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EVOLUTION of the PARANEOPTERA the MAJOR GROUPS of NEOPTERA

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EVOLUTION of the PARANEOPTERA the MAJOR GROUPS of NEOPTERA EVOLUTION of the PARANEOPTERA THE MAJOR GROUPS OF NEOPTERA Polyneoptera • Order Plecoptera • Order Embioptera Holometabola • Order Zoraptera • Order Megaloptera • Order Dermaptera • Order Neuroptera • Order Grylloblattodea • Order Rhaphidioptera • Order Mantophasmatodea • Order Coleoptera • Order Phasmatodea • Order Strepsiptera • Order Orthoptera • Order Hymenoptera • Order Blattodea (incl. termites) • Order Mecoptera • Order Mantodea • Order Siphonaptera • Order Diptera • Order Trichoptera Paraneoptera • Order Lepidoptera • Order Psocoptera • Order Phthiraptera • Order Thysanoptera • Order Hemiptera Paraneoptera • Order Psocoptera - bark lice, book lice • Order Phthiraptera - true or parasitic lice • Order Thysanoptera - thrips • Order Hemiptera - true bugs, aphids, scale insects, cicadas, plant hoppers, tree hoppers, frog hoppers, etc. Phylogenetic Relationships of the Paraneopteran Orders Psocodea Psocoptera Phthiraptera Paraneoptera (=Acercaria) Thysanoptera Condylognatha Hemiptera • Maxillary lacinia slender and Synapomorphies of the Paraneoptera elongate, detached from stipes • Postclypeus enlarged, accommodating similarly enlarged cibarial dialator muscles Psocoptera • Labial palps reduced or lost Psocodea • Tarsomeres 3 or less • Cerci absent • Only one abdominal ganglionic mass • Winged forms with abdominal trichobothria Phthiraptera Paraneoptera Thysanoptera (=Acercaria) Condylognatha Hemiptera • Unique complement of sclerotizations in the cibarium - a Synapomorphies of the Psocodea water-vapor uptake system composed of ovoid "lingual sclerites" connected by a Y- shaped "filamentous duct" to the Psocoptera cibarial sclerite. Psocodea • Rupture-facilitating cuticular modifications (escape mechanism) on the basal part of the antennal flagellomeres - a collarlike fold of the epicuticle and exocuticle (but not the endocuticle) at base of Phthiraptera flagellar segments • Cardo lost • Postclypeus bulbous, protruding • Ovipositor simplified Paraneoptera Thysanoptera (=Acercaria) Condylognatha Hemiptera Synapomorphies of the Condylognatha Psocodea Psocoptera • Opisthognathous head • Expanded hypopharyngeal apodemes • Dorsal shift of anterior tentorial pits • Narrow labrum • Unicondylar mandibular & lacinial Phthiraptera stylets Paraneoptera Thysanoptera (=Acercaria) Condylognatha Hemiptera Mouthparts Evolution of mouthparts within the Paraneoptera (see Fig. 8.2 Grimaldi & Engel) lacinia Psocoptera • Maxillary lacinia slender and elongate, detached from stipes • Postclypeus enlarged, accommodating similarly enlarged cibarial dialator muscles • Labial palps reduced or lost PSOCOPTERA (=Corrodentia) - bark lice, book lice PSOCOPTERA - bark lice, book lice “SYNAPOMORPHIES” Head: • Bulging eyes; bulging postclypeus • Antennal flagellum slender Thorax: • Prothorax small, pterothorax large • Prophragma and mesophragma suppressed or reduced; dorsal longitudinal muscles insert on strongly arched mesonotum • Hind coxa with “Pearman’s organ” with adjacent tympanum - stridulatory Wings: • Forewing CuA with fork near margin, “areola postica” • Wings coupled at rest by blunt projection of pterostigma of forewing hooks to fore margin of hing wing • In flight, small hook on vein CuP couples forewing to hind wing But none of these exclusive to all psocopterans!! PSOCOPTERA - bark lice, book lice CHARACTERISTICS • Very small, soft bodied, < 5mm • Active, fast running • Long, slender antennae • Fore wing > hind wing; wings held rooflike over body; venation reduced; wings may be absent • Face bulging, head depressed • No cerci • Tarsi 2–3-segmented • Wing and body of some species covered with scales; some brightly colored other PSOCOPTERA “bark lice” Hypothesis of Lyall 1985 PSOCOPTERA Liposcelidae “book lice” PHTHIRAPTERA true lice - obligate • Brachypterous or apterous ectoparasites of • Body flattened, including head • Hind femora enlarged birds and mammals • Fusion of meso- and metanotum • Loss of abdominal spiracles 1 and 2 Also supported by recent • Reduction or loss of labial palps molecular sequence data and • Prognathous head additional morphological • Eyes reduced or lost characters Summary of the phylogeny of Psocodea based on mophological (Yoshizawa & Johnson 2006) and molecular data (Johnson et al. 2004) shows evidence for paraphyly of Psocoptera. These analyses indicate that parasitic lice evolved from within the Psocoptera suborder Troctomorpha. Phylogenetic relationships among major lineages of the Psocodea inferred from mitochondrial genome sequences. From Li et al (2013). Result? Psocoptera almost certainly PARAPHYLETIC, especially when the book lice, family