Ultrastructure of aedeagus and spermatheca of Chaetocnema major (Jacquelin du Val) (Chrysomelidae: Galerucinae: Alticini) by scanning electron microscope
Year 2023,
Volume: 4 Issue: 2, 116 - 132, 14.11.2023
Havva Ekmekci
,
Huseyin Özdikmen
,
Neslihan Bal
,
Damla Amutkan Mutlu
,
Zekiye Suludere
Abstract
The paper presents unknown ultrastructure of aedeagus and spermatheca of Chaetocnema major (Jacquelin du Val) which is the type species of the subgenus Chaetocnema (Majoroides) Özdikmen and the single species of the subgenus occurring in Türkiye, are studied and figured under both stereo microscope and Scanning Electron Microscope. Aedeagi and spermathecae have so far only been studied in the genus Chaetocnema Stephens based on stereo microscopic studies. Since no ultrastructural studies with SEM have been conducted on male and female genitalia of Chaetocnema species so far, this is the first study for this purpose. Accordingly, this study mainly aims to reveal and obtain new diagnostic characters by ultrastructural and detailed examinations of the aedeagus and spermathecae of Chaetocnema major. As is known, spermathecae of the genus Chaetocnema, like most genera of the tribe Alticini, are partially homogeneous and do not provide good diagnostic characters at the species level. Studies on the aedeagus have revealed that, despite the partial homogeneity of the spermatheca, the structure of the aedeagus shows quite a wide diversity. On the other hand, for the genus Chaetocnema, sperm sheath morphology at the subspecies level appears to be more relevant and useful than external morphological features. While the results from the present study mainly support these views, the female genitalia (spermathecae) was found to be insignificant and non-diagnostic, but important for diagnosis at the aedeagus species level, and further, investigation revealed new diagnostic characters for both Ultrastructural morphologies aedeagus and spermathecae by examining in SEM.
References
- 1. Döberl M. Alticinae. In: Löbl, I., Smetana, A. (Eds) Catalogue of Palearctic Coleoptera. Vol. 6. Apollo Books, Stenstrup, 2010; 491-563.
- 2. Konstantinov AS, Baselga A, Grebennikov VV., Prena J, Lingalfelter SW. Revision of the Palearctic Chaetocnema species. (Coleoptera: Chrysomelidae: Galerucinae: Alticini). Pensoft, Sofia, 2011; 363 pp.
- 3. Ekiz AN, Şen İ, Aslan EG, Gök A. Checklist of leaf beetles (Coleoptera: Chrysomelidae) of Türkiye, excluding Bruchinae. Journal of Natural History, 2013; 47(33-34), 2213-2287.
- 4. Özdikmen H. Chorotype identification for Turkish Chrysomeloidea (Coleoptera) Part VIII – Chrysomelidae: Alticinae. Munis Entomology & Zoology, 2014; 9(1), 325-375.
- 5. Özdikmen H. A review: A new subgeneric arrangement of the genus Chaetocnema Stephens (Chrysomelidae: Galerucinae: Alticini) with new subgenera based on spermathecal structures. Munis Entomology & Zoology,2021; 16(1), 41-105.
- 6. White RE. A revision of the genus Chaetocnema of America north of Mexico (Coleoptera: Chrysomelidae). Contributions of the American Entomological Institute, 1996; 29(1), 1-158.
- 7. Ruan Y, Yang X, Konstantinov AS, Prathapan KD, Zhang M. Revision of the Oriental Chaetocnema species (Coleoptera, Chrysomelidae, Galerucinae, Alticini). Zootaxa, 2019; 4699(1), 001-206.
- 8. Ruan Y, Konstantinov AS, Ge S, Yang X. Revision of the Chaetocnema picipes species-group (Coleoptera, Chrysomelidae, Galerucinae, Alticini) in China, with descriptions of three new species. Zookeys, 2014a; 387, 11-32.
