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

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 following 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.


MATERIALS AND METHODS
The specimens of the type species of the genera Serboduvalius, Rascioduvalius, Curcicia, Javorella, and Duvalius were analyzed in laboratories of the Institute of Zoology, University of Belgrade -Faculty of Biology, Belgrade, Serbia.These were dissected and thoroughly studied.Dry specimens were adhered onto paper labels.Male genital structures were fixed in a medium composed of Canada balsam and xylol.All taxonomically important morphological characters were studied for comparison.Additional literature data were taken into account for the comparison of some taxa and the conclusions presented herein (Jeannel, 1928;Pretner, 1963;Magrini, 1998;Ćurčić et al., 2001, 2003b, 2003c;Pavićević & Popović, 2001;Ćurčić and Brajković, 2003;Lohaj et al., 2013).

Synonyms -Subgenus Duvaliotes
Distribution -The genus Serboduvalius, comb.n. is presently known only from a few caves and deep soil in southwestern Serbia and eastern Montenegro (Fig. 1).
Remarks -This genus differs from all other phenetically close genera by a set of correlative morphological features as listed in Ćurčić et al. (2001).This set of characteristics is specific to species belonging to the genus, and is not evident within other Duvalius and Duvalius-like taxa.Lohaj et al. (2013) singled out certain its morphological features and mentioned that they are present among other Duvalius taxa, but the characters need to be studied in a complex way.They overlooked the fact that the set of correlative morphological features of the genus is unique and do not exist in any other trechine genera.
We maintain the opinion of Pavićević & Popović (2001), who claimed that it would be inappropriate and even pretentious to establish a new subgenus until the area of Serbia were better covered (when some related species might possibly be discovered).Janák & Moravec (2008) claimed that the recently described trechine Duvalius-like genera from Serbia probably represent either groups of species or subgenera.The solution of the taxonomic problem lies in a better knowledge of the diversity of the genus Duvalius in the Balkans, and descriptions of additional new species in the area studied will enable us to define the final status of the trechine genera analyzed.The recent finding and description of the third Serboduvalius species, Serboduvalius gejzadunayi, comb.n. (Lohaj et al., 2013), which shares the basic correlative morphological characteristics of Serboduvalius, comb.n. (presence of reduced eyes, deep and complete frontal furrows, smooth genae, and 3-4 pairs of elytral discal setae, the second pair of elytral discal setae closer to the first pair than to the third pair of these setae, the same position of the humeral umbilicate setae, and the presence of a similar gutter-formed copulatory piece which is bilobed at the base) confirmed the fact that the genus is valid.The additional trechine species is added into the genus Serboduvalius, comb.n. as well (S.droveniki, comb.n.), since it possesses all leading characteristics of the genus.
The genus Serboduvalius, comb.n. probably belongs to an old and separate phyletic lineage that originated during Tertiary (Ćurčić et al., 2001).The taxon is both an endemic and relict of southwestern Serbia, eastern Montenegro and the Balkan Peninsula.
Other localities -None.
Description and diagnosis -As presented in the paper of Lohaj et al. (2013).
