Oryctes

This article is about the genus of beetles. For the genus of plants, see Oryctes (plant).

Oryctes
Oryctes sp.
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Polyphaga
Infraorder: Scarabaeiformia
Superfamily: Scarabaeoidea
Family: Scarabaeidae
Subfamily: Dynastinae
Tribe: Oryctini
Genus: Oryctes
Hellwig, 1798[1][2]

Oryctes[1] is the economically most important genus of rhinoceros beetles in the subfamily Dynastinae (family: Scarabaeidae) and includes serious pests of palm trees.[3] A total of 47 species have been assigned to the genus, including 2 fossil ones. The extant species are widely distributed in Africa, as well as in Europe, Asia and the Pacific. No established populations are known from the Americas.[4][5] The species are typically between 30 and 60 mm long, dark brown to black, with a robust body and a strong cuticle. They can be distinguished from other genera in the tribe Oryctini by the structure of the pronotum, the horn on the head, the mouthparts and other features.[4][6]

Type species: Scarabaeus nasicornis Linnaeus, 1758, a synonym of Oryctes nasicornis[7]

Distribution

Distribution ranges of the species in the genus Oryctes (blue outline) and that of Oryctes nasicornis, the type species (brown outline)

Most of the extant species of Oryctes are found in Africa with only 8 species in Asia and the Pacific, 3 in the Near East and 1 in Europe.[5] The 33 African species are mainly distributed in sub-Saharan countries, with 18 of them found in Madagascar and surrounding islands like Réunion, Mayotte, Mauritius, the Comoros and the Island of Rodrigues. Eight species (Oryctes amberiensis, O. anguliceps, O. augias, O. clypealis, O. colonicus, O. dollei, O. politus and O. ranavalo) are endemic to Madagascar and occur nowhere else.[5]

O. nasicornis is the only European species. It is widespread in most European countries and has been also reported from some of the neighbouring regions like parts of northern Africa, the Near East and Turkey.[5] Two species (Oryctes agamemnon and O. elegans) are common in the Near East region. A third one, Oryctes richteri, has been reported only from Iran. Oryctes rhinoceros is widespread in southern Asia and the Pacific. Other Asia/Pacific species include Oryctes ata in Turkmenistan, O. centaurus in Papua New Guinea, O. forceps in northern India ("British Bootan, Padong"), O. gnu (Malaysia, Indonesia, Thailand, the Philippines, and southern China), O. heros in Timor, O. hisamatsui in Japan, and O. nudicauda in Myanmar.[4][8][5] No established populations are known from the Americas. Two extinct species, described from German deposits, are also sometimes assigned to Oryctes.[5]

Description

Frontal view of Oryctes nasicornis male with some characteristic features of the genus Oryctes labelled
Tip of abdomen of female Oryctes rhinoceros with arrow pointing at stridulation area

The species of Oryctes can be distinguished from other genera in the subfamily Dynastinae and the tribe Oryctini by a combination of different morphological features. These include:[9][4][6][10]

  • Medium-sized to large rhinoceros beetles, typically between 30 and 60 mm long, and dark reddish brown to black.
  • The head has a single horn in males. Females have a short horn or a knob.
  • The antennae have 10 segments with the segments of the terminal club having a similar length in both males and females.
  • The clypeus often has a central notch which can be quite deep, the clypeus may be broad.
  • The mandibles are broad without an outer tooth, their ends are rounded or obliquely cut off.
  • The pronotum of males has a large central depression (concavity) usually covering more than half the pronotum, typically with two knobs or humps (but not horns) near the middle of the hind margin of the concavity. Some species have only one knob and Oryctes nasicornis has three. The morphology of the sides of the pronotum is also characteristic. Females have a smaller and shallower depression with no or small knobs at the hind margin.
  • The dorsal part of the pre-last abdominal segment, the propygidium, has a stridulatory area, both in males and females. Adults can communicate by rubbing the tips of their elytra over it.

Within a given species, there can be a considerable variation in size, length of the horn and structure of the pronotum. In some species, such variation may be due to the suitability of the breeding substrate. However, in the type species, Oryctes nasicornis, such variation is often genetic and is linked to different subspecies.[9][4]

The subgenera and individual species of Oryctes are characterized, among others by:[4]

  • the length of the adult beetle,
  • the size and shape of the horn,
  • the structure of the clypeus,
  • the detailed structure of the pronotum,
  • the teeth on the foretibia,
  • the teeth on the tip of the hind tibia,
  • the shape of the elytra,
  • the presence/arrangement of pits or grooves on the elytra.

