''Eleodes'' (commonly known as pinacate beetles or desert stink beetles) is a genus of

darkling beetles Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is the Latin genus, generic name that Carl Linnaeus assigned to some flour be ...

, in the family

Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is the Latin generic name that Carl Linnaeus assigned to some flour beetles ...

. They are endemic to western

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

ranging from southern

Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...

to central

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

with many species found along the Mexico-United States border. Some species have been introduced to

Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...

. The name ''pinacate'' is

Mexican Spanish Mexican Spanish () is the variety of dialects and sociolects of the Spanish language spoken in Mexico and its bordering regions. Mexico has the largest number of Spanish speakers, more than double any other country in the world. Spanish is spo ...

, derived from the

Nahuatl Nahuatl ( ; ), Aztec, or Mexicano is a language or, by some definitions, a group of languages of the Uto-Aztecan language family. Varieties of Nahuatl are spoken by about Nahuas, most of whom live mainly in Central Mexico and have smaller popul ...

(Aztec) name for the insect, ''pinacatl'', which translates as "black beetle". ''Eleodes'' species range from about in length and are black in color with some having a reddish tint on their abdomen. The

seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...

e of some species such as '' Eleodes osculans'' collect debris and give the insect a brown color, a similar effect occurs in the species '' Eleodes mirabilis'' giving it the appearance of having whitish stripes on its abdomen. Due to the number of species and their large range these beetles have a fairly varied appearance throughout the many species. All produce

quinone The quinones are a class of organic compounds that are formally "derived from aromatic compounds benzene.html" ;"title="uch as benzene">uch as benzene or naphthalene] by conversion of an even number of –CH= groups into –C(=O)– groups with ...

or similar substances as a deterrent to predators, and many will stand on their heads to spray it. They are typically found in the arid desert regions of their range but can also be found in forests and grasslands. All ''Eleodes'' species are flightless as their Elytron, elytra are fused together and their second pair of wings is very reduced and

vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...

Ecology

Most members of the genus ''Eleodes'' are primarily

detrivores Detritivores (also known as detrivores, detritophages, detritus feeders or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...

feeding on plant and animal debris, some species are more inclined to consume living plants which is especially true for species native to the plains of

such as '' Eleodes hispilabris'' and '' Eleodes opaca''. The

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 ...

e of those species and others are considered

pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...

as they feed on the roots of crops. Species found in dry regions such as members of the subgenus ''Eleodes'' are usually much larger than what is expected for most insects, this is due the lack of water in the area leading to these beetles evolving larger bodies in order to hold more of it. Water loss is also the reason that the Elytron, elytra are fused in ''Eleodes''. As expected, species found in forests, such as members of the subgenus '' Blapylis'' are smaller, closer to the size of most other insects. ''Eleodes'' are generally more active at night, some species are strictly

nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...

while others can and will be active during the day. As these beetles are fairly long lived when it comes to insects they must adapt to the different seasons. During the fall they are diurnal as it is warmest during the day. When winter comes they take shelter and become inactive as to not freeze in the cold. When spring arrives they go back to being active in the day until the summer when they switch to being active at night to survive the deadly high temperatures of the desert. ''Eleodes'' take shelter almost anywhere that can protect them from the hot and cold, this can be wood or rocks but is often rodent burrows. Every night when these beetles scavenge for food they pick a new shelter for the day which if not suitable for protection against the elements, will be abandoned in search of a new one. One subgenus, '' Caverneleodes,'' lives exclusively in caves, cave-like environments, or rock crevices, these beetles have reduced eyes and only leave their hideouts to

scavenge Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...

for food.

Chemical defense

Beetles within ''Eleodes'' and other genera within

exude a foul smelling odor and a very distasteful liquid from

gland A gland is a Cell (biology), cell or an Organ (biology), organ in an animal's body that produces and secretes different substances that the organism needs, either into the bloodstream or into a body cavity or outer surface. A gland may also funct ...

s to ward off predators. This liquid is most commonly a form of

. While not being

poison A poison is any chemical substance that is harmful or lethal to living organisms. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figurati ...

ous the liquid does usually deter predators, the success of this defense mechanism has led to the evolution of a

Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimicry, mimic each other's honest signal, honest aposematism, warning signals, to their mutuali ...