Distribution -This species is presently known from a cave in the vicinity of the town of Rožaje, eastern Montenegro.  ) and what is the precise position of setae of the humeral umbilicate series?Furthermore, the presence of complete frontal furrows is not characteristic for Neoduvalius taxa, so the new species cannot be associated with it.Additionally, the presence of three pairs of elytral discal setae is very rare in Neoduvalius species, while it is observed in all Serboduvalius species.The presence of four pairs of elytral discal setae (in some specimens of Serboduvalius starivlahi, comb. n., S. gejzadunayi, comb. n., and S. droveniki, comb. n.) is not common in Neoduvalius species.Additionally unclear was the presence/ absence of eyes in Serboduvalius gejzadunayi, comb.n. by Lohaj et al. (2013).At first it was recorded that the eyes are completely reduced (a dark spot is registered in a few specimens) (Lohaj et al., 2013: 96), and then the authors reported that the eyes in the species are totally absent (Lohaj et al., 2013: 99).However, small eye spots can be seen in the species in Figs. 1 and 3 presented in the cited paper (Lohaj et al., 2013: 95).The authors wrongly named the type locality as Duboki Potok Cave instead of Pećina u Dubokom Potoku Cave in the Distribution chapter (Lohaj et al., 2013: 98), the village where the cave is situated (Dinje Biševo instead of Donje Biševo) in the chapter Topographic location and ecology (Lohaj et al., 2013: 98), and erroneously determined the pseudoscorpion species caught at the same locality (Neobisium ninae Ćurčić, Dimitrijević & Tomić was described from the type locality vs. Neobisium umbratile Beier, 1938) (Ćurčić et al., 2007, 2008;Lohaj et al., 2013: 98).

Description and diagnosis.
-As presented in the papers of Guéorguiev et al. (2000) and Ćurčić et al. (2001).
Distribution.-This species is presently known from a cave in the vicinity of the town of Ivanjica, Southwestern Serbia.
Remarks.-Presence of reduced eyes, deep and complete frontal furrows, longitudinal fissure on fore tibias, smooth genae, 3-4 pairs of elytral discal setae, a gutter-formed copulatory piece, and position of both the second pair of elytral discal setae (closer to the first than to the third pair of elytral discal setae) and the humeral umbilicate setae (as stated in description of the genus; Ćurčić et al., 2001: 53) confirm the fact that this form belongs to the genus Serboduvalius, comb.n., as was originally stated by Ćurčić et al. (2001).Totally wrong is the opinion by Lohaj et al. (2013), who stated that these two species are probably conspecific on the basis of morphological differences.A dozen observed differences among the species are listed in the paper of Ćurčić et al. (2001), and additional drawings showing some of them are included there as well.Other localities -None.

SERBODUVALIUS DRAGACEVENSIS
Description and diagnosis -As presented in the paper of Ćurčić et al. (2001).
Distribution This species is presently known from a cave in the Dragačevo Mts., southwestern Serbia.
Remarks -Presence of reduced eyes, deep and complete frontal furrows, longitudinal fissure on fore tibias, smooth genae, three pairs of elytral discal setae, a gutter-formed copulatory piece, and position of both the second pair of elytral discal setae (closer to the first than to the third pair of elytral discal setae) and the humeral umbilicate setae (as stated in description of the genus; Ćurčić et al., 2001: 53) confirm that this species belongs to the genus Serboduvalius, comb.n., as was originally stated by Ćurčić et al. (2001).Furthermore, the precise distance between type localities of Serboduvalius starivlahi, comb.n. and S. dragacevensis, comb.n. is 12.5 km (not around 10 km, as was stated by Lohaj et al., 2013: 101).(Magrini, 1998;Lohaj et al., 2013).

Synonyms -Subgenus Duvalites
Distribution -The genus Rascioduvalius, comb.n. is currently known only from few caves and deep soil in western Serbia.
