ZooKeys 278: 91–104, doi: 10.3897/zookeys.278.4650
A new species of the genus Duvalius sg. Neoduvalius from Montenegro with taxonomical remarks on the genus Duvalius (Coleoptera, Carabidae, Trechini)
Roman Lohaj 1,†, Dávid Čeplík 2,‡, Ján Lakota 3,§
1 nstitute of Forensic Sciences, Sklabinská 1, SK-812 72 Bratislava, , Slovakia
2 Južná trieda 13, 040 01 Košice, Slovakia
3 Generála Miloša Vesela 5304/1, SK-034 01 Ružomberok, Slovakia

Corresponding author: Roman Lohaj (rlohaj@gmail.com)

Academic editor: A. Casale

received 9 January 2013 | accepted 5 March 2013 | Published 21 March 2013

(C) 2013 Roman Lohaj. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.


Duvalius (sg. Neoduvalius) gejzadunayi sp. n. from Pećina u Dubokom potoku cave ( Donje Biševo village near Rožaje, Montenegro), the first known representative of this subgenus from the territory of Montenegro is described, illustrated and compared with the related species of the subgenus Neoduvalius Müller, 1913. This new species is characterised by depigmented, medium sized body, totally reduced eyes, deep and complete frontal furrows, 3–4 pairs of discal setae in third elytral stria, as well as by the shape of aedeagus. Data on the distribution and the ecology of this remarkable species, as well as a check-list of the subgenus Neoduvalius are also provided. Recently described genera Serboduvalius Ćurčić, S. B. Pavićević & Ćurčić, B.P.M., 2001, Rascioduvalius Ćurčić, S. B. Brajković, Mitić & Ćurčić, B.P.M., 2003, Javorella Ćurčić, S. B. Brajković, Ćurčić, B.P.M. & Mitić, 2003 and Curcicia Ćurčić, S. B. & Brajković, 2003 are regarded as junior synonyms of the genus Duvalius Delarouzée.


Duvalius (Neoduvalius) gejzadunayi sp. n. , Serboduvalius, Rascioduvalius, Javorella, Curcicia, subterranean environment, Coleoptera, Carabidae, Trechinae, Duboki potok cave, Rožaje, Montenegro, Serbia, taxonomy, new synonymy


The Dinaric mountain chain, and the Balkan peninsula as a whole, are an impressive hotspot of subterranean biodiversity (see, as examples, Pretner 1969, 1973, 1977, Bedek et al. 2006, Gottstein et al. 2002, Jalžić 1984, 1994, Jalžić et al. 2010). Despite the fact that the cave fauna of this region has systematically been studied for more than one and half centuries new taxa are frequently described. Intensive biospeleological research performed in the Dinaric karst region during last twenty years has led to the discovery of many new subterranean trechine taxa, species and also genera (Monguzzi 1993, Casale and Guéorguiev 1994, Casale and Jalžić 1999, Quéinnec 2008, Quéinnec and Pavićević 2008, Quéinnec et al. 2008, Lohaj and Jalžić 2009, Lohaj and Lakota 2010, Lakota et al. 2010, Casale et al. 2012, Lohaj and Mlejnek 2012).

During biospeleological research in the caves of the vicinity of Rožaje (Montenegro) undertaken by members of association Biospeleologica Slovaca (D. Čeplík, G. Dunay, J. Lakota and R. Lohaj) in 2009, a new species of the trechine genus Duvalius belonging to the sg. Neoduvalius was discovered. This new species is described below.

Material and methods

The morphological features of beetles were examined using Olympus SZ 60 and MBS 10 stereo-microscopes. Male and female genitalia were dissected, cleaned and mounted in Euparal® on transparent slides under the examined specimens. Macrophotographs were taken using a Canon 5D mark II camera. Photographs of genitalia were taken using microscope Leitz Ergolux with Nikon Colpix E4500 digital camera attached and were completed using Helicon Focus software program.


