User:Monster Iestyn/Insect systematics

Notes for improving various articles for higher-level insect groups.

Pterygota

Main article: Pterygota

Pterygota	Ephemeroptera (mayflies) Odonata (dragonflies and damselflies) Neoptera

Palaeoptera problem

The relationships between the three main clades of Pterygota – Odonata, Ephemeroptera and Neoptera – are currently unresolved. Three hypotheses have been proposed:

Palaeoptera hypothesis:

Pterygota	Palaeoptera Ephemeroptera Odonata Neoptera

Metapterygota hypothesis:

Pterygota	Ephemeroptera Metapterygota Odonata Neoptera

Chiastomyaria hypothesis:

Pterygota	Odonata Chiastomyaria Ephemeroptera Neoptera

Hypothesis	Hypothesis 1 (Palaeoptera)	Hypothesis 2 (Metapterygota)	Hypothesis 3 (Chiastomyaria)
Cladogram	Pterygota Palaeoptera Ephemeroptera Odonata Neoptera	Pterygota Ephemeroptera Metapterygota Odonata Neoptera	Pterygota Odonata Chiastomyaria Ephemeroptera Neoptera
Possible synapomorphies	inability to flex wings back over the abdomen similar wing base sclerites bristle-like antennae aquatic larvae	suppression of imaginal molts absence of the axillar-furcal muscle basalar-sternal muscles missing terminal filum secondary mandibular articulation as ball-and-socket joint modified mandibular muscles no subimago	direct sperm transfer differences in wing base structure appearance and function indirect flight musculature copulation with aedeagus

Exopterygota

Main article: Exopterygota

I haven't found a source explaining what's going on here yet, but there seems to be two different definitions for this group:

• Palaeoptera + Polyneoptera + Paraneoptera, i.e. all winged insects where wings develop outside the body (sources include ). This was also the original definition used by Sharp, 1898, except Sharp placed Mallophaga and Anoplura (chewing and sucking lice respectively) instead in "Anapterygota" (secondarily wingless insects), which also included Siphonaptera (fleas).
• Polyneoptera + Paraneoptera, i.e. A division of Neoptera where wings develop outside the body.

To put it in other words, sometimes Odonata (dragonflies and damselflies) and Ephemeroptera (mayflies) are included in Exopterygota, and sometimes they are not...

Polyneoptera

Main article: Polyneoptera

One of the major groups of Neoptera.

Also known as 'lower Neoptera'

has incomplete metamorphosis (Polyneoptera was formerly part of Exopterygota)

evidence according to Kjer et al. (2016): tegmina, enlarged anal field (vannus) of hind wing [see Insect wing#Fields], euplantulae

'Kristensen's comb' refers to the polytomy of Kristensen (1981), in which a polytomy is given for the "lower Neoptera" (= most of Polyneoptera, with Zoraptera placed in Paraneoptera)

Proposed clades in Polyneoptera:

• Dictyoptera (= Blattodea [including Isoptera] + Mantodea] [supported in consensus according to Kjer et al. (2016)]
• Eukinolabia (= Embioptera + Phasmatodea)
• Haplocercata (= Dermaptera + Zoraptera)
• Mystroptera (= Embioptera  +  Zoraptera) [rejected by Letsch & Simon (2013)]
• Paurometabola (= Polyneoptera excluding Plecoptera) [strongly rejected according to Kjer et al. (2016)]
• Orthopterida (= Orthoptera  +  Phasmatodea) [rejected by Letsch & Simon (2013)]
• Xenonomia (= Grylloblattodea + Mantophasmatodea) [= Notoptera] [supported in consensus according to Kjer et al. (2016)]

Kjer et al. (2016) phylogeny

Full phylogeny (Figure 5):

	Chelicerata
Mandibulata	Myriapoda Pancrustacea "Crustacea" "Crustacea" "Crustacea" Hexapoda Elipura Collembola Protura Cercophora Diplura Insecta Archaeognatha Dicondylia Zygentoma Pterygota Odonata Ephemeroptera Neoptera Polyneoptera Haplocercata Zoraptera Dermaptera Plecoptera Orthoptera Xenonomia Mantophasmatodea Grylloblattodea Eukinolabia Embioptera Phasmatodea Dictyoptera Mantodea Blattodea Eumetabola Condylognatha Thysanoptera Hemiptera Psocodea Holometabola Hymenoptera Aparaglossata Neuropteroidea Neuropterida Raphidioptera Megaloptera Neuroptera Coleopterida Coleoptera Strepsiptera Mecopterida Antliophora Diptera Siphonaptera Mecoptera Amphiesmenoptera Trichoptera Lepidoptera

