Exoskeleton vs. Endoskeleton: biomechanical trade-offs
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1 Exoskeleton vs. Endoskeleton: biomechanical trade-offs.. Ratio: surface area (square) volume (cube) Hercules beetle (Dynastes hercules) strength of skeleton Shrew (Sorex sp.) a giant spider size of organism Exoskeleton vs. Endoskeleton The exception proves the rule, in aquatic arthropods. but this is not possible) 1
2 Other advantages of the exoskeleton The necessity of molting allows complete metamorphosis The cuticle includes a lipid monolayer The cuticle is impact resistant What about six legs?? Hexapoda may not be monophyletic Hexapoda (six legs) Insecta Dicondylia Pterygota Neoptera Eumetabola (** Exopterygota ) Crustacea? Myriapoda? Collembola Protura Diplura Archaeognatha Zygentoma Ephemeroptera Odonata Plecoptera Polyneoptera** Paurometabola (orthopteroids) Endopterygota (Holometabola) Paraneoptera (hemipteroids)** 2
3 Three classes of Entognathous hexapods: Collembola, Protura, Diplura Class Collembola (springtails): Class Diplura: oral cone (collophore) Class Protura: 12 Entognatha: Class Collembola (springtails, snow fleas) 6000 species families 4 subclasses (Lower Devonian) Not a true insect: 6-segmented abdomen 3
4 Entognatha: Class Protura ( proturans ) 730 species 3-8 families 2 subclasses (Devonian) Not an insect: no antennae anamorphosis Entognatha: Class Diplura ( diplurans ) Not an insect: antennal flagellum w/ intrinsic muscles glandular cerci 800 species 5-9 families 2 subclasses (Carboniferous) 4
5 The three major evolutionary innovations of the Class Insecta: wings, wing-folding, & complete metamorphosis ( 27 orders, total) Hexapoda Insecta (Ectognatha) Dicondylia 1. Pterygota = aquatic immatures 2. Neoptera complete metamorphosis Eumetabola (** Exopterygota ) Crustacea? Myriapoda? Collembola Protura Diplura Archaeognatha Zygentoma Ephemeroptera Odonata Plecoptera Polyneoptera** Paurometabola (orthopteroids) 3. Endopterygota (Holometabola) Paraneoptera (hemipteroids)** 3 ocelli order Archaeognatha (= Microcoryphia, Machiloidea) compound eye antenna cercus maxillary palp (7-segmented) labial palp (3-seg) order Zygentoma (Thysanura) style on coxa exsertile vescicle maxillary palp abdominal styli (seg. 2-9) median caudal filament (MCF) ovipositor molar region incisor region mandibles (mono- vs dicondylic) condyle incisor region molar region condyle condyle style cercus median caudal filament median caudal filament style (8 & 9) ovipositor cercus subclass Apterygota 5
6 Apterygota: Archaeognatha (bristletails) 350 species, 2 families; Devonian (?) leg-like maxillary & labial palps Monura? 5 mm Permian Dasyleptidae (Elmo fauna, Kansas) median caudal filament Families: Meinertellidae & Machilidae Apterygota: order Zygentoma (silverfish & firebrats) 370 species, 3-5 families 2 condyles Thermobia sp. Lepisma sp. Tricholepidion gertschi lives w/ w.na. carpenter ants) 6
