Flower chafer explained

Flower chafers are a group of scarab beetles comprising the subfamily Cetoniinae. Many species are diurnal and visit flowers for pollen and nectar, or to browse on the petals. Some species also feed on fruit. The group is also called fruit and flower chafers, flower beetles and flower scarabs. There are around 4,000 species, many of them still undescribed.

Ten tribes are presently recognized: Cetoniini, Cremastocheilini, Diplognathini, Goliathini, Gymnetini, Phaedimini, Schizorhinini, Stenotarsiini, Taenioderini, and Xiphoscelidini. The former tribes Trichiini and Valgini were elevated in rank to subfamily.[1] The tribe Gymnetini is the biggest of the American tribes, and Goliathini contains the largest species, and is mainly found in the rainforest regions of Africa.

Description

Adult flower chafers are usually brightly coloured beetles, often metallic, and somewhat flattened in shape. The insertions of the antennae are visible from above, while the mandibles and labrum are hidden by the clypeus. The elytra lack a narrow membranous margin and are truncate to expose the pygidium. The abdominal spiracles are diverging so that several lie on the abdominal sternites with at least one exposed. The fore coxae are conical and produced ventrally, while the mid coxae are transverse or only slightly oblique. The mesothoracic epimera is visible from above. The tarsi are each equipped with a pair of simple (not forked) tarsal claws of subequal size.[2] [3]

A feature possessed by adults of many flower chafers, especially Cetoniini, is lateral emargination of the elytra.[4]

Larvae are stout-bodied and very hairy with short legs. The head is partly covered by the prothorax. Each antenna has the apical segment as wide as the penultimate segment. The galea and lacinia are used to form a mala. The anal cleft is transverse. The mandible has a ventral stridulating area. The labrum is symmetrical with a deeply pigmented notch on each side of the midline.

Biology

Adult cetoniines are herbivorous, being found on flowers (from which they consume nectar and pollen), tree sap and rotting fruit. Larvae generally live and feed in decaying plant matter (including decaying wood) or soil. In captivity, cetoniine larvae will feed on soft fruit.

Many species in the tribe Cremastocheilini are known to be predaceous, feeding on hymenopteran larvae or soft-bodied nymphs of Auchenorrhyncha. Spilophorus spp. have been noted feeding on the nesting material and excrement of South African passerine birds,[5] while Spilophorus maculatus has been recorded feeding on Oxyrhachis sp. nymphs[6] and Hoplostomus fuligineus is known to feed on the brood of honey bees in South Africa and the pupae of the wasp Belonogaster petiolata. Campsiura javanica feeds on the larvae of Ropalidia montana in southern India.[7] Cremastocheilus stathamae feeds on ants of the genus Myrmecocystus.[8]

In terms of movement, adults are considered some of the best flyers among beetles. They can hover above and land on flowers or fruit. When threatened by predators, they escape by either performing a rush take off or by falling toward the ground and then flying before impact. Many cetoniines fly with their elytra closed, as their hindwings can unfold and slide out under the elytra during flight (thanks to the emargination of the elytra).

Larvae of some taxa can crawl on their backs using their tergal folds, which are covered in strong bristles. Others crawl on their legs.

Systematics and taxonomy

The following list contains the genera and subtribes in ten tribes of subfamily Cetoniinae, according to Catalogue of Life and Scarabaeidae of the World (2023).

