Phylum Chordata. very diverse phylum but considerably less diverse than eg. arthropods or molluscs



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48,000 species Phylum Chordata very diverse phylum but considerably less diverse than eg. arthropods or molluscs also many fewer species than most other major phyla most advanced phylum of animal kingdom; more complex anatomy eg nervous system eg. circulatory system one to which we belong along with fish, amphibians reptiles, birds and other mammals some of the largest or most massive animals major identifying characteristics: 1. Notochord flexible rodlike structure enclosed by a fibrous sheath extends the length of the body in larva and/or adult provides basic support and serves as main axis for Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 1

muscle attachments to permit fishlike undulatory movements first part of skeleton to form in embryo in primitive chordates the notochord persists through life in most chordates the notochord is replaced by a vertebral column of bone remnants of the notochord remain as intervertebral discs 2. Dorsal tubular nerve cord in most invert groups; nerve cord is ventral & paired with paired ganglia along it s length in chordates the nerve cord is a single dorsal hollow nerve cord front end usually enlarged to form brain 3. Pharyngeal (gill) slits slit-like opening sleading from throat to outside first evolved as a filter feeding apparatus still used by some to filter water for food in others as gills in some groups they are only found in embryo and lost as adults 4. Endostyle or Thyroid Gland specific kind of tissue found only in chordates was originally part of the feeding apparatus Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 2

endostyle secretes mucus and traps food inside the pharyngeal cavity eg. lamprey larva in most chordates the same tissue has become an endocrine gland in the neck region that helps control metabolism 5. Post-anal tail in aquatic chordates it provides motility especially designed for propulsion inwater especially for larval forms and fish fish later added fins to increase efficiency in terrestrial chordates it became a tail for leverage and balance, not for movement in humans tailbones is its remnant Origin of Phylum oldest known chordate fossil is from the Ediacaran (600-543 MY) in Australia 6 cm long Three major Subphyla: Subphylum: Urochordata (tunicates) Subphylum: Cephalochordata (lancelets) Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 3

Subphylum: Vertebrata (vertebrates) Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 4

Subphylum Urochordata (tunicates, sea squirts) 1600 sp all marine widely distributed in all marine waters at all depths size ranges from microscopic to 1 ft in diameter most are bag-like sessile suspension feeders as adults they often live in large colonies attached to rocks and other hard surfaces on the ocean floor adult tunicates seem to lack the major characteristics of the phylum adults resemble sponges more than they do other chordates most are sessile lack a coelom pump water through siphons but tadpole-like larva has typical chordata features adults have tough, nonliving, tunic covering body Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 5

secreted by mantle forms 2 siphons one group, salps, are barrel shaped pelagic animals with transparent gelatinous bodies live singly or in colonial chains up to several meters long Feeding & Digestion filter feeders incurrent siphon pharynx (branchial sac) slits atrium excurrent siphon water is drawn in through the incurrent siphon into a basket-like pharynx (branchial sac) mucus is secreted by glandular endostyle in groove along base of pharyngeal basket uses mucus and cilia to move food toward mouth complete digestive tract food moves from the pharynx into the stomach for processing Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 6

the filtered water passes into the atrium and out the excurrent siphon Respiration pharynx also serves as a respiratory organ Circulation simple open circulatory system with small ventral heart and 2 major blood vessels blood is pumped through sinuses within the animal s tissues blood alternately flows in each directions Nervous System nerve with ganglia and plexus of nerve fibers Excretion no specialized excretory organs nitrogens wastes are in the form of crystals deposited in the tunic Reproduction & Development all tunicates are hermaphrodites with single ovary and testis Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 7

fertilization produces an elongated free-swimming larva = tadpole larva larva reveals true chordate nature has all 5 major chordate characteristics has digestive system but doesn t feed swims about for hours to days, until it finds a suitable substrate to settle on attaches to substrate by adhesive discs once firmly attached the larva undergoes an exceptionally rapid metamorphosis into the adult: in just 10-15 seconds the tail, notochord, nerve cord, brain sensory structures and muscles are crushed into a mass of tissue over the next few hours, the internal organs c hange their positions, the siphons adult nervous system develop loses its tail and most chordate features and becomes an adult Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 8

some believe vertebrates arose from a neotenous tunicate Human Impacts of Tunicates antiviral, antitumor including possible treatment for malignant melanoma the most dangerous form of skin cancer Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 9

Suphylum Cephalochordata (lancelets) only 29 species (5 species in US) closest living relatives to vertebrates they display several intermediate characteristics between invertebrates and vertebrates the adult animal has all major chordate characteristics yet lacks the internal skeleton of vertebrae and bone that most vertebrates have burrowers and swimmer slender, translucent, laterally compressed, fishlike or eel-like body ventral side of body is flattened bears 2 folds o skin = metapleural folds while they can swim they spend most of their time partially buried in sandy bottoms of coastal waters with their heads protruding 3-7 cm long Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 10

often found with tails buried in sand instead of tunic, outer body is covered by soft epithelium have fish-like fins with reinforcing fin rays internal structure is very simple with with basic chordate characteristics: springy notochord for support supports body while swimming or burrowing Movement with well developed V -shaped bundles of swimming muscles = myotomes (=myomeres) provide fish-like movement by contracting against notochord also have dorsal hollow nerve cord Feeding & Digestion are filter feeders mouth surrounded by oral hood with tentacles (=oral cirri) an a wheel organ Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 11

cilia on these structures draw in water and food pharynx strains food from water food is trapped in mucous and enters the mout for processing digestive system is similar to tunicates but slightly more developed has hepatic caecum (or liver) as accessory digestive organ food is drawn into the intestine by mucus and cilia water passes through pharynx into atrium and out the atriopore Circulation closed circulatory system circulatory system similar to fish but no heart Respiration some respiration occurs through the pharynx but most gas exchange occurs by diffusion through the ventral body wall Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 12

