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Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Classification of Animal Kingdom Professor Paras Nath College of Agriculture, Fisheries & Forestry Fiji National University, Koronivia It is believed that at some point in time two distinct lines evolved from the very earliest cell forms. They were a group without a nuclear envelope enclosing the nuclear material, the prokaryotes, and a group with a nuclear envelope enclosing a true nucleus, the eukaryotes. All animal phyla are multicellular, eukaryotic, heterotrophic organisms. Being extremely vast and diverse, this kingdom is classified below using flow chart: Kingdom: Animalia Subkingdom: Metazoa Sub-kingdom: Protozoa Branch: Eumetazoa Phylum - Protozoa Branch: Parazoa Phylum - Porifera Grade: Radiata Grade: Bilateria Phyla Divisions Coelenterata Ctenophora Deuterostomia Protostomia Subdivision Subdivisions Enterocelous Coelomata Phyla Lophophorate Coelomata Phyla Chaetognatha Phoronida Echinodermata Bryozoa Pogonophora (Ectoprocta) Hemichordata Brachiopoda Chordata Schizocoelous Coelomata Phyla Priapulida Sipunculida Mollusca Echiurida Annelida Tardigrada Onychophora Arthropoda Pentastomida Fig. 2. Major groups in animal Pseudocoelomata Phyla Acoelomata Phyla Acanthocephala Platyhelminthes Entoprocta Mesozoa Supper– phylum Rhynchocoela Aschelminthes (Nemeertines) Phyla Rotifera Gastrotricha Kinorhyncha Nematoda Nematomorpha Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology There is variation in the classification of animals proposed by various authors. The classification proposed by Hymen (1940) with little modification is briefly described hereunder: Kingdom: Animalia This is the largest group of the animal classification which includes entire fauna (animal population) of the world. It is divided into two sub-kingdoms. (1) Protozoa and (2) Metazoa The contrasting features of the subkingdoms are given hereunder: 1. Kingdom: Animalia Subkingdom: Protozoa Subkingdom: Metazoa Microscopic, unicellular animals. Usually large multi-cellular animals. 2. Structure simple organelles. with sub-cellular Structure complex with strongly marked cellular differentiation. 3. Grade of organization protoplasmic. Grade of organization cellular, cell-tissue, tissue=organ-system. 4. Little physiological division of labour. Physiological marked. 5. Life cycle including more than one Life cycle comprising more than one generation is universal. generation is rare. 6. Asexual reproduction universal and Asexual reproduction occurs only in lower sexual reproduction rare. metazoan. All reproduce sexually. 7. Conjugation occurs between adults Conjugation occurs between uni-nucleate (hologamy). sperm and ovum (syngamy). 8. When a cell divides, the daughter cells The fertilized egg repeatedly divides and the become separate as independent resulting daughter cells remain cohered and animals. differentiated to form a distinct body. 9. The form of individuals may vary even The form of a body is definite for all the in the same species. members of a species. 10. Natural death does not take place due Natural death takes place; hence termed to lack of a body: hence often termed mortal. immortal. division of labour well Sub-kingdom: I. Protozoa Protozoa are a diverse group of unicellular eukaryotic organisms, many of which are motile.About 50,000 species are reported. Originally, protozoa had been defined as unicellular protists with animal-like behaviour, e.g., movement. Protozoa were regarded as the partner group of protists to protophyta, which have plant-like behaviour, e.g. photosynthesis. Following the Greek root of the name, the singular form is protozoon/proʊtəˈzoʊ.ɒn/(protos=first, zoon = animal). Its use has, however, partially been replaced by the word protozoan, which was originally only used as an adjective. In the same manner the plural form protozoans is sometimes being used instead of protozoa. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology In general, protozoa are referred to as animal-like protists because of movement (motility). However, both protozoa and protists are paraphyletic groups (not including all genetic relatives of the group). For example, Entamoeba is more closely related to humans than to Euglena. "Protozoa" is considered an outdated classification in more formal contexts. However, the term is still used in children's education. While there is no exact definition for the term protozoa, it is often referred to as a unicellular heterotrophicprotist, such as the amoeba and ciliates. The term algae are used for microorganisms that photosynthesize. However, distinction between protozoa and algae is often vague. For example, the alga Dinobryon has chloroplasts for photosynthesis, but it can also feed on organic matter and is motile. Protozoa is sometimes considered a subkingdom. It was traditionally considered a phylum under Animalia referring to unicellular animals, with Metazoa referring to multicellular animals. Characteristics Protozoa commonly range from 10 to 52 micrometers, but can grow as large as 1 mm, and are seen easily by microscope. The largest protozoa known are the deep-sea dwelling xenophyophores, which can grow up to 20 cm in diameter. They were considered formerly to be part of the protista