Kingdom Animalia: Phylum Platyhelminthes Introduction: After the radiata, the remaining portion of the eumetazoans belong to the

bilateria---animals with a bilateral symmetry. This advanced symmetry was actually foreshadowed by the existence of a planula in the coelenterates, where the larvae is elongated along the antero-posterior axis. Bilateral symmetry is actually correlated with motility, in contrast with radial symmetry that is more often times associated with a sessile mode of existence. Furthermore, it is in the bilateria that cephalization starts to develop and is already evident in the most primitive, the basal stock of all bilateria, the platyhelminthes or flatworms. History: The flatworms were known from the ancient times, since some of these, the tapeworms in particular, are human intestinal parasites. Linnaeus initially placed all the invertebrates except arthropods in one class, the Vermes; and the worms he placed in one order, the Intestina. Later on, it was recognized that some of these worms are segmented while others are not. The unsegmented worms were called Entozoa since most of the species known at that time were intestinal parasites. In 1851, the orders under which are the flatworms were finally united into one class---Platelmia, and today is known as the Platyhelminthes. Definition: The platyhelminthes are tripoblastic, acoelomate, dorseventrally flattened metazoans with bilateral symmetry; with a well developed organ systems as compared with the coelenterates. I. General Morphology and Physiology Form. platyhelminths are dorsoventrally flattened, oval to elongate, bilaterally symmetrical; size ranges from microscopic to more than 60 cm long, ex. Lake Baikal triclad, Cephalus arecepta Body wall. they are tripoblastic (with 3 body layers), acoelomate, no body cavity but instead the space between the body walls and internal organs are filled with loose tissue called parenchyma Integumentary System. consists of an epidermis which may be cellular or acellular; may be ciliated or covered by a thick cuticle Muscular System. beneath the epidermis; consists of an outer circular and inner longitudinal layer Digestive System. generally sac-like and incomplete (the mouth is the only opening by which food and solid wastes pass through; entirely absent in some species Excretory System. protonephridial system with flame cells or solenocytes Reproductive System. almost entirely hermaphroditic, but individuals rarely self-fertilize; sperm flagellate; some possess 9+1 instead of the usual 9+2 metazoan construction Nervous System. 1. network of nerve fibers with cephalization and with several longitudinal nerve cords 2. sense organs are present usually in free-living forms or free-living phases of ciliated ectodermal pits

parasitic forms; these may be eyes, ostocysts, taste cells, tentacles and

II. Taxonomy of Platyhelminthes The flatworms can be divided into four classes: the Turbellaria, the Trematoda, the Monogenea and the Cestoda. Turbellarians are mostly free-living, while the last three are entirely parasitic. Because of their varying modes of existence, each class has developed independently specialized morphology and physiology that is best understood if treated separately. A. Class Turbellaria. mostly free-living with digestive system; body usually covered with cilia; rhabdites typically present; suckers absent with rare exceptions; life cycles simple External morphology. from oval to elongate; with head projections: short, pharyngeal or dorsal tentacles (Temnocephala) or lateral projections called auricles in freshwater planarians; coloration is mostly in shades of black, brown or gray and some marine turbellarians are brightly colored Habitat. largely tropical but a few are temperate; primarily aquatic, majority are marine; some are pelagic, most are benthic on sand or mud, on vegetation and under stones; many are meiofauna; freshwater forms also exist but are confined strictly to very humid damp or wet areas; they usually hide under logs or stones at day time and only come out at night to feed Body wall. a. epidermis i. outermost layer of ciliated epidermis bearing surface microvilli; cilia are limited to the ventral surface in triclads and polyclads; few have syncytial epidermis like rhabdocoels and temnocephalids; most have basal lamina beneath except acoels and nemertodermatids ii. provisions for intracellular support include: terminal web. weblike sheet of actin filaments within the epidermis basal lamina. with calcareous spicules in kalyptorynch rhabdocoels mesh of ciliary rootlets in acoels parenchyma

iii. numerous gland cells are characteristic of turbellarian epidermis, although most are situated in the mesenchyme and only the necks extend through the epidermis; these may function to provide adhesion, cover the substratum while moving and capture and envelope prey; there are several gland systems: - duogland adhesive systems. present in majority of turbellarians; provides temporary adhesion to the substratum especially for interstitialspecies; each complex may project from the surface as a papilla, and contains the following:

viscid gland. provides adhesive material releasing gland. emits substance that breaks the attachment of the viscid material and enables the animal to break loose from the substratum anchor cell. serves as the attachment surface through which the glands pass and open production used for locomotion and adhesion in larvae

- frontal gland. a primitive feature; for defense and slime - caudal gland - marginal ring

iv. aside from the gland systems, other structures which serve the same purpose are: muscular suckers. as in large species such as polyclads and temnocephalids and ectocommensals on crustaceans adhesive cilia. in the acoel Paratomella rubra

v. rhabdoids. membrane-bound rod-shaped secretions that form mucus; most common type is rhabdite characterized by a specific, layered ultrastructure in archoophoran macrostomids and polyclads and in neoophorans vi. nematocyts derived from eaten hydroids are used for defense (a) (b) musculature. beneath the epidermis and is of smooth type; in some species, is of epitheliomuscular type; outer circular, middle diagonal and inner longitudinal layers; dorsoventral strands connect the 2 surfaces of the body wall parenchyma. connective tissue between body wall musculature and gut; composed of mesenchymal cells in a fibrous extracellular matrix; extracellular matrix is fluid and forms pseudocoel in freshwater catenulids and is absent in acoels; types of cells found: epidermal replacement cells. Necessary because of an absence of mitosis in adult epidermis; each contains a cluster of centrioles which later become ciliary basal bodies neoblasts. totipotent cells important in wound healing and regeneration fixed parenchymal cells. large branched cells that make contact with and interjoins other cells and tissues; gap junctions suggest role in physiology of turbellarians chromatophores. parenchymal cells with hemoglobin. color the body red and functions in 3. Locomotion ciliary locomotions. small turbellarians muscular undulations. swimming in large polyclads, use of ventral sole in triclads and ditaxical shuffling in other polyclads retraction movements. some interstitial turbellarians peristaltic movements. other interstitial turbellarians mucus thread as suspension bridge. terrestrial triclads somersaulting movements. temnocephalids that are ectocommensals on crustaceans oxygen storage

5. Nutrition. the gut is a blind sac with the mouth serving as portals of entry and exit; may be predaceous, scavengers or grazers; digestion is first extracellular with pharygeal enzymes and endopeptidase from the intestines, then intracellular by the phagocytic cells where endopeptidase continue their action mouth. along the midventral axis; absent in catenulid Paracatenula urania pharynx. a ciliated tube which can be: i. simple. most primitive condition; in macrostomids and catenulids; absent in acoels ii. plicate or folded. lead to the development of a muscular pharynx that can be retracted or extended; in poly- and triclads iii. bulbous pharynx. derived from the plicate condition through reduction of pharyngeal cavity and separation by a muscular septum, this can also be protruded; in rhabdocoels and predatory, commensal or parasitic species intestines. extend directly from the pharynx and may be: i. simple sacs. small turbellarians such as macrostomids, catenulids and rhabdocoels ii. long tubes with lateral extensions or diverticula. serve to increase the surface area for digestion and absorption; single layered and composed of phagocytic and gland cells and ciliated in primitive forms iii. syncytial mass. in acoels which lack a permanent gut cavity 6. Nervous system and Sense organs. The primitive plan is a subepithelial nerve net plus 3 or 4 pairs of nerve cords interconnected by commissures: dorsal or dorsolateral, lateral or marginal, ventrolateral and ventral. A brain containing a statocyst terminates the anterior portion. Advance forms deviate from this plan through a reduction of some pairs of nerve cord which is characteristic of the annelid-arthropod line. longitudinal nerve cords and development of a priminent ventral