Cetoniini

Authority: Leach, 1815

  1. Aethiessa Burmeister, 1842
  2. Anatona Burmeister, 1842
  3. Anelaphinis Kolbe, 1892
  4. Aphelinis Antoine, 1987
  5. Atrichelaphinis Kraatz, 1898
  6. Atrichiana Distant, 1911
  7. Badizoblax Thomson, 1877
  8. Centrantyx Fairmaire, 1884
  9. Cetonia Fabricius, 1775
  10. Chewia Legrand, 2004
  11. Chiloloba Burmeister, 1842
  12. Cosmesthes Kraatz, 1880
  13. Cosmiophaena Kraatz, 1898
  14. Dischista Burmeister, 1842
  15. Dolichostethus Kolbe, 1892
  16. Elaphinis Burmeister, 1842
  17. Enoplotarsus Lucas, 1859
  18. Erlangeria Preiss, 1902
  19. Gametis Burmeister, 1842
  20. Gametoides Antoine, 2005
  21. Glycosia Schoch, 1896
  22. Glycyphana Burmeister, 1842
  23. Gymnophana Arrow, 1910
  24. Hemiprotaetia Mikšić, 1963
  25. Heterocnemis Albers, 1852
  26. Heterotephraea Antoine, 2002
  27. Lorkovitschia Mikšić, 1968
  28. Marmylida Thomson, 1880
  29. Mireia Ruter, 1953
  30. Niphobleta Kraatz, 1880
  31. Pachnoda Burmeister, 1842
  32. Pachnodoides Alexis & Delpont, 2002
  33. Pachytephraea De Palma & Malec, 2020
  34. Paleopragma Thomson, 1880
  35. Paranelaphinis Antoine, 1988
  36. Paraprotaetia Moser, 1907
  37. Pararhabdotis Kraatz, 1899
  38. Parastraella Antoine, 2005
  39. Paraxeloma Holm, 1988
  40. Parelaphinis Marais & Holm, 1989
  41. Phaneresthes Kraatz, 1894
  42. Phonotaenia Kraatz, 1880
  43. Phoxomeloides Schoch, 1898
  44. Podopholis Moser, 1915
  45. Podopogonus Moser, 1917
  46. Pogonopus Arrow, 1910
  47. Polystalactica Kraatz, 1882
  48. Protaetia Burmeister, 1842
  49. Protaetiomorpha Mikšić, 1968
  50. Pseudotephraea Kraatz, 1882
  51. Reineria Mikšić, 1968
  52. Rhabdotis Burmeister, 1842
  53. Rhabdotops Krikken, 1981
  54. Rhyxiphloea Burmeister, 1842
  55. Ruteraetia Krikken, 1980
  56. Simorrhina Kraatz, 1886
  57. Somalibia Lansberge, 1882
  58. Stalagmosoma Burmeister, 1842
  59. Systellorrhina Kraatz, 1895
  60. Tephraea Burmeister, 1842
  61. Thyreogonia Reitter, 1898
  62. Trichocephala Moser, 1916
  63. Tropinota Mulsant, 1842
  64. Walsternoplus Allsopp, Jákl & Rey, 2023
  65. Xeloma Kraatz, 1881
  1. Chlorixanthe Bates, 1889
  2. Euphoria Burmeister, 1842
  1. Acrothyrea Kraatz, 1882
  2. Alleucosma Schenkling, 1921
  3. Amaurina Kolbe, 1895
  4. Analleucosma Antoine, 1989
  5. Cyrtothyrea Kolbe, 1895
  6. Discopeltis Burmeister, 1842
  7. Grammopyga Kolbe, 1895
  8. Heteralleucosma Antoine, 1989
  9. Homothyrea Kolbe, 1895
  10. Leucocelis Burmeister, 1842
  11. Lonchothyrea Kolbe, 1895
  12. Mausoleopsis Lansberge, 1882
  13. Mecaspidiellus Antoine, 1997
  14. Molynoptera Kraatz, 1897
  15. Molynopteroides Antoine, 1989
  16. Oxythyrea Mulsant, 1842
  17. Paleira Reiche, 1871
  18. Paralleucosma Antoine, 1989
  19. Phoxomela Schaum, 1844
  20. Pseudalleucosma Antoine, 1989
  21. Pseudooxythyrea Baraud, 1985

