# Cassidinae

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{{Short description|Subfamily of beetles}}
{{Automatic taxobox
| image = Hispid beetle.jpg
| image_caption = ''[Platypria](/source/Platypria)'' sp.
| taxon = Cassidinae
| authority = [Gyllenhal](/source/Leonard_Gyllenhaal), 1813
| subdivision_ranks = Tribes
| subdivision_ref = <ref name="Borowiec"/>
| subdivision = <small>(m): monogeneric</small>
*[Alurnini](/source/Alurnini)
*[Anisoderini](/source/Anisoderini)
*[Aproidini](/source/Aproidini) (m)
*[Arescini](/source/Arescini)
*[Aspidimorphini](/source/Aspidimorphini) 
*[Basiprionotini](/source/Basiprionotini)
*[Bothryonopini](/source/Bothryonopini)
*[Callispini](/source/Callispini)
*[Callohispini](/source/Callohispini) (m)
*[Cassidini](/source/Cassidini)
*[Chalepini](/source/Chalepini) (= Uroplatini)
*[Coelaenomenoderini](/source/Coelaenomenoderini)
*[Cryptonychini](/source/Cryptonychini)
*[Cubispini](/source/Cubispini)
*[Delocraniini](/source/Delocraniini) (m)
*[Dorynotini](/source/Dorynotini)
*[Eugenysini](/source/Eugenysini)
*[Eurispini](/source/Eurispini)
*[Exothispini](/source/Exothispini) (m)
*[Goniocheniini](/source/Goniocheniini)
*[Gonophorini](/source/Gonophorini)
*[Hemisphaerotini](/source/Hemisphaerotini)
*[Hispini](/source/Hispini)
*[Hispoleptini](/source/Hispoleptini) (m)
*[Hybosispini](/source/Hybosispini)
*[Imatidiini](/source/Imatidiini) (= Cephaloleiini)
*[Ischyrosonychini](/source/Ischyrosonychini) (= Physonotini)
*[Leptispini](/source/Leptispini)
*[Mesomphaliini](/source/Mesomphaliini) (= Stolaini)
*[Notosacanthini](/source/Notosacanthini)
*[Omocerini](/source/Omocerini)
*[Oncocephalini](/source/Oncocephalini)
*[Promecothecini](/source/Promecothecini)
*[Prosopodontini](/source/Prosopodontini) (m)
*[Sceloenoplini](/source/Sceloenoplini)
*[Spilophorini](/source/Spilophorini) (= Oediopalpini)
}}

The '''Cassidinae''' ('''tortoise and leaf-mining beetles''') are a [subfamily](/source/subfamily) of the leaf beetles, or [Chrysomelidae](/source/Chrysomelidae). The antennae arise close to each other and some members have the pronotal and [elytral](/source/Elytron) 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](/source/tribe_(biology)) Hispini, are notable for the spiny outgrowths to the [pronotum](/source/pronotum) and [elytra](/source/Elytron).

==Description==
The "cassidoids" have a rounded outline with the edges of the [pronotum](/source/pronotum) and [elytra](/source/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 colour due to water movements within the translucent [cuticle](/source/Arthropod_cuticle). All members of the subfamily have the mouthparts reduced into a cavity in the head capsule, the legs have four segmented [tarsi](/source/Arthropod_leg). The hispoids have [larvae](/source/Larva) that are ecological diverse as [leaf miners](/source/Leaf_miner) or cryptic exophagous feeders, while the cassidoids feed freely externally on the plant surfaces. Within cassidoids, the tortoise beetles are diagnosed by the larvae recycling their feces and cast [exuviae](/source/exuviae) into a discrete structure held on the caudal processes (or, urogomphi). [Fecal shields](/source/Fecal_shield) are thought to provide protection from the sun, wind, heat, predators, parasites, and/or [parasitoids](/source/Parasitoid) though experimental evidence is mixed.<ref>Chaboo, C.S., S. Adam, K. Nishida, L. Schletzbaum. 2023. Architecture, construction, retention, and repair of fecal shields in three tribes of tortoise beetles (Insecta: Coleoptera: Chrysomelidae Cassidinae: Cassidini, Mesomphaliini, Spilophorini). ZooKeys Special Issue, Research on Chrysomelidae 9. ZooKeys 1177:87–146. DOI: 10.3897/zookeys.1177.102600</ref><ref>{{cite journal|journal= Chemoecology| year=2009| volume =19| issue=1| pages=63–66| title=Faecal shield chemical defence is not important in larvae of the tortoise beetle ''Chelymorpha reimoseri'' (Chrysomelidae: Cassidinae: Stolaini)|author1=Bottcher, A |author2=Jorge Paulo Zolin |author3=Flávia Nogueira-de-Sá |author4=José Roberto Trigo | doi=10.1007/s00049-009-0006-x| bibcode=2009Chmec..19...63B| s2cid=23072397}}</ref>