 The nervous system lacks ganglia except in the brain. Sense organs include: i. pigment-cup eyes. orientation to light; most turbellarians are negatively phototactic; polyclads and land planarians

ii. statocyst. gravity receptors; consists of a capsule that encloses a fluid-filled cavity and a central concretion called statolith; most conspicuous but occurs only in acoels, nemertodermatids, proseriates and some catenulids iii. ciliary receptors. mechanoreceptors; concentrated on tentacles, auricles and body margins iv. ciliary or microvilli pits. chemoreceptors used in locating food; catenulids 7. Excretion and osmoregulation. protonephridial type (flame cell) which consists of: (a) (b) (c) (d) branched tubules terminating in a number of blind capillaries tubule cell and cap cell bearing 2 or more flagella terminate these branches; the 2 cells actually interdigitate their flagella forming a barrel-shaped region which are very thin areas which are the entrance into the system nephridiophores terminate at the other end of the system and serves as excretory pores by which fluid wastes are voided into the exterior through the epidermis evidences tend to support the claim that the system is more of osmoregulatory than excretory: 8. Reproduction a. Asexual i. transverse fission / budding. freshwater planarians in temperate regions reproduce by fission in summer and sexually during fall with shorter day length and lower temperature; with complete separation where the 2 parts regenerate the missing portions, and with incomplete separation that the individuals may remain attached (each called a zooid) to form a chain, each zooid detaches from the chain once it has developed sufficiently (paratomy), Catenula, Stenostomum, Microstomum ii. architomy. simultaneous fragmentation of the body into several pieces, Phagocata iii. regeneration. freshwater planarians iv. parthenogenesis. b. Sexual. all turbellarians, except for a few parasitic species, are hermaphrodites, reproducing by way of copulation and internal fertilization; the basic and primitive plan is as follows: i. male. single pair of testes --> sperms ducts --> seminal vesicle --> penis bulb (bears gland/prostatic vesicles and armed with a stylet) --> penis (lies in male genital canal, which opens through male gonophore into posterior ventral surface) ammonia is easy to remove (across body surface) considering the small size of the turbellarians the system is most well-developed in freshwater species and are least developed or even absent in marine forms

ii. female. single pair of ovaries --> oviduct --> bursa (short term sperm storage center) --> vagina (terminates in female gonophore, surrounded by cement glands) iii. types of ovaries serve as basis for distinguishing 2 levels of organization in Turbellaria: archoophoran (single ovary) and neoophoran (an ovary and vitellaria) iv. more advanced forms may exhibit any of these variations: more than a pair of testes/ovaries muscular penis without stylet cirrus (eversible copulatory organ) multiple male parts (for defensive purpose) ovary may differentiate into vitellarium (only produces eggs) and yolk gland (only produces yolk-filled nurse cells) and unite into an ovovitellarium seminal receptacle (long term sperm storage center) uterus (temporary storage center for ripe eggs) egg capsules a single gonophore and genital atrium into which both male and female systems open v. Hypodermic impregnation occurs when penis with a stylet is rammed into the body wall of the partner and sperm is injected into the parenchyma. vi. Sperm morphology is diverse: uniflagellate 9x2+2 sperm of Nemertodermatida; sperm of some Acoela without free flagella (two 9x2+2 axonemes incorporated into cell); biflagellated sperm of Polycladida and Rhabdocoela, both with 9x2+1 axonemes. vii. Development is lecithotrophic and direct and with spiral determinate cleavage; stereogastrula by epiboly; most are withoutlarvae but 2 forms exist: Muellers larva (8 armed) and Goettes larva (4 armed) which are ciliated and swim for a few days before settling down and feeding 9. Interspecific Relationships Bdelloura. ectocommensal triclad on book gill of horseshoe crabs Acoel sp. harbon green zoochlorellae or golden zooxanthellae or diatoms in their parenchyma Fecampiidae. rhabdocoel endoparasitic in hemocoel of crustaceans

 Some species parasitic in gut and body cavities of molluscs, crustaceans, echinoderms and skin of fishes. 10. Taxonomy of Turbellaria. The modern classification divides them into three major groups, which may or may not be closely related to each other. These are the Acoelomorpha (Nemertodermatida and Acoela), Catenulida (Catenulida) and Rhabditophora (remaining orders). The first two groups have archoophoran organization, as do the primitive members of the Rhabditophora (i.e., the Macrostomida, Haplopharyngida, and Polycladida); the rest of the Rhabditophora have the neoophoran organization. Class Trematoda - entirely parasitic flatworms with digestive system; cilia absent in adults but instead a well-developed cuticle is present; possess orally and ventrally located adhesive organs; life cycles complex; descriptions mainly for digeneans. External morphology. size ranges from 0.2 mm to 6.0 cm in length; dorsoventrally flattened, some thick and fleshy, others are long and threadlike; an oral and, if present, a ventral sucker (acetabulum) that prevent dislodgement and aid in feeding. Habitat. many infect the gut or gut derivatives of their definitive host; lungs, bile ducts, pancreatic ducts and intestines are common sites Body wall. body covered by a nonciliated cytoplasmic syncytium, the tegument (provides protection against hosts enzymes in gut-inhabiting species, site of gas exchange, absorbs glucose and amino acids in endoparasites); beneath are consecutive layers of circular, longitudinal and diagonal muscle Locomotion. aquatic larval stages swim with the help of either cilia, tails, etc depending on the stage of the species adults are less mobile once they reach their habitat in their respective hosts, only crawling or wriggling to reach their target or sometimes just to shift positions Nutrition. The mouth is surrounded by a powerful, muscular sucker, which is used for attachment and aids the transport of food into the mouth.

The bulbous pharynx ingests cells and cell fragments, mucus, tissue fluids, or blood of the host on which the parasite feeds. The pharynx passes into a short esophagus leading to two blind intestinal caeca that extend posteriorly along the length of the body. Digestion is primarily extracellular in the caeca. Respiration. facultative anaerobes; amount of oxygen utilized in respiration depends on the location within the host and also on the develomental stage of parasite Nervous system and sensory organs. A pair of anterior cerebral ganglia from which longitudinal nerve cords extend posteriorly, of which the ventral pair is the most highly developed. The fluke body surface has a variety of sensory papillae, and ocelli occur in many miracidia and some cercariae. Excretion and osmoregulation. Protonephridia: a pair of longtitudinal collecting ducts empty into a single posterior bladder and nephridiophore. The nephridia excrete water and waste metabolites, such as unwanted iron from hemoglobin in blood-feeding worms. Reproduction. similar to neoophoran turbellarians but egg production is many times a. The generalized reproductive system is as follows: i. male: 2 testes --> sperm ducts --> external seminal vesicle --> cirrus sac (with internal seminal vesicle, prostate glands, cirrus) --> genital atrium --> gonopore ii. female: 1 ovary --> oviduct (with duct from seminal vesicle, commont duct from right and left vitellaria) --> ootype (surrounded by Mehlis gland) --> uterus --> genital atrium --> gonopore iii. In some digeneans, a vestigial vagina (Laurers canal) is present. b. During copulation, sperm exchange is mutual, and cross fertilization is the general rule, although self-fertilization is known to occur in rare cases. c. Cirrus of one worm is inserted via the gonopore into the uterine opening of the other worm, and sperm are ejaculated. The prostate gland provides semen for sperm survival. Sperm travel up the uterus through the ootype to be store in the seminal receptacle. d. Once released from the ovary, eggs are fertilized either in the oviduct or within the ootype. Because digenean eggs are ectolecithal, the yolk is supplied by the yolk cells. Egg capsules pass along the entire length of the uterus where embryonic development begins and eventually are released from the gonopore. e. Some variations to this basic plan include: dilation of the vitelline duct to form a vitelline reservoir one or two vaginae may be present greater than in free-living flatworms

f. Generalized life cycle: eggs (in feces) --> miracidium (ciliated free-swimming larva) --> sporocyst (with germ cells; inside 1IH) --> daughter sporocysts or redia (with germ cells) --> cercariae (inside 2IH) --> (encysts into) metacercariae --> metacercariae (excysts inside DH) --> adult (sexual maturity at target organ)

Class Cestoidea. entirely endoparasitic flatworms devoid of a digestive tract; cilia are absent in adults but instead possess a well-developed tegument; typically with more than one host per life cycle The cestodes are composed of two subclasses that differ considerably, the eucestoda and the cestodaria. The discussion of these two groups will be done separately to avoid confusion.