Cremastocheilini

Authority: Burmeister & Schaum, 1841

  1. Aspilochilus Rojkoff, 2013
  2. Aspilus Westwood, 1874
  3. Protochilus Krikken, 1976
  1. Arielina Rossi, 1958
  2. Astoxenus Péringuey, 1907
  3. Basilewskynia Schein, 1957
  4. Coenochilus Schaum, 1841
  1. Centrochilus Krikken, 1976
  2. Clinterocera Motschulsky, 1857
  3. Cremastocheilus Knoch, 1801
  4. Cyclidiellus Krikken, 1976
  5. Cyclidinus Westwood, 1874
  6. Cyclidius MacLeay, 1838
  7. Genuchinus Westwood, 1874
  8. Lissomelas Bates, 1889
  9. Paracyclidius Howden, 1971
  10. Platysodes Westwood, 1874
  11. Psilocnemis Burmeister, 1842
  1. Cymophorus Kirby, 1827
  2. Myrmecochilus Wasmann, 1900
  3. Rhagopteryx Burmeister, 1842
  1. Genuchus Kirby, 1825
  2. Meurguesia Ruter, 1969
  3. Problerhinus Deyrolle, 1864
  1. Goliathopsis Janson, 1881
  1. Heterogenius Moser, 1911
  2. Pseudastoxenus Bourgoin, 1921
  1. Chthonobius Burmeister, 1847
  2. Lissogenius Schaum, 1844
  1. Brachymacroma Kraatz, 1896
  2. Campsiura Hope, 1831
  3. Macromina Westwood, 1874
  4. Pseudopilinurgus Moser, 1918
  1. Nyassinus Westwood, 1879
  1. Anatonochilus Péringuey, 1907
  2. Oplostomus MacLeay, 1838
  3. Placodidus Péringuey, 1900
  4. Scaptobius Schaum, 1841
  1. Callynomes Mohnike, 1873
  2. Centrognathus Guérin-Méneville, 1840
  3. Parapilinurgus Arrow, 1910
  4. Periphanesthes Kraatz, 1880
  5. Pilinurgus Burmeister, 1842
  6. Priska Jákl, 2018
  1. Spilophorus Westwood, 1848
  1. Telochilus Krikken, 1975
  1. Lecanoderus Kolbe, 1907
  2. Trichoplus Burmeister, 1842
  1. Pseudoscaptobius Krikken, 1976
  2. Trogodes Boheman, 1857

Diplognathini

Authority: Burmeister, 1842

  1. Anoplocheilus MacLeay, 1838
  2. Anthracophora Burmeister, 1842
  3. Anthracophorides Moser, 1918
  4. Apocnosoides Antoine, 2001
  5. Charadronota Burmeister, 1842
  6. Conradtia Kolbe, 1892
  7. Diphrontis Gerstaecker, 1882
  8. Diplognatha Gory & Percheron, 1833
  9. Eriulis Burmeister, 1842
  10. Hadrodiplognatha Kraatz, 1898
  11. Heteropseudinca Valck Lucassen, 1933
  12. Lamellothyrea Krikken, 1980
  13. Metallopseudinca Valck Lucassen, 1933
  14. Niphetophora Kraatz, 1883
  15. Nselenius (Holm & Perissinotto, 2004)
  16. Odontorrhina Burmeister, 1842
  17. Parapoecilophila Hauser, 1904
  18. Phonopleurus Moser, 1919
  19. Pilinopyga Kraatz, 1888
  20. Porphyronota Burmeister, 1842
  21. Pseudinca Kraatz, 1880
  22. Stethopseudinca Valck Lucassen, 1933
  23. Tetragonorrhina Kraatz, 1896
  24. Trichostetha Burmeister, 1842
  25. Triplognatha Krikken, 1987
  26. Trymodera Gerstaecker, 1867
  27. Uloptera Burmeister, 1842

