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Here we present new findings of the hymenopteran Diphyus quadripunctorius (Müller, 1776) from three cave localities in Serbia. We examined a total of five overwintering female specimens collected during 2016, 2019 and 2020 in caves – Devojačka Pećina Cave, Sisevačka Pećina Cave in eastern Serbia and Ogorelička Pećina Cave in southeastern Serbia. Additionally, a short description of D. quadripunctorius females is provided. Its phenology, tentative host association and adaptations to cave life are also discussed., {"references":["Baird, K., & Shaw, M. R. (2019). Overwintering behaviour of Diphyus quadripunctorius (Müller) (Hymenoptera: Ichneumonidae, Ichneumoninae) in south-east Scotland. The Entomologist's Monthly Magazine, 155(4), 217-225. Broad, G. R., Shaw, M. R., & Fitton, M. G. (2018). Ichneumonid wasps (Hymenoptera: Ichneumonidae): their classification and biology. RES handbooks for the identification of British insects, volume 7, part 12. Royal Entomological Society, London, 417 pp. Chappuis, P.-A., & Jeannel, R. (1951). Biospeologica LXXII. Énumération des grottes visitées 1927-1949 (huitième série). Archives de zoologie expérimentale et générale, 88, 81-230. Constantineanu, M. I. (1959). Familia Ichneumonidae: subfamilia Ichneumoninae: tribul Ichneumonini Stenopneusticae. Fauna Republicii Populare Romîne, vol. 9, fasc. 4. Editura Academiei Republicii Populare Romîne, Bucharest, 1248 pp. Ćurčić, B. P. M., Juberthie, C., & Decu, V. (2014). Serbia. pp. 7-46. I Ćurčić, B. P. M. (Ed.): Cave fauna of Serbia, Montenegro, and Macedonia. Monographs, vol. 16. Institute of Zoology, University of Belgrade - Faculty of Biology & Serbian Biological Society, Belgrade, 259 pp. Decu, V., Casale, A., Scaramozzino, P. L., Lopez, F., & Tinaut, A. (1998). Hymenoptera. pp. 1015-1024. In Juberthie, C., & Decu, V. (Eds.): Encyclopaedia biospeologica. Tome II. Société de biospéologie, Bucharest-Moulis, 839-1373 pp. Gauld, I. D. (1984). An introduction to the Ichneumonidae of Australia. British Museum of Natural History, London, 413 pp. https://www.facebook.com/groups/insectserbia/permalink/1206342092902201 https://www.facebook.com/groups/insectserbia/permalink/507737822762635 Legrand, P., Vanderplanck, M., & Verheggen, F. J. (2019). Comparison of the sex pheromone composition of Harmonia axyridis originating from native and invaded areas. Insects, 10(10), 326. Moldovan, O. (2004). Biodiversity in terrestrial cave habitats. pp. 295-299. In Gunn, J. (Ed.): Encyclopaedia of caves and karst sciences. Fitzroy Dearborn, New York, 1940 pp. Nešić, D., & Pavićević, D. (2009). Rezultati novih istraživanja Devojačke pećine (Gaura Fećilor). Zaštita prirode, 60(1-2), 419-429. Nešić, D., Pavićević, D., Zatezalo, A., Mijatović, M., & Grubač, B. (2008). Rezultati kompleksnih istraživanja Ogoreličke pećine. Zaštita prirode, 59(1-2), 51-66. Quicke, D. L. (2015). The braconid and ichneumonid parasitoid wasps: biology, systematics, evolution and ecology. Wiley-Blackwell, Hoboken, 681 pp. Remy, P. A. (1953). Description des grottes yougoslaves (Herzégovine, Dalmatie, Crna Gora et ancien Sandjak de Novi Pazar). Glasnik Prirodnjačkog muzeja srpske zemlje, Serija B, 5-6, 175-233. van Achterberg, C. (2013). Fauna Europaea: Ichneumonoidea. Fauna Europaea version 2017.06. https://fauna-eu.org [Accessed on 15.11.2020]. Vander Meer, R. K., Breed, M. D., Winston, M., & Espelie, K. E. (2019). Pheromone communication in social insects: ants, wasps, bees, and termites. CRC Press, Boca Raton, 384 pp. Vas, Z., & Kutasi, C. (2016). Hymenoptera from caves of Bakony Mountains, Hungary – an overlooked taxon in hypogean research. Subterranean Biology, 19, 31-39. Yu, D. S., van Achterberg, C. V., & Horstmann, K. (2012). Taxapad 2012, Ichneumonoidea 2011. Database on flash-drive. Ottawa, Canada. Žanel, R., & Stanković, S. (1924). Prilog poznavanju pećinske faune i pećina u Srbiji. Glas Srpske kraljevske akademije, 113, 91-107."]}

