensis (Jeannel, 1923), comb. n., R. zlatiborensis Ćurčić, Brajković & Ćurčić, 2005, comb. n., Curcicia bolei (Pretner, 1963), comb. n., Javorella javorensis Ćurčić, Brajković & Ćurčić, 2003, comb. n., J. suvoborensis (Pavićević & Popović, 2001), comb. n., and J. suvodolensis Ćurčić, Brajković & Ćurčić, 2003, comb. n.

Key words: Coleoptera; Carabidae; taxonomy; revision; Duvalius; Serboduvalius; Rascioduvalius; Curcicia; Javorella; cave- dwelling fauna; Serbia; Montenegro

INTRODUCTION droveniki Magrini, 1998, an endogean species from Recently Lohaj et al. (2013) described a new trechine eastern Montenegro (Magrini, 1998). It is our opin- from a cave in eastern Montenegro – ion that Duvalius (Biharotrechus) droveniki belongs Duvalius (Neoduvalius) gejzadunayi Lohaj, Čeplík to the genus Serboduvalius as well. Additionally, Lo- & Lakota, 2013. However, a thorough analysis of all haj et al. (2013) erroneously included the following important taxonomic traits of this taxon and a com- species in the subgenus Neoduvalius Müller, 1913, parison with other members belonging to the genus genus Duvalius, without any scientific explanation Duvalius Delarouzée, 1859, and its phenetically close and taxonomic review: Serboduvalius starivlahi genera has uncovered that it actually belongs to the (Guéorguiev, Ćurčić & Ćurčić, 2000), S. dragaceven- genus Serboduvalius Ćurčić, Pavićević & Ćurčić, sis Ćurčić, Pavićević & Ćurčić, 2001, Rascioduvalius 2001, whose representatives were previously known cvijici (Jeannel, 1923), R. stopicensis (Jeannel, 1923) from some caves in southwestern Serbia (Guéorguiev and R. zlatiborensis Ćurčić, Brajković & Ćurčić, 2005. et al., 2000; Ćurčić et al., 2001). After additional de- We refute such superfluous analysis by Lohaj et al. tailed analysis, we found that the abovementioned (2013) and maintain the opinion that the species be- species is actually closest to Duvalius (Biharotrechus) long to the genera Serboduvalius and Rascioduvalius

1675 1676 S. B. ĆURČIĆ ET AL.

Ćurčić, Brajković, Mitić & Ćurčić, 2005, as was pre- RESULTS AND DISCUSSION viously stated by Ćurčić et al. (2001, 2003c, 2005), Guéorguiev (2007), Zagmajster (2007) and Janák & Moravec (2008). Moreover, Lohaj et al. (2013) syno- FAMILY CARABIDAE LATREILLE, 1802 nymized a few trechine genera from Serbia without any scientific argumentation (Serboduvalius, Rasci- SUBFAMILY BONELLI, 1810 oduvalius, Javorella Ćurčić, Brajković & Ćurčić, 2003, and Curcicia Ćurčić & Brajković, 2003). We are pro- TRIBE TRECHINI BONELLI, 1810 posing the re-erection to the genus level according to the studies of Ćurčić et al. (2001, 2003a, 2003b, GENUS SERBODUVALIUS ĆURČIĆ, 2003c, 2005), Ćurčić & Brajković (2003), and other PAVIĆEVIĆ & ĆURČIĆ, 2001, COMB. N. authors (Guéorguiev, 2007; Zagmajster, 2007; Janák & Moravec, 2008). Apart from the aforementioned facts, the paper of Lohaj et al. (2013) contains nu- Etymology – After Serbia, itsterra typica. merous mistakes and erroneous facts of both techni- cal and scientific nature. Therefore, a need for a thor- Type species – Serboduvalius dragacevensis Ćurčić, ough revision of the paper arose and we present it in Pavićević & Ćurčić, 2001, comb. n. the current study. Other species – S. starivlahi (Guéorguiev, Ćurčić & MATERIALS AND METHODS Ćurčić, 2000), comb. n., S. gejzadunayi (Lohaj, Čeplík & Lakota, 2013), comb. n., and S. droveniki (Magrini, The specimens of the type species of the generaSer - 1998), comb. n. boduvalius, Rascioduvalius, Curcicia, Javorella, and Duvalius were analyzed in laboratories of the Insti- Synonyms – Subgenus Duvaliotes Jeannel, 1928 tute of Zoology, University of Belgrade – Faculty of (part.): Magrini, 1998: 129; subgenus Neoduvalius Biology, Belgrade, Serbia. These were dissected and Müller, 1913 (part.): Guéorguiev et al., 2000: 227; Lo- thoroughly studied. Dry specimens were adhered haj et al., 2013: 100; subgenus Biharotrechus Bokor, onto paper labels. Male genital structures were 1922 (part.): Moravec et al., 2003: 299. fixed in a medium composed of Canada balsam and xylol. All taxonomically important morphological Old combinations – Duvalius (Duvaliotes): Magrini, characters were studied for comparison. Additional 1998: 129; Duvalius (Neoduvalius): Guéorguiev et al., literature data were taken into account for the com- 2000: 227; Lohaj et al., 2013: 100; Duvalius (Biharo- parison of some taxa and the conclusions present- ): Moravec et al., 2003: 299. ed herein (Jeannel, 1928; Pretner, 1963; Magrini, 1998; Ćurčić et al., 2001, 2003b, 2003c; Pavićević & Type locality – Mala Pećina Cave, village of Rti, near Popović, 2001; Ćurčić and Brajković, 2003; Lohaj et Kotraža, Dragačevo Mts., southwestern Serbia. al., 2013). Type series – Mala Pećina Cave, village of Rti, near A Carl Zeiss Stemi 2000 binocular stereomicro- Kotraža, Dragačevo Mts., southwestern Serbia, 30 scope and Carl Zeiss Axioskop 40 microscope were April 2001, leg. S. Ćurčić, B. Mitić & S. Makarov used for examining the morphological characteris- (holotype male, allotype female, and a paratype tics of the trechine ground . Additionally, the male); idem, 20 August 2001 (five paratype males original illustrations of all the type species from lit- and a paratype female). erature sources were compared (Jeannel, 1928; Pret- ner, 1963; Magrini, 1998; Ćurčić et al., 2001, 2003b, Diagnosis – As presented in the paper of Ćurčić 2003c; Lohaj et al., 2013). et al. (2001). REMARKS ON SOME TRECHINE GROUND BEETLE TAXA FROM THE BALKAN PENINSULA 1677