Goliathini

Authority: Latreille, 1829

  1. Dicronocephalus
  2. Platynocephalus
  1. Fornasinius Bertoloni, 1852
  2. Goliathus Lamarck, 1801
  3. Hegemus Thomson, 1881
  4. Hypselogenia Burmeister, 1840
  1. Gariep Péringuey, 1907
  2. Ichnestoma Gory & Percheron, 1833
  3. Karooida Perissinotto, 2020
  4. Mzansica Perissinotto, 2020
  1. Anagnathocera Arrow, 1922
  2. Anisorrhina Westwood, 1842
  3. Asthenorhella Westwood, 1874
  4. Asthenorhina Westwood, 1843
  5. Bietia Fairmaire, 1898
  6. Caelorrhina Hope, 1841
  7. Cheirolasia Westwood, 1842
  8. Chloresthia Fairmaire, 1905
  9. Chlorocala Kirby, 1828
  10. Chondrorrhina Kraatz, 1880
  11. Compsocephalus White, 1845
  12. Cosmiomorpha Saunders, 1852
  13. Cyphonocephalus Westwood, 1842
  14. Desfontainesia Alexis & Delpont, 1999
  15. Dicellachilus Waterhouse, 1905
  16. Dicheros Gory & Percheron, 1833
  17. Dicronorhina Hope, 1837
  18. Diphyllomorpha Hope, 1843
  19. Dymusia Burmeister, 1842
  20. Euchloropus Arrow, 1907
  21. Eudicella White, 1839
  22. Eutelesmus Waterhouse, 1880
  23. Gnathocera Kirby, 1825
  24. Gnorimimelus Kraatz, 1880
  25. Hemiheterorrhina Mikšić, 1974
  26. Herculaisia Seilliere, 1910
  27. Heterorhina Westwood, 1842
  28. Ingrisma Fairmaire, 1893
  29. Ischnoscelis Burmeister, 1842
  30. Jumnos Saunders, 1839
  31. Lansbergia Ritsema, 1888
  32. Lophorrhina Westwood, 1842
  33. Lophorrhinides Perissinotto, Clennell & Beinhundner, 2019
  34. Mawenzhena Alexis & Delpont, 2001
  35. Mecynorhina Hope, 1837
  36. Moseriana Ruter, 1965
  37. Mystroceros Burmeister, 1842
  38. Narycius Dupont, 1835
  39. Neomystroceros Alexis & Delpont, 1998
  40. Neophaedimus Lucas, 1870
  41. Neoscelis Schoch, 1897
  42. Pedinorrhina Kraatz, 1880
  43. Plaesiorrhina Westwood, 1842
  44. Petrovitzia Mikšić, 1965
  45. Priscorrhina Krikken, 1984
  46. Pseudodiceros Mikšić, 1974
  47. Pseudotorynorrhina Mikšić, 1967
  48. Ptychodesthes Kraatz, 1883
  49. Raceloma Thomson, 1877
  50. Rhamphorrhina Klug, 1855
  51. Rhinarion Ruter, 1965
  52. Rhomborhina Hope, 1837
  53. Scythropesthes Kraatz, 1880
  54. Smicorhina Westwood, 1847
  55. Spelaiorrhina Lansberge, 1886
  56. Stephanorrhina Burmeister, 1842
  57. Taurhina Burmeister, 1842
  58. Tmesorrhina Westwood, 1841
  59. Torynorrhina Arrow, 1907
  60. Trichoneptunides Legrand, 2001
  61. Trigonophorinus Pouillaude, 1913
  62. Trigonophorus Hope, 1831

Gymnetini

Authority: Kirby, 1827

  1. Blaesia Burmeister, 1842
  2. Halffterinetis Morón & Nogueira, 2007
  1. Allorrhina Burmeister, 1842
  2. Amazula Kraatz, 1882
  3. Amithao Thomson, 1878
  4. Argyripa Thomson, 1878
  5. Astroscara Schürhoff, 1937
  6. Badelina Thomson, 1880
  7. Balsameda Thomson, 1880
  8. Chiriquibia Bates, 1889
  9. Clinteria Burmeister, 1842
  10. Clinteroides Schoch, 1898
  11. Cotinis Burmeister, 1842 - (Green June Beetles)
  12. Desicasta Thomson, 1878
  13. Guatemalica Neervoort Van De Poll, 1886
  14. Gymnephoria Ratcliffe, 2019
  15. Gymnetina Casey, 1915
  16. Gymnetis MacLeay, 1819
  17. Hadrosticta Kraatz, 1892
  18. Heterocotinis Martinez, 1948
  19. Hologymnetis Martinez, 1949
  20. Hoplopyga Thomson, 1880
  21. Hoplopygothrix Schürhoff, 1933
  22. Howdenypa Arnaud, 1993
  23. Jansonia Schürhoff, 1937
  24. Macrocranius Schürhoff, 1935
  25. Madiana Ratcliffe & Romé, 2019
  26. Marmarina Kirby, 1827
  27. Neocorvicoana Ratcliffe & Mico, 2001
  28. Pseudoclinteria Kraatz, 1882
  29. Stethodesma Bainbridge, 1840
  30. Tiarocera Burmeister, 1842

