Tone Novak

• University of Maribor

Autophagy is a highly conserved metabolic process that regulates cellular homeostasis and energy supply by degrading dysfunctional and excess cell constituents and reserve materials into products that are reused in metabolic and biosynthetic pathways. Macroautophagy is the best studied form of autophagy in invertebrates. Starvation is a common stre...

In terms of taxonomy and systematics the harvestman genus Trogulus Latreille, 1802 is one of the most complex arachnid taxa in Europe. Although many problems have been resolved and a number of new species were described since 2008, so far, the diversity within several species subgroups within the genus has not been satisfactorily disentangled. One...

Most subterranean habitats, especially caves, are considered extreme environments, mainlybecause of the limited and erratic food supply and constant darkness. In temperate regions, manyclimatic conditions, such as temperature and air humidity, are periodically less adverse or even morefavourable in caves than the harsh seasonal weather on the surfa...

Amilenus aurantiacus overwinter in diapause, a natural starvation period, in hypogean habitats. The structure of spherites in the midgut diverticula (MD) and Malpighian tubules (MT) has been studied comparatively by light microscopy and TEM to detect eventual differences in mineral consumption in the beginning and at the end of the starvation perio...

The cave crickets Troglophilus neglectus Krauss, 1879 and T. cavicola (Kollar, 1833) originating from caves in Slovenia were surveyed for gregarines. Two gregarine species were identified. Gregarina troglophili (Golemansky & Lipa, 1991) occurred in both cave cricket species, T. cavicola being reported a new host species. Besides, a new species, Gre...

Nemastoma bidentatum Roewer, 1914 is a complex of closely related litter-dwelling harvestmen, characterized by a penis glans with two pairs of lateral lancet-like spines, and a femur IV with a saw-like series of pointed tubercles. Here we a) revise the hitherto known taxa within the N. bidentatum complex, and b) analyze in detail the relations amon...

In arachnids, the Malpighian tubules (MTs), coxal glands and stercoral pockets are capable of collecting and removing excreta from the body. The presence of the MTs among Opiliones was evidenced for the first time in Amilenus aurantiacus in 2015. Individuals undergo a winter diapause subterranean habitats. Here, we provided the morphological and cy...

Two closely related, syntopic species, Leiobunum rupestre and L. subalpinum, spend the day at apparently randomly distributed resting sites. In this preliminary research, we studied differences in their microclimatic preferences at a locality in the Pohorje Mts., Slovenia. We measured air temperature and relative humidity at individual resting site...

In this work, we investigated the relationship between habitat types (HTs) and selected environmental factors in the highlands at altitudes of 1800–2558 m in the Kamnik-Savinja (KS) Alps in Slovenia. For 275 sampling sites, we identified seven HTs in their typical form and 11 ecotones, and provided field and modelled data for 14 environmental facto...

Protura is a small class of Hexapoda, generally poorly known, and rather scarce data on its occurrence in caves are scattered throughout literature on fauna in caves and in some papers on Protura. Although the cave-dwelling fauna is relatively well studied in Slovenia, published records on Protura are rare. In this paper, data on the occurrence and...

Nemaspela Šilhavý, 1966 (Opiliones: Nemastomatidae) is a genus of exclusively troglobiotic harvestmen species inhabiting caves in the Crimea, Caucasus and Balkan Peninsula. In this paper, Nemaspela borkoae sp. nov., recently found in four caves in Montenegro, is described. The new species is characterized by its small body, 1.5–2.1 mm long, and ver...

Fauna of two pegmatitecaves in Slovenia is presented.

Troglobionts are organisms that are specialized for living in a subterranean environment. These organisms reside prevalently in the deepest zones of caves and in shallow subterranean habitats, and complete their entire life cycles therein. Because troglobionts in most caves depend on organic matter resources from the surface, we hypothesized that t...

A faunistic review of the harvestmen (Opiliones) of the Submediterranean region of Slovenia is represented and the current taxonomlcal and zoogeographical problems of some species are discussed.

In the present paper, further findings of harvestmen in the Sub Mediterranean region of Slovenia are presented, and comments on some actual problems with the species are added. With the exception of the possible discovery of a troglobiotic species in the region, all species living there have presumably been recorded. With the exclusion of Trogulus...

The European cave spider, Meta menardi, is a representative of the troglophiles, i.e. non-strictly subterranean organisms. Our aim was to interpret the cytological results from an ecological perspective, and provide a synthesis of the hitherto knowledge about M. menardi into a theory of key features marking it a troglophile. We studied ultrastructu...

