Chaetognaths, Echinoderms, and Hemichordates

1. Chaetognaths, Echinoderms, and Hemichordates Chapter 22

2. Deuterostomes • Deuterostome characteristics: • Radial, indeterminate cleavage • Formation of the mouth from a second opening • Enterocoelous coelom development • Chaetognaths are placed outside both protostome & deuterostome groups.

3. Phylum Chaetognatha • The arrow worms, phylum Chaetognatha, are all marine, planktonic organisms. • Some deuterostome embryological characters. • Molecular works suggests they are protostomes. • Currently not considered to be part of either group.

4. Clade Ambulacraria • Superphylum Ambulacraria contains two deuterostome phyla • Echinodermata and Hemichordata • Members share a three-part (tripartite) coelom, similar larval forms, and an axial complex (specialized metaneprhidium). • Xenoturbella is the sister taxon to Ambulacraria. • Now considered to be a Phylum: Xenoturbellida

5. Phylum Echinodermata • Echinoderms include sea stars, brittle stars, sea urchins, crinoids, sea cucumbers. • Entirely marine • Lack ability to osmoregulate. • Almost entirely benthic. • Nonsegmented.

6. Phylum Echinodermata • Five extant classes of echinoderms are currently recognized.

7. Phylum Echinodermata – Characteristics • Spiny endoskeleton of plates • Water vascular system • Pedicellariae • Dermal branchiae (skin gills) • Pentaradial symmetry in adults

8. Phylum Echinodermata - Symmetry • Echinoderms are bilaterally symmetrical as larvae. • This means their ancestors were bilaterally symmetrical.

9. Phylum Echinodermata - Symmetry • As adults they show secondary radial symmetry – pentaradial (5 parts). • Perhaps an adaptation for sessile living in early echinoderms. • Crinoids

10. Phylum Echinodermata - Symmetry • Today’s echinoderms are mostly motile. • Many are still radial. • Some have again become superficially bilateral (skeletal & organ systems still pentaradial). • Sea cucumbers. • A few sea urchins. • No well defined head or brain.

11. Phylum Echinodermata - Deuterostomes • Echinoderms have a true coelom with deuterostome development. • Radial, indeterminate cleavage. • Enterocoelous – the mesoderm lined coelom develops from outpocketing of the primitive gut. • Formation of the mouth at the end of the embryo opposite the blastopore.

12. Water Vascular System • Echinoderms have a water vascular system derived from part of the coelom. • A system of canals and specialized tube feet that functions in: • Locomotion • Food gathering • Respiration • Excretion

13. Water Vascular System • The water vascular system opens to the outside through small pores in the madreporite.

14. Water Vascular System • Canals of the water vascular system lead to the tube feet. • Tube feet may have suckers, allowing the echinoderm to move while remaining firmly attached to the substrate – important in areas with lots of wave action.

15. Endoskeleton • Echinoderms have an endoskeleton of calcareousossicles often with spines. • Endoskeleton is covered by an epidermis. • Some have a very substantial endoskeleton (sea urchins), others have only a few scattered dermal ossicles (some sea cucumbers).

16. Development • Eggs (which may be brooded or laid as benthic egg masses) hatch into bilateral, free-swimming larvae. • The type of larva is specific to each echinoderm class. • Class Asteroidea • Bipinnaria • Brachiolaria • Class Ophiuroidea • Ophiopluteus • Class Echinoidea • Echinopluteus • Class Holothuroidea • Auricularia • Class Crinoidea • Doliolaria

17. Development • Metamorphosis involves a reorganization into a radial juvenile. • Left/right becomes oral/aboral.

18. Class Asteroidea • Class Asteroidea includes sea stars. • Common on rocky shores and coral reefs, some found on sandy substrates.

19. Class Asteroidea • Sea stars have arms (rays) arranged around a central disc. • The body is flattened, flexible, and covered with a ciliated, pigmented epidermis.

20. Class Asteroidea • The mouth is on the underside of the sea star. • Ambulacral grooves stretch out from the mouth along each ray. • Tube feet border each groove.

21. Class Asteroidea • The aboral surface is often rough and spiny. • Around the base of each spine there are pincerlike pedicellariae that keep the surface free of debris and sometimes help with food capture.

22. Class Asteroidea • Skin gills are soft epidermis covered projections of the coelom that extend between ossicles and serve a respiratory function.