Subclass Eucestoda. External morphology. the body of the adult can be divided into: i. scolex. the head which may bear several types of holdfast organs to maintain its position inside the hosts gut such as suckers, grooves, hooks, spines, glands or simple or absent altogether. Sucker-like organs of attachment are mainly of three types: acetabulum. muscular, cup-shaped, usually four bothridia. muscular, highly mobile projections with leaf-like margins bothria. usually 2 (up to 6) shallow pits or longer grooves In acetabular scolices, hooks are usually borne anterior to the suckers, on a protrusible cone-shaped extension called rostellum. ii. neck. narrow region behind the scolex, its posterior region contains germinal cells that give rise to the strobila; if neck is absent similar present behind the scolex. cells are

iii. strobila. the body of the tapeworm which is actually composed of a series of segments called proglottids; each contains at least a set of both male and female reproductive organs, and according to the degree of development of these may be classified into: immature. sexually non-functional mature. functional gravid/ripe. filled with eggs

A gravid proglottid upon reaching the end of the strobila may detach (apolysis) or may remain until senile or exhausted (pseudopolysis/anapolysis). In apolysis, the eggs are released either through the gonopore or through breaks in the proglottid. In anapolysis the eggs are released upon disintegration of proglottid. If a proglottid detaches when still immature and grows to maturity in the gut, it is called hyperapolysis. Structurally, a proglottid is called craspedote if the posterior edge of one proglottid overlaps the anterior portion of its succeeding proglottids and acraspedote if no fold or overlap is present. Habitat. all eucestodes are endoparasites in the gut of vertebrates, with rare exceptions in the coelomic organs Body wall. I. II. an outer well-developed cuticle, more properly termed tegument since it is a living layer; has outer plasma membrane thrown into projections called microtrichia which serve to increase the absorptive surface area.

underneath this layer are the outer circular and inner longitudinal muscle layers; in addition, a secondary parenchyma musculature of longitudinal, transverse and dorsoventral fibers encloses the interior parenchyma. Locomotion. ciliated larvae are free-swimming (but not all); adults exhibit minimal locomotory capabilities, although they squirm about in the hosts gut Nutrition. no GIT; acquires nourishment by active transport through the tegument Respiration. facultative anaerobes Nervous system and sense organs. an anterior nerve mass or brain lies in the scolex two lateral longitudinal nerve cords extend posteriorly through the strobila (but there may be pairs of dorsal, ventral and accessory lateral cords) ring commissures/interproglottidal commissures connect these longitudinal nerve cords in each proglottid forming a ladder-like appearance

Excretion and osmoregulation. There are two main parts of canals running along the length of the strobila: the ventrolateral pair which is a bit larger in diameter, and the dorsolateral pair A transverse canal joins the ventrolateral pair at the posterior margin of each proglottid. Flame cells scattered in the parenchyma empty their contents into collecting ductules/tubules that connect to these longitudinal cords. Unlike the turbellarians and trematodes, the protonephridial system of cestodes have been found to be excretory ( microvilli were foundlining the excretory ducts which may function in the resorption of certain electrolytes, etc.); and osmoregulation seems to be taken cared of by the tegument which can take amino acids by active transports to counteract the changing osmolarity of the few surrounding medium

Reproduction. tapeworms are monoecious, with a few very rare exceptions; most species possess a complete set of reproductive organs per proglottid, but some may have two pairs, a few have one male and two female systems in each proglottid; most species exhibit androgyny/protandry, a few exhibit gynandry/protogyny i. the basic plan is as follows:

male: one to many testes each has a vas deferens --> unite to form vas deferens --> external seminal vesicle --> ejaculatory duct &/or internal seminal vesicle (both of which are inside the cirrus sac) muscular cirrus --> genital atrium female: ovary --> oviduct joined by ducts from vitelline glands --> ootype surrounded by Mehlis gland --> two canals: uterus, a blind sac, and seminal receptacle --> vagina --> common gonopore

ii. If two or more worms are present, cross-fertilization is the rule; however, as often the case, only one worm is present so self-fertilization either within the same proglottid or with other proglottid occurs. In forms without vagina, hypodermic impregnation is the method of fertilization. iii. Generalized life cycle: (same or egg --> oncosphere (hexacanth (6-hooked) larva; inside IH) --> different hosts) metacestode (parenteral/extraintestinal site) --> adult

iv. There are two basic variations: Infective stage oncosphere differentiating metacestode metacestode w/ free-swimming larva coracidium procercoid pleocercoid w/out free-swimming larva oncosphere cysticercoid

DETAILED TAXONOMY OF PHYLUM PLATYHELMINTHES Class: Turbellaria Subclass: Archoophora orders that reflect a more primitive level of organization. Vitellaria absent; eggs entolecithal; cleavage spiral. Order Nemertodermatida. small marine species similar to acoels but possessing uniflagellate sperms and an epithelial digestive tract. Statocyst usually with two statoliths. ex. Nemertoderma Order Acoela. small marine flatworms, usually less than 2 mm in length. Statocyst with one statolith. Mouth and sometimes a simple pharynx present, but no digestive cavity. Protonephridia absent. Gonads often not bounded by a cellular wall. Oviducts absent. ex. Amphisclopos, Anaperus, Afronta, Polychoerus, Convoluta, Archaphanostoma, Pseudaphanostoma, Diopisthoporus, Paratomella. A few species are commensal within the intestine of various echinoderms. Order Catenulida. mostly small, freshwater species having a simple pharynx; ciliated saclike intestine; and unpaired gonads, with the male gonopore dorsal above the pharynx. Statocyts sometimes present, usually with one statolith. No male gonoducts. ex. Stenostomum, Catenula Order Macrostomida. small marine and freshwater species having a simple saclike ciliated intestine, a simple pharynx, and one pair of ventrolateral nerve cords. ex. Macrostomum, Microstomum Order Polyclada. marine flatworms of moderate size, averaging 3-20 mm in length, with a greatly flattened and more or less oval shape. A pair of anterior marginal or dorsal tentacles may be present. Many are brightly colored. Intestine elongated and centrally located, with many highly branched diverticula. Plicate pharynx either an anteriorly directed tube or pendant from the roof of the pharyngeal cavity. Eyes numerous. ex. Gnesioceros, Leptoplana, Notoplana, Stylochus, Prostheceraeus, Pseudoceros Order Lecithoepitheliata. marine and freshwater species in which ovary produces eggs surrounded by follicle-like yolk cells. The mouth and complex pharynx are at the anterior end of body, intestine simple. ex. Prorhynchus Subclass: Neoophoran orders that reflect an advanced level of organization. Vitellaria present; eggs ectolecithal; and development greatly modified from the spiral pattern. Order Prolecithophora. usually small. marine and freshwater species having a plicate or bulbous pharynx and a simple intestine. Ovary produces eggs and follicle-like yolk cells. ex. Plagiostomum, Hydrolimax Order Proseriata. small, mostly marine turbellarians, including many interstitial forms. Statocyst with one statolith. Pharynx is plicate and tubular, but is not branched. ex. Otoplana, Monocelis, Nemertoplana Order Neorhabdocoela. a large group of small marine and freshwater species, having a bulbous pharynx, a simple straight intestine, and one pair of nerve cords. Suborder Typhloplanoida. mouth usually located in the middle and pharynx oriented at right angles to long axis of body. Contains marine and freshwater, free-living species. ex. Mesostoma Suborder Dalyellioida. mouth typically at anterior end of body and pharynx oriented parallel to long axis of body. Contains marine and freshwater species, some of which are commensal and parasitic on and within snail, clams, sea urchins, and sea cucumbers. ex. Anoplodiera, Syndesmis, Kronborgia Suborder Kalyptorhynchia. mouth anterior to middle of body. Anterior protrusible, muscular proboscis often bears cuticularized hooks or teeth. Contains mostly marine interstitial species. ex. Gyratrix, Gnathorhynchus Suborder Temnocephalida. commensal and parasitic on crustaceans, molluscs, and turtles. Posterior ventral surface provided with an adhesive disc, and anterior margin bears finger-like projections, by which worm moves leechlike on its host. ex. Temnocephala