family. Protozoa exist throughout aqueous environments and soil, occupying a range of trophic levels.Protozoa are single-celled animals that feed primarily on bacteria, but also eat other protozoa, soluble organic matter, and sometimes fungi. They are several times larger than bacteria – ranging from 1/5000 to 1/50 of an inch (5 to 500 µm) in diameter. As they eat bacteria, protozoa release excess nitrogen that can then be used by plants and other members of the food web. Motility and digestion Tulodens are 2 of the slow-moving form of protozoa. They move around with whiplike tails called flagella, hair-like structures called cilia, or foot-like structures called pseudopodia. Others do not move at all. Protozoa may absorb food via their cell membranes, some, e.g., amoebas, surround food and engulf it, and yet others have openings or "mouth pores" into which they sweep food, and that engulfing of food is said to be phagocytosis. All protozoa digest their food in stomach-like compartments called vacuoles. Pellicle The pellicle is a thin layer supporting the cell membrane in various protozoa, protecting them and allowing them to retain their shape, especially during locomotion, allowing the organism to be more hydrodynamic. They vary from flexible and elastic to rigid. Although somewhat stiff, the pellicle is also flexible and allows the protist to fit into tighter spaces. In ciliates and Apicomplexa, it is formed from closely packed vesicles called alveoli. In euglenids, it is formed from protein strips arranged spirally along the length of the body. Examples of protists with a pellicle are the euglenoids and the paramecium, a ciliate. The pellicle consists of many bacteria that adhere to the surface by their attachment pili. Thus, attachment pili allow the organisms to remain in the broth, from which they take nutrients, while they congregate near air, where the oxygen concentration is greatest. Ecological role As components of the micro- and meiofauna, protozoa are an important food source for microinvertebrates. Thus, the ecological role of protozoa in the transfer of bacterial and algal production to successive trophic levels is important. As predators, they prey upon Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology unicellular or filamentous algae, bacteria, and microfungi. Protozoa are both herbivores and consumers in the decomposer link of the food chain. They also control bacteria populations and biomass to some extent. Protozoa such as the malaria parasites (Plasmodium spp.), trypanosomes and leishmania, are also important disease causing agents in humans. Life cycle Some protozoa have life stages alternating between proliferative stages (e.g., trophozoites) and dormant cysts. As cysts, protozoa can survive harsh conditions, such as exposure to extreme temperatures or harmful chemicals, or long periods without access to nutrients, water, or oxygen for a period of time. Being a cyst enables parasitic species to survive outside of a host, and allows their transmission from one host to another. When protozoa are in the form of trophozoites (Greek, tropho = to nourish), they actively feed. The conversion of a trophozoite to cyst form is known as encystation, while the process of transforming back into a trophozoite is known as excystation. Protozoa can reproduce by binary fission or multiple fission. Some protozoa reproduce sexually, some asexually, while some use a combination, (e.g., Coccidia). An individual protozoan is hermaphroditic. What do protozoa do? Protozoa play an important role in mineralizing nutrients, making them available for use by plants and other soil organisms. Protozoa (and nematodes) have a lower concentration of nitrogen in their cells than the bacteria they eat. (The ratio of carbon to nitrogen for protozoa is 10:1 or much more and 3:1 to 10:1 for bacteria.) Bacteria eaten by protozoa contain too much nitrogen for the amount of carbon protozoa need. They release the excess nitrogen in the form of ammonium (NH4+). This usually occurs near the root system of a plant. Bacteria and other organisms rapidly take up most of the ammonium, but some is used by the plant. The explanation of mineralization and immobilization is given in Fig.1a.below: Fig.2a. Path ways showing mineralization and immobilization of nutrients in soil. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Another role that protozoa play is in regulating bacteria populations. When they graze on bacteria, protozoa stimulate growth of the bacterial population (and, in turn, decomposition rates and soil aggregation.) Exactly why this happens is under some debate, but grazing can be thought of like pruning a tree – a small amount enhances growth, too much reduces growth or will modify the mix of species in the bacterial community. Protozoa are also an important food source for other soil organisms and help to suppress disease by competing with or feeding on pathogens. Where are protozoa? Protozoa need bacteria to eat and water in which to move, so moisture plays a big role in determining which types of protozoa will be present and active. Like bacteria, protozoa are particularly active in the rhizosphere next to roots. Typical numbers of protozoa in soil vary widely – from a thousand per teaspoon in low fertility soils to a million