Phytophaga

Clade of beetles

Phytophaga
Typical Chrysomeloidea (left) and Curculionoidea (right)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Suborder:	Polyphaga
Infraorder:	Cucujiformia
Clade:	Phytophaga

Phytophaga is a clade of beetles within the infraorder Cucujiformia consisting of the superfamilies Chrysomeloidea and Curculionoidea that are distinctive in the plant-feeding habit combined with the tarsi being pseudotetramerous or cryptopentamerous, where the fourth tarsal segment is typically greatly reduced or hidden by the third tarsal segment. The Cucujoidea are a sister to the Phytophaga.^[1]^[2] In some older literature the term Phytophaga was applied only to the Chrysomeloidea.

Families in the Phytophaga

Phytophaga

Chrysomeloidea

	Megalopodidae

	Cerambycidae

Chrysomelidae

Curculionoidea

	Nemonychidae

	Anthribidae

Belidae

Attelabidae

	Brentidae

	Curculionidae

Phytophaga phylogeny

The diversification of species within the Phytophaga is thought to be associated with the speciation within the Angiosperms. The plant-feeding habit may have been a shift from microfungal, spore-feeding (on strobili and cycads) and saprotrophic habits.^[3] With nearly 125,000 described species they are the second largest phytophagous lineage of insects after the order Lepidoptera.^[4] GH45s are only encoded by the genomes possessed by the Phytophaga beetles. The derived G45s from Phytophaga degrade 3 main substances: amorphous cellulose, xyloglucan and glucomannan. It was also composed of fungal sequences, acquired by gene transfer from fungi.^[5]

References

^ Zhang, Shao-Qian; Che, Li-Heng; Li, Yun; Dan Liang; Pang, Hong; Ślipiński, Adam; Zhang, Peng (2018). "Evolutionary history of Coleoptera revealed by extensive sampling of genes and species". Nature Communications. 9 (1): 205. Bibcode:2018NatCo...9..205Z. doi:10.1038/s41467-017-02644-4. PMC 5768713. PMID 29335414.
^ Marvaldi, Adriana E.; Duckett, Catherine N.; Kjer, Karl M.; Gillespie, Joseph J. (2009). "Structural alignment of 18S and 28S rDNA sequences provides insights into phylogeny of Phytophaga (Coleoptera: Curculionoidea and Chrysomeloidea)". Zoologica Scripta. 38: 63–77. doi:10.1111/j.1463-6409.2008.00360.x. hdl:11336/92765. S2CID 84345520.
^ Farrell, B. D. (1998-07-24). ""Inordinate Fondness" Explained: Why Are There So Many Beetles?". Science. 281 (5376): 555–559. doi:10.1126/science.281.5376.555. PMID 9677197.
^ Haddad, Stephanie; McKenna, Duane D. (2016). "Phylogeny and evolution of the superfamily Chrysomeloidea (Coleoptera: Cucujiformia)". Systematic Entomology. 41 (4): 697–716. doi:10.1111/syen.12179. S2CID 87055542.
^ Busch, André; Danchin, Etienne G. J.; Pauchet, Yannick (2019-05-10). "Functional diversification of horizontally acquired glycoside hydrolase family 45 (GH45) proteins in Phytophaga beetles". BMC Evolutionary Biology. 19 (1): 100. doi:10.1186/s12862-019-1429-9. ISSN 1471-2148. PMC 6509783. PMID 31077129.

Extant Coleoptera families

Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Subclass: Pterygota
Infraclass: Neoptera
Superorder: Holometabola

Suborder Archostemata
Crowsoniellidae (Crowsoniella relicta) Cupedidae (reticulated beetles) Jurodidae (Sikhotealinia zhiltzovae) Micromalthidae (telephone-pole beetle) Ommatidae

Suborder Adephaga

Extant families	Amphizoidae (trout-stream beetles) Aspidytidae Carabidae (ground beetles) Cicindelidae (tiger beetles) Dytiscidae (predaceous diving beetles) Gyrinidae (whirligig beetles) Haliplidae (crawling water beetles) Hygrobiidae Meruidae (Meru phyllisae) Noteridae (burrowing water beetles) Trachypachidae (false ground beetles)

Suborder Myxophaga
Hydroscaphidae (skiff beetles) Lepiceridae Sphaeriusidae Torridincolidae

Suborder Polyphaga

Bostrichiformia

Bostrichoidea	Bostrichidae (auger beetles) Dermestidae (skin beetles) Endecatomidae Jacobsoniidae (Jacobson's beetles) Nosodendridae (wounded-tree beetles) Ptiniidae (furniture beetles, death watch beetles, spider beetles)
Derodontoidea	Derodontidae (tooth-necked fungus beetles)