Order Tricladida. relatively large, marine, freshwater, and terrestrial turbellarians. Pharynx is plicate, tubular and posteriorly directed; gut has three branches. Among marine species, Bdelloura is commensal on the book gills of horseshoe crabs. The freshwater species are known as planarians and include Planaria, Dendrocoelum, Procerodes, Dugesia, Phagocata, Polycelis, Procotyla. Land planarians include Bipalium, Orthodemus, Geoplana. Class: Trematoda Subclass Digenea. with characters of the class; digeneans can be grouped according Group 1: Gymnocephalous cercariae. large head with simple tail Family Fasciolidae. first intermediate host (1IH) snail Lymnea philippinensis, second IH (2IH) any water plant such as kangkong or water crest, definitive host (DH) in the liver and bile ducts of man (sheep and related cattle are reservoir hosts (RH)), ex. Fasciola hepatica (sheep liver fluke). Belongs to the Order: Strigeidida. Family Heterophyidae. smallest flukes; with an additional 3rd sucker called genital sucker or gonotyl (because it is on top of the genitalia); 1IH snail, 2IH fish Mugil tilapia of any other suitable fish, DH intestines of man (other fish-eating mammals serve as RH), ex. Heterophyes heterophyes. Belongs to the Order: Opisthorchiida. Group 2: Echinostome cercariae. with simple tail but the head possesses a collar of spines Family Echinostomatidae. oral sucker is surrounded by a collar of spines the number and arrangement of which is of taxonomic importance; 1IH snail Gyraulus convexiusculus, 2IH snail Pila luzonica, DH man, ex. Echinostoma ilocanum. Belongs to Order: Echinostomida. Group 3: Lophocercous cercariae. tail has fins Family Opisthorchidae. 1IH snail Bulinus, 2IH Cyprinoid fishes, DH man and other fish-eating mammals, ex. Clonorchis sinensis (Chinese liver fluke). Belongs to Order: Echinostomida. to the morphological type of their cercariae:

Group 4: Xiphidiocercariae. mouth bears a stylet for penetrating hosts Family Troglotrematidae. 1IH snail Antemelania asperata, 2IH mountain crab Parathelpusa Philippina , DH man and other crab-eating mammals, ex. Paragonimus westermani (lung fluke); Belongs to Order: Troglotrematidae. Group 5: Furcocercous cercariae. with forked tail Family Schistosomatidae. dioecious; only IH snail Oncomelania bupensis quadrasi , DH man and the habitat is the hepatic portal vein, ex. Schistosoma japonicum. Belongs to Order: Strigeidida. NOTE: Treat the Order as per classification but with no description, Pechinick is deficient on this part. Subclass Aspidogastrea. distinguishing feature is the adhesive organ, which is either single septate sucker covering the entire ventral surface or a longitudinal row of suckers; gut contains a single intestinal cecum; reproductive system similar to Digenea but there is only one testis; mostly endoparasites in the gut of fish and reptiles and in the pericardial and renal cavities of bivalve molluscs; life cycle involves one or two hosts, ex. Aspidogaster conchicola, Cotyaspis insignis (in freshwater mussels)

Class Monogenea. mostly ectoparasites of aquatic vertebrates, especially fishes, but amphibians and reptiles are also hosts; with an anterior sucker or prohaptor and a posterior sucker or opisthaptor that bears hooks and suckers; life cycle has no intermediate host and one egg (by way of ciliated larva, the oncomiracidium) give rise to one adult worm; with aerobic metabolism; neoophoran reproductive system but usually with single testis and extensive vitellaria. Subclass Monopisthocotyles. with simple opisthaptor; single vagina, single uterus with one egg, ex. Gyrodactylus in the cloaca of frogs and gills of freshwater fishes Subclass Polyopisthocotyles. complex opisthaptor; paired vagina and uterus, ex. Polystoma integerrimum in the bladder of frogs and toads and shows synchronization of parasite life cycle with that of the amphibian host Class: Cestoda Subclass Eucestoda. There are 12 orders under eucestoda largely based on the differences in scolex morphology, 5 of which are entirely parasitic in elasmobranchs. At this level, only those of medical importance will be considered. Order Cyclophyllidea. scolex usually with 4 suckers, rostellum may or may not be present, if present may or may not bear hooks; genital pores mostly lateral; uterine pore absent; only one IH (vertebrates, insects, mites, annelids and molluscs), parasitic in birds and mammals Taenia solium (pork tapeworm). scolex with rostellum and hooks; trilobed ovary; 7-12 thick lateral branches on each side of the main uterine stem, IH pig and man which harbors cysticercous cellulosae, DH man Taenia saginata (beef tapeworm). scolex without rostellum and hooks; bilobed ovary, 15-30 uterine branches; IH cattle and rarely man which harbor cysticercous bovis, DH man Taenia pisiformis. occurs in cats and dogs with rabbits as IH Order Pseudophyllidea. scolex with 2 bothria, with or without hooks; genital pore may either be lateral, dorsal or ventral; uterine pore present, dorsal or ventral; egg usually operculate, containing a coracidium; with 2 IH Diphyllobothrium latum (broad fish tapeworm). finger-shaped scolex with dorsal and ventral bothria; proglottids are wider than long; common genital pore opens midventrally; with coiled rosette-like uterus; 1IH copepod crustaceans Cyclops, Diatomus which harbor the procercoid, 2IH freshwater fishes which harbor the plerocercoid, DH man, dog, cat and less frequently 22 other fish-eating mammals

Other ORDERS mentioned in Pechinik are: Order: Tetraphylidea Order: Trypanorhyncha Order: Caryophyllidea Order: Spathebothriidea Subclass Cestodaria. no scolex but there may be small proboscis-like organ at anterior end; typically oblong , elongate, rounded posterior end sometimes forming a crenulated rosette or with frilled edges; gonopore near posterior end; monozoic orwith a single set of reproductive organs (as opposed to polyzoic eucestodes); ovary posterior, vitelline glands follicular, lateral; uterus N-shaped or looped, uterine pore near anterior end; testes in two lateral preovarian fields; the oncosphere is ciliated and called a lycophora; IH are invertebrates and DH are elasmobranchs and holocephali and chelonia, particularly in the coelom and gut, ex. Gyrocotyle fimbrata Kingdom Animalia: Phylum Nematoda Introduction After the Platyhelminthes, which are acoelomates, next in line among the bilateria would be the pseudocoelomates. These include the phyla Rotifera, Gastrotrichia, Kinorhyncha, Nematoda, Nematomorpha and Acanthocephala. Among these groups, the nematodes are the most important in terms of number of species, economic, ecological and most of all, medical significance since a large number of these worms are parasitic in both humans and animals. History All the phyla mentioned above were originally considered to be under one phylum, the Aschelminthes, which according to the classic work of Linnaeus was under the Entozoa, order Intestina, class Vermes. Later, it was realized that the groups belonging under phylum Aschelminthes had little in common with each other, and each of them was raised to phylum level. The nematodes in particular were separated from other Vermes by Rudolphi in 1608. Definition The nematoda are pseudocoelomate roundworms having a cylindrical body that tapers at both ends. They are bilaterally symmetrical, with cuticle as body covering, a complete digestive system, dioecious, and show considerable dimorphism. General Morphology and Physiology: I. Form. elongate and cylindrical, tapering at both ends, bilateral and ranges from less than a II. Body wall and Integumentary system. A. tripoblastic, pseudocoelomate, outermost non-cellular cuticle which can be 1. 2. 3. further divided into three regions: mm to more than a meter in length

cortical region. covered by ultra thin layer of lipid; external cortical layer is of a keratin-related material; inner cortical layer is made of collagen middle/homogenous/matrix region. also of collagen basal lamella. a layer of fine fibrous material B. the cuticle may have markings or ornamentations like shallow punctuations, deeper pores, spines of varying complexity, and lateral or sublateral thickenings called alae, which may be cervical or cephalic, caudal or longitudinal

