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Protostome Animals Protostomes • Biologists estimate that there are > 10 million arthropod species • A hectare of pasture in England may contain 45 million beetles • In tropical rainforests, 40% of total mass of organisms are beetles and other insects • Ants: estimates are 1 million billion individuals • Mollusks: > 93,000 species Protostomes include some of the most familiar and abundant animals on Earth Protostomes: consumers, predators, and scavengers in many marine ecosystems The Relative Abundance of Animal Lineages Major protostome phyla Using the Fossil Record Ch elic era t es Major non-protostome phyla Arthropoda s an ce ds ta apo s i ru r C My Insects Nematoda all of the protostome phyla appear very early in the history of animal evolution two major events occurred in the fossil record of protostomes: • extinction of the trilobites (250 mya) • appearance of insects Mollusca Platyhelminthes Annelida Ectoprocta Chordata Echinodermata Cnidaria Porifera Other phyla hvordan man kan gøre jordnøddesmør fuglefoder Overview of Protostome Evolution 1. - An Overview of Protostome Evolution undergo spiral cleavage after fertilization during gastrulation, the initial pore becomes the mouth - coelom arises when openings form within blocks of mesodermal tissue Protostomes Protostome development Coelom - have 3 embryonic tissues have a coelom: exceptions - 3. some exhibit segmentation; Annelids, Arthropods 4. monophyletic group 1. Lophotrochozoans • • grow by adding to body elements flatworms, rotifers, annelids, molluscs 2. Ecdysozoans • • grow by molting nematodes & arthropod, etc. Ch ord ata po da Art hro Ec hin od erm ata On ych op ho ra Ta rdig rad a Ne ma tod a Mo llu sca inth es lm he Pla ty An ne lid a Ro tife ra mo rph a no ph ora Ecdysozoa Ac oe lo ida ria Cte Cn Po rife ra Lophotrochozoa are bilaterally symmetrical Protostomes: 2 main lineages: Protostomes: A Monophyletic Group Comprising Two Major Lineages There are 22 phyla of protostomes, but the eight major phyla shown account for about 99.5% of the known species 2. Once the protosome pattern of development had evolved, protostomes split into the lophotrochozoans and ecdysozoans Triploblasty Bilateral symmetry 2 major subgroups exist within the protostomes What Is a Lophotrochozoan? 14 phyla of lophotrochozoans include: molluscs, annelids, flatworms Lophotrochozoan: • presence of a feeding structure: lophophore • a type of larva: trochophore (common to many phyla) • Lophophore: specialized structure where the mouth is in the center of a ring of tentacles: functions in suspension feeding Lophotrochozoans Have Distinctive Traits Lophophores function in suspension feeding in adults. Food particles Water current Anus Mouth Tentacles have ciliated cells along surface Gut What Is a Lophotrochozoan? Lophotrochozoans Have Distinctive Traits Trochophore larvae swim and feed. 14 phyla of lophotrochozoans include: molluscs, annelids, flatworms Lophotrochozoan: • Trocophores: type of larva common to several phyla of lophotrochozoa • common in marine mollusks and annelids • may represent a feeding and a dispersal stage in the life cycle ring of cilia along the middle Mouth Cilia used in locomotion and feeding Anus What Is an Ecdysozoan? • The primary contrast between lophotrochozoans and ecdysozoans involves their methods of growth • Ecdysozoans: grow by molting—shedding of cuticle or exoskeleton • most prominent of the 7ecdysozoan phyla are roundworms (Nematoda) and the arthropods (Arthropoda) Ecdysozoans Exoskeleton: • is a firm, nonliving covering that is difficult to penetrate • exoskeletons pose a problem: they cannot grow as the animal body inside them grows solution evolved in the ancestors of the ecdysozoans • they shed outgrown exoskeleton • the presence of the exoskeleton required the evolution of new forms of movement and respiration Ecdysozoans • some have wormlike bodies covered by flexible exoskeletons called cuticles • a thin cuticle allows the exchange of gases, minerals, and water across the body surface, but restricts the animal to moist habitats • bodies are supported primarily by hydrostatic skeletons Ecdysozoan • Explaining the evolution of body coverings: hypothesis: support of hypothesis: - How Do Body Plans Vary among Phyla? Themes in the Diversification of Protostomes all protostomes • protostomes have diverged into 22 different phyla that are recognized by distinctive body plans or specialized mouthparts - are bilaterally symmetrical and triploblastic • Result: millions of different species - undergo embryonic development in a similar way • What triggered this diversification? consider evolutionary innovations - most protostome phyla have wormlike bodies How Do Body Plans Vary among Phyla? The Nature of the Coelom Varies among Protostomes No coelom Pseudocoelom to m es da • acoelomate body plan – reversion to lack of body cavity (for example, platyhelminthes (flatworms) are