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Dynastes hercules

Dynastes hercules , Photo: Michael Lahanas

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Polyphaga
Infraordo: Scarabaeiformia
Superfamilia: Scarabaeoidea

Familia: Scarabaeidae
Subfamilia: Dynastinae
Tribus: Dynastini
Genus: Dynastes
Species: Dynastes hercules
Subspecies (12): D. h. bleuzeni – D. h. ecuatorianus – D. h. hercules – D. h. lichyi – D. h. morishimai – D. h. occidentalis – D. h. paschoali – D. h. reidi – D. h. septentrionalis – D. h. takakuwai – D. h. trinidadensis

In synonymy (1): D. h. tuxtlaensis
Name

Dynastes hercules (Linnaeus, 1758)
Original combination: Scarabaeus hercules
Type locality: 'America'

Synonyms

Scarabaeus scaber Linnaeus, 1758

Primary references

Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio Decima, Reformata. Tomus I. Holmiæ (Stockholm): impensis direct. Laurentii Salvii. 824 pp. DOI: 10.5962/bhl.title.542 BHL Reference page. [first availability, see p. 345]

Links

AnimalBase
Hercules
ION (subspecies list)
Catálogo electrónico de los organismos presentes en Colombia
Mystery behind the strongest creature in the world en:Institute of Physics (2008-03-11)
ZooBank

Vernacular names
беларуская: Жук-геркулес
Deutsch: Herkuleskäfer
English: Hercules beetle
français: Dynaste Hercule
italiano: Scarabeo Ercole
português: Besouro hércules
русский: Жук-геркулес
svenska: Herkulesbagge
ไทย: ด้วงกว่างเฮอร์คิวลิส

The Hercules beetle (Dynastes hercules) is a species of rhinoceros beetle native to the rainforests of Central America, South America, and the Lesser Antilles. It is the longest extant species of beetle in the world, and is also one of the largest flying insects in the world.

Etymology

Dynastes hercules is known for its tremendous strength and is named after Hercules, a hero of classical mythology who is famed for his great strength.
Taxonomy

D. hercules has a complex taxonomic history and has been known by several synonyms. It is in the subfamily Dynastinae (rhinoceros beetles) in the larger family Scarabaeidae (commonly known as scarab beetles). Not counting subspecies of D. hercules, seven other species are recognized in the genus Dynastes.
Subspecies

Several subspecies of D. hercules have been named,[1][2] though still some uncertainty exists as to the validity of the named taxa.[3][4]

Dynastes hercules ecuatorianus Ohaus, 1913
Dynastes hercules hercules (Linnaeus, 1758)
Dynastes hercules lichyi Lachaume, 1985
Dynastes hercules morishimai Nagai, 2002
Dynastes hercules occidentalis Lachaume, 1985
Dynastes hercules paschoali Grossi & Arnaud, 1993
Dynastes hercules reidi Chalumeau, 1977 (= baudrii Pinchon, 1976)
Dynastes hercules septentrionalis Lachaume, 1985 (= tuxtlaensis Moron, 1993)
Dynastes hercules takakuwai Nagai, 2002
Dynastes hercules trinidadensis Chalumeau & Reid, 1995 (= bleuzeni Silvestre and Dechambre, 1995)

Description

Adult body sizes (not including the thoracic horn) vary between 50 and 85 mm (2.0 and 3.3 in) in length and 29 and 42 mm (1.1 and 1.7 in) in width.[5] Male Hercules beetles may reach up to 173 mm (7 in) in length (including the horn), making them the longest species of beetle in the world, if jaws and/or horns are included in the measurement.[3][6] The size of the horn is naturally variable, more so than any variation of the size of legs, wings, or overall body size in the species. This variability results from developmental mechanisms that coincide with genetic predisposition in relation to nutrition, stress, exposure to parasites, and/or physiological conditions.[7]