- 9. Ruan Y, Konstantinov AS., Ge S, Yang X. Revision of Chaetocnema semicoerulea species-group (Coleoptera, Chrysomelidae, Galerucinae, Alticini) in China, with descriptions of three new species. Zookeys, 2014b;463, 57-74.
- 10. Ataş F, Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. A SEM study on aedeagus and spermatheca of Cassida seraphina Ménétries, 1836 (Coleoptera: Chrysomelidae: Cassidinae) from Türkiye. Munis Entomology & Zoology, 2019a; 14(2), 395-411.
- 11. Ataş F, Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. A SEM study on aedeagus and spermatheca of Cassida hablitziae Motschulsky, 1838 (Coleoptera: Chrysomelidae: Cassidinae) from Türkiye. Munis Entomology & Zoology, 2019b; 14(2), 519-529.
- 12. Bal N, Özdikmen H. Aedeagus structure of Cassida prasina Illiger, 1798 (Coleoptera: Chrysomelidae: Cassidinae) in Scanning Electrone Microscope (SEM). KSU Tarım ve Doğa Dergisi - KSU Journal of Agriculture and Nature, 2020; 23: 748-753.
- 13. Özdikmen H, Bal N. The structure of spermatheca in the subgenus Cassida (Cassida) Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Fresenius Environmental Buletin,2021;30(5), 5270-5291.
- 14. Özdikmen H, Coral Şahin D, Bal N, A new species of Cassida Linnaeus, 1758 from Türkiye (Chrysomelidae: Cassidinae). Microscopy Research and Technique, 2020a; 83, 1171-1177.
- 15. Özdikmen H, Bal N, Amutkan Mutlu D,Suludere Z. A SEM study on aedeagus and spermatheca of Cassida nebulosa Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) from Türkiye. Munis Entomology & Zoology, 2020b;15(1), 252-261.
- 16. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. Comparative ultrastructural analysis of six subgenera of Cassida Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) based on spermatheca of the type species and its taxonomic significance. Transactions of the American Entomological Society, 2021a; 147(1), 67-99.
- 17. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. The structure of spermathecae in the subgenus Cassida (Onychocassis) Spaeth in Spaeth & Reitter, 1926 (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Munis Entomology & Zoology, 2021b; 16(2), 972-984.
- 18. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. Ultrasturucture of aedeagus and spermatheca of type species of Hypocassida Weise, 1893 (Chrysomelidae: Cassidinae: Cassidini) and their taxonomic significance. International Journal of Tropical Insect Science, 2022a; 42, 2337-2349.
- 19. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. Comparative ultrastructural analysis to seven subgenera of Cassida Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) based on aedeagus of the type species and its taxonomic significance. Transactions of the American Entomological Society, 2022b;148(1), 65-112.
- 20. Sahlberg J. Coleoptera Mediterranea Orientalia, Quae in Aegypto, Palaestina, Syria, Caramania atque in Anatolia Occidentali anno 1904. Öfversigt af Finska Vatenskaps–Societetens Förhandlingar, 1913; 55A(19), 1-281.
- 21. Tölg F. Feststellungen von Nahrungspflanzen kleinasiatischer Halticinen und Ceuthorrhynchinen. Koleopterologische Rundschau, 1938; 24(6), 234-241.
- 22. Csiki, E, Heikertinger F. Chrysomelidae, Halticinae, in: Junk, W. and Schenkling, S., Coleoterorum catalogus, ‘s–Gravenshage(1911–1940), pars 59, 1-215.
- 23. Kral J. Wissenschaftlichen Ergebnis der zoologischen Expedition des Nationalmuseums in Prag nach der Türkei, Coleoptera, Alticidae. Acta Entomologica Musei Nationalis Pragae, 1967; 37, 261-268.
- 24. Medvedev L. A List of Chrysomelidae collected by Dr. W. Wittmer in Türkiye (Coleoptera). Revue suisse de Zoologie,1970; 77, 309-319.
- 25. Aslan İ, Gruev B, Özbek H. A preliminary review of the subfamily Alticinae (Coleoptera, Chrysomelidae) in Türkiye. Turkish Journal of Zoology, 23(1999), 373-414.
