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EVOLUTION WITHIN THE : , , Holometabola NEOPTERA Ephemeroptera (25 orders)

PTERYGOTA THE MAJOR GROUPS OF NEOPTERA Polyneoptera • Order • Order • Order Holometabola • Order Dermaptera • Order • Order Grylloblattodea • Order • Order Mantophasmatodea • Order Rhaphidioptera • Order • Order Coleoptera • Order • Order • Order • Order • Order Isoptera • Order • Order Mantodea • Order Siphonaptera • Order Diptera • Order Trichoptera • Order Paraneoptera • Order • Order Phthiraptera • Order Thysanoptera • Order THE MAJOR GROUPS OF NEOPTERA Polyneoptera • Order Plecoptera • Order Embioptera • Order Zoraptera Holometabola • Order Dermaptera • Order Megaloptera • Order Grylloblattodea • Order Neuroptera • Order Mantophasmatodea • Order Rhaphidioptera • Order Phasmatodea • Order Coleoptera • Order Orthoptera • Order Strepsiptera • Order Blattodea • Order Hymenoptera • Order Isoptera • Order Mecoptera • Order Mantodea • Order Siphonaptera • Order Diptera • Order Trichoptera • Order Lepidoptera Paraneoptera • Order Psocoptera • Order Phthiraptera • Order Thysanoptera • Order Hemiptera • Are the 3 major groups of Neoptera each monophyletic? • What are the phylogenetic relationships among these 3 groups?

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? Hypothesis of Boudreaux 1979

Enlarged hind wing anal region (but reductions and aptery occur) Polyneoptera (=Polyneopterata)

Wing flexion Many, see Grimaldi & Engel and future lectures Paraneoptera (=Acercarida)

Complete & others, see Grimaldi & Engel and future lectures Holometabola Phalloneopterata (=Endopterygida) • Jugal bar • Male gonopod morphogenesis similar • Also supported by recent molecular evidence Hypothesis Hennig 1981 Plecoptera

Uniquely primitive and may be sister taxon to all other Neoptera!

Paurometabola (=Polyneoptera, exclusive of Plecoptera)

• Terrestrial mode of life • Rigid tegmina with posteromotorism • Cluster of accessory glands in male genitalia

Paraneoptera

Holometabola (=) Hypothesis Kristensen 1981, 1991 Plecoptera Embioptera lower Neoptera Zoraptera Dermaptera Grylloblattodea (Mantophasmatodea) Phasmatodea Orthoptera Blattodea Isoptera Mantodea

Paraneoptera

Holometabola (=Endopterygota) Working hypothesis as proposed by Boudreaux and presented by Grimaldi & Engel

Polyneoptera

Enlarged hind wing anal region (but reductions and aptery occur)

Paraneoptera

Holometabola What are the relationships among the orders of POLYNEOPTERA? (not considering Kristensen’s assessment) Plecoptera PLECOPTERIDA Embioptera

Zoraptera

Dermaptera

Grylloblattodea

Mantophasmatodea

Phasmatodea ORTHOPTERIDA Orthoptera

Blattodea

DICTYOPTERA Isoptera

Synthesis of various hypotheses Mantodea (see Grimaldi & Engel) Plecoptera PLECOPTERIDA Embioptera

• Ovipositor lost • Head prognathous • Suppression of male styli Zoraptera • Fusion of premental lobes (glossa + paraglossa) • Anal lobe of hind wing lost • Neuroanatomical traits • Cerci reduced to 1 orDermaptera 2 segments • Wings dehiscent • Hind femora enlarged, with distinctive musculature • Communal behaviorGrylloblattodea • Others (see Grimaldi & Engel) Mantophasmatodea

Phasmatodea ORTHOPTERIDA Orthoptera

• Enlarged hind wing Blattodea anal region

DICTYOPTERA Isoptera

Mantodea Plecoptera PLECOPTERIDA Embioptera

Zoraptera

Dermaptera

Grylloblattodea

Mantophasmatodea

Phasmatodea ORTHOPTERIDA Orthoptera

• Second valvula reduced, with developmentBlattodea of gonoplac as functional ovipositor • Enlarged precostal field in forewingIsoptera DICTYOPTERA Mantodea Plecoptera PLECOPTERIDA Embioptera

Zoraptera

Dermaptera • Future lecture Grylloblattodea

Mantophasmatodea

Phasmatodea ORTHOPTERIDA Orthoptera

Blattodea • Future lecture Isoptera DICTYOPTERA Mantodea