Dawn of the Arthropods: An Introduction to 85% The Arthropods • First arthropods Early Cambrian 600 mya – All environments, diverse ecologies – Tiny mites + crustacea < 1 mm --- Japanese spider crabs 3 m long – 85% of described animal taxa – 3 - 100 million species Tardigrades and Onychophorans Similar to Arthropods • chitinous cuticles Phylum Tardigrada • ~900 species • 50 – 1200 µm •Surface films of freshwater on terrestrial plants, mosses and lichens •abundant (3 million m-2) Phylum Tardigrada Defining characteristic: mouthparts: protrusible, oral stylet for piercing plant and animal tissues to obtain fluids Phylum Tardigrada • chitinous cuticle lining outer surface and gut • arthropod-like striated muscles • hemocoel • gas exchange across body surface • NO RESPIRATORY STRUCTURES • lack cilia • 4 pairs of clawed appendages (non-jointed) Phylum Tardigrada • nervous system similar to arthropods • paired ventral nerve cord Phylum Tardigrada video • offspring develop as miniature adults • male deposits sperm into female seminal receptacle • cuticle never calcified • cryptobiosis – dehydrate with environmental extremes • gonochoristic •ANABIOSIS: reduced metabolism •CRYPTOBIOSIS: extreme anabiosis Phylum Onychophora “velvet worms” Defining Characteristics: • 2nd pair of appendages modified into jaws • 3rd pair of appendages form oral papillae Phylum Onychophora “velvet worms” Defining Characteristics: • slime glands discharge adhesive through oral papillae • hydrostatic skeleton Phylum Onychophora • ~ 100 species • moist terrestrial habitat • thin, non-waxy cuticle • nocturnal Phylum Onychophora • carnivores, herbivores, omnivores • predators shoot proteinaceaous glue from oral papillae • some use glue as defense video Phylum Onychophora • circular, longitudinal, and diagonal muscles • 1 pair feeding appendages • no jointed appendages • hydrostatic skeleton • pair of nephridia / segments Phylum Onychophora • Ocelli; no compound eyes, photonegative • Sperm morphology similar to oligochaetes and leeches •Mostly dioecious •Oviparous, viviparous, or ovoviviparous •Direct development Phylum Arthropoda • 6 classes • 80 orders • 2400 families • Loss of motile cilia • Segmented, jointed, sclerotized chitinous exoskeleton • Flexible joints, hemocoel, Sense organs, ecdysis, specialized segments (head, thorax, abdomen) Phylum Arthropoda: Pechenik Classification Subphyla: 1) Trilobitomorpha (extinct) 1) Class Trilobita 2) Chelicerata 1) Class Merostomata 2) Class Arachnida 3) Class Pycnogonida 4) Class Eurypterida (extinct) 3) Mandibulata 1) Class Myriapoda 1) Order Chilopoda/Diplopoda 2) Class Hexapoda 1) Subclass Entognatha 2) Subclass Insecta 3) Class Crustacea Arthropoda Exoskeleton • water impermeable • procuticle – mostly chitin •EXOSKELETON LIMITS CONSTANT GROWTH Arthropoda: Support and Locomotion • exoskeleton • Muscles connect body segments + joints • Exoskeleton across joints NOT absent, but thin and flexible; Phylum Arthropoda • • metamerism most taxa have fused and modified specialized segments Molting • exoskeleton covering except for sensory hairs, gland openings, spiracles • parts of gut also lined with cuticle • ecdysis – loss of exoskeleton Molting • new cuticle secreted first, but then hardened • period of vulnerability • under neural and hormonal control Arthropoda Nervous System and Sense Organs • Brain • Tactile receptors = bristles or setae Arthropoda Nervous System and Sense Organs • • • • Brain Tactile receptors = bristles or setae Proprioception necessary with jointed appendages Stretch receptors span joints and provide knowledge of limb positions Arthropoda Nervous System and Sense Organs • • • • • • • • Brain Tactile receptors = bristles or setae Proprioception necessary with jointed appendages Stretch receptors span joints and provide knowledge of limb positions Three photoreceptors: simple ocelli, complex lensed ocelli, compound eyes Compound eyes: ommitidia Ommitidium: nerve tracts and field of vision Visual fields of neighboring ommitidia overlap – movements detected by each – compound eyes suitable for detection of subtle movement Arthropoda Nervous System and Sense Organs • • • • • • • • Brain Tactile receptors = bristles or setae Proprioception