METAMERIC, EUCOELOMATE ECDYSOZOANS: NON
Transcription
METAMERIC, EUCOELOMATE ECDYSOZOANS: NON
METAMERIC, EUCOELOMATE ECDYSOZOANS: NON-ARTHROPODS “ARTHROPOD COUSINS” AS ALWAYS, PHYLOGENY FIRST! Kingdom: Animalia Subkingdom: Eumetazoa Subkingdom: Parazoa Phylum: Porifera Phylum: Placozoa Clade: Bilateria Clade: Radiata Phylum: Cnidaria Phylum: Ctenophora Division: Protostomia Superphylum: Lophotrochozoa Division: Deuterostomia Superphylum: Ecdysozoa PHYLUM: ONYCHOPHORA Distinguishing characteristics of Phylum Onychophora: These are the “velvet worms” Body centipede-like, but without jointed legs (“lobopods”) Organisms equipped with two oral “slime glands” that shoot strands of mucus (over 1 ft!) in order to entangle prey PHYLUM: TARDIGRADA Distinguishing characteristics of Phylum Tardigrada: These are the “water bears” Minute animals found on aquatic an marine vegetation Excretory system similar to Malpighian tubules of insects Can contract into a state of suspended animation called cryptobiosis ONYCHOPHORA & TARDIGRADA: TRAITS SHARED WITH ARTHROPODS Onychophora and Tardigrada share with Arthropods: metameric body plan outer cuticle shed by ecdysis (Ecdysozoa) body cavity & fluid called hemocoel & hemolymph Historically, it was thought that “myriapods” (e.g. centipedes) evolved directly from velvet worms, and that insects evolved directly from myriapods Evidence shows all arthropods are more closely related to each other than they are to either Onychophora or Tardigrada, although they all may form a monophyletic group called Panarthropoda PHYLUM ARTHROPODA: BEGINNINGS? Tullymonstrum, a “Tully monster” PHYLUM ARTHROPODA: BEGINNINGS? Opabinia, an Opabiniid PHYLUM ARTHROPODA: BEGINNINGS? Anomalocaris, a proto-arthropod Zebra Swallowtail Marbled Orb Weaver Presented by Praying Mantis ROB Brownish-Gray Fishing Spider PHYLUM: ARTHROPODA Distinguishing characteristics of Phylum Arthropoda: These are the arthopods! Inverterbrates with jointed legs All have chitinous exoskeleton All have a hemocoel and blood fluid called hemolymph PHYLUM ARTHROPODA: FUN FACTS Arthropoda is the largest phylum in the animal kingdom Over 900,000 described species Could be as many as 5,000,000 more Artrhopods outnumber 200,000,000 to 1! PHYLUM ARTHROPODA: WHY SO MANY ARTHROPODS? Chitinous exoskeleton gives arthropods selective advantage in harsh environments Evolved a variety of defenses to avoid predation LOTS of offspring— r-selected Effectively infinite number of niches allows small animals to adapt & radiate Monarch Butterfly (toxic) Viceroy Butterfly (not toxic) Dogbane Leaf Beetle PHYLUM ARTHROPODA: CLASSIFICATION & TAXONOMY There are “currently” 5 subphyla of arthropods: Subphylum: Trilobitomorpha* Subphylum: Chelicerata Subphylum: Crustacea** Subphylum: Hexapoda Subphylum: Myriapoda * = extinct ** = paraphyletic NOT TRILOBITES, BUT…????? (e.g. Dickinsonia & Sprigginia SUBPHYLUM: TRILOBITOMORPHA Fossil forms only Body divided into three regions: cephalon thorax pygidium Thorax & pygidium further divided into 3 longitudinal lobes: median or central lobe 2 pleural lobes SUBPHYLUM: TRILOBITOMORPHA SUBPHYLUM: CHELICERATA Distinguishing characters: All chelicerates possess a medial pair of feeding appendages called chelicerae Chelicerae may be modified into either jaws, fangs, or as part of a capitulum Chelicerae are usually flanked laterally by a second pair of appendages called pedipalps SUBPHYLUM CHELICERATA: GENERAL FORM & FUNCTION The body of chelicerates is divided into 2 regions: Prosoma (cephalothorax) – Contains head, feeding, & sensory structures Opisthosoma (abdomen) – Contains all other major organs & structures Breathing is often accomplished by either book gills or book lungs A few arachnids have spiracles that open into tracheae Excretory function is accomplished