Organismal Biology #23
LOPHOTROCHOZOA

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The Lophotrochozoa are a diverse group of phyla with similar ribosomal RNA sequences.
The annelid worms and mollusks are two large phyla in this group; primitive members of both these phyla share a ciliated larval stage called a trochophore.
Mollusks are a large and diverse group of animals; familiar mollusks include snails, clams, squids, and octopuses.  
The Annelida are worms with thorough segmentation of the body.
The phoronids, bryozoans, and brachiopods are all filter-feeders that share a ciliated feeding organ called a lophophore.

The Lophotrochozoa are a diverse group of phyla. Most have a true coelom of the schizocoel type, but a few have only a pseudocoel. Ribosomal RNA sequences show these various phyla to be related. Phyla placed in this group include:
  • Phylum Rhynchocoela or Nemertea: "Proboscis worms" with a long, stick-like proboscis or "evert" that can be withdrawn by turning it inside out.
  • Phylum Rotifera: Rotifers, with a wheel-like crown of cilia at one end.
  • Phylum Acanthocephala: Spiny-headed, parasitic worms.
  • Phylum Annelida: Segmented worms, described below.
  • Several smaller phyla related to the Annelida:
    1. Sipunculida: Marine worms with tentacles around the mouth.
    2. Echiurida: Sausage-shaped worms with a tube-like extension in front of the mouth.
    3. Chaetognatha: Arrow worms, with dart-shaped bodies, abundant in marine plankton.
    4. Gastrotricha: Gastrotrichs.
    5. Pogonophora: Deep-sea tube-dwelling worms.
  • Phylum Mollusca: Snails, clams, squids, etc., described below.
  • Four phyla (Entoprocta, Phoronida, Bryozoa, Brachiopoda) characterized by a ciliated feeding organ called a lophophore.
Animal family tree


Lophophore-bearing phyla:
A Lophophore is a crown of ciliated tentacles that help gather suspended food particles.
The cilia trap these particles and bring them to the mouth, a form of filter-feeding.
The animal can withdraw its lophophore if conditions are muddy or if predators threaten.

Similarities of phoronids, bryozoans, and brachiopods:
  • All have lophophores.
  • True coelom, used as a hydrostatic skeleton.
  • Simple, U-shaped digestive tube, complete with mouth and anus.
  • Benthonic (bottom-dwelling), either mobile or sessile (attached).
Phylum Phoronida (phoronid worms): Tube-dwelling worms with a lophophore surrounding the mouth. Coelom is used in burrowing: muscle contraction builds up pressure in the coelom, which swells sideways and pushes sediment aside. Probably related to ancestry of Bryozoa and Brachiopoda.

Phylum Bryozoa ("moss animals"): Largest and most successful lophophorate group. Members quite varied. All are small, aquatic animals living in colonies. Many colonies are polymorphic, containing several dissimilar types of individuals. Ancestry is probably close to Phoronida.

Phylum Brachiopoda: Probably derived from phoronid ancestors by addition of a two-part shell as an aid in burrowing. Shell has two unequal valves; axis of symmetry bisects center of each valve. Valves are connected by muscles only (class Inarticulata) or by a hinge (class Articulata). Muscular stalk (pedicle) attaches animal to the bottom. Fossil record shows that brachiopods were more abundant and more diverse during the Paleozoic Era (530 million to 300 million years ago).

Phylum Entoprocta: A small group of sessile (attached) animals with a lophophore and a simple, U-shaped digestive tract. They differ from the previous 3 phyla in two ways: they have a pseudocoel instead of a true coelom, and their lophophore surrounds the anus as well as the mouth.

Evolution of Lophotrochozoa


Phylum Mollusca (mollusks):
Animals with a true coelom of the schizocoel type, usually bearing a shell composed mostly of calcium carbonate and secreted by a mantle. The mantle is always withdrawn at the rear to form a mantle cavity, which contains anus and gills. Primitive mollusks and gastropods use a tongue-like radula with embedded teeth to scrape encrusted algae from rock surfaces.
  • Class Monoplacophora (primitive mollusks): Mollusks with a simple dome-shaped or low conical shell. Muscles, blood vessels, and other structures segmentally arranged. Digestive tract simple.
  • Class Gastropoda (snails and slugs): Body usually undergoes asymmetrical torsion (twisting and coiling). One-piece (univalve) shell, usually coiled. Most species herbivorous. Well-developed head, sense organs, and nervous system. Locomotion typically by creeping on a muscular foot.
  • Class Polyplacophora (chitons): Simple, flattened body, with shell divided into several overlapping plates that permit some flexibility. Head small but radula well-developed and used in feeding.
  • Class Bivalvia or Pelecypoda (clams, etc.): Body usually symmetrical, narrowly compressed from side to side. Two-piece (bivalve) shell; left and right halves are often mirror images (except at hinge). Many species filter-feed, straining small particles from the water. Head and sense organs poorly developed. Muscular foot hatchet-shaped (flattened side-to-side, like a wedge), often used in burrowing. (The old name "Pelecypoda" means "hatchet-foot".)
  • Class Scaphopoda (tusk-shells): Small mollusks with tusk-like shells. Mantle cavity runs for entire length of shell along posterior margin; water passes through mantle cavity, exiting through hole at the top.
  • Class Cephalopoda (octopuses, squids, nautiloids, etc.): Body usually symmetrical. One-piece shell is symmetrically curved or coiled in median plane, or often lost. Most species are actively swimming predators. Head very well-developed, with sense organs (especially eyes), brain, and beak. Muscular foot subdivided into numerous tentacles. Body doubled over, with mantle cavity (originally rear) tucked beneath head and opening forward. Frequent "ink glands" that secrete dark, inky fluid to confuse predators.


Phylum Annelida: Segmented worms. Complete digestive tract (with both mouth and anus) runs nearly the entire length of the body. Outer covering of chitin is thin, flexible, and prevents fluid loss. The true coelom and most internal organs are segmentally arranged. Blood cicrulates in closed vessels only. Advanced excretory organs (nephridia) are present. Some ability to regenerate missing parts after injury.
All annelids exhibit Metamerism, a division of the body into numerous similar segments.
  • Class Polychaeta: Largest group, mostly marine. Sense organs and nervous system highly developed; several setae (bristles) per segment.
  • Class Oligochaeta (earthworms): Poorly developed head; only one pair of setae per segment. Important to soil because their digestive wastes leave behind soil nutrients and their tunnels let air reach plant roots.
  • Class Hirudinea (leeches): Mostly parasitic, live in fresh water, attach to the outside of animals and suck blood. Leeches have degenerate anatomy: fewer sense organs, fewer segments, etc.
Locomotion in annelids (controlled separately in each segment):
  • Each segment contains a walled-off portion of the body cavity.
  • Muscles parallel to the body axis can shorten segments; these segments swell and anchor into the surrounding sand or soil.
  • Muscles perpendicular to the body axis will lengthen body segments and cause them to push forward.
  • Nervous system produces rhythmic waves of shortening and waves of lengthening among the segments.
  • Small bristles (setae) may help anchor the shortened segments.
  • REVIEW:         Study guide and vocabulary

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