23. Class Asteroidea • The lower part of the stomach can be everted through the mouth during feeding.

24. Class Asteroidea • The upper part of the stomach connects to a pair of digestive glands (pyloric ceca) in each arm.

25. Class Asteroidea - Feeding • Most sea stars are carnivorous; feeding on molluscs, crustaceans, polychaetes, echinoderms, other inverts & sometimes small fish.

26. Class Asteroidea - Reproduction • Most sea stars have separate sexes with a pair of gonads in each ray. • Fertilization is external.

27. Class Asteroidea - Regeneration • Echinoderms can regenerate lost parts. • Sea stars can readily replace an arm if it is lost. • This may take several months. • They can also cast off an injured arm.

28. Class Asteroidea - Regeneration • Some species can even regenerate an entire individual from a broken off arm. • Usually, a small piece of the central disc must be included. • Linckia can regenerate a whole new individual from a broken arm with no central disc attached.

29. Concentricycloidea • The two species of sea daisies were described for the first time in 1986. • They are tiny (< 1 cm), have no arms and the tube feet are arranged around the periphery of the disc. • Once considered a separate class, they are highly derived sea stars.

30. Class Ophiuroidea • Brittle stars (Class Ophiuroidea) are the largest group of echinoderms. • Abundant in all benthic marine environments – even the abyssal sea bottom. • Brittle stars have very slender arms.

31. Class Ophiuroidea • No pedicillariae or skin gills. • Madreporite is on the oral surface. • Tube feet have no suckers, their primary function is to aid in feeding.

32. Class Ophiuroidea • Brittle stars move using their arms rather than tube feet. http://youtu.be/BWOdssnzsMY http://youtu.be/4Texm2eTmSc

33. Class Echinoidea • Class Echinoidea includes sea urchins and sand dollars.

34. Class Echinoidea • The endoskeleton is well developed in echinoids. • Dermal ossicles have become close-fitting plates that form the test. http://www.jaxshells.org/test.htm

35. Class Echinoidea • Echinoids lack arms, but still show the pentamerous plan in the five ambulacral areas with pores in the test for the tube feet.

36. Class Echinoidea • Most echinoids are “regular” having a hemispherical shape, radial symmetry, and medium to long spines. • Regular urchins move using their tube feet with some help from spines.

37. Class Echinoidea • “Irregular” echinoids include the sand dollars and heart urchins that include some species that have become bilateral. • Spines are usually short and are used in locomotion.

38. Class Echinoidea • Some urchins have very reduced tests, and bright coloration. • The pedicellariae in these species contain painful toxins.

39. Class Echinoidea • Echinoids live in all seas from the intertidal to the deep sea. • Urchins usually prefer rocky substrate, while sand dollars and heart urchins like to burrow into sandy substrate.

40. Class Echinoidea • Echinoids have a complex chewing mechanism called Aristotle’s lantern. • Teeth are attached here. • Sea urchins are usually omnivorous feeding mostly on algae.

41. Class Echinoidea • Sand dollars use their short spines to move sand & its organic contents to the sides, the food particles drop between the spines, and ciliated tracts on the oral side carry the particles to the mouth.

42. Class Holothuroidea • Sea cucumbers (Class Holothuroidea) are elongated along the oral/aboral axis. • Bilateral • Ossicles are greatly reduced in most species.

43. Class Holothuroidea • The body wall is usually leathery with tiny ossicles embedded in it, but can be very thin.

44. Class Holothuroidea • Oral tentacles are modified tube feet located around the mouth. • Food particles are gathered by the oral tentacles. • Tentacles are put into the pharynx one by one so food can be sucked off.

45. Class Holothuroidea • Sea cucumbers move using ventral tube feet and waves of contraction along the muscular body wall.

46. Class Holothuroidea • Sea cucumbers have a very unusual defense mechanism: • They are able to cast out part of their viscera. • The lost parts regenerate. • Some have organs of Cuvier that can be expelled in the direction of an enemy. • These tubules become long and sticky, sometimes containing toxins.

47. Class Crinoidea • Crinoids include sea lilies and feather stars. • At metamorphosis, juveniles become sessile and stalked. • Adults are free-moving in some species. • Long, many branched arms.

48. Class Crinoidea • Crinoids use their tube feet and mucus nets to feed on small organisms that are passed to their ciliated ambulacral grooves.

49. Phylogeny • Echinoderms are probably derived from bilateral ancestors. • Pentaradial symmetry may have been an adaptation to a sessile existence. • Some forms then become mobile. • Some mobile forms are secondarily bilateral.

50. Phylum Hemichordata • Hemichordates (acorn worms) are marine animals that have gill slits and a rudimentary notochord – however, the notochord is not homologous with the notochord in vertebrates.