- 26. Gruev B. Actualized checklist of Alticinae (Coleoptera: Chrysomelidae) in the Balkan Peninsula. Travaux Scientifiques de l'Universite de Plovdiv Animalia, 2001; 37, 9-28.
- 27. Aslan EG, Gök A. Host–plant relationships of 65 flea beetles species from Türkiye, with new associations (Coleoptera: Chrysomelidae: Alticinae). Entomological News,2006; 117(3), 297-308.
- 28. Özdikmen H. A comprehensive contribution for leaf beetles of Türkiye with a zoogeographical evaluation for all Turkish fauna (Coleoptera: Chrysomelidae). Munis Entomology & Zoology, 2011; 6(2), 540-638.
- 29. Bal N, Özdikmen H, Coral Şahin D. Twenty-six new flea beetles for the fauna of Çankırı province in Türkiye (Chrysomelidae: Galerucinae: Alticini). Munis Entomology & Zoology, 2018; 13(2), 527-537.
- 30. Kasap H, Crowson RA, The male reproductive organs of Bruchidae and Chrysomelidae (Coleoptera). Turkish Journal of Entomology, 1979; 3, 199-216.
- 31. Mann, JS, Crowson RA. On the internal male reproductive organs and their taxonomic significance in the Leaf Beetles (Coleoptera: Chrysomelidae). Entomologia Generalis, 1983; 9, 75-99.
- 32. Suzuki K. Comparative morphology of the internal reproductive system of Chrysomelidae (Coleoptera). In: Jolivet P, Petitpierre E, Hsiao TH (Eds) Biology of Chrysomelidae. Kluwer Academic Publishers, Dordrecht, Boston and London, 1988; 317-355.
- 33. Arnqvist G. The evolution of animal genitalia: distinguishing between hypotheses by single species studies. Biological Journal of the Linnaean Society, 1997; 60, 365-379.
- 34. Hernández JM, Ortuño VM. Estudio de la genitalia femenina en Iberodorcadion Breuning, 1943 y comentarios sobre su valor taxonómico (Coleoptera: Cerambycidae). Graellsia, 1992; 48, 91-97.
- 35. Hernández JM. La genitalia femenina en el género Corymbia Des Gozis, 1886 (Coleóptera, Cerambycidae). Elytron,1993; 7, 99-104.
- 36. Ferronato EMO. A cápsula da espermateca dos Eumolpinae (Coleoptera: Chrysomelidae) associados a Theobroma cacao L. (Sterculiaceae) do Sul da Bahia. Anais da Sociedade Entomológica do Brasil, 2000; 29, 57-63.
- 37. Kısmalı Ş. İzmir ili ve çevresinde kültür bitkilerinde zarar yapan Chrysomelinae ve Halticinae (Coleoptera: Chrysomelidae) alt familyalarına ait türler, tanınmaları, konukçuları, yayılışları ve kısa biyolojileri üzerinde araştırmalar. Ege Üniversitesi Ziraat Fakültesi Dergisi, Seri A, 1973; 10(2), 341-378.
- 38. Snodgrass RE. Principles of Insect Morphology. McGraw-Hill Book Co., New York and London, 1935; 667 pp.
- 39. De Marzo, L. Biodiversita della spermateca nei coleotteri. Atti Accademia Nazionale Italiana di Entomologia Anno, 2008; LVI, 69-96.
- 40. Pascini TV., Martins GF. The insect spermatheca: an overview. Zoology, 2017; 121, 56-71.
- 41. Rodríguez-Mirón GM, Zaragoza-Caballero S. Revisión taxonómica del género Mastostethus (Coleoptera: Megalopodidae) en México y descripción de dos especies nuevas. Revista Mexicana de Biodiversidad,2017, 88, 312-334.
- 42. Matsumura Y, Suzuki K. Comparative morphology of internal reproductive systems in leaf beetles of the Donaciinae and Criocerinae (Coleoptera: Chrysomelidae) and its implication for the phylogeny. Zootaxa, 2008; 1845, 1-32.