Kao odgovor na terestrični način života, zglavkari, a naročito insekti, su razvili niz adaptacija koje su omogućile najpre opstanak, a potom i kolonizaciju gotovo svih ekoloških niša u novoj sredini. Insekti su ekstremno diverzifikovana grupa organizama zahvaljujući svakako i činjenici da poseduju veliki biohemijski potencijal. Sekreti koje produkuju insekti imaju najrazličitije namene: u pitanju su iritanti i/ili repelenti protiv predatora, otrovi za imobilizaciju i ubijanje plena, feromoni, itd. Svi pomenuti hemijski produkti, sa ulogom u intra- i interspecijskim interakcijama su najčešće sekundarni metaboliti, tj. produkti poreklom od jedinjenja koja učestvuju u primarnim biohemijskim procesima nastalim različitim hemijskim transformacijama. Veliki asortiman antipredatorske odbrane trčuljaka podrazumeva morfološke, ponašajne i biohemijske strategije. Hemijska odbrana je izuzetno efikasna kod adultnih trčuljaka i podrazumeva izbacivanje produkata pigidijalnih žlezda u vidu odbrambenih sekreta. Pigidijalne žlezde kao parne dorzalne invaginacije telesnog zida u posteriornom delu abdomena su univerzalno prisutne kod Carabidae i ostalih pripadnika nadporodice Caraboidea. Svaka žlezda se sastoji od agregacije sferičnih ili manje-više izduženih sekretornih lobusa (acinusa), u kojima su sekretorne ćelije raspoređene oko centralnog (aksijalnog) sabirnog lumena. Sekreti se dalje sprovode dugim, glavnim sabirnim kanalom do rezervoara za skladištenje. Zidovi rezervoara su obloženi glatkom muskulaturom i promenljive su debljine. Rezervoar se sužava u cilindričnu cev (eferentni kanal), koja se otvara u spoljašnju sredinu. Komponente sekreta detektovane hemijskim analizama moguće je grupisati u deset glavnih kategorija: (1) hinoni, (2) aldehidi, (3) alkoholi, (4) estri, (5) fenoli, (6) karboksilne kiseline, (7) ketoni, (8) monoterpeni, (9) nitrili i (10) ugljovodonici. Ciljevi ove doktorske disertacije usmereni su na dopunjavanje i proširivanje znanja o građi egzokrinih pigidijalnih žlezda i hemijskom sastavu njihovih sekreta kod trčuljaka. U ovoj studiji su analizirane adultne jedinke 27 vrsta trčuljaka iz 13 rodova i pet potporodica sakupljene na teritorijama Srbije i Crne Gore... In response to the terrestrial lifestyle, arthropods (especially insects) developed a series of adaptations that firstly enabled the survival, and then the colonization of almost all ecological niches in the new environment. Insects are an extremely diversified group also due to the fact that they have a great biochemical potential. Secretions produced by insects have a variety of uses: irritants and/or repellents against predators, immobilization and killing of pray, pheromones, etc. All of the mentioned chemical products with the role in intra- and interspecfic interactions are usually secondary metabolites, i.e., products originated from compounds that participate in primary biochemical processes by different chemical transformations. A large assortment of antipredator defense of ground beetles involves morphological, behavioral and biochemical strategies. An extremely effective way of defending against predators in adult beetles is via chemicals which are discharged as pygidial gland products in the form of defense secretions. Pygidial defensive glands are dorsal body wall invaginations in the posterior part of the abdomen, and are universally present in Carabidae and other members of the Caraboidea superfamily. Each gland consists of an aggregation of spherical or more or less elongate secretory lobes (acini), in which secretory cells are arranged around the central (axial) collecting lumen. Secretions are further transported by a long main collecting canal to a storage reservoir. The reservoir walls are coated with smooth muscles of variable thicknesses. The reservoirs are continuing into cylindrical tubes (efferent ducts) which open into the environment. Chemical components of the pygidial secretions in Carabidae can be grouped into ten main categories: (1) quinones, (2) aldehydes, (3) alcohols, (4) esters, (5) phenols, (6) carboxylic acids, (7) ketones, (8) monoterpenes, (9) nitriles, and (10) hydrocarbons. The goals of this doctoral dissertation are focused on complementing and expanding knowledge on the morphological structure of exocrine pygidial glands and the chemical composition of their secretions in ground beetles...

