Metamerism (Segmentation) Body divided into longitudinal series of similar repeared units (segments) Characteristic of many bilateral phyla,

a, especially annelids, arthropods, and chordates Molluscs show primitively, annelidmollusc link? Acron, pygidium (or telson), variable number of intermediate

Near-Perfect Segmentation Everything (appendages, muscles, coelom, nerves, blood vessels, reproductive system, excretory system) repeated in every segment Primary advantage of segmentation divides body into compartments Regulate more or less independently Annelids exhibit closest to perfect

Framework for Specialization Regional specialization Some seen in annelids where? Processes of regional specialization
Restriction Divergence Fusion

Burrowing Theory R. B. Clark Offered as an explanation of significance of segmentation

True Segmentation Why annelids but not cestodes? Segments are true repetitions New cestode proglottids form anteriorly Annelid segments in front of pygidium segments are derived characteristic in cestodes

Phylum Annelida Segmented Roundworms Metamerism Eucoelomate Bilateral symmetry Protostome Closed circulatory system Most < 10 mm long, typical range 2 mm 7 cm; Australian earthworm ~ 3 m

Chaetae Chitinous bristles Extend from epidermis Primarily traction

Evolution and Significance of Segmentation Read this section Some information already covered (regional specialization) Burrowing theory

Annelida Diversity >12,000 species Three major classes

Oligochaeta Polychaeta Hirudinea

Some debate on phylogeny (more later)

Class Oligochaeta 3,500 species Includes earthworm Occupy most major habitats Most freshwater and terrestrial Marine species (~200) return to ocean Oligochaete is an oligochaete

Class Oligochaeta Uniform mode of feeding Burrowing detritivores Digestive system adapted for this
(organs and typhlosole)

Also of note Produce cellulase and chitinase Calciferous gland aid in gas exchange??

Circulatory System Matches general annelid circulatory system Hearts number varies 1 7 Hearts and dorsal vessel contain valves Oligochaeta with extensive integumental capillaries (gas exchange)

Reproduction Asexual reproduction common, especially aquatic forms Some no evidence of sex Regeneration if enough body intact anteriorly Direct development Eggs deposited in cocoon

Class Hirudinea About 600 species, most abundant in north temperate zones Freshwater (majority), marine, and terrestrial Most are carnivores Smallest ~1 cm, most 2 5 cm, one Amazon species 30 cm long Smallest individuals larger than smallest oligochaetes and polychaetes

Class Hirudinea Many common features - Oligochaeta Lack parapodia Lack head appendages Hermaphroditic Direct development Eggs deposited in cocoon

Hirudinea Evolution Probably evolved from freshwater oligochaete Originally may have been ectocommensal

Class Polychaeta Most diverse of annelids >8,000 species Most are marine Diversity evident 86 families Can be split into two former taxonomic groups
Errantia active crawlers and swimmers Sedentaria burrowers Adaptations reflect lifestyle

See Fig 13-35

See Fig 13-47

See Fig 13-52

See Fig. 13-48

 These adaptations are a reflection of the life-style to which these organisms have adapted A wide range of modes of feeding (in contrast to oligochaetes and leeches Wide range of reproductive strategies Big point of reproduction trochophore larva