- 43. Reid CAM. The Australian species of the tribe Zeugophorini (Coleoptera: Chrysomelidae: Megalopodinae). General and Applied Entomology, 1989; 21, 39-47.
- 44. Biondi M.Revision of the species of Chaetocnema from Madagascar (Coleoptera Chrysomelidae Alticinae). European Journal of Entomology,2001; 98, 233-248.
- 45. Borowiec L, Świętojańska J. Revision of Cassida litigiosa group from southern Africa (Coleoptera: Chrysomelidae: Cassidinae). Annales Zoologici, 2001; 51, 153-184.
- 46. Borowiec L, Skuza M. The structure of spermatheca in the genus Chelymorpha Chevrolat, 1837 (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Annales Zoologici,2004; 54, 439-451.
- 47. Borowiec L, Opalinska S. The structure of spermathecae of selected genera of Stolaini and Eugenysini (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Annales Zoologici,2007; 57, 463-479.
- 48. Borowiec L, Pomorska J. The structure of the spermathecae of the genus Stolas (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini) and its taxonomic significance. Annales Zoologici, 2009; 59, 201-221.
- 49. Rodríguez-Mirón GM., Zaragoza-Caballero S. Revisión taxonómica del género Mastostethus (Coleoptera: Megalopodidae) en México y descripción de dos especies nuevas. Revista Mexicana de Biodiversidad, 2017, 88, 312-334.
Chaetocnema major (Jacquelin du Val) (Chrysomelidae: Galerucinae: Alticini)’ün aedeagus ve spermateka'sının taramalı elektron mikroskobu ile ince yapısı
Year 2023,
Volume: 4 Issue: 2, 116 - 132, 14.11.2023
Havva Ekmekci
,
Huseyin Özdikmen
,
Neslihan Bal
,
Damla Amutkan Mutlu
,
Zekiye Suludere
Abstract
Bu makale, Chaetocnema major (Jacquelin du Val) türünün aedeagus ve spermatekasının bilinmeyen ultrastrüktürünü sunmaktadır. Chaetocnema (Majoroides) Özdikmen alt cinsinin tip türleri ve Türkiye'de bulunan tek türü hem stereo mikroskop hem de Taramalı Elektron Mikroskobu altında incelenmiş ve figüre edilmiştir. Aedeagi ve spermathecae şimdiye kadar sadece Chaetocnema Stephens cinsinde stereo mikroskobik çalışmalara dayanarak incelenmiştir. Chaetocnema türlerinin erkek ve dişi genital organları üzerinde bugüne kadar SEM ile ultrastrüktürel bir çalışma yapılmadığından, bu çalışma bu amaca yönelik ilk çalışmadır. Buna göre, bu çalışma Chaetocnema major'un aedeagus ve spermateka'sının ultrastrüktürel ve detaylı incelemeleri ile yeni diagnostik karakterleri ortaya çıkarmayı ve elde etmeyi amaçlamaktadır. Bilindiği gibi, Chaetocnema cinsinin spermatekaları, Alticini kabilesinin çoğu cinsi gibi, kısmen homojendir ve tür düzeyinde iyi teşhis karakterleri sağlamaz. Aedeagus üzerine yapılan çalışmalar, spermatekanın kısmi homojenliğine rağmen, aedeagusun yapısının oldukça geniş bir çeşitlilik gösterdiğini ortaya koymuştur. Öte yandan, Chaetocnema cinsi için, alt tür düzeyinde sperm kılıfı morfolojisi, dış morfolojik özelliklerden daha ilgili ve yararlı görünmektedir. Mevcut çalışmadan elde edilen sonuçlar temel olarak bu görüşleri desteklerken, dişi genital organın (spermateka) önemsiz ve tanısal olmadığı, ancak aedeagus tür düzeyinde tanı için önemli olduğu bulunmuştur ve ayrıca, SEM'de incelenerek hem Ultrastrüktürel morfolojiler aedeagus hem de spermathecae için yeni tanısal karakterler ortaya çıkarılmıştır.