Remarks -This genus differs from all other phenetically close genera by a set of correlative morphological features as listed in Ćurčić et al. (2003c).This set of characteristics is specific to species belonging to the genus, and is not evident within other Duvalius and Duvalius-like taxa.Lohaj et al. (2013) singled out certain its morphological features and mentioned that they are present among other Duvalius taxa, but the characters need to be studied in a complex way.They overlooked the fact that the set of correlative morphological features of the genus is unique and do not exist in any other trechine genera.Therefore, we refute its synonymization and the inclusion of the species within Duvalius (Neoduvalius).This fact is confirmed by the opinion of Pavićević & Popović (2001), who mentioned that the criteria according to which Rascioduvalius cvijici, comb.n. and R. stopicensis, comb.n. are included into the subgenus Neoduvalius of Duvalius are not satisfactory.However, these authors proposed that the taxa should be transferred into the subgenus Duvalius s. str.Jeannel (1928) himself claimed that Rascioduvalius cvijici, comb.n. has a particular position in the trechine systematics.He was not sure whether it belongs to the subgenus Duvalius (= Duvalites) or Neoduvalius of Duvalius.The same author even admitted that the mentioned species deserves an isolated generic status due to the presence of complete frontal furrows (Jeannel, 1928).Additionally, Pavićević & Popović (2001) said that it would be inappropriate and even pretentious to establish a new subgenus until the area of Serbia were better covered (when some related species might possibly discovered).Janák & Moravec (2008) claimed that the recently described trechine Duvalius-like genera from Serbia probably represent either groups of species or subgenera.The solution of the taxonomic problem lies in a better knowledge of the diversity of the genus Duvalius in the Balkans.Descriptions of additional new species in the area studied will enable us to define the final status of the trechine genera analyzed.The recent finding and description of the third Rascioduvalius species, R. zlatiborensis, comb.n., which shares the basic correlative morphological characteristics of Rascioduvalius, comb.n. (presence of reduced eyes with depigmented ommatidia, deep and complete frontal furrows, longitudinal fissure on fore tibias, curved pre-ocular furrow, hairy genae, two pairs of elytral discal setae, the same position of the humeral umbilicate setae, and the presence of a similar bifid gutter-formed copulatory piece) confirmed the fact that the genus is valid.
The genus Rascioduvalius, comb.n. probably belongs to an old and separate phyletic lineage which originated during the Tertiary (Ćurčić et al., 2003c).The taxon is both an endemic and relict of western Serbia and the Balkan Peninsula.(JEANNEL, 1923), COMB.N.
Other localities -None.
Distribution -This species is endogean and can be found under stones in forests on Mt.Murtenica, western Serbia.
Remarks -The presence of reduced eyes with depigmented ommatidia, deep and complete frontal furrows, longitudinal fissure on fore tibias, curved preocular furrow, hairy genae, two pairs of elytral discal setae, position of the humeral umbilicate setae (as stated in description of the genus; Ćurčić et al., 2003c: 484), and presence of a bifid gutter-formed copulatory piece, confirm the fact that this species belongs to the genus Rascioduvalius, comb.n., as previously stated by Ćurčić et al. (2003c).Jeannel (1928) and Lohaj et al. (2013) proposed its including in the subgenus Neoduvalius, genus Duvalius, but this is wrong because the species possesses complete frontal furrows (vs.incomplete ones are present in Neoduvalius species).Our opinion is supported by Pavićević & Popović (2001), who mentioned that the transfer is under doubt.Lohaj et al. (2013) erroneously stated that Rascioduvalius cvijici, comb.n., R. stopicensis, comb.n., and R. zlatiborensis, comb.n., are probably conspecific (inhabiting both caves and MSS on Mts. Zlatibor and Murtenica, western Serbia) on the basis of morphological differences, but without any argumentation.A dozen observed differences among the species are listed in the papers of Jeannel (1923Jeannel ( , 1928) ) and Ćurčić et al. (2003c, 2005), along with drawings showing some of them.Additionally, the distance between the type localities of Rascioduvalius cvijici, comb.n. (village of Bela Reka, Mt.Murtenica) and R. zlatiborensis, comb.n. (Markova Pećina Cave, village of Gornji Ljubiš, Mt.Zlatibor) is around 8 km (not around 2 km, as was stated by Lohaj et al., 2013).
Other localities -None.
Distribution -This species is troglobitic and can be found under stones in the Stopića Pećina Cave on Mt.Zlatibor, western Serbia.
