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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: Cucujiformia
Cladus: Phytophaga
Superfamilia: Chrysomeloidea

Subfamilia: Cassidinae
Tribus (after Borowiec & Świętojańska): Alurnini – Anisoderini – Aproidini – AresciniAspidimorphiniBasiprionotini – Bothryonopini – Callispini – Callohispini – CassidiniChalepini – Coelaenomenoderini – Cryptonychini – Cubispini – DelocraniniDorynotiniEugenysini – Eurispini – Exothispini – Goniocheniini – Gonophorini – HemosphaerotiniHispini – Hispoleptini – Hybosispini – Imatidiini – Ischyrosonychini – Leptispini – Mesomphaliini – NotosacanthiniOmoceriniOncocephalini – Promecothecini – Prosopodontini – SceloenopliniSpilophorini – †Oposispini

? †Oligocassida


Cassidinae Gyllenhal, 1813: 434

Type genus: Cassida Linnaeus, 1758.

Hispinae Gyllenhal, 1813: 448

Primary references

Gyllenhal, L. 1813. Insecta Suecica. Classis I. Coleoptera sive Eleuterata. Tomi I, Pars III. Scaris: L.J. Leverentz, [2] + 730 + [2] pp. BHL. Reference page.

Additional references

Borowiec, L. 1999. A world catalogue of the Cassidinae (Coleoptera: Chrysomelidae). Biologica Silesiae, Wroclaw.
Borowiec, L. 2009. New records of Neotropical tortoise beetles (Coleoptera: Chrysomelidae: Cassidinae). Genus 20(4): 615–722. PDF.
Borowiec, L. & Sekerka, L. 2010b. Subfamily Cassidinae Gyllenhal, 1813. Pp. 368–390. In: Löbl, I. & Smetana, A. (eds.) Catalogue of Palaearctic Coleoptera. Volume 6. Chrysomeloidea. Apollo Books, Stenstrup, Denmark. 924 pp. ISBN 978-87-88757-84-2. Google Books. Reference page.
Borowiec, L. & Świętojańska, J. 2014. 2.7.2 Cassidinae Gyllenhal, 1813. Pp. 198–217 in: Leschen et al., 2014. Full chapter 2.7 (with link to PDF) Reference page.
Flinte, V., Borowiec, L., de Freitas, S., Viana, J.H., Fernandes, F.R., Nogueira-De-Sá, F., de Macedo, M.V. & Monteiro, R.F. 2009. Tortoise beetles of the State of Rio de Janeiro, Brazil (Coleoptera: Chrysomelidae: Cassidinae). Genus 20(4): 571–614. PDF.
Hincks, W.D. 1952. The genera of the Cassidinae (Coleoptera: Chrysomelidae). Transactions of the Royal Entomological Society of London 103: 327–358. DOI: 10.1111/j.1365-2311.1952.tb01061.x Paywall.
Nadein, K.S., Perkovsky, E.E. & Moseyko, A.G. 2015. New Late Eocene Chrysomelidae (Insecta: Coleoptera) from Baltic, Rovno and Danish ambers. Papers in Palaeontology 2(1): 117–137. DOI: 10.1002/spp2.1034 Paywall. PDF. Reference page.
Peng, L-F., Li, J-L., Hou, Y-M. & Zhang, X. 2018. Descriptions of immature stages of Octodonta nipae (Maulik) (Coleoptera, Chrysomelidae, Cassidinae, Cryptonychini). ZooKeys 764: 91–109. DOI: 10.3897/zookeys.764.24168. Reference page.
Sekerka, L., Jia, F.L., Pang, H. & Boroweic, L. 2016. Cassidinae (Coleoptera: Chrysomelidae) types deposited at Sun Yat-sen University, Guangzhou, China. Zootaxa 4084(1): 50–78. DOI: 10.11646/zootaxa.4084.1.2 Paywall. ResearchGate Open access.Reference page.
Shameem, K.M., Prathapan, K.D., Nasser, M. & Chaboo, C.S. 2016. Natural history of Javeta pallida Baly, 1858 on Phoenix palms in India (Chrysomelidae, Cassidinae, Coelaenomenoderini). In: Jolivet, P., Santiago-Blay, J. & Schmitt, M. (eds.) Research on Chrysomelidae 6. ZooKeys 597: 39–56. DOI: 10.3897/zookeys.597.6876 Open access. Reference page.
Simões, M.V.P., Husemann, M. & Sekerka, L. 2021. A Catalog of the Tortoise Beetle (Coleoptera: Chrysomelidae: Cassidinae) Collection Deposited in the Zoological Museum Hamburg (ZMH). The Coleopterists Bulletin 75(1): 191–210. DOI: 10.1649/0010-065X-75.1.191 Paywall. ResearchGate Open access. Reference page.
Staines, C.L. 2009. The Hispine Beetles (Coleoptera: Chrysomelidae: Cassidinae) of the Caribbean Basin with a Key to the Species of Hispaniola. Annals of Carnegie Museum 78(1): 17–28. DOI: 10.2992/007.078.0102 Paywall.
Staines, C.L. 2010. Nomenclatural notes on Chalepini and Sceloenopliini (Coleoptera: Chrysomelidae: Cassidinae. Insecta Mundi (0122): 1–2. PDF.
Stephens, J.F. 1831–1832. Illustrations of British entomology; or, a synopsis of indigenous insects: containing their generic and specific distinctions; with an account of their metamorphoses, times of appearance, localities, food, and economy, as far as practicable. Mandibulata, Vol. IV. London: Baldwin & Cradock, 413 pp., pls. 20–23. [published in parts: pp. 1–366, 1831; 367–413, 1832] BHL Reference page. [see pages 364, 365]