Thanks
This study is derived from the first author's doctoral thesis.
References
- 1. Döberl M. Alticinae. In: Löbl, I., Smetana, A. (Eds) Catalogue of Palearctic Coleoptera. Vol. 6. Apollo Books, Stenstrup, 2010; 491-563.
- 2. Konstantinov AS, Baselga A, Grebennikov VV., Prena J, Lingalfelter SW. Revision of the Palearctic Chaetocnema species. (Coleoptera: Chrysomelidae: Galerucinae: Alticini). Pensoft, Sofia, 2011; 363 pp.
- 3. Ekiz AN, Şen İ, Aslan EG, Gök A. Checklist of leaf beetles (Coleoptera: Chrysomelidae) of Türkiye, excluding Bruchinae. Journal of Natural History, 2013; 47(33-34), 2213-2287.
- 4. Özdikmen H. Chorotype identification for Turkish Chrysomeloidea (Coleoptera) Part VIII – Chrysomelidae: Alticinae. Munis Entomology & Zoology, 2014; 9(1), 325-375.
- 5. Özdikmen H. A review: A new subgeneric arrangement of the genus Chaetocnema Stephens (Chrysomelidae: Galerucinae: Alticini) with new subgenera based on spermathecal structures. Munis Entomology & Zoology,2021; 16(1), 41-105.
- 6. White RE. A revision of the genus Chaetocnema of America north of Mexico (Coleoptera: Chrysomelidae). Contributions of the American Entomological Institute, 1996; 29(1), 1-158.
- 7. Ruan Y, Yang X, Konstantinov AS, Prathapan KD, Zhang M. Revision of the Oriental Chaetocnema species (Coleoptera, Chrysomelidae, Galerucinae, Alticini). Zootaxa, 2019; 4699(1), 001-206.
- 8. Ruan Y, Konstantinov AS, Ge S, Yang X. Revision of the Chaetocnema picipes species-group (Coleoptera, Chrysomelidae, Galerucinae, Alticini) in China, with descriptions of three new species. Zookeys, 2014a; 387, 11-32.
- 9. Ruan Y, Konstantinov AS., Ge S, Yang X. Revision of Chaetocnema semicoerulea species-group (Coleoptera, Chrysomelidae, Galerucinae, Alticini) in China, with descriptions of three new species. Zookeys, 2014b;463, 57-74.
- 10. Ataş F, Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. A SEM study on aedeagus and spermatheca of Cassida seraphina Ménétries, 1836 (Coleoptera: Chrysomelidae: Cassidinae) from Türkiye. Munis Entomology & Zoology, 2019a; 14(2), 395-411.
- 11. Ataş F, Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. A SEM study on aedeagus and spermatheca of Cassida hablitziae Motschulsky, 1838 (Coleoptera: Chrysomelidae: Cassidinae) from Türkiye. Munis Entomology & Zoology, 2019b; 14(2), 519-529.
- 12. Bal N, Özdikmen H. Aedeagus structure of Cassida prasina Illiger, 1798 (Coleoptera: Chrysomelidae: Cassidinae) in Scanning Electrone Microscope (SEM). KSU Tarım ve Doğa Dergisi - KSU Journal of Agriculture and Nature, 2020; 23: 748-753.
- 13. Özdikmen H, Bal N. The structure of spermatheca in the subgenus Cassida (Cassida) Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Fresenius Environmental Buletin,2021;30(5), 5270-5291.
- 14. Özdikmen H, Coral Şahin D, Bal N, A new species of Cassida Linnaeus, 1758 from Türkiye (Chrysomelidae: Cassidinae). Microscopy Research and Technique, 2020a; 83, 1171-1177.
- 15. Özdikmen H, Bal N, Amutkan Mutlu D,Suludere Z. A SEM study on aedeagus and spermatheca of Cassida nebulosa Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) from Türkiye. Munis Entomology & Zoology, 2020b;15(1), 252-261.