necessary with jointed appendages Stretch receptors span joints and provide knowledge of limb positions Three photoreceptors: simple ocelli, complex lensed ocelli, compound eyes Compound eyes: ommitidia Ommitidium: nerve tracts and field of vision Visual fields of neighboring ommitidia overlap – movements detected by each – compound eyes suitable for detection of subtle movement Visual System • ocelli or compound eyes • light sensitive surface with light-absorbing pigment Circulation and Gas Exchange • Gas exchange structures = invaginations in cuticle • Inwardly branching trachea: hemocoel via spiracles • gas exchange with hemolymph Circulatory System • heart with ostia = open circulatory system • hemolymph Arthropoda Digestive System • Foregut- ingestion, transport, storage, mechanical digestion • Midgut- enzyme production, chemical digestion, absorption • Hindgut- water absorption, preparation of fecal material Arthropoda Excretion and Osmoregulation • Nephridia and malpighian tubules • nephridial pores excrete concentrated nitrogenous waste products • Urine from malpighian tubules to gut, concentrates urine (reabsorb water) • Terrestrial arachnids, myriapods, and insects excrete predominantly uric acid Arthropoda Repro and Development • Dioecious • Fertilization internal, parental care common Class Trilobita • 4000 described species • extinct • dorsoventrally flattened • 3 sections • biramous appendages Crustacea • Terrestrial and aquatic • All depths in marine, brackish, and freshwater • > 67,000 described species, likely 5-10x that number • Diverse form, size, and habitat • 5 classes, 34 orders Crustacea Subclass Malacostraca - Order Isopoda (pillbugs) - Order Amphipoda (scuds) - Order Euphausiacea (krill) - Order Stomatopoda - Order Decapoda (crabs, shrimp, lobsters, hermit crabs, crayfish) Subclass Branchiopoda – brine (monkeys) Subclass Ostracoda - ostracods Subclass Copepoda – copepods Subclass Pentastomida – nasal parasites Subclass Cirripedia (maxilopoda) – barnacles Characteristics • Head = 5 segments, trunk divided into thorax and abdomen • Carapace or cephalic shield • Appendages multi-articulate; either uniramous or biramous • Mandibles are modified limbs that function as jaws • Gas exchange by diffusion across specialized surfaces Characteristics • Nephridia • Simple and compound eyes in at least one stage – Compound eyes on stalk • Gut with digestive cecae • Nauplius larvae, either mixed or direct development • 2 pair antennae Crustacean Bauplan • Thorax – anterior segments fused = cephalon • Maxillipeds: additional mouthparts • Number of segments in thorax varies • Thorax appendages = pereopods – swimming, walking, gas exchange, feeding, defense – ultiarticulate and biramous Basic Crustacean Bauplan • Abdomen – Segments – Number of segments used in ID • Appendages = pleopods – Biramous, flap-like – swimming – Culminate in telson • Anus • caudal rami • w/uropods (last pair of abdominal appendages) forms tail fan Adult Crustacean Crustacean Bauplan • Nauplius Larvae – Single, median, simple eye – 3 pairs of sectioned, functional limbs • Become antennules, antennae, and mandibles Circulation • Open circulatory system – Dorsal ostiate heart – Internal organs bathed in fluid – Simple heart and vessels in most • Sessile species no heart; pumping vessels • Blood – Variety of cell types • Dissolved hemoglobin or hemocyanin • Explosive cells release a clotting agent at injury sites Heart Shape • Heart long and tubular; to postcephalic region • Or, globular, box shape, in thorax; association with thoracic gills Gas Exchange • Aquatic – Small organisms = diffusion – Concealed gills for protection, prevent dessication – External gills • Modified thoracic limbs • Gills are thin; maximize gas exchange • Most species beat gills to maintain flow Gas Exchange • Terrestrial – Cutaneous Respiration • Membranes on legs of some species – Gills • Concealed – Pseudotrachea • Internal blind sacs to outside through small pores • Air in sacs, gas exchange with blood • Internal gills moist Feeding Strategies • • • • • • • • • • Water currents Hydraulic vacuum Filter feeding Feeding basket Passive Twirling antennae Direct manipulation Sand grazers or Sand lickers