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''Leptodirus'' is a
cave A cave or cavern is a natural void in the ground, specifically a space large enough for a human to enter. Caves often form by the weathering of rock and often extend deep underground. The word ''cave'' can refer to smaller openings such as sea ...
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
in the family
Leiodidae Leiodidae is a family of beetles with around 3800 described species found worldwide. Members of this family are commonly called round fungus beetles due to the globular shape of many species, although some are more elongated in shape. They are g ...
. The
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
contains only the single
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
''Leptodirus hochenwartii''. It is a true
troglobite A troglobite (or, formally, troglobiont) is an animal species, or population of a species, strictly bound to underground habitats, such as caves. These are separate from species that mainly live in above-ground habitats but are also able to live u ...
, endemic to
Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, an ...
n,
Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...
n and, partly,
Italian Italian(s) may refer to: * Anything of, from, or related to the people of Italy over the centuries ** Italians, an ethnic group or simply a citizen of the Italian Republic or Italian Kingdom ** Italian language, a Romance language *** Regional Ita ...
caves.


Biology and ecology

''Leptodirus hochenwartii'' is a true troglobite, adapted to the subterranean life and unable to survive in the outside environment. As a result, it possesses typical troglobiotic features, such as elongated
leg A leg is a weight-bearing and locomotive anatomical structure, usually having a columnar shape. During locomotion, legs function as "extensible struts". The combination of movements at all joints can be modeled as a single, linear element ca ...
s and antennae, the absence of
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expres ...
s, the absence of
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
in the
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
, as well as the
anophthalmia Anophthalmia, (Greek: ἀνόφθαλμος, "without eye"), is the medical term for the absence of one or both eyes. Both the globe (human eye) and the ocular tissue are missing from the orbit. The absence of the eye will cause a small bony orbi ...
(absence of eyes). However, the most striking features are the slender
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
, hence the specific name (''leptos''=slender, ''deiros''=neck), and the domed elytrae which cover the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
completely and give the animal its peculiar round appearance. This adaptation (so-called "false physogastry") allows the animal to store wet air under elytrae and use it for breathing in drier areas. Another typical feature is a specific
receptor Receptor may refer to: * Sensory receptor, in physiology, any structure which, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and responds to a ...
(the ''Hamann organ'') on antennae which helps the animal to perceive air humidity level. It lives predominantly in large and cold caves where the temperature does not exceed . Its
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
is largely unknown, but the specimens were seen feeding on
organic material Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
, both animal and vegetable origin, which come from outside environment via percolating water or
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s’ and
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s’
guano Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
, and carcasses of different cave animals. Even less is known about its life history. The only study done on ''L. hochenwartii'' so far showed that, as is the case in most specialised cave Leptodirini, females lay a small number of relatively large
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
which take a long time to develop. The number of
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
l
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
s is reduced to only one, and the larvae do not feed before
moulting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
. The maximum period of activity of adults is still unknown. As usual in troglobites, environmental stability and the absence of sunlight have brought about the loss of
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., Endogeny (biology), endogeno ...
, whereas seasonal rhythm is affected by rainfall patterns.


Research history

The animal was first found in 1831 by Luka Čeč, an assistant to the lamplighter in the
Postojna Cave Postojna Cave ( sl, Postojnska jama; german: Adelsberger Grotte; it, Grotte di Postumia) is a long karst cave system near Postojna, southwestern Slovenia. It is the second-longest cave system in the country (following the Migovec System) as wel ...
system in southwestern
Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, an ...
, when exploring new inner cave portions discovered some year before. He gave the specimen to count Earl Franz von Ho(c)henwart who was unable to determine the species, and gave it in turn to
Ferdinand Jožef Schmidt Ferdinand Joseph Schmidt (also as Ferdinand Jožef Schmidt or Ferdinand Jožef Šmit) (20 February 1791, Sopron – 16 February 1878, Ljubljana) was an Austro-Hungarian businessman, naturalist and explorer who was among the pioneers of biospeleolo ...
, a naturalist and
entomologist Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
from
Ljubljana Ljubljana (also known by other historical names) is the capital and largest city of Slovenia. It is the country's cultural, educational, economic, political and administrative center. During antiquity, a Roman city called Emona stood in the ar ...
. Schmidt recognized the beetle as a new species and described it in the article under the name "Beitrag zu Krain's Fauna" (''Contribution to the fauna of
Carniola Carniola ( sl, Kranjska; , german: Krain; it, Carniola; hu, Krajna) is a historical region that comprised parts of present-day Slovenia. Although as a whole it does not exist anymore, Slovenes living within the former borders of the region sti ...
'') which appeared in the Carniolan paper ''Illyrisches Blatt'' in 1832. He named it ''Leptodirus'' (which means "slender-necked (beetle)") ''hochenwartii'' after the donor, and also gave it the common Slovene name ''drobnovratnik'' and
German German(s) may refer to: * Germany (of or related to) **Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ger ...
''Enghalskäfer'', both related to its typical slender thorax. The article represents the first formal description of a cave animal, since the
olm The olm or proteus (''Proteus anguinus'') is an aquatic salamander in the family Proteidae, the only exclusively cave-dwelling chordate species found in Europe. In contrast to most amphibians, it is entirely aquatic, eating, sleeping, and br ...
(''Proteus anguinus''), described in 1768 by
Josephus Nicolaus Laurenti Josephus Nicolaus Laurenti (4 December 1735, Vienna – 17 February 1805, Vienna) was an Austrian naturalist and zoologist of Italian origin. Laurenti is considered the auctor of the class Reptilia (reptiles) through his authorship of ' (1768) ...
, was not recognized as a cave animal at the time. In 1856 the
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
n
entomologist Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
Viktor Motchoulski described a new ''Leptodirus'' species, named ''L. schmidti'', currently recognized as a subspecies of ''L. hochenwartii''. Subsequent research by Schmidt and other naturalists revealed further previously unknown cave inhabitants, which aroused considerable interest among cave researchers. For this reason, the discovery of ''L. hochenwartii'' (along with the olm) is considered the starting point of biospeleology as a
scientific discipline The branches of science, also referred to as sciences, scientific fields or scientific disciplines, are commonly divided into three major groups: *Formal sciences: the study of formal systems, such as those under the branches of logic and mat ...
.


