Leptodirus
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''Leptodirus'' is a
cave Caves or caverns are natural voids under the Earth's Planetary surface, surface. Caves often form by the weathering of rock and often extend deep underground. Exogene caves are smaller openings that extend a relatively short distance undergrou ...
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
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 (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
contains only the single
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
''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, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
n,
Croatia Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...
n and, partly,
Italian Italian(s) may refer to: * Anything of, from, or related to the people of Italy over the centuries ** Italians, a Romance ethnic group related to or simply a citizen of the Italian Republic or Italian Kingdom ** Italian language, a Romance languag ...
caves.


Biology and ecology

''Leptodirus hochenwartii'' is a true troglobite, adapted to 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 cap ...
s and antennae, the absence of
wing A wing is a type of fin that produces both Lift (force), lift and drag while moving through air. Wings are defined by two shape characteristics, an airfoil section and a planform (aeronautics), planform. Wing efficiency is expressed as lift-to-d ...
s, the absence of
pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...
in the
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, Exoskeleton, shell, germ or Peel (fruit), rind. Etymology The term is derived from ''integumentum'', which is Latin for "a coverin ...
, and
anophthalmia Anophthalmia (Greek: ἀνόφθαλμος, "without eye") is the medical term for the absence of one or both eyes. Both the globe and the ocular tissue are missing from the orbit. The absence of the eye will cause a small bony orbit, a constricte ...
(absence of eyes). However, the most striking features are the slender
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of 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 di ...
, hence the specific name (''leptos''=slender, ''deiros''=neck), and the domed elytrae which cover the
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
completely and give the animal its peculiar round appearance. This adaptation (so-called "false physogastry") allows the animal to store wet air under its 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 neurite structure that, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and respond ...
(the ''Hamann organ'') on its 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 natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...
is largely unknown, but the specimens were seen feeding on
organic material Organic matter, organic material or natural organic matter is 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 come fro ...
, both animal and vegetable origin, which come from outside environment via percolating water or
bat Bats are flying mammals of the order Chiroptera (). 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 birds, flying with their very long spread-out ...
s’ and
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s’
guano Guano (Spanish from ) is the accumulated excrement of seabirds or bats. Guano is a highly effective fertiliser due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. Guano was also, to a le ...
, 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 An egg is an organic vessel in which an embryo begins to develop. Egg, EGG or eggs may also refer to: Biology * Egg cell, the female reproductive cell (gamete) in oogamous organisms Food * Eggs as food Places * Egg, Austria * Egg, Switzerland ...
which take a long time to develop. The number of
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
l
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
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 a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
. 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 oscillation that repeats roughly every 24 hours. Circadian rhythms can refer to any process that originates within an organism (i.e., Endogeny (biology), endogenous) and responds to the env ...
, 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 (; ; ) 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 well as one of its top tourism sites. The caves were created by the P ...
system in southwestern
Slovenia Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
, 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, a
naturalist Natural history is a domain of inquiry involving organisms, including animals, fungi, and plants, in their natural environment, leaning more towards observational than experimental methods of study. A person who studies natural history is cal ...
and
entomologist Entomology (from Ancient Greek ἔντομον (''éntomon''), meaning "insect", and -logy from λόγος (''lógos''), meaning "study") is the branch of zoology that focuses on insects. Those who study entomology are known as entomologists. In ...
from
Ljubljana {{Infobox settlement , name = Ljubljana , official_name = , settlement_type = Capital city , image_skyline = {{multiple image , border = infobox , perrow = 1/2/2/1 , total_widt ...
. 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 ( ; ; ; ) 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 still tend to identify with its traditional parts Upp ...
'') 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, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
''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 which is the only species in the genus ''Proteus'' of the family Proteidae and the only exclusively cave-dwelling chordate species found in Europe; the family's other extant g ...
(''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 country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
n
entomologist Entomology (from Ancient Greek ἔντομον (''éntomon''), meaning "insect", and -logy from λόγος (''lógos''), meaning "study") is the branch of zoology that focuses on insects. Those who study entomology are known as entomologists. In ...
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 Biospeleology, also known as cave biology, is a branch of biology dedicated to the study of organisms that live in caves and are collectively referred to as troglofauna. Biospeleology as a science History The first documented mention of a ...
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 (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Leptodirus''. It is
endemic Endemism is the state of a species being found only 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 foun ...
to western
Dinaric Alps The Dinaric Alps (), also Dinarides, are a mountain range in Southern Europe, Southern and Southcentral Europe, separating the continental Balkan Peninsula from the Adriatic Sea. They stretch from Italy in the northwest through Slovenia, Croatia ...
, from
Inner Carniola Inner Carniola ( ; ) is a traditional region of Slovenia, the southwestern part of the larger Carniola region. It comprises the Hrušica (plateau), Hrušica karst plateau up to Postojna Gate, bordering the Slovenian Littoral (the Goriška, Gor ...
(
Slovenia Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
) north to
Velebit Velebit (; ; ) 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 northwest near ...
(
Croatia Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...
) south. Six
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
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 Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...
. The subspecies ''L. h. reticulatus'' are found in
Slovenia Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
,
Croatia Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...
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) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
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 an inventory of the global conservation status and extinction risk of biological sp ...
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 (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