TL total body length (measured from the anterior margin of clypeus to the apex of elytra)

L overall length, from apex of mandibles to apex of elytra, measured along the suture

HL head length (measured from the anterior margin of the clypeus to the neck constriction)

HW maximum width of head

HL/HW ratio length of head/ maximum width of head

PL pronotum length (measured along the median line)

PW maximum width of pronotum, as greatest transverse distance

PL/PW ratio length of pronotum/maximum width of pronotum

EL elytral length (as linear distance measured along the suture from the elytral base to the apex)

EW maximum width of elytra

EL/EW ratio length of elytra/maximum width of elytra


HPM collection of Hrvatski Prirodoslovni Muzej, Zagreb, Croatia (B. Jalžić)

NMP collection of Národní Muzeum Praha, Czech republic

ZSM collection of Zoologische Staatssammlung Munich, Germany

CAC private collection of Achille Casale, Torino, Italy

CDC private collection of Dávid Čeplík, Košice, Slovakia

CGD private collection of Gejza Dunay, Kráľovce, Slovakia

CAG private collection of Artur Gitzen, Neuhofen, Germany

CJL private collection of Ján Lakota, Ružomberok, Slovakia

CRL private collection of Roman Lohaj, Pezinok, Slovakia

CVZ private collection of Vladimír Zieris, Pardubice, Czech republic

hw handwritten

p printed

Forward slash indicates separate labels

The following species have been also studied:

Duvalius (Neoduvalius) eurydice (Schaufuss, 1881): 1♂ labelled: „Croatia (hw)/Coll. Geittner (p)“ (ZSM); 1♀ „05028 (p)/Raduć. (p) 25.VII. (hw)“ (ZSM); 1♂ „Croatia (p)/eurydice (hw)/Sammlung Dr.K.Daniel (p)“ (ZSM); 1♂ „Croatia (p)/Collection Strasser (p)“ (ZSM).

Duvalius (Neoduvalius) guidononveilleri Janák & Moravec, P., 2008: 1♂ Paratype labelled: „Jugoslavija, Srbija, Divčibare–sedlo, 750-900 m, 20.5.1991, V.Zieris lgt.“ (CVZ).

Duvalius (Neoduvalius) klimeschi (Winkler, 1914): 1♀ labelled: „Dinar Alpen Troglav–Geb. A.Winkler (p)/Neoduv. troglavensis Wkl. (hw)“ (ZSM).

Duvalius (Neoduvalius) kodrici (Scheibel, 1938): 1♂1♀ labelled: „Pljesevica grupe NO Lika Cro, L.Weirather, Insbruck (p)/Höhle Nr. (p) 658 (hw) grundbuch, Weirather (p)/Neoduvalius opermanni Schbl. (hw)“ (ZSM, CAC)

Duvalius (Neoduvalius) langhofferi (Csiki, 1913): 1♀ labelled: „Ogulin, Tamnica potok, Donje Dubrave, 22.06.2008, B. Jalžić (hw)“ (HPM); 1♀ „Tamnica potok, D.Dubrave, Kordun, 2.03.1985, leg. Jalžić (hw)“ (HPM); 1♀ „Kordun, Mateško selo, Mijatova jama,  15. trav. 1979, Rada“ (HPM); 1♀ „Kordun, Mateško selo, Mijatova jama“ (HPM).

Duvalius (Neoduvalius) opermanni Scheibel, 1933: 2♂♂ labelled: „CROATIA, Ogulin, Slunj, Stara Kršlja, Dumenčića špilja cave, 21.5.2012 resp. 19.9.2012, J. Lakota, B. Jalžić lgt.“ (CJL, HMP).