complex among '' Eleodes acuticauda'', '' Eleodes dentipes'', and '' Coelocnemis magna'', the latter of which while not being within ''Eleodes'' is a still distasteful prey item. All three previously mentioned species look strikingly similar leading to a strengthened defense against potential predators. Another Müllerian mimicry complex exists between '' Eleodes scabrosa'' and '' Eleodes tuberculata.'' The "head-standing" behavior exhibited by these beetles is aposematic. By leaving their presence and identity unambiguous to potential predators, it means that more individuals can survive as predators will avoid beetles with the "head-standing" behavior. While this defense is generally effective in deterring predators, they are still preyed upon by many species, such as

owls Owls are birds from the Order (biology), order Strigiformes (), which includes over 200 species of mostly Solitary animal, solitary and Nocturnal animal, nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vis ...

foxes 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 ...

coyote The coyote (''Canis latrans''), also known as the American jackal, prairie wolf, or brush wolf, is a species of canis, canine native to North America. It is smaller than its close relative, the Wolf, gray wolf, and slightly smaller than the c ...

skunk Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or gi ...

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 ...

turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...

s, etc.

Fossil record and evolution

Fossils of '' Eleodes acuticauda'', '' Eleodes osculans'', '' Eleodes carbonaria'', '' Eleodes granulata'', '' Eleodes gracilis'', and others have been found in the

La Brea Tar Pits La Brea Tar Pits comprise an active Paleontological site, paleontological research site in urban Los Angeles. Hancock Park was formed around a group of tar pits where natural Bitumen, asphalt (also called asphaltum, bitumen, or pitch; ''brea'' ...

, Carpinteria Tar Pits, and the

Snowmastodon Site The Snowmastodon site, also known as the Ziegler Reservoir fossil site, is the location of an important Ice Age fossil excavation near Snowmass Village, Colorado. Fossils were first discovered on October 14, 2010, during the construction of a ...

which are all dated to the

Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...

epoch. ''Eleodes'' split off from the rest of Amphidorini at least 8 to 15 million years ago in the Miocene, although this estimate is likely on the lower side due to low taxon sampling.

Taxonomy

''Eleodes'' is the largest

New World The term "New World" is used to describe the majority of lands of Earth's Western Hemisphere, particularly the Americas, and sometimes Oceania."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: ...

genus of

darkling beetle Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is the Latin generic name that Carl Linnaeus assigned to some flour beetles ...