Fig. 1. Distribution of species of the genus Serboduvalius. Square - S. dragacevensis; triangle - S. starivlahi; circle - S. gejzadunayi; pen- tagon - S. droveniki.

Distribution – The genus Serboduvalius, comb. n. is that they are present among other Duvalius taxa, but presently known only from a few caves and deep soil the characters need to be studied in a complex way. in southwestern Serbia and eastern Montenegro (Fig. They overlooked the fact that the set of correlative 1). morphological features of the genus is unique and do not exist in any other trechine genera. Remarks – This genus differs from all other pheneti- cally close genera by a set of correlative morphologi- We maintain the opinion of Pavićević & Popović cal features as listed in Ćurčić et al. (2001). This set (2001), who claimed that it would be inappropriate of characteristics is specific to species belonging to and even pretentious to establish a new subgenus un- the genus, and is not evident within other Duvalius til the area of Serbia were better covered (when some and Duvalius-like taxa. Lohaj et al. (2013) singled out related species might possibly be discovered). Janák certain its morphological features and mentioned & Moravec (2008) claimed that the recently described 1678 S. B. ĆURČIĆ ET AL. trechine Duvalius-like genera from Serbia probably a cave in the vicinity of the town of Rožaje, eastern represent either groups of species or subgenera. The Montenegro. solution of the taxonomic problem lies in a better knowledge of the diversity of the genus Duvalius in Remarks – After detailed analysis of morphologi- the Balkans, and descriptions of additional new spe- cal features of Duvalius (Neoduvalius) gejzadunayi cies in the area studied will enable us to define the fi- (presence of reduced eyes, deep and complete frontal nal status of the trechine genera analyzed. The recent furrows, longitudinal fissure on fore tibias, smooth finding and description of the third Serboduvalius genae, 3-4 pairs of elytral discal setae, position of the species, Serboduvalius gejzadunayi, comb. n. (Lohaj second pair of elytral discal setae and the humeral et al., 2013), which shares the basic correlative mor- umbilicate setae, and similar kind of copulatory phological characteristics of Serboduvalius, comb. n. piece), we concluded that this species actually be- (presence of reduced eyes, deep and complete frontal longs to the genus Serboduvalius, comb. n., together furrows, smooth genae, and 3-4 pairs of elytral discal with two previously described species from south- setae, the second pair of elytral discal setae closer to western Serbia and another species from eastern the first pair than to the third pair of these setae, the Montenegro. The paper of Lohaj et al. (2013) con- same position of the humeral umbilicate setae, and tains a description of the species, but the diagnosis the presence of a similar gutter-formed copulatory is incomplete since the authors surprisingly did not piece which is bilobed at the base) confirmed the fact compare the new species with the other phenetically that the genus is valid. The additional trechine spe- close species Serboduvalius droveniki, comb. n. (liv- cies is added into the genus Serboduvalius, comb. n. ing in deep soil in the vicinity of the type locality of S. as well (S. droveniki, comb. n.), since it possesses all gejzadunayi, comb. n.) and S. dragacevensis, comb. n. leading characteristics of the genus. (the type species of the genus Serboduvalius, comb. n.). The following remains unclear: what criteria was The genus Serboduvalius, comb. n. probably be- used by the authors to include the species within Du- longs to an old and separate phyletic lineage that valius (Neoduvalius) and what is the precise position originated during Tertiary (Ćurčić et al., 2001). The of setae of the humeral umbilicate series? Further- taxon is both an endemic and relict of southwestern more, the presence of complete frontal furrows is Serbia, eastern Montenegro and the Balkan Penin- not characteristic for Neoduvalius taxa, so the new sula. species cannot be associated with it. Additionally, the presence of three pairs of elytral discal setae is very SERBODUVALIUS GEJZADUNAYI (LOHAJ, rare in Neoduvalius species, while it is observed in all ČEPLÍK & LAKOTA, 2013), COMB. N. Serboduvalius species. The presence of four pairs of elytral discal setae (in some specimens of Serbodu- Old combination – Duvalius (Neoduvalius) gejzadu- valius starivlahi, comb. n., S. gejzadunayi, comb. n., nayi: Lohaj et al., 2013, 94. and S. droveniki, comb. n.) is not common in Neodu- valius species. Additionally unclear was the presence/ Type locality – Pećina u Dubokom Potoku Cave, vil- absence of eyes in Serboduvalius gejzadunayi, comb. lage of Donje Biševo, 1 180 m.a.s.l., near Rožaje, east- n. by Lohaj et al. (2013). At first it was recorded that ern Montenegro. the eyes are completely reduced (a dark spot is regis- tered in a few specimens) (Lohaj et al., 2013: 96), and Other localities – None. then the authors reported that the eyes in the species are totally absent (Lohaj et al., 2013: 99). However, Description and diagnosis – As presented in the paper small eye spots can be seen in the species in Figs. 1 of Lohaj et al. (2013). and 3 presented in the cited paper (Lohaj et al., 2013: 95). The authors wrongly named the type locality as Distribution – This species is presently known from Duboki Cave instead of Pećina u Dubokom REMARKS ON SOME TRECHINE GROUND BEETLE TAXA FROM THE BALKAN PENINSULA 1679