Cassidinae (in traditional sense)

Cassidinae of the world – an interactive manual (Coleoptera: Chrysomelidae) (by Lech Borowiec and Jolanta Świętojańska)
Cassidinae Gyllenhal, 1813
Subfamily Cassidinae (Chrysomelidae) – atlas of leaf beetles of Russia

Hispinae (in traditional sense)

Hispinae Gyllenhal, 1813
Subfamily Hispinae (Chrysomelidae) – atlas of leaf beetles of Russia
Staines, C.L. 2015. Catalog of the Hispines of the World (Coleoptera: Chrysomelidae: Cassidinae). Online.
Hispines of the World by C.L. Staines.
Indian Species of Chrysomelidae

Vernacular names
Deutsch: Schildkäfer
English: Tortoise Beetles
français: Casside
日本語: カメノコハムシ亜科
русский: Щитоноски

The Cassidinae (tortoise and leaf-mining beetles) are a subfamily of the leaf beetles, or Chrysomelidae. The antennae arise close to each other and some members have the pronotal and elytral edges extended to the side and covering the legs so as to give them the common name of tortoise beetles. Some members, such as in the tribe Hispini, are notable for the spiny outgrowths to the pronotum and elytra.


The "cassidoids" have a rounded outline with the edges of the pronotum and elytra expanded, spreading out to cover the legs and head. They are often colourful and metallic, with ornate sculpturing; a few species have the ability to change the colour due to water movements within the translucent cuticle. All members of the subfamily have the mouthparts reduced into a cavity in the head capsule, the legs have four segmented tarsi. The hispoids have larvae that are leaf miners, while the cassidoids feed on the plant surfaces, sometimes covering their bodies with faecal shields. Although fecal shields are thought to provide defense, no evidence exists for such a role.[2]

A few species in two closely related tribes (Mesomphaliini and Eugenysini, putative sister taxa) show maternal care of larvae.[3][4] These species can be viewed as subsocial or parasocial, with evidence pointing to there being two evolutionary origins of subsociality within this one lineage.[5][6]

