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ARTHROPODS and the Origins of Insects ARTHROPODA and Related Phyla

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ARTHROPODS and the Origins of Insects ARTHROPODA and Related Phyla ARTHROPODS and the origins of insects ARTHROPODA and related phyla Phylum Annelida Phylum Gastrotricha Phylum Nematoda Phylum Nematomorpha Phylum Priapulida Phylum Kinorhyncha Phylum Loricifera Phylum Onychophora Phylum Tardigrada Phylum Arthropoda Phylum Annelida - segmented worms Marine, freshwater, & terrestrial segmented worms. Cosmopolitan, very diverse. Polychaeta (mostly marine), Oligochaeta (earthworms etc.) and Hirudinea (leeches). 1 mm - 3 m. 15,000 species. Phylum Gastrotricha Microscopic (0.06-3 mm) unsegmented free-living, aquatic worms. Marine and freshwater. Mieofauna and periphyton. Very abundant. Microphagous detritivores. Ca. 750 species. Phylum Nematoda - roundworms Microscopic to very long (up to 100 cm) unsegmented worms. Free-living and parasitic (intestinal roundworms, hookworms, pinworms, trichinosis), many pathogens of animals and plants (important to agriculture). Ubiquitous, very abundant. Marine, freshwater, terrestrial. Ca. 80,000 species. Phylum Nematomorpha - horsehair worms Very long, thin unsegmented worms (1 cm - 1 m). Immature forms parasitic in insects, sowbugs; adults occur in freshwater. Ca. 350 species. Phylum Priapulida Marine segmented worms (several mm to 15 cm), with extensible, spiny proboscis. Ca. 15 species known. Predaceous. Live in mud on bottom of shallow seas (esp. cold oceans). Phylum Kinorhyncha Microscopic marine segmented worms. Bristly, spiny, with crown of curved spines on head. Mouth bears piercing stylets. Feed on diatoms, protozoans, detritus. Muddy bottoms and meiofauna. Ca. 150 species. Phylum Loricifera Microscopic marine animals (1 micron -1 mm). Mieofauna. Only discovered in 1983. 25 species. Phylum Onychophora - velvet worms Terrestrial, wormlike. Elongate. Segmented, each with pair of short legs. Antenna-like papillae on head. Possess tracheae. Live on forest floor of tropical and southern temperate rainforest forest. Predaceous. Spit quick-hardening protein-based glue from oral papillae to ensnare prey. Feed on snails, worms, insects, sowbugs. Tear open prey with jaws, inject saliva, suck out digested tissue. Mostly on southern continents. Ca. 100 species. Phylum Tardigrada - water bears Microscopic, segmented animals (200-500 microns), with clawed legs. Found in moss, lichens, freshwater, marine. Feed on algae, other tiny arthropods. Can withstand very harsh or extreme environments in dormant, cryptobiotic, state. 840 species. ARTHROPOD EVOLUTION The TRADITIONAL view (Haeckel, Snodgrass, Weygoldt, Nielsen, Brusca & Brusca and many others) ARTICULATA - segmented animals poda Annelida OnychophoraTardigrada Chelicerata Crustacea Myria Hexapoda (insects) Segmented body: each segment with •coelomic cavities •neuromeres •nephridia •appendages Annelida (=Arthropoda of some authors) Onychophora PANARTHROPODA Tardigrada Chelicerata Crustacea Myriapoda Hexapoda Exoskeleton (chitin + protein) which can be molted ARTHROPODA (=Euarthropoda of some authors) poda Annelida OnychophoraTardigrada Chelicerata Crustacea Myria Hexapoda • Tagmosis - regional specialization of body segments •Sclerotization or calcification of the exoskeleton •Exoskel. of articulated plates •Jointed appendages •Head with 5 pairs of appendages •Compound eyes •Etc. The 4 major groups of extant (EU)ARTHROPODA Chelicerata (horseshoe crabs, arachnids, sea spiders) Mandibulata Crustacea (shrimps, crabs, etc.) Atelocerata (=Tracheata) Myriapoda (centipedes, millipedes) Hexapoda (insects) See Table 3.2 Grimaldi & Engel; Tree of Life for more details Annelida Onychophora Tardigrada Chelicerata MANDIBULATA Crustacea Myriapoda Hexapoda •Mandibles •Highly developed compound eyes ATELOCERATA poda Annelida OnychophoraTardigrada Chelicerata Crustacea Myria Hexapoda •Loss of 2nd antennae •Tentorium •Tracheae •Malpighian tubules poda Annelida OnychophoraTardigrada Chelicerata Crustacea Myria Hexapoda PATTERN OF TAGMOSIS UNIQUE! Head: 5-6 fused segments with labrum, mandibles, maxillae, labium Thorax: 3 segments, 3 pairs of 6- segmented legs Abdomen: 11-segments, no limbs except for terminal genitalia and cerci The fourth stage in the development of the body form (Fig. 24 D) differentiates the insects from all other arthropods. It is well shown in the embryo of many insects (Fig. 23 C). In this stage the trunk segments back of the protocephalon (Prc) become segregated into three regions. The first may be called the gnathal region (Gn), since its appendages are destined to become feeding organs; the second is the thoracic region (Th), set apart as the locomotor