Potoku Cave in the Distribution chapter (Lohaj et species are listed in the paper of Ćurčić et al. (2001), al., 2013: 98), the village where the cave is situated and additional drawings showing some of them are (Dinje Biševo instead of Donje Biševo) in the chapter included there as well. Topographic location and ecology (Lohaj et al., 2013: 98), and erroneously determined the pseudoscorpion SERBODUVALIUS DRAGACEVENSIS ĆURČIĆ, species caught at the same locality (Neobisium ninae PAVIĆEVIĆ & ĆURČIĆ, 2001, COMB. N. Ćurčić, Dimitrijević & Tomić was described from the type locality vs. Neobisium umbratile Beier, 1938) Old combination. – Duvalius (Neoduvalius) dra- (Ćurčić et al., 2007, 2008; Lohaj et al., 2013: 98). gacevensis: Lohaj et al., 2013, 101.

SERBODUVALIUS STARIVLAHI (GUÉORGUIEV, Type locality – Mala Pećina Cave, village of Rti, near ĆURČIĆ & ĆURČIĆ, 2000), COMB. N. Kotraža, Dragačevo Mts., southwestern Serbia.

Old combinations – Duvalius (Neoduvalius) stariv- Other localities – None. lahi: Guéorguiev et al., 2000, 227. Description and diagnosis – As presented in the paper Duvalius (Neoduvalius) starivlahi: Lohaj et al., 2013, of Ćurčić et al. (2001). 102. Distribution This species is presently known from a Type locality. – Hadži-Prodanova Pećina Cave, vil- cave in the Dragačevo Mts., southwestern Serbia. lage of Raščići, near Ivanjica, Southwestern Serbia. Remarks – Presence of reduced eyes, deep and com- Other localities. – None. plete frontal furrows, longitudinal fissure on fore tib- ias, smooth genae, three pairs of elytral discal setae, a Description and diagnosis. – As presented in the pa- gutter-formed copulatory piece, and position of both pers of Guéorguiev et al. (2000) and Ćurčić et al. the second pair of elytral discal setae (closer to the (2001). first than to the third pair of elytral discal setae) and the humeral umbilicate setae (as stated in description Distribution. – This species is presently known from of the genus; Ćurčić et al., 2001: 53) confirm that this a cave in the vicinity of the town of Ivanjica, South- species belongs to the genus Serboduvalius, comb. n., western Serbia. as was originally stated by Ćurčić et al. (2001). Fur- thermore, the precise distance between type locali- Remarks. – Presence of reduced eyes, deep and com- ties of Serboduvalius starivlahi, comb. n. and S. dra- plete frontal furrows, longitudinal fissure on fore tib- gacevensis, comb. n. is 12.5 km (not around 10 km, as ias, smooth genae, 3-4 pairs of elytral discal setae, a was stated by Lohaj et al., 2013: 101). gutter-formed copulatory piece, and position of both the second pair of elytral discal setae (closer to the SERBODUVALIUS DROVENIKI first than to the third pair of elytral discal setae) and (MAGRINI, 1998), COMB. N. the humeral umbilicate setae (as stated in descrip- tion of the genus; Ćurčić et al., 2001: 53) confirm the Old combinations – Duvalius (Duvaliotes) droveniki: fact that this form belongs to the genus Serboduval- Magrini, 1998, 129. ius, comb. n., as was originally stated by Ćurčić et al. (2001). Totally wrong is the opinion by Lohaj et al. Duvalius (Biharotrechus) droveniki: Moravec et al., (2013), who stated that these two species are prob- 2003, 299. ably conspecific on the basis of morphological dif- ferences. A dozen observed differences among the Type locality. – Mt. Hajla, Prokletije, near Rožaje, 1680 S. B. ĆURČIĆ ET AL.

