Tortoise Beetle
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The Cassidinae (tortoise and leaf-mining beetles) are a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
of the leaf beetles, or
Chrysomelidae The beetle family Chrysomelidae, commonly known as leaf beetles, includes over 37,000 (and probably at least 50,000) species in more than 2,500 genera, making it one of the largest and most commonly encountered of all beetle families. Numerous s ...
. 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 The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
Hispini, are notable for the spiny outgrowths to the
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on e ...
and
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
.


Description

The "cassidoids" have a rounded outline with the edges of the
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on e ...
and
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
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 A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
. 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 A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
that are ecological diverse as
leaf miners A leaf miner is any one of numerous species of insects in which the larval stage lives in, and eats, the leaf tissue of plants. The vast majority of leaf-mining insects are moths ( Lepidoptera), sawflies ( Symphyta, a paraphyletic group whi ...
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 In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have molted. The exuviae of an animal can be important to biologists as they can often be ...
into a discrete structure held on the caudal processes (or, urogomphi). Fecal shields are thought to provide protection from the sun, wind, heat, predators, parasites, and/or
parasitoids In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
though experimental evidence is mixed. A few species in two closely related tribes ( Mesomphaliini and Eugenysini, putative
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
) show
paternal care In biology, paternal care is parental investment provided by a male to his own offspring. It is a complex social behavior in vertebrates associated with animal mating systems, life history traits, and ecology. Paternal care may be provided in co ...
of
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
.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. These species can be viewed as
subsocial Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother was ...
, with evidence pointing to there being two evolutionary origins of subsociality within this one lineage.


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, 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. 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. 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. ''BioLib'' currently (March 2024) includes the following tribes:


Aspidimorphini

Auth. Chapuis, 1875; genera: # '' Aspidimorpha'' Hope, 1840 # '' Conchyloctenia'' Spaeth, 1902 # '' Hybosinota'' Spaeth, 1909 # '' Laccoptera'' Boheman, 1855 # '' Mahatsinia'' Weise, 1910 # '' Nilgiraspis'' Spaeth, 1932


Basiprionotini

Auth. Gressitt, 1952; genera: # '' Androya'' Spaeth, 1911 # '' Basiprionota'' Chevrolat, 1837 # '' Cassidopsis'' Fairmaire, 1899 # '' Craspedonta'' Chevrolat, 1837 # '' Epistictina'' Hincks, 1950 # '' Megapyga'' Boheman, 1850 # '' Metriopepla'' Fairmaire, 1882 # '' Pseudandroya'' Spaeth, 1952 ; Cassidini ; Delocranini ; Dorynotini ; Eugenysini ; Goniocheniini ; Hemisphaerotini


Imatidiini

Auth. Hope, 1840 (= Cephaloleiini); genera: # '' Aslamidium'' Borowiec, 1984 # '' Calliaspis'' Dejean, 1837 # '' Cyclantispa'' Sekerka, 2014 # '' Demotispa'' Baly, 1858 # '' Imatidium'' Fabricius, 1801 # '' Katkispa'' Sekerka, 2014 # '' Lechispa'' Sekerka, 2014 # '' Parentispa'' Sekerka, 2014 # '' Parimatidium'' Spaeth, 1938 # '' Pseudostilpnaspis'' Borowiec, 2000 # '' Spaethaspis'' Hincks, 1952 # '' Stilpnaspis'' Weise, 1905 # '' Weiseispa'' Sekerka, 2014 # '' Windsorispa'' Sekerka, 2014


Notosacanthini

Auth. Gressitt, 1952 # '' Herminella'' Spaeth, 1913 # '' Hermosacantha'' Borowiec & Świetojańska, 2014 # '' Notosacantha'' Chevrolat, 1837 ; Omocerini ; Physonotini = synonyms Ischyrosonychini and Physonotini ; Spilophorini (= Oediopalpini)