Remarks -The presence of reduced eyes with depigmented ommatidia, deep and complete frontal furrows, longitudinal fissure on fore tibias, curved preocular furrow, hairy genae, two pairs of elytral discal setae, position of the humeral umbilicate setae (as stated in description of the genus; Ćurčić et al., 2003c: 484), and the presence of a bifid gutter-formed copu-latory piece, confirm that this species belongs to the genus Rascioduvalius, comb.n., as previously stated by Ćurčić et al. (2003c).Jeannel (1928) and Lohaj et al. (2013) proposed its inclusion as a subspecies of the species cvijici in the subgenus Neoduvalius, genus Duvalius, but this is wrong because it possesses complete frontal furrows (vs.incomplete ones, present in Neoduvalius species).Our opinion is supported by Pavićević & Popović (2001), who mentioned that the transfer is under doubt.The clear differences among Rascioduvalius cvijici, comb.n., R. stopicensis, comb.n., and R. zlatiborensis, comb.n. are mentioned in the papers of Jeannel (1923Jeannel ( , 1928) ) and Ćurčić et al. (2003c, 2005), along with accompanying drawings, and these are distinguished as three well-separated species.
Other localities -None.
Description and diagnosis -As presented in the paper of Ćurčić et al. (2005).
Remarks -The presence of reduced eyes with depigmented ommatidia, deep and complete frontal furrows, longitudinal fissure on fore tibias, curved pre-ocular furrow, hairy genae, two pairs of elytral discal setae, and the position of the humeral umbilicate setae (as stated in description of the genus; Ćurčić et al., 2003c: 484) confirm that this species belongs to the genus Rascioduvalius, comb.n., as originally stated by Ćurčić et al. (2003c).Type species -Curcicia bolei (Pretner, 1963), comb.n.

Synonyms
Distribution -The genus Curcicia, comb.n. is currently known only from a cave in eastern Serbia.
Remarks -This genus differs from all other phenetically close genera (including Duvalius) by a set of correlative morphological features as listed in Ćurčić & Brajković (2003).This set of characteristics is specific to species belonging to the genus, and is not evident within other Duvalius and Duvalius-like taxa.Lohaj et al. (2013) singled out certain of its morpho-logical features and mentioned that they are present among other Duvalius taxa, but the characters need to be studied in a complex way.They overlooked the fact that the set of correlative morphological features of the genus is unique and do not exist in any other trechine genera.Therefore, we refute its synonymization and inclusion in the species within Duvalius.Janák & Moravec (2008) are of the opinion that the recently described trechine Duvalius-like genera from Serbia probably represent either groups of species or subgenera.The solution of the taxonomic problem lies in a better knowledge of the diversity of the genus Duvalius in the Balkans.Descriptions of additional new species in the area studied will enable us to define the final status of all the trechine genera there.
The genus Curcicia, comb.n. probably belongs to an old and separate phyletic lineage which originated during the Tertiary (Ćurčić & Brajković, 2003).The taxon is both an endemic and relict of eastern Serbia and the Balkan Peninsula.
Other localities -None.
Description and diagnosis -As presented in the paper of Pretner (1963).
Distribution -This species is troglobitic and inhabits the Prekonoška Pećina Cave on the Svrljiške Planine Mts., eastern Serbia.
Remarks -The absence of eyes, presence of hypertrophied head, deep and complete frontal furrows, smooth genae, absence of longitudinal fissure on fore tibias, presence of two pairs of elytral discal setae, hind pronotal angles that are not pointed, position of the humeral umbilicate setae (as stated in the description of the genus; Ćurčić & Brajković, 2003: 28P), and presence of a bifid copulatory piece, confirm that this species belongs to the genus Curcicia, comb.n., as stated by Ćurčić & Brajković (2003).Lohaj et al. (2013) erroneously proposed including the species in the genus Duvalius.It is of interest to underline that even Pretner (1963) and Pavićević & Popović (2001) claimed that Curcicia bolei, comb.n. has an isolated position in the subgenus Duvalius and could not be associated with any of the existing congeners.This indicates that the authors were aware of the supraspecific position of the taxon (Ćurčić and Brajković, 2003).
