Figure 6 - uploaded by Slavko Polak
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Iconography annexed to the original description of the Stagobius troglodytes published in " Bidrag til den underjordiske Fauna " , 1849.
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... Since the initial description of the first cave-dwelling organism in 1768, interest in this scientific field has surged significantly in Europe and the USA, and more recently, in tropical regions (Mammola, 2019;Polak, 2005). Substantial efforts have been directed toward enhancing our understanding of the biogeographical, ecological, and evolutionary processes that govern subterranean ecosystems (Lienhard & Ferreira, 2013;Mamolla et al., 2019Mamolla et al., , 2020Souza-Silva et al., 2021;Recknagel & Trontelj, 2022). ...
In caves, the absence of natural light in deeper regions starkly contrasts with the entrance areas, which still exhibit a light gradient extending into the cave interior. This interplay with the structural gradient of the environment potentially exerts distinct influences on invertebrate communities residing in different cave light zones. To investigate this, we formulated a hypothesis positing that communities within distinct cave light zones respond differently to habitat structure and microclimatic conditions. Our approach involved a spatial multi‐scale sampling of invertebrates and the application of statistical analyses to contrast the responses of communities inhabiting photic and aphotic zones. Photic zone richness is influenced by factors such as air moisture, resource availability, root presence, and shelter diversity. In contrast, the richness of communities in aphotic zones is shaped by resource availability, the presence of roots, branches, and distance from the cave entrance. As expected, the richness in the photic zone surpasses that of the deeper regions, highlighting the challenges faced by invertebrates attempting to establish themselves in the aphotic zone. The species composition of faunal communities varied predictably from the entrance to the aphotic zone, and the three most important factors driving this variation were geographic distance, humidity, and distance from the entrance. The composition between these cave zones differs significantly, primarily due to the high number of obligate cave species predominantly inhabiting the aphotic region. Indeed, communities associated with different cave zones exhibit distinct responses to resource, microclimatic, and structural variables.
Abstract in Portuguese is available with online material.
... When Austro-Hungarian entomologist Ferdinand Schmidt in 1832 described the first beetle species adapted to caves in Postojna Cave, Slovenia (Schmidt 1832;Polak 2005), he also revealed the existence of subterranean insect life stable and suitable. Until then unsuspected, this ability to adapt to underground life is now rich in many examples and one now recognizes a real "hidden" underground diversity where all the major phyla are now represented (Sendi et al. 2020). ...
Planthoppers are an interesting and contrasting model among insects for studying the subterranean environments. Their morphological and ethological adaptations to the underground conditions (complete darkness, lower temperatures, high hygrometry, stability of environmental constants, rarefied food sources, etc.), and their worldwide distribution in both temperate and tropical areas make them an interesting model among invertebrates. In this review, we highlight why cave planthoppers study matters, with particular emphasis on the Cixiidae. The two hypotheses proposed, the ‘climatic relict hypothesis’ and the ‘adaptive shift’, are not sufficient enough to clearly understand and explain the drivers to cavernicoly. Phylogenetic analyses approaches might help to better document and increase our knowledge on such peculiar environments. The singularity of the distribution pattern of the adaptation to cavernicoly in planthoppers raises also interesting questions to investigate and suggest contrasting scenarios to explore further, particularly should the Cixiidae be defined as a subtroglophile lineage?
... This largely changed with the annexation of modern Bosnia and Herzegovina territories, including Popovo Polje and Vjetrenica, by the Austro-Hungarian Empire (1878) [19][20][21][22]. Only a few decades before that, the first subterranean animal, Leptodirus hochenwartii Schmidt, 1832, from the Postojna Cave was described, and speleobiology-the biology of subterranean habitats-was born [23]. Southward extension of the empire suddenly enabled naturalists and admirers of subterranean caves to sample specialized fauna in Vjetrenica and other caves in Popovo Polje [24]. ...
... For a cave whose entrance is characterized by winds reaching up to 89 km/h (Roman Ozimec, personal data), there is no wonder it bears a name meaning "a windy place" in local languages [9,10,12,23]. Similar to the polje, the Vjetrenica Cave System developed in the Mesozoic limestones, predominantly during the Cretaceous age, stretching in the NW-SE direction, and is situated in the outskirts of the Zavala Village in Popovo Polje, Bosnia and Herzegovina (42.8458, 17.9838) [55]. ...