Duvalius (Neoduvalius) reitteri (Miller, 1881): 1♂ labelled: „Croatia Stava. (hw)/Reitteri Mil.. (hw)/Coll. Mihók (p)/Neoduvalius reitteri Mill. (hw) Det.Dr.Bokor E. (p)“ (ZSM); 1♀ „Senjsko Bilo Cro., Winkler (p)“ (ZSM); 1♂ „Senjsko Bilo Cro., Winkler (p)/Anophthalmus reitteri Mill. (hw)“ (ZSM); 1♂ „A.Reitteri Mit. Croatien (hw)“ (ZSM); 1♂ „Ostrovica Geb. 8.7.89 (hw)/Reitteri (hw)“ (ZSM); 1♂ „Mogorice (hw)/Croatia (hw)/Reitteri Mill. (hw)/ Sammlung Dr.K.Daniel (p)“ (ZSM); 1♀ „Croatien Likaer Grotten Reitter 79 (p)/Sammlung Müller (p) (ZSM); 8♂♂, 7♀♀: „Bosnia, 1.8.2009, Bosanski Petrovac, Dragišića pećina cave, individually, D.Čeplík, R. Lohaj lgt.“ (CDC, CRL); 6♂♂, 7♀♀: „Croatia, Lika, Gospić, Mogorić willage env., Pčelina špilja cave, 22.5.2012, individually, D.Čeplík, B. Jalžić, J. Lakota, R. Lohaj lgt.“ (HPM, CDC, CJL, CRL).

Duvalius (Neoduvalius) schatzmayri (J. Müller, 1912): 1♀ labelled: „Mračna pećina Prolog Bos. Novak 17.8.89 (p)/Neoduvalius schatzmayri J. Müll. (hw) det. F. Stöcklein 1953 (p)“ (ZSM).

Duvalius (Neoduvalius) starivlahi Guéorguiev, B.V., Čurčić, S.B. & Čurčić, B.P.M., 2000: 1♀ labelled: Serbia, Hadži Prodanova Pećina cave, Raščići, Ivanjica, Leg. B. Mitić, 29.05.2003“ (CRL), 1♂ the same data, but Leg. S. Čurčić, 28.05.2008 (CAG).

Duvalius (Neoduvalius) styx (Apfelbeck, 1904): 1♂ labelled: „Mrzla pec. Plitvic. jez. (hw)/Weir. VII.39 H.658 (hw)“ (ZSM).

Material examined.

Holotype male labelled as follows: „MONTENEGRO, Rožaje, Donje Biševo, Pećina u Dubokom potoku cave, 17.4.–11.8. 2009 traps, R.Lohaj, D.Čeplík & G.Dunay lgt.“ (white label, printed) / „HOLOTYPUS Duvalius (Neoduvalius) gejzadunayi sp. n. R. Lohaj, D. Čeplík & J.Lakota det. 2011“ (red label, printed), (NMP). Paratypes: 6 ♂♂ 7 ♀♀, the same data as Holotype (CDC, CGD, CAG, CJL, CRL, CVZ). All paratypes are labelled with white, printed locality labels and with red printed labels „PARATYPUS Duvalius (Neoduvalius) gejzadunayi sp.nov. R. Lohaj, D.Čeplík & J.Lakota det. 2011“.


A medium sized (L 5.5–6.3 mm), glabrous, depigmented, anophthalmous trechine species with the character states of the genus Duvalius Delarouzée, 1859 of subgenus Neoduvalius J. Müller, 1913 (Fig. 1). Colour reddish-brown, pronotum cordiform, transverse, head large and rounded, with deep, complete frontal furrows (Fig. 3), elytral stria 3 with 3–4 discal setigerous punctures. Close to Duvalius (Neoduvalius) starivlahi Guéorguiev, Ćurčić, S.B. & Ćurčić, B.P.M, 2000, from which is distinguished by several different morphological features (see: Discussion).


TL 5.2–6.0 mm (HT 5.3 mm). Colour reddish-brown, legs, antennae and palpi paler. Glabrous, shiny, head and pronotum with distinct isodiametric microsculpture, microsculpture of elytra with isodiametric and transverse meshes.