A few species in two closely related tribes ([Mesomphaliini](/source/Mesomphaliini) and [Eugenysini](/source/Eugenysini), putative [sister taxa](/source/sister_taxa)) show [paternal care](/source/Parental_care) of [larvae](/source/larvae).<ref name="Chaboo, C.S. 2014">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.</ref><ref>{{cite book|page=[https://archive.org/details/encyclopediaento00capi/page/n2211 2148]|editor=Capinera, JL|edition=2| title= Encyclopedia of Entomology|year=2008|url=https://archive.org/details/encyclopediaento00capi|url-access=limited|publisher=Springer|chapter=Leaf beetles (Coleoptera: Chrysomelidae)|author=Flowers, RW}}</ref> These species can be viewed as [subsocial](/source/subsocial), with evidence pointing to there being two evolutionary origins of subsociality within this one lineage.<ref name="Chaboo, C.S. 2014"/><ref>{{cite journal| last1 = Leocádio | first1 = Michele| last2 = Simões | first2 = Marianna V. P. | last3 = Sekarka | first3 = Lukas | last4 = Shrago | first4 = Carlos G. | last5 = Mermudes | first5 = José R. M. | last6 = Windsor | first6 = Donald M. | date = 2020 | title = Molecular systematics reveals the origins of subsociality in tortoise beetles (Coleoptera, Chrysomelidae, Cassidinae) | journal = [Systematic Entomology](/source/Systematic_Entomology) | volume = 45 | issue = 4 | pages = 894–910 | url = https://www.researchgate.net/publication/341462065 |doi=10.1111/syen.12434| bibcode = 2020SysEn..45..894L| s2cid = 219450159 }}</ref>

==Taxonomy==
Cassidinae 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](/source/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.<ref name=chaboo>{{cite journal|title=Biology and phylogeny of the Cassidinae Gyllenhal sensu lato (tortoise and leaf-mining beetles) (Coleoptera: Chrysomelidae)|author=Chaboo, CS| journal=Bulletin of the American Museum of Natural History| volume=305|pages=1–250| year=2007| doi=10.1206/0003-0090(2007)305[1:BAPOTC]2.0.CO;2|s2cid=83469991 | url=https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1477&context=entomologyfacpub| doi-access=free}}</ref>

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.<ref name="Borowiec"/> 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.<ref name="Borowiec"/>

The subfamily names Cassidinae and Hispinae are both founded by Gyllenhal in the same 1813 book, but following the [Principle of the First Reviser](/source/Principle_of_the_First_Reviser), Chen in this case,<ref>{{cite journal|author= Chen, S. H.|year=1940| title=Attempt at a new classification of the leaf beetles| journal=Sinensia| volume=11| pages=451–481}}</ref>  priority is given to the name Cassidinae.<ref name=chaboo />