The Western Balkan’s Vjetrenica Cave in southern Bosnia and Herzegovina is renowned for high richness of subterranean species. However, the data on its fauna have been published only in monographs printed in a small number of copies, making them hardly accessible to the wider scientific community. To overcome this issue, we compiled the data from published monographs with the data from our own recent field surveys. Further, as they are connected via water channels or small crevices in bedrock, we defined the Vjetrenica Cave System as a system comprising Vjetrenica and Bjelušica Caves and Lukavac Spring. Altogether, 93 troglobiotic, i.e., obligate subterranean aquatic (48) and terrestrial (45), taxa were reported for the system, verifying the Vjetrenica Cave System as the second richest locality in subterranean biodiversity in the world. The global uniqueness of the system is also reflected in the fact that as many as 40 troglobiotic species were described from the system. Finally, we reviewed the factors endangering this unique subterranean community and questioned whether it will withstand human-induced changes and pressures due to infrastructural development in southern Bosnia and Herzegovina.
... The difficulty of precisely investigating subterranean biotremology has restricted the capability and opportunity to conduct such studies. The first subterranean insect was not even reported until 1831 (Polak 2005). The arthropod biodiversity of underground environments remains largely unexplored (Ficetola et al. 2018;Rillig et al. 2019). ...
Subterranean arthropods would be ideal candidates for biotremological studies except that soil is a heterogeneous mixture of porous and solid materials with poorly characterized mechanical properties, which makes such studies difficult in situ. However, increased awareness of the impacts of subterranean arthropod herbivory on above-ground biota and the rapid development of modern electronic sensors and computer systems have encouraged efforts to conduct subterranean arthropod biotremological studies in recent years. In addition, impetus to address economic damage caused by subterranean arthropod pests has prompted multiple studies on the intra- and interspecific vibrational communication and incidental movement activities of subterranean pests. Many of these studies have been conducted on subterranean Cicadoidean, Ensiferan, Scarabaeoid, and Curculionid species that are a focus of this chapter. Studies on Lepidoptera, social insects, and Araneae also are discussed here in economic and ecological context, particularly with respect to multimodal communication.
... News of the discovery spread and in the next years, many eminent naturalists came to study the fauna of Postojnska jama, describing cave species of other animal taxa. The snowball effect triggered by the description of L. hochenwartii is considered the starter of speleobiology and PPCS the "cradle of speleobiology" [10]. The pace of new discoveries by numerous scientists in caves of the PPCS (see detailed review of the beginnings in [10]) produced the first checklists of the PPCS already in the 19 th century. ...
... The snowball effect triggered by the description of L. hochenwartii is considered the starter of speleobiology and PPCS the "cradle of speleobiology" [10]. The pace of new discoveries by numerous scientists in caves of the PPCS (see detailed review of the beginnings in [10]) produced the first checklists of the PPCS already in the 19 th century. An important milestone in cataloguing was made by Benno Wolf, who collected all data on animals in caves of the world, and compiled them in Animalium Cavernarum Catalogus, issued between 1934 and 1938 [11]. ...
The Postojna-Planina Cave System (PPCS) in central Slovenia is a globally exceptional site of subterranean biodiversity, comprised of many interconnected caves with cumulative passage length exceeding 34 km. Two rivers sink into the caves of the PPCS, called the Pivka and Rak, and join underground into Unica River, which emerges to the surface. The studies of fauna of PPCS began in the 19th century with the first scientific descriptions of specialized cave animals in the world, making it “the cradle of speleobiology”. Currently, the species list of PPCS contains 117 troglobiotic animal species belonging to eight phyla, confirming its status as the richest in the world. Of these, 47 species have been scientifically described from the PPCS, and more than 10 await formal taxonomic descriptions. We expect that further sampling, detailed analyses of less studied taxa, and the use of molecular methods may reveal more species. To keep the cave animals’ checklist in PPCS up-to-date, we have supplemented the printed checklist with an online interface. As the revised checklist is a necessary first step for further activities, we discuss the importance of PPCS in terms of future research and conservation.
... During the past two centuries, following the establishment of speleobiology -the biology of subterranean habitats -in 1832 (Polak 2005;Sket 2012), an immense effort has been invested in the discovery of specialized subterranean species. By the early 2000s, almost 1000 species adapted for living in these habitats, which seem simple and resource deprived, have been recognized in the western Dinaric Karst alone (Sket et al. 2004). ...