Stolaini

Auth. Hincks, 1952 # '' Acromis'' Chevrolat, 1837 # '' Amythra'' Spaeth, 1913 # '' Anacassis'' Spaeth, 1913 # '' Anepsiomorpha'' Spaeth, 1913 # '' Botanochara'' Dejean, 1837 # '' Chelymorpha'' Chevrolat, 1837 # '' Convexocoleus'' Shin, 2013 # '' Cyrtonota'' Chevrolat, 1837 # '' Echoma'' Chevrolat, 1837 # '' Elytrogona'' Chevrolat, 1837 # ''
Eutheria Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...
'' Spaeth, 1909 # '' Hilarocassis'' Spaeth, 1913 # '' Mesomphalia'' Hope, 1839 # '' Nebraspis'' Spaeth, 1913 # '' Ogdoecosta'' Spaeth, 1909 # '' Omaspides'' Chevrolat, 1837 # '' Paraselenis'' Spaeth, 1913 # '' Phytodectoidea'' Spaeth, 1909 # '' Poecilaspidella'' Spaeth, 1913 # '' Stoiba'' Spaeth, 1909 # ''
Stolas Stolas may refer to: *Stolas (demon), a demon in the ''Ars Goetia'' *Stolas (band), an American post-hardcore band *'' Stolas: Book of Angels Volume 12'', a 2009 Masada Quintet album * ''Stolas'' (beetle), a genus of beetles in the family Chrysomel ...
'' Billberg, 1820 # '' Terpsis'' Spaeth, 1913 # '' Trilaccodea'' Spaeth, 1902 # '' Xenicomorpha'' Spaeth, 1913 # '' Zatrephina'' Spaeth, 1909 In addition, Borowiec & Świętojańska (2014)''Borowiec, L.; Świętojańska, J. 2014: 2.7.2 Cassidinae Gyllenhal, 1813. Pp. 198–217 in: Leschen et al., 2014 included: ; Aproidini # '' Aproida'' Pascoe, 1863 ; Chalepini (= Uroplatini) ; Cryptonychini ;
Cubispini Cubispini is a tribe of leaf beetles from the Neotropical realm. It is classified within either the Eumolpinae or the Cassidinae. It was established by Francisco Monrós in 1954 to contain the genus ''Cubispa'', which was originally placed in the ...
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, priority is given to the name Cassidinae. Deloyala guttata - Mottled Tortoise Beetle.jpg, ''
Deloyala guttata ''Deloyala guttata'', the mottled tortoise beetle, is a species of tortoise beetle in the family Chrysomelidae. It is found in the Caribbean, Central America, North America, and South America. Larva carry shields made from exuviae and frass ...
'', mottled tortoise beetle, showing typical shape and patches of metallic coloration Hispella testacea - 2012-10-16.webm, ''Hispella testacea'' in copula (Hispini) Leaf beetle (Chalepus sanguinicollis).JPG, Leaf beetle ''Chalepus sanguinicollis'',
Grand Cayman Grand Cayman is the largest of the three Cayman Islands and the location of the territory's capital, George Town, Cayman Islands, George Town. In relation to the other two Cayman Islands, it is approximately 75 miles (121 km) southwest of L ...
Cassidinae, Stolas cf conspersa, Matutu MG.jpg, Blue and orange tortoise-beetle ''Stolas cf. conspersa'' from Brazil Tortoise beetle (Stolas lebasii).jpg, ''Stolas lebasii'',
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
Tortoise beetle (Stolas lebasii) 2.jpg, ''Stolas lebasii'',
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
Plagiometriona clavata.jpg, '' Plagiometriona clavata'' Charidotella sexpunctata larva.jpg, '' Charidotella sexpunctata'', larva protected by fecal shield Charidotella sexpunctata adult.jpg, ''Charidotella sexpunctata'', adult


See also

* List of Cassidinae genera


References


External links


Hispines of the world


{{Taxonbar, from=Q2120908 Polyphaga subfamilies Articles containing video clips Taxa named by Leonard Gyllenhaal