Biology

Immature stages of Oryctes nasicornis

Species of Oryctes typically breed in decaying wood like rotting tree trunks or other decaying organic matter. While a given species might have a preference for a certain type of wood, there seems to be some flexibility. Females of several species readily lay eggs in old sawdust heaps near sawmills or compost heaps in gardens. For example, decaying coconut trunks are the preferred and traditional breeding sites for Oryctes rhinoceros in Asia and decaying oak bark and wood for Oryctes nasicornis in Europe, but both species often breed in old sawdust heaps, compost heaps and other decaying organic matter. In some species, the larvae can also feed on the living tissue of tree trunks and roots. The immature stages include the egg stage, three larval stages and the pupal stage. The development from egg to adult may be completed in about half a year, but can also extend over several years.[11][12][13][14][15][16]

Oryctes adults are nocturnal and are often attracted to light.[17] Except for the pest species, little else is known about the adult behaviour of many species, including their feeding habits. Adult beetles from the subfamily Dynastinae typically wound the bark of tree trunks (bark-carving) and feed on the sap which is excreted from the damaged trunk. Such a behaviour is also suspected for some species of Oryctes, while others might not feed at all in the adult stage.[18][16] As described in the next section, the adults of pest species feed by boring into living palms.

Agricultural pests

Coconut palm in Hawaii, heavily damaged by Oryctes rhinoceros attacks.

A number of Oryctes species cause damage to palm trees and are regarded as agricultural pests.[19][6] The four species discussed in the following paragraphs are the most destructive:

Adult beetles of Oryctes rhinoceros and Oryctes monoceros attack coconut and oil palms in the Asia/Pacific region, and in Africa respectively. The beetle bore into the centre of the palm crown, starting at one of the middle leaf axils. They then feed on the very young, still developing leaves (fronds), usually cutting off a significant part of the leaflets or parts of the whole fronds. Triangular cuts on the fronds are typical signs of Oryctes attacks. Damaged palms have a reduced nut production and young palms often die after heavy attacks. The larval stages of both species do not cause any damage.[11][12]

Oryctes elegans and Oryctes agamemnon are pests of date palms in the Near East and in northern Africa. In both species the larval as well as the adult stage can cause significant damage. The adults of Oryctes elegans bore into the stalks of the date bunches, often causing the bunches to break off. They may also bore into the base of the fronds, causing these to break off. The larvae of O. elegans live in the palm crown and at the base of the palms feeding on dead old fronds and other decaying matter but may also damage live tissue. For example, they may invade and damage the top part of the trunk or feed on the roots and the new shoots at the base of the palm.[20][14][21]

Oryctes agamemnon has a similar biology, but in this species, the larval stages cause most of the damage. The larvae can be very destructive when they destroy the respiratory roots near the base of the date palm which weakens the tree and can cause the palm to fall down after heavy attacks. Damage to the new shoots at the palm base which are used for planting can be also severe. Damage by larvae is particularly serious in Tunisia and Algeria which were invaded by O. agamemnon starting in the 1990s. In the Near East, adults of O. agamemnon also infest the stalks of the fruit bunches like O. elegans. However, such damage was not observed in Tunisia and Algeria.[22][14][23]

Taxonomy and species

For many years, the authorship of the genus Oryctes has been cited as "Illiger, 1798" who described the genus in his book "Verzeichniss der Käfer Preussens" [Catalogue of the beetles of Prussia].[24] However, the name was first used by Hellwig in a note a few months earlier who wrote that he initiated the work on the beetles of Prussia, but did not have time to complete it and handed it over to Illiger.[1] Illiger worked under him for several years and the book was apparently a collaborative effort by both authors. However, Hellwig is now regarded as the sole author of Oryctes since he made the name first available and mentioned the names of several species (like rhinoceros or nasicornis) that are part of the genus.[2]

In 1888, Paul Oppenheim introduced the genus Oryctites when he described the fossil species Oryctites fossilis and Oryctites is sometimes cited as a synonym of Oryctes.[25][5] However, the genus name Oryctites has been considered to be taxonomically invalid and there are doubts if the species fossilis belongs to Oryctes.[26]

A number of subgenera have been described under the genus Oryctes and the following species are currently (2025) recognized:[4][5][27]