Dynastes hercules is highly sexually dimorphic, with only males exhibiting the characteristic horns (one on the head, and a much larger one on the prothorax).[8] The body of males is black with the exception of the elytra, which can have shades of olive-green. They have a black suture with sparsely distributed black spots elsewhere on the elytra.[5] They have a slightly iridescent coloration to their elytra, which varies in color between specimens and may be affected by the humidity of the local environment in which they develop.[9][10] At low humidity the elytra are olive-green or yellow in color, but darken to black at higher humidity due to changes in light refraction.[11]

Females of D. hercules have punctured elytra which are usually entirely black, but sometimes have the last quarter of the elytra colored in the same way as the males.[5]

Female

Male

Hercules beetle illustration from The Naturalist's Miscellany (1789-1813) by George Shaw (1751-1813).

Distribution and habitat

Populations of D. hercules may be found from southern Mexico to Bolivia in mountainous and lowland rain forests. Known populations include the Lesser Antilles, Trinidad and Tobago, Brazil, Ecuador, Colombia, and Peru.[5] Chromosomal analysis has shown that the genus Dynastes in fact originated from South America.[12]
Life cycle
Larva of D. hercules

Not much is known about the life cycle in the wild, but much evidence has been gained through observations of captive-bred populations.[5] The mating season for adults typically occurs during the rainy season (July to December). Females have an average gestation period of 30 days from copulation to egg-laying, and may lay up to 100 eggs on the ground or on dead wood. The eggs have an incubation period of approximately 27.7 days before they hatch. [11] Once hatched, the larval stage of the Hercules beetle may last up to two years in duration, where it will go through 3 metamorphosis stages, also known as instars.[5] The larvae have a yellow body with a black head. The larvae can grow up to 11 cm (4.5 in) in length and weigh more than 100 grams.[11] In laboratory conditions at 25 ± 1°C, the first instar stage lasts an average of 50 days, the second stage an average of 56 days, and the third an average of 450 days. After the third instar stage, the pupal stage lasts about 32 days, where it will transition into an adult. Adult beetles can live for three to six months in captivity.[5]
Diet and behavior
Diet

The larvae of the Hercules beetle are saproxylophagous, meaning that they feed on rotting wood; they reside in same during their two-year developmental stage. [5][9] The adult Hercules beetle feeds on fresh and rotting fruit.[9][10] They have been observed feeding on peaches, pears, apples, and grapes in captivity.[10][13]
Behavior

Within their native rain forest habitats, the adult beetles, which are nocturnal, forage for fruit at night and hide or burrow within the leaf litter during the day.[11] The adult D. hercules beetles are capable of creating a 'huffing' sound, generated by stridulating their abdomen against their elytra to serve as a warning to predators.[11][14] Like most insects, communication within the species is a mix of chemoreception, sight, and mechanical perception. Experiments on D. hercules have shown that a male placed in the vicinity of a female will immediately orient towards her and seek her out, suggesting chemical communication through strong sexual pheromones.[14]
Combat behavior between males

It has been observed in wild habitat and in captivity that male D. hercules will engage in combat to win possession and mating rights to a female. [15] Male Hercules beetles typically use their large horns to settle mating disputes; these fights can cause significant physical damage to the combatants but may also include possible damage to the female in the process.[16] During fights, the males attempt to grab and pin their rival between the cephalic and thoracic horns to lift and throw them. The successful male wins mating rights with the female, though the beetles remain polygynandrous.[11]
Physical strength

Reports suggest the Hercules beetle is able to carry up to 850 times its body mass. Actual measurements on a much smaller (and relatively stronger: see square-cube law) species of rhinoceros beetle shows a carrying capacity only up to 100 times their body mass, at which point they can barely move.[17]
Relationship to humans

Dynastes hercules does not negatively affect human activities, either as an agricultural pest or disease vector. The beetles can be kept as pets.[11]
Relationship to the environment

Dynastes hercules is a beneficial contributor to the rain forest ecosystem, primarily during their larval stage where they are saproxylophagous. Feeding on rotting wood assists with biodegradation and cycling nutrients in the environment.[5]
See also

