AMPHIESMENOPTERA - the “garment wings” The Major Groups of Holometabola Holometabola (=Endopterygota) Order Hymenoptera Superorder Neuropterida Order Raphidioptera Order Megaloptera Order Neuroptera Order Coleoptera Order Strepsiptera Superorder Mecopterida (= Panorpida) Antliophora Order Mecoptera - scorpionflies Order Siphonaptera - fleas Order Diptera - true flies Amphiesmenoptera Order Trichoptera - caddisflies Order Lepidoptera - butterflies & moths AMPHIESMENOPTERA - the “garment wings” SYNAPOMORPHIES • Adult prelabium fused with hypopharynx • Pronotum with paired setose “warts” • Wings with extensive covering of setae • Anal veins of the forewing apparently looping up into a double-Y configuration • Presence of paired glands opening on sternum V • Male genital segment IX with tergum and sternum fused, forming a closed ring Hypothetical amphiesmenopteran, from Kristensen (1984) TRICHOPTERA - caddisflies The Major Groups of Holometabola Holometabola (=Endopterygota) Order Hymenoptera Superorder Neuropterida Order Raphidioptera Order Megaloptera Order Neuroptera Order Coleoptera Order Strepsiptera Superorder Mecopterida (= Panorpida) Antliophora Order Mecoptera - scorpionflies Order Siphonaptera - fleas Order Diptera - true flies Amphiesmenoptera Order Trichoptera - caddisflies Order Lepidoptera - butterflies & moths Diversity of Recent hexapods as proportion of named species. (from Grimaldi & Engel 2005. Evolution of the Insects. Cambridge Univ. Press.) Trichoptera, a minor order! The 7th largest Hexapod order! Culicomorpha/Psychodomorpha 16,669 Trichoptera 14,291 Amphibia 6,347 Hydrachnidia 6,007 Odonata 5,680 Hydradephaga 5,560 Cypriniformes 3,497 Siluriformes 3,093 Ephemeroptera 3,046 Hydrophiloidea 2,800 0 1800 3600 5400 7200 9000 10800 12600 14400 16200 18000 Number of described species for world’s largest, monophyletic freshwater animal taxa. (from Malm, et al. 2013, Syst. Ent. 38:459-473) Trichoptera taxonomic diversity Extant Fossil Total Species 14,548 685 15,233 Genera 616 125 741 Families 49 12 61 (from Morse, JC, Trichoptera World Checklist, 2015 http://www.clemson.edu/cafls/departments/esps/database/trichopt/) 10 SYNAPOMORPHIES • Larvae aquatic, no open spiracles, respiration epidermal, often by filamentous abdominal gills • Larval tentorium reduced • Larval antennae small • Larval abdomen without prolegs SYNAPOMORPHIES • Adult with haustellum (prelabium + hypopharynx) • Adult mandibles reduced, without articulation TRICHOPTERA - caddisflies Diagnostic characteristics Adults • Mothlike, usually drably colored, soft bodied, 2-40 mm • Wings covered with hairs, occasionally with scales; held roof-like over body; some brachypterous; one species with micropteryous females • Antennae long, filiform • Mouthparts reduced, but palps well developed; prelabium fuses with hypopharynx to form well developed haustellum DIAGNOSTIC CHARACTERS • Larvae aquatic, with cases/retreats CLASSIFICATION (Holzenthal et al. 2011) Order Trichoptera Suborder Annulipalpia Superfamily Philopotamoidea (2 families) Superfamily Psychomyioidea (7 families) Superfamily Hydropsychoidea (1 family) Suborder Integripalpia Superfamily Glossosomatoidea (1 family) Superfamily Hydroptiloidea (2 families) Superfamily Rhyacophiloidea (2 families) Infraorder Brevitentoria Superfamily Leptoceroidea (7 families) Superfamily Sericostomatoidea (12 families) Superfamily Tasimioidea (1 family) Infraorder Plenitentoria Family Kokiriidae Superfamily Limnephiloidea (6 families) Superfamily Phryganeoidea (7 families) BIOLOGY Annulipalpia (10 families) Fixed retreat makers and net-spinners. Feed on fine detritus, some predaceous. Hydropsychidae Psychomyiidae Annulipalpia (cont.) Dipseudopsidae Polycentropodidae Integripalpia, Rhyacophiloidea (2 families) Free-living larvae - cool running water, predators Hydrobiosidae Rhyacophilidae Integripalpia, Glossosomatoidea (1 family) Tortoise-case makers - cases, grazers Glossosomatidae Integripalpia, Hydroptiloidea (2 families) Purse-case makers or microcaddisflies, < 