Systematic ecological investigations were carried out in the alpine cave Ivačičeva jama (2450 m) near the Meteorological Station at Kredarica (Triglav Mountains, Triglav National Park = TNP, NW Slovenia, YU) once each season from October 1979 until July 1980. The average annual temperature and air humidity were low (-1.0°C, 90 %) and the average ig...

The spermatophores in the sironid species Cyphophthalmus duricorius duricorius JOSEPH, 1868 are presented. They were found in three females in Slovenia. The spermatophores consist of a bulb-like terminal portion, a tubular and a basal portion, and are very similar to those in C. serbicus (HADŽI, 1973).

Abstract - Since 1996, intensive faunal research has taken place in Friuli-Venezia Giulia in the regions of Udine, Pordenone and Gorizia, organised by the Museo Friulano di Storia Naturale in Udine. In addition, some opilionids from other regions have been deposited in the Museum. In this contribution, 46 opilionid species from 112 localities are c...

In Europe there are about 10 Nelima species living especially in the Mediterranean region (Martens 1969, 1978). Until 1998, N. sempronii Szalay, 1951, and N. doriae (Canestrini, 1871) had been the only Nelima species registered in Slovenia (Novak et Gruber 2000). N. sempronii lives in Central Europe and in the Apennines, but in the Eumediterranean...

The European cave spider, Meta menardi, is among the most common troglophile species inhabiting the cave entrance zone in Europe, where prey is scarce in winter. Spiders feed only if prey is available; otherwise, they are subjected to long-term winter starvation. We carried out a four-month winter starvation of M. menardi under controlled condition...

The homologous and phylogenetically old scent glands of harvestmen—also called defensive or repugnatorial glands—represent an ideal system for a model reconstruction of the evolutionary history of exocrine secretion chemistry (“phylogenetic chemosystematics”). While the secretions of Laniatores (mainly phenols, benzoquinones), Cyphophthalmi (naphth...

During the growth period, in surface habitats, spiders catch enough prey to feed normally. In contrast, in the cave entrance zone, prey may be relatively scarce. Meta menardi inhabits this cave section, resulting in temporary starvation. We studied structural changes in the midgut epithelial cells of M. menardi during a short-term and a medium-term...

The herald moths, Scoliopteryx libatrix, overwinter in hypogean habitats. The ultrastructure of their fat body (FB) cells and Malpighian tubule (MT) epithelial cells was studied by light microscopy and transmission electron microscopy, and essential biometric and biochemical measurements were performed. The FB was composed of adipocytes and sparse...

Dr. Jürgen Gruber lenkte über Jahrzehnte die Arachnologie an der 3. Zoologischen Abteilung des Naturhistorischen Museums in Wien. Gemeinsam mit Konrad Thaler (Innsbruck) und Friedrich Barth (Wien) bildeten sie das Triumvirat an hauptberuflichen Arachnologen (excl. Acarologen) in Österreich. Auch die Pensionierung vor nunmehr 15 Jahren konnte Gruber...

Subterranean habitats are known for their rich endemic fauna and high vulnerability to disturbance. Many methods and techniques are used to sample the biodiversity of terrestrial invertebrate fauna in caves, among which pitfall trapping remains one of the most frequently used and effective ones. However, this method has turned out to be harmful to...

In nature conservation, providing new, advanced knowledge of protected habitats and biota often requires extensive sampling, which contradicts the main goal of minimizing damage and disturbance. How to avoid this innate collision was the main concern in our study. Highland habitats are suitable for addressing this problem, since they are relatively...

Subterranean habitats are known for their rich endemic fauna and high vulnerability to disturbance. Many methods and techniques are used to sample the biodiversity of terrestrial invertebrate fauna in caves, among which pitfall trapping remains one of the most frequently used and effective ones. However, this method has turned out to be harmful to...

We present a simple, manageable and cheap method for accelerated disturbance and, consequently, detection of thanatotic and soil-mimetic arachnids and other arthropods. The method combines sieving a substrate in the field, followed by heating and lighting the dispersed sample by a handheld electric light in the laboratory. The name Hot-Dancing allu...

During winter, cave cricket larvae undergo dormancy in subterranean habitats; this dormancy is termed diapause in second year Troglophilus cavicola larvae because they mature during this time, and termed quiescence in T. neglectus, because they mature after dormancy. Here we used electron microscopy to analyze ultrastructural changes in the epithel...