List of largest insects

References

"Hercules Beetle, Dynastes hercules". BioLib.cz. 2017. Archived from the original on 2018-12-06. Retrieved 2017-07-09.
Huang, J.P. (15 September 2017). "The Hercules beetles (subgenus Dynastes, genus Dynastes, dynastidae) A revisionary study based on the integration of molecular, morphological, ecological, and geographic analyses". Miscellaneous Publications, Museum of Zoology, University of Michigan. 206: 1–32 – via ResearchGate.
Ratcliffe, B.C.; Cave, R.D. (2015). "The dynastine scarab beetles of the West Indies (Coleoptera: Scarabaeidae: Dynastinae)". Bulletin of the University of Nebraska State Museum. 28: l–346.
Huang, J., Knowles, L. The species versus subspecies conundrum: quantitative delimitation from integrating multiple data types within a single bayesian approach in Hercules beetles. Systematic Biology, vol. 65, 2015, 15p.
Keller, O.; Cave, R. D. Cave (2016). "Hercules Beetle Dynastes hercules (Linnaeus, 1758) (Insecta: Coleoptera: Scarabaeidae)" (PDF). University of Florida, IFAS Extension. Archived (PDF) from the original on 2017-08-29. Retrieved 2017-05-20.
"Largest species of beetle". Guinness World Records. Archived from the original on 2017-05-09. Retrieved 2017-05-20.
Lavine, L.; Gotoh, H.; Brent, C. S.; Dworkin, I.; Emlen, D. J. (2015). "Exaggerated Trait Growth in Insects". Annual Review of Entomology. 60: 453–472. doi:10.1146/annurev-ento-010814-021045. PMID 25341090.
Huang, J. (2016). "Parapatric genetic introgression and phenotypic assimilation: testing conditions for introgression between Hercules beetles (Dynastes, Dynastinae)". Molecular Ecology. 25 (21): 5513–5526. doi:10.1111/mec.13849. hdl:2027.42/134423. PMID 27661063. S2CID 22612044.
Rassart, M.; Colomer, J. F.; Tabarrant, T.; Vigneron, J. P. (2008). "Diffractive hygrochromic effect in the cuticle of the hercules beetle Dynastes hercules". New Journal of Physics. 10 (3): 033014. Bibcode:2008NJPh...10c3014R. doi:10.1088/1367-2630/10/3/033014.
Hinton, H. E.; Jarman, G. M. (1973). "Physiological colour change in the elytra of the Hercules beetle, Dynastes hercules". Journal of Insect Physiology. 19 (3): 533–539. doi:10.1016/0022-1910(73)90064-4..
Toussaint, A. (2015). "Dynastes hercules (Hercules Beetle)" (PDF). The Online Guide to the Animals of Trinidad and Tobago (University of West Indies). Archived (PDF) from the original on 18 February 2018. Retrieved 17 February 2018.
Dutrillaux B.; Dutrillaux A.-M. (2013). "A South American Origin of the Genus Dynastes (Coleoptera: Scarabaeidae: Dynastinae) Demonstrated by Chromosomal Analyses". Cytogenetic and Genome Research. 147 (1): 37–42. doi:10.1159/000351210. PMID 23735513. S2CID 28561729.
Krell, F., Krell, V. Longevity of the Western Hercules beetle, D. grantii Horn (Coleoptera: Scarabaeidae: Dynastinae). The Coleopterists Bulletin, vol. 69, 2015, 1p.
Kulikowski, A. "Dynastes hercules". Animal Diversity Web. Archived from the original on 2017-05-22. Retrieved 2017-05-20.
Beebe, W. (1947). "Notes on the Hercules Beetle, Dynastes hercules (Linn.), at Rancho Grande, Venezuela, with Special Reference to Combat Behavior". Department of Tropical Research, New York Zoological Society. 32: 109–116.
"Hercules Beetles". University of Kentucky Entomology. 2008. Archived from the original on 2017-03-30. Retrieved 2017-05-19.
Kram, R. (1996). "The Journal of Experimental Biology" (PDF). 199: 609–612. Archived (PDF) from the original on 2017-08-10. Retrieved 2017-05-19.

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