第28章無脊椎動物

Phylum Annelida comprises the true, segmented worms. These animals are found in marine, terrestrial, and freshwater habitats, but the presence of water or humidity is a critical factor for their survival in terrestrial habitats. The annelids are often called “segmented worms” due to their key characteristic of metamerism, or true segmentation. Approximately 22,000 species have been described in phylum Annelida, which includes polychaete worms (marine annelids with multiple appendages), and oligochaetes (earthworms and leeches). Some animals in this phylum show parasitic and commensal symbioses with other species in their habitat.

Annelids display bilateral symmetry and are worm-like in overall morphology. The name of the phylum is derived from the Latin word annulus, which means a small ring, an apt description of the ring-like segmentation of the body. Annelids have a body plan with metameric segmentation, in which several internal and external morphological features are repeated in each body segment. Metamerism allows animals to become bigger by adding “compartments,” while making their movement more efficient. The overall body can be divided into head, body, and pygidium (or tail). During development, the segments behind the head arise sequentially from a growth region anterior to the pygidium, a pattern called teloblastic growth. In the Oligochaetes, the clitellum is a reproductive structure that generates mucus to aid sperm transfer and also produces a “cocoon,” within which fertilization occurs; it appears as a permanent, fused band located on the anterior third of the animal (Figure 28.29).

The epidermis is protected by a collagenous, external cuticle, which is much thinner than the cuticle found in the ecdysozoans and does not require periodic shedding for growth. Circular as well as longitudinal muscles are located interior to the epidermis. Chitinous bristles called setae (or chaetae) are anchored in the epidermis, each with its own muscle. In the polychaetes, the setae are borne on paired appendages called parapodia.

Most annelids have a well-developed and complete digestive system. Feeding mechanisms vary widely across the phylum. Some polychaetes are filter-feeders that use feather-like appendages to collect small organisms. Others have tentacles, jaws, or an eversible pharynx to capture prey. Earthworms collect small organisms from soil as they burrow through it, and most leeches are blood-feeders armed with teeth or a muscular proboscis. In earthworms, the digestive tract includes a mouth, muscular pharynx, esophagus, crop, and muscular gizzard. The gizzard leads to the intestine, which ends in an anal opening in the terminal segment. A cross-sectional view of a body segment of an earthworm is shown in Figure 28.30; each segment is limited by a membranous septum that divides the coelomic cavity into a series of compartments.

Most annelids possess a closed circulatory system of dorsal and ventral blood vessels that run parallel to the alimentary canal as well as capillaries that service individual tissues. In addition, the dorsal and ventral vessels are connected by transverse loops in every segment. Some polychaetes and leeches have an open system in which the major blood vessels open into a hemocoel. In many species, the blood contains hemoglobin, but not contained in cells. Annelids lack a well-developed respiratory system, and gas exchange occurs across the moist body surface. In the polychaetes, the parapodia are highly vascular and serve as respiratory structures. Excretion is facilitated by a pair of metanephridia (a type of primitive “kidney” that consists of a convoluted tubule and an open, ciliated funnel) that is present in every segment toward the ventral side. Annelids show well-developed nervous systems with a ring of fused ganglia present around the pharynx. The nerve cord is ventral in position and bears enlarged nodes or ganglia in each segment.

Annelids may be either monoecious with permanent gonads (as in earthworms and leeches) or dioecious with temporary or seasonal gonads (as in polychaetes). However, cross-fertilization is preferred even in hermaphroditic animals. Earthworms may show simultaneous mutual fertilization when they are aligned for copulation. Some leeches change their sex over their reproductive lifetimes. In most polychaetes, fertilization is external and development includes a trochophore larva, which then metamorphizes to the adult form. In oligochaetes, fertilization is typically internal and the fertilized eggs develop in a cocoon produced by the clitellum; development is direct. Polychaetes are excellent regenerators and some even reproduce asexually by budding or fragmentation.