Taxonomy and range

''Leptodirus hochenwartii'' is the only species in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''Leptodirus''. It is
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to western
Dinaric Alps The Dinaric Alps (), also Dinarides, are a mountain range in Southern and Southcentral Europe, separating the continental Balkan Peninsula from the Adriatic Sea. They stretch from Italy in the northwest through Slovenia, Croatia, Bosnia and Herz ...
, from
Inner Carniola Inner Carniola ( sl, Notranjska; german: Innerkrain) is a traditional region of Slovenia, the southwestern part of the larger Carniola region. It comprises the Hrušica karst plateau up to Postojna Gate, bordering the Slovenian Littoral (the ...
(
Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, an ...
) north to
Velebit Velebit (; it, Alpi Bebie) is the largest, though not the highest, mountain range in Croatia. The range forms a part of the Dinaric Alps and is located along the Adriatic coast, separating it from Lika in the interior. Velebit begins in the nor ...
(
Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...
) south. Six
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
are currently recognized in this
range Range may refer to: Geography * Range (geographic), a chain of hills or mountains; a somewhat linear, complex mountainous or hilly area (cordillera, sierra) ** Mountain range, a group of mountains bordered by lowlands * Range, a term used to i ...
: * ''Leptodirus hochenwartii hochenwartii'' Schmidt, 1832 * ''Leptodirus hochenwartii schmidti'' (Motschoulsky, 1856) * ''Leptodirus hochenwartii reticulatus'' J. Müller, 1906 * ''Leptodirus hochenwartii pretneri'' Müller, 1926 * ''Leptodirus hochenwartii croaticus'' Pretner, 1955 * ''Leptodirus hochenwartii velebiticus'' Pretner, 1970 Of those, two subspecies (''hochenwartii'' and ''schmidti'') are only found in Slovenia, and three (''pretneri'', ''croaticus'' and ''velebiticus'') are only found in
Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...
. The subspecies ''L. h. reticulatus'' are found in
Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, an ...
,
Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...
and in the Trieste Karst (in Italy), where was originally found in Grotta Noè.


Conservation

Although
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
has not evaluated its
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
, due to its limited range and slow reproduction, ''L. hochenwartii'' is considered rare and vulnerable, despite the fact that individual density in some caves can be high. The main threats are illegal and massive collecting and pollution of the caves. As a consequence, the species is included in the Slovenian
Red list The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biologi ...
of threatened species (category R).''Slovenian official gazette'' (2002)
no. 82, Tuesday 24 September 2002
Additionally, it is included in the Annex II of the
EU Habitats Directive The Habitats Directive (more formally known as Council Directive 92/43/EEC on the Conservation of natural habitats and of wild fauna and flora) is a directive adopted by the European Community in 1992 as a response to the Berne Convention. The E ...
(92/43/EEC).''EU Habitats directive''
(1992). European Commission
On this basis, 15 areas of conservation (pSCI) are established in Slovenia which include the majority of known localities.


Notes


References

{{Taxonbar, from1=Q12033027, from2=Q2300017 Leiodidae Beetles of Europe Beetles described in 1832 Cave beetles Postojna Cave Monotypic beetle genera