Key to the species of the genus Chaetomargoreicheia 1a. Submedian pronotal discal setae absent, with two pairs of sublateral pronotal discal setae and about 30 less distinct teeth on each lateral elytral margin .............................................................................................................. C. lakotai (Magrini & Bulirsch, 2005) 1b. With two pairs of submedian pronotal discal setae, one pair of sublateral pronotal discal setae or these lacking and less than 30 more distinct teeth on each lateral elytral margin............................................................................................................................ 2 2a. With shorter TL (2.60 mm), one pair of sublateral pronotal discal setae, posteriorly broadened subovoid elytra and about 25 teeth on each lateral elytral margin ..................................................................................................................... C. zoufali (Reitter, 1913) 2b. With longer TL (3.09 mm), no sublateral pronotal discal setae, oval elytra widest at middle and 27–28 teeth on each lateral elytral margin.................................................................................................................................................................. C. gljevensis sp. n.

Ćurčić, Srećko, Pavićević, Dragan, Vesović, Nikola, Rađa, Tonći (2018): A new hypogean species of the genus Chaetomargoreicheia Magrini & Bulirsch, 2005 (Carabidae: Scaritinae: Clivinini) from Croatia. Zootaxa 4438 (1): 183-188, DOI: https://doi.org/10.11646/zootaxa.4438.1.11

Chaetomargoreicheia gljevensis, sp. n. (Figs. 1–6) Type material. Holotype ♂, Jama u Ljučici Cave, village of Gljev, near Sinj, Mt. Kamešnica, 470 m a.s.l., Dalmatia, S Croatia, 22.xi.2016, TR (p) HOLOTYPE Chaetomargoreicheia gljevensis sp. n. Ćurčić, Pavićević, Vesović & Rađa, 2017 (red p) (IZFB). Description. Relatively small, robust, very convex, TL 3.09 mm, EW 0.975 mm. Body rusty-brown, antennae and mouthparts rusty-yellow, legs rusty-red (front legs somewhat darker) (Fig. 1). Integument shiny, with fine polygonal microsculpture. Head robust, narrow, elongate, HL/HW 1.58, narrower and longer than pronotum, HW 0.475 mm (Fig. 1). Anterior margin of clypeus slightly concave, while anterior margin of labrum relatively straight. Labrum with five setae, of which two terminal being the longest. Frontal furrows short, broad, deep, posteriorly divergent. Eyes missing. Cheeks convex and glabrous. Head integument shiny, with barely visible microsculpture. Chaetotaxy: with one pair of clypeal setae and two pairs of supraorbital setae. Antennae elongate, narrow, pubescent, AL 1.16 mm, TL/AL 2.66. Antennomere 1 subcylindrical, somewhat widened basally, A1 0.11 mm, antennomere 2 the longest, slightly curved, A2 (0.18 mm) slightly longer than A3 (0.08 mm) and A4 (0.09 mm) combined, A5 0.09 mm, antennomeres 6–10 moniliform, slightly longer than wide, similar to antennomere 5, terminal antennomere sub-ovoid, A11 0.14 mm (Table 1). Mandibles long, sicklelike. Terminal maxillary palpomere rudimentary, maxillary palps of medium length, slightly pubescent. Pronotum convex, slightly wider than long, PL/PW 0.965, regularly narrowed posteriorly, PW 0.725 mm, widest slightly prior to the middle (Figs. 1 and 2; Table 1). Anterior pronotal margin straight. Lateral pronotal margins arcuate, extended from fore angles to base as a very distinct prebasal groove. Fore pronotal angles prominent, pointed, almost right, hind pronotal angles broadly rounded. Median impression narrow, deep, not reaching anterior pronotal margin and prebasal groove, somewhat widened posteriorly. Lateral marginal gutter very narrow, regular. Proepisterna visible from above basally. Peduncle long and very wide. Disc with polygonal microsculpture. Chaetotaxy: with four marginal setiferous pores slightly out of lateral marginal furrow on each side. First seta situated at the level of anterior pronotal margin, second seta at sixth of pronotal length, third seta prior to the middle of pronotum, and fourth seta at the level of 7/10 of pronotal length. First seta being the shortest. Two submedian discal setae on each side, sublateral setae absent. Legs robust, of medium length (Fig. 1). Each protibia with three outer apical spines, of which distalmost being the longest. Apical spur and distalmost spine of almost equal length, slightly curved. First protarsomere elongate. HT 0.46 mm. First metatarsomere very elongate. Elytra convex, oval, widest around the middle, EL 1.56 mm, EW 0.975 mm, EL/EW 1.60 (Fig. 1; Table 1). Base bordered, sloping, straight in the middle. Shoulders broadly rounded. Suture depressed. Prescutellar setiferous punctures isolated. Lateral marginal gutter very wide, lateral margin with a number (28 on the left side, 27 on the right side) of small