  1. Oryctes agamemnon Burmeister, 1847 - Arabian rhinoceros beetle[28]
  2. Oryctes amberiensis Sternberg, 1910[29]
  3. Oryctes anguliceps Fairmaire, 1901[30]
  4. Oryctes ata Semenov & Medvedev, 1932[31]
  5. Oryctes augias (Olivier, 1789)[32]
  6. Oryctes boas (Fabricius, 1775)[33][34]
  7. Oryctes borbonicus Dechambre, 1982[35]
  8. Oryctes capucinus Arrow, 1937[36]
  9. Oryctes centaurus Sternberg, 1910[29]
  10. Oryctes cherlonneixi Dechambre, 1996[37]
  11. Oryctes chevrolatii Guérin-Méneville, 1844[38]
  12. Oryctes clypealis Fairmaire, 1897[39]
  13. Oryctes colonicus Coquerel, 1852[40]
  14. Oryctes comoriensis Fairmaire, 1893[41]
  15. Oryctes congonis Endrödi, 1969[42]
  16. Oryctes curvicornis Sternberg, 1910[29]
  17. Oryctes dollei Fairmaire, 1897[43]
  18. Oryctes elegans Prell, 1914 - date palm fruit stalk borer[44]
  19. Oryctes erebus Burmeister, 1847[28]
  20. Oryctes forceps Dechambre, 1980[8]
  21. †Oryctes fossilis (Oppenheim, 1888)[25]
  22. Oryctes gigas Castelnau, 1840[45]
  23. Oryctes gnu Mohnike, 1874 - Malaysian rhinoceros beetle[46]
  24. Oryctes gracilis Prell, 1934[47]
  25. Oryctes heros Endrödi, 1973[4]
  26. Oryctes hisamatsui Nagai, 2002[48]
  27. Oryctes latecavatus Fairmaire, 1891[49]
  28. Oryctes mayottensis Dechambre, 1982[35]
  29. Oryctes minor Waterhouse, 1876[50]
  30. Oryctes monardi Beck, 1942[51]
  31. Oryctes monoceros (Olivier, 1789) - African rhinoceros beetle[32]
  32. Oryctes nasicornis (Linnaeus, 1758) - type species (as Scarabaeus nasicornis L.: the European rhinoceros beetle)[52]
  33. Oryctes nudicauda Arrow, 1910[53]
  34. Oryctes ohausi Minck, 1913[54]
  35. Oryctes owariensis (Palisot de Beauvois, 1806)[55]
  36. †Oryctes pluto Weyenbergh, 1869[56]
  37. Oryctes politus Fairmaire, 1901[30]
  38. Oryctes prolixus Wollaston, 1864[57]
  39. Oryctes pyrrhus Burmeister, 1847[28]
  40. Oryctes ranavalo Coquerel, 1852[40]
  41. Oryctes rhinoceros (Linnaeus, 1758) - coconut rhinoceros beetle, Asiatic rhinoceros beetle[52]
  42. Oryctes richteri Petrovitz, 1958[58]
  43. Oryctes sahariensis Bruneau de Miré, 1960[59]
  44. Oryctes simiar Coquerel, 1852[40]
  45. Oryctes sjoestedti Kolbe, 1905[60]
  46. Oryctes tarandus (Olivier, 1789)[32]
  47. Oryctes vicinus Gahan, 1900[61]