==Tribes and selected genera==
The following list of tribes and selected genera has been collated from ''BioLib''<ref>[https://www.biolib.cz/en/taxon/id12210/ BioLib.cz: subfamily Cassidinae Gyllenhal, 1813] (retrieved 11 August 2025)</ref> and Borowiec & Świętojańska (2014):<ref name="Borowiec">Borowiec, L.; Świętojańska, J. 2014: 2.7.2 Cassidinae Gyllenhal, 1813. Pp. 198–217 in: Leschen et al., 2014</ref>
*Tribes formerly included in Hispinae:
**[Alurnini](/source/Alurnini) {{Au|Chapuis, 1875}}
**[Anisoderini](/source/Anisoderini) {{Au|Chapuis, 1875}}
**[Aproidini](/source/Aproidini) {{Au|Weise, 1911}}<ref name="Borowiec"/><small>Pascoe, 1863</small><ref>Chaboo, C.S., V.E. Sandoval-Gómez, M. Hopper, & G.B. Monteith. 2024. Biology of Aproida balyi Pascoe, 1863 (Coleoptera: Chrysomelidae: Cassidinae: Aproidini) on its host plant, Eustrephus latifolius R. Rr. ex Ker-Gawl (Asparagaceae) in Australia. Insecta Mundi 1074: 1-28.</ref>
**[Arescini](/source/Arescini) {{Au|Chapuis, 1875}}
**[Bothryonopini](/source/Bothryonopini) {{Au|Chapuis, 1875}}
**[Callispini](/source/Callispini) {{Au|Chapuis, 1875}}
**[Callohispini](/source/Callohispini) {{Au|Uhmann, 1960}}
**[Chalepini](/source/Chalepini) {{Au|Weise, 1910}} (= Uroplatini)<ref name="Borowiec"/>
**[Coelaenomenoderini](/source/Coelaenomenoderini) {{Au|Weise, 1911}}
**[Cryptonychini](/source/Cryptonychini) {{Au|Chapuis, 1875}}<ref name="Borowiec"/>
**[Eurispini](/source/Eurispini) {{Au|Chapuis, 1875}}
**[Exothispini](/source/Exothispini) {{Au|Weise, 1911}}
**[Gonophorini](/source/Gonophorini) {{Au|Chapuis, 1875}}
**[Hispini](/source/Hispini) {{Au|Gyllenhal, 1813}}
**[Hispoleptini](/source/Hispoleptini) {{Au|Chapuis, 1875}}
**[Hybosispini](/source/Hybosispini) {{Au|Weise, 1910}}
**[Imatidiini](/source/Imatidiini) {{Au|Hope, 1840}}  (= Cephaloleiini)
**[Leptispini](/source/Leptispini) {{Au|Fairmaire, 1868}}
**[Oncocephalini](/source/Oncocephalini) {{Au|Chapuis, 1875}}
**†Oposispini {{Au|Uhmann, 1939}}
***†''Oposispa'' {{Au|Uhmann, 1939}}
****†''Oposispa scheelei'' {{Au|Uhmann, 1939}} (Baltic amber)
***†''Sucinagonia'' {{Au|Uhmann, 1939}}
****†''Sucinagonia javetana'' {{Au|Uhmann, 1939}} (Baltic amber)
**[Promecothecini](/source/Promecothecini) {{Au|Chapuis, 1875}}
**[Prosopodontini](/source/Prosopodontini) {{Au|Weise, 1910}}
**[Sceloenoplini](/source/Sceloenoplini)  {{Au|Uhmann, 1930}}
**[Spilophorini](/source/Spilophorini) {{Au|Chapuis, 1875}} (= Oediopalpini)
===Tribes formerly included in Cassidinae ''sens. st.''===
*[Aspidimorphini](/source/Aspidimorphini) {{Au|Chapuis, 1875}}; selected genera:
** ''[Aspidimorpha](/source/Aspidimorpha)'' {{Au|Hope, 1840}}
*[Basiprionotini](/source/Basiprionotini) {{Au|Gressitt, 1952}}; selected genera:
** ''[Basiprionota](/source/Basiprionota)'' <small>Chevrolat, 1836</small>
** ''[Epistictina](/source/Epistictina)'' <small>Hincks, 1950</small>
** ''[Metriopepla](/source/Metriopepla)'' <small>Fairmaire, 1882</small>
*[Cassidini](/source/Cassidini) {{Au|Gyllenhal, 1813}}; selected genera:
** ''[Agroiconota](/source/Agroiconota)'' {{Au|Spaeth, 1913}}
** ''[Cassida](/source/Cassida)'' {{Au|Linnaeus, 1758}} - '''type genus'''
** ''[Charidotella](/source/Charidotella)'' {{Au|Weise, 1896}}
** ''[Chiridopsis](/source/Chiridopsis)'' {{Au|Spaeth, 1922}}
*[Cubispini](/source/Cubispini) {{Au|Barber, 1946}}<ref name="Borowiec"/>
** ''[Cubispa](/source/Cubispa)'' {{Au|Barber, 1946}}
*Delocranini {{Au|Spaeth, 1929}}: monogeneric ''[Delocrania](/source/Delocrania)'' {{Au|Guérin, 1844}}
*[Dorynotini](/source/Dorynotini) {{Au|Monrós & Viana, 1949}}; genera:
** ''[Dorynota](/source/Dorynota)'' {{Au|Chevrolat, 1836}}
** ''[Heteronychocassis](/source/Heteronychocassis)'' {{Au|Spaeth, 1915}}
** ''[Omoteina](/source/Omoteina)'' {{Au|Chevrolat, 1836}}
** ''[Paranota](/source/Paranota)'' {{Au|Monros & Viana, 1949}}
** ''[Paratrikona](/source/Paratrikona)'' {{Au|Spaeth, 1923}}
*[Eugenysini](/source/Eugenysini) {{Au|Hincks, 1952}}; all genera:
# ''[Agenysa](/source/Agenysa)'' {{Au|Spaeth, 1905}}
# ''[Eugenysa](/source/Eugenysa)'' {{Au|Chevrolat, 1836}}
# ''[Miocalaspis](/source/Miocalaspis)'' {{Au|Weise, 1899}}
*[Goniocheniini](/source/Goniocheniini) {{Au|Spaeth, 1942}}; all genera:
# ''[Chlamydocassis](/source/Chlamydocassis)'' {{Au|Spaeth, 1952}}
# ''[Goniochenia](/source/Goniochenia)'' {{Au|Weise, 1896}}
# ''[Herissa](/source/Herissa)'' {{Au|Spaeth, 1909}}
# ''[Polychalma](/source/Polychalma)'' {{Au|Barber & Bridwell, 1940}}
# ''[Zeugonota](/source/Zeugonota)'' {{Au|Spaeth, 1913}}
*[Hemisphaerotini](/source/Hemisphaerotini) {{Au|Monrós & Viana, 1951}}
** ''[Hemisphaerota](/source/Hemisphaerota)'' {{Au|Chevrolat, 1836}}
*[Ischyrosonychini](/source/Ischyrosonychini) {{Au|Chapuis, 1875}} (= Physonotini {{Au|Spaeth, 1942}})
** ''[Physonota](/source/Physonota)'' {{Au|Boheman, 1854}}
*[Mesomphaliini](/source/Mesomphaliini) {{Au|Hope, 1840}} (= Stolaini {{Au|Hincks, 1952}})<ref>Borowiec L (1999) A world catalogue of the Cassidinae (Coleoptera: Chrysomelidae). ''Biologica Silesiae'', Wrocław, 476 pp.</ref> selected genera:
** ''[Acromis](/source/Acromis)'' {{Au|Chevrolat, 1836}}
** ''[Chelymorpha](/source/Chelymorpha)'' {{Au|Chevrolat, 1836}}
** ''[Mesomphalia](/source/Mesomphalia)'' {{Au|Hope, 1839}}
** ''[Stolas](/source/Stolas_(beetle))'' {{Au|Billberg, 1820}}
*[Notosacanthini](/source/Notosacanthini) {{Au|Gressitt, 1952}}
# ''[Herminella](/source/Herminella)'' <small>Spaeth, 1913</small>
# ''[Hermosacantha](/source/Hermosacantha)'' <small>Borowiec & Świetojańska, 2014</small>
# ''[Notosacantha](/source/Notosacantha)'' <small>Chevrolat, 1836</small><ref>Monteith, Geoff B., Sandoval-Gómez, Vivian E., & Caroline S. Chaboo. 2021. Natural history of the Australian tortoise beetle Notosacantha dorsalis (Waterhouse, 1877) (Coleoptera: Chrysomelidae: Cassidinae: Notosacanthini) with summary of the genus in Australia. The Australian Entomologist 48(4): 329–354.</ref>
*[Omocerini](/source/Omocerini) {{Au|Hincks, 1952}}; selected genera:
** ''[Omocerus](/source/Omocerus)'' {{Au|Chevrolat, 1835}}
** ''[Polychalca](/source/Polychalca)'' {{Au|Chevrolat, 1836}}