per teaspoon in some highly fertile soils. Fungal-dominated soils (e.g. forests) tend to have more testate amoebae and ciliates than other types. In bacterial-dominated soils, flagellates and naked amoebae predominate. In general, high claycontent soils contain a higher number of smaller protozoa (flagellates and naked amoebae), while coarser textured soils contain more large flagellates, amoebae of both varieties, and ciliates.Protozoa play an important role in nutrient cycling by feeding intensively on bacteria (Fig.7b.,1c.,1d. and1e.). Flagellates have one or two flagella which they use to propel or pull their way through soil. Ciliates are the largest of the protozoa and the least numerous. They consume up to ten thousand bacteria per day, and release plant available nitrogen. Ciliates use the fine cilia along their bodies like oars to move rapidly through soil. Fig.3b.The speck-like bacteria next to the oval protozoa and large, angular sand particle. Fig.4c.Bacteria ingested by an amoeba. Fig.6d.A flagellum extending from the protozoan on the left and the tiny specks are bacteria. Fig.5e.Ciliates showing the fine cilia along their bodies like oars to move rapidly through soil. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Nematodes and Protozoa Protozoa and bacterial-feeding nematodes compete for their common food resource: bacteria. Some soils have high numbers of either nematodes or protozoa, but not both. The significance of this difference to plants is not known. Both groups consume bacteria and release NH4+. Soil Dwelling Vampires Most protozoa eat bacteria, but one group of amoebae, the vampyrellids, eat fungi. The perfectly round holes drilled through the fungal cell wall, much like the purported puncture marks on the neck of a vampire’s victim, are evidence of the presence of vampyrellid amoebae. The amoebae attach to the surface of fungal hyphae and generate enzymes that eat through the fungal cell wall. The amoeba then sucks dry or engulfs the cytoplasm inside the fungal cell before moving on to its next victim. Vampyrellids attack many fungi including root pathogens, such as Gaeumannomyces graminis, shown in the following Fig.1f. This fungus attacks wheat roots and causes take-all disease. Fig.1f.Vampyrellids attacking fungal root pathogen (Gaeumannomyces graminis). Classification All protozoal species are assigned to the kingdom Protista in the Whittaker classification. The protozoa are then placed into various groups primarily on the basis of how they move. The groups are called phyla (singular, phylum) by some microbiologists and classes by others. Members of the four major groups are illustrated in Fig. 1. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig.1.An array of protozoa showing representatives of the four major groups. Protozoa were previously often grouped in the kingdom of Protista, together with the plant-like algae and fungus-like slime molds. As a result of 21st-century systematics, protozoa, along with ciliates, mastigophorans, and apicomplexans, are arranged as animallike protists. Protozoa are unicellular organisms and are often called the animal-like protists because they subsist entirely on other organisms for food. Most protozoa can move about on their own. Amoebas, trypanosomesandparameciaare all examples of animal-like protists (Figs. 2 to 5). Protozoa are classified into three groups based on their shape: Ciliates are the largest and move by means of hair-like cilia. They eat the other two types of protozoa, as well as bacteria. Amoebae also can be quite large and move by means of a temporary foot or “pseudopod.” Amoebae are further divided into testate amoebae (which make a shell-like covering) and naked amoebae (without a covering). Flagellates are the smallest of the protozoa and use a few whip-like flagella to move. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig. 4. Paramecium aurelia, the best known of all ciliates Fig.5. Structure of a Paramecium aurelia showing various body features. The kingdom Protista or Protoctista now includes about 80,000 species of singlecelled organisms that have the genetic material enclosed in a nucleus and have membrane bound organelles, namely, golgi body, mitochondria, lysosomes, centrosome etc. The kingdom includes both plant Protista (=Protophyta) and animal Protista (=Protozoa). In modern classifications Protozoa is considered as a convenient name for unicellular animals and is not given any rank or status. Modified Sleigh’s Systemof Classification(by A. Pechenik, 2002) Kingdom: Protozoa (14 phyla) 1. Phylum: Ciliophora(ciliates). 2. Phylum:Amoebozoa(Amoebas). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology 3. Phylum:Radiozoa(radiolarians). 4. Phylum:Heliozoa (sun animalcules). Flagellated Protozoa 5. 6. 7. 8. 9. Phylum:Trichozoa (Trichomonas). Phylum:Euglenozoa (Euglena). Phylum:Dinozoa (Dinoflagellates). Phylum:Choanozoa (Choanoflagellates). Phylum:Metamonada (Giardia). Spore-Forming Protozoa 10. Phylum:Apicomplexa (=Sporoza), includes three subphyla. Honorary Protozoa 11. 12. 13. 