v Digestive system: mouth only (protostomes) v How does gas exchange take place? v What are protonephridia for?

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Biosc 41 Announcements 10/15 v Quiz today- Phyla Platyhelminthes and Mollusca v Lecture- Phylum Arthropoda v Lab- Phylum Arthropoda v Mon s lecture will finish any arthropod info we don t get to today, have arthropod quiz, and do an exam review activity v Mon s lab will finish arthropod lab material (+ crayfish dissection) Quick Platyhelminthes Review v What type of symmetry do animals in Phylum Platyhelminthes have? v Digestive system: mouth only (protostomes) v How does gas exchange take place? v What are protonephridia for? v What is the class of free-living flatworms? An example? v What are the classes of parasitic flatworms? Example? Quick Mollusca Review v Features and structures universal amongst molluscs: Mantle- secretes shell in shelled molluscs; sometimes portions of mantle form siphon(s) Foot- contains sensory cells; adapted in various ways in different classes Visceral mass- internal organs consistent between classes, but arranged in different orientations Circulatory system includes heart and is open in all but cephalopods (which have a closed circulatory system) Nervous system includes nerve cords and centralized brain tissue Digestive system includes radula, well-developed digestive gland, and complete gut Heart and kidneys function as excretory system Quick Mollusca Review v Four major classes: Class Polyplacophora- Chitons Class Gastropoda- snails, nudibranchs, slugs Life cycle includes trochophore & veliger larvae Veliger stage involves torsion Class Bivalvia- clams, oysters, mussels Life cycle includes trochophore & veliger larvae Class Cephalopoda- squid, octopus, cuttlefish, nautilus Mantle has been modified to include a siphon for locomotion Foot has been modified into tentacles Closed circulation and complex brain Quick Mollusca Review Quiz 5 v What is a possible adverse effect of suspension feeding? v Molluscs are the animal group with the largest number of recent extinctions v What features distinguish bivalves from brachiopods (Phylum Brachiopoda)? 1. What is the class of free-living flatworms? And an example? 2. What are the two classes of parasitic flatworms? And an example of each? 3. Identify 3 of the 4 major classes of Phylum Mollusca 4. What are three features distinguishing bivalve molluscs from brachiopods (Phylum Brachiopoda)? 5. What is a possible adverse effect of suspension feeding? 1

10/15/14 More Relationships An Introduction to the Invertebrates, Arthropods, Part One Reference: Chapter 33 "Don't accept the chauvinistic tradition that labels our era the age of mammals. This is the age of arthropods. They outnumber us by any criterion by species, by individuals, by prospects for evolutionary continuation." Stephen Jay Gould, 1988 Arthropod = jointed leg Eumetazoan protostome, bilateral symmetry, triploblastic, coelomate (mostly) Defining characteristics Segmented body Some segments are fused by tagmosis into tagmata Exoskeleton made of chitin Secreted by epidermis Molting (Ecdysis à Ecdysozoa) Exoskeleton shed regularly during life or only during larval development No cilia in larvae or adults Most successful in terms of diversity and sheer numbers >1,000,000 species! Phylum Arthropoda general body plan Body divided into segments (somites) Regionally fused into specialized groups by tagmosis (i.e., 5 segments form head) Internal cavity à hemocoel (not coelom!) No internal segmentation (no septa contrast w/annelids) Each body segment has a pair of jointed appendages Arthropod Origins The arthropod body plan consists of a segmented body, hard exoskeleton, and jointed appendages This body plan dates to the Cambrian explosion (535 525 million years ago) Early arthropods show little variation from segment to segment Tagmosis allows for specialization & diversity Diversity of body form possible because of specialization of segments, regions and appendages Tagmosis = segments grouped together Specialized for particular functions for greater efficiency Head, thorax and abdomen are tagmata Regions specialized for performing different tasks 2