by means of Malpighian tubules Book lungs of scorpion SUBPHYLUM CHELICERATA: CLASSIFICATION & TAXONOMY There are 3 classes in Subphylum Chelicerata: Class: Merostomata Class: Pycnogonida Class: Arachnida CLASS: MEROSTOMATA Distinguishing characteristics of Class Merostomata: These are the horsheshoe crabs (extant) and the sea scorpions (extinct) Opisthosoma terminates in a spike-like telson CLASS MEROSTOMATA: HORSESHOE CRAB BLOOD The hemolymph of horseshoe crabs uses copper as an oxygen carrier instead of iron (hemocyanin), so “blood” is blue Horseshoe crab “blood” is harvested because it contains amoebocytes that react with certain bacterial toxins LAL - test CLASS MEROSTOMATA: MATING ORGY CLASS: PYCNOGONIDA Distinguishing characteristics of Class Pygnogonida: These are the sea spiders Opisthosoma is highly reduced No special respiratory or excretory structures 4 to 6 pairs of walking legs CLASS: ARACHNIDA Distinguishing characteristics of Class Arachnida: These are the spiders, scorpions, ticks & chiggers, daddy-longlegs, and others Adults almost always have 4 pairs of walking legs CLASS ARACHNIDA: CLASSIFICATION & TAXONOMY There are 12 orders of Class Arachnida—here are 6 common ones: Order: Araneae Order: Scorpiones Order: Opiliones Order: Acari Order: Solifugae Order: Thelyphonida ORDER: ARANEAE Characteristics of Order Araneae: These are the spiders Chelicerae bear hollow fangs attached to venom glands Usually web-spinning organs called spinnerets at posterior of abdomen All are carnivorous, but habits highly variable Black-and-yellow argiope ORDER: ARANEAE Grass Spider ORDER: ARANEAE Rabid Wolf Spider ORDER: ARANEAE Goldenrod Crab Spider ORDER: ARANEAE Ozark (or “Missouri”) Tarantula ORDER: ARANEAE Latrodectus mactans Loxoceles reclusa ORDER: SCORPIONES Characteristics of Order Scorpiones: These are the scorpions Pedipalps modified into pincers Opisthoma divided into preabdomen & postabdomen Postabdomen terminates in a sting Book lungs only ORDER: OPILIONES Characteristics of Order Opiliones: These are the daddy- long-legs (or harvestmen if you’re a weirdo yank) No constriction between prosoma and opisthosoma Legs long and slender THESE ARE NOT SPIDERS !!! Closest relatives are mites ORDER: ACARI Characteristics of Order Acari: These are the mites, ticks, and “chiggers” Prosoma and opisthosoma fused Parasitic forms have modified chelicerae and pedipalps forming a capitulum that terminates in a barbed rostrum ORDER: SOLIFUGAE Characteristics of Order Solifugae: These are the camel spiders, sun spiders, or wind scopions Body spider-like, but chelicerae enlarged and lacking hollow fangs; pedipalps absent Spinnerets absent ORDER: THELYPHONIDA Characteristics of Order Thelyphonida: These are the whip- scorpions or “vinegaroons” Pedipalps large, curved, & pincer-like Opisthosoma terminates in flagellalike “tail” capable of firing acetic acid SUBPHYLUM: CRUSTACEA Characteristics: Only arthropod group with two pairs of antennae 1 pair of mandibles, and 2 pairs of maxillae (usually) Appendages are biramous (= two branches) Diversity of Crustacea SUBPHYLUM CRUSTACEA: PHYLOGENETIC ISSUES “Crustacea” is paraphyletic with Hexapoda Together, crustaceans and hexapods form a monophyletic clade called Pancrustacea Current thought is that hexapods (e.g. insects) evolved directly from a malacostracan crustacean ancestor Beyond this, primary hypothesis is that Pancrustacea, Myriapoda, and Chelicerata are indeed distinct groups; and that they diverged from 1 or 2 ancestral “protoarthropod” groups at roughly the same time… SUBPHYLUM “CRUSTACEA” CLASSIFICATION & TAXONOMY There are 5 classes in Subphylum Crustacea— the 3 most common are: Class: Branchiopoda Class: Maxillopoda Class: Malacostraca CLASS: BRANCHIOPODA Distinguishing characteristics of Class