14. Phylum:Labyrinthomorpha (Slime-nets). Phylum:Opalozoa (Opalina). Phylum:Microsporidea (now transferred to fungi). Phylum:Myxozoa (now shifted to kingdom Animalia) Kingdom:Chromista, Slime-nets (Labyrinthomorpha) and Opalina (Opalinata) have been brought under this new kingdom. Subkingdom: II. Metazoa The subkingdom Metazoa comprises multicellular animals as per the old classification. According to Whittaker Animals are multicellular, eukaryoticorganisms of the kingdomAnimalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their lives. Most animals are motile, meaning they can move spontaneously and independently. All animals must ingest other organisms or their products for sustenance (Heterotrophic). Most known animal phyla appeared in the fossil record as marine species during the Cambrian explosion, about 542 million years ago. Animals are divided into various subgroups, including birds, mammals, amphibians, reptiles, fish and insects.It is divided into two branches. 1. Branch: Parazoa 2. Branch: Eumetazoa The contrasting features of the two branches are given in the following tables: Subkingdom: Metazoa Branch: Parazoa Branch: Eumetazoa 1. Animals sessile. Animals mostly mobile. 2. Tissues absent or poorly defined. Tissues well defined. 3. No organs. Organs well marked. 4 No mouth and digestive tract. Mostly with mouth and digestive tract. 5. Body surface porous. Body surface not porous. 6. One to many internal cavities lined by Body cavities not lined by choanocytes. choanocytes. 7. Physiological division of labour not well Physiological division of labour well marked. marked. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Branch: I. Parazoa The Parazoa are an ancestral subkingdom of animals, literally translated as "beside the animals". Parazoans differ from their choanoflagellate ancestors in that they are not microscopic and have differentiated cells. However, they are an outgroup of the animal phylogenetic tree being that they are multicellular and do not have tissues or organs. The only surviving parazoans are the sponges (Fig.6 and 7), which belong to the phylum Porifera, and the Trichoplax in the phylum Placozoa. Fig.6. Underwater photograph of a marine sponge. Fig.7. Different species of sponges. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Porifera The animals of the phylumPorifera (/pɒˈrɪfərə/; meaning "pore bearer" are sponges (Fig.8). They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl (formerly known as mesenchyme)sandwiched between two thin layers of cells. Sponges have unspecialized cells that can transform into other types and which often migrate between the main cell layers and the mesohyl in the process. Sponges do not have nervous, digestive or circulatory systems. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes. They are mostly irregular with a system of water canals and pores. Internal surface lined with choanocytes. They are sessile, marine, a few in freshwater and are solitary or colonial. They display no body symmetry (are asymmetrical); all other animal groups display some sort of symmetry. There are currently 5000 species of sponges, 150 of which are freshwater. Larvae are planktonic and adults are sessile. Fig. 9.Leucosolenia: A. An individual, and B. Its L. S. Fig.8. Stove-pipe sponge (Aplysina archeri) showing pink variation. Branch: II Eumetazoa Eumetazoa (Greek: εὖ [eu], well + μετά [metá], after + ζῷον [zóon], animal) is a clade comprising all major animal groups except sponges, placozoa, and several other obscure or extinct life forms, such as Dickinsonia. This includes radially, biradially or bilaterally symmetrical animals. The clade is usually held to contain at least Ctenophora, Cnidaria, and Bilateria. Whether mesozoans and placozoans belong is in dispute. Characteristics of eumetazoans include true tissues organized into germ layers, and an embryo that goes through a gastrula stage. A Lancelet (or Amphioxus) specimen of Branchiostoma lanceolatum (Pallas, 1774) collected in coarse sand sediments (600 µm) on the Belgian continental shelf is shown in Fig.10. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig.10.A Lancelet (or Amphioxus) specimen of Branchiostoma lanceolatum (Pallas, 1774) The eumetazoans are a major group of animals in the Five Kingdoms classification comprising all animals except the sponges, placozoans and mesozoans and are divided in the following two grades: 1. Radiata 2. Bilateria Their contrasting features are given in the following table: Branch: Eumetazoa Grade: Radiata Grade: Bilateria 1. Body radially or biradially Body bilaterally symmetrical. Some time radials. symmetrical. Some time bilateral. 2. Bilateral symmetry is a secondary Radial symmetry is a secondary adaptation. adaptation. 3. Organ-systems are incipient, i.e. not Organ-systems well marked. well marked. 4. Mesoderm is not developed. Mesoderm is well developed. 5. Coelomic cavity invariably absent. Either no coelomic cavtty or pseudocoelom or well developed true coelom. 6. Tentacles with nematocysts, comb Tentacles, if present, without nematocysts. No comb rows in some. rows. 7. Principal external opening of Principal external openings of digestive cavity are digestive cavity is mouth. mouth and anus. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Grade: I. Radiata In the early 19th century, Georges Cuvier united ctenophores and cnidarians in the Radiata. Thomas Cavalier-Smith, in 1983, redefined Radiata as a subkingdom consisting of Porifera, Myxozoa, Placozoa, Cnidaria and Ctenophora. Lynn Margulis and K. V. Schwartz later redefined Radiata in their Five Kingdom classification, this time including only Cnidaria and Ctenophora.The animals of this group are diploblastic with radial symmetry (radiate) and bear tentacles (tentaculate). Their digestive cavity opens externally through mouth. Phylum: Coelenterata (With hollow intestine) or Cnidaria (Nrettle- bearing). About 10,000 species. Mouth encircled by tentacles bearing nematocysts. Body cavity as coelenteron. Sessile or free swimming. Solitry or colonial. Marine or freshwater. Example: Aurelia, Hydra, Obelia, Corals, etc. Phylum-Ctenophora (Comb-jellies). About 90 species. Symmetry biradial. Two tentacles and eight longitudinal rows of ciliated combs, free swimming and marine. Examples: Coeloplana, Pleurobrachia, Ctenoplana, etc. Fig.11. Aurelia sp. Fig.12. Coeloplana sp. Grade: II. Bilateria The bilateria/ˌbaɪləˈtɪəriə/are all animals having a bilateral symmetry, i.e. they have a front and a back end, as well as an upside and downside. In contrast, radially symmetrical animals like jellyfish have a topside and downside, but no front and back. The bilateria are a major group of animals, including the majority of phyla but not sponges, cnidarians, placozoans and ctenophores. For the most part, Bilateria embryos display three different germ layers, called the endoderm, mesoderm, and ectoderm. From this they are called triploblastic. Nearly all are bilaterally symmetrical, or approximately so; the most notable exception is the echinoderms, which achieve near-radial symmetry as adults, but are bilaterally symmetrical as larvae. Except for a few phyla (i.e. flatworms and gnathostomulids), the Bilateria have complete digestive tracts with separate mouth and anus. Some Bilateria lack body cavities (acoelomates, i.e. Platyhelminthes, Gastrotricha and Gnathostomulida), while others display primary body cavities (deriving from the blastocoel, as pseudocoel) and/or secondary cavities (that appear de novo, for example the coelom). This grade is subdivided into two divisions: 1. Protostomia 2. Deuterostomia Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Their contrasting characters are listed in the following table. Grade: Bilateria 1. Divison: Protostomia Mouth arises from blastopore or from the anterior margin of blastopore. Division: Deuterostomia Mouth arises anteriorly some distance away from blastopore. 2. Coelom absent or Pseudocoelom (a Coelom developed as enterocoel. persistent blastocoel) or coelom developed as schizocoel. 3. Cleavage spiral and determinate. Cleavage radial and indeterminate. Division I: Protostomia Protostomia (from Greek meaning "mouth first") is a clade of animals. Most animals have bilateral symmetry and three germ layers. The major distinctions between deuterostomes and protostomes are found in embryonic development.The division comprises animals in which the mouth arises from or near blastopore.The protostomes were so named because it used to be thought that in their embryos the dent formed the mouth while the anus was formed later, at the opening made by the other end of the gut. Fig.13.ACaribbean Reef Squid, an example of a protostome. It is subdivided into following four subdivisions: 1. 2. 3. 4. Acoelomata, Paeuudocoelomata, Schizocoelous coelomata Lophophorate coelomata Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Subdivision (i): Acoelomata No body or coelom. Space between body wall and digestive cavity is occupied by mesenchyme.An acœlomate animal is characterized by the absence of coelom and thus the internal organs derived from mesoderm.This is called a blastocoelienne cavity. In these animals, the digestive system is very simple and has only one hole, food intake is mainly through the skin.They also have a bilateral symmetry of the body. These animals are part of triploblastic that is to say that their embryo has three germ layers.The third sheet, the mesoderm, in this case has allowed the appearance of the muscles, which results in an autonomous locomotion and concentration of sensory organs at the front of the body (in other words, cephalization). These groups of animals have many species parasites and are the cause of many diseases, often fatal diseases such as tapeworm, the liver fluke, the schistosomiasis, the schistosomiasis or onchocerciasis (river blindness). Phylum: Platyhleminthes The flatworms, known in scientific literature as Platyhelminthes or Plathelminthes (from the Greek πλατύ, platy, meaning "flat" andἕλμινς (root: ἑλμινθ -), helminth-, meaning worm) are a phylum of relatively simple bilaterian, unsegmented, soft-bodied invertebrateanimals. Unlike other bilaterians, they have no body cavity, and no specialized circulatory and respiratoryorgans, which restrict them to having flattened shapes that allow oxygen and nutrients to pass through their bodies by diffusion (Fig. 14). The digestive cavity has only one opening for both the ingestion (intake of nutrients) and egestion (removal of undigested wastes); as a result, the food cannot be processed continuously. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig.14. a. A free-living planarian (Dugesia). Fig.14. b.A fluke worm (Probolitrema). Platyhelminthes consists of the unsegmented dorsoventrally flattenedflatworms, which includes both free-living and parasitic species. They have bilateral symmetry, and can move by using layers of muscles, or in some species, by gliding along a slime trail using cilia. Flatworms are slightly less developed than segmented worms due to their lack of a circulatory system and complete digestive system. Instead, flatworms absorb nutrients through their skin and excrete wastes using specialized "flame cells." Some flatworms have primitive light-sensing "eyes" that allow them to move either towards or away from light, while other species have different types of sensors on their bodies, including chemical, balance, and water movement receptors. Most species of flatworms reproduce either sexually or asexually.Examples: Fasciola, Planaria, Taenia, etc. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Mesozoa (Middle animals). The Mesozoa are enigmatic, minuscule, worm-like parasites of marine invertebrates. As of 2012 it was still unclear whether they are degenerate platyhelminthes (flatworms) or truly-primitive, basalmetazoans. Generally, these tiny, elusive creatures consist of a somatoderm (outer layer) of ciliated cells surrounding one or more reproductive cells. Decades ago, Mesozoa were classified as a phylum. Molecularphylogeny studies, however, have shown that the mysterious mesozoans are polyphyletic. That is, they consist of at least two unrelated groups.The Mesozoa are a small phylum of small and poorly understood animals. They have very simple bodies, often consisting of less than 50 cells. All known species are internal parasites of marine invertebrates.About 50 species described. No organs. Body with an outer single layer of cells enclosing reproductive cells. Examples: Rhopalura, Dicyema, etc. Fig. 15. a.Rhopalura. Phylum: Rhynchocoela or Nemertinea (Ribbon worms). About 750 species. Body dorsoventrally flattened with both mouth and anus. Mostly marine, few terrestrial and freshwater, Examples: Cerebratulus, Lineus, etc. Subdivision (ii): Pseudocoelomata Pseudocoelomate (s ´´dōsē´l māt´´) are a group of invertebrates with a three-layered body that has a fluid-filled body cavity (pseudocoelom) between the endoderm and the Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology mesoderm (the innermost and middle tissue layers). Body cavity is a pseudocoelom which is a persistent blastocoel, not lined by mesoderm. The pseudocoelom is contrasted with the coelom of mollusks, annelid worms, and the more complex animals (including humans and other vertebrates). Pseudocoelomates lack a circulatory system, and the pseudocoelom itself lacks the endothelial lining of a coelom. The hydrostatic pressure of the pseudocoelom gives the body a supportive framework that acts as a skeleton. Nematodes or roundworms, rotifers, acanthocephalans (spiny-headed worms), kinorhynchs and nematomorphs or horsehair worms are pseudocoelomates. Phylum: Acanthocephala Acanthocephala (Greek ἄκανθος, akanthos, thorn + κεφαλή, kephale, head) is a phylum of minute parasitic worms known as acanthocephalans, thorny-headed worms, or spiny-headed worms, characterized by the presence of a protrusible proboscis with recurved spines, which it uses to pierce and hold the gut wall of its host. Acanthocephalans are highly adapted to a parasitic mode of life, and have lost many organs and structures through evolutionary processes. Acanthocephalans lack a mouth or digestive cavity. This is a feature they share with the cestoda (tapeworms), although the two groups are not closely related. Adult stages live in the intestines of their host and uptake nutrients which have been digested by the host, directly, through their body surface. Acanthocephalans have complex life cycles, involving at least two hosts, which may include invertebrates, fishes, amphibians, birds, and mammals. About 1150 species have been described.Example: Pomphorhynchus,Acanthocephalus, etc. Fig.16. Adult Pomphorhynchus in abluefish (Pomatomus saltatrix). Fig.17. Morphological features of Corynosoma wegeneri (Phylum Acanthocephala). The endoparasitic thorny-headed worms (1150 species) require two hosts to complete the life cycle. The juveniles are parasitic in crustaceans and insects. Adults live in the Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology digestive tract of vertebrates, especially fish. The body of the adult is elongate and composed of a trunk, a short head region and a proboscis covered with recurved spines with which the worms embed themselves in the host’s gut wall. The proboscis is not used for feeding, only for attachment, since like tapeworms; acanthocephalans have no gut and absorb nutrients directly from the host’s gut through the tegument. Eggs are passed into the environment; they develop further when ingested by the intermediate host. The juvenile acanthor bears an anterior crown of hooks used to penetrate the tissues of the intermediate host, where it undergoes developmental changes to form an infective cystacanth. When the intermediate host is eaten by the definite host, the cystacanth excysts and attaches to the gut wall and develops into an adult. Fig.18. Rhadinorhynchussp. Fig.19. Polymorphus spp. Fig. 21. The largest acanthocephalan, Macracanthorhyncus hirudinaceous. Fig.20.Profillicolus bolulus in the Fig.22. Lateral and face view of proboscis of intestine of the eider duck. Hypoechinorhynchus thermaceri Buron. Rhadinorhynchus sp.