7 Apterygota are probably paraphyletic, not monophyletic ( Apterygota (blue bar) is probably not a clade) outgroup (monocondylic) dicondylic mandible? Monura (extinct; condyly uncertain) Archaeognatha (=Microcoryphia) Zygentoma (=Thysanura) Dicondylia Pterygota (other insects) monophyletic: an ancestral species and all its descendants (a clade). paraphyletic: a monophyletic group from which one or more subsidiary clades (monophyletic groups) is excluded. The origin of wings: Paleoptera Hexapoda Insecta Dicondylia Pterygota = aquatic immatures Neoptera complete metamorphosis Eumetabola (** Exopterygota ) Crustacea? Myriapoda? Collembola Protura Diplura Archaeognatha Zygentoma Ephemeroptera Odonata Plecoptera Polyneoptera** Paurometabola (orthopteroids) Endopterygota (Holometabola) Paraneoptera (hemipteroids)** 7
8 The origin of wings: Paleoptera Ancestral groups all have aquatic immatures: Paleoptera Neoptera Holometabola Ephemeroptera Odonata Plecoptera Megaloptera Pterygota, Paleoptera: Ephemeroptera (mayflies) > 3000 species, 25 families; suborders & superfamilies uncertain Adult adult naiad (nymph) Naiad (immature) Ancestral features: 11-segmented abdomen cerci & median caudal filament 3 ocelli post-adult molt paired genitalia (!) (as in Archaeognatha & Zygentoma) 8
9 Mayflies aquatic immatures Nymph/naiad: subaquatic long-lived (vs. adult) tracheal gills morphologically diverse Permian mayflies: Protereisma sp. (Protereismatidae) Naiad (immature) (after Kukalova) long antennae 5-segmented tarsus nearly full wing venation subequal wings 9 pairs of gills Adult (after Carpenter) 10th MCF 9
10 Pterygota, Paleoptera: Odonata (dragonflies & damselflies) Two (or three) suborders, 27 families, 600 genera, 6500 extant species s.o. Anisoptera (dragonflies) s.o. Zygoptera (damselflies) caudal gills Thoracic tilt s.o. Anisoptera (dragonflies) s.o. Zygoptera (damselflies) 10
11 Tracheal gills and nymphal mask Anisoptera naiad (rectal gills) mask Zygoptera (caudal gills) mask Damselflies 11
12 Dragonflies Libelluloidea Aeshnoidea Dragonfly progenitors: order Protodonata (Carboniferous, approx 350 mya) 28 inches (70 cm) Meganeuridae: Meganeura Meganeuridae: Megatypus 12
13 Phylogenetic hypotheses for Paleoptera I. Paleoptera are monophyletic (Hennig) Outgroup Neoptera (all other winged insects) Odonata Paleoptera Ephemeroptera II. Paleoptera are paraphyletic (Kristensen) (Boudreaux) Outgroup Outgroup Ephemeroptera Odonata Paleoptera Paleoptera Odonata Ephemeroptera Neoptera (all other winged insects) Neoptera (all others) The Origin of Wing-Folding: Neoptera Hexapoda Insecta Dicondylia Pterygota = aquatic immatures Neoptera complete metamorphosis Eumetabola (** Exopterygota ) Crustacea? Myriapoda? Collembola Protura Diplura Archaeognatha Zygentoma Ephemeroptera Odonata Plecoptera Polyneoptera** Paurometabola (orthopteroids) Endopterygota (Holometabola) Paraneoptera (hemipteroids)** 13
14 The morphological basis of wing-folding (neoptery): Axillary sclerites Paleoptera PALEOPTERA (no (no wing-folding) Neoptera NEOPTERA (wing-folding) 2A+ 3A+ C+ Sc- R+ Rs- MA+ MP- CuA+ CuP- 1A+ Odonata posterior notal process 1A+ 2A+ 3A+ C+ Sc- R+ Rs- MA+ MP- CuA+ Plecoptera Pterygota, Neoptera, Polyneoptera: order Plecoptera (stoneflies) 3500 species, 2 suborders, 16 families adult: adult: naiad: (nymph) naiad (immature) 14
