sign in sign up
<<
Home , Body louse, Mallophaga, Paraneoptera, Psocodea, Psocomorpha, Psocoptera

EVOLUTION of the THE MAJOR GROUPS OF

Polyneoptera • Order • Order Holometabola • Order • Order • Order Dermaptera • Order • Order Grylloblattodea • Order Rhaphidioptera • Order Mantophasmatodea • Order Coleoptera • Order • Order • Order • Order • Order (incl. termites) • Order • Order Mantodea • Order Siphonaptera • Order Diptera • Order Trichoptera Paraneoptera • Order • Order • Order Phthiraptera • Order Thysanoptera • Order Paraneoptera • Order Psocoptera - bark lice, book lice • Order Phthiraptera - true or parasitic lice • Order Thysanoptera - • Order Hemiptera - true bugs, , scale , , 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 • 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 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 of Psocoptera. These analyses indicate that parasitic lice evolved from within the Psocoptera suborder . Phylogenetic relationships among major lineages of the Psocodea inferred from mitochondrial 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 • 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 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, , 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: - 24 families - 5 families Troctomorpha - 8 families • 264 spp., 71 genera, 26 families in U.S. and Canada

PHTHIRAPTERA (including 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 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 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 (). 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 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 ~ /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 parasite cladograms and ~host switch/colonization examples of sorting events ~intrahost speciation ~extinction PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice • Dr. Roger Price, Prof. Emer., University of Minnesota, world famous "Mallophaga" taxonomist. • Amassed huge collection of lice at University of Minnesota. • Pioneered of pocket gopher lice which led to the most well supported evidence of cospeciation in insects. Described 100s of species of bird lice as well.

Price, R. D., Hellenthal, R. A., Palma, R. L., Johnson, K. P., & Clayton, D. H. 2003. Chewing Lice: World Checklist & Biological Overview. Special Publication 24, Illinois Natural History Survey. 501 pp. PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice

Habitat & Habits:

• pubis - human crab or pubic louse. • humanus - human head (subsp. capitus) and (subsp. humanus). Vector of 3 important diseases (infection caused by scratching feces or crushed louse into skin): ~ (, a bacterium) - epidemics during the World Wars, now a problem among the homeless (high fever, severe headache, back and leg pain, fleeting rash, enlarged spleen) ~ (tick borne or louse borne), caused by a spirochaete (, a bacterium) (sudden fever, chills, headaches, muscle or joint aches, and nausea; a rash may also occur) ~ Epidemic (, a bacterium). Louse becomes infective after biting typhus infected person. Has killed millions of people throughout history (headache, sustained high fever, cough, rash, severe muscle pain, chills, falling blood pressure, stupor, sensitivity to light, and delirium). Spreads during times of war, famine, privation. PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice

Diversity & Distribution:

5,000 species world, about 1,100 North America. Many undescribed species. • Amblycera: ca. 1,200 species on birds (rails, storks, hawks, hummingbirds, etc.); ca. 170 species on mammals in South America and Australia (marsupials, caviomorph rodents) • Rhyncophthirina: 3 species, 1 on elephants, 2 on wild pigs • : 2,700 spp. on birds, 380 on mammals (most in Trichodectiae) • Anoplura: 550 spp. all feed on mammals, especially ungulates and squirrels. Human head and body lice are shared with the higher primates and apes. PHTHIRAPTERA (including Mallophaga and Anoplura) True or parasitic lice

Collecting & Preserving:

• Collect from hosts by carefully examining feathers and fur • Record identity of host • Store dry or in alcohol and mount permanently on slides