Hexapods only (with experimental coloring):

Hexapoda

Elipura Collembola Protura Cercophora Diplura Insecta Archaeognatha Dicondylia Zygentoma Pterygota Odonata Ephemeroptera Neoptera Polyneoptera Haplocercata Zoraptera Dermaptera Plecoptera Orthoptera Xenonomia Mantophasmatodea Grylloblattodea Eukinolabia Embioptera Phasmatodea Dictyoptera Mantodea Blattodea Eumetabola Condylognatha Thysanoptera Hemiptera Psocodea Holometabola Hymenoptera Aparaglossata Neuropteroidea Neuropterida Raphidioptera Megaloptera Neuroptera Coleopterida Coleoptera Strepsiptera Mecopterida Antliophora Diptera Siphonaptera Mecoptera Amphiesmenoptera Trichoptera Lepidoptera

Paoliida

Main article: Paoliida

Extinct group of insects from the Palaeozoic. Historically their position and composition was controversial, but recent studies have resolved it as the sister group to Dictyoptera.

Alternatively it has been placed in its own superorder Paoliidea in infraclass Gryllones (= Polyneoptera). (I'm assuming this isn't widely accepted, given the use of a different (Russian?) insect classification and nomenclature based on Rasnitsyn & Quicke (2002)'s History of Insects, where e.g. Scarabaeona = Pterygota, Scarabaeones = all winged insects except Gryllones (= Neoptera + Palaeoptera).)

Sroka et al. (2015) phylogeny

Full phylogeny (Figure 11):

Dicondylia

Zygentoma Paranotalia †Carbotriplurida Pterygota Neopterygota †Palaeodictyopterida †Spilapteridae †Thesoneuridae †Homoiopteridae †Eugerionidae †Dictyoneuridae †Megasecoptera †Permothemistida †Diaphanopterodea Neoptera Hydropalaeoptera †Bojophlebiidae Euhydropalaeoptera Odonatoptera Panephemeroptera †Syntonopteroidea †Miracopteridae †Syntonopteridae †Lithoneuridae †Triplosobidae Ephemerida †Permoplectoptera †Misthodotidae †Protereismatidae †Cretereismatidae †Litophlebioidea †Litophlebiidae †Tintorinidae Heptabranchia †Coxoplectoptera †Mickoleitiidae †Palingeniopsidae †Mesephemeridae Ephemeroptera Siphluriscidae †Hexagenitidae Siphlonuridae

Coleoptera phylogeny

Phylogeny based on Boudinot et al. (2022) (Figure 8?): and Goczał & Beutel (2023)

Coleopterida	Strepsiptera Coleoptera †Tshekardocoleidae Mesocoleoptera †Permocupedidae Metacoleoptera †Rhombocoleidae, †Asiocoleidae, †Triadocupedidae, †Taldycupedidae †Ademosynidae, †Schizophoridae Archostemata Micromalthidae Ommatidae Cupedidae Myxophaga Adephaga Polyphaga

(See also Figure 2 in https://doi.org/10.1111/syen.12623 )

Chrysomeloidea phylogeny

From Haddad et al. (2018) (nuclear genome):

Chrysomeloidea	Chrysomelidae Megalopodidae (paraphyletic) + Oxypeltidae Orsodacnidae Vesperidae Disteniidae Cerambycidae

Nucleotide data Chrysomeloidea Chrysomelidae Megalopodidae (paraphyletic) + Oxypeltidae Orsodacnidae Vesperidae Disteniidae Cerambycidae

Amino acid data Chrysomeloidea Chrysomelidae Megalopodidae (paraphyletic) + Oxypeltidae Vesperidae Orsodacnidae Disteniidae Cerambycidae

From Nie et al. (2020) (mitochondrial genome):

Chrysomeloidea	Chrysomelidae 'lamiine' clade Lepturinae (including Necydalinae) Lamiinae Spondylidinae 'cerambycine' clade Dorcasominae Cerambycinae Prioninae (including Parandrinae) 'Dis‑Ves' clade Disteniidae Vesperidae 'Oxy‑Meg‑Ors' clade Orsodacnidae Megalopodidae Oxypeltidae Cerambycidae s.s.

