NEW CAVE-DWELLING SPECIES OF THE GENUS REMYELLA JEANNEL (LEPTODIRINI, LEIODIDAE, COLEOPTERA) FROM SERBIA

The new leiodid beetle species, Remyella raskae n. sp. and Remyella javorensis n. sp. (both from caves in Southwest Serbia, region of Sandžak), are described and diagnosed. Adult genitalia and other taxonomically important characters are illustrated. The new species studied are clearly distinct from their closest congeners. The new forms are of Tertiary or even Pre-Tertiary age and originated during the Alpine Orogeny, which affected vast areas of the Balkan Peninsula, including the Dinarides, their terra typica. The new species represent endemic relicts inhabiting Southwest Serbia. Thorough analysis indicates that the new Remyella taxa originated in the proto-Balkan region; their present distributions correspond to at least a part of their primordial ranges (and habitats).


INTRODUCTION
The genus Remyella Jeannel, 1931, comprises one species (Remyella scaphoides Jeannel) and five subspecies, which are distributed over a narrow Dinaric area (the Pešter Polje) in Serbia and Montenegro (P e r r e a u , 2000; L ö b l and S m e t a n a , 2004; Ć u r č i ć , 2005).All Remyella taxa are endemic and live only in cave habitats (J e a n n e l , 1931, 1934; W i n k l e r , 1933; P r e t n e r , 1968; G i a c h i n o and E t o n t i , 1995).The following Remyella subspecies are presently known: R. scaphoides borensis Winkler, 1933, inhabiting the Špela Bor Cave (village of Ugao near Tutin, Pešter Polje, Southwest Serbia), the Ledenica Cave (village of Korita near Bijelo Polje, Pešter Polje, Northeast Montenegro), the Špela Hajnit Cave (village of Ugao near Tutin, Pešter Polje, Southwest Serbia), and the Pećina u Vrh Livade Radojeve Cave (village of Korita near Bijelo Polje, Pešter Polje, Northeast Montenegro); R. scaphoides droveniki Giachino & Etonti, 1995, from the Pećina u Anin Kapeš Cave, the Pećina ispod Gluare Cave, the Uleva Pećina Cave, and the Uleva Pećina III Cave (all in the village of Doliće near Sjenica, Pešter Polje, Southwest Serbia); R. scaphoides hussoni Jeannel, 1934, inhabiting the Pećina u Hamidovoj Vrtači Cave (village of Doliće near Tutin, Pešter Polje, Southwest Serbia); R. scaphoides propiformis Winkler, 1933, from the Špela Maja Hajne Cave and the Kršikuće Cave (both in the village of Ugao near Tutin, Pešter Polje, Southwest Serbia); and R. scaphoides scaphoides Jeannel, 1931, inhabiting the Velika Pećina Cave (village of Grgaje near Sjenica, Pešter Polje, Southwest Serbia) (P e r r e a u , 2000; J e a n n e l , 1931, 1934; W i n k l e r , 1933; P r e t n e r , 1968; G i a c h i n o and E t o n t i , 1995; G u é o r g u i e v , 1990 rounded and more setose); shape of spiculum of the eighth ventrite in females (onion-shaped vs. ginkgo leaf-shaped vs. ginkgo leaf-shaped); form of the median lobe (widened below apex, strongly narrowing apically in lateral view, its apex not dragged, and its lateral sides slightly concave apically vs. widened below apex/with sub-parallel sides, strongly/gradually narrowing apically in lateral view, its apex subacute and dragged, and its lateral sides mostly straight apically vs. widened below apex, strongly narrowing apically in lateral view, its apex not dragged, and its lateral sides slightly concave apically); length of the Description.-Medium-sized.Body length: 4.21 mm (3.88-4.45mm).Body scaphoid; elytra elongated, convex, and without physogastry (Fig. 1).Body color yellowish-brown.Tegument shiny.Both head and pronotum with polygonal microsculptures.
Abdomen with seven visible segments in males and six in females.Male abdominal sternite IX (urite) complete, in form of weakly sclerotized pleuro-tergite with membranous area ventrally, carrying a few setae apically.Spiculum of eighth ventrite in females wide, onion-shaped, with thin base and rounded top.
