Flatworms are acoelomate: they lack a body cavity. The rest of the phyla we discuss do have a body cavities; first, Pseudocoelomates: Pseudocoelomates: phylum Nematoda Nematodes are the most successful of several pseudocoelomate phyla • 80-100,000 describes species, probably > 500,000 total • Found in nearly all habitats (marine, freshwater, terrestrial, free living, and parasitic) • Abundant and very ecologically important. • All are very similar in body plan: All fundamental bilaterian traits: TRIPLOBLASTIC, ORGAN SYSTEM level of organization (more complex than flatworms), BILATERAL SYMMETRY • Motile and cephalized -- supporting system is hydrostatic and viscoelastic • External cuticle -- protection, support, etc. -- a major key to the success of this group; molted about 4 times during its maturation • Internal cavity: pseudocoelom, formed from blastocoel Pseudocoelomates: phylum Nematoda Pseudocoelomates: phylum Nematoda Internal cavity: pseudocoelom, formed from blastocoel early in development: Internal cavity: pseudocoelom, formed from blastocoel early in development: Pseudocoelomates: phylum Nematoda Pseudocoelomates: phylum Nematoda Internal cavity: pseudocoelom, formed from blastocoel early in development: Adult body plan of nematodes, with pseudocoelom, no mesodermal tissue around gut. Other major features: ectoderm mesoderm • Eutely (constant cell number) to varying degrees -- makes them important ‘model animals’ pseudocoel Gut (no muscle) Gut lumen Cross-section Chaenorhabditis elegans: adult worm has 959 somatic cells, plus some reproductive cells (gonads) Pseudocoelomates: phylum Nematoda Pseudocoelomates: phylum Nematoda Adult body plan of nematodes, with pseudocoelom, no mesodermal tissue around gut. Other major features: ectoderm mesoderm • Eutely (constant cell number) to varying degrees -- makes them important ‘model animals’ pseudocoel Gut (no muscle) • Complete gut with mouth and anus Basic body plan of adult nematode: Gut lumen Cross-section Functional attributes of nematode body plan: • Cuticle provides leverage for muscle action, and protection; must be molted to grow • no circulatory system, but pseudocoelom provides some convective transport (especially when ‘stirred’ by movement) • complete gut is a major advance: it allows continuous-flow food processing (one-way); also permits specialization of gut regions. • incomplete gut requires batch processing • no muscle in gut, no gut motility. Food ‘pushed’ through by pharynx action. Pseudocoelomates: phylum Nematoda Pseudocoelomates: phylum Nematoda Nematodes are extremely important ecologically Nematodes are extremely important ecologically • scavengers, predators, parasites (in nearly every habitat, often in very large numbers -- millions per cubic meter of soil) • scavengers, predators, parasites (in nearly every habitat, often in very large numbers -- millions per cubic meter of soil) • Many are parasites on plants (including crop plants; causative agents of some very important crop diseases). • Many are parasites on plants (including crop plants; causative agents of some very important crop diseases). • A few parasitize humans: pinworm, hookworm, Trichinella spiralis (causes trichinosis). • A few parasitize humans: Ascaris -- which infects about 25% of humans worldwide Adults in intestine Trichinella cysts in human muscle Eggs shed in feces (eaten) (DON’T eat raw pork!!!) Crawl up trachea to esophagus and swallowed Ascaris lumbricoides cluster passed from a person (cluster is about 4 inches wide) Pseudocoelomates: phylum Nematoda Nematodes are extremely important ecologically • scavengers, predators, parasites (in nearly every habitat, often in very large numbers -- millions per cubic meter of soil) • Many are parasites on plants (including crop plants; causative agents of some very important crop diseases). • A few parasitize humans: filaria such as Wuchereria (mosquito transmission, causes elephantitis) Elephantitis: severe, chronic (permanent) lympho-edema (swelling) Wuchereria in blood smear Juveniles penetrate gut wall; migrate in blood to lungs Eucoelomates -- animals with a ‘true’ coelom: the ‘classic’ phylogeny from morphology and development: The alternate molecular-based view of protostomes, with two main lineages: Ecdysozoans (those that molt) and Lophotrochozoa Eucoelomate protostomates The pseudocoelomate body plan has a complete gut but lacks muscle tissue in the gut (no inherent gut motility). In eucoelomates the coelom is lined with mesoderm, providing muscle in the gut to power gut motility Coeloms and their mesodermal lining develop in two distinct ways. First we consider protostomates: • spiral determinate cleavage • coelom forms by schizocoely: a block of mesoderm splits to form an internal cavity Eucoelomate protostomates Coelom formation by schizocoely: Eucoelomate protostomates Coelom formation by schizocoely: Eucoelomate protostomates Coelom formation by schizocoely: Eucoelomate protostomates Coelom formation by schizocoely: At the end of this process, the internal cavity is completely lined with mesoderm (definition of a true coelom) Mesoderm layer around the gut leads to muscle tissue, lets gut process food more effectively. Eucoelomate protostomates Coelom formation by schizocoely: Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) BY FAR the most successful animals in terms of numbers of species and ecology: • > 1 million described species (mainly insects); probably 520 times as many that are not yet described (along with many fossil forms) • Found in essentially all habitats, usually in great numbers • diverse in form and function Triploblastic eucoelomate protostomes, with segmentation Like nematodes, they have an external cuticle Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) Key features of arthropod body plan: • Reduced coelom in adults • open circulation (hemocoel) • Hard external skeleton (exoskeleton); also called cuticle, made largely of chitin (acetylglucosamine polymer) mixed in a complex way with protein and varying degrees of mineralization • jointed appendages (often highly modified) on segments • segmentation (metamerism: serial repetition of body parts in segments). As in annelids, segments contain subdivisions of organ systems (muscles, nerve nets, etc.) • Tagmosis (grouping--sometimes fusion--of segments into specialized units). In many groups, tagmosis can obscure the underlying segmentation. Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) Exoskeleton is key to arthropod success. Three functional attributes: Exoskeleton is key to arthropod success. Three functional attributes: 1. Movement requires flexible joints in appendages (hence, “jointed feet”). Joints are thin, flexible regions of exoskeleton (the rest of the exoskeleton is usually thick ‘armor’). 2. Exoskeleton provides support as well as protection. Helps arthropods move on land without buoyant support of water. • Exoskeleton also reduces evaporation (another advantage on land) Exoskeletons preadapted early arthropods (which were aquatic) to invade land; they were among the first land animals. Jointed legs, protection, and support are advantages of exoskeletons. • Muscles are connected across joints in antagonistic pairs (one to extend, one to flex) • Hard exoskeleton provides good muscle anchorage and effective movement; attachment arrangement provides variable leverage (fast movement or slower but powerful movement). But, there is a big problem: Exoskelton is largely non-living and it cannot grow. Therefore, it must be molted (shed) in order for animal to get bigger. Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) 3. How to grow with an exoskeleton: molting process Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) 3. How to grow with an exoskeleton: molting process • arthropods grow in bursts • Many arthropods go through a number of molts as they progress from larvae to adults • often change form (metamorphosis) and size Molting may limit maximum size, especially in terrestrial forms (without skeleton, animal is mostly helpless and cannot support itself -- too large and it collapses. So giant ants and crabs wouldn’t work well…. Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) This is about as big as terrestrial arthropods get (some fossil forms were a bit bigger) Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) This is about as big as aquatic arthropods get (some fossil forms were a bit bigger) Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) Multiple repeats of body segments (with appendages): metamerism. Tagmosis (segment fusion and modification) is pronounced in several arthropod classes. • Tagmosis creates specialized body regions comprised of multiple fused segments Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) Arthropod taxonomy is complex and controversial; most prevalent view is of 4 major subgroups: TRILOBITES: (extinct; extremely abundant in Paleozoic, and as fossils); many segments but little specialization of appendages and relatively little tagmosis. • most commonly, body is divided into three regions: head, thorax, abdomen (as in insects) • Many appendages become modified or lost Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) CRUSTACEANS: diverse, primarily aquatic, a few terrestrial (crabs, lobsters, shrimp, barnacles, isopods, etc). • About 40,000 species; abundant and important in marine and freshwater ecosystems. • Most go through a very different larval stage before metamorphosing to adults. • Many appendages that are highly specialized (gills, legs, claws, mandibles, antennae, feeding nets, etc.) • Appendages are biramous; they occur as pairs, often with different functions; in crustaceans, usually legs and gills Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) • Most crustaceans show considerable tagmosis; often there is a fusion of head and thorax into a carapace • Barnacles are a highly derived form that is sessile, living within a protective shell. Don’t look like most crustaceans but have typical jointed arthropod legs, used as feeding nets. Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) UNIRAMIANS: uniramous appendages; extraordinarily successful; millions of species (mostly insects). • Very diverse: centipedes, millipedes, insects; crawling, swimming, burrowing, flying, parasitic. Often highly specialized (especially mouth parts). • Only invertebrate lineage to have evolved flight (probably several times in different insect orders). • Social insects (bees, ants, termites) have the most complex social systems of all animals--including humans. Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) CHELICERATES: spiders, scorpions, horseshoe crabs, etc. Very successful (about 65,000 species, and very ancient. • two major regions (tagmata), anterior (cephalothorax) and abdomen • usually 4 pairs of legs on anterior and two pairs modified as mouth parts. The appendages used as mouth parts are very claw-like (chelicera) and (in spiders) are connected to venom glands. • Life cycles usually simple; eggs hatch into miniature adults (no larval stage in most species). • A major “technology” of spiders is use of silk (for webs, safety lines, etc.). Eucoelomate protostomates: phylum Arthropoda (‘jointed feet’) Uniramian body form is either