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''Lucanus cervus'', known as the European stag beetle, or the greater stag beetle, is one of the best-known species of
stag beetle Stag beetles comprise the family Lucanidae. It has about 1,200 species of beetles in four subfamilies.Smith, A.B.T. (2006). A review of the family-group names for the superfamily Scarabaeoidea (Coleoptera) with corrections to nomenclature and a c ...
(family Lucanidae) in
Western Europe Western Europe is the western region of Europe. The region's extent varies depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the Western half of the ancient Mediterranean ...
, and is the eponymous example of the genus. ''L. cervus'' is listed as Near Threatened by the
IUCN 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 ...
.


Taxonomy

''Lucanus cervus'' is situated in the genus ''Lucanus'' within the family Lucanidae. In the genus there are two subgenera: ''Lucanus'' Scopoli, 1763 and ''Pseudolucanus'' Hope and Westwood, 1845. The species ''L. cervus'' contains four subspecies. The
nominate subspecies In 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), but that can successfully interbreed. ...
''L. cervus cervus'' (Linnaeus, 1758) was established via the original description of the species in 1758. The three latterly added subspecies are ''L. cervus judaicus'' Planet, 1900, ''L. cervus laticornis'' Deyrolle, 1864, and ''L. cervus turcicus'' Sturm, 1843.


Description

The European stag beetle is the largest beetle in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. Their colour is usually black with reddish
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
(and red mandibles in males).
Sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, the males have enlarged mandibles and are larger than the females. Although the male's mandibles seem threatening, they are too weak to be harmful. Nevertheless, females can inflict a painful bite. The resemblance of the male's mandibles to the antlers of a stag, and their use in combat between males, much like with deer, gives the species its scientific and common names. Adult size varies between different areas of its distribution, for example beetles from Spain, Germany, and the Netherlands are larger than those from Belgium or the UK. Males grow up to in length, and females grow between . In Britain, female ''Lucanus cervus'' are often confused with both sexes of the lesser stag beetle (''Dorcus parallelipipedus''). They can be distinguished as lesser stag beetles are smaller, with a completely black exoskeleton, and a larger, squarer head. Another difference is that ''Lucanus cervus'' larvae feed on wood below ground, and ''Dorcus parallelipipedus'' larvae eat wood above ground. File:Lucanus cervus female top view.jpg, ''Lucanus cervus'' female File:Dorcus parallelipipedus m.jpg, ''Dorcus parallelipipedus'' male File:Dorcus parallelipipedus f.jpg, ''Dorcus parallelipipedus'' female