well-expressed denticles diminishing in size from base to apex of elytra. Striae impressed, punctuated, while intervals convex, shiny and with fine polygonal microsculpture. Intervals 2–7 with setae (up to 20 per interval). Setae of umbilicate series elongate. Elytral apex rounded. Male genitalia (Figs. 3–6). Aedeagus relatively large. Median lobe curved, somewhat widened apically, slightly impressed dorsally sub-basally, with a distinct, rounded apex in lateral view (Figs. 3 and 4). Median lobe in dorsal and ventral views gradually narrowing basally, slightly curved to the left and more sclerotized apically (Figs. 5 and 6). Inner sac colored, with a number of small teeth. Copulatory piece consisting of three sclerotized lamellae, one dorsal and two ventral, of which one ventral is longer and broader, with a small digit-like, weakly sclerotized distal end directed upwards (Figs. 3–6). Lamellae mutually merged posteriorly. Basal bulb elongate, pointed apically in left lateral and dorsal views (Figs. 3 and 5), rounded in right lateral and ventral views (Figs. 4 and 6). Parameres straight, elongate, of different shape and size, each with two apical setae of similar length. Right paramere massive, longer and broader, left one shorter and narrower (Figs. 3–6). Female genitalia unknown. Differential diagnosis. C. gljevensis sp. n. is closely related to up-to-now known congeners, C. zoufali and C. lakotai, which are distributed in certain mountain ranges in E Herzegovina and S Montenegro (Fig. 7). The new species differs from C. zoufali in the greater value of TL (3.09 mm vs. 2.60 mm), lack of sublateral pronotal discal setae (0 vs. one pair), shape (oval vs. subovoid) and maximum width of the elytra (widest at the middle vs. widest after the middle) and number of teeth on lateral elytral margins (27–28 on each side vs. 25 on each side) (Reitter, 1913; Holdhaus, 1924; Jeannel, 1957; Magrini & Bulirsch, 2005; Bulirsch & Guéorguiev, 2008). Chaetomargoreicheia gljevensis sp. n. differs from C. lakotai in the greater values of TL (3.09 mm vs. 2.41 mm), HW (0.475 mm vs. 0.41 mm), AL (1.16 mm vs. 0.90 mm), PL (0.70 mm vs. 0.58 mm), PW (0.725 mm vs. 0.59 mm), EL (1.56 mm vs. 1.30 mm), EW (0.975 mm vs. 0.90 mm) and EL/EW (1.60 vs. 1.44), lack of sublateral pronotal discal setae (0 vs. two pairs), presence of submedian pronotal discal setae (two pairs vs. 0), shape of fore pronotal angles (almost right vs. obtuse), shape of the shoulders (sloping, less rounded vs. elevate, more rounded), number and size of teeth on lateral elytral margins (27–28 on each side, more distinct vs. about 30 on each side, less distinct), and shape of the aedeagus in right lateral (median lobe more curved, medially thin, apex longer, basal bulb wider vs. median lobe less curved, medially thick, apex shorter, basal bulb narrower) and ventral views (median lobe gradually narrowing basally, apex curved to the right vs. median lobe widened medially, apex curved to the left) (Table 1) (Magrini & Bulirsch, 2005; Bulirsch & Guéorguiev, 2008). Notes. Bulirsch & Guéorguiev (2008) mentioned that the position of the fifth lateral pronotal setae is unclear in C. zoufali —one seta is almost in the marginal gutter on the left and another seta is rather distinctly removed to disc on the right, so it is not quite clear whether C. zoufali possesses four or five pairs of lateral pronotal setae, i.e., one pair of sublateral pronotal discal setae or lacks the setae. Within the diagnosis of the new species with C. zoufali, we presented the numbers of lateral and sublateral pronotal setae for C. zoufali as reported in the original description and other relevant literature sources (Reitter, 1913; Holdhaus, 1924; Jeannel, 1957; Magrini & Bulirsch, 2005). Etymology. The species is named after the village of Gljev, near Sinj (S Croatia), in which is situated the type locality. Type locality. Jama u Ljučici Cave, village of Gljev, near Sinj, Mt. Kamešnica, 470 m a.s.l., Dalmatia, S Croatia. Distribution, habitat, and bionomy. The type specimen was collected by hand in the middle (20 m from the entrance, 4 m below ground surface), totally dark part of the Jama u Ljučici Cave, below a rock, on tuffy and limestone ground with a relatively low level of humidity (Fig. 8). The cave in question is in its front and medium parts relatively dry (except in rainy season), in spite of the fact that a 1–2-m-thick stone layer occurs above it, but the walls in the posterior part are always wet. Tuff amount in the cave is high, which points out to the existence of a strong hydrological activity in the remote past. We have been tried to gather more specimens by pitfall trapping with rotten meat as bait (which is usually used for collecting cave-dwelling and endogean beetles) within the type locality, but any additional efforts were unsuccessful.