Branchiopoda: These are the water fleas, tadpole shrimp, and fairy shrimp Legs flattened and leaflike; serve as primary respiratory strucure Beyond this, body forms variable CLASS BRANCHIOPODA: CLASSIFICATION & TAXONOMY There are 4 orders of Class Branchiopoda The most common one is Order: Cladocera These are the water fleas Minute crustaceans, resemble fleas CLASS BRANCHIOPODA: ONE OF MY FAVS Tadpole shrimp are pretty rad. CLASS: MAXILLOPODA Distinguishing characteristics of Class Maxillopoda: Ostracods, copepods (cyclops), & others No typical abdominal appendages Adults may retain a unique, nauplius eye between or posterior to compound eyes CLASS MAXILLOPODA: CLASSIFICATION & TAXONOMY There are 6 subclasses of Class Maxillopoda. The two most common are: Subclass: Ostracoda Subclass: Copepoda SUBCLASS: OSTRACODA Characteristics of Subclass Ostracoda: Ostracods Superficially resemble minute clams Body enclosed in a bivalve carapace SUBCLASS: COPEPODA Characteristics of Subclass Copepoda: Copepods or “cyclops” Body paddle-shaped Antennules often longer than other appendages Medial eye cluster gives appearance of a cyclops Free-living forms have extreme ecological importance in both marine and freshwater habitats CLASS: MALACOSTRACA Distinguishing characteristics of Class Malacostraca: These are the “major crustaceans”— crawdads, lobsters, crabs, shrimp, pill bugs Most biodiverse class of crustaceans 19-20 body segments: 5 head, 8 thorax, 6-7 abdominal CLASS MALACOSTRACA: CLASSIFICATION & TAXONOMY There are 4 orders in Class Malacostraca. Three of the most common are: Order: Amphipoda Order: Isopoda Order: Decapoda ORDER: AMPHIPODA Characteristics of Order Amphipoda: These are the beach fleas & scuds Body laterally compressed Carapace absent Uniramous appendages Uropods do not form tail fan as in true shrimp ORDER: AMPHIPODA Giant amphipods measuring over 1 foot (!) have recently been discovered in deep ocean environments. ORDER: ISOPODA Characteristics of Order Isopoda: These are sowbugs, pill bugs, wood lice, or roly-polies 7 pairs of legs Only group of crustaceans that can be truly terrestrial ORDER: DECAPODA Characteristics of Order Decapoda: These are the most familiar crustaceans— crabs, lobsters, crayfish, & shrimp All have 5 pairs of walking legs If you can boil it and eat it, it’s probably a decapod! DECAPOD DIVERSITY DECAPOD DIVERSITY Ozark blind crayfish, Orconectes australis CRAYFISH—A TYPICAL CRUSTACEAN: GENERAL FORM & FUNCTION Body: the body is divided into two major regions—cephalothorax and abdomen Cuticle of head extends posteriorly to form dorsolateral covering called a carapace Carapace may project forward to form nose-like rostrum Abdomen terminates in a telson, flanked by two pairs of uropods (telson + uropods = tail fan) CRAYFISH—A TYPICAL CRUSTACEAN: GENERAL FORM & FUNCTION Appendages: may be either biramous (twoarmed) or uniramous Biramous appendages have a medio-proximal endopod, and a latero-distal exopod that join at a common epipod There are 4 groups of appendages: – – – – sensory appendages mouthparts walking legs swimmerets CRAYFISH—A TYPICAL CRUSTACEAN: GENERAL FORM & FUNCTION Sensory appendages are biramous and include long, lateral anntennae and shorter, medial antennules Mouthparts are biramous and include the innermost, jaw-like mandibles; 2 pairs of maxillae, and 3 pairs of maxillipeds 2nd maxilla bears a flat scaphognathite (gill bailer) to keep water moving over the gills Endopods and exopods are more pronounced in maxillipeds CRAYFISH—A TYPICAL CRUSTACEAN: GENERAL FORM & FUNCTION Walking legs (5 pairs) are uniramous 1st pair of walking legs bear pincers, or chelae Epipods of all walking legs (and 2nd & 3rd maxillipeds) form gills Swimmerets are biramous and found on the abdomen only In males, the first pair of