==Gallery==
<gallery>
Deloyala guttata - Mottled Tortoise Beetle.jpg|''[Deloyala guttata](/source/Deloyala_guttata)'', mottled tortoise beetle, showing typical shape and patches of metallic coloration
Tortoise beetle Aslamidium sp (14746391027).jpg|''[Aslamidium](/source/Aslamidium)'' sp.
Hispella testacea - 2012-10-16.webm|''Hispella testacea'' in copula (Hispini)
Leaf beetle (Chalepus sanguinicollis).JPG|Leaf beetle ''Chalepus sanguinicollis'', [Grand Cayman](/source/Grand_Cayman)
Cassidinae, Stolas cf conspersa, Matutu MG.jpg|Blue and orange tortoise-beetle ''[Stolas](/source/Stolas_(beetle)) cf. conspersa'' from Brazil
Plagiometriona clavata.jpg|''[Plagiometriona](/source/Plagiometriona) clavata''
Chrysomelidae - Mesomphalia turrita.jpg|Museum specimen of ''[Mesomphalia](/source/Mesomphalia) turrita'' (Mesomphaliini)
Charidotella sexpunctata larva.jpg|''[Charidotella sexpunctata](/source/Charidotella_sexpunctata)'', larva protected by fecal shield
Charidotella sexpunctata adult.jpg|''Charidotella sexpunctata'', adult
Sceloenopla maculata 123075318.jpg|''[Sceloenopla](/source/Sceloenopla) maculata'' (Sceloenoplini)
</gallery>

==See also==
* [List of Cassidinae genera](/source/List_of_Cassidinae_genera)

==References==
{{Reflist|2}}

==External links==
*{{Commonscat inline|Cassidinae}}
* [http://entomology.si.edu/collections_coleoptera-hispines.html Hispines of the world]
* [http://www.cassidae.uni.wroc.pl/katalog%20internetowy/index.htm Photo atlas and interactive key to world Cassidinae]

{{Taxonbar|from=Q2120908}}

Category:Cassidinae
Category:Polyphaga subfamilies
Category:Articles containing video clips
Category:Taxa named by Leonard Gyllenhaal

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Adapted from the Wikipedia article [Cassidinae](https://en.wikipedia.org/wiki/Cassidinae) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Cassidinae?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.