center of the insect by the special development of its appendages as locomotory organs; the third is the abdominal region (Ab), on which the appendages are reduced and mostly obliterated. FIG. 24 Diagrams suggesting the evolution of the definitive insect structure from that of a theoretical wormlike ancestor. Ab, abdomen; Gn, gnathal segments; H, definitive head; Lm, labrum; Mth, mouth; Prc, protocephalon (prostomium and at least one somite), or procephalic part of definitive head; Prpt, periproct; Prst, primitive head, or archicephalon (prostomium); Th, thorax. In the fifth stage (Fig. 24 E) the body of the insect attains its final form. The gnathal segments are now united with the protocephalon in the definitive head (H), which thus comprises. the prostomium and four succeeding segments and carries the gnathal appendages as well as the protocephalic appendages. In the Pterygota the thorax acquires wing rudiments in the form of paranotal lobes, two pairs of which eventually become wings, and is finally evolved into a highly perfected locomotor mechanism. The abdomen loses most of its appendages, becoming principally a container of the more important viscera and the seat of respiratory and reproductive activities. ARTHROPOD EVOLUTION: The NEW view (Aguinaldo, Zrzavy, and others) Gastrotricha ECDYSOZOA - molting animals Priapulida Loricifera ? Kinorhyncha Nematomorpha Nematoda Panarthropoda Onychophora Tardigrada ARTHROPODA •Absence of locomatory cilia •Trilayered cuticle (epicuticle, exocuticle, endocuticle) molted via ecdysteroids The 4 major groups of extant ARTHROPODA (= Euarthropoda) Chelicerata (horseshoe crabs, arachnids, sea spiders) Mandibulata Crustacea (shrimps, crabs, etc.) Atelocerata (=Tracheata) Myriapoda (centipedes, millipedes) Hexapoda (insects) See Table 3.2 Grimaldi & Engel; Tree of Life for more details Chelicerata Pycnogonida - sea spiders Xiphosura - horseshoe crabs +Eurypterida - sea scorpions Arachnida Palpigradi - microwhip scorpions Aranea - spiders Amblypigida - whip scorpions Thelyphonida (= Uropygida) - vinegaroons Schizomida - short-tailed whip scorpions Acari - mites, ticks Opiliones - harvestmen Scorpiones - scorpions Pseudoscorpiones - pseudo scorpions Solfugida - sun scorpions, camel spiders Ricinulei - hooded tickspiders See Table 3.2 Grimaldi & Engel; Tree of Life for more details Crustacea Branchiopoda - cladocerans, brine shrimp Remipedia Cephalocaridae - horseshoe shrimp Maxillopoda - barnacles, copepods Ostracoda - ostracods Malacostraca Isopoda - isopods, doodle bugs, roly-polies Amphioda - amphipods, scuds Decapoda - shrimp, crabs, lobsters Stomatopoda - mantis shrimp Mysida - mysids Euphausiacea - krill and several other orders Myriapoda Chilopoda - centipedes Symphyla Pauropoda Diplopoda - millipedes Diplopoda Symphyla Pauropoda Chilopoda MANDIBULATA Traditional hypothesis Atelocerata Atelocerata Myriapoda Hexapoda Crustacea MANDIBULATA New hypothesis Pancrustacea Myriapoda Hexapoda Crustacea Pancrustacea Atelocerata vs Pancrustacea Traditional hypothesis Atelocerata Myriapoda Hexapoda Crustacea Pancrustacea New hypothesis MANDIBULATA Paradoxopoda Mandibulata Pancrustacea Hexapoda Pancrustacea Crustacea Mandibulata Myriochelata Myriapoda Chelicerata – Friedrich, M. & Tautz, D. 1995. rDNA phylogeny of the major extant arthropod classes and the evolution of myriapods. Nature 376, 165–167. – Hwang, U.-W., Friedrich, M., Tautz, D., Park, C. J. & Kim, W. 2001. Mitochondrial protein phylogeny joins myriapods with chelicerates. Nature 413, 154–157. – among other papers Pancrustacea, Mandibulata “The latest word. .” Reiger et al. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature, 463: 1079-1084. • 41 kilobases from 62 single copy nuclear protein-coding genes • 75 arthropod taxa representing every major lineage • 5 outgroup taxa (tardigrades, onychophorans) • 3 independent analyses (parsimony, likelihood, Bayesian) Results • Strong support for Pancrustacea, Mandibulata • “Crustacea” paraphyletic • Xenocarida (Remipedia + Cephalocarida) sister group to Hexapoda Mandibulata, Myriochelata “The latest word. .” Rota-Stabelli et al. 2010. A congruent solution to arthropod phylogeny: phylogenomics, microRNAs and morphology support monophyletic Mandibulata. Proceedings of the Royal Society (B):doi: 10.1098/rspb.2010.0590. Pages 1-9 • 198 protein-coding genes, new sequences for myriapods • micro RNA complements from all major arthropod taxa • 393 morphological characters Results • Strong support for Mandibulata • Myriochelata is a tree-reconstruction artifact caused by saturation and long-branch attraction • common origin of Arthropoda (tardigrades, onychophorans and arthropods) Remipedia Cephalocarida from Reiger et al. 2010.
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