Exoskeleton Structure v Cuticle forms well-developed exoskeleton made up of plates (sclerites) Growth by ecdysis (hormone-induced molting) v Epidermis is a single layer of epithelial cells that secrete the cuticle in layers Outer layer = epicuticle, with water-repellant hydrophobic layers Inner layer = procuticle, protein & chitin Procuticle is hardened by 1. Sclerotization = cross-linking of proteins into 3D matrix 2. Mineralization (in crustaceans) = depositing calcium carbonate in procuticle (i.e., crab shell) The Arthropod Cuticle 1 4 Molting v Stages between molts are termed instars This is when actual tissue growth occurs, although there s no size increase until after the molt Usually applies to immature stages v Frequency of molting is under hormonal control (ecdysone) v Cuticle is weakened enzymatically, then animal crawls out v After molting, animal sucks in air/water to inflate new cuticle which then hardens The pros and cons of the arthropod exoskeleton v Pros 1. Provides protection; hard covering over muscles & viscera 2. Facilitates osmoregulation in terrestrial environment (less water loss through epidermis) 3. No need for hydrostatic skeleton, so body plan could diversify 4. Better leverage than endoskeleton (or no skeleton) for moving muscles attached to limbs v Cons 1. Growth must proceed through a series of molts 2. Chitin exoskeleton is heavy constraint on size Arthropod circulation v Open circulatory system Hemolymph moves through hemocoel, and has amebocytes, pigments, and sometimes clotting factors Muscular heart required since rigid exoskeleton prevents body movements from moving hemolymph Arthropod Respiration v Varies between groups but usually by gills in aquatic habitats and a tracheal system in terrestrial arthropods The tracheal system is composed of tubes that distribute air throughout the body Some arthropods have a specialized lung-like structure called a book lung or book gill Special chamber with platelike structures Enhanced surface area facilitates rapid exchange of CO 2 and O 2 Diffusion of gases across thin areas of the cuticle also contributes to respiration 3

Arthropod Osmoregulation and Waste Excretion v Reuptake of salts and nutrients v Antennal and maxillary glands in crustaceans Produce urine; ammonia also excreted from gills v Malpighian tubules in arachnids and insects Blind tubes extend into hemocoel and empty into gut Produce uric acid which is passed through the anus Solid matter may also be passed through the anus Arthropod Nervous System v Brain is 2-3 ganglia with specific functions v Ganglionated ventral nerve cord v Sense organs (sensilla) protrude out of cuticle v Membranous drums ( ears ) v Chemoreceptors with thin cuticle Arthropod Vision Simple ocelli Complex lensed ocelli Compound eyes made of ommatidia Arthropod Digestive System v Complete, with regional specialization Foregut - food intake, transport, storage, mechanical digestion (jaws, pharynx, gizzard) Midgut - extracellular digestion, nutrient uptake (cecae, digestive gland, hepatopancreas) Hindgut - excretion of undigested material, water reabsorption Arthropod Locomotion v Striated muscle Exoskeleton used as attachment point for muscles v Muscles anchor on inside of cuticle Connect into the jointed appendages on each segment Arthropod locomotion v Jointed appendages (limbs) + specialized muscles to move the limb pieces (podites) Extrinsic muscles connect to body wall Intrinsic muscles are contained entirely inside the limb v Ancestral condition, found in crustaceans, is to have biramous limbs (each limb has 2 branches) Uniramous (1 branch) - insects Biramous - crustaceans 4

Diversity of Arthropod legs Arthropod Taxonomy Seven Subphyla: 1. Trilobites (extinct since Paleozoic) 2. Onychophora (velvet worms) 3. Tartigrada (water bears) 4. Crustacea (crabs, lobsters, shrimp) 5. Chelicerata (spiders, horseshoe crabs, pycnogonids) 6. Myriapoda (centipedes, millipedes) 7. Hexapoda (Insects) Subphylum Trilobitomorpha (Extinct) Cephalon Thorax Pygidium Most common group of fossil arthropods - Once abundant in oceans - Disappeared by Paleozoic (345 million yr ago) Subphylum Onychophora v Onychophora nail bearers or velvet worms 1-3 cm v Transitional group within the Ecdysozoa possess some features not characteristic of arthropods Pseudocoelomate Soft cuticle no exoskeleton! No jointed appendages Legs are hollow stub legs v Found in litter layer & decaying logs Prey on small worms, arthropods 2 8 Subphylum Tartigrada Subphylum Crustacea v Water bears or moss piglets About 0.5 mm Terrestrial and aquatic habitats v true extremophiles: can survive Wide temperature extremes (-80C to >100C) Ionizing radiation Outer space! (for a while) v Capable of anhydrobiosis Able to desiccate to <3% water and stay that way for >10 years v Most occur in marine and freshwater environments v External Features: Distinguishing feature: only arthropods with 2 pairs of antennae Body of 3 tagmata: head, thorax, abdomen Cephalic shield or carapace present to protect dorsum Mandibles, modified limbs, act as jaws; 2 pairs of antennae Limbs are biramous (2 forks) Gills are actually legs modified for gas exchange 5