(Fig.18) and Polymorphus spp. (Fig.19) are showing the spiny proboscis that gives acanthocephalans their name. Profillicolus bolulus(Fig.20) in the intestine of the eider duck shows the ‘orchard effect’: spacing of the parasites so that ‘they do not tread on each other’s toes’. The lateral and face view of the proboscis of Hypoechinorhynchus thermaceriare shown through the electron micrographs in the Fig.22.Acanthocephalans often exist in great numbers in the vertebrate host and can cause much damage to the gut wall.The largest acanthocephalan, Macracanthorhyncus hirudinaceous, causes serious disease in its definitive host, the pig(Fig.21). Beetles act as hosts for the juveniles. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Entoprocta (Moss animals). "Entoprocta", coined in 1870, means "anus inside". The alternative name "Kamptozoa", meaning "bent" or "curved" animals, was assigned in 1929. Some authors use "Entoprocta", while others prefer "Kamptozoa". Most species are colonial, and their members are known as "zooids", since they are not fully independent animals. Zooids are typically 1 millimetre (0.039 in) long but ranging from 0.1 to 7 millimetres (0.0039 to 0.28 in) long. Entoproctaisa phylum of mostly aquatic animals.While the great majority is marine, two species live in freshwater, Loxosomatoides sirindhornae reported in 2004 in central Thailand and Urnatella gracilis found in all the continents except Antarctica. Colonial species are found in all the oceans, living on rocks, shells, algae and underwater buildings. These are solitary or colonial and suspension feeders by means of tentacles, however some species occur commensally on animals that have their own feeding current system (e.g. poriferans).The solitary species, which are marine, live on other animals that feed by producing water currents, such as sponges, ectoprocts and sessile annelids.The digestive tube is U-shaped. Mouth and anus are close together and surrounded by a tentacular crown (Fig.23 and 24).The zooids absorb oxygen and emit carbon dioxide by diffusion, which works well for small animals. Mature individuals are goblet-shaped, on relatively long stalks.Some species eject unfertilized ova into the water, while others keep their ova in brood chambers until they hatch, and some of these species use placenta-like organs to nourish the developing eggs. After hatching, the larvae swim for a short time and then settle on a surface. There they metamorphose, and the larval gut generally rotates by up to 180°, so that the mouth and anus face upwards. Both colonial and solitary species also reproduce by cloning – solitary species grow clones in the space between the tentacles and then release them when developed, while colonial ones produce new members from the stalks or from corridor-like stolons. Most families of entoprocts are colonial, and all but 2 of the 150 reported species are marine. A few solitary species can move slowly.Example: Pedicellina, Loxosomaa, etc. Fig.23. Pedicellina cernua (Pallas, 1774) from Guernsey, a British Crown dependent island in the English Channel. Entire colony × 27. The colony has three growing ends – a; 1–8 – individuals of colony; 1 and 8 are quite immature; 7 is still young, tentacles retracted; 2 is seen in longitudinal section; g – generative organ, and below it the ganglion; m – mouth; r – rectum; s – stomach; between g and r are three embryos in the brood-pouch; the tentacles are Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology retracted; in 5 and 6 the tentacles are expanded; in 6 two embryos are seen within the circle of the tentacles, to the left of them is the rectum, and to the right the mouth; 3 is in the act of losing its calyx, and has already developed the beginning of a new polypide-bud; in 4 the primary calyx has been lost, and the new calyx is clearly marked off from the stalk. Fig.24. Body plan of Entoprocta. Super-phylum: Aschelminthes The members of this Super-phylum are commonly known as sac worms. This group is an assemblage of pseudocoelomates with an anterior mouth, posterior anus and straight digestive tube. The Aschelminthes (also known as Aeschelminthes), closely associated with the Platyhelminthes, are an obsolete phylum of pseudocoelomate and other similar animals that are no longer considered closely related and have been promoted to phyla in their own right. The term Aschelminth is now generally only used as an informal name for any member of the approximately ten different invertebrate phyla formerly included within Aschelminthes. It is considered a polyphyletic group.Although invertebrate experts do not necessarily agree on these categorizations, groups that are generally incorporated into Aschelminthes are briefly described hereunder: Phylum: Rotifera The word "rotifer" is derived from a Latin word meaning "wheel-bearer", due to the corona around the mouth that in concerted sequential motion resembles a wheel (though the organ does not actually rotate).The rotifers (Rotifera, commonly called wheelanimals or wheel animalcules) make up a phylum of microscopic and near-microscopic pseudocoelomateanimals. They were first described by Rev. John Harris in 1696, and other forms were described by Anton van Leeuwenhoek in 