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 very long, thin appendages, with legs II about 15 times as long
as the body. Although very similar, Nemaspela ladae Karaman, 2013 and N. borkoae sp. nov. can be
easily distinguished by the terminally rounded vs conical glans, straight vs conspicuously ventrally bent
pedipalp tarsus on its proximal portion and pedipalp tarsus measuring about ½ vs ⅔ tibia length. Nemaspela
ladae and N. borkoae sp. nov. constitute the western Nemaspela group, both missing the male cheliceral
apophysis present in all species of the eastern Nemaspela group from the Crimea and Caucasus, except in
N. femorecurvata Martens, 2006. However, according to the glans morphology, N. borkoae sp. nov. seems
much more closely related to several species from the Caucasus than to N. ladae from the Balkan Peninsula.
We speculate that N. ladae and N. borkoae sp. nov. might originate from two epigean ancestral lineages.
... But this journey could not be completed before the cognition that karst underground harbours unique life, which emerged through the second part of the 19 th century. The birth of speleobiology was triggered by Ljubljana entomologist Ferdinand J. Schmidt (1791-1878 with his description of the first cave beetle Leptodirus hochenwartii Schmidt 1832 (Schmidt, 1832;Polak, 2005), followed soon by discoveries of other cave-adapted invertebrates in Postojnska jama (Aljančič et al., 1993;Sket, 1993A;Sket, 2012). The hotspot of karst research finally revealed the global hotspot of the subterranean biodiversity recognised today (Sket, 1993A;Sket, 1999;Culver and Sket, 2000;Culver & Pipan, 2009;Sket, 2012). ...
Olm or Proteus (Proteus anguinus Laurenti 1768) was the first taxonomically described cave animal in the world, by J. N. Laurenti, 1768, upon a specimen that was apparently found on the famous lake Cerkniško jezero, Slovenia, yet the existence of this unusual animal in Slovenia had been known long before.
The research on Proteus is one of the oldest Slovenian natural history projects, a 330 year spiritual bond: from the first description by one of pioneers of karst research J. V. Valvasor in 1689, to the renowned naturalists J. A. Scopoli, who was the first researcher to actually examine proteus from the Stična area in 1762. One of the central figures of the early proteus research was Ž. Zois, the first who studied proteus behaviour, and conducted earliest physiological and ecological observations, together with Viennese zoologist Karl von Schreibers. Zois’s work was continued by two researchers of proteus distribution F. J. Hochenwart and H. Freyer, and other researchers of the 19th and 20th century.
For the last 250 years, this mysterious animal has constantly raised scientific and public attention, and gradually became not only an important symbol of Slovenia’s nature, but also a part of its cultural heritage. The zoologically extraordinary Proteus was also an important object in the history of international nature research, puzzling the minds of most prominent naturalists, from Linnaeus, Cuvier and Humboldt, to Lamarck and Darwin.
This article also presents reproductions of the earliest illustrations of proteus: a collection of ten published and unpublished work between 1752 and 1849, a forgotten heritage of the first 100 years of proteus research. In Memoriam Žiga Zois (1747–1819).
... The Postojna-Planina Cave system (PPCS), with 99 species, is the most biologically diverse cave system in the world (Sket 2010). The PPCS is the type locality for many cave-dwelling species (Pretner 1968;Polak 2005); for example, the first cave animal discovered and recognized the beetle Leptodirus hochenwartii. It also includes the first described cave representatives of the cave land snail Zospeum spelaeum, the cave springtail Onychiurus stillicidii, the cave amphipod Niphargus stygius, the cave isopod Titanethes albus, the cave pseudoscorpion Neobisium spelaeum, the cave spider Stalita taenaria, the cave millipede Brachydesmus subterraneus, the only true cave hydrozoan Velkovrhia enigmatica, and others. ...
Slovenia is a biodiversity hotspot due to its location at the junction of four biogeographical regions with different ecological conditions. Because of this, 37.87% of
Slovenia’s area is currently classified as Natura 2000 sites, the highest share among the 28 EU countries. Slovenian flora comprises 3472 vascular taxa and is rich in endemic
species, but unfortunately also rare and threatened species. Especially distinctive is the Illyrian floral element—that is, plants with limited distributions along the Dinaric Alps
from Slovenia to Albania, mainly thermophilic and heliophilic endemic species. The vegetation of Slovenia differs from that of neighboring regions because of its rich flora
as well as its different vegetation history and development after the last ice age. Slovenia is known for its extensive forest cover (58.9%), mostly dominated by beech.