C. the hypodermis lies beneath the cuticle which it secretes; it is syncytial and 4 longitudinal hypodermal cords divide the musculature into quadrants as they project into the pseudocoel III. Muscular system the musculature is composed entirely of longitudinal fibers. A. Nematode musculature can be classified according to the number of rows per 1. holomyarian. no rows or only 2 rows per quadrant 2. meromyarian. few rows (2-5) per quadrant 3. polymyarian. many rows per quadrant B. They are also classified according to the shape and distribution of contractile sarcoplasm: 1. platymyarian. cell ovoid in cross section, contractile portion adjacent to the hypodermis. 2. coelomyarian. cell spindle-shaped and formed into a narrow U-shaped structure, with the contractile portion at the curve of the U and attached to the hypodermis. 3. circomyarian. the contractile fibrils at the periphery entirely encircle the noncontractile portion of the cell. IV. The Coelom The nematode coelom is actually a pseudocoelom and not a true coelom since it is not lined by mesoderm and is actually the remnant of a persistent blastocoel. quadrant:

The pseudocoelom contains pseudocoelomic fluid or haemolymph that functions as the transporting fluid (in the absence of a circulatory system) since it bathes the internal organs such as the GIT and reproductive system, and the muscle cells. It also contains a variety of electrolytes, fats, carbohydrates, and proteins such as albumins, globulins, hemoglobin and several enzymes. A unique type of cell found in the pseudocoelom is the coelomocyte, usually 2-6, ovoid or much branched and found attached to adjacent organs. Their purpose is still obscure but there are hints that they may serve in the accumulation and storage of Vitamin B12 and in the synthesis and secretion of certain proteins. The pseudocoelom also functions as a hydrostatic skeleton, in which muscles will exert pressure as they contract against this enclosed fluid, which will cause the worm to become firm and capable of moving about. Digestive system.

V.

The GIT is complete, the mouth can be surrounded by a maximum of 6 lips. The buccal cavity follow which may be elongate, reduced or absent altogether and is of taxonomic significance; if it is lined by a thick cuticle, a buccal capsule is formed. The pharynx or esophagus is highly muscular with a number of bulbs that are of taxonomic importance. Its lumen is triradiate and lined with cuticle, and radial muscles run from the lumen to the body along the entire length of the esophagus. The esophageal glands that secrete digestive enzymes are also present. The intestine is a single-layered, non-muscular tube extending from the esophagus to the proctodeum; in females it terminates in a rectum and out through the anum while in males the rectum also receives the sperms and is therefore a cloaca.

VI. Excretion and osmoregulation. (either glandular and tubular types) A. glandular type. in free-living worms; involved in secretion of enzymes, proteins or mucoproteins B. tubular type. H- , U- or inverted U-shaped complex consisting of 2 lateral canals running along the length of the body with a transverse canal joining near the anterior end and opens to the exterior through a midventral excretory duct and pore; variations depend on reduction in the number of tubules C. renette cells. many parasitic species have a pair of large, granular subventral gland cells associated with the transverse ducts and are thought to be secretory in function VI. Nervous system and sense organs. intraepithelial and located within the epidermis, pharynx and hindgut. A. brain is a circumpharyngeal nerve ring, attached to it are paired ventral, lateral ventral being the largest and dorsal cephalic ganglia, dorsal pairs being the smallest and

ventral cord reaches the posterior end, it gives rise to two branches that extends dorsally to encircle the rectum, forming the rectal commissure or posterior nerve ring. principal sensilla include: 1. papillae. labial and cephalic papillae are low projections of the cuticle on the lips and head; outer labial and cephalic are mechanoreceptors, inner labial is a chemoreceptor 2. setae. elongated bristles on both head and body; touch receptors that cause the animal to withdraw from the stimulus

3. amphids. blind, pouchlike or tubelike invaginations of the cuticle; chemoreceptor well developed in marine species 4. phasmids. pair of unicellular glands which open separately on either side of the tail; chemoreceptor well developed in parasitic species 5. ocelli. in marine and freshwater nematodes, function is uncertain 6. stretch receptors. in epidermal cords of nematodes; probably regulate locomotor movements VIII. Reproductive System. most are dioecious but hermaphrodites are not uncommon A. sexual dimorphism is known: males are often smaller than females, posterior end of male may be curled like a hook or broadened into a bursa B. basic plan of male reproductive system is as follows: male: 1-2 testes --> sperm duct --> seminal vesicle --> vas deferens --> ejaculatory duct (prostate glands) --> cloaca

Accessory structures related to sperm deposition consists of a pair of sclerotized, acellular copulatory spicules that pry open the vagina during copulation. A gubernaculum or telomon (which are dorsal and ventral sclerotizations of the spicules out of the spicular sheaths. C. basic plan of female reproductive system is as follows: female: 1-6 ovaries --> oviduct --> spermatheca/seminal receptacle --> uterus --> Females of some species produce a pheromone to attract males. The number of reproductive tracts per female and their position relative to each other are given descriptive terms: ovijector --> vagina --> vulva cloacal wall, respectively) serves to guide the exsertion of the

1. monodelphic. 1 ovary per uterus with the vulva almost always near the uterus 2. didelphic. 2 ovaries and uteri 3. polydelphic. more than 2 uteri and associated structures 4. amphidelphic. 2 uteri converge from opposite directions 5. prodelphic. parallel and converge from an anterior direction D. 6. opisthodelphic. converge from a posterior direction Fertilization is internal through copulation --> fertilized eggs are released --> eggs embryonate--> eggs hatch into first larval stage L1 --> L1 molts 4 times going through L2, L3 and L4 --> adult

E. The infective stage in parasitic species is either the embryonated egg or L3 larva. Taxonomy of Phylum Nematoda Class Aphasmida/Adenophorea. variously shaped amphids behind the lips; amphids generally well developed; caudal and hypodermal glands common; exretory system without lateral canals but usually a single, ventral renette cell or entirely absent; deirids or cervical papillae always absent; mostly free-living Order Trichocephalida. Family Trichinellidae. anterior end more slender than posterior end; esophagus very slender, capillary-like tube embedded within one or more rows of large, glandular cells called stichocytes; both sexes with a single gonad; eggs with polar plugs, ex. Trichuris trichura (whipworm). in colon of man with the anterior end embedded into the mucosa, infective stage (IS) embryonated egg which is barrel-shaped. Order Euoplida Order Dorylaimida Order Mermithida Order Dioctophymata. stout, often very large; esophageal glands highly developed and multinucleate; male with ball-shaped muscular copulatory bursa without rays; both sexes with single gonad; eggs deeply scultured or pitted, ex. Dioctophyma renale (kidney worm). adults in kidneys of dogs, eggs out through urine, into the water and eaten by fish where it develops up to L3, dog eats infected fish, L4 and adult in dog (zoonotic) Class Phasmida/Secernentea. possess porelike amphids in the lateral lips; amphids generally poorly developed; caudal and hypodermal glands absent; excretory system with one or two lateral canals with or without renette cells; deirids commonly present; mostly parasitic Order Rhabditida. tiny worms commonly with 6 lips; muscular esophagus divided into an anterior corpus, median isthmus, and posterior bulb, with a pseudobulb often present between corpus and isthmus; bulb usually absent in parasitic stages; tail of both sexes conical; gubernaculum usually present; parasitic generations parthenogenetic or hermaphroditic, alternating with gonochoristic, free-living generations, ex. Turbatrix aceti (vinegar eel). free-living found in fermenting juices Order Strongylida. commonly long and slender; esophagus swollen posteriorly but lacks a bulb; male with well developed copulatory bursa supported by rays; eggs thin-shelled and rarely develops beyond morula when laid, ex. Ancylostoma duodenale (human hookworm). equipped with buccal capsule and large teeth for hanging on to intestinal mucosa, sucks blood, eggs passed through feces, hatches into rhabditiform larva, which is free-living, transforms into filariform larva L3 and penetrates skin of DH, through blood as L4 and reaches intestines to become adult Order Ascaridida. Family Ascarididae. commonly large, stout worms; usually with 3 lips, less often 2 or none, muscular occasionally with one or two appendices at the sculptured uterine layer, unembryonated when laid, ex. Ascaris lumbricoides (human giant intestinal roundworm). adults lay eggs in small intestines, out through feces, embryonation in the soil, bloodstream, L2 to L3 in lungs, L3 out of respiratory tract and gets swallowed to become L4 and finally adult in small intestine Family Oxyuridae. medium-sized to small; commonly with sharply pointed tails; lips if present usually 3, alae well-developed; esophagus with well-developed posterior bulb eggs thin-shelled and usually fully embryonated when laid; life cycle usually direct, ex. Enterobiu vermicularis (human pinworm). adults in colon, gravid females crawl out and lay eggs or burst along perianal folds which cause the characteristic itching (pruritus ani), thin-shelled D-shaped eggs quickly embryonate, spread around by incessant scratching, tossing in bed, licking of infected fingers, inhalation of eggs, theses are swallowed and hatch in the small intestine and mature in the colon; retroinfection also occurs: the eggs hatch and larve crawl back through the anus up to the colon Family Anisakidae. Order Camallanata. lips absent; buccal capsule present, or absent, or replaced by large bilateral sclerotized valves; esophagus long and distinctly divided into anterior muscular and posterior glandular portion; ovoviviparous; anus and vulva may be atrophied in females, ex. Dracunculus medinensis (guinea worm). adults in subcutaneous tissue, gravid females burst to release L1 which cause violent local immune reaction and a blister results, this eventually ruptures and releases L1 out of the host; L1 reaches freshwater and is ingested by copepods Cyclops and Diaptomus where they develop into L2 and L3; infected copepods are ingested by man while drinking and L3 breaksout, migrates through duodenum, mesenteries, abdominal muscles, and reaches subcutaneous tissue where they molt to L4 and finally the adult Order Spirurida. mouth simple, lips absent, buccal capsule absent in most species, esophagus divided into anterior muscular and posterior glandular portions; oviparous or voviviparous, ex. Wuchereria bancrofti (Bancrofts filaria). adults in the major lymph vessels and nodes; females release the larvae called microfilariae; these L1 are swept into the blood stream mainly through the thoracic duct; these get ingested by a suitable mosquito host during a blood meal, inside of which develops into a sausage stage L2 and a filariform L3; when another blood meal is in progress, they escape through the proboscis and enters the host through the puncture wound made by the mosquito, and migrate to mature into adults in the major lymphatics; blocking of the lymph vessels result in elephantiasis usually of the extremities, breasts, and male genitalia, IH in the Philippines are Aedes finlaya poecilus (urban), Anopheles minimus flavirostris (rural), Culex fatigans, and Mansonia uniformis Lecture Notes in Zoology 111 Kingdom Animalia: Phylum Annelida INTRODUCTION:

The annelida, just like the platyhelminthes and nematodes are commonly referred to as worms, although biologically speaking of a different kind. With the inclusion of some minor phyla into this taxon, the external morphology of the organisms comprising this group is more diverse. Annelids or segmented worms are eucoleomates. Metamerism or segmentation evolved in this in this phylum probably as an adaptation for peristaltic burrowing in soft substrate. The following is a general account of the phylum. .

HISTORY: The annelids were included in the vermes with other known worms. Later, they were finally separated from the unsegmented worms by Cuvier in 1799 and designated the Annelids by Lamarck. DEFINITIONS: The annelids are triploblastic, with a bilateral symmetry with parts or the whole length of the body segmented. GENERAL MORPHOLGY AND PHYSIOLOGY:$$ I. Form: A. shape elongate B. symmetry - bilateral C. external features: Metamerism is limited to the trunk region only; the head or acron or prostomium is not considered a segment, so is the pygidium or terminal segment, where new segments are formed. Each segment, with its coelom separated by a transverse septum, is called a metamere that possesses its own nervous, circulatory and excretory components.

II. Integumentary System: Non-chitinous cuticle with chitinous, paired lateral bristles or- setae that increase traction while moving through substratum. III. Muscular System: With an outer circular and inner longitudinal muscles Contraction and relaxation of these muscle layers changes the shape of each segment and thereby creates the characteristic peristaltic wave IV. Coelom:

The schiziocoelom is separated per segment by transverse septa. Each septum is composed of two layers of peritonium; one from the anterior segment and the other from the posterior segment.

V. Digestive System: complete, straight tube mouth anus digestion is extracellular VI. Respiratory System: respiration is through the cuticle VII. Excretory System: nephridial tubules one pair/segment

VIII. Circulatory System: closed type with blood flowing in well developed blood vessels IX. Nervous System: anterior dorsal brain which consists of two pairs of ganglia: the suprapharyngeal ganglion and the subpharyngeal ganglion, both of which are united by the circumpharyngeal commisure a long single or double ventral nerve cord with ganglionic swellings continue posteriorly, with lateral nerves emerging at strategic points to coincide with the segmentations. X. Reproductive System: both monoecious and dioecious forms exist

TAXONOMY AND CLASSIFICATION OF THE ANNELIDA: The new classification of the annelids have included two minor phyla namely the Pogonophora and Echiura. The oligochaetes and hirudinids were merged into a single class and the polychaetes in a separate taxon group. This new grouping is based on the presence of true segmentations and a common larval form. The special morphology and physiology will be discussed as each group is presented. A. Class Polychaeta: Marine annelids with many setae situated on fleshy lateral outgrowths called parapodium; conspicuous segmentation, intersegmental septa present, large coelom, usually a well developed head, bearing appendages; sexes are separate; and a free swimming trochophore larvae. 1. 2. 3. Habitat most are marine, very few are brackish, freshwater or terrestrial (burrowing) Form size ranges from less than 1 mm to about 3 mm External features with dorsal preoral prostomium which bears numerous sensory structure such as eyes, antennae and ventrolateral palps with ventral postoral peristomium mouth is located between the prostomium (which forms the anterior margin) and the peristomium (which forms its lateral and ventral margin) this entire structure is referred to as the head body segments possess paired lateral appendages extending from the body segments called to parapodia which bear setae setae maybe (a) defensive hollow and contain poisons; (b) for locomotion the parapodia may function in locomotion. Respiration, sensory perception, excretion and reproduction

4.

Integumentary System: - a thin layer of cuticle overlying a single layer of columnar epithelium which is connected to the muscle layers by connective tissue

5. Muscular System: outer circular muscle layer inner longitudinal layer which is thicker and bundles and two divided into two dorsolateral bundles and two ventrolateral bundles oblique muscles are also present especially at level of the parapodia 6. Coelom: lined by peritoneum and divided by septa 7. Digestive System: mouth proboscis/'pharynx/bucca1 cavity short esophagus stomach intestine rectum the modes of feeding and respective food acquiring structures differ due to the different modes of existence; a) raptorial (jaws) b) herbivore / browsers/ omnivores/ scavengers c) non-selective deposit feeders (proboscis) d) selective deposit feeders (tentacles) e) filter feeders (radioles)

8. Respiration mostly diffusion through the general body surface may also be aided by gills which may or may not be associated with the parapodia. If associated with the parapodia, these are usually borne on the dorsal cirri or brachial lobe of the notopodium in filter feeders, the radioles also serve as organs for gaseous exchange 9. Circulation: (variable) forms without a circulatory system rely on the coelomic fluid some possess partially open type with large dorsal sinus some possess a closed system: with vessels and blood. . . a) dorsal blood vessel which flows anteriorly network of vessels which connect the dorsal blood vessel with the ventral blood vessel ventral blood vessel which flows posteriorly b) in each segment: the ventral blood vessel divides into the ventral intestinal and ventral parapodial vessels. These continue as the dorsal intestinal and the dorsal parapodial vessels respectively and drain into the dorsal blood vessel c) contractile vessels or pseudoheart are present d) blood contains amoebocyte or respiratory pigments which may be: i. hemoglobin ---- red ii. hemerythrin ---- red iii. chlorocruorin ---- green 10. Nervous System: the brain: a bilobed ganglion in the prostomium ventral nerve cord = connected to the brain via the circumpharyngeal or circumesophageal commisure which is ganglionated; and bears segmental ganglia which gives rise to lateral nerves to each segment polychaete sense organs include the following: a) eyes - located on the prostomium and ranges from 2 to 4 pairs b) nuchal organs - a pair of ciliated sensory pits /slits that are often reversib1e and located in the head region - composed of ciliated columnar sensory cells, sometimes with gland cells - functions as chemoreceptors which are important in the detection of food c) statocvsts associated with the circumpharyngeal connectives, functions for orientation d) ciliated sense organs on the trunk segments and are composed of tubercles, ridges, or bends dorsal organs e) dispersed receptors: - on parapodial and head appendage 11. Excretion and 0smoregulation: either of the protonepriridial or metanephridial type (a) protonephridial type solenocytes (b) metanephridial type nephrostome which is a funnel shaped heavily ciliated structure either one pair / segment or one pair for the whole animal the nephridiopore always open to the segment immediately posterior to the segment where it is situated; and opens to the exterior in the region of the neuropodium accessory excretory structures: (a) chloragogen tissues adjacent to the intestinal and vascular walls and contains greenish (coproporphyrin) and brownish (phaeopnorbiae) pigments which may actually be excretory products (b) coelomocytes (c) intestinal wall