Aedeagus small, weakly sclerotized, with straight median lobe in lateral view, narrowing distally, with flattened apex curved dorsally (Fig. 2).Median lobe wide and stout dorsally, narrowing gradually at first and then suddenly to form a subacute apex distally.Paramerae somewhat curved exteriorwards and narrowing distally, longer than median lobe (Fig. 2).Parameral setae subequal and equidistant: one preapical dorsal, one proximal dorsal, and one ventral.Tegumen well-sclerotized, somewhat elongated, with hyaline rounded ventral lamina.Inner sac leaving median lobe ventrally and exceeding its length, carrying two weak sclerifications subapically.Endophalus in form of two median sclerified phanerae and a basal phanera at the place of ductus insertion in the inner sac.
Gonostyli straight.Each stylus with a single apical seta, three inner setae, and one outer seta.Two inner setae close-set, while the third inner seta is well-distanced.Outer seta situated at level of first inner seta.Spermatheca small, completely hyaline, not sclerified, elongately curved, with rounded dilated apex.Ductus long and thin; accessory gland in hyaline form.
Bionomy and distribution.-This species was found under stones and on walls in the middle part of the Pećina na Vrelu Raške Cave near Novi Pazar, Pešter Polje, Southwest Serbia.We noticed that the new species prefers wet cave walls and floor around the bed of the Raška River, which runs through the cave.This beetle feeds on filtrated organic matter found on the cave's wet walls and floor.
The new species probably belongs to an old phyletic lineage of Tertiary (or Pre-Tertiary) origin.This species is both relict and endemic to Southwest Serbia, like other known Remyella taxa inhabiting some cave habitats in a limited area of the Dinarides in the Western Balkans (G u é o r g u i e v , 1977).
-After Mt.Javor (Southwest Serbia), its terra typica.We maintain that the genus Remyella is in need of a new revision that will definitely show the real status of all its congeners.Most present subspecies of this genus each deserve a higher (species) rank in future classification.However, we still lack sufficient data on some mostly morphological and anatomical characters (shape of the head, length and shape of certain antennomeres, shape of the urite and spiculum, shape of the aedeagus, length and form of the paramerae, form of the inner sac with the copulatory piece, and form of the female genitalia) to prove our assertion.Only after detailed study based on a larger sample of the analyzed leptodirines will it be possible to ascertain the real status of all taxa belonging to the genus Remyella.
and III; antennomere III 1.77 (1.56-1.94)times longer than antennomere II.Antennomeres IV-VI longer than the preceding ones, antennomere IV being somewhat longer than the others.Antennomere VIII shorter and narrower than antennomeres VII and IX.
Legs very long and thin, with femora thickened basally.Protibiae apically curved exteriorwards, with setose apical border and inner polydentate spine each.Meso-and metatibiae straight.Protarsi pentamerous and not dilated in males.
Abdomen with seven visible segments in males and six in females.Male abdominal sternite IX (urite) complete, in form of weakly sclerotized pleuro-tergite with membranous area ventrally, carrying a few setae apically.Spiculum of eighth ventrite in females wide, ginkgo leaf-shaped, with thin base and rounded top.
Aedeagus small, weakly sclerotized, with straight median lobe in lateral view, narrowing distally, with flattened apex curved dorsally (Fig. 4).Median lobe wide and stout dorsally, narrowing gradually at first and then suddenly to form a subacute apex distally.Paramerae somewhat curved exteriorwards and narrowing distally, longer than median lobe (Fig. 4).Parameral setae subequal and equidistant: one preapical dorsal, one proximal dorsal, and one ventral.Tegumen well-sclerotized, somewhat elongated, with hyaline rounded ventral lamina.Inner sac leaving median lobe ventrally and exceeding its length, carrying two weak sclerifications subapically.Endophalus in form of two median sclerified phanerae and a basal phanera at the place of ductus insertion in the inner sac.
Gonostyli straight.Each stylus with a single apical seta, three inner setae, and one outer seta.Two inner setae close-set, while the third inner seta is well-distanced.Outer seta situated at level of first inner seta.Spermatheca small, completely hyaline, not sclerified, elongately curved, with rounded dilated apex.Ductus long and thin; accessory gland in hyaline form.