s comprising around 200 species divided up into the following 17 subgenera. Subgenera: * Subgenus Amphidora Eschscholtz, 1829 **'' Eleodes littoralis'' Eschscholtz, 1829 **'' Eleodes nigropilosa'' LeConte, 1851 **'' Eleodes subdeplanta'' Blaisdell, 1943 * Subgenus Ardeleodes **'' Eleodes tibialis'' Blaisdell, 1909 * Subgenus Blapylis Horn, 1870 **'' Eleodes alticola'' Blaisdell, 1925 **'' Eleodes aristata'' Somerby, 1977 **'' Eleodes bishopensis'' Somerby and Doyen, 1976 **'' Eleodes blanchardii'' Blaisdell, 1909 **'' Eleodes brunnipes'' Casey, 1890 **'' Eleodes caseyi'' Blaisdell, 1909 **'' Eleodes clavicornis'' Eschscholtz, 1829 **'' Eleodes consobrina'' LeConte, 1851 **'' Eleodes constricta'' LeConte, 1858 **'' Eleodes cooperi'' Somerby and Doyen, 1976 **'' Eleodes cordata'' Eschscholtz, 1829 **'' Eleodes fuchsii'' Blaisdell, 1909 **'' Eleodes hoppingii'' Blaisdell, 1909 **'' Eleodes hornii'' Blaisdell, 1909 **'' Eleodes hybrida'' Blaisdell, 1917 **'' Eleodes inculta'' LeConte, 1861 **'' Eleodes kaweana'' Blaisdell, 1933 **'' Eleodes lariversi'' Somerby and Doyen, 1976 **'' Eleodes lecontei'' Horn, 1870 **'' Eleodes manni'' Blaisdell, 1917 **'' Eleodes nana'' Blaisdell, 1909 **'' Eleodes neotomae'' Blaisdell, 1909 **'' Eleodes novoverrucula'' Boddy, 1957 **'' Eleodes nunenmacheri'' Blaisdell, 1918 **'' Eleodes oregona'' Blaisdell, 1941 **'' Eleodes orophila'' Somerby, 1977 **'' Eleodes panamintensis'' Somerby, 1977 **'' Eleodes parvicollis'' Eschscholtz, 1829 **'' Eleodes patulicollis'' Blaisdell, 1932 **'' Eleodes pimelioides'' Mannerheim, 1843 **'' Eleodes planata'' Eschscholtz, 1829 **'' Eleodes producta'' Mannerheim, 1843 **'' Eleodes propinqua'' Blaisdell, 1918 **'' Eleodes robinetti'' Boddy, 1957 **'' Eleodes rotundipennis'' LeConte, 1857 **'' Eleodes scabripennis'' LeConte, 1859 **'' Eleodes scabriventris'' Blaisdell, 1932 **'' Eleodes scabrosa'' Eschscholtz, 1829 **'' Eleodes schlingeri'' Somerby and Doyen, 1976 **'' Eleodes schwarzii'' Blaisdell, 1909 **'' Eleodes snowii'' Blaisdell, 1909 **'' Eleodes spilmani'' Somerby and Doyen, 1976 **'' Eleodes strumosa'' Blaisdell, 1932 **'' Eleodes subvestita'' (Blaisdell, 1939) **'' Eleodes tenebrosa'' Horn, 1870 **'' Eleodes triplehorni'' Somerby and Doyen, 1976 **'' Eleodes trita'' Blaisdell, 1917 **'' Eleodes tuberculata'' Eschscholtz, 1829 **'' Eleodes versatilis'' Blaisdell, 1921 **'' Eleodes volcanensis'' Somerby, 1977 **'' Eleodes wakelandi'' Somerby, 1977 * Subgenus Caverneleodes Triplehorn, 1975 ** '' Eleodes easterlai'' Triplehorn, 1975 ** '' Eleodes gruta'' Aalbu, Smith & Triplehorn, 2012 ** '' Eleodes guadalpensis'' Aalbu, Smith & Triplehorn, 2012 ** '' Eleodes labialis'' Triplehorn, 1975 ** '' Eleodes leptoscelis'' Triplehorn, 1975 ** '' Eleodes microps'' Aalbu, Smith & Triplehorn, 2012 ** '' Eleodes reddelli'' Triplehorn, 2007 ** '' Eleodes rugosifrons'' Triplehorn & Reddell, 1991 ** '' Eleodes sprousei'' Triplehorn & Reddell, 1991 ** '' Eleodes thomasi'' Aalbu, Smith & Triplehorn, 2012 ** '' Eleodes wheeleri'' Aalbu, Smith & Triplehorn, 2012 ** '' Eleodes wynnei'' Aalbu, Smith & Triplehorn, 2012 * Subgenus Chaseleodes Thomas, 2015 **'' Eleodes connata'' Solier, 1848 **'' Eleodes curta'' Champion, 1884 * Subgenus Cratidus LeConte, 1862 **'' Eleodes osculans'' LeConte, 1851 **'' Eleodes ursus'' Triplehorn, 1996 * Subgenus Discogenia LeConte, 1866 **'' Eleodes acutangula'' Blaisdell, 1921 **'' Eleodes marginata'' Eschscholtz, 1829 **'' Eleodes scabricula'' LeConte, 1858 * Subgenus

Eleodes ''Eleodes'' (commonly known as pinacate beetles or desert stink beetles) is a genus of darkling beetles, in the family Tenebrionidae. They are endemic to western North America ranging from southern Canada to central Mexico with many species fou ...