- 16. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. Comparative ultrastructural analysis of six subgenera of Cassida Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) based on spermatheca of the type species and its taxonomic significance. Transactions of the American Entomological Society, 2021a; 147(1), 67-99.
- 17. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. The structure of spermathecae in the subgenus Cassida (Onychocassis) Spaeth in Spaeth & Reitter, 1926 (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Munis Entomology & Zoology, 2021b; 16(2), 972-984.
- 18. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. Ultrasturucture of aedeagus and spermatheca of type species of Hypocassida Weise, 1893 (Chrysomelidae: Cassidinae: Cassidini) and their taxonomic significance. International Journal of Tropical Insect Science, 2022a; 42, 2337-2349.
- 19. Özdikmen H, Bal N, Amutkan Mutlu D, Suludere Z. Comparative ultrastructural analysis to seven subgenera of Cassida Linnaeus, 1758 (Coleoptera: Chrysomelidae: Cassidinae) based on aedeagus of the type species and its taxonomic significance. Transactions of the American Entomological Society, 2022b;148(1), 65-112.
- 20. Sahlberg J. Coleoptera Mediterranea Orientalia, Quae in Aegypto, Palaestina, Syria, Caramania atque in Anatolia Occidentali anno 1904. Öfversigt af Finska Vatenskaps–Societetens Förhandlingar, 1913; 55A(19), 1-281.
- 21. Tölg F. Feststellungen von Nahrungspflanzen kleinasiatischer Halticinen und Ceuthorrhynchinen. Koleopterologische Rundschau, 1938; 24(6), 234-241.
- 22. Csiki, E, Heikertinger F. Chrysomelidae, Halticinae, in: Junk, W. and Schenkling, S., Coleoterorum catalogus, ‘s–Gravenshage(1911–1940), pars 59, 1-215.
- 23. Kral J. Wissenschaftlichen Ergebnis der zoologischen Expedition des Nationalmuseums in Prag nach der Türkei, Coleoptera, Alticidae. Acta Entomologica Musei Nationalis Pragae, 1967; 37, 261-268.
- 24. Medvedev L. A List of Chrysomelidae collected by Dr. W. Wittmer in Türkiye (Coleoptera). Revue suisse de Zoologie,1970; 77, 309-319.
- 25. Aslan İ, Gruev B, Özbek H. A preliminary review of the subfamily Alticinae (Coleoptera, Chrysomelidae) in Türkiye. Turkish Journal of Zoology, 23(1999), 373-414.
- 26. Gruev B. Actualized checklist of Alticinae (Coleoptera: Chrysomelidae) in the Balkan Peninsula. Travaux Scientifiques de l'Universite de Plovdiv Animalia, 2001; 37, 9-28.
- 27. Aslan EG, Gök A. Host–plant relationships of 65 flea beetles species from Türkiye, with new associations (Coleoptera: Chrysomelidae: Alticinae). Entomological News,2006; 117(3), 297-308.
- 28. Özdikmen H. A comprehensive contribution for leaf beetles of Türkiye with a zoogeographical evaluation for all Turkish fauna (Coleoptera: Chrysomelidae). Munis Entomology & Zoology, 2011; 6(2), 540-638.
- 29. Bal N, Özdikmen H, Coral Şahin D. Twenty-six new flea beetles for the fauna of Çankırı province in Türkiye (Chrysomelidae: Galerucinae: Alticini). Munis Entomology & Zoology, 2018; 13(2), 527-537.
- 30. Kasap H, Crowson RA, The male reproductive organs of Bruchidae and Chrysomelidae (Coleoptera). Turkish Journal of Entomology, 1979; 3, 199-216.
- 31. Mann, JS, Crowson RA. On the internal male reproductive organs and their taxonomic significance in the Leaf Beetles (Coleoptera: Chrysomelidae). Entomologia Generalis, 1983; 9, 75-99.