Chrysomelidae phylogeny

From Nie et al. (2019) (mitochondrial genome):

Chrysomelidae	'eumolpine' clade Cryptocephalinae Lamprosomatinae Eumolpinae Cassidinae Spilopyrinae 'sagrine' clade Criocerinae Donaciinae Bruchinae Sagrinae Synetinae 'chrysomeline' clade Galerucinae Alticinae Chrysomelinae (excluding Timarcha) Timarcha

Xenonomia

After figures and supplementary material of

Notes from Appendix S1:

• Tenuivenaptera Béthoux, Cui & Wipfler, 2024 = All species in Dataset S1 + Blattogryllopterida
• Blattogryllopterida Cui, 2012 = Blattogryllidae sensu Storozhenko & Aristov (2014) (not monophyletic according to main article!!) + Icaroptera
• Icaroptera Béthoux, Cui & Wipfler, 2024 = extant Grylloblattidae

Figure 2F, Figure S3:

(Note: is Aristovia being called Burmablattogryllus in Figure S3??)

(Excluding Mantophasmatodea)

Tenuivenaptera	†Fabrecia Blattogryllopterida †Permoblattogryllus †Plesioblattogryllus †Aristovia Icaroptera Gallosiana

(Including Mantophasmatodea, possible placements)

Xenonomia Mantophasmatodea Karoophasma Austrophasma Grylloblattodea? †Fabrecia †Permoblattogryllus †Plesioblattogryllus †Aristovia Gallosiana

Xenonomia †Fabrecia Mantophasmatodea Karoophasma Austrophasma Grylloblattodea? †Permoblattogryllus †Plesioblattogryllus †Aristovia Gallosiana

Xenonomia †Fabrecia †Permoblattogryllus Mantophasmatodea Karoophasma Austrophasma Grylloblattodea? †Plesioblattogryllus †Aristovia Gallosiana

Odonata phylogeny

Odonata according to Bechly:

Odonata

Zygoptera (see below) Epiproctophora †Isophlebioptera †Euthemistidae (incertae sedis) †Parazygoptera (sedis mutabilis) †Sphenophlebiidae †Euparazygoptera †Asiopteridae †Triassolestoidea †Cyclothemistidae †Triassolestidae †Selenothemistidae (sedis mutabilis) †Isophlebiida (sedis mutabilis) †Archithemistidae †Isophlebioidea †Campterophlebiidae †Isophlebiidae Euepiproctophora Epiophlebiidae Anisopteromorpha †Erichschmidtiidae (incertae sedis) †Heterophlebioptera †Liassophlebiidae †Heterophlebiidae Trigonoptera †Stenophlebioptera †Gondvanogomphidae †Stenophlebiidae Pananisoptera †Liassogomphidae †Juragomphidae Neoanisoptera †Aeschnidiidae Anisoptera (see below)

Anisoptera

(Grey notes are my attempts to match the extant families in Bechly's classification with the equivalent families/subfamilies/etc under the World Odonata List classification)

Anisoptera

Petalurida †Protolindeniidae Petalurodea †Cretapetaluridae Petaluroidea †Aktassiidae Petaluridae (= Petaluridae) Euanisoptera Exophytica Cavilabiata Cordulegastrida Zoraenidae (= Cordulegastridae) Cordulegastridae (= Cordulegastridae) Cristotibiata Neopetaliidae (= Neopetaliidae) Brachystigmata †Nannogomphidae Eubrachystigmata †Hemeroscopidae Neobrachystigmata Chlorogomphida †Araripechlorogomphidae Chlorogomphoidea Chloropetaliidae (= Chlorogomphidae) Chlorogomphidae (= Chlorogomphidae) Paneurypalpidomorpha †Juracorduliidae Eurypalpidomorpha †Valdicordulioidea †Valdicorduliidae †Araripephlebiidae Eurypalpidiformia †Eocorduliidae Paneurypalpida †Araripelibellulidae Eurypalpida (= Libelluloidea) (see below) Gomphides (= Gomphoidea) †Araripegomphidae (incertae sedis) Progomphidae (= Gomphidae: Gomphoidinae) Desmoproctida Lindeniidae (= Gomphidae: Lindeniinae) Oligophlebiata Hagenioidea (= Gomphidae: Hageniinae) †Proterogomphidae Hageniidae Brevicubitalia Zonophoridae (= Gomphidae: Gomphoidinae) Gomphida Epigomphidae (= Gomphidae: Gomphinae: Gomphini) Gomphidae (= Gomphidae: Gomphinae: Gomphini + Onychogomphinae + Octogomphini) Aeshnoptera †Cymatophlebiella (incertae sedis) †Mesuropetaloidea †Mesuropetalidae †Liupanshaniidae Aeshnomorpha Austropetaliida Archipetaliidae (= Austropetaliidae: Archipetaliinae) Austropetaliidae (= Austropetaliidae: Austropetaliinae) Panaeshnida †Progobiaeshnidae Aeshnida †Cymatophlebioidea †Rudiaeschnidae †Cymatophlebiidae Paneuaeshnida †Paracymatophlebiidae Euaeshnida †Eumorbaeschnidae Neoaeshnida Gomphaeschnidae (= Aeschnidae: Brachytroninae: Gomphaeschnini) Aeshnodea Allopetaliidae (= Aeschnidae: Brachytroninae: Gomphaeschnini) Euaeshnodea Brachytronidae (= Aeschnidae: Brachytroninae: Brachytronini) Aeshnoidea Telephlebiidae (= Aeschnidae: Brachytroninae: Gomphaeschnini) Aeshnidae (= Aeschnidae: Aeshninae)

