Biology:Agriotes

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Agriotes is a very large genus of beetles in the family of click beetles or Elateridae. More than 350 species have been described and are currently recognized as valid, see the list of Agriotes species. The natural habitat of many species is grassland.[1][2] Breeding takes place in the soil, like in other groups of click beetles, where the larvae or wireworms feed on seeds and roots.[3] The genus is widely distributed in Europe, Asia (including the Near East), northern Africa and the Americas.[4][5] Compared to related genera of click beetles, species of Agriotes can be mainly recognized by structures on the pronotum and the head (see below).[6][7][8] The larvae of common species, can be harmful to crops and damage cereals like maize, forage crops like alfalfa, or root crops like potatoes and sugar beet.[9][10] Three Agriotes species are invasive and have spread to North America, where they also have become serious agricultural pests and are suppressing and displacing native species.[11][12]

Biology and ecology

Like many other click beetles, the species of the genus Agriotes breed in the soil and the larvae (wireworms) are elongated with a hardened cuticle. Details of their biology and ecology are mainly known for pest species. The general habitat of many species appears to be grassland.[13][3][2] Further, Agriotes species are often common in higher altitudes, in areas of higher precipitation during the winter and in acidic, humus-rich soils.[10][3]

For the more common and harmful Agriotes species, the food plants have been studied. The larvae are polyphagous and feed on seeds, roots and tubers, like those of grasses, cereals and other crops, while the adult beetles feed on the leaves of grasses. In addition, the larvae also feed on germinating seeds and on the stem of young seedlings.[14][15][16]

Agriotes female beetles lay their eggs in batches in the soil, close to the surface. The eggs hatch after several weeks. The larvae develop slowly, usually involving more than 10 instars and the whole life cycle from egg to egg extends over several years.[17][16]

Distribution

Distribution ranges of the species in the genus Agriotes in Europe, Asia and Africa. The blue outline delimitates the distribution of all species and the brown outline that of Agriotes sputator, the type species of the genus.

Species of the genus Agriotes are widely distributed around the world and often live in mountainous areas. However, distribution records often mention the species under different genera like the basionyms. In Europe, the genus is found in most areas except for the most northern parts of Norway, Sweden, Finland and Russia.[4][3][5] Actually, Agriotes sputator, the type species of the genus is also found in most parts of Europe.[18][19]

In Africa, species of Agriotes like Agriotes sordidus are found in the North-West, in Morocco, Algeria, Tunisia and as far south as the Canary Islands.[6][20] Further, there are some records from Egypt and Agriotes sikorae has been described from Madagascar, but with no records of Agriotes species from other parts of sub-Saharan Africa.[21]

Some parts of Western Asia have a high diversity of Agriotes species and Turkey has probably the highest number of Agriotes species of any country. A checklist published in 2025 lists 92 Agriotes species from Turkey (of which 2 are currently not recognized as valid for this genus).[22] Agriotes species are also common in countries neighbouring Turkey and in Central Asia, but the genus appears to be absent in Saudi Arabia and southern parts of Iraq, Iran and Afghanistan.[4] In the remaining parts of Asia, there are few records from Siberia, but the genus is present in all parts of China, the Koreas and in central and southern parts of Japan.[23][4] Further south, Agriotes species have been recorded in Myanmar, Thailand, Vietnam, Laos, Cambodia and mainland Malaysia (in some cases described under different genera).[24][25][23][26] In addition, Agriotes australasiae has been described from Aru Islands, an isolated location in Indonesia, near New Guinea.[27]

Distribution ranges of the species in the genus Agriotes in the Americas. The blue outline delimitates the distribution of the native species and the red outline that of 3 invasive pest species, Agriotes lineatus, A. obscurus and A. sputator.

In North America, Agriotes species have been described from Canada to Mexico, with the largest number of species reported from Mexico, especially from the southern states of Chiapas, Oaxaca and Vera Cruz.[28][29] The genus is also common in southern parts of Canada and northern parts of the U.S. Further, it is present in most central parts of the U.S., but is rare or absent in many southern parts, except for California. For example, Agriotes imperfectus has been described by LeConte from southern California.[30] The genus might also be present in other southern states of the U.S. like Arizona, Texas and Alabama.[5] However, those records appear to be unconfirmed.