V letih 2000-2001 smo raziskovali favno in ekološke razmere v Železni jami pri Dobu. V jami smo našli 53 vrst, dominantne so bile: Meta menardi, Amilenus auriantiacus, Troglophilus neglectus, T. cavicola, Laemosthenes schreibersi, Ceuthmonocharis robici, Limonia nubeculosa in nedoločena vrsta foride. Troglomorfozni taksoni so bili Zospeum sp., C. r...

Two phalangiid species, Amilenus aurantiacus (Simon, 1881) and Gyas annulatus (Olivier, 1791) (Opiliones, Phalangiidae), spend their winter dormancy in large numbers in subterranean habitats. In caves they rest on the walls and ceilings of the entrance sections. Both are considered trogloxenes, or the latter are sometimes classified as troglophiles...

The cave cricket Troglophilus neglectus regularly overwinters for 4-5 months in hypogean habitats. Winter dormancy is a natural starvation period, providing the opportunity to study autophagy under natural conditions. We aimed to evaluate the autophagic activity in adipocytes and urocytes of the fat body in three time frames: directly before overwi...

Izvleček UDK 595.142:551.442(497.4Huda luknja) Tone Novak, Csaba Csuzdi, Franc Janžekovič, Tanja Pipan, Dušan Devetak & Saša Lipovšek: Preživetje površinskega deževnika Dendrodrilus rubidus tenuis (Oligochaeta: Lum-bricidae) v podzemeljskem okolju V vzorcih curka prenikajoče vode v Hudi luknji pri Doliču je bila peregrina vrsta deževnika dendrodril...

The peregrine earthworm Dendrodrilus rubidus tenuis was regularly sampled in a percolating water drip originating in a habitat inaccessible to humans in the Huda luknja pri Dolicu cave, Slovenia. The reconstruction of this habitat includes both a larger passage with bat colonies supplying wet bat guano sediments, on which the earthworms feed, and n...

The harvestmen Amilenus aurantiacus overwinter in diapause in hypogean habitats. The midgut diverticula have been studied microscopically (light microscopy, TEM) and biochemically (energy-storing compounds: lipids and glycogen) to analyze changes during this programmed starvation period. Throughout the investigated period, the epithelium of the mid...

Most organisms are able to survive shorter or longer exposure to sub-zero temperatures. Hypothetically, trogloxenes characterized as not adapted, and troglophiles as not completely adapted to thermally stable subterranean environment, have retained or partially retained their ability to withstand freezing, while most troglobionts have not. We teste...

A highly specialized, endemic troglobiotic harvestman, Hadzinia ferrani sp. n., recently found in the Ferranova buža cave in the karst of Mt. Ulovka in central Slovenia is described. It is characterized by its small, 1.1−1.4 mm long body and very long, thin appendages, with leg II more than 20-times as long as the body. Although very similar to Nem...

Juvenile harvestmen Gyas annulatus overwinter in dormancy in hypogean habitats for 4-5 months. The ultrastructure of the autophagic structures in their midgut epithelium cells was studied by light-, transmission electron microscopy (TEM) and immunofluorescence microscopy (IFM) during this non-feeding period. Before overwintering (November), autopha...

Although expected since 1980s, no eyeless troglobitic nemastomatid harvestman (Opiliones: Nemastomatidae) species has been known in Slovenia. Here we introduce a highly specialized, endemic troglobitic species Hadzinia ferrani Novak & Kozel, 2014, recently found in the cave Ferranova buža near Vrhnika to the regional entomologists. It is characteri...

Both regular and irregular, i.e., sporadic, absence of the ventral spur (ventral tooth) on the basal cheliceral article has been reported in Oligolophinae. In this contribution, a female Lacinius ephippiatus with one missing ventral spur, and the other partly reduced, on the basal cheliceral article is described. For comparison, we studied chelicer...

Eleven of the most important terrestrial invertebrate species in Slovenian caves were analyzed for differences in their fresh and dry biomass, energy content and carbon bulk during winter. These data were combined with the species abundance in 54 caves and adits in order to estimate their organic carbon bulk and carbon input into these habitats. In...

For the understanding of shrew communities, their coexistence and competition, the knowledge of diet partitioning among sympatric shrew species is crucial. In this study, the trophic niches of three coexisting Sorex species: S. alpinus, S. araneus and S. minutus were compared for the first time. Shrews were taken from a montane forest habitat in no...

Until recently in Slovenia, the presence of invertebrates in the shrew diet had not been investigated. Here we present a list of invertebrates and other ingested items in five shrew species: Sorex alpinus, S. araneus, S. minutus, Crocidura suaveolens and Neomys fodiens in Slovenia. These shrews fed mostly on arthropods and earthworms. Among the art...