Nervous System hollow nerve cord above notochord anterior end of nerve cord = very small brain almost no cephalization pairs of spinal nerves innervate each myotome single ocellus at front of head Excretion simple excretory system of nephridia similar to those in annelids Reproduction & Development all are dioecious with males and females larvae resemble adults but are covered by cilia cilia are used for swimming and drawing food toward the mouth retain chordate features in adult resemble small fish Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 13

Subphylum Vertebrata General most complex (?advanced) group of animals larger, more active need more food one explanation for their diversity and dominance is that, in general, vertebrate species have a lot more genes amphioxus has as many genes as an ant or fly eg flies 10,000 genes eg. annelids 13,000 genes but mice and humans ~20,000 genes fish became the 1st true vertebrates oldest known fossils of a vertebrate: 560 MY old 2.5 long found in Australia a similar 530 MY old fossil was found in China while only 1 invertebrate group (the insects) developed the powers of flight all vertebrate groups have developed some ability to fly or at least glide through the air Major Characteristics: 1. internal jointed skeleton of bone or cartilage Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 14

an endoskeleton permits unlimited growth much more efficient design is a living skeleton grows with animals (not a case) doesn t need to shed regularly probably began as cartilage then later became calcified into bone cartilage grows fast to form initial skeleton a hardened skeleton is also ideal for muscle attachments especially in areas of high mechanical stress real bone emerged as external protective dermal plates protected the head and brain and anterior part of the body eg. ostracoderms, placoderms these bony plates later became modified into scales of some fish Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 15

since bone is living tissue it also becomes important as a mineral reservoir (esp. phosphorus & calcium) eg humans: calcium needed for: muscle contractions nerve impulses clotting secretions heart beat etc in the most primitive vertebrates its not much more than a cartilage rod = notocord skeletons of lampreys and sharks and rays and some bony fish (eg. sturgeon) remains mainly cartilage in adults in most vertebrates it is divided into: axial skeleton braincase - surrounds brain vertebral column ribcage appendicular skeleton (limbs) jointed appendages: pectoral & pelvic eg. fins, legs, wings, 2. Segmented skeletal muscles (myotomes) Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 16

became W shaped instead of V shaped as in amphioxus provided more control over body movements 3. complex skin multilayered: epidermis, dermis contains numerous of sensory receptors glands (oil, sweat, wax, scent, poison, etc) keratin structures: scales, hair, feathers 4. more efficient digestive system digestion shifts from moving food by cilia and mucus to using muscular contractions (= peristalsis) to move food through GI tract vertebrates were the first to add an acid producing stomach that more effectively digests proteins additional digestive glands: pancreas & liver improve digestive efficiency Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 17

5. efficient respiratory systems closely tied to circulation of blood the original function of the pharyngeal slits to filter water for food becomes functional gills 6. increasingly efficient circulatory system with pumping heart (2,3, or 4 chambered) ventral heart most inverts have dorsal heart closed circuits of arteries and veins RBC s (erythrocytes) containing hemoglobin for efficient distribution of oxygen to tissues 7. most complex and best developed nervous system of all animals usually well developed head with sense organs and brain (cephalization lifestyle shift from filter feeding to predation increased emphasis on brain and senses better sensory and motor integration Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 18

senses: CNS = brain & spinal cord central processing and coordination PNS = nerves (eg. cranial nerves, spinal nerves) conduct impulses to brain from sense organs and from brain to muscles and glands complex eyes inner ears for sound and balance improved taste and smell lateral line for water vibrations electroreceptors to detect prey 8. Improved efficiency of excretory system paired true kidneys (most cephalochordates had none) collect and get rid of metabolic wastes & toxins greater role in salt and water balance 9. almost all are dioecious and reproduce only sexually Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 19

Origin and Evolution of Chordates similarities with echinoderms & hemichordates: radial cleavage deuterostomes same coelom formation oldest known fossils: a. Pikaia (Burgess Shale; middle Cambrian, ~500 MY) 5 cm long; V shaped myotomes probably a cephalochordate b. Haikonella (early Cambrian, 530 MY) chordate features: notochord, phayrnx, dorsal nerve cord also some vertebrate features but not a vertebrate: pharyngeal muscles paired eyes enalrged brain urochordates are probably the most primitive surviving group but adults are too specialized Neoteny? larval form achieves sexual reproduction; accelerated development of reproductive organs Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 20

cephalochordates are clearly similar to ancestral vertebrate form most primitive vertebrates were jawless fossil agnathans predate all gnathostomes in fossil record (>500 MY) oldest group = conodonts conodonts are clearly related to group represented by living lampreys and hagfish lampreys may have evolved from ostracoderms: small bottom dwellers covered with bony plates jawless extinct probably endoskeleton of cartilage not bone Classification of Vertebrates the classes of vertebrates are often grouped into clades according to criteria of major evolutionary significance: A. jaws present or absent agnatha = jawless gnathostomes = mouth with jaws B. fins versus walking legs Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 21

pisces = paired fins for swimming tetrapods = paired limbs for terrestrial locomotion C. offspring develop within a fluid-filled sac (=amnion) anamniotes = do not develop within fluid filled sac; eg fishes and amphibians amniotes = do develop within a fluid filled sac; eg. reptiles, birds, mammals from most primitive to most advanced: jawless fish 35sp; >500MY fishes sharks and rays 850sp; bony fish 21,000sp; 420MY amphibians 4300sp; 360MY reptiles 7000sp; 280MY birds 9700sp; 150MY mammals 4600sp; 200MY Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 22