References

  1. ^ a b c Hellwig, J.C.L. (1798). "II. Ankündigungen neuer Bücher". Intelligenzblatt der allgemeinen Literatur-Zeitung. 1798 (13): 100–101.
  2. ^ a b Alonso-Zarazaga, Miguel Angel; Krell, Frank-Thorsten (2011). "Change of authorship of Aphodius and Oryctes to Hellwig, 1798 (Insecta: Coleoptera: Scarabaeidae)". Zootaxa. 3060: 67–68. doi:10.11646/zootaxa.3060.1.5.
  3. ^ Royals, Hanna R.; Gilligan, Todd M.; Brodel, Charles F. "Coconut Rhinoceros Beetles - Oryctes spp" (PDF). USDA-APHIS-PPQ National Identification Service (NIS). Retrieved 24 December 2024.
  4. ^ a b c d e f g h Endrödi, S. (1973). "Monographie der Dynastinae. 5. Tribus: Oryctini (Coleoptera, Lamellicornia, Melolonthidae)" (PDF). Entomologische Arbeiten aus dem Museum Georg Frey. 24: 1–87.
  5. ^ a b c d e f g h "Oryctes Hellwig, 1798". GBIF - Global Biodiversity Information Facility. Retrieved 9 January 2025.
  6. ^ a b c "Oryctes Illiger, 1798". An Early Screening Tool for Exotic Scarab Pests to Australia. Retrieved 24 December 2024.
  7. ^ Latreille, P.A. (Pierre André) (1810). Considérations générales sur l’ordre naturel des animaux composant les classes des crustacés, des arachnides, et des insectes. Paris: F. Schoell. pp. 1–444 [194, 428].
  8. ^ a b Dechambre, Roger-Paul (1980). "Description de nouveaux Oryctes [Col. Scarabaeoidea Dynastidae]". Bulletin de la Société entomologique de France. 85 (1–2): 20–22.
  9. ^ a b Endrödi, Sebö (1938). "Die paläarktischen Rassenkreise des Genus Oryctes (Ill.)" (PDF). Archiv für Naturgeschichte (NF). 7: 53–96.
  10. ^ Royals, Hanna R.; Gilligan, Todd M.; Brodel, Charles F. "Coconut rhinoceros beetles - Oryctes spp" (PDF). USDA-APHIS-PPQ National Identification Services (NIS). Retrieved 4 January 2025.
  11. ^ a b Catley, A. (1969). "The coconut rhinoceros beetle Oryctes rhinoceros (L) [Coleoptera: Scarabaeidae: Dynastinae]". PANS. 15 (1): 18–30. doi:10.1080/04345546909415075.
  12. ^ a b Bedford, G.O. (1980). "Biology, ecology, and control of palm rhinoceros beetles". Annual Review of Entomology. 25: 309–339. doi:10.1146/annurev.en.25.010180.001521.
  13. ^ Ansorge, Horst J. (2003). "Massenvorkommen von Nashornkäfern Oryctes nasicornis (Linné, 1758) in Gerberlohe im mittelalterlichen Stralsunder Rathaus (Col., Scarabaeidae)" (PDF). Entomologische Nachrichten und Berichte. 47 (3–4): 153–156.
  14. ^ a b c Bedford, Geoffrey O.; Al-Deeb, Mohammad Ali; Khalaf, Mohammed Zaidan; Mohammadpour, Kazem; Soltani, Rasmi (2015). "Chapter 5: Dynastid Beetle Pests". In Wakil, Waqas; Romeno Faleiro, Jose; Miller, Thomas A. (eds.). Sustainable Pest Management in Date Palm: Current Status and Emerging Challenges (PDF). Springer Cham. pp. 73–108. doi:10.1007/978-3-319-24397-9_5.
  15. ^ "Nashornkäfer (Oryctes nasicornis (Linnaeus, 1758))". Insekten Sachsen. Retrieved 4 January 2025.
  16. ^ a b Kamber, Lea. "Nashornkäfer (Oryctes nasicornis)" (PDF). Kanton Bern. Retrieved 4 January 2025.
  17. ^ Ehsine, M’hammed; Belkadhi, Mohamed Sadok; Chaieb, Mohammed (2014). "Seasonal and nocturnal activities of the rhinoceros borer (Coleoptera: Scarabaeidae) in the north Saharan oases ecosystems". Journal of Insect Science. 14 (1) 256. doi:10.1093/jisesa/ieu118.
  18. ^ Ichiishi, Wataru; Shimada, Shinpei; Motobayashi, Takashi; Abe, Hiroaki (2019). "Completely engaged three-dimensional mandibular gear-like structures in the adult horned beetles: reconsideration of bark-carving behaviors (Coleoptera, Scarabaeidae, Dynastinae)". ZooKeys. 813: 89–110. doi:10.3897/zookeys.813.29236.
  19. ^ Hurpin, B.; Fresneau, M. (1969). "Contribution a l'étude de Oryctes elegans [Col. Dynastidae]". Annales de la Société entomologique de France (N.S.). 5 (3): 595–612. doi:10.1080/21686351.1969.12277916.