Head large, rounded, neck markedly distinct, HL 0.85- 0.97 mm (HT 0.87 mm), HW 1.05–1.22 mm (HT 1.08 mm), slightly narrower than pronotum, index HL/HW 0.79 - 0.81 (HT 0, 80), glabrous, with distinct isodiametric microsculpture. Frontal furrows deep, complete, reaching neck constriction. Head with two pairs of long supraorbital setae, anterior pair behind middle of head length, posterior pair at hind part of head near the neck. Mandibles relatively long and slender, acutely pointed, the right one with tridentate basal teeth. Clypeus with 2 pairs of setae, labrum with three pairs. Eyes completely reduced, in some specimens present as a dark spot. Antennae long and slender, reaching almost half of elytral length, covered with dense decumbent pubescence, antennomere 3 longest, scape and antennomeres 5–10 nearly equally long.

Pronotum glabrous, slightly transverse, with maximum width in anterior fifth, PL 0.95–1.08 mm (HT 1.00 mm), PW 1.13–1.40 mm (HT 1.25 mm), index PL/PW 0.75–0.81 (HT 0.80), on base distinctly narrower than on anterior margin. Sides rounded, before hind angles sinuate, anterior angles rounded, obtuse, posterior sharply pointed. Lateral furrows well developed, deep, with two pairs of setae; anterolateral setae situated in the anterior fifth, basolateral pair before hind angles. Median furrow weakly marked, visible in middle of pronotum.

Elytra glabrous, elongate, almost parallel-sided, with maximum width in middle, EL 2.80–3.25 mm (HT 3.00 mm), EW 1.75–2.10 mm (HT 1.85 mm), index EL/EW 1.58–1.70 (HT 1.62), apically rounded. Shoulders well defined, forming obtuse angle, scutellum small, flat; single pair of basal scutellar setigerous pores present. Elytral striae 1–4 well developed, deep, striae 5 –7 vanished, reduced to rows of foveae. Elytral stria 3 with 3–4 discal setigerous punctures (formula 3 + 3, 3 + 4, 4 + 3 or 4 + 4) and pair of apical ones. Umbilicate series consists of 9 setae on both elytra, formula 4 humeral + 2 middle + 3 apical, humeral group of umbilicate pores aggregated. Ventrites 4–6 glabrous, each with pair of setae on their posterior margins, anal ventrite with pair of setae in males and females.

Legs long, slender, densely pubescent, protibiae with a deep longitudinal furrow on their dorsal side. First two tarsomeres of male protarsi distinctly dilated and toothed at their internal margins. Tarsal claws long and slender, pointed at apex.

Aedeagus (Figs 5, 6) 0.91–1.00 mm long (HT 0.97 mm), median lobe in lateral aspect regularly curved and moderately narrowed apically. Endophallus with copulatory piece widely bilobed both at base and apex, with a reduced bundle of scales in middle. Apex obtuse, widely rounded dorsaly. Parameres slender, length of parameres about half of length of aedeagus, each paramere with four thick apical setae.

Female genitalia: (Fig. 7): without peculiar features, apical segments of gonostyli elongated and slender, regularly curved, at apex pointed, with two dorsal and two ventral spines.


Patronymic, dedicated to our dear friend Gejza Dunay (Kráľovce, Slovakia), member of our biospeleological expeditions on Balkans and one of the discoverers of this new species.


So far known only from the type locality, Duboki potok cave near Rožaje, Montenegro.

Topographic location and ecology.

Rožaje town is situated in the easternmost part of Montenegro and is surrounded by high rugged mountains exceeding 2000 m, notably Bjelasica Mts to the west, Hajla Mts and Suva planina Mts to the south and Mokra Gora Mts to the east. To the north is the more open Pešter Polje hill country.