10/15/14 Subphylum Crustacea Generalized Crustacean Body Plan Marine arthropods video (~10 min): http://shapeoflife.org/video/phyla/marine-arthropodssuccessful-design Subphylum Crustacea- Internal Features Hemocoel Major body cavity; filled with hemolymph Coelom restricted to compartment around gonads Muscular System Striated muscles form a large part of the body Capable of fast, strong movement Subphylum Crustacea Internal Features Circulatory system Open - no system of veins to separate blood from interstitial fluid Hemolymph exits heart through arteries and passes through hemocoel and gills to return to the heart via sinuses Hemolymph may be colorless, reddish, or bluish Contains ameboid cells that may help prevent clotting Biramous leg Subphylum Crustacea Internal Features Respiratory System Very small crustaceans conduct gas exchange across thin areas of the cuticle Larger crustaceans have gills Subphylum Crustacea Internal Features Excretory System Nitrogenous wastes (mostly ammonia) excreted across thin areas of cuticle in the gills Antennal or Maxillary Gland ( green glands ) Excrete urine Function is to regulate ionic and osmotic composition of body fluids Hemocyanin and/or hemoglobin are respiratory pigments 6

10/15/14 Subphylum Crustacea Internal Features Subphylum Crustacea Internal Features Digestive System Complete gut Stomach often consists of more than one chamber Nervous and Sensory Systems Fused ganglia form brain Double ventral nerve cord has a pair of ganglia for each somite (segment) Tactile hairs on the body Chemical sensing of taste and smell occurs in hairs on antennae and mouth Statocysts controls balance Compound eyes; often on stalks Gastric stomach includes gastric mill for grinding Pyloric stomach Subphylum Crustacea Reproduction & Life Cycles Diverse reproductive modes Barnacles are hermaphrodites, but cross-fertilize Some ostracods reproduce parthenogenetically (eggs don t require fertilization) Most crustaceans brood their eggs Branchiopoda & Ostracoda Maxillopoda (Copepods, Barnacles) Malacostraca (Isopods, Decapods [Crab, shrimp, lobsters]) Most common life cycle involves metamorphosis Egg hatches into nauplius larva, which molt and goes through several instars Segments and appendages are added in successive molts Classes of Subphylum Crustacea - Branchiopoda & Ostracoda Classes of Subphylum Crustacea Small, aquatic or marine crustaceans Important in food webs Many have a bivalve carapace that encloses the body Example Daphnia (Class Branchiopoda) Class Maxillopoda Subclass Copepoda Copepods form important links in aquatic food webs Subclass Cirripedia Barnacles are a group of mostly sessile crustaceans But start life as a motile, nauplius larva They have a cuticle that is hardened into a shell Hermaphroditic one of the few arthropods that are 7

10/15/14 Class Malacostraca (shrimp, lobsters, crabs, isopods) Largest class of crustacean and most diverse Over 20,000 species worldwide Ecologically and commercially important Class Malacostraca Order Decapoda ( 10 feet ) Crabs, shrimp, lobsters One or more pairs of walking legs modified to form pincers (chelae) Many species have bright coloration and/or camouflage Subphylum Chelicerata Class Malacostraca Order Isopoda ( same feet ) Class Malacostraca Order Stomatopoda ~ 65,000 spp. Body of 2 tagmata: cephalothorax + abdomen 1st pair of appendages = chelicerae (look like fangs) No antennae Gas exchange by book gills, book lungs or tracheae Separate sexes Classes: Pycnogonida (sea spiders) Merostomata (horseshoe crabs) Arachnida (spiders) Terrestrial, marine and freshwater species Terrestrial species are the only crustacean to have become fully terrestrial ( pill bugs ) Restricted to moist environments Marine species include a few bizarre parasites of fish- and very large species! Mantis Shrimp Voracious predators Some species are ambush hunters and snag prey with raptorial limbs Others have forelimbs modified into clubs that carry a powerful concussive force Class Pycnogonida: Sea spiders Marine, intertidal to abyssal depths; worldwide distribution About 1,000 species Benthic, live on seaweeds or other invertebrates Sucking proboscis on 1st head segment used to feed on softbodied invertebrates Males: brood eggs on ovigers, special leg appendages Females: hollow legs filled with eggs 8