Agriotes species have been recorded in most Central American countries and in some Caribbean Islands.[28][31] In South America, Agriotes species have been found in French Guiana, Uruguay, Argentina and central and southern parts of Chile (again, often reported under different genera).[32][33][34][35][36][37]

Description

Prothorax of Agriotes lineatus in side view, showing its shape and the course of the lateral ridge of the pronotum, marked in green on the right side (pronotum yellow and prosternum red).

The species of the genus Agriotes range in length from 3 and 13 mm. They are rarely larger than 13 mm. The species can be recognized by a combination of different characters. Often, the shape of the pronotum, especially the ridges on the sides of the pronotum are cited as the most important feature. Other characteristic features involve the head, the antennae, the elytra and the claws.[6][8]

The pronotum is longer than wide, sometimes only slightly. The lateral ridges are curved in an S-like manner with the part near the head (left side in the images at the right) bending into the ventral side of the pronotum. Therefore, from above the lateral ridges are only visible at the rear end of the pronotum and are hidden together with the sternum in the middle and in the front. In some species the lateral ridges are interrupted in the middle.[6][38][7][8]

Head of Agriotes obscurus in frontal view.

On the head, the mouthparts are pointing downwards (hypognathous). The ridges above the base of the antennae are poorly developed, especially towards the mouthparts, and often do not reach the clypeus, in contrast to the closely related genus Ectinus.[38][7]

The larval stages also have characteristic morphological features which can be used for identifying many species.[39] Further, identification by molecular methods is possible.[18]

Agricultural pests

Larva of Agriotes lineatus placed on a patch of moss.

Three Agriotes species, Agriotes lineatus, A. obscurus and A. sputator have been accidentally introduced into North America and have become invasive.[40] The larval stages of these and of several other species in Europe are serious agricultural pests, damaging cereals and other field crops. The larvae attack the seeds, roots, seedlings, stems and tubers, reducing the growth of the crops and in severe cases causing plant death. The main crops affected are maize, other cereals, alfalfa and root crops like sugar beet and potatoes.[3][9][41]

Vernon Pitfall Trap for catching Agriotes beetles. The lid of the trap has been opened and shows granules treated with the pheromone lure taped to the underside of the lid.

Pest species can be monitored using pheromone traps, which target flying beetles, or using pitfall traps, which target beetles walking on the ground. Agriotes females often produce a mixture of pheromones and these mixtures are typically species specific and can be used only for one species.[42] The pitfall traps can be made more attractive by combining them with a pheromone lure.[43]

In the past, farmers have relied on persistent organochlorine pesticides like lindane and aldrin for managing Agriotes pest species. However, due to their harmful effects on the environment, alternatives have been and are still being developed. So far, the most promising method is to switch to an alternative, less attractive crop if damaging levels of wireworms are detected.[44][45] Other methods include using less sensitive crops and timing tilling and irrigation in order to destroy eggs and young wireworm larvae in the top soil layer by desiccation.[45][46] Further, mating disruption with pheromones and biological control methods like the use of fungal pathogens have been studied.[41][43]

In Europe, maize is the crop most often attacked by Agriotes species, but the damage rarely reaches significant levels and an Integrated Pest Management (IPM) approach has been advocated. This means pest populations are monitored and management is only initiated once a threshold level has been reached.[47]

Taxonomy and species

The click beetle genus Agriotes has been named and described repeatedly, probably due to Agriotes species being rather common in Europe and in North America. Initially, the genus was named twice in 1829 when Eschscholtz first named the genus Agriotes and a few months later it was named again by Dillwyn as Hemirhypus.[48][49] The following year, it was named and described in detail once more by Stephens as Cataphagus.[50] Several more descriptions followed in the coming years, see the taxobox above.

The name Agriotes originates from the Greek word agrios (ἄγριος) which means wild and agriotes has been translated as wilderness.[51] The genus Agriotes is very large and more than 350 species are currently accepted as valid (status 2026), see the complete list of Agriotes species. The following short list includes the most common species and the most serious agricultural pests:

Identification of the individual species relies mainly on the size, colour, the presence of hairs on different body parts as well as the detailed structures of the elytra, pronotum and antennae.[52][53][8]