Distribution and habitat

''Lucanus cervus'' is widespread across Europe, though it is absent from Ireland. In Germany it is widespread, mainly in the south. In
Hungary Hungary is a landlocked country in Central Europe. Spanning much of the Pannonian Basin, Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia and ...
this species is widespread in the hilly and mountainous areas. In
Romania Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
it is widespread, especially in the hilly areas with sun exposed slopes. It also occurs in the European part of Turkey. In Italy it is mainly distributed in northern and central regions. In
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
and
Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in continental Europe, Portugal borders Spain to its north and east, with which it share ...
it is present only in the northern half of each country. In Britain it is largely confined to the south-east of England, where it is widespread. This species is now extinct in
Latvia Latvia, officially the Republic of Latvia, is a country in the Baltic region of Northern Europe. It is one of the three Baltic states, along with Estonia to the north and Lithuania to the south. It borders Russia to the east and Belarus to t ...
; it also disappeared from
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
around 1970, but was reintroduced in 2013. It is also found in
Caucasus The Caucasus () or Caucasia (), is a region spanning Eastern Europe and Western Asia. It is situated between the Black Sea and the Caspian Sea, comprising parts of Southern Russia, Georgia, Armenia, and Azerbaijan. The Caucasus Mountains, i ...
,
Asia Minor Anatolia (), also known as Asia Minor, is a peninsula in West Asia that makes up the majority of the land area of Turkey. It is the westernmost protrusion of Asia and is geographically bounded by the Mediterranean Sea to the south, the Aegean ...
,
Syria Syria, officially the Syrian Arab Republic, is a country in West Asia located in the Eastern Mediterranean and the Levant. It borders the Mediterranean Sea to the west, Turkey to Syria–Turkey border, the north, Iraq to Iraq–Syria border, t ...
and west
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
. Its range is currently hen?increasing only 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 ...
and
Slovakia Slovakia, officially the Slovak Republic, is a landlocked country in Central Europe. It is bordered by Poland to the north, Ukraine to the east, Hungary to the south, Austria to the west, and the Czech Republic to the northwest. Slovakia's m ...
. ''Lucanus cervus'' has been associated with a range of trees including those in the
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
oak An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
(''Quercus''), lime (''
Tilia ''Tilia'' is a genus of about 30 species of trees or bushes, native throughout most of the temperateness, temperate Northern Hemisphere. The tree is known as linden for the European species, and basswood for North American species. In Great Bri ...
''),
beech Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of Mesophyte, mesophytic forests) Eurasia and North America. There are 14 accepted ...
(''Fagus''),
willow Willows, also called sallows and osiers, of the genus ''Salix'', comprise around 350 species (plus numerous hybrids) of typically deciduous trees and shrubs, found primarily on moist soils in cold and temperate regions. Most species are known ...
(''Salix'') and certain species in other genera including black poplar (''
Populus nigra ''Populus nigra'', the black poplar, is a species of Populus sect. Aigeiros, cottonwood poplar, the type species of section ''Aigeiros'' of the genus ''Populus'', native to Europe, southwest and central Asia, and northwest Africa.Flora Europaea' ...
''), ash (''
Fraxinus excelsior ''Fraxinus excelsior'', known as the ash, or European ash or common ash to distinguish it from other types of ash, is a flowering plant species in the olive family Oleaceae. It is native throughout mainland Europe east to the Caucasus and Alb ...
''), horse-chestnut (''
Aesculus hippocastanum ''Aesculus hippocastanum'', the horse chestnut, is a species of flowering plant in the maple, Sapindus, soapberry and lychee family Sapindaceae. It is a large, deciduous, synoecious (hermaphroditic-flowered) tree. It is also called horse-chestnu ...
''), wild cherry (''
Prunus avium ''Prunus avium'', commonly called wild cherry, sweet cherry or gean is a species of Prunus subg. Cerasus, cherry, a flowering plant in the rose family, Rosaceae. It is native to Eurasia and naturalized elsewhere. It is an ancestor of ''Prunus c ...
''), and common walnut (''
Juglans regia ''Juglans regia'', known by various common names including the common walnut, English walnut, or Persian walnut amongst other names, is a species of walnut. It is native to Eurasia in at least southwest and central Asia and southeast Europe, ...
''). Larval development sites were found in proximity of dead wood of the allochthonous ''
Quercus rubra ''Quercus rubra'', the northern red oak, is an oak tree in the red oak group (''Quercus'' section ''Lobatae''). It is a native of North America, in the eastern and central United States and southeast and south-central Canada. It has been intro ...
''.


Life cycle


Life cycle

Adults appear during late May to the beginning of August, being most active in the evenings. Females lay about 30 eggs in a piece of decaying wood deep in the soil. Stag beetle larvae, which are blind and shaped like a letter "C", feed on rotting wood in a variety of places, tree stumps, old trees and shrubs, rotting fence posts, compost heaps, and leaf mould. The larvae have a cream-coloured, soft, transparent body with six orange legs, and an orange head which is very distinct from the very sharp brown pincers. They have combs in their legs which they use for communication (
stridulation Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...
) with other larvae. The larvae go through several
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 (stages), taking several years to become
pupae A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
. Males use their large mandibles to fight with each other in competition for females, intending to throw their opponent off the log or branch they are on. The work of entomologist Charlie Morgan during the late 1970s discovered that the pupae of the stag beetle live in the soil for about 3 months, then emerge in summer to awkwardly fly off to mate. Adults suck up the liquids of
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
, fallen fruit, and tree
sap Sap is a fluid transported in the xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a s ...
. Their slow, lumbering flight, usually at dusk, makes a distinctive low-pitched buzzing sound. The males fly more readily than the females.