  20. ^ Khalaf, M.Z.; Shbar, A.K.; Al-Seria, M.H.; Sami, R.A.; Naher, F.H. (2011). "Some aspects of biology and control methods of fruit stalk borer Oryctes elegans Prell (Coleoptera: Scarabaeidae)". Journal of Agricultural Science and Technology. A1: 142–147.
  21. ^ Atwa, Atwa A. (2018). "Biological control potential of two Steinernematid species against the date fruit stalk borer (Oryctes elegans Prell, Coleoptera: Scarabaeidae)". Journal of Insect Science. 18 (3) 26. doi:10.1093/jisesa/iey060.
  22. ^ Soltani, Rasmi; Ikbel, Chaieb; Ben Hamouda, Med Habib (2008). "Descriptive study of damage caused by the rhinoceros beetle, Oryctes agamemnon, and its influence on date palm oases of Rjim Maatoug, Tunisia". Journal of Insect Science. 8 (1) 57. doi:10.1673/031.008.5701.
  23. ^ Chouia, Abdelouahed; Guerfi, Zoubir; Sadine, Salah Eddine (2018). "Contribution to the study of a new date palm pest Oryctes agamemnon in the palm groves of El-Oued, Algeria" (PDF). Tunesian Journal of Plant Protection. 13 (SI): 159–170.
  24. ^ Illiger, Johann Karl Wilhelm (1798). Verzeichniss der Käfer Preussens. Halle, Germany: Verlag Gebauer. pp. 1–510.
  25. ^ a b Oppenheim, Paul (1888). "Die Insectenwelt des lithographischen Schiefers in Bayern" (PDF). Palaeontographica. 34: 215–247 [238].
  26. ^ Krell, Frank-Thorsten (2000). "The fossil record of Mesozoic and Tertiary Scarabaeoidea (Coleoptera : Polyphaga)". Invertebrate Taxonomy. 14: 871–905. doi:10.1071/IT00031.
  27. ^ "Oryctini Mulsant, 1842". BioLib.cz. Retrieved 18 November 2021.
  28. ^ a b c Burmeister, Herrmann (1847). Handbuch der Entomologie. Volume 5 - Coleoptera Lamellicornia, Xylophila et Pectinicornia (PDF). Berlin, Germany: Theod. Chr. Fr. Enslin. pp. 1–584 [197, 201].
  29. ^ a b c Sternberg, Chr. (1910). "Neue Dynastiden-Arten" (PDF). Annales De La Société Entomologique De Belgique. Bruxelles. 54: 91–102.
  30. ^ a b Fairmaire, L. (1901). "Description de quelques Coléoptères receuillis par M. le docteur Decorse dans le sud de Madagascar, Plateau de L'Androy". Notes from the Leyden Museum. 23: 65–85.
  31. ^ Semenov, A.P.; Medvedev, S. (1932). "Species generis Oryctes Ill. Faunae rossicae et mesasiaticae". Annuaire Museum De Zoologique De l’Academie Des Sciences URSS. 32: 481–502.
  32. ^ a b c Olivier, [Guillaume Antoine] (1789). "3. Scarabé Scarabaeus". Entomologie, ou histoire naturelle des insectes, avec leurs caractères génériques et spécifiques, leur description, leur synonymie, et leur figure enluminée. Vol. I Coléoptères (PDF). Paris, France: Baudouin. pp. 1–190 [36–39].
  33. ^ Fabricius, Io. Christ. (1775). Systema Entomologiae. Flensburgi et Lipsiae [Flensburg and Leipzig]: Officina Libraria Kortii. pp. 1–832 [8].
  34. ^ Prell, H. (1936). "Beiträge zur Kenntnis der Dynastinen (XVI) (Col.). Die Dynastinen der Fabriciusschen Sammlung im Zoologischen Museum der Universität Kiel" (PDF). Berliner Entomologische Zeitschrift. 1936 (3–4): 179–190. doi:10.1002/mmnd.193619360303.
  35. ^ a b Dechambre, R.P. (1982). "Nouveaux Oryctes de la région malgache". Revue Française d’Entomologie (Nouvelle-Serie). 4 (4): 176–178.
  36. ^ Arrow, Gilbert J. (1937). "Systematic notes on beetles of the subfamily Dynastinae, with descriptions of a few new species in the British Museum collection". Transactions of the Entomological Society London. 86 (3): 35–57. doi:10.1111/j.1365-2311.1937.tb00246.x.
  37. ^ Dechambre, R.P. (1996). "Une nouvelle espèce d'Oryctes du Gabon". Revue Française d’Entomologie (Nouvelle-Serie). 18 (1): 17–18.
  38. ^ Guérin-Méneville, F.E. (1884). Iconographie du Règne Animal de G.Cuvier. Insectes (PDF). Paris: J.B. Baillière. pp. 1–576 [85].
  39. ^ Fairmaire, L. (1897). "Matériaux pour la faune coléoptérique de la région Malgache (4e note)". Annales De La Société Entomologique De Belgique. 41: 363–406 [380].