References

  1. ↑ Vernon, Robert S.; van Herk, Willem G.; Blackshaw, Roderick P.; Shimizu, Yoko; Clodius, Markus (2014). "Mark–recapture of Agriotes obscurus and Agriotes lineatus with dense arrays of pheromone traps in an undisturbed grassland population reservoir". Agricultural and Forest Entomology 16 (3): 217–226. doi:10.1111/afe.12045. 
  2. ↑ 2.0 2.1 Blackshaw, Rod P.; Vernon, Robert S.; Thiebaud, Florent (2018). "Large scale Agriotes spp. click beetle (Coleoptera: Elateridae) invasion of crop land from field margin reservoirs". Agricultural and Forest Entomology 20 (1): 51–61. doi:10.1111/afe.12228. 
  3. ↑ 3.0 3.1 3.2 3.3 3.4 Staudacher, Karin; Schallhart, Nikolaus; Pitterl, Peter; Wallinger, Corinna; Brunner, Nina; Landl, Marion; Kromp, Bernhard; Glauninger, Johann et al. (2013). "Occurrence of Agriotes wireworms in Austrian agricultural land". Journal of Pest Science 86: 33–39. doi:10.1007/s10340-011-0393-y. 
  4. ↑ 4.0 4.1 4.2 4.3 Löbl, Ivan; Smetana, Ales (2007). Catalogue of Palaearctic Coleoptera - Volume 4 - Elateroidea - Derodontoidea - Bostrichoidea - Lymexyloidea - Cleroidea - Cucujoidea. Stenstrup (Denmark): Apollo Books. pp. 1–935. https://brill.com/display/title/23984. 
  5. ↑ 5.0 5.1 5.2 "Agriotes Eschscholtz, 1829". https://www.gbif.org/species/1162125. 
  6. ↑ 6.0 6.1 6.2 6.3 Schwarz, Otto (1891). "Revision der paläarktischen Arten der Elateriden-Gattung Agriotes Eschsch. [Revision of the Palaearctic species of the genus Agriotes Eschsch., Elateridae"]. Deutsche entomologische Zeitschrift 1891 (I): 81–114. https://www.zobodat.at/pdf/Deutsche-Ent-Zeitschrift_1891_0081-0114.pdf. 
  7. ↑ 7.0 7.1 7.2 "Unterfamilie Agriotinae". 24 September 2020. https://coleonet.de/coleo/texte/agriotinae.htm. 
  8. ↑ 8.0 8.1 8.2 8.3 "Gattung Agriotes Eschsch.". 16 September 2023. https://coleonet.de/coleo/texte/agriotes.htm. 
  9. ↑ 9.0 9.1 Furlan, Lorenzo (2014). "IPM thresholds for Agriotes wireworm species in maize in Southern Europe". Journal of Pest Science 87 (4): 609–617. doi:10.1007/s10340-014-0583-5. PMID 25395918. Bibcode: 2014JPesS..87..609F. 
  10. ↑ 10.0 10.1 Kozina, A.; Lemic, D.; Bazok, R.; Mikac, K.M.; Mclean, C.M.; Ivezić, M.; Igrc Barčić, J. (2015). "Climatic, edaphic factors and cropping history help predict click beetle (Coleoptera: Elateridae) (Agriotes spp .) abundance". Journal of Insect Science 15 (1). doi:10.1093/jisesa/iev079. 
  11. ↑ Vernon, Bob; LaGasa, Eric; Philip, Hugh (2001). "Geographic and temporal distribution of Agriotes obscurus and A. lineatus (Coleoptera: Elateridae) in British Columbia and Washington as determined by pheromone trap surveys.". Journal of the Entomological Society of British Columbia 98: 257–265. https://www.biodiversitylibrary.org/partpdf/213947. 
  12. ↑ Singleton, Kendal; van Herk, Willem G.; Saguez, Julien; Scott, Ian; Gries, Regine; Gries, Gerhard (2023). "Sex pheromone of Nearctic Agriotes mancus and its similarity to that of three Palearctic Agriotes invasive in North America". Agricultural and Forest Entomology 25 (3): 468–476. doi:10.1111/afe.12568. 
  13. ↑ Byers, R.A.; Barker, G.M. (2000). "Soil dwelling macro-invertebrates in intensively grazed dairy pastures in Pennsylvania, New York and Vermont". Grass and Forage Science 55 (3): 253–270. doi:10.1046/j.1365-2494.2000.00223.x. 
  14. ↑ Furlan, Lorenzo; Bonetto, Christian; Patalano, Giampiero; Lazzeri, Luca (2004). "Potential of biocidal meals to control wireworm populations". Agroindustria 3 (3): 313–316. 