Lifespan

Stag beetles spend most of their life as
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
and can take from 3 to 7 years for them to pupate, but the lifespan of the adults is only a few weeks.


Subspecies

The species ''Lucanus cervus'' contains four subspecies, with ''Lucanus cervus cervus'' being the
nominate subspecies In 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), but that can successfully interbreed. ...
. The three latterly added subspecies are ''L. cervus judaicus'', ''L. cervus laticornis'', ''L. cervus turcicus'', and ''L. cervus akbesianus''. File:Lucanus cervus cervus3.JPG, ''Lucanus cervus cervus'' File:Lucanus cervus judaicus 3.JPG, ''Lucanus cervus judaicus'' File:Lucanus cervus turcicus (Bulgaria) TP11.jpg, ''Lucanus cervus turcicus'' File:Lucanus cervus akbesianus1.JPG, ''Lucanus cervus akbesianus'' File:Lucanus cervus laticornis.JPG, ''Lucanus cervus laticornis'' (middle)


Behaviour and ecology

The natural reaction of the beetle to an approaching large object is to remain motionless, making them a good photographic subject.


Daily activity, use of space and detectability

In a primary ancient forest in northern Italy, stag beetle males were less elusive than females. Males were more frequently radio-tracked in flight, females mostly underground and in the proximity of deadwood. Males were mostly observed flying at sunset, and resting or walking on standing trees during the day. The combination of air temperature and humidity determined the optimal weather conditions for male flights. Flying at sunset, mostly performed by males, significantly increased the detectability of the species. Stag beetles are shown to be more active during the first part of their adult lives. Males were more prone to disperse than females but the home range size did not differ between the sexes. Dividing the flight season in three intervals, the most active individuals were recorded during the first and the second ones (first and second half of June).


Predators

Natural predators of ''L. cervus'' in Britain include
cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...
s,
fox Foxes are small-to-medium-sized omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull; upright, triangular ears; a pointed, slightly upturned snout; and a long, bushy tail ("brush"). Twelve species ...
es,
badger Badgers are medium-sized short-legged omnivores in the superfamily Musteloidea. Badgers are a polyphyletic rather than a natural taxonomic grouping, being united by their squat bodies and adaptions for fossorial activity rather than by the ...
s,
carrion crow The carrion crow (''Corvus corone'') is a passerine bird of the family Corvidae, native to western Europe and the eastern Palearctic. Taxonomy and systematics The carrion crow was one of the many species originally described by Carl Linnaeus ...
s,
magpie Magpies are birds of various species of the family Corvidae. Like other members of their family, they are widely considered to be intelligent creatures. The Eurasian magpie, for instance, is thought to rank among the world's most intelligent c ...
s,
woodpeckers Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar and the extreme polar regions. ...
,
kestrel The term kestrel (from , derivative from , i.e. ratchet) is the common name given to several species of predatory birds from the falcon genus ''Falco''. Kestrels are most easily distinguished by their typical hunting behaviour which is to hover ...
s, and false black widows; these tend to strike at the most vulnerable stage in the beetle's life cycle, when adults are seeking to mate and lay eggs. In the case of magpies, they have been observed in the field as waiting for emergence on a single site, subsequently consuming the beetle's abdomen. In Italy, the main predator is the
hooded crow The hooded crow (''Corvus cornix''), also colloquially called just hoodie, is a Eurasian bird species in the genus '' Corvus''. Widely distributed, it is found across Northern, Eastern, and Southeastern Europe, as well as parts of the Middle E ...
(''Corvus cornix'').