  40. ^ a b c Coquerel, Ch. (1852). "Observations entomologiques sur divers insectes recueillis à Madagascar". Annales De La Société Entomologique De France. 10: 359–405.
  41. ^ Fairmaire, L. (1893). "Coléoptères des îles Comores" (PDF). Annales De La Société Entomologique De Belgique. 37: 521–555 [531].
  42. ^ Endrödi, S. (1969). "Oryctes congonis n.sp. aus Afrika". Revue de Zoologie et de Botanique africains. 80 (3–4): 397–402.
  43. ^ Fairmaire, L. (1897). "Matériaux pour la faune coléoptérique de la région Malgache (3e note)". Annales De La Société Entomologique De Belgique. 41: 92–119 [108].
  44. ^ Prell, H. (1914). "Beiträge zur Kenntnis der Dynastinen X (Col.)" (PDF). Entomologische Mitteilungen. 3 (7/8): 197–226 [210].
  45. ^ Castelnau, [F.] (1840). Histoire Naturelle des Insectes Coléoptères (PDF). Paris: P. Duménil. pp. 1–563 [114].
  46. ^ Mohnike, [O.] (1874). "[No title - notes on new and rare beetles from the families Cetonidae and Dynastidae]". Verhandlungen des Naturhistorischen Vereines der preussischen Rheinlande. 31. section: Antropology, Zoology and Anatomy, pp.261–262.
  47. ^ Prell, Heinrich (1934). "Beiträge zur Kenntnis der Dynastinen (XII). Beschreibungen und Bemerkungen" (PDF). Entomologische Blätter, Krefeld. 30: 55–60.
  48. ^ Nagai, S. (2002). "A new species and a new record of the Dynastid beetle from the Ryukyu Islands, Southwest Japan". Japanese Journal of Systematic Entomology. 8 (1): 45–48.
  49. ^ Fairmaire, L. (1891). "Contribution à l'étude de la faune entomologique de S.Thomé". O Instituto: Revista Scientifica E Literária, Coimbra. 39: 112–116.
  50. ^ Waterhouse, Chas. O. (1876). "New species of Coleoptera from the Island of Rodriguez, collected by the Naturalists accompanying the Transit-of-Venus Expedition" (PDF). The Annals and Magazine of Natural History, Including Zoology, Botany and Geology. London. 4 (18): 105–121.
  51. ^ Beck, P. (1942). "Coléoptères lamellicornes d'Angola: Dynastinae". Revue Suisse de Zoologie. 49: 571–577.
  52. ^ a b Linnaeus, Carolus (1758). Systema Naturae. Stockholm, Sweden: Laurentius Salvius. pp. 1–824 [346].
  53. ^ Arrow, G.J. (1910). The Fauna of British India (Including Ceylon and Burma) Lamellicornia I. (PDF). London, U.K.: Taylor & Francis. pp. 1–322 [277].
  54. ^ Minck, Paul (1913). "Beiträge zur Kenntnis der Dynastinen. 2.Afrikanische Oryctiden" (PDF). Entomologische Rundschau. 30: 16.
  55. ^ Palisot de Beauvois, A.M.F.J. (1806). Insectes receuillis en Afrique et en Amerique, dans les royaumes d'Oware et de Benin, à Saint-Dominique et dans les Etats-Unis, pendant les années 1786-1797. Paris, France: Fain et Compagnie. pp. 1–276 [41].
  56. ^ Weyenbergh, H. (1869). "Sur les Insectes fossiles du Calcaire Lithographique de la Bavière, qui se trouvent au Musée Teyler" (PDF). Archives De Musée Teyler. 2: 1–48 [36].
  57. ^ Wollaston, T. Vernon (1864). Catalogue of the Coleopterous Insects of the Canaries in the collection of the British Museum (PDF). London, U.K.: Taylor and Francis. pp. 1–648 [202].
  58. ^ Petrovitz, Rudolf (1958). "Beitrag zur Kenntnis der Scarabaeiden-Fauna des Iran" (PDF). Stuttgarter Beiträge zur Naturkunde. 8: 1–12.
  59. ^ Bruneau de Miré, Ph. (1960). "Les insectes "rhinocéros" du Palmier Dattier (Col. Scarabaeidae, Trib. Dynastini)". Journal d'agriculture tropicale et de botanique appliquée. 7 (4–5): 241–255.
  60. ^ Kolbe, H. (1905). "Dynastiden aus Kamerun, gesammelt von Herrn Professor D:r Yngve Sjöstedt" (PDF). Arkiv För Zoologi. 2 (18): 1–20.
  61. ^ Gahan, C.J. (1900). "The Expedition to Sokotra. XI. Descriptions of the new genera and species of Coleoptera". Bulletin of the Liverpool Museum. 3: 8–13.
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