Phaedimini

Authority: Schoch, 1894

  1. Hemiphaedimus Mikšić, 1972
  2. Phaedimus Waterhouse, 1841
  3. Philistina MacLeay, 1838
  4. Theodosia Thomson, 1880

Schizorhinini

Authority: Burmeister, 1842

  1. Agestrata Eschscholtz, 1829
  2. Ischiopsopha Gestro, 1874
  3. Lomaptera Gory & Percheron, 1833
  4. Macronota Hoffmannsegg, 1817
  5. Megaphonia Schürhoff, 1933
  6. Morokia Janson, 1905
  7. Mycterophallus Neervoort Van De Poll, 1886
  8. Thaumastopeus Kraatz, 1885
  1. Anacamptorrhina Blanchard, 1842
  2. Aphanesthes Kraatz, 1880
  3. Aurum Hutchinson & Moeseneder, 2019
  4. Axillonia Krikken, 2018
  5. Bisallardiana Antoine, 2003
  6. Chalcopharis Heller, 1902
  7. Chlorobapta Kraatz, 1880
  8. Chondropyga Kraatz, 1880
  9. Clithria Burmeister, 1842
  10. Diaphonia Newman, 1840
  11. Dichrosoma Kraatz, 1885
  12. Digenethle Thomson, 1877
  13. Dilochrosis Thomson, 1878
  14. Eupoecila Burmeister, 1842
  15. Grandaustralis Hutchinson & Moeseneder, 2013
  16. Hemichnoodes Kraatz, 1880
  17. Hemipharis Burmeister, 1842
  18. Kerowagia Delpont, 1996
  19. Lenosoma MacLeay, 1863
  20. Lyraphora Kraatz, 1880
  21. Macrotina Strand, 1934
  22. Metallesthes Kraatz, 1880
  23. Microdilochrosis Jákl, 2009
  24. Microlomaptera Kraatz, 1885
  25. Micropoecila Kraatz, 1880
  26. Navigator Moeseneder & Hutchinson, 2016
  27. Neoclithria Neervoort Van De Poll, 1886
  28. Neorrhina Thomson, 1878
  29. Octocollis Moeseneder & Hutchinson, 2012
  30. Panglaphyra Kraatz, 1880
  31. Peotoxus Krikken, 1983
  32. Poecilopharis Kraatz, 1880
  33. Pseudoclithria Neervoort Van De Poll, 1886
  34. Rigoutorum Hutchinson & Moeseneder, 2022
  35. Schizorhina Kirby, 1825
  36. Schochidia Berg, 1898
  37. Stenopisthes Moser, 1913
  38. Storeyus Hasenpusch & Moeseneder, 2010
  39. Tafaia Valck Lucassens, 1939
  40. Tapinoschema Thomson, 1880
  41. Territonia Krikken, 2018
  42. Trichaulax Kraatz, 1880

Stenotarsiini

Authority: Kraatz, 1880

  1. Anochilia Burmeister, 1842
  2. Epistalagma Fairmaire, 1880
  1. Chromoptilia Westwood, 1842
  2. Descarpentriesia Ruter, 1964
  1. Bricoptis Burmeister, 1842
  2. Coptomia Burmeister, 1842
  3. Coptomiopsis Pouillaude, 1919
  4. Eccoptomia Kraatz, 1880
  5. Euchilia Burmeister, 1842
  6. Euryomia Burmeister, 1842
  7. Heterocranus Bourgoin, 1919
  8. Hiberasta Fairmaire, 1901
  9. Hyphelithia Kraatz, 1880
  10. Liostraca Burmeister, 1842
  11. Micreuchilia Pouillaude, 1917
  12. Micropeltus Blanchard, 1842
  13. Pareuchilia Kraatz, 1880
  14. Pygora Burmeister, 1842
  15. Pyrrhopoda Kraatz, 1880
  16. Vieuella Ruter, 1964
  1. Doryscelis Burmeister, 1842
  2. Epixanthis Burmeister, 1842
  3. Hemiaspidius Krikken, 1982
  4. Pararhynchocephala Paulian, 1991
  5. Parepixanthis Kraatz, 1893
  6. Pseudepixanthis Kraatz, 1880
  7. Rhynchocephala Fairmaire, 1883
  1. Euchroea Burmeister, 1842
  1. Heterophana Burmeister, 1842
  2. Oxypelta Pouillaude, 1920
  3. Pogoniotarsus Kraatz, 1880
  4. Pogonotarsus Burmeister, 1842
  5. Zebinus Fairmaire, 1894
  1. Heterosoma Schaum, 1844
  2. Plochilia Fairmaire, 1896
  1. Bonoraella Ruter, 1978
  2. Celidota Burmeister, 1842
  3. Cyriodera Burmeister, 1842
  4. Dirrhina Burmeister, 1842
  5. Hemilia Kraatz, 1880
  6. Lucassenia Olsoufieff, 1940
  7. Melanchroea Kraatz, 1900
  8. Moriaphila Kraatz, 1880
  9. Pantolia Burmeister, 1842
  10. Tetraodorhina Blanchard, 1842
  1. Parachilia Burmeister, 1842
  1. Callipechis Burmeister, 1842
  2. Ischnotarsia Kraatz, 1880
  3. Rhadinotaenia Kraatz, 1900
  4. Stenotarsia Burmeister, 1842
  5. Vadonidella Ruter, 1973