Sand-dwelling antlion larvae are well known predators with a highly specialized prey-catching strategy, which either construct efficient pitfall traps or bury themselves in the sand ambushing prey on the surface. Although habitat selection in antlions has been extensively studied, little is known about the role substrate density plays in such behav...

Terrestrial animals in subterranean habitats are often classified according to their degree of morphological or ecological specialization to the subterranean environment. The commonly held view is that, as distance into a cave increases, the frequency of morphologically specialized, i.e., troglomorphic, species or ecological specialization will inc...

Originally described by Meštrov in 1962, hypotelminorheic habitats are superficial subterranean drainages, typically less than a meter or so in depth, that emerge at small seepage springs. These are persistent wet spots, typically with blackened leaves in small depressions. There may be no flow during dry periods, but the underlying clay retains wa...

Trogulus banaticus Avram, 1971 is characterised and recorded as new for Slovenia. This species was previously mistaken for T. coriziformis C. L. Koch, 1839 and T. graecus Dahl, 1903 which were later rejected from the Slovenian fauna. T. banaticus is compared with the similar, and partly sympatric, T. tingiformis C. L. Koch, 1847 with which it has o...

The cave crickets Troglophilus cavicola and Troglophilus neglectus are the most widely distributed European species of the family Rhaphidophoridae. Their life cycles span two years. They overwinter twice in caves in 4-6 months lasting diapause, T. cavicola in warmer microhabitats. In caves, older T. cavicola undergo sexual maturation, while T. negl...

The impact of grazing has been discussed in many organisms, and it has been shown that it has considerable influence on the structural variation of vegetation, resulting in its tussocky appearance. Such spatial heterogeneity results in plants formations that facilitate other species by providing safe sites against predation and physical stress. On...

Three hymenopteran species – the ichneumonids Amblyteles armatorius and Diphyus quadripunctorius, and the proctotrupid Exallonyx longicornis – are most frequently cited from all over Europe as executing their inactive hypogean ecophase in caves. Yet, little is known about their environmental requirements during their summer quiescence and/or winter...

Hypogean habitats are relatively simple exhibiting low diversity, low production and relative constancy of environmental factors, and are therefore appropriate for studying species coexistence in situ. We investigated the coexistence of two closely related, similarly sized orb-weaving spider species, Meta menardi and Metellina merianae, living synt...

Measurements of thermo- and hygropreference in Leiobunum roseum were carried out in the rocky-scree torrent gorge of Golobnica on the Trnovski gozd Karst Plateau in western Slovenia. This is one of the habitats of the endemic Slovenian Apiaceae plant Hladnikia pastinacifolia, and the southernmost habitat of L. roseum. This species has been confirme...

Periodical changes in the structure of spherites in the Malpighian tubule cells of the cave cricket Troglophilus neglectus were studied to elucidate their role during the cricket's life cycle in natural circumstances. Special interest was given to the dormant overwintering period when we hypothesized that the primary role of spherites is to supply...

Phalangium opilio is the most widespread and one of the most common harvestman species in anthropogenic environments. A preliminary field experiment was carried out in Slovenia testing its preferences for different substrates. A two metres high rectangular tower with walls constituted of vertical bands of concrete, twice fired tile, wood and styrof...

Phalangium opilio is the most widespread and one of the most common harvestman species in anthropogenic environments. A preliminary field experiment was carried out in Slovenia testing its preferences for different substrates. A two metres high rectangular tower with walls constituted of vertical bands of concrete, twice fired tile, wood and styrof...

In this case study we present an application of different analytical electron microscopic methods in biology, to elucidate their usefulness in such investigations. Using analytical electron microscopy, spherites in the digestive gland cells of the helicid snail Chilostoma lefeburiana were examined at three stages: just before the non-feeding period...

The fauna and community ecology of percolation water was studied using monthly samples of 12 drips in the Medvedji rov in the cave Huda luknja. This is the largest cave system in the so-called isolated karst which consists of limestone patches of <1 to 10 km2 in size in central and northeastern Slovenia. Huda luknja is developed in one of these pat...

We sampled 54 caves and galleries in north and east Slovenia methodically for ecological data and the invertebrate fauna. The snow fly Chionea austriaca, recorded in 37% of the cavities, proved indifferent with regard to most of the abiotic factors, but occurred in significantly higher individual numbers in caves on north and east slopes. The subte...