Pećina u Dubokom Potoku cave is situated at 1180 m near the village of Dinje Biševo, 8km north of Rožaje, on the left side of a deep clough valley with the Duboki Potok (English = „deep creek“) draining it. The Duboki Potok is one of the left-side confluents of Ibar river, which flows from this area to the Black Sea. The cave is inactive (without water course) with entrance of ca 1 × 1, 5 m, followed by spacious hall with wet, black,  clay-like humus floor, but deeper in the cave this changes to an ochre-brown sticky mud. The cave floor is littered throughout with stones, stalagmites and stalactites occur only in the furthest recesses of the cave. Total length of the cave is nearly 60 m. Type series of Duvalius gejzadunayi sp. n. was collected by traps baited with cheese and meat, placed in various parts of the cave.

Associated fauna

Araneae (det. A. Mock, Košice, Slovakia)

    Dysderidae sp.

Pseudoscorpionidea (det. R. Ozimec, Zagreb, Croatia)


        Neobisium (Blothrus) umbratile Beier, 1938

Diplopoda (det. A. Mock, Košice, Slovakia)


        Melogona broelemenni (Verhoeff)

Collembola (det. Ľ. Kováč, Košice, Slovakia)

    Entomobryidae, Entomobryinae:

        Heteromurus nitidus (Templeton, 1835)

        Verhoeffiella media (Loksa et Bogojević, 1967)

    Entomobryidae, Tomocerinae:

        Tomocerus sp.


        Arrhopalites principalis Stach, 1945


        Megalothorax sp.



        Rozajella jovanvladimiri Ćurčić SB et al., 2007 (type locality)

    Staphylinidae, Pselaphinae:

        Bryaxis sp.

Figures 1–4.

1, 3 Duvalius (Neoduvalius) gejzadunayi sp. n., Holotype male, Duboki potok cave 2, 4 Duvalius (Neoduvalius) starivlahi Guéorguiev et al. 2000, Topotype male, Hadži Prodanova pećina cave 1, 2 habitus, dorsal view 3, 4 detail of head and pronotum.

Figures 5–7.

Duvalius (Neoduvalius) gejzadunayi sp. n., 5 aedeagus, left lateral aspect 6 aedeagus, dorsal aspect 7 female genitalia, ventral aspect


Subgenus Neoduvalius includes 20 described species (see Check-list bellow), small to medium sized (4–9 mm), depigmented, glabrous, with large head, prominent temporae, strongly reduced or totally absent eyes and third elytral interval with 2–5 setigerous pores, distributed in subterranean habitats from Croatia (Mala Kapela, Lika and Plješevica) through Bosnia & Herzegovina and Serbia to Montenegro (Rožaje).

Duvalius gejzadunayi sp. n. is closely related to Duvalius starivlahi Guéorguiev et al. 2000, described from Hadži-Prodanova pećina cave near Ivanjica, southwest Serbia. Both species are unique within subgenus Neoduvalius, characterised by peculiar combination of characters: (1) head with deep and complete frontal furrows, (2) elytral disc with 3–4 (exceptionally 5) setigerous pores. Complete frontal furrows are, except these two mentioned species, present only in Duvalius (Neoduvalius) cvijici Jeannel, which possess 2 elytral setigerous pores. Species with 3 elytral discal setae are shared also with Duvalius (Neoduvalius) schatzmayri Müller and Duvalius (Neoduvalius) petraeus Knirsch, but both species possess short, incomplete frontal furrows.