swimmerets are modified into club-like copulatory structures Figure 14.21 CRAYFISH—A TYPICAL CRUSTACEAN: GENERAL FORM & FUNCTION CRAYFISH—A TYPICAL CRUSTACEAN: FEEDING & DIGESTION Mouthparts guide food to mandibles, and then into the mouth Digestive system: 1) 2) 3) 4) 5) 6) mouth esophagus cardiac stomach (w/ gastric mill) pyloric stomach intestine anus Gastric mill in cardiac stomach of a crayfish Digestive gland secretes digestive enzymes into the pyloric stomach CRAYFISH—A TYPICAL CRUSTACEAN: EXCRETORY SYSTEM Excretion is accomplished by means of two labyrinth-like green glands, internal to and just below the bases of the antennae Nitrogenous waste is expelled from the green glands through an excretory pore Interior of green glands CRAYFISH—A TYPICAL CRUSTACEAN: RESPIRATION & CIRCULATION Arthropods have an open circulatory system: Water enters branchial chamber and brings oxygen over gills Oxygen-rich hemolymph leaves gills and enters pericardial sinus Hemolymph enter heart through heart ostia, which pumps hemolymph out through arteries CRAYFISH—A TYPICAL CRUSTACEAN: NERVOUS SYSTEM Nervous system consists of: suprapharyngeal ganglion subpharyngeal ganglion circumpharyngeal connectives ventral nerve cord CRAYFISH—A TYPICAL CRUSTACEAN: REPRODUCTION Most crustaceans are dioecious Gametes are produced in a gonad— testis or ovary Gametes travel through a gonoduct—sperm duct or oviduct Gametes exit through a gonopore Fertilization is external Female with brood ????? SUBPHYLUM: HEXAPODA Distinguishing characteristics of Subphylum Hexapoda: These are the insects and their relatives All have 3 pairs of uniramous, thoracic walking legs Body segments (somites) grouped into three tagmata—head, thorax, & abdomen SUBPHYLUM HEXAPODA: SEGMENTATION VS. TAGMATIZATION A body segment (metamere or somite) is simply one out of a number of repeated body units. Head Thorax A tagma is a group of somites that performs a distinct set of functions: Head = sensory Thorax = flight & Abdomen locomotion Abdomen = all other major body functions The plural of tagma is tagmata SUBPHYLUM HEXAPODA: CLASSIFICATION & TAXONOMY There are 2 (or 3) classes in Subphylum Hexapoda: Class: Entognatha Class: Insecta NON-INSECT HEXAPODS: CLASS ENTOGNATHA Characteristics of Class Entognatha: These are the non- insect hexapods—e.g. springtails, etc. All have internalized, stylet-like mouthparts Mandibles are monocondylic (1 point of articulation) CLASS ENTOGNATHA: ORDER COLLEMBOLA Distinguishing characteristics of Order Collembola: These are the “springtails” Posterior abdomen bears a forked furcula for jumping Ventral structure called a collophore aids in water intake & excretion CLASS: INSECTA Characteristics of Class Insecta: These are the insects! Hexapods with external mouthparts (usually) Mandibles are dicondylic—2 points of muscle attachment CLASS INSECTA: GENERAL STRUCTURE & FUNCTION Head bears: 1 pair of antennae 2 compound eyes 3 simple eyes (ocelli) 3 pairs of mouthparts – mandibles – maxillae – labia CLASS INSECTA: GENERAL STRUCTURE & FUNCTION CLASS INSECTA: GENERAL STRUCTURE & FUNCTION Thorax bears: legs wings (usually 2 pair) – Rapid wingbeats possible due to: 1) asynchronous nerve impulses 2) fibrillar flight muscles CLASS INSECTA: GENERAL STRUCTURE & FUNCTION INSECT LEGS Cursorial = running Saltatorial = jumping Fossorial = digging Raptorial = predatory CLASS INSECTA: GENERAL STRUCTURE & FUNCTION Abdomen bears: Respiratory structures (spiracles) Copulatory / genital structures Anus Female ichneumon wasp ovipositing CLASS INSECTA: SENSORY & NERVOUS SYSTEM Nervous system resembles that of other arthropods: In insects, ganglia & other cephalic nerve masses usually referred to as a brain Brain connects to ventral nerve cord An insect brain CLASS INSECTA: SENSORY & NERVOUS SYSTEM