Eurypalpida

Synthemistidae (= Synthemistidae) Neolamellida Gomphomacromiidae (= Corduliidae: Gomphomacromiinae or Libelluloidea incertae sedis) Valvulida Pseudocorduliidae (= Corduliidae: Gomphomacromiinae or Libelluloidea incertae sedis) Trichodopalpida Macromiidae (= Macromiidae) Laterocarinida Austrocorduliidae (= Corduliidae: Gomphomacromiinae or Libelluloidea incertae sedis) Mediocostida Idionychidae (= Corduliidae: Idionychinae or Libelluloidea incertae sedis) Longiansida Idomacromiidae (= Corduliidae: Idomacromiinae or Libelluloidea incertae sedis) Liberaponsida Cordulephyidae (= Corduliidae: Cordulephyinae + Neophyinae or Libelluloidea incertae sedis) Haplohamulida Oxygastridae (= Corduliidae: Gomphomacromiinae or Libelluloidea incertae sedis) Italoansida Corduliidae (= Corduliidae: Corduliinae or Corduliidae s.str.) Anauriculida Hemicorduliidae (= Corduliidae: Corduliinae or Corduliidae s.str.) Libellulida Urothemistidae (= Libellulidae: Urothemistinae) Libellulidae (= Libellulidae)

Zygoptera

Zygoptera

†Steleopteridae (incertae sedis) †Eosagrionidae (incertae sedis) Euzygoptera Lestomorpha Hemiphlebiidae Lestiformia †Cretacoenagrionidae Eulestiformia Chorismagrionidae Lestida Perilestidae Lestodea Synlestidae Lestinoidea Megalestidae Lestidae Coenagrionomorpha Hypolestidae (sedis mutabilis) Megapodagrionidae (sedis mutabilis) Coenagrioniformia (sedis mutabilis) Platystictidae Coenagrionida Pseudostigmatoidea Coryphagrionidae Pseudostigmatidae Coenagrionodea Platycnemidoidea "Platycnemididae" Protoneuridae Coenagrionidae Caloptera †Sieblosiidae Eucaloptera Amphipterygida Pseudolestidae (incertae sedis) Diphlebiidae Amphipterygoidea Thaumatoneuridae Amphipterygidae Calopterygomorpha Libellaginidae Chlorocyphidae Calopterygiformia Euphaeida Polythoridae Epallagoidea †Zacallitidae Epallagidae Calopterygida Heliocharitidae Calopterygoidea Caliphaeidae Calopterygidae

Odonatoptera

Odonatoptera

†Geroptera †Eugeropteridae Neodonatoptera †"Eomeganisoptera" †"Erasipteridae" Euodonatoptera †Meganisoptera †Namurotypidae †Meganeuromorpha †Paralogidae (sedis mutabilis) †Kargalotypidae (sedis mutabilis) †Kohlwaldiidae (sedis mutabilis) †Meganeuridae (sedis mutabilis) Odonatoclada †Lapeyriidae †Campylopterodea (incertae sedis) †Campylopteridae Nodialata †Protanisoptera †Polytaxineuridae †Ditaxineuromorpha †Permaeschnidae †Ditaxineurida †Callimokaltaniidae †Ditaxineuroidea †Hemizygopteridae †Ditaxineuridae Discoidalia Stigmoptera †Protozygoptera †Permagrionoidea †Permagrionidae †Permolestidae †Archizygoptera †Permephallagidae (sedis mutabilis) †Kennedyidae (sedis mutabilis) †Protomyrmeleontoidea (sedis mutabilis) †Batkeniidae †Protomyrmeleontidae Panodonata †Tarsophlebiidae Odonata (see above) †Triadophlebioptera †Triadotypomorpha †Triadotypidae †Pirouetiidae †Triadophlebiomorpha †Zygophlebioidea †Xamenophlebiidae †Zygophlebiidae †Triadophlebiida †Mitophlebiidae †Triadophlebioidea †Triadophlebiidae †Paurophlebiidae

Diptera phylogeny

• Diptera