12. Reproductive System: (a) asexual - rarely occurs - maybe by budding, fragmentation and regeneration - few are monoecious; the anterior abdominal segments produce the eggs while the posterior segments produce the sperms - majority are dioecious - the gonads are not distinct gonads but are simply masses of developing germs cells or gametes and are restricted to certain segments only - the gametes are shed into the coelom where they remain until maturity - as soon as the coelom becomes filled, the gametes may exit via any of the following structures/ methods: i. coleomoduct / gonoducts

(b) sexual

ii. iii. iv. v. vi.

nephridia coelomoducts and nephridia anal rupture rupture copulation

Some special reproductive phenomena in polychaetes: 1. 2. epitoky formation of a reproductive individual called epitoke where the worm is normally divided into the atokus portion and the highly modified epitokus portion Swarming simultaneous swimming of epitolus individuals to the surface to shed sperms and eggs; the females release a pheromone that stimulates the sperm release, sperms in the water in turn stimulate the release of the eggs

13. Embryogeney:

Telolecithal spiral cleavage gastrulation by invagination, epiboly, or both a trochophore larvae results; free-swimming, top-shaped individual, with several areas of cilia; i. prototroch girdle of ciliated cells ii. telotroch cilia just above the anus iii. metatroch cilia just below the mouth; between the prototroch and telotroch

14. Classification of the Polychaeta (from Barnes) The following account of the polychaetes will be limited to the more common and representative forms. A special group, the pogonophorans belonging to the Family Siboglinidae, will be given highlight after the taxonomy of the polychaetes. Subclass Errantia segments are numerous and similar - well-developed parapodia with acicula and setae - head with sensory structures - pharynx with jaws or teeth - swimming, crawling, burrowing and tube-dwelling forms

Family Aphroditidae Sea mice. Long setae forming a felt-like covering of the dorsal surface e.g. Aphrodita Family Polynoidea and Sigalionidae scale worms. The dorsal surface bears scales called elytra e.g Lepidonotus, Polynoe, Sigalion Family Amphinomidae Fire worms, crawling polychaetes with brittle poisonous setae, e.g. Amphinome, Hermodice Family Nereidae clam worms, four eyes and four pairs of peristomial cirri, pharynx contains a pair of jaws, large crawling species, e.g. Nereis Family Eunicidae Palolo worms. Tubiculous species containing eyes and setae, e.g. Eunice, Palola Family Myzostomidae greatly flattened commensals and parasites of echinoderms, particularly crinoids e.g. Myzostome Subclass Sedentaria - body commonly displays regional differentiation - parapodia reduced without acicula or compound setae - prostomium without sensory appendages but the head is commonly provided with palps, tentacles and other structures for feeding - no teeth or jaws present

Family Arenicolidae Lugworms, sedentary burrowers without head appendages e.g. Arenicola Family Terebellidae prostomium bears long, non-retractile tentacles, tuberculous or sedentary burrowers, e.g. Terebella FamilySabellidae fanworms or feather dusters worms, lives in non-calcareous tubes e.g. Sabella Family Serpulidae - fanworms or feather dusters worms, forms calcareous tubes, may burrow inside scleractine skeletons e.g. Spirobranchus (Christamas tree worm) The Pogonophorans ( Family Siboglinindae) History Originally a separate phylum, it was first included in the Phylum Annelida despite the unusual morphology it exhibits. Morphologically, it shares the same traits of the segmented worms on the following criteria: body wall musculature, excretory system, nervous system, circulatory system and pigment, presence of metamerism and setae, coelom and embryological developmemt stages. This is further strengthened by similarities in the base sequence of nuclear and mitochondrial DNA. Recent studies on DNA parsimony and comparison of DNA elongation factor shows that the pogonophorans are derived polychaetes. This means that the pogonophorans evolutionary ancestor was a polychaete. General morphology and Physiology 1. Form 2. 3. 4. External morphology: (a) cephalic lobe with ciliated tentacles and a glandular region (b) trunk with paired and unpaired papillae with tow regions of cilia and rings of setae (c) opisthosoma composed of 6 to 25 segments that bears setae A non-calcareous tube houses the organism that is embedded in the substratum.

Muscular outer circular and inner longitudinal muscle layer Coelom schizocoelom Digestive System possess no digestive system; the following are some proposal how the organism is able to acquire and/or generate its food (a) (b) (c) The tentacle capture, digest and absorb food and nutrition Absorbs dissolve organic matter (DOM) from sea water and soft muddy substratum (for the members of subfamily Perviata) Chemotrophic process assisted by bacteria in the trophosome; the following are facts that supports this theory the trophosomes has a density of 106 bacteria per gram

5. 6. 7. 8. 9.

the organism is found near volcanic vents hypothesized to be the source of hydrogen sulfide and methane, or near decomposing bodies of whales the needed material are collected by the tentacles and brought to the trophosome proposed reaction : CO2 + 4H2S + O2 [CH2O]n + 4S + 3H2O most likely applicable for the members of Subfamily Vestiminifera

Respiratory System via the cephalic lobe Excretory System One pair of excretory organ of undetermined type Circulatory System closed type with hemoglobin as the oxygen binding pigment Nervous System possess a brain with ventral nerve cord Reproductive System The male and female are present as separate individuals The gametes are produced by distinct gonads Development stages: fertilized eggs spiral cleavage trochophore larvae

10.

Taxonomy

Subfamily Perviata ( Frenulata) small, thin pogonophorans that lacks a vestimentum and obstruaculum anteriorly, e.g Siboglinum, Oligobranchia, Polybranchia Subfamily Obsturata ( Vestimentifera) Large, thick-bodied pogonophorans up to 2 meters long bearing both vestimentum and obstruaculum anteriorly e.g. Riftia, Ridgeia, Lamellibranchia, Tevnia

CLASS Clitellata all members possess a clitellum; a modified segment with a large concentration of albumin secreting gland cells (usually located in the anterior half of the organism and rarely more than 10 segments Subclass Oligochaeta The members are mostly terrestrial and freshwater annelids with no parapodia and possess few setae; conspicuous segmentation; presence of intersegmental septa; a large coelom; no district head with appendages; hermaphroditic 1. External features Metamerism well-developed Absence of a parapodia Prostomium without appendages Setae present but fewer than those of polychaetes Integumentary System Thin cuticle Epidermis with mucus secreting gland cells Usually with 4 setal sacs per segment located ventrolateral Muscular System Presence of a well-developed outer circular layer and an inner longitudinal layer Coelom In each segment, it is connected to the outside by a sphinctered midorsal pore located in the intersegmental furrows. These exudes coelomic fluid which keeps the integument moist Digestive System Most are scavengers and feed on dead organic matter Mouth buccal cavity muscular pharynx which secretes saliva of mucus and enzymes, and can be everted to serve as an adhesive pad or as a modified pump esophagus which may be modified into a crop (for storage) and gizzard (for grinding food), equipped with calciferous glands for maintaining Ca +2 and CO-3 levels intestines which can secrete cellulase and chitinase, and in terrestrial forms a typhosole or dorsal invagination to increase the digestive surface area anus Respiratory System Diffusion of gasses occur through the general body surface Gills are present in aquatic forms but these are very rare Circulatory System Basically similar to polychaetes with lateral vessels connecting the dorsal and ventral blood vessels Pseudohearts with valves are present Excretory and Osmoregulation Always metanephridia: funnel shaped nephrostome tubules bladder nephridiopore Although most oligochaetes possess typical metanephridia called holonephridia, some may have multiple nephridia called meronephridia as a result of the division of the embryonic nephrostome, others may have instead branched nephridia Metanephridia may open to the outside or may open into the various parts of the GIT, an enteronephric trait Chloragogen tissues surround the intestines and dorsal blood vessels. These are yellowish cells chiefly the center for glycogen and fat synthesis and storage, deamination of protein, ammonia formation and urea synthesis