Pholeuonopsis (Pholeuonopsis) sljivovicensis, sp. n. (Figs. 1–9) Etymology. After the village of Šljivovica (near Čajetina, Mt. Zlatibor, western Serbia), where the type locality of the new species is situated—the Cave by the Užice-Mt. Tara road. Material examined. Holotype male, seven paratype males, and 17 paratype females collected from pitfall traps with rotten meat as bait in the Cave by the Užice-Mt. Tara road, village of Šljivovica, near Čajetina, 642 m a.s.l., Mt. Zlatibor, western Serbia, 18.09 - 02.11.2013, leg. M. Petković & F. Bosco; a paratype male and a paratype female collected by hand, idem, 18.09. 2013, leg. D. Antić & S. Ćurčić. The type specimens are deposited in the collection of the Institute of Zoology, University of Belgrade - Faculty of Biology, Belgrade, Serbia (IZFB- 14 / 77- 103). Diagnosis. The new species clearly differs from all existing Pholeuonopsis taxa from underground habitats and deep soil in Serbia, Bosnia and Herzegovina, and Montenegro. It is phenetically closest to the three species from Serbia, Pholeuonopsis (Pholeuonopsis) magdelainei, P. (P.) cvijici and P. (P.) zlatiborensis. However, there are numerous distinctions between the new species and these three species, and they are presented below. Pholeuonopsis (Pholeuonopsis) sljivovicensis sp. n. is easily distinguished from P. (P.) magdelainei by its longer body size (3.92 mm in males, 4.08 mm in females vs. 3.80 mm), the antennal length in males (extending beyond the mid-elytra level vs. reaching two-thirds of the elytra), form of the antennomere VIII (more elongate vs. less elongate), shape of the lateral pronotal margins anteriorly (more rounded vs. less rounded), maximum width of the pronotum (slightly before the mid-portion vs. at the hind pronotal angles), shape of the hind pronotal angles (short, less protruding backwards vs. elongate, more protruding backwards), height of the mesosternal carina (high vs. low), shape of the elytra (inversely ovate vs. oval), size of the scutellum (large vs. small), shape of the parameres apically (more widened vs. less widened), position of the parameral setae (almost equidistant vs. the two upper setae being close-set), form of the gonostyli (less curved vs. more curved), position of the outer gonostyl seta (at the level between the two inner setae vs. at the level of the second inner seta), and form of the spermatheca (more curved vs. less curved) (Jeannel 1924; Ćurčić et al. 2006; current paper). Pholeuonopsis (Pholeuonopsis) sljivovicensis sp. n. is easily distinguished from P. (P.) cvijici by its somewhat longer body size (3.62–4.04 mm in males, 3.86–4.42 mm in females vs. 3.81–3.90 mm in males, 3.76–4.09 mm in females), the antennal length (extending beyond the mid-elytra level in males, while not exceeding the mid-elytra level in females vs. extending beyond the mid-elytra level in both sexes), form of the antennomeres VIII (more elongate vs. less elongate) and XI (ovoid vs. sub-elliptical), combined pronotum and elytra/antenna length ratio (1.36 in males and 1.53 in females vs. 1.33 in males and 1.46 in females), the pronotum length/width ratio (0.81 vs. 0.75), shape of the lateral pronotal margins anteriorly (more rounded vs. less rounded), maximum width of the pronotum (slightly before the mid-portion vs. at the hind pronotal angles), shape of the hind pronotal angles (short, less protruding backwards vs. elongate, more protruding backwards), shape of the posterior pronotal margin in males (somewhat convex medially vs. straight), height and shape of the mesosternal carina (high, more rounded, with slightly convex anterior edge vs. low, more acute, with more convex anterior edge), shape of the lateral elytral margins anteriorly (almost straight vs. somewhat impressed), the elytra length/width ratio (1.53 vs. 1.49), length of certain protarsomeres (protarsomere I slightly shorter than the following two protarsomeres combined vs. protarsomere I longer than the following two protarsomeres combined), form of the median lobe (dorsal view) distally (thickened, with more rounded apex vs. not thickened, with more pointed apex), length and shape of the parameres (slightly shorter than the median lobe, more widened distally, sigmoidly curved in lateral view vs. much shorter than the median lobe, less widened distally, regularly arcuate in lateral view), form of the basal bulb in dorsal view (more elongate vs. less elongate), shape of the median lobe in lateral view (less curved, more widened basally vs. more curved, less widened basally), form of the