Cassidinae

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.

Cassidinae
Platypria sp.
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Suborder:	Polyphaga
Infraorder:	Cucujiformia
Family:	Chrysomelidae
Subfamily:	Cassidinae Gyllenhal, 1813
Tribes[1]
(m): monogeneric Alurnini Anisoderini Aproidini (m) Arescini Aspidimorphini Basiprionotini Bothryonopini Callispini Callohispini (m) Cassidini Chalepini (= Uroplatini) Coelaenomenoderini Cryptonychini Cubispini Delocraniini (m) Dorynotini Eugenysini Eurispini Exothispini (m) Goniocheniini Gonophorini Hemisphaerotini Hispini Hispoleptini (m) Hybosispini (m) Imatidiini (= Cephaloleiini) Ischyrosonychini (= Physonotini) Leptispini Mesomphaliini (= Stolaini) Notosacanthini Omocerini Oncocephalini Promecothecini Prosopodontini (m) Sceloenoplini Spilophorini (= Oediopalpini)

Description

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 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 ecological diverse as leaf miners 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 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 though experimental evidence is mixed.^[2][3]

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

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.^[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]

The the following list of tribes and selected genera has been collated from BioLib^[9] and Borowiec & Świętojańska (2014):^[1]

Alurnini Chapuis, 1875

Anisoderini Chapuis, 1875

Aproidini Weise, 1911^[1]

1. Aproida Pascoe, 1863^[10]

Aspidimorphini

Auth. Chapuis, 1875; selected genera:

• Aspidimorpha Hope, 1840
• Laccoptera Boheman, 1855

Basiprionotini

Auth. Gressitt, 1952; selected genera:

• Androya Spaeth, 1911
• Basiprionota Chevrolat, 1837
• Epistictina Hincks, 1950
• Metriopepla Fairmaire, 1882

Bothryonopini

Auth. Chapuis, 1875; all genera:

1. Bothryonopa Guérin-Méneville, 1840
2. Macrispa Baly, 1858
3. Neodownesia Gressitt, 1953
4. Pistosia Weise, 1905
5. Wallacea (beetle) Baly, 1859

Callispini

Auth. Chapuis, 1875; selected genera:

• Amblispa Baly, 1858
• Callispa Baly, 1858

Cassidini

Auth. Gyllenhal, 1813; selected genera:

• Agroiconota Spaeth, 1913
• Cassida Linnaeus, 1758 - type genus
• Charidotella Weise, 1896
• Chiridopsis Spaeth, 1922

Chalepini Weise, 1910 (= Uroplatini)^[1]

• Baliosus Weise, 1905
• Chalepus Thunberg, 1805

Coelaenomenoderini

Auth. Weise, 1911; genera:

1. Balyana Péringuey, 1898
2. Bulolispa Gressitt & Samuelson, 1990
3. Coelaenomenodera Blanchard, 1845
4. Cyperispa Gressitt, 1957
5. Enischnispa Gressitt, 1957
6. Heterrhachispa Gressitt, 1957
7. Javeta Baly, 1858
8. Pharangispa Maulik, 1929
9. Rhabdotohispa Maulik, 1913

Cryptonychini Chapuis, 1875^[1]

Cubispini Barber, 1946^[1]

Delocranini Spaeth, 1929

1. Delocrania Guérin, 1844

Dorynotini

Auth: Monrós & Viana, 1949; genera:

1. Dorynota Chevrolat, 1837
2. Heteronychocassis Spaeth, 1915
3. Omoteina Chevrolat, 1837
4. Paranota Monros & Viana, 1949
5. Paratrikona Spaeth, 1923

Eugenysini

Auth: Hincks, 1952; genera:

1. Agenysa Spaeth, 1905
2. Eugenysa Chevrolat, 1837
3. Miocalaspis Weise, 1899

Eurispini

Auth: Chapuis, 1875; genera:

1. Eurispa Baly, 1858
2. Leucispa Chapuis, 1875
3. Squamispa Maulik, 1928

Goniocheniini

Auth: Spaeth, 1942; genera:

1. Chlamydocassis Spaeth, 1952
2. Goniochenia Weise, 1896
3. Herissa Spaeth, 1909
4. Polychalma Barber & Bridwell, 1940
5. Zeugonota Spaeth, 1913

Hemisphaerotini Monrós & Viana, 1951

• Hemisphaerota Chevrolat, 1837

Gonophorini

Auth: Chapuis, 1875; genera:

1. Agoniella Weise, 1911
2. Agonita Strand, 1942
3. Aspidispa Baly, 1869
4. Downesia Baly, 1858
5. Gonophora Chevrolat, 1837
6. Klitispa Uhmann, 1940
7. Micrispa Gestro, 1897
8. Monagonia Uhmann, 1931
9. Wallacispa Uhmann, 1940

Hispini

Auth. Gyllenhal, 1813; genera:

1. Acmenychus Weise, 1905
2. Asamangulia Maulik, 1915
3. Callanispa Uhmann, 1959
4. Cassidispa Gestro, 1899
5. Chrysispa Weise, 1897
6. Dactylispa Weise, 1897
7. Dicladispa Gestro, 1897
8. Dorcathispa Weise, 1901
9. Hispa Linnaeus, 1767
10. Hispellinus Weise, 1897
11. Jambhala (beetle) Würmli, 1975
12. Lesageana L. Medvedev, 2003
13. Phidodontina Uhmann, 1938
14. Philodonta Weise, 1906
15. Platypria Guérin-Ménéville, 1840
16. Pleurispa Weise, 1901
17. Polyconia Weise, 1905
18. Pseudispella Kraatz, 1895
19. Rhadinosa Weise, 1905
20. Rhodtrispa Chen & Tan, 1964
21. Sinispa Uhmann, 1940
22. Thomispa Würmli, 1975
23. Thoracispa Chapuis, 1875
24. Trichispa Chapuis, 1875
25. Unguispa Uhmann, 1954

Imatidiini

Auth. Hope, 1840 (= Cephaloleiini); genera:

1. Aslamidium Borowiec, 1984
2. Calliaspis Dejean, 1837
3. Cyclantispa Sekerka, 2014
4. Demotispa Baly, 1858
5. Imatidium Fabricius, 1801
6. Katkispa Sekerka, 2014
7. Lechispa Sekerka, 2014
8. Parentispa Sekerka, 2014
9. Parimatidium Spaeth, 1938
10. Pseudostilpnaspis Borowiec, 2000
11. Spaethaspis Hincks, 1952
12. Stilpnaspis Weise, 1905
13. Weiseispa Sekerka, 2014
14. Windsorispa Sekerka, 2014

Ischyrosonychini Chapuis, 1875 (synonym Physonotini Spaeth, 1942)

Leptispini

Auth. Fairmaire, 1868; genera:

1. Leptispa Baly, 1858
2. Ovotispa L. Medvedev, 1992

Mesomphaliini

Auth. Hope, 1840 (synonym Stolaini Hincks, 1952)^[11]

1. Acromis Chevrolat, 1837
2. Amythra Spaeth, 1913
3. Anacassis Spaeth, 1913
4. Anepsiomorpha Spaeth, 1913
5. Botanochara Dejean, 1837
6. Chelymorpha Chevrolat, 1837
7. Convexocoleus Shin, 2013
8. Cyrtonota Chevrolat, 1837
9. Echoma Chevrolat, 1837
10. Elytrogona Chevrolat, 1837
11. Eutheria Spaeth, 1909
12. Hilarocassis Spaeth, 1913
13. Mesomphalia Hope, 1839
14. Nebraspis Spaeth, 1913
15. Ogdoecosta Spaeth, 1909
16. Omaspides Chevrolat, 1837
17. Paraselenis Spaeth, 1913
18. Phytodectoidea Spaeth, 1909
19. Poecilaspidella Spaeth, 1913
20. Stoiba Spaeth, 1909
21. Stolas Billberg, 1820
22. Terpsis Spaeth, 1913
23. Trilaccodea Spaeth, 1902
24. Xenicomorpha Spaeth, 1913
25. Zatrephina Spaeth, 1909

Notosacanthini

Auth. Gressitt, 1952

1. Herminella Spaeth, 1913
2. Hermosacantha Borowiec & Świetojańska, 2014
3. Notosacantha Chevrolat, 1837^[12]

Omocerini Hincks, 1952

Selected genera:

• Omocerus Chevrolat, 1835
• Polychalca Chevrolat, 1837

Oncocephalini

Auth. Chapuis, 1875; all genera:

1. Chaeridiona Baly, 1869
2. Oncocephala Agassiz, 1846
3. Prionispa Chapuis, 1875

Promecothecini

Auth. Weise, 1911

1. Promecispa Weise, 1909
2. Promecotheca Blanchard, 1853

Sceloenoplini

Auth. Uhmann, 1930

1. Acentroptera Guérin-Méneville, 1844
2. Ocnosispa Weise, 1910
3. Pseudispa Chapuis, 1875
4. Sceloenopla Dejean, 1835
5. Serratispa Staines, 2002

Spilophorini Chapuis, 1875 (= Oediopalpini)

• Calyptocephala Chevrolat, 1836

Gallery

• 
  Deloyala guttata, mottled tortoise beetle, showing typical shape and patches of metallic coloration
• Hispella testacea in copula (Hispini)
• 
  Leaf beetle Chalepus sanguinicollis, Grand Cayman
• 
  Blue and orange tortoise-beetle Stolas cf. conspersa from Brazil
• 
  Stolas lebasii, Panama
• 
  Stolas lebasii, Panama
• 
  Plagiometriona clavata
• 
  Charidotella sexpunctata, larva protected by fecal shield
• 
  Charidotella sexpunctata, adult

See also

• List of Cassidinae genera