insect evolution Bio153: lecture 17 deuterostomes and the rise of chordates • • • invertebrate terrestrial invasion • transition from invertebrate ⇒ vertebrate • • insect diversification • showed “bursts” of diversification from wingless insects…. 1. winged insects 2. folded wing insects 3. metamorphosis 4. hardened 1st pair of wings most successful group ever more spp. of beetles than of all vertebrates! Hexapoda: 6 legs no legs on abdominal segments 1) evolution of wings - not “limbs” (bird, bat) possibly: - accessory of respiratory system? - fanning increased gas exchange - during Jurassic (O2 levels ↑ / CO2↓): dragon flies: 75 cm wingspans! • an adaptation, but also an encumbrance! 2) folded wings - streamlined - crawl through vegetation, burrow… 3) metamorphosis - larva, pupa, adult - ↓ competition between adult and offspring – exploit more environments 1 • beetles: appear ~ 230 mya • quickly rose to 40% of all insects • why so successful? 4) hardened 1st pair of wings - protection from mechanical damage predators parasites echinoderms (starfish, sea urchins etc.) • abundant in Paleozoic: crinoids • strictly marine – reduced in Permian extinction deuterostomes • • • • echinoderms, hemichordates & chordates blastopore = anus coelomate, triploblastic radial cleavage (cell fate determined late) • unique to echinoderms : pentamerous symmetry (5-sided) • radial symmetry is a derived trait (Echinoderms are in Bilateria) • larvae are bilaterally symmetrical 2 chordate characters: 1. notochord • stiff rod derived from embryonic gut • mesodermal without notochord 2. pharyngeal gill slits • used in filter feeding in early chordates • slits persist in aquatic vertebrates • become parts of jaw, inner ear in terrestrial vertebrates with notochord 3. dorsal hollow nerve cord • derived from ectoderm • “flipped” from annelid, arthropod arrangement hemichordates (closely related to chordates) • • acorn worms only 1 chordate character (pharyngeal gill slits) 4. post-anal tail • extension of notochord, musculature • important in locomotion chordates 1. urochordates (tunicates) 2. cephalochordates (lancelets) 3. vertebrates all chordates show the 4 chordate characters at some point in life cycle 3 1. • • • urochordates (sea squirts, tunicates) all 4 chordate characters as larvae lost in adult – sessile lifestyle heterochrony: descendent retains larval form 2. cephalochordates (e.g. amphioxus): all 4 characters • • chordate origins still obscure ~ 540 mya (Cambrian) steps in vertebrate evolution: 1. pre-vertebrate – suspension feeder (similar to amphioxus) 2. agnathan – no jaws; muscular pump to move food current 3. gnathostome – vertebrate with jaws probable vertebrate phylogeny: agnathans lamprey cartilaginous fish ray-finned fish lobe-finned fish lungfish tetrapods • agnathans: e.g. hagfish • no bones, no jaws; cartilaginous skeleton • only vertebrate with partially open circulatory system • several hearts to help move blood (cardinal hearts + caudal heart + branchial heart) jaws arose from gill arches: 1. both are bars of tissue that hinge and bend forward 2. both derived from neural crest 3. attached muscles have same embryonic development 4 ray-finned fish: • comprise half of all verts • ~20,000 spp. • swim bladders • gas exchange through gills • some gulp air • cartilaginous fish: e.g. sharks, rays • lack swim bladders • heterocercal tail: gives lift • complex sense organs lungfish: 3 species (tropical) lobe-finned fish: • one of 2 extant lineages with jointed limbs • loss of swim bladder • flourished in Devonian; thought extinct until 1938 - bi-lobed sac attached to floor of pharynx - highly vascularized - gulps air -similar to fish that gave rise to amphibians the terrestrial invasion (part 3) • late Devonian (~ 360 mya) •Panderichthys (extinct) – type of lobefinned fish • shallow ponds are stagnant – little O2 (lots of decaying algae) • gills not effective • Panderichthys: - bi-lobed sac in pharynx probably could swallow air (like lungfish) - lobed fins to push on substrate Ichthyostega: • dorsoventrally flattened • dorsally placed eyes suggest adaptation to life in shallow ponds Devonian/ Carboniferous Greenland 5 Ichthyostega: 1) powerful limbs and feet 2) vertebra are interlocking – prevents twisting of backbone 3) strong rib cage – prevents crushing of internal organs April 6th, 2006 why live on land? • limbs evolved from fins • Hox genes in fish expressed in tetrapod limbs 1) more O2 2) less competition 3) less predation (esp. in reproduction New adaptations: • neck! – shoulder free; arms and head have greater movement • new sensory organs – lower jaw senses vibration 6
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