5 mm. Only instar 5 builds case. Feed on algae. Hydroptilidae, Ptilocolepidae Integripalpia, Infraorders Brevitentoria, Plenitentoria (39 families) Tube-case makers. Many feeding strategies Limnephilidae Phryganeidae Leptoceridae Beraeidae Calamoceratidae Helicopsychidae Brachycentridae Caddisflies in ecosystems Diagrammatic cross-section of a stream, showing caddisfly microhabitat diversification. (from Wiggins 2004,Caddisflies, the Underwater architects. Univ. Toronto Press.) LIFE CYCLE • 1 generation/year in temperate regions; perhaps continuous production in tropics • 5 larval instars • Pupation in fixed, sealed cocoons of modified case or retreat • Pupation 2-4 weeks, pharate adult cuts out of cocoon and swims rapidly to surface • Adults short-lived; sometimes drink water • Most are noctural or crepuscular • Mate on ground or vegetation, with courtship rituals and swarming HABITATS Trichoptera Biogeographic Regions (de Moor & Ivanov 2008. Hydrobiologia 595: 393-407) 1 West Nearctic 2 East Nearctic 3 West Palaearctic 4 Boreal Palaearctic 5 South Palaearctic 6 East Palaearctic 7 Beringian 8 Neotropical 9 Afrotropical 10 Cape South African 11 Oriental 12 Australasian 13 Patagonian COLLECTING and PRESERVING • Adults are attracted to ultraviolet lights • Collect adults with cyanide (KCN) kill jars, mount on pins; or alcohol pan traps COLLECTING and PRESERVING • Diurnal species can be netted or aspirated off substrate. • Malaise traps set adjacent to stream also collect diurnal species or those not attracted to lights. • Larvae in Kahle's fluid (15 parts by volume 95% EtOH, 1 part glacial acetic acid, 6 parts formalin, 30 parts distilled water). But NOT for DNA extraction, use 95% EtOH instead. • Do not disassociate larvae from their cases. • Metamorphotypes very valuable for associating larvae- pupae-adults. WHY DO I STUDY TRICHOPTERA? • I love insects! • I love rivers and streams, especially in tropical mountains! • Amazing larvae! • Very diverse, but not overwhelming • Beautiful adults • Male genitalia morphology complex • I like to draw • Many new species • Easy to collect, prepare, and curate • Literature accessible - catalogs • Nomenclatural problems largely resolved TRICHOPTERA - caddisflies Additional Information • Tree of Life Trichoptera pages: http://tolweb.org/Trichoptera/8230 • Morse, 1997. Phylogeny of Trichoptera. Annual Review of Entomology 42: 427-50. • Holzenthal, R.W., R.J. Blahnik, A.L. Prather, K.M. Kjer. 2007. Order Trichoptera Kirby 1813 (Insecta), Caddisflies. Zootaxa 1668: 639-698 http://www.mapress.com/zootaxa/taxa/Trichoptera.html • Kjer, K.M., R.J. Blahnik, and R.W. Holzenthal. 2001. Phylogeny of Trichoptera (Caddisflies): Characterization of Signal and Noise Within Multiple Datasets. Systematic Biology 50(6): 781-816. • Kjer, K.M, R.J. Blahnik, and R.W. Holzenthal. 2002. Phylogeny of caddisflies (Insecta, Trichoptera). Zoologica Scripta 31: 83-91. • Holzenthal, R.W., R.J. Blahnik, K.M. Kjer, and A.L. Prather. 2007. An update on the phylogeny of caddisflies (Trichoptera). Pages 143-153. In, J. Bueno-Soria, R. Barba-Alvarez, and B.J. Armitage (editors). Proceedings of the 12th International Symposium on Trichoptera. The Caddis Press. Columbus, Ohio. TRICHOPTERA - caddisflies Additional Information (cont.) • Wiggins, G.B. (1996). Larvae of the North American caddisfly genera (Trichoptera). Toronto: University of Toronto Press. • Wiggins, G.B. (2004). Caddisflies, the underwater architects. Toronto: University of Toronto Press. • Holzenthal, R.W., R.E. Thomson, and B. Ríos-Touma. 2015. Order Trichoptera. Pages 965-1002. In: Thorp, J., Rogers, D.C. (Eds.), Ecology and General Biology: Thorp and Covich'sFreshwater Invertebrates, Academic Press. • Holzenthal, R.W., J.C. Morse, and K.M. Kjer. 2011. Order Trichoptera Kirby, 1813. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 1348: 209-211. • Trichoptera World Checklist: http://entweb.clemson.edu/database/trichopt/index.htm • Trichoptera Literature Database: http://www.trichopteralit.umn.edu/about.