Among the trogulid harvestman Trogulus nepaeformis (Scopoli 1763) collected from sixty localities in Slovenia and northeastern Italy we found many individuals with evidence of old leg injuries. In a few localities, injured individuals accounted for 30% of the total. In non-autotomizing trogulids, in vivo damage to leg articles is characterized by r...

A new species of Cyphophthalmi belonging to the Laurasian family Sironidae is described and illustrated on the basis of external anatomy and genitalia. Siro crassus sp. n., known from leaf litter from two beech forest localities in Eastern Slovenia, is a relatively large (2.2–2.6 mm in length), long-legged epigean species, in which the patella of l...

A new species of Cyphophthalmi belonging to the Laurasian family Sironidae is described and illustrated on the basis of external anatomy and genitalia. Siro crassus sp. n., known from leaf litter from two beech forest localities in Eastern Slovenia, is a relatively large (2.2–2.6 mm in length), long-legged epigean species, in which the patella of l...

The authors present a short historical faunistic, zoogeographical and ecological review of research on harvestmen (Opiliones) of the territory of presentday Slovenia, and discuss some actual ecological and nature conservational questions. Till recent, 64 species plus two subspecies of Opiliones inhabiting Slovenia are recorded, most of these are Al...

A critical review of the literature on harvestmen from Bosnia and Herzegovina was performed, and data from major collections of harvestmen in Bosnia and Herzegovina have been listed to provide an overview of recent knowledge on the topic. Where needed, comments and/or corrections have been added to avoid further confusion. At the moment, 54 valid s...

Critical review of the literature on harvestmen from Croatia was made, and data from major collections of harvestmen in Croatia are listed to provide an overview of recent knowledge on the topic. Where needed comments and/or corrections have been added to avoid further confusion. Altogether, 64 valid species and one subspecies have been firmly iden...

The structure of the midgut gland and its changes in different seasons have been examined in the harvestmen Gyas annulatus and Gyas titanus (Arachnida: Opiliones: Phalangiidae). In both species, in the epithelium of the midgut gland two different types of cells are present: secretory and digestive ones. The secretory cells are characterized by plen...

Gyas annulatus and G. titanus (Arachnida: Opiliones: Phalangiidae) were studied comparatively with special interest in their hypogean ecophase. Direct comparisons were made in two caves where both species overwintered, and in the laboratory. Their chilling hardiness was tested, and the dynamics of lipid and glycogen stocks was studied. Juveniles ov...

In 2000-2001, the fauna of and ecological conditions in the Železna jama cave near Dob in the isolated karst area of Domžale-Moravče (east of the town of Domžale, central Slovenia), were investigated. Fifty-three species were found in the cave, the dominant ones being Meta menardi, Amilenus auriantiacus, Troglophilus neglectus, T. cavicola, Laemost...

Harvestmen (Arachnida: Opiliones) in human settlements of Slovenia were investigated examining opilionid collections deposited in Slovenia, and additional intensive investigations in 29 bigger urban centres. Altogether, data from 58 towns and villages have been treated. So far, 42 of 65 species and subspecies living in Slovenia have been found in t...

On the basis of 33 locality records of Gyas titanus Simon, 1879 (Opiliones, Phalangiidae) it is stated that the species more or less continuously inhabits the western and middle Balkans. Thus the Alpine-Balkans-Carpathian range of the species is continuous.

A faunistic review of the harvestmen (Opiliones) of the Central European zoogeographical region in Slovenia is presented and some remarks on species are added.

In the years 1977-2001, the fauna of and ecological conditions in 55 cavities – caves and artificial tunnels – in northern and central Slovenia were systematically investigated. Zoogeographically, this is the meeting point of the Alpine, Pannonian and Dinaric biome. This article lists 321 terrestrial genera, 456 species and 100 subspecies belonging...

UDC 556.34:59(497.4-18) Tanja Pipan, Vesna Navodnik, Franc Janžekovič & Tone No- vak: Studies of the fauna of percolation water of Huda luknja, a cave in isolated karst in northeast Slovenia The fauna and community ecology of percolation water was studied using montly samples of 12 drips in te Medvedji rov in te cave Huda luknja. This is te largest...

This comparative study provides information on the physiological relevance of spherites in three species of various higher taxa during overwintering: the helicid snail Chilostoma lefeburiana, the harvestman Gyas annulatus and the cave cricket Troglophilus neglectus. They all spend a 4−6 months inactive winter ecophase in caves and other hypogean ha...