15 Pterygota, Neoptera, Polyneoptera: Order Grylloblattodea (ice or rock crawlers) > 33 species, 5 genera, 1 family (Grylloblattidae) Discovered in 1913 (Walker 1914) as a living fossil ) ovipositor cercus Pterygota, Neoptera, Polyneoptera: Order Mantophasmatodea (gladiators, heelwalkers) ~24 living species, 3 families, 8 genera (first recognized in 2002!) 15
16 Pterygota, Neoptera, Polyneoptera: order Dictyoptera, s.o. Blattodea (cockroaches) 4500 species, 6 9 families Blattella germanica Periplaneta sp. Sc C Sc CuP C R Cockroaches, past & present: R CuA M Blattoptera: Archimylacridae (Carboniferous) Sc R 1 Rs M CuA CuP 1A CuA CuP M anal fan 300 my-old fossil, 4 long Carboniferous (central Asia) 16
17 Pterygota, Neoptera, Polyneoptera: order Dictyoptera, s.o. Isoptera (termites) 3100 species, 7 families (all eusocial) isopterous wings workers nests (colonies) Mastotermes darwiniensis (Mastotermitidae) Termites: Social cockroaches imago eggs in ootheca anal lobe Indotermes soldier (Termitidae) Synapomorphies: 1. Eusociality 2. True castes Amitermes hastatus (primitivetermitidae) worker tertiary queen 2ndary queen primary queen 17
18 Pterygota, Neoptera, Polyneoptera: order Dictyoptera, s.o. Mantodea (mantises) 2400 species, 15 families ootheca Pterygota, Neoptera, Polyneoptera: order Phasmatodea (stick & leaf insects) 3000 species, 2 suborders, several (?) families order Dermaptera (earwigs) 2200 species, 3 suborders, 11 families s.o. Hemimerina s.o. Forficulina Timema sp. order Embioptera (webspinners) 360 species, 2 suborders, 8 families s.o. Arixenina 18
19 Pterygota, Neoptera, Polyneoptera: Order Orthoptera (grasshoppers, crickets & katydids) 22,000 species, 2 suborders, many families suborder Caelifera (11,000 spp): Acrididae (short-horned grasshopper) suborder Ensifera (11,000 spp): crickets, katydids, long-horned grasshoppers ear ear Specialized Neoptera without complete metamorphosis: the Paraneoptera Hexapoda Insecta Dicondylia Pterygota = aquatic immatures Neoptera complete metamorphosis Eumetabola (** Exopterygota ) Crustacea? Myriapoda? Collembola Protura Diplura Archaeognatha Zygentoma Ephemeroptera Odonata Plecoptera Polyneoptera** Paurometabola (orthopteroids) Endopterygota (Holometabola) Paraneoptera (hemipteroids)** 19
20 (orthopteroid) order Zoraptera (39 spp) Pterygota, Neoptera, [Eumetabola], Paraneoptera: the smaller hemipteroid orders Psocoptera: book lice (5500 spp) order Thysanoptera: thrips (6000 spp) order Psocodea: lice (11,000 spp) Phthiraptera: s.o. Mallophaga, biting lice (4450 spp) s.o. Anoplura, sucking lice (550 spp) Pterygota, Neoptera, [Eumetabola], Paraneoptera: more thrips (Thysanoptera) soldier caste, Carcinothrips leai (Australia) 20
21 Pterygota, Neoptera, Paraneoptera: Order Hemiptera (bugs) 3 (or 4) suborders, >133 families, >2000 genera, 100,000 species (size: 1mm - 110mm!) s.o. Auchenorrhyncha (40,000 spp): cicadas and hoppers Membracidae: tree hoppers Cicadellidae: leaf hoppers Cicadidae: cicadas s.o. Sternorrhyncha (25,000 spp): aphids, scales, whiteflies, other plant lice Fulgoroidea: plant hoppers s.o. (?) Coleorrhyncha (34 spp) Peloridiidae Gondwanan distrib. s.o. Heteroptera (30,000 spp): true bugs Lygaeidae (seed bugs) Coreidae (squash bugs) Pentatomidae (stink bugs) Miridae (plant bugs) (heteropterous wing) Nepidae (water bugs) 21
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