copulatory piece in dorsal view (with two basal short straight structures and one apical sub-triangular structure vs. with two basal and two median long, straight structures, without apical sub-triangular structure), shape of the male abdominal sternite IX (urite) (more oval vs. sub-ovate), form of the gonostyli (less curved vs. more curved), and shape of the spermatheca (more elongate, somewhat narrower vs. less elongate, somewhat thickened) (Ćurčić & Brajković 2002; Ćurčić et al. 2006; current paper). Finally, Pholeuonopsis (Pholeuonopsis) sljivovicensis sp. n. is easily distinguished from P. (P.) zlatiborensis by its longer body size (3.92 mm in males, 4.08 mm in females vs. 3.60 mm in males, 3.90 mm in females), the presence/absence of eye-spots (absent vs. present), the antennal length (extending beyond the mid-elytra level in males, while not exceeding the mid-elytra level in females vs. extending beyond the mid-elytra level in both sexes), form of the antennomere VIII (more elongate vs. less elongate), combined pronotum and elytra/antenna length ratio (1.36 in males and 1.53 in females vs. 1.33 in males and 1.43 in females), shape of the pronotum both anteriorly (more rounded vs. less rounded) and posteriorly (less constricted vs. more constricted), height and form of the mesosternal carina (somewhat lower, more rounded vs. somewhat higher, more pointed), shape of the elytra (inversely ovate vs. ovate), maximum width of the elytra (somewhat anterior to the mid-level in both sexes vs. at the mid-level in males and somewhat anterior to the mid-level in females), the female/male elytral width ratio (1.07 vs. 1.11), shape of the lateral elytral margins anteriorly (almost straight vs. inconspicuously impressed), form of the median lobe (dorsal view) distally (sub-parallel vs. gradually narrowing proximally), shape of the median lobe apex dorsally (shorter, more rounded vs. longer, more pointed), form of the parameres in dorsal view (somewhat curved, more widened distally vs. almost straight, less widened distally), position of the parameral setae (almost equidistant vs. the two lower setae being close-set), form of the basal bulb in dorsal view (more elongate vs. less elongate), form of the copulatory piece in dorsal view (with two basal short, straight structures and one apical subtriangular structure vs. with two basal short, curved structures, without apical sub-triangular structure), form of the male abdominal sternite IX (urite) (more elongate vs. less elongate), form of the gonostyli (somewhat curved vs. almost straight), number of the inner gonostyl setae (two vs. three), distribution of the outer gonostyl seta (situated at the level between the two inner setae vs. situated at the level between the second and third inner setae), and shape of the spermatheca (more curved, somewhat narrower vs. less curved, somewhat wider) (Ćurčić et al. 2006; current paper). Description. Medium-sized. Body length: 3.92 mm (3.62–4.04 mm) in males, 4.08 mm (3.86–4.42 mm) in females. Body pholeuonoid, elliptic, elongated, from yellowish-brown to reddish-brown (Fig. 1). Integument shiny, pubescent, microsculptured. Head elongate, slightly longer than wide, without eyes or eye-spots (Fig. 1). An inconspicuous occipital carina present. Head dorsally covered with numerous densely distributed small punctures and small yellowish erect setae. Vertex with a shallow impression. Labrum emarginate, with a few long setae. Penultimate maxillary palpomere well widened apically, the ultimate one being narrow, short, narrowing apically. Antennae moderately long, slender, widening distally (Fig. 1). Combined pronotum and elytra length to antenna length ratio: 1.36 (males) and 1.53 (females). Antennae inserted at about mid-head level, shorter than the body (Fig. 1), extending beyond the mid-elytra level in males, while not exceeding the mid-elytra level in females. Antennomere I widened apically, shorter than antennomere II. Antennomere II longer than antennomere III. Antennomere VIII the shortest, elongately sub-spherical. Antennomeres I–VI moderately thickening apically, and antennomeres VII and IX–XI well broadened. Distalmost antennomere ovoid, longer than the penultimate one. Pronotum small, sub-bell-shaped, wider than long (length/width ratio 0.81), widest slightly before the midportion, covered with dense, short, yellowish laid setae and impressed punctures (Fig. 1). The lateral margins bordered, sigmoidly shaped. Posterior pronotal margin shorter than the base of elytra. Both anterior and posterior pronotal margins