Class Merostomata: Horseshoe crabs v Inhabit shallow marine waters v Burrow just under sand surface, prey on buried animals like bivalves v Small chelicerae v Limited distribution: Limulus polyphemus restricted to east coast of North America v Distinctive telson, or tail spine Class Arachnida: Scorpions, spiders, mites, ticks v Arachnids have an abdomen and a cephalothorax, which has six pairs of appendages: the chelicerae, the pedipalps, and four pairs of walking legs v Gas exchange by trachaea or book lungs v No compound eyes but have ocelli (often multiple) v Many spiders produce silk, a liquid protein, from specialized abdominal glands Figure 33.33 Subphylum Myriopoda = lots of legs Digestive gland Intestine Heart Stomach Brain Eyes v Myriapods are terrestrial, and have jaw-like mandibles v Classes: Diplopoda (millipedes)- have two legs per segment on each side; slow but powerful Ovary Poison gland Anus Spinnerets Gonopore (exit for eggs) Silk gland Book lung Sperm receptacle Chelicera Pedipalp Chilopoda (centipedes)- have one leg per segment on each side; fast but not as powerful Class Diplopoda - Millipedes v Eat decaying leaves and plant matter v Have many legs, with two pairs per trunk segment v Non-venomous, but some can secrete noxious chemicals Class Chilopoda - Centipedes v Carnivores v One pair of legs per trunk segment v Most are venomous 9

Subphylum Hexapoda = 6 legs- Insects and relatives v More species than all other forms of life combined Live in almost every terrestrial habitat and in fresh water Internal anatomy includes several complex organ systems v Insects diversified several times Following the evolution of flight Adaptation to feeding on gymnosperms Expansion of angiosperms v Insect and plant diversity declined during the Cretaceous extinction, but has been increasing in the 65 million years since Figure 33.35 Abdomen Anus Vagina Malpighian tubules Thorax Head Ovary Tracheal tubes Compound eye Heart Antennae Dorsal artery Crop Cerebral ganglion Nerve cords Mouthparts Insect Success: the power of flight v Flight is one key to the great success of insects v An animal that can fly can escape predators, find food, and disperse to new habitats much faster than organisms that can only crawl Insect Development v Many insects undergo metamorphosis during their development v Incomplete (hemimetabolous) metamorphosis Young are called nymphs Resemble adults but smaller and go through a series of molts until they reach full size v Complete (holometabolous) metamorphosis Larval stages known by such names as maggot, grub, or caterpillar Larval stage looks entirely different from adult stage Usually involves some sort of pupation Insect Development Figure 33.37 Hemimetabolous - insects hatch looking much like adults. Nymphs gradually attain adult form. Holometabolous - young very different from adults Pupal stage metamorhosis into adult (a) Larva (caterpillar) (b) Pupa (c) Later-stage pupa (d) Emerging adult (e) Adult 10

Insect Life Habit Summary of Terrestrial Arthropods v Most insects have separate sexes and reproduce sexually Individuals find and recognize members of their own species by bright colors, sound, or odors v Insects may be predators, herbivores, microbivores, parasites Insect parasites that prey on other insects are referred to parasitoids, because they kill their host v Some insects are beneficial as pollinators, while others are harmful as carriers of diseases, or pests of crops v Ants, bees and termites have evolved eusociality Decentralised, self-organized colony with division of labor and physical differences between castes v Insects are classified into more than 30 orders Terrestrial arthropods video (~14 min): http://shapeoflife.org/video/phyla/terrestrial-arthropodsconquerors 11