Mechanoreception is accomplished by means of sensilla A sensillum is either a hairlike projection or individual body cells attached to a nerve ending Although a large number of sensilla may be found on the antennae, sensilla are generally found all over the body Diagram showing antenna with sensilla CLASS INSECTA: SENSORY & NERVOUS SYSTEM Chemoreception is accomplished by bundles of sensory cells located in sensory pits Olfactory (smelling) sensory pits are generally concentrated on the antennae Gustatory (tasting) sensory pits are generally located on feeding palps, but may also be located on feet (tarsi) Some insects, like butterflies, can “taste” with their feet. CLASS INSECTA: SENSORY & NERVOUS SYSTEM Visual reception is accomplished by both compound eyes and ocelli Compound eyes are comprised of hundreds or thousands of individual facets called ommatidia – Ommatidia act together to sense shape & color Ocelli sense light changes CLASS INSECTA: SENSORY & NERVOUS SYSTEM Auditory reception is accomplished using either hair sensilla or tympanal organs Tympani may be located on the thorax, abdomen, or legs Tympani present in grasshoppers, crickets, katydids, cicadas, and butterflies & moths CLASS INSECTA: FEEDING & DIGESTION Insects may be either herbivorous or carnivorous Digestive system: Mouth Pharynx Esophagus Crop Gizzard Gastric caeca & stomach Intestine Rectum Anus Some carnivorous insects are parasitoids CLASS INSECTA: FEEDING & DIGESTION CLASS INSECTA: EXCRETION Excretion is accomplished by means of special structures called Malpighian tubules Tubules at the end of midgut absorb posassium acid urate from hemocoel, and then pass urate down to rectum Water & potassium are reabsorbed by the rectum Uric acid is expelled out the anus with solid waste CLASS INSECTA: EXCRETION CLASS INSECTA: CIRCULATION Insects have an open circulatory system consisting of a single blood vessel: Dorsal aorta pumps hemolymph into hemocoel via peristaltic waves Beyond this, hemolymph circulates freely in the hemocoel Hemolymph enters heart again through ostia CLASS INSECTA: CIRCULATION CLASS INSECTA: RESPIRATION Most insects breathe using a tracheal system Air enters tracheae through abdominal spiracles Tracheae branch to become tracheoles Tracheoles deliver oxygen to individual cells throughout the body Abdominal spiracles in a caterpillar CLASS INSECTA: REPRODUCTION Insects are dioecious, and fertilization is almost always internal Exception: some primitive forms deposit sperm encased in a spermatophore, and leave it to be “found” by a female Male system: testes vas deferens seminal vesicle (w/ accessory glands) ejaculatory duct aedeagus genital opening Female system: ovaries lateral oviduct common oviduct (w/ accessory glands) vagina vulva; separate spermatheca for sperm storage CLASS INSECTA: REPRODUCTION Male Female CLASS INSECTA: REPRODUCTION Spermatophore Beetles in copula, aedeagus extended CLASS INSECTA: GROWTH & DEVELOPMENT Insects have evolved 3 different growth & development strategies: Ametabolous develoment – juveniles are minature adults Hemimetabolous development – juveniles are called nymphs; no pupal stage Holometabolous development – Juveniles are called larvae; pupal development HEMIMETABOLOUS DEVELOPMENT Dog-day Cicada HOLOMETABOLOUS DEVELOPMENT CLASS INSECTA: CLASSIFICATION & TAXONOMY There are 29 orders in Class Insecta. Know these 12: Ametabolous orders: – Order: Thysanura Hemimetabolous orders: – – – – – – – Order: Ephemeroptera Order: Odonata Order: Mantodea Order: Blattodea Order: Orthoptera Order: Phasmatodea Order: Hemiptera Holometabolous orders: – – – – Order: Coleoptera Order: Lepidoptera Order: Diptera Order: Hymenoptera AMETABOLOUS INSECTS ORDER: THYSANURA Characteristics of Order Thysanura: These are the silverfish & firebrats Body