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Nervous System and sense organs Brain is located in the 3rd segment (more posterior than in the polychaetes) Two ventral nerve cords fused and gives rise to form 3 to 4 pairs of lateral nerve cords per segment Sense organs include the following (a) tactile free nerve endings (b) chemoreceptive tubercles (c) photoreceptive dermal light sense photoreceptors scattered all over the terminal segments and dorsal portion (d) eyes are absent except in a few aquatic forms with pigment-cup eyes Reproduction (a) Asexual transverse fission into 2 or more parts fragmentation and regeneration (b) Sexual - all are hermaphroditic with distinct gonads in limited number of reproductive segments usually with one or two male segments followed by 1 female segment at the anterior half of the body in these segments, the paired testes and ovaries project into the coelom where immature gametes are released in the seminal receptacle and ovisacs respectively which are also paired the gonoducts (vas efferentia and oviducts) may fused and exit as one duct ventrally pseudopenis and/or prostrate glands and seminal receptacles clitellum secretes mucus for copulation and cocoon formation, it also secretes albumin for the eggs copulation is the rule in oligochaetes with mutual sperm transfer by inserting the male genitalia into the spermatheca self-fertilization and parthenogenesis is also known top occur

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Classification of the Oligochaetes (from Barnes) The following is an abbreviated account of the Oligochaetes: Order Lumbricuda 4 pairs of setae per segment, male gonopores open in the same segment contaning the testes. Freshwater forms e.g. Lumbricus Order Tubificida septal bundles usually with 2 or more setae, rarely absent, with one male segment followed by on female segment, male gonopores in segments immediately in front of or behind the testicular segment. Marine and freshwater, with some species widely distributed in poorly oxygenated and polluted waters e.g. Tubifex sp. Order Haplotaxida basically two male segments followed by two female segments, male gonopores one or two segments behind the testicular segments e.g. Lumbricus terestris, Pheretima benguetensis, Megascolides

Subclass Hirudinea The members are called the leeches. These annelids are usually dorso-ventrally flattened, with a prostomium and 32 body segments; two suckers one surrounding the mouth and the other at the posterior end, no setae nor parapodia; hermaphroditic, coelom is mall due to the growth of mesenchymal cells. 1. 2. Habitat predominant freshwater forms, with primaey invasion into land and secondary invasion into the sea External features: Dorsoventrally flattened With smaller anterior sucker that surrounds the mouth and a larger disc shaped posterior sucker Metamerism has been reduced to 34 permanent segments but obscured by secondary external annulations No setae and parapodia Clitellum is composed of segments IX, X and XI The body is divided into five regions (a) Head or cephalic region prostomium plus segments I to VI, with dorsal eyes and anterior sucker (b) Preclitellar region segments VI to IX (c) Clitellar region segments IX to XI with midventral male and female gonopore (d) Trunk region segments XII to XXIV (e) Terminal or posterior region segments XXV to XXXIV, fused together to form a large ventral posterior sucker

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Integumentary System cuticle with a single layer of epidermis, connective dermis with pigment cells large unicellular gland cells Muscular System outer circular, middle oblique and inner longitudinal muscle cords which are attached end to end at the suckers; dorsoventral muscle strands are also present Coelom no distinct coelom, septa is lost, the cavity is invaded by connective tissue, also chloragogen and botryoidal tisues thereby leaving coelomic sinuses Circulatory System sinuses or blood vascular systems; contractile lateral longitudinal channels with extracellular hemoglobin Respiratory System gills for aquatic forms but otherwise thru general body surface with the help of undulations Digestive System Mouth may lead to either (a) proboscis/sucking pharynx highly muscular with a triangular lumen lined with cuticle and quipped with salivary glands that secrete an anticoagulat (hirudin) or (b) jaw/no proboscis/pumping pharynx composed of 3 large oval blade like jaws, one dorsal and 2 ventrolateral Either will lead to the esophagus stomach or crop which maybe a straight tube or possess from one to eleven diverticula intestines which maybe simple or with pairs of lateral caeca rectum dorsal anus just in front of the posterior sucker Leeches are predatory, but 75% are blood sucking extoparasites which only have exopeptidase and bacterial flora to help breakdown high molecular eight proteins, lipids and carbohydrates

9. Excretion and Osmoregulation Leeches possess from 10 to 17 pairs of metanephridia Non-ciliated nephrostome and tubule except for the posterior section Ventrolateral nephridiopore A bladder maybe present (as in oligochaetes) Multiple funnels maybe present 10. Nervous system and sense organs

The brain is in the 10th anf 11th segments Paired ventral cords with fused segmental ganglia Sense organs includes (a) From 2 to 10 eyes (b) Sensory papillae with bristle (c) Dispersed sensory cells, photoreceptors, free nerve endings

11. Reproduction Asexual reproduction and regeneration absent All are hermaphroditic With distinct gonads (a) Male from 4 to many paired testes vas efferens vas deferens epididymis ejaculatory duct single median male gonopore either spermatophore or penis with prostrate glands which exist in segment X (b) Female only 1 pair of ovaries (between the anterior testes and atrium) inside the ovisacs paired oviducts fuses into a vagina female gonopore in segment XI Copulation follows either by: (a) Insertion of the penis (b) Hypodermic impregnation of spermatophore After which the worms separate clitellum becomes conspicuous secretes cocoon cocoon filled with albumin fertilized eggs are released into cocoon as it slips over the female gonopore direct development of eggs into adults 12. Classification Order Acanthaobdellida primitive, with setae and compartmentalized coelom in the first five segments Order Rhynchobdellida strictly aquatic leeches with a proboscis, the circulatory system is separate from the coelomic sinuses. Includes the fish leeches and ectoparasites of many invertebrates and vertebrates Order Gnathobdellida aquaic or terrestrial leeches having a nonreversible pharynx and three pairs of jaws. Fuve annula per segment Family Hirudinidae chiefly amphibious or aquatic blood sucking leeches e.g. Hirudo medicinalis (medicinal leech), Limnotes (carabao leech) Family Haemadipsidae terrestrial tropical leeches of the Australasian region attacking chiefly warm-blooded vertebrates e.g. Haemadipsa Order Pharyngobdellida all with non-protrusible pharynx; teeth are lacking, although one or two stylets maybe present. Primarily aquatic with some semiterrestrial forms Class Echiura Taxonomy: Order Echiura contains nearly 95% of all echiurans Family Bonellidae the members occur over a remarkable depth range from the intertidal zone to the abyss. All species show pronounced sexual dimorphism, with females typically being at least 20 times larger than males. The males live symbiotically on or inside the female e.g. Beonellia Family Echiuridae the largest of the echiurid family, holding nearly 60% of all species, mostly species in shallow waters and typically show little or no sexual dimorphis e.g. Echiuris Order Xenopneusta - atypical echiurans having an unusually short proboscis, an open circulatory system & an enlarged cloacal region that serves in gas exchange e.g. Urechis Order Heteromyota contains a single familywith only one species found only in Japan. The proboscis can be over 1 meter long, more than 3 time longer than the rest of the animals. Unlike other echiurans, the members of this species have hundreds of unpaired nephridia in the coelomic cavity Currently with 140 members described Present in sandy or muddy burrows, rock crevices of shallow waters Their size varies from a few millimeters (male) up to 8 cm (usually females) External morphology: proboscis, trunk and a ring of setae near the anus The proboscis can be extended and the ventral side is ciliated, used to draw the food into the mouth Possess a single coelom with one to a hundred metanephridia Possess anal sacs that collects coelomic fluid discharged to anus The male and female are separate individuals They possess no gonads; the gametes are produced from the peritoneal lining and released via the nephridiopore Fertilization is external with eggs developing into a trochophore larvae Males are smaller, with no proboscis, no circulatory system and with a degenerated GIT The male dwells in the body of the female specifically the nephridia A female may harbor an average of 20 male individuals inside her Among the larvae, maleness requires contact with the proboscis of an adult female; otherwise the larvae becomes a female