Duvalius (Neoduvalius) starivlahi and Duvalius (Neoduvalius) gejzadunayi sp. n. can be easily separated using the following key:

1(2) Head elongate, subglobose, frontal furrows very deep. Eyes reduced, composed of 8–20 ommatidia (Fig. 4). Aedeagus strongly arcuate, forming an angle of 90o, basal bulb large, apex from dorsal view sinuated and bent (see Guéorguiev et al. 2000). TL 5.1–5.7 mm. Serbia, Ivanjica Duvalius (Neoduvalius) starivlahi Guéorguiev et al. 2000
2(1) Head rounded, frontal furrows shallower, eyes totally absent (Fig. 3). Aedeagus strongly elongate, apex from dorsal view widely rounded (Figs 5, 6). TL 5.2–6 mm. Montenegro, Rožaje Duvalius (Neoduvalius) gejzadunayi sp. n.
Taxonomical notes on the genus Duvalius

During the period 2001–2003, four new Serbian genera: Serboduvalius Ćurčić SB et al., 2001 (based on Serboduvalius dragacevensis Ćurčić SB et al., 2001 and Duvalius Neoduvalius) starivlahi Guéorguiev et al. 2000)); Rascioduvalius Ćurčić SB et al., 2003 (based on Duvalius (Neoduvalius) cvijici Jeannel, 1923); Javorella Ćurčić SB et al., 2003 (based on Duvalius Duvalius s.str.) suvoborensis Pavićević and Popović, 2001) and Curcicia Ćurčić SB and Brajković, 2003, (based on Duvalius Duvalius s.str.) bolei Pretner, 1963) were erected based on species formerly included in the genus Duvalius s.l. (Ćurčić et al. 2001, Ćurčić et al. 2003a, 2003b, Ćurčić and Brajković 2003).

The aforementioned genera were based on morphological features that are present among many other species of various subgenera of Duvalius s.l., namely: flattened and reduced eyes (Serboduvalius, Rascioduvalius, Javorella);anophthalmy (Curcicia); presence of darkened eye border (Serboduvalius, Rascioduvalius, Javorella), presence of deep and complete frontal furrows (Serboduvalius, Rascioduvalius, Javorella), or incomplete frontal furrows (Curcicia), convex or flat genae which are glabrous (Serboduvalius, Javorella, Curcicia);or finely pubescent (Rascioduvalius); first male tarsomere almost twice as long as wide or longer than wide (all genera); presence of 2 pairs of discal elytral setae (Rascioduvalius, Javorella, Curcicia); or 3 (rarely 2–4) pairs of discal elytral setae (Serboduvalius); specific position of humeral setae (all genera, note: humeral group of umbilical series in all these taxa is formed by 4 aggregated setae, as in all other Duvalius representatives); presence of longitudinal furrows on protibiae (Serboduvalius, Rascioduvalius, Javorella); absence of such furrow (Curcicia); specific shape of copulatory piece of median lobe of aedeagus (all genera).

Based on a thorough literature research in combination of specimen examination we conclude that the type species designated for genera Serboduvalius, Rascioduvalius, Javorella and Curcicia do not exhibit any autapomorphies or synapomorphies, respectively, which satisfactorily separate them from other representatives of Duvalius s.l. and warrant erection of new taxa. Therefore the following new synonyms are proposed:

Duvalius Delarouzée, 1859

= Serboduvalius Ćurčić, S. B., Pavićević & Ćurčić, B.P.M., 2001, syn. nov.

= Rascioduvalius Ćurčić, S. B., Brajković, Mitić & Ćurčić, B.P.M., 2003, syn. nov.

= Javorella Ćurčić, S. B. Brajković, Ćurčić, B.P.M., & Mitić, 2003, syn. nov.

= Curcicia Ćurčić S. B. & Brajković, 2003, syn. nov.

Check-list of the genus Duvalius sg. Neoduvalius.

Duvalius Delarouzée, 1859: 65 type species Duvalius raymondi Delarouzée, 1859, subgenus Neoduvalius Müller, J., 1913: 180 type species: Anophthalmus Reitteri L. Miller, 1881

Locality data given here is recorded in the language of the original text.