Bionomy and distribution.-This species was found on the wet floor and both under and on stones in the posterior part of the Baždarska Pećina Cave, village of Ursule, Mt.Javor, near Sjenica, Southwest Serbia.This beetle feeds on filtrated organic matter found on the cave's wet floor and rocks.
The new species probably belongs to an old phyletic lineage of Tertiary (or Pre-Tertiary) origin.This species is both relict and endemic to Mt. Javor in Southwest Serbia.The endemic differentiation of Remyella and its related genera on the Balkan Peninsula was facilitated by the great Alpine Orogeny, paleoclimatic events, and subsequent evolution of the underground karstic relief, which yielded many new epigean and hypogean niches suitable for the preservation of this old and autochthonous fauna.
Ultrastructure Research, Faculty of Life Sciences, University of Vienna) for their valuable help in taking digital photos of the analyzed beetles.Last but not least, we also appreciate the help of Dr. Heinrich Schönmann (Natural History Museum, Vienna), who loaned us different Remyella taxa important for comparison with the new species.This study was financially supported by the Serbian Ministry of Science (Grant No. 143053).
). Remyella raskae n. sp. and Remyella javorensis n. sp.The present study gives descriptions and diagnoses of the new species of the genus Remyella.The diagnosis of Remyella raskae n. sp. is based on thorough analysis of the type series of six males and four females, collected during 2005 in the Pećina na Vrelu Raške Cave near Novi Pazar, Pešter Polje, Southwest Serbia.The diagnosis of Remyella javorensis n. sp. is based on thorough analysis of the type series of seven males and 27 females, collected during 2005 in the Baždarska Pećina Cave, village of Ursule, Mt.Javor, near Sjenica, Southwest Serbia.MATERIAL AND METHODS The specimens of Remyella raskae n. sp. were collected by hand in the Pećina na Vrelu Raške Cave near Novi Pazar, Pešter Polje, Southwest Serbia, while those of Remyella javorensis n. sp. were collected (also by hand) in the Baždarska Pećina Cave, village or Ursule, Mt.Javor near Sjenica, Southwest Serbia.
median lobe (not reaching level of proximal dorsal parameral seta vs. mostly reaching level of proximal dorsal parameral seta vs. reaching level of proximal dorsal parameral seta); distribution of the parameral setae (all setae equidistant vs. ventral param- eral seta situated mostly at level between two dorsal setae/closer to preapical dorsal seta, rarely closer to proximal dorsal seta vs. setae equidistant); form of the inner sac (with well-sclerotized phanerae, its outer part carrying two weak sclerifications sub-apically vs. mostly with weakly-sclerotized phanerae, its outer part rarely with two weak sclerifications subapically vs. with well-sclerotized phanerae, its outer part carrying two weak sclerifications sub-apically); form of the tegumen (relatively elongated vs. relatively elongated/rounded vs. relatively elongated); position of the outer gonostyl seta (situated at level of the first inner seta vs. situated at level between the first and second inner setae vs. situated at level of the first inner seta); form of the spermatheca (more elongated, slightly widening distally vs. not so elongated, less widening distally vs. less curved, tube-shaped, with a small rounded top); and species distribution (Pešter Polje vs. Pešter Polje vs. Mt.Javor) (J e a n n e l , 1931, 1934; W i n k l e r , 1933; G i a c h i n o and E t o n t i , 1995) (Figs.1-4).
Material examined.-Holotypemale, from the Baždarska Pećina Cave, village of Ursule, Mt.Javor, near Sjenica, Southwest Serbia, 25.8.2005, leg.S. Ćurčić; six paratype males and 27 paratype females, same data as holotype, leg.S. Ćurčić, B. Ćurčić, and N. Ćurčić.All type specimens are deposited in the collection of the Center for Biospeleology of Southeast Europe, Belgrade, Serbia (CBSEE-06/27-60).Diagnosis.-The new species clearly differs from all its congeners.There are two species phenetically most similar to it, both from underground habitats in Southwest Serbia and Northeast Montenegro: R. scaphoides and R. raskae n. sp.The numerous distinctions between the three analyzed species are presented above in the diagnosis of Remyella raskae n. sp.