somewhat convex medially. Fore pronotal angles rounded, obtuse, and the hind ones acute, pointed, prominent, somewhat protruding backwards. Pronotal disc weakly convex, with two basal and two median shallow impressions. Mesosternal carina high, sub-triangular, obtuse, carrying a few teeth and a number of setae on the posterior edge (Fig. 2). The anterior edge slightly convex, while the posterior one somewhat concave. Metasternum without carina. Elytra elongate, inversely ovate (length/width ratio 1.53), wider in females than in males (female/male elytral width ratio 1.07) (Fig. 1). The maximum width reached somewhat anterior to mid-elytral level in both sexes. The lateral margins well-chitinized, almost straight anteriorly. Marginal furrows relatively wide, shallow, gradually narrowing towards elytral apex. Humeri obtuse, rounded. Elytral disc convex, gradually declining anteriorly, but steeply declining posteriorly, with numerous short, yellowish laid setae and impressed punctures. Scutellum large, triangular. Elytral apex rounded. Legs long, slender, with the femora thickened basally (Fig. 1). Tibiae somewhat curved, moderately thickening apically. Meso- and metatibiae with a few fine spines each. Protarsi 4 -segmented in both sexes. The first protarsomere nearly three times as long as wide, slightly shorter than the following two protarsomeres combined. Tarsal claws long, thin, sharply pointed. Male genitalia as presented in Figs. 3–5. Aedeagus medium-sized, well-sclerotized, stout (Figs. 3 and 4). Basal bulb well-developed, rounded in dorsal view (Fig. 3), while huge in lateral view (Fig. 4). Median lobe in dorsal view thickened and sub-parallel distally, somewhat narrowing proximally, sub-apically rounded, apically with a pointed apex that is somewhat rounded anteriorly (Fig. 3). Median lobe longer than parameres. Median lobe in lateral view curved in the anterior portion, gradually narrowing apically, with a pointed beak-like apex (Fig. 4). Parameres long, thin, somewhat curved exteriorwards dorsally and sigmoidly curved laterally, narrowing apically, but with a widened apex each (Figs. 3–5). Each paramere with three apical, almost equidistant setae (Fig. 5). Inner sac elongated, tubular. Copulatory piece consisting of a basal phanera, two basal sclerotized short, straight structures and one apical sub-triangular structure, along with two weakly sclerotized bands (Fig. 3). Male abdominal sternite IX (urite) well-developed, oval (Fig. 6). Female genitalia (Figs. 7 and 8). Gonostyli elongated, thin, gradually narrowing distally, somewhat curved (Fig. 7). Each stylus with a single apical seta, two inner setae and one outer seta. The outer seta situated at the level between the two inner setae. Spermatheca small, unique, hook-like, strongly curved, somewhat narrowing apically, well-sclerotized (Fig. 8). Female abdominal segment VIII large, transverse, with a short narrow anterior process, setose (Fig. 9). Bionomy and distribution. The new species was found under rocks and in pitfall traps in the posterior, totally dark part of the Cave by the Užice-Mt. Tara road, village of Šljivovica, near Čajetina, Mt. Zlatibor, western Serbia (Fig. 10). The species prefers wet clay cave walls and floor, where it probably feeds on filtrated organic matter from trickling water. The type locality of the new species is inhabited by the blind leiodid species— Leptinus testaceus Müller, 1817, which usually lives in nests of mammals, birds and hymenopterans, but is frequently found in caves as well (Perreau 2000). The new species probably belongs to an old phyletic lineage of Mesogeid (alpine ranges of southern Europe and Africa) origin, like some other Pholeuonopsis taxa from Serbia (Ćurčić & Brajković 2002; Ćurčić et al. 2006). It belongs to the eastern Mesogeid range (southern Balkan Peninsula and Asia Minor), with the initial activity in the Cretaceous and the main tectonic phase in the Eocene-Miocene (Guéorguiev 1977). The species is both relict and endemic to the Dinarides in western Serbia, like other Serbian congeners inhabiting some cave habitats (Jeannel 1924; Guéorguiev 1977; Ćurčić & Brajković 2002; Ćurčić et al. 2006). The endemic differentiation of Pholeuonopsis spp. and the relative taxa on the Balkan Peninsula was facilitated by the great Alpine Orogeny, paleoclimatic events, and subsequent evolution of the underground karstic relief, which yielded numerous new epigean and hypogean niches suitable for the preservation of the old and autochthonous fauna (Ćurčić et al. 2012, 2013, 2014 a).