Parasites

Mites in the order
Monogynaspida Mesostigmata is an order of mites belonging to the Parasitiformes. They are by far the largest group of Parasitiformes, with over 8,000 species in 130 families. Mesostigmata includes parasitic as well as free-living and predatory forms. They ca ...
(Suborder Uropodina) are
phoretic Phoresis or phoresy is a temporary Commensalism, commensalistic relationship when an organism (a phoront or phoretic) attaches itself to a host organism solely for travel. It has been seen in tick, ticks and mite, mites since the 18th century, ...
(attached for the purpose of transportation) and have been observed attached to ''L. cervus'' as deutonymphs on the membrane of the joint between head and pronotum.


Relationship with humans


Protection

''Lucanus cervus'' is listed as
Near Threatened A near-threatened species is a species which has been Conservation status, categorized as "Near Threatened" (NT) by the International Union for Conservation of Nature (IUCN) as that may be vulnerable to Endangered species, endangerment in the ne ...
by the
IUCN 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 ...
. Although this species is widely distributed in Europe, it is in significant decline in the north and central part of its range and future trends of European forests will pose serious threats to this species, thus making the species close to qualifying for Vulnerable. ''Lucanus cervus'' is registered in the second appendix of the
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 ...
of the
European Union The European Union (EU) is a supranational union, supranational political union, political and economic union of Member state of the European Union, member states that are Geography of the European Union, located primarily in Europe. The u ...
from 1992, which requires that member states set aside
special areas of conservation A special area of conservation (SAC) is defined in the European Union's Habitats Directive (92/43/EEC), also known as the ''Directive on the Conservation of Natural Habitats and of Wild Fauna and Flora''. They are to protect the 220 habitats and ap ...
. The species is also registered in the third appendix of the Convention on the Conservation of European Wildlife and Natural Habitats (Berne convention) of 1982 and Schedule 5 of the UK's
Wildlife and Countryside Act 1981 The Wildlife and Countryside Act 1981 (c. 69) is an act of Parliament in the United Kingdom implemented to comply with European Council Directive 79/409/EEC on the conservation of wild birds. In short, the act gives protection to native species ...
.


Log pyramids

Log pyramids, also called stumperies, are vertical log piles where the logs are buried upright, providing plenty of underground rotten broadleaf wood for stag beetle larvae to feed on, as their natural food sources are in decline.


Gallery

File:Lucanus cervus cervus David Goliath.JPG, Size variation of males File:Cerf-volant MHNT Ventre.jpg, Underside File:Lucanus cervus turcicus (Bulgaria) TP54.jpg, Antenna File:Lucanus cervus turcicus (Bulgaria) TP40.jpg, Mouthparts – the galea (feathery) and the labial palps File:Hirschkäfer im Flug im FFH-Gebiet Reliktwald Lampertheim.jpg, Flying File:Hirschkäfer Paarung.jpg, Mating File:Lucanus cervus akbesianus Ei.JPG, Egg File:Pupa of Lucanus cervus akbesianus 3.JPG, Pupa File:Lucanus Cervus fighting.webm, Video of males fighting


References


External links

* *
stagbeetle.info Research site ran
by
Royal Holloway Royal Holloway, University of London (RH), formally incorporated as Royal Holloway and Bedford New College, is a public university, public research university and a constituent college, member institution of the federal University of London. It ...
, University of London
''Biology of the Stag Beetle''
translated from the Spanish article "''de lo poco conocido y lo mucho por conocer''"
3D model of ''Lucanus cervus''

LIFE Rosalia: Lucanus cervus
{{Taxonbar, from=Q190491 Beetles of Europe
cervus ''Cervus'' is a genus of deer that primarily are native to Eurasia, although one species occurs in northern Africa and another in North America. In addition to the species presently placed in this genus, it has included a whole range of other s ...
Beetles described in 1758 Taxa named by Carl Linnaeus Habitats Directive species