brown or shiny silver with three long caudal filaments Only dicondylic, ametabolous group of insects A silverfish HEMIMETABOLOUS INSECTS ORDER: EPHEMEROPTERA Characteristics of Order Ephemeroptera: These are the mayflies Nymphs aquatic with 3 caudal filaments Adults have highly veined membranous wings; abdomen with 2 or 3 caudal filaments ORDER: ODONATA Characteristics of Order Odonata: These are the dragonflies & damselflies compound eyes occupying most of head Long, slender abdomen Nymphs are aquatic predators ODONATA DIVERSITY ORDER: MANTODEA Characteristics of Order Mantodea: These are the praying mantids (mantises) Front legs raptorial and armed with spines Thorax elongate Hindwings broad and fan-like MANTODEA DIVERSITY ORDER: BLATTODEA Characteristics of Order Blatodea: These are the cockroaches Body oval and flattened Head concealed by shieldlike prothorax Abdomen bearing 2 posterior cerci Roaches can vector diseases like salmonella and hepatitis. ORDER: ORTHOPTERA Characteristics of Order Orthoptera: These are the grasshoppers, katydids, & crickets Hindlegs modified for Eastern lubber grasshopper jumping Forewings narrow & leathery; hindwings broad & membranous True katydid ORDER: PHASMATODEA Characteristics of Order Phasmatodea: These are the stick insects Body long and stick- like Wings absent Cerci present ORDER: HEMIPTERA Characteristics of Order Hemiptera: These are the “true bugs” plus cicadas, aphids, & hopping bugs Piercing-sucking mouthparts Only half of forewing membranous in some groups (“true bugs”) HEMIPTERA DIVERSITY HOLOMETABOLOUS INSECTS: ORDER COLEOPTERA Characteristics of Order Coleoptera: These are the beetles Forewings hardened and covering abdomen; (wing covers = elytra) Most diverse order of animals on the planet Different species of lightning beetles (Lampyridae) have different flash patterns. COLEOPTERA DIVERSITY COLEOPTERA DIVERSITY: LOCAL STUFF Tiger beetle June beetle Longhorn beetle American burying beetle THE BEATLES ORDER: LEPIDOPTERA Characteristics of Order Lepidoptera: These are the moths & butterflies Both pairs of wings covered with microscopic scales Mouthparts, when present, modified into a coiled sucking tube (maxillae) Eastern tiger swallowtail LEPIDOPTERA DIVERSITY ORDER: DIPTERA Characteristics of Order Diptera: These are the mosquitoes, gnats, crane flies, & true flies All have a single pair of wings (if present) Common house fly ORDER: DIPTERA Striped horse fly ORDER: DIPTERA Hover fly ORDER: HYMENOPTERA Characteristics of Order Hymenoptera: These are the wasps, bees, & ants Two pairs of membranous wings with veins reduced Female ovipositor may be modified into stinger Invertebrate group with the highest number of eusocial taxa Leaf-cutter ants use leaves to farm a fungus as their food source. ORDER: HYMENOPTERA A cicada killer wasp ORDER: HYMENOPTERA A “cow killer” velvet ant SUBPHYLUM: MYRIAPODA Distinguishing characteristics of Subphylum Myriapoda: These are the centipedes and millipedes Body divided into two regions—head & trunk 4 pairs of uniramous mouthparts 4 classes—2 primary: – – Class: Diplopoda Class: Chilopoda CLASS: DIPLOPODA Characteristics of Class Diplopoda: These are the millipedes Bodies usually round in cross section 2 pairs of legs per trunk segment Slow moving scavengers Some are equipped with poison glands that produce cyanides The Kentucky flat millipede is poisonous, but not venomous. CLASS: CHILOPODA Characteristics of Class Chilopoda: These are the centipedes Bodies usually flat in cross section 1 pair of legs per trunk segment First pair of legs modified into poison claws (venomous) Fast-moving predators The giant red-headed centipede is the largest centipede in the Ozarks. THE IMPORTANCE OF ARTHROPODS: A LESSON IN ECOLOGY Oak decline—villain bug, or a perfect storm?