Eastern Montenegro. 1923: 10; Duvalius (Neoduvalius): Winkler, 1926: 260; Jeannel, 1928: 553; Moravec et al., 2003: 307; Lo- Other localities. – None. haj et al., 2013: 100; Duvalius (Duvalius): Pavićević & Popović, 2001: 4. Description and diagnosis. – As presented in the pa- per of Magrini (1998). Type locality – Village of Bela Reka, 1 100 m.a.s.l., Mt. Murtenica, western Serbia. Distribution. – This species is endogean and is pres- ently known from soil on Mt. Hajla, Prokletije, East- Type series – From under stones in a forest, village ern Montenegro. of Bela Reka, 1 100 m.a.s.l., Mt. Murtenica, western Serbia, June 1923, leg. R. Jeannel (holotype male and Remarks. – Presence of reduced eyes, deep and com- allotype female). plete frontal furrows, longitudinal fissure on fore tib- ias, smooth genae, 3-4 pairs of elytral discal setae, a Diagnosis – As presented in the paper of Ćurčić et al. gutter-formed copulatory piece, and similar position (2003c). of both the second pair of elytral discal setae (closer to the first than to the third pair of elytral discal se- Distribution – The genus Rascioduvalius, comb. n. is tae) and the humeral umbilicate setae (as stated in currently known only from few caves and deep soil description of the genus; Ćurčić et al., 2001: 53) con- in western Serbia. firm that this species actually belongs to the genus Serboduvalius, comb. n. Closest to S. gejzadunayi, Remarks – This genus differs from all other pheneti- comb. n. according to form of the aedeagus and the cally close genera by a set of correlative morphologi- copulatory piece and other morphological features cal features as listed in Ćurčić et al. (2003c). This set (Magrini, 1998; Lohaj et al., 2013). of characteristics is specific to species belonging to the genus, and is not evident within other Duvalius GENUS RASCIODUVALIUS ĆURČIĆ, and Duvalius-like taxa. Lohaj et al. (2013) singled out BRAJKOVIĆ, MITIĆ & ĆURČIĆ, 2003, COMB. N. certain its morphological features and mentioned that they are present among other Duvalius taxa, but Etymology – After Rascia, the old name for Serbia, its the characters need to be studied in a complex way. terra typica. They overlooked the fact that the set of correlative morphological features of the genus is unique and Type species – Rascioduvalius cvijici (Jeannel, 1923), do not exist in any other trechine genera. Therefore, comb. n. we refute its synonymization and the inclusion of the species within Duvalius (Neoduvalius). This fact Other species – R. stopicensis (Jeannel, 1923), comb. is confirmed by the opinion of Pavićević & Popović n. and R. zlatiborensis Ćurčić, Brajković & Ćurčić, (2001), who mentioned that the criteria according to 2005, comb. n. which Rascioduvalius cvijici, comb. n. and R. stopi- censis, comb. n. are included into the subgenus Ne- Synonyms – Subgenus Duvalites Jeannel, 1920 (part.): oduvalius of Duvalius are not satisfactory. However, Jeannel, 1923: 10; subgenus Neoduvalius Müller, these authors proposed that the taxa should be trans- 1913 (part.): Winkler, 1926: 260; Jeannel, 1928: 553; ferred into the subgenus Duvalius s. str. Jeannel (1928) Moravec et al., 2003: 307; Lohaj et al., 2013: 100; sub- himself claimed that Rascioduvalius cvijici, comb. n. genus Duvalius Delarouzée, 1859 (part.): Pavićević & has a particular position in the trechine systematics. Popović, 2001: 4. He was not sure whether it belongs to the subgenus Duvalius (= Duvalites) or Neoduvalius of Duvalius. Old combinations – Duvalius (Duvalites): Jeannel, The same author even admitted that the mentioned REMARKS ON SOME TRECHINE GROUND BEETLE TAXA FROM THE BALKAN PENINSULA 1681 species deserves an isolated generic status due to the Duvalius (Neoduvalius) cvijici cvijici: Moravec et al., presence of complete frontal furrows (Jeannel, 1928). 