Taenioderini

Authority: Mikšić, 1976

  1. Anocoela Moser, 1914
  2. Chalcothea Burmeister, 1842
  3. Chalcotheomima Mikšić, 1970
  4. Clerota Burmeister, 1842
  5. Glyptothea Bates, 1889
  6. Glyptotheomima Mikšić, 1976
  7. Hemichalcothea Mikšić, 1970
  8. Microchalcothea Moser, 1910
  9. Paraplectrone Mikšić, 1985
  10. Penthima Kraatz, 1892
  11. Plectrone Wallace, 1867
  12. Pseudochalcothea Ritsema, 1882
  13. Pseudochalcotheomima Mikšić, 1985
  1. Bacchusia Mikšić, 1976
  2. Bombodes Westwood, 1848
  3. Carneluttia Mikšić, 1976
  4. Coilodera Hope, 1831
  5. Costinota Schürhoff, 1933
  6. Eumacronota Mikšić, 1976
  7. Euremina Westwood, 1867
  8. Euselates Thomson, 1880
  9. Gnorimidia Lansberge, 1887
  10. Ixorida Thomson, 1880
  11. Macronotops Krikken, 1977
  12. Meroloba Thomson, 1880
  13. Pleuronota Kraatz, 1892
  14. Stenonota Fairmaire, 1889
  15. Taeniodera Burmeister, 1842
  16. Xenoloba Bates, 1889

Xiphoscelidini

Authority: Krikken, 1984

  1. Aporecolpa Lansberge, 1886
  2. Callophylla Moser, 1916
  3. Heteroclita Burmeister, 1842
  4. Ischnostomiella Krikken, 1978
  5. Meridioclita Krikken, 1982
  6. Myodermidius Bourgoin, 1920
  7. Neoclita Perissinotto, 2017
  8. Oroclita Krikken, 1982
  9. Plochiliana Ruter, 1978
  10. Protoclita Krikken, 1978
  11. Rhinocoeta Burmeister, 1842
  12. Scheinia Ruter, 1957
  13. Xiphoscelis Burmeister, 1842
  14. Xiphosceloides Holm, 1992

References

External links

Notes and References

  1. Bouchard . P. . Bousquet . Y. . Davies . A. . Alonso-Zarazaga . M. . Lawrence . J. . Lyal . C. . Newton . A. . Reid . C. . Schmitt . M. . Ślipiński . A. . Smith . A. . 2011 . Family-group names in Coleoptera (Insecta) . ZooKeys . 88 . 1–972 . 10.3897/zookeys.88.807 . 3088472 . 21594053 . free.
  2. Web site: Australian Faunal Directory . 2022-12-08 . biodiversity.org.au . en.
  3. Web site: Subfamily Cetoniinae - Fruit and Flower Chafers . 2022-12-08 . bugguide.net.
  4. Šípek . Petr . Fabrizi . Silvia . Eberle . Jonas . Ahrens . Dirk . 2016 . A molecular phylogeny of rose chafers (Coleoptera: Scarabaeidae: Cetoniinae) reveals a complex and concerted morphological evolution related to their flight mode . Molecular Phylogenetics and Evolution . en . 101 . 163–175 . 10.1016/j.ympev.2016.05.012. 27165937 .
  5. Book: Shipley. A. E.. Marshall. Guy A. K.. The Fauna of British India. 1910. Taylor and Francis. London, Calcutta and Bombay. 201.
  6. A remarkable predacious cetoniid, Spilophorus maculatus (Gory & Percheron), from southern India (Coleoptera: Scarabaeidae). Ghorpade, K.D.. The Coleopterists Bulletin. 29. 4. 1975. 226–230. 3999660.
  7. Beetles (Coleoptera, Scarabaeidae) in a social wasp nest (Hymenoptera, Vespidae) in India. Jeanne, Robert L. & J. H. Hunt. Entomologist's Monthly Magazine. 1992. 128. 139–141. 2014-10-28. https://web.archive.org/web/20160304055501/http://www4.ncsu.edu/~jhhunt/JeanneBeetles.pdf. 2016-03-04. dead.
  8. Cazier, M.A. & Marjorie Statham. 1962. The behaviour and habits of the myrmecophilous scarab, Cremastocheilus stathamae Cazier, with notes on other species (Coleoptera: Scarabaeidae).. J. New York Entomol. Soc. . 70. 125–149.

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