Distribution -The genus Javorella, comb.n. is currently known from two caves and deep soil in southwestern and western Serbia.
Remarks -This genus differs from all other phenetically close genera (including Duvalius) by a set of correlative morphological features as listed in Ćurčić et al. (2003b).This set of characteristics is specific to species belonging to the genus, and is not evident within other Duvalius and Duvalius-like taxa.Lohaj et al. (2013) singled out certain of its morphological features and mentioned that they are present among other Duvalius taxa, but the characters need to be studied in a complex way.They overlooked the fact that the set of correlative morphological features of the genus is unique and do not exist in any other trechine genera.Therefore, we refute its synonymization and inclusion in the species within Duvalius.Janák & Moravec (2008) are of the opinion that the recently described trechine Duvalius-like genera from Serbia probably represent either groups of species or subgenera.
The genus Javorella, comb.n. probably belongs to an old and separate phyletic lineage which originated during the Tertiary (Ćurčić et al., 2003b).The taxon is both an endemic and relict of southwestern and western Serbia and the Balkan Peninsula.Other localities -None.

JAVORELLA JAVORENSIS
Description and diagnosis -As presented in the paper of Ćurčić et al. (2003b).
Distribution -This species is troglobitic and inhabits the Pećina pod Kapilijama Cave on Mt.Javor, southwestern Serbia.
Remarks -The presence of reduced eyes with depigmented ommatidia, pigmented eye border, deep and complete frontal furrows, smooth genae, longitudinal fissure on fore tibias, two pairs of elytral discal setae, position of the humeral umbilicate setae (as stated in description of the genus; Ćurčić et al., 2003b: 16), and the presence of a unifid gutter-formed copulatory piece, confirm the fact that this species belongs to the genus Javorella, comb.n., as originally stated by Ćurčić et al. (2003b).Lohaj et al. (2013)  Other localities -None.
Description and diagnosis -As presented in the paper of Pavićević & Popović (2001).
Distribution -This species is troglobitic and inhabits the Cave in Brezaci on Mt.Suvobor, western Serbia.
Remarks -The presence of reduced eyes with depigmented ommatidia, pigmented eye border, deep and complete frontal furrows, smooth genae, longitudinal fissure on fore tibias, two pairs of elytral discal setae, position of the humeral umbilicate setae (as stated in the description of the genus; Ćurčić et al., 2003b: 16), and the presence of an unifid gutterformed copulatory piece, confirm that this species belongs to the genus Javorella, comb.n., as stated by Ćurčić et al. (2003b).This species was previously erroneously included in the subgenus Duvalius (genus Duvalius) by Pavićević & Popović (2001), as well as in the genus Duvalius (without assignment to the subgenus) by Lohaj et al. (2013).Even Pavićević and Popović (2001) think that the species has an isolated position in the subgenus Duvalius and cannot be associated with any of existing species groups.
Other localities -None.
Description and diagnosis -As presented in the paper of Ćurčić et al. (2003a).
Distribution -This species is endogean and can be found under stones in valleys on Mt.Javor, southwestern Serbia.
Remarks -The presence of reduced eyes with depigmented ommatidia, pigmented eye border, deep and complete frontal furrows, smooth genae, longitudinal fissure on fore tibias, two pairs of elytral discal setae, and the position of the humeral umbilicate setae (as stated in the description of the genus; Ćurčić et al., 2003b: 16) confirm that this species belongs to the genus Javorella, comb.n., as originally stated by Ćurčić et al. (2003a).Lohaj et al. (2013) erroneously proposed including all the Javorella species in the genus Duvalius.
Pavićević & Popović (2001)d including the species in the subgenus Neoduvalius, genus Duvalius, in spite the fact that it has complete frontal furrows (vs.incomplete ones are present in Neoduvalius species).Pavićević & Popović (2001)also do not support such an opinion.