Tapinopterus (Tapinopterus) cognatus pancici Ćurčić and Ilić, ssp. n. (Figs. 7���12) Type localities. By the road Jelak Reserve - village of Brzeće, Mt. Kopaonik, 1,100 m a.s.l., Jo��anička Banja, C Serbia; village of Brzeće, Mt. Kopaonik, 1,100 m a.s.l., Jo��anička Banja, C Serbia; village of Đerekari, Mt. Kopaonik, 800 m a.s.l., Brus, C Serbia; village of Razbojna, Mt. Kopaonik, 600 m a.s.l., Blace, C Serbia. Type material. Holotype ♂, by the road Jelak Reserve���village of Brzeće, Mt. Kopaonik, 1,100 m a.s.l., Jo��anička Banja, C Serbia, 28.VI���09.VIII. 2014, from pitfall traps, leg. N. Ilić (p) HOLOTYPE Tapinopterus (Tapinopterus) cognatus pancici ssp. n. Ćurčić & Ilić, 2015 (red p) (IZFB) (Fig. 7). Paratypes: 5 ♂♂ 3 ♀♀, same data as for holotype; 1 ♀, village of Brzeće, Mt. Kopaonik, 1,100 m a.s.l., Jo��anička Banja, C Serbia, 03.VIII. 1996, from pitfall traps, leg. N. Ilić; 4 ♀♀, village of Đerekari, Mt. Kopaonik, 800 m a.s.l., Brus, C Serbia, 03.VIII. 1996, from pitfall traps, leg. N. Ilić; 1 ♂ 1 ♀, village of Razbojna, Mt. Kopaonik, 600 m a.s.l., Blace, C Serbia, 03.VIII. 1996, from pitfall traps, leg. N. Ilić (p) PARATYPE Tapinopterus (Tapinopterus) cognatus pancici ssp. n. Ćurčić & Ilić, 2015 (red p) (IZFB, cNI). Description. BL M 10.53 mm (R 10.10���10.98 mm), BW M 3.46 mm. Head, pronotum, and elytra dark brown, shiny, antennae of similar color, legs and abdomen piceous brown, palpi somewhat paler. Head large, robust, somewhat wider than long (ratio HL/HW M 0.92), smooth, with a fine isodiametric microsculpture. Antennae moderately long, somewhat exceeding elytral humeri, with dense, decumbent pubescence from antennomere 4. Pronotum weakly transverse, somewhat wider than long, ratio PL/PW M 0.90, PL M 2.60 mm (R 2.44���2.75 mm), PW M 2.90 mm (R 2.75���3.05 mm), PB M 1.93 mm (R 1.82���2.07 mm), with rounded lateral margins, smooth, with a fine isodiametric microsculpture, widest slightly before first third. Anterior margin straight or somewhat sinuated, with prominent, at apex rounded angles, basal margin straight or slightly sinuated. Posterior angles prominent, acute, almost rectangular. Pronotal base with single deep longitudinal posterolateral impression on each side of midline, reaching basal third of pronotum. Midline distinct, not reaching basal or anterior margins. Both midline and posterolateral impressions with transversal wrinkles. Elytra oval, with an isodiametric microsculpture, ratio EW/EL M 0.57, with maximum width around at middle, with distinct, weakly punctate striae. Disc relatively flattened basally, while somewhat convex apically. Shoulders prominent, rounded. Second elytral striae with one setigerous pore each situated at the level of �� elytral length. Median lobe of aedeagus in dorsal view relatively long, straight, with apical blade wide, curved to left (Fig. 8), in lateral view arcuate, sub-apically abruptly broadened, with an apex abruptly bent upwards (Fig. 9). Right non-conchoid paramere elongate, of almost equal width over its whole length, curved (Fig. 10), while left conchoid broad, ear-like (Fig. 8). Male abdominal sternite IX (urite) well-developed, ovoid (Fig. 11). Gonocoxites IX long, gradually narrowing distally, somewhat curved, basally joined with massive gonosubcoxites IX (Fig. 12). Etymology. The subspecies is named after Academician Prof. Dr. Josif Pančić, a famous Serbian naturalist and the first president of the Serbian Royal Academy, whose mausoleum is located in the highest peak of Mt. Kopaonik, the terra typica. Diagnosis. The diagnostic differences between Tapinopterus (T.) cognatus pancici ssp. n. and the morphologically and geographically closest taxa are previously presented in the differential diagnosis of T. (T.) cognatus vlasinensis ssp. n. Distribution and habitats. The type specimens have been collected at different altitudes (600���1,100 m a.s.l.) in mixed deciduous-coniferous forests at a few sites on Mt. Kopaonik., Published as part of ��ur��i��, Sre��ko, Ili��, Nastas, Vesovi��, Nikola & Anti��, Dragan, 2015, Two new high-altitude subspecies of Tapinopterus (Tapinopterus) cognatus (Dejean, 1831) (Coleoptera: Carabidae: Pterostichinae) from Serbia, pp. 197-200 in Zootaxa 4034 (1) on pages 199-200, DOI: 10.11646/zootaxa.4034.1.11, http://zenodo.org/record/243749

Ćurčić, Srećko, Ilić, Nastas, Vesović, Nikola, Antić, Dragan (2015): Two new high-altitude subspecies of Tapinopterus (Tapinopterus) cognatus (Dejean, 1831) (Coleoptera: Carabidae: Pterostichinae) from Serbia. Zootaxa 4034 (1): 197-200, DOI: http://dx.doi.org/10.11646/zootaxa.4034.1.11