  15. ↑ Chaton, P.F.; Lempérière, G.; Tissut, M.; Ravanel, P. (2008). "Agriotes larvae (Coleoptera: Elateridae): A trial of seed coating to control larval populations with the insecticide fipronil". Pesticide Biochemistry and Physiology 90 (2): 97–105. doi:10.1016/j.pestbp.2007.09.001. 
  16. ↑ 16.0 16.1 Sufian, Muhammad (2013). Biology, monitoring and management of economically important wireworm species (Coleoptera: Elateridae) in organic farming (PhD thesis). Bonn: Rheinischen Friedrich-Wilhelms-Universität. Retrieved 24 September 2026.
  17. ↑ Rawlins, W.A. (1940). "Biology and control of the wheat wireworm, Agriotes mancus Say". Bulletin of the Cornell University Agricultural Experiment Station 738: 1–30. https://ecommons.cornell.edu/bitstream/1813/30591/2/1940%20CUAES%20Bull%20738.pdf. 
  18. ↑ 18.0 18.1 Staudacher, K.; Pitterl, P.; Furlan, L.; Cate, P.C.; Traugott, M. (2011). "PCR-based species identification of Agriotes larvae". Bulletin of Entomological Research 101 (2): 201–210. doi:10.1017/S0007485310000337. PMID 21034519. 
  19. ↑ "Agriotes sputator (Linnaeus, 1758)". https://www.gbif.org/species/5878766. 
  20. ↑ Zapata de la Vega, José Luis; Sánchez-Ruiz, Antonio (2012). "Catálogo actualizado de los Elatéridos de la Península Ibérica e Islas Baleares (Coleoptera: Elateridae)". Arquivos Entomolóxicos 6: 115–271. https://www.aegaweb.com/arquivos_entomoloxicos/ae06_2012_zapata_de_la_vega_sanchez_ruiz_catalogo_elateridae_pen_iberica_baleares.pdf. 
  21. ↑ Schwarz, Otto (1896). "Neue Elateriden aus Afrika [New Elateridae from Africa"]. Deutsche entomologische Zeitschrift 40 (1–2): 93–96. https://www.biodiversitylibrary.org/item/103427#page/102/mode/1up. 
  22. ↑ Gülperçin, Nilay; Tezcan, Serdar (2025). "Click beetles: An updated Checklist from Türkiye". Munis Entomology & Zoology 20 (supplement): 4110–4141. https://www.munisentzool.org/Issue/abstract/click-beetles-an-updated-checklist-from-turkiye_14560. 
  23. ↑ 23.0 23.1 Platia, Giuseppe (2007). "Contribution to the knowledge of the Agriotini of China. Genera Agriotes Eschscholtz, Ectinus Eschscholtz, Tinecus Fleutiaux and Rainerus gen. n. (Coleoptera, Elateridae, Agriotini)". Boletín Sociedad Entomológica Aragonesa 41: 7–42. 
  24. ↑ Fleutiaux, E. (1918). "Nouvelles contributions a la faune de l 'Indo-chine française (Coleoptera Serricornia) [New contributions to the fauna of French Indochina (Coleoptera Serricornia)"]. Annales de la Société Entomologique de France 87: 175–278 [266]. doi:10.1080/21686351.1918.12279609. https://www.biodiversitylibrary.org/item/36393#page/284/mode/1up. 
  25. ↑ Fleutiaux, E. (1939). "Les Élatérides de l'Indochine française: Septième Partie, XVII. Subfam. Agriotinae [The Elateridae of French Indochina: 7th part, XVII. Subfam. Agriotinae"]. Annales de la Société Entomologique de France 108 (3–4): 121–148. doi:10.1080/21686351.1939.12278988. https://doi.org/10.1080/21686351.1939.12278988. 
  26. ↑ Platia, Giuseppe (2021). "New species and geonemic records of the genera Agonischius Candeze, Vuilletus Fleutiaux, Agriotes Eschscholtz, Ectinus Eschscholtz, Tinecus Fleutiaux, Rainerus Platia and Chatanayus Fleutiaux from China, South East Asia, Indonesia and Malaysia (Coleoptera, Elateridae, Elaterini and Agriotini)". Boletin de la Sociedad Entomologica Aragonesa 68: 335–354. https://descargas.sea-entomologia.org/Boletin68/335354BSEA68.pdf. 