2003, 307. Additionally, Pavićević & Popović (2001) said that it would be inappropriate and even pretentious to es- Duvalius (Neoduvalius) cvijici cvijici: Lohaj et al., tablish a new subgenus until the area of Serbia were 2013, 101. better covered (when some related species might pos- sibly be discovered). Janák & Moravec (2008) claimed Type locality – Village of Bela Reka, 1 100 m.a.s.l., Mt. that the recently described trechine Duvalius-like Murtenica, western Serbia. genera from Serbia probably represent either groups of species or subgenera. The solution of the taxonom- Other localities – None. ic problem lies in a better knowledge of the diversity of the genus Duvalius in the Balkans. Descriptions Description and diagnosis – As presented in the pub- of additional new species in the area studied will en- lications of Jeannel (1923, 1928) and Ćurčić et al. able us to define the final status of the trechine genera (2003c). analyzed. The recent finding and description of the third Rascioduvalius species, R. zlatiborensis, comb. Distribution – This species is endogean and can be n., which shares the basic correlative morphological found under stones in forests on Mt. Murtenica, characteristics of Rascioduvalius, comb. n. (presence western Serbia. of reduced eyes with depigmented ommatidia, deep and complete frontal furrows, longitudinal fissure Remarks – The presence of reduced eyes with depig- on fore tibias, curved pre-ocular furrow, hairy genae, mented ommatidia, deep and complete frontal fur- two pairs of elytral discal setae, the same position of rows, longitudinal fissure on fore tibias, curved pre- the humeral umbilicate setae, and the presence of a ocular furrow, hairy genae, two pairs of elytral discal similar bifid gutter-formed copulatory piece) con- setae, position of the humeral umbilicate setae (as firmed the fact that the genus is valid. stated in description of the genus; Ćurčić et al., 2003c: 484), and presence of a bifid gutter-formed copula- The genus Rascioduvalius, comb. n. probably be- tory piece, confirm the fact that this species belongs longs to an old and separate phyletic lineage which to the genus Rascioduvalius, comb. n., as previously originated during the Tertiary (Ćurčić et al., 2003c). stated by Ćurčić et al. (2003c). Jeannel (1928) and The taxon is both an endemic and relict of western Lohaj et al. (2013) proposed its including in the sub- Serbia and the Balkan Peninsula. genus Neoduvalius, genus Duvalius, but this is wrong because the species possesses complete frontal fur- RASCIODUVALIUS CVIJICI (JEANNEL, 1923), rows (vs. incomplete ones are present in Neoduval- COMB. N. ius species). Our opinion is supported by Pavićević & Popović (2001), who mentioned that the transfer Old combinations – Duvalius (Duvalites) cvijici cvi- is under doubt. Lohaj et al. (2013) erroneously stated jici: Jeannel, 1923, 10. that Rascioduvalius cvijici, comb. n., R. stopicensis, comb. n., and R. zlatiborensis, comb. n., are probably Duvalius (Neoduvalius) cvijici cvijici: Winkler, 1926, conspecific (inhabiting both caves and MSS on Mts. 260. and Murtenica, western Serbia) on the basis of morphological differences, but without any argu- Duvalius (Neoduvalius) cvijici cvijici: Jeannel, 1928, mentation. A dozen observed differences among the 553. species are listed in the papers of Jeannel (1923, 1928) and Ćurčić et al. (2003c, 2005), along with drawings Duvalius (Duvalius) cvijici cvijici: Pavićević & showing some of them. Additionally, the distance Popović, 2001, 4. between the type localities of Rascioduvalius cvijici, 1682 S. B. ĆURČIĆ ET AL. comb. n. (village of Bela Reka, Mt. Murtenica) and R. latory piece, confirm that this species belongs to the zlatiborensis, comb. n. (Markova Pećina Cave, village genus Rascioduvalius, comb. n., as previously stated of Gornji Ljubiš, Mt. Zlatibor) is around 8 km (not by Ćurčić et al. (2003c). Jeannel (1928) and Lohaj et around 2 km, as was stated by Lohaj et al., 2013). al. (2013) proposed its inclusion as a subspecies of the species cvijici in the subgenus Neoduvalius, genus RASCIODUVALIUS STOPICENSIS (JEANNEL, Duvalius, but this is wrong because it possesses com- 1923), COMB. N. plete frontal furrows (vs. incomplete ones, present in Neoduvalius species). Our opinion is supported by Old combinations – Duvalius (Duvalites) cvijici stopi- Pavićević & Popović (2001), who mentioned that the censis: Jeannel, 1923, 11. transfer is under doubt. The clear differences among Rascioduvalius cvijici, comb. n., R. stopicensis, comb. Duvalius (Neoduvalius) cvijici stopicensis: Winkler, n., and R. zlatiborensis, comb. n. are mentioned in 1926, 260. the papers of Jeannel (1923, 1928) and Ćurčić et al. (2003c, 2005), along with accompanying drawings, Duvalius (Neoduvalius) cvijici stopicensis: Jeannel, and these are distinguished as three well-separated 1928, 554. species.