Liposcelidae, are included. Liposcelidae - book lice • Cosmopolitan, 6 genera, 100 species, most in Liposcelis • Minute, pale • Found in tight, concealed spaces, under bark, compost, leaf litter, some in ant nests • Also damp paper, cardboard, books - feed on mold and organic glue; also dead insect specimens - minor museum pest • Also mammal and bird nests - nidicolous - where they feed on nest detritus; even among feathers and fur of some birds and mammals - phoretic association • Apteryous or brachypterous These characteristics perhaps precursors of an ectoparasitic lifestyle! “PSOCOPTERA” - bark lice, book lice Habitat & Habits: • In addition to nidicolous species, found on branches, trunks of trees and shrubs, rock outcrops, galleries of wood boring insects, nests of ants, bees, paper wasps, termites, caves, dead leaves, etc. • Feed by using laciniae as pick to scrape algae, lichen, fungal hyphae, films of yeast, detritus “PSOCOPTERA” - bark lice, book lice Habitat & Habits: • Some live in colonies in silken sheets of webbing on trunks of trees - silk from labial glands • Parthenogenetic and viviparous species found • Some Liposcelidae live in houses or in stored grain • Some display drumming courtship behavior; in one species males display by standing on head! “PSOCOPTERA” - bark lice, book lice Collecting and Preserving: • Use small natural bristle brush dipped in alcohol • Collect off substrate; also beating and Berlese funnel. • Preserve in 80% alcohol, pin scaled species. • Must mount on slides for detailed determinations (based on microscopic structures of genitalia, laciniae, claws, hypopharynx). “PSOCOPTERA” - bark lice, book lice Diversity & Distribution: • 4,400 species world, mostly in tropics - very poorly known; 3 suborders: Psocomorpha - 24 families Trogiomorpha - 5 families Troctomorpha - 8 families • 264 spp., 71 genera, 26 families in U.S. and Canada PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice SYNAPOMORPHIES - highly synapomorphic! • Obligate ectoparasites of birds and mammals • Development of hydropyle and operculum in egg; cemented to host hair or feather • Head with very limited movement • Loss of dorsal tentorial arms • Virtual loss of discrete abdominal ganglia; fused with 3rd thoracic • Reduction or loss of lacineal stylets • Great reduction of maxillae • Reduction of antennal flagellum to 3 flagellomeres (plus pedicel) • Loss of wings • Loss of ocelli • Reduction of compound eyes a few ommatidia, usually lost • Reduction to 3 (from 4) nymphal instars • Loss of metathoracic spiracle PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice Other characteristics: • Generally small, flattened • Head prognathous • Wingless • Antennae short, 3-5-segmented • Eyes small • Legs short • Cerci lacking • Mouthparts chewing; sucking in suborder Anoplura and withdrawn in short beak when not in use Phylogenetic Relationships of the Suborders of Phthiraptera Amblycera “Mallophaga” chewing lice Ischnocera Rhyncophthirina The order Mallophaga is PARAPHYLETIC! Anoplura sucking lice see Grimaldi & Engel Table 8.1 for defining characters Evolution of mouthparts within the Paraneoptera (see Fig. 8.2 Grimaldi & Engel) lacinia Phthiraptera Amblycera Evolution of mouthparts within the Paraneoptera (see Fig. 8.2 Grimaldi & Engel) (absent) Phthiraptera Anoplura • Loss of laciniae • Hypopharynx and labium developed as piercing stylets PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice Habitat & Habits: • Feed on keratin in hair and feathers or dermal material, sebaceous secretions, although some regularly take blood. • Members of suborder Anoplura are obligate ectoparasites of eutherian (placental) mammals => highly successful blood feeders (hematophagy). Other suborders are obligate ectoparasites of birds and mammals, in feathery and hairy coats. Almost all bird groups have lice; among mammals, only monotremes, anteaters, armadillos, whales and porpoises, sea cows, and bats do not. PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice Habitat & Habits: • Usually host specific and spend entire life cycle on host; some restricted to particular body regions (e.g., head and body louse vs. pubic louse of humans) • Cement eggs - "nits”- directly to hair or feathers; eggs large relative to female body • Interesting parallels between host species evolution and louse evolution => cospeciation or parallel cladogenesis - evident through statistical congruence between cladograms of host and their parasites. • How close the congruence is depends on several factors or sorting events: ~ cospeciation ~ host switch/colonization ~ intrahost speciation ~ extinction Host Parasite intrahost speciation host switch/colonization c o s p e c i a t i o n extinction? Host Parasite Sorting events: Congruence between host and ~cospeciation
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