1 Duvalius (Neoduvalius) bradycephalus Jeannel, 1928: 549 (Duvalius, TL: Herzégovine)

= Duvalius (Neoduvalius) hercegovinensis Knirsch, 1926c: 62 (Neoduvalius, TL: »Herzegovina«)

Distribution: Bosnia & Herzegovina

2 Duvalius (Neoduvalius) cvijici cvijici Jeannel, 1923: 10 (Duvalites, TL: Murtenica planina, vers 1100m. d’alt., en forêt, comm. de Bela Rjeka, département d’Užice (Serbie occidentale))

Distribution: Serbia (Užice, Murtenica plateau)

Duvalius (Neoduvalius) cvijici stopicensis Jeannel, 1923: 11 (Duvalites, TL: grotte Stopića pećina, à Rožanstvo, département d’Užice (Serbie occidentale))

Distribution: Serbia (Užice, Zlatibor Mts)

3 Duvalius (Neoduvalius) cuniculinus Knirsch, 1929: 86 (Neoduvalius, TL: Crna-gora (Herc.)

Distribution: Bosnia & Herzegovina (Glavatićevo)

4 Duvalius (Neoduvalius) dragacevensisĆurčić , S.B., Pavičevć & Ćurčić, B.P.M., 2001: 53 (Serboduvalius, TL: Mala Pećina Cave, village Rti, near Kotraža, the Dragačevo Mts, southwestern Serbia)

Distribution: Serbia (Ivanjica, Dragaćevo Mts)

Note: This species was described based on set of 9 specimens (7 males, 2 females) collected in Mala pećina cave near willage Kotraža. This locality is situated ca 10 km from Hadži-Prodanova pećina cave, type locality for Duvalius (Neoduvalius) starivlahi Guéorguiev et al. Based on presented morphological differencies it is very probable that these two species are conspecific.

5 Duvalius (Neoduvalius) eurydice Schaufuss, 1881: 86 (Anophthalmus, TL: in cavernis Croatiae [=Špilja u Lici (=Špilje Like) (Bedek et al. 2006: 73)])

Distribution: Croatia (Lika)

6 Duvalius (Neoduvalius) gejzadunayi sp. n., TL: Montenegro, Rožaje, Donje Biševo, Pećina u Dubokom potoku cave

Distribution: Montenegro (Rožaje)

7 Duvalius (Neoduvalius) guidononveilleri Janák & Moravec, P., 2008: 12 (Duvalius, TL: Jugoslavija, Srbija: Maljen, Divčibare-sedlo, 700-800m)

Distribution: Serbia (Maljen plateau)

8 Duvalius (Neoduvalius) humerosus Knirsch, 1926: 63 (Neoduvalius, TL: Prenj-planina in cca. 1400 m)

Distribution: Bosnia & Herzegovina (Prenj Mts)

9 Duvalius (Neoduvalius) klimeschi Winkler, 1914: 171 (Trechus, TL: Troglavgebiet (Dinarische Alpen) an der dalmatinisch-bosnischen Grenze in zirka 1500 Meter Höhe)

Distribution: (Croatia, Bosnia & Herzegovina)–Mt. Troglav

10 Duvalius (Neoduvalius) kodrici Scheibel, 1938: 221 (Neoduvalius, TL: Plitvička–Plješevica, Kočevlje (Gotschee))

Distribution: Croatia–Mala Kapela Mts, Plješevica Mts (Pretner 1973: 171)

11 Duvalius (Neoduvalius) langhofferi Csiki, 1913: 386 (Anophthalmus, TL: Croatia: in antro prope Josipdol a Dom [=Špilja u Mekoti (=Plandište jama) (Bedek et al. 2006: 73)])

Distribution: Croatia (Ogulin)

12 Duvalius (Neoduvalius) neumanni Müller, J., 1911 1 (Trechus, TL: Höhle „Dragišica“ bei Petrovac (Bosnien) [=Dragišići pećina])

Distribution: Bosnia & Herzegovina (Bosanski Petrovac, Dragišići)

13 Duvalius (Neoduvalius) opermanni Scheibel, 1933: 241 (Duvalius, TL: Höhle nächst Rakovica in Kroatien [=Dumenčića špilja (Bedek et al. 2006: 21)])