Duvalius (Duvalius) cvijici stopicensis: Pavićević & RASCIODUVALIUS ZLATIBORENSIS ĆURČIĆ, Popović, 2001, 4. BRAJKOVIĆ & ĆURČIĆ, 2005, COMB. N.

Duvalius (Neoduvalius) cvijici stopicensis: Moravec et Old combination – Duvalius (Neoduvalius) zlatibo- al., 2003, 307. rensis: Lohaj et al., 2013, 102.

Duvalius (Neoduvalius) cvijici stopicensis: Lohaj et Type locality – Markova (= Ršumska) Pećina Cave, al., 2013, 101. village of Gornji Ljubiš, Mt. Zlatibor, western Serbia.

Type locality – Stopića Pećina Cave, village of Other localities – None. Rožanstvo, Mt. Zlatibor, near Užice, western Serbia. Description and diagnosis – As presented in the paper Other localities – None. of Ćurčić et al. (2005).

Description and diagnosis – As presented in the pub- Distribution – This species is troglobitic and inhabits lications of Jeannel (1923, 1928) and Ćurčić et al. the Markova (= Ršumska) Pećina Cave on Mt. Zlati- (2003c). bor, western Serbia.

Distribution – This species is troglobitic and can be Remarks – The presence of reduced eyes with dep- found under stones in the Stopića Pećina Cave on igmented ommatidia, deep and complete frontal Mt. Zlatibor, western Serbia. furrows, longitudinal fissure on fore tibias, curved pre-ocular furrow, hairy genae, two pairs of elytral Remarks – The presence of reduced eyes with depig- discal setae, and the position of the humeral um- mented ommatidia, deep and complete frontal fur- bilicate setae (as stated in description of the genus; rows, longitudinal fissure on fore tibias, curved pre- Ćurčić et al., 2003c: 484) confirm that this species ocular furrow, hairy genae, two pairs of elytral discal belongs to the genus Rascioduvalius, comb. n., as setae, position of the humeral umbilicate setae (as originally stated by Ćurčić et al. (2003c). Lohaj et stated in description of the genus; Ćurčić et al., 2003c: al. (2013) proposed including the species in the sub- 484), and the presence of a bifid gutter-formed copu- genus Neoduvalius, genus Duvalius, in spite the fact REMARKS ON SOME TRECHINE GROUND BEETLE TAXA FROM THE BALKAN PENINSULA 1683 that it has complete frontal furrows (vs. incomplete logical features and mentioned that they are present ones are present in Neoduvalius species). Pavićević & among other Duvalius taxa, but the characters need Popović (2001) also do not support such an opinion. to be studied in a complex way. They overlooked the fact that the set of correlative morphological features GENUS CURCICIA ĆURČIĆ & BRAJKOVIĆ, 2003, of the genus is unique and do not exist in any other COMB. N. trechine genera. Therefore, we refute its synonymi- zation and inclusion in the species within Duvalius. Etymology – After Prof. Dr. Božidar Ćurčić, a famous Janák & Moravec (2008) are of the opinion that the Serbian zoologist. recently described trechine Duvalius-like genera from Serbia probably represent either groups of spe- Type species – Curcicia bolei (Pretner, 1963), comb. cies or subgenera. The solution of the taxonomic n. problem lies in a better knowledge of the diversity of the genus Duvalius in the Balkans. Descriptions of Other species – None. additional new species in the area studied will enable us to define the final status of all the trechine genera Synonyms – Subgenus Duvalius Delarouzée, 1859 there. (part.): Pretner, 1963: 144; Pavićević & Popović, 2001: 3; Moravec et al., 2003: 301; genus Duvalius The genusCurcicia , comb. n. probably belongs to Delarouzée, 1859 (part.): Lohaj et al., 2013: 100. an old and separate phyletic lineage which originated during the Tertiary (Ćurčić & Brajković, 2003). The Old combinations – Duvalius (Duvalius): Pretner, taxon is both an endemic and relict of eastern Serbia 1963: 144; Pavićević & Popović, 2001: 3; Moravec et and the Balkan Peninsula. al., 2003: 301; Duvalius: Lohaj et al., 2013: 100. CURCICIA BOLEI (PRETNER, 1963), COMB. N. Type locality – Prekonoška Pećina Cave, village of Prekonoga, near Svrljig, Svrljiške Planine Mts., east- Old combinations – Duvalius (Duvalius) bolei: ern Serbia. Pretner, 1963, 144; Pavićević & Popović, 2001, 3; Moravec et al., 2003, 301; Duvalius bolei: Lohaj et al., Type series – Prekonoška Pećina Cave, village of 2013, 100. Prekonoga, near Svrljig, Svrljiške Planine Mts., east- ern Serbia, 26 May 1959, leg. E. Pretner (holotype Type locality – Prekonoška Pećina Cave, village of male). Prekonoga, near Svrljig, Svrljiške Planine Mts., east- ern Serbia. Diagnosis – As presented in the paper of Ćurčić & Brajković (2003). Other localities – None.