It includes both the former subfamily "Hispinae" (leaf-mining beetles), as well as the former more narrowly defined subfamily Cassidinae (familiar as tortoise beetles) which are now split into several tribes that include the tribe Cassidini, and in all include over 125 genera. The traditional separation of the two groups was based essentially on the habitats of the larvae and the general shapes of the adults. The name Cassidinae for the merged subfamily is considered to have priority.[7]

The former grouping of "Hispinae" (sometimes called leaf-mining beetles, or "hispoids") included the tribes Alurnini, Anisoderini, Aproidini, Arescini, Bothryonopini, Callispini, Callohispini, Cephaloleiini, Chalepini, Coelaenomenoderini, Cryptonychini, Cubispini, Eurispini, Exothispini, Gonophorini, Hispini, Hispoleptini, Hybosispini, Leptispini, Oediopalpini, Oncocephalini, Promecothecini, Prosopodontini, Sceloenoplini and Spilophorini.[1] Most members of these tribes are elongated, slightly flattened beetles with parallel margins, and antennal bases close together on their small heads. They often have punctate elytra and pronotum, sometimes with spines both on and along the edges. The former grouping of Cassidinae (sometimes called tortoise beetles, or "cassidoids") included the tribes Aspidimorphini, Basiprionotini, Cassidini, Delocraniini, Dorynotini, Eugenysini, Goniocheniini, Hemisphaerotini, Mesomphaliini, Notosacanthini, Omocerini and Physonotini.[1]

The subfamily names Cassidinae and Hispinae are both founded by Gyllenhal in the same 1813 book, but following the Principle of the First Reviser, Chen in this case,[8] priority is given to the name Cassidinae.[7]

See also

List of Cassidinae genera


Borowiec, L.; Świętojańska, J. 2014: 2.7.2 Cassidinae Gyllenhal, 1813. Pp. 198–217 in: Leschen et al., 2014
Bottcher, A; Jorge Paulo Zolin; Flávia Nogueira-de-Sá; José Roberto Trigo (2009). "Faecal shield chemical defence is not important in larvae of the tortoise beetle Chelymorpha reimoseri (Chrysomelidae: Cassidinae: Stolaini)". Chemoecology. 19 (1): 63–66. doi:10.1007/s00049-009-0006-x. S2CID 23072397.
Chaboo, C.S., F.A. Frieiro-Costa, J. Gómez-Zurita, R. Westerduijn. 2014. Subsociality in leaf beetles (Coleoptera: Chrysomelidae: Cassidinae, Chrysomelinae). Journal of Natural History 48: 1–44.
Flowers, RW (2008). "Leaf beetles (Coleoptera: Chrysomelidae)". In Capinera, JL (ed.). Encyclopedia of Entomology (2 ed.). Springer. p. 2148.
Chaboo, C.S., F.A. Frieiro-Costa, J. Gómez-Zurita, R. Westerduijn. 2014. Subsociality in leaf beetles (Coleoptera: Chrysomelidae: Cassidinae, Chrysomelinae). Journal of Natural History 48: 1–44.
Leocadio, Michele; Sekarka, Lukas; Simoes, Marianna; Shrago, Carlos. "Molecular systematics reveals the origins of subsociality in tortoise beetles (Coleoptera, Chrysomelidae, Cassidinae): Evolution of subsociality in Cassidinae". doi:10.1111/syen.12434. S2CID 219450159.
Chaboo, CS (2007). "Biology and phylogeny of the Cassidinae Gyllenhal sensu lato (tortoise and leaf-mining beetles) (Coleoptera: Chrysomelidae)". Bulletin of the American Museum of Natural History. 305: 1–250. doi:10.1206/0003-0090(2007)305[1:BAPOTC]2.0.CO;2. S2CID 83469991.
Chen, S. H. (1940). "Attempt at a new classification of the leaf beetles". Sinensia. 11: 451–481.

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