Cucujiformia

Chrysomeloidea	Cerambycidae (longhorn beetles) Chrysomelidae (leaf beetles) Disteniidae Megalopodidae Orsodacnidae Oxypeltidae Vesperidae
Cleroidea	Acanthocnemidae (Acanthocnemus nigricans) Biphyllidae (false skin beetles) Byturidae (fruitworm beetles) Chaetosomatidae Cleridae (checkered beetles) Lophocateridae Mauroniscidae Melyridae (soft-wing flower beetles) Metaxinidae (Metaxina ornata) Peltidae Phloiophilidae (Phloiophilus edwardsi) Phycosecidae Prionoceridae Protopeltidae Rentoniidae Rhadalidae Thanerocleridae Thymalidae Trogossitidae (bark-gnawing beetles)
Coccinelloidea	Akalyptoischiidae Alexiidae Anamorphidae Bothrideridae (dry bark beetles) Cerylonidae (minute bark beetles) Coccinellidae (lady beetles, or God's cows) Corylophidae (minute fungus beetles) Discolomatidae Endomychidae (handsome fungus beetles) Eupsilobiidae Euxestidae (well polished beetles) Latridiidae (minute brown scavenger beetles) Murmidiidae Mycetaeidae Teredidae
Cucujoidea	Agapythidae (Agapytho foveicollis) Boganiidae Cavognathidae Cryptophagidae (silken fungus beetles) Cucujidae (flat bark beetles) Cybocephalidae Cyclaxyridae Erotylidae (pleasing fungus beetles) Helotidae Hobartiidae Kateretidae (short-winged flower beetles) Laemophloeidae (lined flat bark beetles) Lamingtoniidae (Lamingtonium binnaberrense) Monotomidae (root-eating beetles) Myraboliidae Nitidulidae (sap beetles) Passandridae (parasitic flat bark beetles) Phalacridae (shining flower beetles) Phloeostichidae Priasilphidae Propalticidae Protocucujidae Silvanidae (silvanid flat bark beetles) Smicripidae (palmetto beetles) Sphindidae (dry-fungus beetles) Tasmosalpingidae
Curculionoidea (weevils)	Anthribidae (fungus weevils) Attelabidae (leaf-rolling weevils) Belidae (primitive weevils) Brentidae (straight snout weevils, New York weevil) Caridae Curculionidae (true weevils, bark beetles, ambrosia beetles) Nemonychidae (pine flower weevils)
Lymexyloidea	Lymexylidae (ship-timber beetles)
Tenebrionoidea	Aderidae (ant-like leaf beetles) Anthicidae (ant-like flower beetles) Archeocrypticidae (cryptic fungus beetles) Boridae (conifer bark beetles) Chalcodryidae Ciidae (minute tree-fungus beetles) Melandryidae (false darkling beetles) Meloidae (blister beetles) Mordellidae (tumbling flower beetles) Mycetophagidae (hairy fungus beetles) Mycteridae (palm and flower beetles) Oedemeridae (false blister beetle) Perimylopidae, or Promecheilidae Prostomidae (jugular-horned beetles) Pterogeniidae Pyrochroidae (fire-coloured beetles) Pythidae (dead log bark beetles) Ripiphoridae (wedge-shaped beetles) Salpingidae (narrow-waisted bark beetles) Scraptiidae (false flower beetles) Stenotrachelidae (false longhorn beetles) Synchroidae (synchroa bark beetles) Tenebrionidae (darkling beetles) Tetratomidae (polypore fungus beetles) Trictenotomidae Ulodidae Zopheridae (ironclad beetles, cylindrical bark beetles)

Elateriformia

Buprestoidea	Buprestidae (jewel beetles, or metallic wood-boring beetles) Schizopodidae
Byrrhoidea	Byrrhidae (pill beetles) Callirhipidae (cedar beetles) Chelonariidae (turtle beetles) Cneoglossidae Dryopidae (long-toed water beetles) Elmidae (riffle beetles) Eulichadidae (forest stream beetles) Heteroceridae (variegated mud-loving beetles) Limnichidae (minute mud beetles) Lutrochidae (travertine beetles) Psephenidae (water-penny beetles) Ptilodactylidae
Dascilloidea	Dascillidae (soft bodied plant beetles) Rhipiceridae (cicada beetle, cicada parasite beetles)
Elateroidea	Artematopodidae (soft-bodied plant beetles) Brachypsectridae (Texas beetles) Cantharidae (soldier beetles) Cerophytidae (rare click beetles) Elateridae (click beetles) Eucnemidae (false click beetles) Jurasaidae Lampyridae (fireflies) Lycidae (net-winged beetles) Omethidae (false fireflies, long-lipped beetles) Phengodidae (glowworm beetles) Rhagophthalmidae Sinopyrophoridae Throscidae (false metallic wood-boring beetles)
Rhinorhipoidea	Rhinorhipidae (Rhinorhipus tamborinensis)
Scirtoidea	Clambidae Decliniidae (Declinia relicta) Eucinetidae (plate-thigh beetles) Scirtidae

Scarabaeiformia

Scarabaeoidea

Belohinidae (Belohina inexpectata)
Bolboceratidae
Diphyllostomatidae (false stag beetles)
Geotrupidae (dor beetles)
Glaphyridae (bumble bee scarab beetles)
Glaresidae (enigmatic scarab beetles)
Hybosoridae (scavenger scarab beetles)
Lucanidae (stag beetles)
Ochodaeidae (sand-loving scarab beetles)
Passalidae (betsy beetles)
Pleocomidae (rain beetles)
Scarabaeidae (scarabs)
Trogidae (hide beetles)

Staphyliniformia

Histeroidea	Histeridae (clown beetles) Sphaeritidae (false clown beetles) Synteliidae
Hydrophiloidea	Epimetopidae Georissidae (minute mud-loving beetles) Helophoridae Hydrochidae Hydrophilidae (water scavenger beetles) Spercheidae
Staphylinoidea	Agyrtidae (primitive carrion beetles) Hydraenidae Leiodidae (round fungus beetles) Ptiliidae (feather-winged beetles) Silphidae (carrion beetles) Staphylinidae (rove beetles)