Distribution – The genus Curcicia, comb. n. is cur- Description and diagnosis – As presented in the paper rently known only from a cave in eastern Serbia. of Pretner (1963).

Remarks – This genus differs from all other pheneti- Distribution – This species is troglobitic and inhabits cally close genera (including Duvalius) by a set of the Prekonoška Pećina Cave on the Svrljiške Planine correlative morphological features as listed in Ćurčić Mts., eastern Serbia. & Brajković (2003). This set of characteristics is spe- cific to species belonging to the genus, and is not evi- Remarks – The absence of eyes, presence of hyper- dent within other Duvalius and Duvalius-like taxa. trophied head, deep and complete frontal furrows, Lohaj et al. (2013) singled out certain of its morpho- smooth genae, absence of longitudinal fissure on 1684 S. B. ĆURČIĆ ET AL. fore tibias, presence of two pairs of elytral discal se- Diagnosis – As presented in the paper of Ćurčić tae, hind pronotal angles that are not pointed, posi- et al. (2003b). tion of the humeral umbilicate setae (as stated in the description of the genus; Ćurčić & Brajković, 2003: Distribution – The genus Javorella, comb. n. is cur- 28P), and presence of a bifid copulatory piece, con- rently known from two caves and deep soil in south- firm that this species belongs to the genus Cur- western and western Serbia. cicia, comb. n., as stated by Ćurčić & Brajković (2003). Lohaj et al. (2013) erroneously proposed Remarks – This genus differs from all other pheneti- including the species in the genus Duvalius. It is of cally close genera (including Duvalius) by a set of interest to underline that even Pretner (1963) and correlative morphological features as listed in Ćurčić Pavićević & Popović (2001) claimed that Curcicia bo- et al. (2003b). This set of characteristics is specific lei, comb. n. has an isolated position in the subgenus to species belonging to the genus, and is not evident Duvalius and could not be associated with any of the within other Duvalius and Duvalius-like taxa. Lohaj existing congeners. This indicates that the authors et al. (2013) singled out certain of its morphological were aware of the supraspecific position of the taxon features and mentioned that they are present among (Ćurčić and Brajković, 2003). other Duvalius taxa, but the characters need to be studied in a complex way. They overlooked the fact GENUS JAVORELLA ĆURČIĆ, BRAJKOVIĆ & that the set of correlative morphological features of ĆURČIĆ, 2003, COMB. N. the genus is unique and do not exist in any other trechine genera. Therefore, we refute its synonymi- Etymology – After Mt. Javor, its terra typica. zation and inclusion in the species within Duvalius. Janák & Moravec (2008) are of the opinion that the Type species – Javorella javorensis Ćurčić, Brajković & recently described trechine Duvalius-like genera Ćurčić, 2003, comb. n. from Serbia probably represent either groups of spe- cies or subgenera. Other species – J. suvoborensis (Pavićević & Popović, 2001), comb. n. and J. suvodolensis Ćurčić, Brajković The genusJavorella , comb. n. probably belongs to & Ćurčić, 2003, comb. n. an old and separate phyletic lineage which originated during the Tertiary (Ćurčić et al., 2003b). The taxon Synonyms – Subgenus Duvalius Delarouzée, 1859 is both an endemic and relict of southwestern and (part.): Pavićević & Popović, 2001: 1; genus Duvalius western Serbia and the Balkan Peninsula. Delarouzée, 1859 (part.): Lohaj et al., 2013: 100. JAVORELLA JAVORENSIS ĆURČIĆ, BRAJKOVIĆ Old combinations – Duvalius (Duvalius): Pavićević & & ĆURČIĆ, 2003, COMB. N. Popović, 2001: 1; Duvalius: Lohaj et al., 2013: 100. Old combination – Duvalius javorensis: Lohaj et al., Type locality – Pećina pod Kapilijama Cave, village of 2013, 100. Trudovo, near Nova Varoš, Mt. Javor, southwestern Serbia. Type locality – Pećina pod Kapilijama Cave, village of Trudovo, near Nova Varoš, Mt. Javor, southwestern Type series – Pećina pod Kapilijama Cave, village of Serbia. Trudovo, near Nova Varoš, Mt. Javor, southwestern Serbia, 20 August 2003, leg. S. Ćurčić (holotype male Other localities – None. and four paratype females); idem, August 2002, leg. S. Ćurčić & B. Mitić (allotype female and a paratype Description and diagnosis – As presented in the paper female). of Ćurčić et al. (2003b). REMARKS ON SOME TRECHINE GROUND BEETLE TAXA FROM THE BALKAN PENINSULA 1685