  27. ↑ Blanchard, Émile (1853). Voyage au Pôle Sud et dans l'Océanie sur les corvettes l'Astrolabe et la Zélée, execute par ordre du roi pendant les années 1837-1838-1839-1840, sous le commandement de M. J. Dumont-d'Urville – Zoologie – Tome quatrième – Description des insects [Voyage to the South Pole and in Oceania with the corvettes Astrolabe and Zélée, on the order of the king in the years 1837-1840 and under the command of J. Dumont-d'Urville – Zoology – Vol. 4 – Description of insects]. Paris: Gide et J. Baudry. pp. 1–422 [90]. https://www.biodiversitylibrary.org/item/254562#page/102/mode/1up. 
  28. ↑ 28.0 28.1 Candèze, E. (1863). Monographie des Élatérides [Monograph of the Elateridae]. 4. Liége (Belgium): H. Dessain. pp. 1–534. https://www.biodiversitylibrary.org/item/36954#page/368/mode/1up. 
  29. ↑ Champion, George C. (1896). Biologia Centrali-Americana – Insecta – Coleoptera [The biology of Central America – Insecta – Coleoptera]. III Part I. London: R.H. Porter. pp. 511–533 [Agriotes]. https://www.biodiversitylibrary.org/item/14604#page/533/mode/1up. 
  30. ↑ LeConte, John L. (1884). "Short studies of North American Coleoptera (No. 2)". Transactions of the American Entomological Society 12 (3–4): 1–31. https://www.biodiversitylibrary.org/page/7481898#page/11/mode/1up. 
  31. ↑ Candèze, [Ernest Charles Auguste] (1878). "Élatérides nouveaux, cinquième partie [New Elateridae, 5th part"]. Annales de la Société Entomologique de Belgique 21: clxxxix–cxcix. https://www.biodiversitylibrary.org/item/46150#page/393/mode/1up. 
  32. ↑ Candèze, Ernest (1874). "Révision de la Monographie de Élatérides - Premier fascicule [Revision of the monograph of Elateridae – first part"]. Mémoires de la Société Royale des Sciences de Liége – 2e sér. 4 (1): 1–218 [154]. https://www.biodiversitylibrary.org/item/87670#page/200/mode/1up. 
  33. ↑ Candèze, Ernest (1881). "Élatérides nouveaux, Troisième fascicule [New Elateridae, 3rd part"]. Mémoires de la Société Royale des Sciences de Liège. 2e série 9 (1): 1–117 [20]. https://www.biodiversitylibrary.org/item/204142#page/54/mode/1up. 
  34. ↑ Fairmaire, Léon (1883). "Coléoptères de Magellan et de Santa Cruz [The Coleoptera of the provinces of Magellan and Santa Cruz"]. Annales de la Société Entomologique de France 3, 6th series (4): 483–506 [492]. https://www.biodiversitylibrary.org/item/101301#page/496/mode/1up. 
  35. ↑ Schwarz, Otto (1904). "Neue Elateriden aus Süd-Amerika [New Elateridae from South America"]. Deutsche Entomologische Zeitschrift 1904: 49–80 [80]. https://www.biodiversitylibrary.org/item/103303#page/86/mode/1up. 
  36. ↑ Fleutiaux, Ed. (1907). "Révision des Elateridae du Chili [Revision of the Elateridae of Chile"]. Revista Chilena de Historia Natural 11: 160–232. https://archive.org/details/revistachilenad111907muse/page/160/mode/2up. 
  37. ↑ Fleutiaux, E. (1942). "Sur certains Hémirhipides (Col. Elateridae) [About certain Hemirhipini (Col. Elateridae)]". Annales de la Société Entomologique de France 111: 91–116 [112]. doi:10.1080/21686351.1942.12279003. 
  38. ↑ 38.0 38.1 Douglas, Hume B.; Etzler, Frank E.; Johnson, Paul J.; Hammond, H.E. James (2024). "Matrix-based key to the click beetle genera of Canada and USA with a summary of habitat use (Coleoptera, Elateridae)". ZooKeys 1200: 75–144. 
  39. ↑ Furlan, Lorenzo; Benvegnù, Isadora; Bilò, María Fabiana; Lehmhus, Jörn; Ruzzier, Enrico (2021). "Species identification of wireworms (Agriotes spp.; Coleoptera: Elateridae) of agricultural importance in Europe: A new "horizontal identification table"". Insects 12 (6). doi:10.3390/insects12060534. PMID 34201325. 
  40. ↑ Rashed, Arash; van Herk, Willem G. (2024). "Pest elaterids of North America: New insights and opportunities for management". Annual Review of Entomology 69: 1–20. doi:10.1146/annurev-ento-120220-123249. 