Distribution: Croatia (Rakovica, Slunj)

14 Duvalius (Neoduvalius) petraeus Knirsch, 1927: 51 (Neoduvalius, TL: Muharnica-planina. Süd.-Bosn.)

Distribution: Bosnia & Herzegovina (Muharnica Mts)

15 Duvalius (Neoduvalius) reitteri Miller, 1881: 203 (Anophthalmus, TL: Grotte bei Mogorice, Südkroatien, im Likaner Reg. [=Pčelina špilja (Buljmize, Mogorić, Gospić) (Bedek et al. 2006: 46)])

= Duvalius (Neoduvalius) acherontius Schaufuss, 1881: 36 (Anophthalmus, TL: in cavernis Croatiae) synonymy in Heyden, 1883: 10

Distribution: Croatia (Lika), Bosnia & Herzegovina (Bosnia)

16 Duvalius (Neoduvalius) schatzmayri Müller, J., 1912: 297 (Trechus, TL: Höhle des Prologgebirges an der bosnisch-dalmatinischen Grenze [=Mračna pećina (Pretner 1973: 172)])

Distribution: Bosnia & Herzegovina/Croatia (Dalmatia ): Prolog Mts

17 Duvalius (Neoduvalius) starivlahiGuéorguiev, B.V., Ćurčić, S.B. & Ćurčić, B.P.M., 2000: 227 (Duvalius, TL: Hadži-Prodanova Pećina Cave, v. Raščići, near Ivanjica, cca 650 m)

Distribution: Serbia (Ivanjica)

18 Duvalius (Neoduvalius) styx Apfelbeck, 1904: 139 (Trechus, TL: Bosnien, Höhle bei Vacar Vakuf (Zentral-Bosnien))

Distribution: Bosnia & Herzegovina (Mrkonjic Grad), Croatia (Mala Kapela Mts, Plitvicka jezera, Medvedjak) (Pretner 1973: 172)

19 Duvalius (Neoduvalius) vranensis Breit, 1904: 28 (Trechus, TL: Höhle des Vran-Gebirges in der Herzegovina [=Mijatova jama (Hauser, 2011: 117)])

Distribution: Bosnia & Herzegovina (Vran Mts)

20 Duvalius (Neoduvalius) zlatiborensis Ćurčić S. B., Brajković, Ćurčić, B. P. M., 2005: 65 (Rascioduvalius, TL: Markova (=Ršumska) Pećina cave, village of Gornji Ljubiš, Mt. Zlatibor, Western Serbia)

Distribution: Serbia (Užice, Zlatibor Mts)

Note: This species was described based on two females collected in Markova pećina cave, which is very close (ca 10 km) to Stopića pećina cave, type locality for Duvalius (Neoduvalius) cvijici stopicensis Jeannel and ca 2 km to Murtenica planina Mts., type locality for Duvalius (Neoduvalius) cvijici cvijici Jeannel. Based on presented morphological differencies it is very probable that all these taxa are conspecific, inhabiting both caves and MSS of the region of Zlatibor and Murtenica Mts.


Professor Achille Casale (University of Sassari, Italy) is greatly acknowledged for his critical comments to the manuscript. Authors thank Pavel Krásenský (Chomutov, Czech republic) and Dušan Beňo (Banská Bystrica, Slovakia) for the habitus photos. Special thanks are dedicated to Jon Cooter (Oxford, UK) for the language revision of the text and two anonymous reviewers for their comments on the manuscript. We also thank our colleagues and friends for enabling us to study material from institutional and or private collections, namely Michael Balke and Arnaud Faille (Zoologische Staatssammlung Munich, Germany), Achille Casale (Torino, Italy), Artur Gitzen (Neuhofen, Germany), Branko Jalžić (Hrvatski Prirodoslovni Muzej, Zagreb, Croatia) and Vladimír Zieris (Pardubice, Czech republic).

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