Distribution – This species is troglobitic and inhabits Duvalius) by Pavićević & Popović (2001), as well as the Pećina pod Kapilijama Cave on Mt. Javor, south- in the genus Duvalius (without assignment to the western Serbia. subgenus) by Lohaj et al. (2013). Even Pavićević and Popović (2001) think that the species has an isolated Remarks – The presence of reduced eyes with depig- position in the subgenus Duvalius and cannot be as- mented ommatidia, pigmented eye border, deep and sociated with any of existing species groups. complete frontal furrows, smooth genae, longitudinal fissure on fore tibias, two pairs of elytral discal setae, JAVORELLA SUVODOLENSIS ĆURČIĆ, position of the humeral umbilicate setae (as stated BRAJKOVIĆ & ĆURČIĆ, 2003, COMB. N. in description of the genus; Ćurčić et al., 2003b: 16), and the presence of a unifid gutter-formed copulato- Old combination – Duvalius suvodolensis: Lohaj et ry piece, confirm the fact that this species belongs to al., 2013, 100. the genus Javorella, comb. n., as originally stated by Ćurčić et al. (2003b). Lohaj et al. (2013) erroneously Type locality – From under stones, Suvodol valley, proposed the inclusion of all of the Javorella species village of , near Nova Varoš, Mt. Javor, south- in the genus Duvalius. western Serbia.

JAVORELLA SUVOBORENSIS (PAVIĆEVIĆ & Other localities – None. POPOVIĆ, 2001), COMB. N. Description and diagnosis – As presented in the paper Old combinations – Duvalius (Duvalius) suvoboren- of Ćurčić et al. (2003a). sis: Pavićević & Popović, 2001, 1; Duvalius suvobo- rensis: Lohaj et al., 2013, 100. Distribution – This species is endogean and can be found under stones in valleys on Mt. Javor, south- Type locality – Cave in Brezaci, village of Brezaci, Ra- western Serbia. jac, near Valjevo, Mt. Suvobor, western Serbia. Remarks – The presence of reduced eyes with depig- Other localities – None. mented ommatidia, pigmented eye border, deep and complete frontal furrows, smooth genae, longitudi- Description and diagnosis – As presented in the paper nal fissure on fore tibias, two pairs of elytral discal of Pavićević & Popović (2001). setae, and the position of the humeral umbilicate se- tae (as stated in the description of the genus; Ćurčić Distribution – This species is troglobitic and inhabits et al., 2003b: 16) confirm that this species belongs to the Cave in Brezaci on Mt. Suvobor, western Serbia. the genus Javorella, comb. n., as originally stated by Ćurčić et al. (2003a). Lohaj et al. (2013) erroneously Remarks – The presence of reduced eyes with dep- proposed including all the Javorella species in the ge- igmented ommatidia, pigmented eye border, deep nus Duvalius. and complete frontal furrows, smooth genae, longi- tudinal fissure on fore tibias, two pairs of elytral dis- Acknowledgments – This study was financially supported by cal setae, position of the humeral umbilicate setae the Serbian Ministry of Education, Science, and Technologi- (as stated in the description of the genus; Ćurčić et cal Development (Grant No. 173038). al., 2003b: 16), and the presence of an unifid gutter- formed copulatory piece, confirm that this species REFERENCES belongs to the genus Javorella, comb. n., as stated by Ćurčić et al. (2003b). This species was previously er- Ćurčić, S. B. and M. M. Brajković (2003). Curcicia, a new genus of roneously included in the subgenus Duvalius (genus endemic ground beetles (Trechini, Carabidae, Coleoptera) 1686 S. B. ĆURČIĆ ET AL.

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