- 32. Suzuki K. Comparative morphology of the internal reproductive system of Chrysomelidae (Coleoptera). In: Jolivet P, Petitpierre E, Hsiao TH (Eds) Biology of Chrysomelidae. Kluwer Academic Publishers, Dordrecht, Boston and London, 1988; 317-355.
- 33. Arnqvist G. The evolution of animal genitalia: distinguishing between hypotheses by single species studies. Biological Journal of the Linnaean Society, 1997; 60, 365-379.
- 34. Hernández JM, Ortuño VM. Estudio de la genitalia femenina en Iberodorcadion Breuning, 1943 y comentarios sobre su valor taxonómico (Coleoptera: Cerambycidae). Graellsia, 1992; 48, 91-97.
- 35. Hernández JM. La genitalia femenina en el género Corymbia Des Gozis, 1886 (Coleóptera, Cerambycidae). Elytron,1993; 7, 99-104.
- 36. Ferronato EMO. A cápsula da espermateca dos Eumolpinae (Coleoptera: Chrysomelidae) associados a Theobroma cacao L. (Sterculiaceae) do Sul da Bahia. Anais da Sociedade Entomológica do Brasil, 2000; 29, 57-63.
- 37. Kısmalı Ş. İzmir ili ve çevresinde kültür bitkilerinde zarar yapan Chrysomelinae ve Halticinae (Coleoptera: Chrysomelidae) alt familyalarına ait türler, tanınmaları, konukçuları, yayılışları ve kısa biyolojileri üzerinde araştırmalar. Ege Üniversitesi Ziraat Fakültesi Dergisi, Seri A, 1973; 10(2), 341-378.
- 38. Snodgrass RE. Principles of Insect Morphology. McGraw-Hill Book Co., New York and London, 1935; 667 pp.
- 39. De Marzo, L. Biodiversita della spermateca nei coleotteri. Atti Accademia Nazionale Italiana di Entomologia Anno, 2008; LVI, 69-96.
- 40. Pascini TV., Martins GF. The insect spermatheca: an overview. Zoology, 2017; 121, 56-71.
- 41. Rodríguez-Mirón GM, Zaragoza-Caballero S. Revisión taxonómica del género Mastostethus (Coleoptera: Megalopodidae) en México y descripción de dos especies nuevas. Revista Mexicana de Biodiversidad,2017, 88, 312-334.
- 42. Matsumura Y, Suzuki K. Comparative morphology of internal reproductive systems in leaf beetles of the Donaciinae and Criocerinae (Coleoptera: Chrysomelidae) and its implication for the phylogeny. Zootaxa, 2008; 1845, 1-32.
- 43. Reid CAM. The Australian species of the tribe Zeugophorini (Coleoptera: Chrysomelidae: Megalopodinae). General and Applied Entomology, 1989; 21, 39-47.
- 44. Biondi M.Revision of the species of Chaetocnema from Madagascar (Coleoptera Chrysomelidae Alticinae). European Journal of Entomology,2001; 98, 233-248.
- 45. Borowiec L, Świętojańska J. Revision of Cassida litigiosa group from southern Africa (Coleoptera: Chrysomelidae: Cassidinae). Annales Zoologici, 2001; 51, 153-184.
- 46. Borowiec L, Skuza M. The structure of spermatheca in the genus Chelymorpha Chevrolat, 1837 (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Annales Zoologici,2004; 54, 439-451.
- 47. Borowiec L, Opalinska S. The structure of spermathecae of selected genera of Stolaini and Eugenysini (Coleoptera: Chrysomelidae: Cassidinae) and its taxonomic significance. Annales Zoologici,2007; 57, 463-479.
- 48. Borowiec L, Pomorska J. The structure of the spermathecae of the genus Stolas (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini) and its taxonomic significance. Annales Zoologici, 2009; 59, 201-221.
- 49. Rodríguez-Mirón GM., Zaragoza-Caballero S. Revisión taxonómica del género Mastostethus (Coleoptera: Megalopodidae) en México y descripción de dos especies nuevas. Revista Mexicana de Biodiversidad, 2017, 88, 312-334.