  41. ↑ 41.0 41.1 Paluch, Maximilian (2021). Biological control of wireworms (Agriotes spp.) in potato cultivation using the entomopathogenic fungus Metarhizium brunneum: Factors that influence the effectiveness of mycoinsecticide formulations (PDF) (PhD thesis). Darmstadt: Technische Universität Darmstadt. Retrieved 1 October 2025.
  42. ↑ Tóth, Miklós; Furlan, Lorenzo; Yatsynin, Venyamin G.; Ujváry, István; Szarukán, István; Imrei, Zoltán; Tolasch, Till; Francke, Wittko et al. (2003). "Identification of pheromones and optimization of bait composition for click beetle pests (Coleoptera: Elateridae) in Central and Western Europe". Pest Management Science 59 (4): 417–425. doi:10.1002/ps.629. PMID 12701702. Bibcode: 2003PMSci..59..417T. 
  43. ↑ 43.0 43.1 van Herk, Willem G.; Kabaluk, J. Todd; Vernon, Robert S.; Richard, Romain; Benz, Philipp (2023). "Disorientation of male Agriotes click beetles in the presence of granulate pheromones - a case for mating disruption". Arthropod-Plant Interactions 17 (4): 485–506. doi:10.1007/s11829-023-09988-3. Bibcode: 2023APInt..17..485V. 
  44. ↑ Barsics, Fanny; Haubruge, Eric; Verheggen, François J. (2013). "Wireworms' management: An overview of the existing methods, with particular regards to Agriotes spp. (Coleoptera: Elateridae)". Insects 4 (1): 117–152. doi:10.3390/insects4010117. 
  45. ↑ 45.0 45.1 Furlan, Lorenzo; Kreutzweiser, David (2015). "Alternatives to neonicotinoid insecticides for pest control: case studies in agriculture and forestry". Environmental Science and Pollution Research 22 (1): 135–147. doi:10.1007/s11356-014-3628-7. PMID 25273517. Bibcode: 2015ESPR...22..135F. 
  46. ↑ Johnson, Scott N.; Anderson, Eric A.; Dawson, Gregory; Griffiths, D. Wynne (2008). "Varietal susceptibility of potatoes to wireworm herbivory". Agricultural and Forest Entomology 10 (2): 167–174. doi:10.1111/j.1461-9563.2008.00372.x. 
  47. ↑ Furlan, Lorenzo; Vasileiadis, Vasileios P.; Chiarini, Francesca; Huiting, Hilfred; Leskovšek, Robert; Razinger, Jaka; Holb, Imre J.; Sartori, Erica et al. (2017). "Risk assessment of soil-pest damage to grain maize in Europe within the framework of Integrated Pest Management". Crop Protection 97: 52–59. doi:10.1016/j.cropro.2016.11.029. Bibcode: 2017CrPro..97...52F. 
  48. ↑ Eschscholtz, Johann Friedrich (1829). "Eintheilung der Elateriden in Gattungen [Division of the Elateridae into genera"]. Entomologisches Archiv 2: 31–35. https://species.wikimedia.org/wiki/Template:Eschscholtz,_1829. 
  49. ↑ Dillwyn, L.W. (1829). Memoranda relating to coleopterous insects, found in the neighbourhood of Swansea. Swansea: W.C. Murray & D. Rees. pp. 1-75. https://books.google.de/books?id=8KZgAAAAcAAJ&printsec=frontcover&redir_esc=y#v=onepage&q&f=false. 
  50. ↑ Stephens, James Francis (1830). Illustrations of British Entomology. Mandibulata, vol. III. London: Baldwin and Cradock. pp. 1–374 [247]. https://www.biodiversitylibrary.org/item/97135#page/5/mode/1up. 
  51. ↑ "Merriam Webster Dictionary". https://www.merriam-webster.com/dictionary/Agriotes. 
  52. ↑ Becker, Edward C. (1956). "Revision of the Nearctic species of Agriotes (Coleoptera: Elateridae)". The Memoirs of the Entomological Society of Canada 88 (S1): 5–101. doi:10.4039/entm8801fv. 
  53. ↑ Telfer, Mark G.. "Joy's keys to Elateridae, Eucnemidae and Throscidae". https://quelestcetanimal-lagalerie.com/wp-content/uploads/2016/11/Joy%E2%80%99s-keys-to-Elateridae-Eucnemidae-and-Throscidae.pdf. 

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Wikidata ☰ Q1442513 entry



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