Eschscholtz, 1829 ** '' Eleodes acuta'' (Say, 1824) ** '' Eleodes acuticauda'' LeConte, 1851 ** '' Eleodes adumbrata'' Blaisdell, 1925 ** '' Eleodes armata'' LeConte, 1851 ** '' Eleodes curvidens'' Triplehorn & Cifuentes-Ruiz, 2011 ** '' Eleodes dentipes'' Eschscholtz, 1829 ** '' Eleodes discincta'' Blaisdell, 1925 ** '' Eleodes eschscholtzii'' Solier, 1848 ** '' Eleodes femorata'' LeConte, 1851 ** '' Eleodes fiski'' Triplehorn, 2015 ** '' Eleodes gracilis'' LeConte, 1858 ** '' Eleodes grandicollis'' Mannerheim, 1843 ** '' Eleodes hispilabris'' (Say, 1824) ** '' Eleodes loretensis'' Blaisdell, 1923 ** '' Eleodes mexicana'' Blaisdell, 1943 ** '' Eleodes mirabilis'' Triplehorn, 2007 ** '' Eleodes moesta'' Blaisdell, 1921 ** '' Eleodes muricatula'' Triplehorn, 2007 ** '' Eleodes obscura'' (Say, 1824) ** '' Eleodes rossi'' Blaisdell, 1943 ** '' Eleodes rugosa'' Perbosc, 1839 ** '' Eleodes samalayucae'' Triplehorn, 2007 ** '' Eleodes sanmartinensis'' Blaisdell, 1921 ** '' Eleodes scyroptera'' Triplehorn, 2007 ** '' Eleodes spinipes'' Solier, 1848 ** '' Eleodes sponsa'' LeConte, 1858 ** '' Eleodes subcylindrica'' Casey, 1890 ** '' Eleodes suturalis'' (Say, 1824) ** '' Eleodes tenuipes'' Casey, 1890 ** '' Eleodes vanduzeei'' Blaisdell, 1923 * Subgenus Heteropromus Blaisdell, 1909 **'' Eleodes veterator'' Horn, 1874 * Subgenus Litheleodes Blaisdell, 1909 **'' Eleodes arcuata'' Casey, 1884 **'' Eleodes aspera'' LeConte, 1866 **'' Eleodes corvina'' Blaisdell, 1921 **'' Eleodes extricata'' (Say, 1824) **'' Eleodes granulata'' LeConte, 1857 **'' Eleodes hirtipennis'' Triplehorn, 1964 **'' Eleodes letcheri'' Blaisdell, 1909 **'' Eleodes papillosa'' Blaisdell, 1917 **'' Eleodes subtuberculata'' Walker, 1866 * Subgenus Melaneleodes Blaisdell, 1909 ** '' Eleodes tricostata'' (Say, 1824) ** '' Eleodes neomexicana'' Blaisdell, 1909 ** '' Eleodes pedinoides'' LeConte, 1858 ** '' Eleodes wenzeli'' Blaisdell, 1925 ** '' Eleodes parowana'' Blaisdell, 1925 ** '' Eleodes halli'' Blaisdell, 1941 ** '' Eleodes carbonaria'' (Say, 1824) ** '' Eleodes quadricollis'' Eschscholtz, 1829 ** '' Eleodes anthracina'' Blaisdell, 1909 ** '' Eleodes humeralis'' LeConte, 1857 ** '' Eleodes rileyi'' Casey, 1891 ** '' Eleodes rufipes'' Pierre, 1976 * Subgenus Metablapylis Blaisdell, 1909 **'' Eleodes aalbui'' Triplehorn, 2007 **'' Eleodes californica'' Blaisdell, 1929 **'' Eleodes delicata'' Blaisdell, 1929 **'' Eleodes dissimilis'' Blaisdell, 1909 **'' Eleodes nevadensis'' Blaisdell, 1909 **'' Eleodes nigrina'' LeConte, 1858 * Subgenus Omegeleodes Triplehorn & Thomas, 2011 ** '' Eleodes debilis'' LeConte, 1858 * Subgenus Promus LeConte, 1862 **'' Eleodes anachronus'' Triplehorn, 2010 **'' Eleodes bidens'' Triplehorn, 2007 **'' Eleodes brucei'' Triplehorn, 2007 **'' Eleodes calcarata'' Champion, 1884 **'' Eleodes composita'' Casey, 1891 **'' Eleodes erratica'' Champion, 1884 **'' Eleodes exarata'' Champion, 1884 **'' Eleodes fusiformis'' LeConte, 1858 **'' Eleodes goryi'' Solier, 1848 **'' Eleodes hoegei'' Champion, 1884 **'' Eleodes insularis'' Linell, 1899 **'' Eleodes knullorum'' Triplehorn, 1971 **'' Eleodes longicornis'' Champion, 1884 **'' Eleodes madrensis'' Johnston, 2015 **'' Eleodes montana'' Champion, 1884 **'' Eleodes opaca'' (Say, 1824) **'' Eleodes spiculifera'' Triplehorn, 2007 **'' Eleodes spinolae'' Solier, 1848 **'' Eleodes striolata'' LeConte, 1858 **'' Eleodes subnitens'' LeConte, 1851 **'' Eleodes watrousi'' Triplehorn, 2007 * Subgenus Pseudeleodes Blaisdell, 1909 ** '' Eleodes caudifera'' LeConte, 1858 ** '' Eleodes granosa'' LeConte, 1866 ** '' Eleodes inornata'' Johnston, 2016 ** '' Eleodes leechi'' Tanner, 1961 ** '' Eleodes longipilosa'' Horn, 1891 ** '' Eleodes pilosa'' Horn, 1870 ** '' Eleodes spoliata'' Blaisdell, 1933 ** '' Eleodes tribulus'' Thomas, 2005 * Subgenus Steneleodes Blaisdell, 1909 **'' Eleodes angulata'' (Eschscholtz, 1829) **'' Eleodes angusta'' Eschscholtz, 1829 **'' Eleodes blapoides'' Eschscholtz, 1829 **'' Eleodes coarctata'' Champion, 1885 **'' Eleodes corrugans'' Triplehorn, 2007 **'' Eleodes distincta'' Solier, 1848 **'' Eleodes forreri'' Champion, 1884 **'' Eleodes gigantea'' Mannerheim, 1843 **'' Eleodes glabricollis'' Champion, 1884 **'' Eleodes gravida'' (Eschscholtz, 1829) **'' Eleodes hepburni'' Champion, 1884 **'' Eleodes innocens'' LeConte, 1866 **'' Eleodes laevigata'' Solier, 1848 **'' Eleodes longicollis'' LeConte, 1851 **'' Eleodes mutilata'' Blaisdell, 1921 **'' Eleodes olida'' Champion, 1892 **'' Eleodes ornatipennis'' Blaisdell, 1937 **'' Eleodes peropaca'' Champion, 1892 **'' Eleodes platypennis'' Triplehorn, 2007 **'' Eleodes ponderosa'' Champion, 1884 **'' Eleodes punctigera'' Blaisdell, 1935 **'' Eleodes ruida'' (Say, 1835) **'' Eleodes sallaei'' Champion, 1885 **'' Eleodes solieri'' Champion, 1885 **'' Eleodes stolida'' Champion, 1885 **'' Eleodes sulcatula'' Champion, 1884 **'' Eleodes tenebricosa'' Gemminger, 1870 **'' Eleodes tessellata'' Champion, 1892 * Subgenus Tricheleodes Blaisdell, 1909 ** '' Eleodes hirsuta'' LeConte, 1861 ''

Incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...

'': **'' Eleodes aequalis'' (Say, 1835) **'' Eleodes amaura'' Champion, 1892 **'' Eleodes barbata'' Wickham, 1918 **'' Eleodes brevicollis'' Gemminger, 1870 **'' Eleodes cylindrica'' (Herbst, 1799) **'' Eleodes dilaticollis'' Champion, 1884 **'' Eleodes ebenina'' (Solier, 1848) **'' Eleodes elongatula'' Eschscholtz, 1829 **'' Eleodes impolita'' (Say, 1835) **'' Eleodes maura'' (Say, 1835) **'' Eleodes melanaria'' Eschscholtz, 1829 **'' Eleodes obliterata'' (Say, 1835) **'' Eleodes polita'' Champion, 1892 **'' Eleodes rotundicollis'' (Eschscholtz, 1829) **'' Eleodes scapularis'' Champion, 1884 **'' Eleodes segregata'' Champion, 1892 **'' Eleodes striata'' (Guérin-Méneville, 1834) **'' Eleodes sulcata'' (Eschscholtz, 1829)

Notes

References

External links

Desert USA – Stinkbugs
{{Taxonbar, from=Q6938538 Tenebrionidae genera Gran Desierto de Altar Fauna of the Sonoran Desert Taxa named by Johann Friedrich von Eschscholtz