Slovenian fauna comprises more than 21,500 continental taxa and at least 1600 marine taxa. Slovenia is known for its Karst region, with a rich subterranean fauna, of which many species were first discovered and described in Slovenia. Three out of five species of large carnivores found in Europe are present in Slovenia: the brown bear, gray wolf, and Eurasian lynx. The extensive forest cover also makes possible a great diversity of mesofaunal carnivores and other mammals.
... The first cave-dwelling invertebrate known to science was the beetle Leptodirus hochenwartii, described by Ferdinand Schmidt in 1832 ( Fig. 6.1). The only cave dweller to be described earlier than this was the European cave salamander, Proteus anguinus Laurenti -and it was originally thought not to be a cave inhabitant but rather a denizen of the periodically dry Slovenian karst lake, Cerkniško jezero (Polak 2005). The story of the collection of L. hochenwartii, its taxonomic description, and the interpretation of its unique morphology is a convenient way to frame the general discussion about insects in caves. ...
... Luka Čeč, a Slovenian cave guide and discoverer of most of what is now the tourist route in the world-famous show cave Postojnska jama, found a specimen of what came to be described as L. hochenwartii in the Calvary section of the cave. He gave the specimen to Franz Josef von Hochenwart, who was preparing a written guide to Postojnska jama (Polak 2005). He in turn gave the specimen to the entomologist Ferdinand Schmidt, who erected a new genus for this unusual species. ...
... The Danish zoologist Jorgen Christian Schiödte visited Schmidt and Postojnska jama in 1845, and described a number of new species, including the beetle Stagobius troglodytes. Stagobius troglodytes is a synonym for L. hochenwartii, but it was not until 1849 that Schiödte finally took the lesser-known Schmidt's description seriously (Polak 2005). ...
The first cave‐dwelling invertebrate known to science was the beetle Leptodirus hochenwartii, described by Ferdinand Schmidt in 1832. This chapter presents a discussion of L. hochenwartii. It describes the variety of subterranean spaces, including but not limited to caves, that insects inhabit, the ecological roles of insects in subterranean habitats, and morphological and life‐history adaptations of insects to subterranean life. The L. hochenwartii has many of the features associated with adaptation to subterranean life, including reduced pigment, loss of eyes, and appendage elongation. The chapter shows a number of specialized cave insects, soil insects, and generalists at a milieu souterrain superficiel (MSS) site in Teno, Tenerife, Canary Islands. It also describes probable modes of successful colonization of subterranean spaces, taxonomic and geographic patterns of subterranean insect biodiversity and human utility and protection of cave insects.
... Hochenwart gave this unusual beetle to the famous Carnolian entomologist Ferdinand J. Schmidt (1791Schmidt ( -1878, an expert on beetles, who immediately recognized the beetle as a true subterranean animal with a special adaptation to the subterranean environment. He described the beetle as a new species and new genus, Leptodirus hochenwartii (Leiodidae) (Fig. 222 -223), the species was described in "Beitrag zu Krain' s Fauna" which appeared in the Carnolian paper Illyrisches Blatt on 21 st January 1832 (POLAK, 2005). Thus the first subterranean beetle was discovered, described and the amazing adventure of the study of subterranean beetles and insects and their life history and indeed the science of biospeleology had started. ...
... Thus the first subterranean beetle was discovered, described and the amazing adventure of the study of subterranean beetles and insects and their life history and indeed the science of biospeleology had started. The discovery of the first cave beetle raised great interest among the European naturalists of that time and continues to the present time where it is regarded as one of the most interesting and attractive branches of entomology (POLAK, 2005). ...
... Among the first zoologists that visited Carniola and its caves was Jorgen Christian Schiødte (1815Schiødte ( -1884 Schaum (1819Schaum ( -1865, Jacob Sturm (1771Sturm ( -1848 and Gustav Joseph from Germany, Rodolphe Khevenhüller -Metch and Rudolf Ignaz Schiner (1813Schiner ( -1873 from the Kingdom of Austria (POLAK, 2005). As time passed, other species and genera were described: in 1844 Anophthalmus schmidtii (Carabidae) by Jacob in 1846Heinrich Karl Küster (1807-1876 described Laemostenus schreibersi (Carabidae); in 1852 Ludwig Miller (1820Miller ( -1897 described Bathysciotes khevenhuelleri (Leiodidae) and in 1855 the first troglobitic Pselaphine Machaerites spelaeus. ...