acoelomate) De ut er os Ar th ro po Ta rd ig ra da da at o O ny ch op ho Ne m ol lu sc a ra Ecdysozoa M el id a es th in An n m el yh at Pl Ac oe lo m or ph a R ot ife ra Lophotrochozoa Reduced coelom Pseudocoelom • pseudocoelom, which forms from an opening that originates between the ectoderm and mesoderm layers, arose independently in rotifers and nematodes Reduced coelom Coelom Pseudocoelom No coelom Pseudocoelom • radical changes occurred in coelom formation How Do Body Plans Vary among Phyla? • A fully functioning coelom has two roles: (1) providing space for fluids to circulate among organs (2) providing a hydrostatic skeleton for movement • In arthropods and molluscs, the coelom is drastically reduced…….Why??? Evolution of Arthropods • some wormlike ecdysozoans developed a layer of protective material, chitin, a strong, flexible, waterproof polysaccharide • a rigid body does not allow wormlike movement; such animals require appendages that can be manipulated by muscles Arthropod Body Plan Arthropods Have Specialized Body Plans Arthropod body plan (external view) Arthropods • have segmented bodies that are organized into prominent regions: head, thorax, and abdomen Tagma Head Thorax Abdomen • distinguished by jointed limbs and exoskeleton made of chitin • locomotion is based on muscles that apply force against the exoskeleton to move legs or wings (hydrostatic skeleton is unnecessary) • have a hemocoel that provides space for internal organs and circulation of fluids Success of Arthropods • this exoskeleton made aquatic arthropods excellent candidates for invading terrestrial environments • arthropods are the dominant animals on Earth, approaching 2 million described species Arthropods and Their Relatives Onychophorans - have a flexible cuticle that contains chitin, and reduced coelom - have soft, fleshy, unjointed legs that form from outgrowths of the body - recent molecular evidence links the onychophorans (phylum Onychophora) to arthropod lineages Exoskeleton (covers body) Jointed limbs Segmented body Fossil Arthropods trilobites (phylum Trilobita) • were once the dominant line of arthropods before becoming extinct by the Paleozoic era • heavily armored, had jointed appendages Arthropods and Their Relatives Water bears: • (phylum Tardigrada) also have fleshy, unjointed legs and reduced coelom • are extremely small, and lack circulatory systems and gas exchange organs • when dry conditions occur, the animal shrinks to a small barrel-shaped object that can survive for a least a decade Molluscan Body Plan Molluscs Have Specialized Body Plans Mollusc body plan (internal view) • The molluscan body plan is based on three major components: (1) foot: a large muscle located at the base of the animal and usually used in movement (2) visceral mass: the region containing the internal organs and external gill (3) mantle: a tissue layer that covers the visceral mass and that secretes a shell in some species • In molluscs, the coelom's functions are replaced by the visceral mass and the muscular foot Variation Among Body Plans of Wormlike Phyla • The wormlike protostomes are characterized by similar body plans but specialized mouthparts or feeding structures Gill Mantle (secretes shell) Visceral mass (internal organs and external gill) Muscular “foot” Some Protostome Phyla with Wormlike Bodies Have Specialized Mouthparts Echiurans (“spoon worms”) Mouth Proboscis • Echiurans: spoon worms – suspension feeders • Priapulids: penis worms – sit and wait predators • Nemerteans: ribbon worms – active predators Some Protostome Phyla with Wormlike Bodies Have Specialized Mouthparts Priapulids (“penis worms”) Everted throat Mouth The toothed throat everts (turns insideout), grabs the prey, and then retracts Food particles are trapped in mucus in the “gutter” of the proboscis and then moved to the mouth by cilia Some Protostome Phyla with Wormlike Bodies Have Specialized Mouthparts Nemerteans (“ribbon worms”) Mouth Proboscis The barb-tipped proboscis everts (from a different opening than the mouth) and spears or entangles the prey, and then retracts it toward the mouth Evolution of Protostomes • most protostome evolution took place in oceans • many different lineages of animals evolved feeding structures designed to extract small prey from the water, most likely because this was the only type of food available during much of animal evolution Evolution of Protostomes • external body coverings probably evolved as a result of the selective pressures resulting from predation • body coverings evolved independently in lophotrochozoan and ecdysozoan lineages • although greater complexity has evolved in many animal lineages, several lineages that have remained simple have been very successful Evolution of Protostomes • flowing water brings food with it: thus the sessile lifestyle evolved repeatedly during lophotrochozoan and ecdysozoan evolution • competition for space is a consequence of the sessile lifestyle