Linnean rank. kingdom Animalia Animalia Animalia phylum Arthropoda Chordata Chordata class Insecta Reptilia Mammalia order

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Class exercise: what is an insect? Microevolution Changes in gene frequencies in a population, also known as change at or below the species level Macroevolution Insect Macroevolution Large-scale changes over a geological time period (i.e. extended microevolution) Changes at or above the species level Linnean rank kingdom Animalia Animalia Animalia phylum Arthropoda Chordata Chordata class Insecta Reptilia Mammalia order Diptera Ornithischia (Predentata) Primates family Diopsidae Ceratopsidae Hominidae genus Teleopsis Triceratops Homo species dalmanni horridus sapiens Diversity of Form Function Taxon a group at any taxonomic level. 1

Insect in amber 40MYA 2 big take homes: The major innovations in insect evolution Great radiations of insects Four major innovations in insect evolution Tracheal system / impermeable exoskeleton - allowed insects to colonize l First wings allowed insects to disperse better, colonize new habitats, escape from predators Neoptery allowed insects to live in rougher habitats because their wings were protected. Holometaboly allowed larvae to specialize on growth while adults specialized on reproduction. Insects have been around for over 400 million years (before dinosaurs). The first major innovation in arthropod evolution The ability to live out of the water The development of the tracheal system Impermeable exoskeleton 2

Insects are arthropods phylum: Arthropoda They share many features with the annelids (phylum Annelida) onychophorans (phylum Onychophora) Annelida Scientists are continually working to improve phylogenies This is the latest, is found in the textbook. Onychophora Onychophora Insects are arthropods phylum: Arthropoda Extant arthropods include: Chelicerates Arachnids, horseshoe crabs Crustaceans Myriapods Crabs, shrimp Centipedes, millipedes Textbook Paleoptera Hemimetabola And Exopterygote Neoptera Holometabola Endopterygote Paleoptera Hemimetabola And Exopterygote Neoptera Jumping Bristletail (Archaeognatha: Machilidae) Stebbins Reserve, Solano County, California, USA Very similar to the first insects! Holometabola Endopterygote Wheeler et al. 2001. Cladistics 17, 113-169. photograph Alex Wild 2003 3

Late Devonian ~350 MYA into the Carboniferous Period (300 mya) The second major evolutionary step: FLIGHT insects appeared! Including both extinct extant groups Paleopteran group (ʺold wingʺ) Extinct Order Paleodictyoptera (52 cm wingspan) Extant Order Ephemeroptera Neopteran groups Extant Orders Orthoptera Blattodea Flight evolved only once in insects With the colonization of the skies The first great radiation of insects occurred Fossil Paleodictyoptera Note patterned wings long cerci Wingspan of up to ½ a meter! 260 million years or so ago over what are today the prairies of central Kansas. http://www2.southwind.net/~royb/fossil_insects.html 4

Order Meganisoptera Meganuera monyi Brongniart Paleoptera Predatory insects Aquatic immature stage Wing span at 75 cm Elevated levels (30 40%) of oxygen levels allowed for this gigantism http://www.kheper.auz.com/gaia/biosphere/arthropods/insecta/meganisoptera.htm Textbook Hemimetabola And Exopterygote Neoptera Holometabola Endopterygote Wings Wings are a principal distinguishing factor in the hierarchy of insects ( ʹpteraʹ of order names). The third major innovation Carboniferous Period (300 mya) The evolution of a wing flexion mechanism Flight has provided insects with dispersal capabilities is in part responsible for the radiation of insect species. Neopterous or new-winged Primitively apterygote vs. Secondarily apterygote Order Archeaognatha Order Orthoptera Textbook Paleoptera Hemimetabola And Exopterygote Neoptera Holometabola Endopterygote 5

The fourth major evolutionary step: Carboniferous Period (~300 mya) The evolution of holometaboly (complete metamorphosis) Permian Period (250 mya) Gymnosperms (conifers) became dominate plant form. To date, fossil remains belonging to thirty Insect Orders have been detected. These include the extant Orders Odonata, Plecoptera, Embioptera, Diptera, Thysanoptera, Hemiptera, Coleoptera, Mecoptera, numerous additional extinct orders. End of the Permian The Permian-Triassic boundary was a time of major extinction Sugarcane grub adult larva Photo: Lyle Buss Holometabolous insects comprise nearly 85% of all insects! Permian Period (250 mya) At the end of the Permian was the biggest of the mass extinctions 95% of all marine life was killed 70% of all l families became extinct This reduced the diversity in taxa within surviving insect orders 6

Mesozoic Era Breakup of Pangaea Gymnosperms, seed ferns dominated. Known as Age of the Reptiles. Rise of the Hymenoptera Lepidoptera Mesozoic Era Cretaceous Period (145 mya) Beginning of the Angiosperms (flowering plants). Coevolution The process of reciprocal evolution between interacting species, driven by natural selection Increase in plant diversity resulted in the second great insect radiation Produced all of the extant orders most of the families that we know today Great radiations of insects Flight Angiosperm plants The orders of insects are fairly clear However, relationship among orders higher groups is more contentious Resemblance does not necessarily indicate relatedness 7

Resemblance does not necessarily indicate relatedness Why might different species resemble one another? Why might different species resemble one another? Why might different species resemble one another? Homoplasy Convergent Evolution Homoplasy Belidae (Coleoptera) Lycidae (Coleoptera) Order: Neuroptera Order: Mantodea Homoplasy Similar or identical traits that have arisen independently in distinct species Textbook Order: Neuroptera Order: Mantodea 8

Why might different species resemble one another? Homoplasy Homology Coreidae Leptoglossus Hemiptera: Coreidae Leptoglossus spp. Order Hemiptera suborder Heteroptera 9