Predators Parasitism Water Currents • Thoracic limbs for swimming and creating suspension feeding currents • Water drawn into space • Particles trapped by setae • moved to food groove and toward head Hydraulic vacuum • Mouth appendages = paddles • Water containing food drawn into interlimb space • Food particles are not filtered, but captured in small parcels of water • Individual algal cells are captured this way Filter feeding • Sessile crustaceans have feathery cirri to filter feed • food up to one mm • = detritus, bacteria, algae and various zooplankton • Some can coil cirrus around large prey in a tentacle fashion Filter feeding in slow water • Extend pairs of cirri like a fan • Sweep rhythmically through water Filter feeding in fast water • Allow water to run through filter • video Passive feeding • Use cirri to passively strain • Burrow into sand with anterior facing upward • Extend cirri to capture bacteria, protists and phytoplankton • Antennae brush food towards mouth Twirl antennae • Create spiraling currents that bring food toward mouth • Food entangled in setae near base of mouth, brushed in Direct manipulation • Manipulation by mouthparts, pereopods and subchelate anterior legs Sand grazers or Sand lickers • Brush sand grains with setose mouthparts • Select individual sand grain, rotate and tumble against mouthparts to remove organic material Predator • Grab prey with chelae pereopods • Tear, grind and shear with mouthparts • Hunters or ambushers use raptorial subchelae to stab, club or smash prey • Some hold prey in cage using endopods; others inject and suck out tissues Snapping Shrimp • Use large cheliped to snap close: produces loud popping sound and “shock” wave • Pressure wave stuns prey, pull into burrow Digestive system • Foregut – Lined with cuticle that is continuous with exoskeleton, molted – Short pharynx-esophagus, stomach – Stomach = chambers for storage, grinding and sorting Midgut intestine Length varies with shape size, diet digestive ceca Hindgut Short, to anus Excretion and Osmoregulation • Ammonia by nephridia and gills • nephridial excretory organs as antennal glands (green glands) or maxillary glands • Inner blind end is coelomic remnant of nephridium = sacculus Other osmoregulation • Sacculus - Actively remove and secrete material from blood into excretory lumen • metabolic waste removal and water and ion balance • Thin areas of cuticle – Gill surfaces • terrestrial isopods: ammonia diffuses from the body as gas Nervous System and Sense Organs • CNS • Brain: three fused ganglia • Protocerebrum • Deutocerebrum • Tritocerebrum • Primitive nervous system = ladderlike Nervous System and Sense Organs • variety of sensory receptors • innervated setae or sensilla: contain mechanoreceptors and chemoreceptors • Propioceptors • Animals in Class Malacostraca: statocysts Nervous System and Sense Organs • Two rhabdomeric photoreceptors: • Median simple eyes • Lateral compound eyes • Most possess both, either simultaneously or during development • Naupliar eye = primitive, secondarily lost Nervous System and Sense Organs • • • • • Lateral compound eyes Lack visual acuity Discern shapes, patterns and movement Color vision in some Lacking in many taxa Nervous System and Sense Organs • Underwater vision • Problems with angular distribution of light, lower intensity, and narrow range of wavelengths than in air • Solution: Mount eyes on stalks, increase information available to eyes. Increases field of view, and binocular range Nervous System and Sense Organs • Complex Endocrine and Neurosecretory Systems • Not well known • Molting, chromatophore activity, and reproduction under hormonal and neurosecretory control • Bioluminescence in several groups Reproduction And Development • Exploit virtually every life history scheme imaginable • Usually dioecious • Hermaphroditism in remipedes, cephalocarids, cirripedes, few decapods • Parthenogenesis common among branchiopods and certain ostracods Reproduction And Development • Gonads paired structures in trunk • Pair of gonoducts from gonads to genital pores on trunk segment • Male pair of penes, or single fused median penis • Female include seminal receptacles • Most Crustacea Copulate in Pairs