1703. Most rotifers are around 0.1– 0.5 mm long (although their size can range from 50 μm to over 2 millimeters), and are common in freshwater environments throughout the world with a few saltwater species; for example, those of genus Synchaeta. Some rotifers are free swimming and truly planktonic, others move by inchworming along a substrate, and some are sessile, living inside tubes or gelatinous holdfasts that are attached to a substrate. About 2200 species of rotifers have been described. Anterior end with a ciliated crown.Pharynx with internal jaws. Examples: Philodina, Rotatoria, etc. About 25 species are colonial (e.g., Sinantherina semibullata), either sessile or planktonic. Rotifers are an important part of the freshwater zooplankton, Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology being a major foodsource and with many species also contributing to the decomposition of soil organic matter. Most species of the rotifers are cosmopolitan, but there are also some endemic species, like Cephalodella vittata to Lake Baikal. Recent barcoding evidence, however, suggests that some 'cosmopolitan' species, such as Brachionus plicatilis, B. calyciflorus, Lecane bulla, among others, are actually species complexes. Fig.25. Colony of Rotifers (Sinantheria socialis, Family: Flosculariidae) on Egeria densa (North German Lake). Fig.26. Scanning electron micrographs showing morphological variation of bdelloid rotifers and their jaws. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Gastrotricha The gastrotrichs (from Greek γαστήρ, gaster ["belly"], and θρίξ, thrix ["hair"]), often called hairy backs or hairy stomach worms, are a phylum of microscopic (0.06-3.0 mm), pseudocoelomateanimals abundant in fresh water and marine environments. Most fresh water species are part of the periphyton and benthos. Marine species are found mostly interstitially in between sediment particles, while terrestrial species live in the water films around grains of soil. The common name "hairy back" apparently arises from a mistranslation of "gastrotrich;" the better common name for all gastrotrichs is "hairy belly," which refers to ventral cilia present in most species. "Hairy back" should be limited to the large Genus Chaetonotus, whose members usually have backs covered with hair-like spines. Gastrotrichs are bilaterally symmetric, with a transparent body and a flat underside. Many species have a pair of short projections at the posterior end (Fig.27, Fig.28 and Fig.29). The body is covered with cilia, especially about the mouth and on the ventral surface, and has two terminal projections with cement glands that serve in adhesion. This is a double-gland system where one gland secretes the glue and another secretes a de-adhesive to sever the connection. Like many microscopic animals, their locomotion is primarily powered by hydrostatics. About 150 species described. Ventral surface is flattened andciliated. Cuticle is with spines, plates or scales. These are found in freshwater and marine water bodies. Examples: Chaetonotus, Macrodasys, etc. Fig.27. Fig.29. Fig.28. Fig.27. & 28.Photographs of two species of gastrotrich.Fig.29. A generalized Gastrotrich body parts. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Kinorhyncha Kinorhyncha (ki-no-RINK-a) is formed from two Greek roots that mean "moving by the snout". Kinorhyncha (Gr. κίνηω, kīneō 'move' + ρυνχος, rhynchos 'snout') is a phylum of small (1 mm or less) marine pseudocoelomateinvertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. The reference is to the way the animal moves by everting its mouth cone.They are called spiny-crown worm, jaw-moving worms,mud dragons.Kinorhynchans are segmented, limbless animals, with a body consisting of a head, neck, and a trunk of eleven segments. Cuticle segmented and with recurved spines. Unlike some similar invertebrates, they do not have external cilia, but instead have a number of spines along the body, plus up to seven circles of spines around the head. These recurved spines are used for locomotion, withdrawing the head and pushing forward, then gripping the substrate with the spines while drawing up the body.Anterior end is spiny. About 180 species are known. Exampels: Echinoders, Pycnophyses, etc. Fig.30. Body plan ofKinorhyncha and some other images of spiny-crown worm. Phylum: Nematoda The nematodes/ˈnɛmətoʊdz/or roundworms comprise the phylumNematoda.This is a phylum of worms, having a long, round, and generally smooth body; the roundworms. They are mostly parasites, in plants and animals, but some are free-living in soil or water. This phylum is also called Nematoidea. They are a diverse animal phylum inhabiting a very broad range of environments. Nematode species can be difficult to distinguish; and although over 28,000 have been described, of which over 16,000 are parasitic, the total number of nematode species has been estimated to be about 1 million. Unlike cnidarians and flatworms, nematodes have tubular digestive systems with openings at both ends. Nematodes are slenderand cylindrical worms havingradial or biradial arrangement of structures around the mouth. They are typically less than 2.5 mm (0.098 in) long. The smallest nematodes are microscopic, while free-living species can reach as much as 5 cm (2.0 in), and some parasitic species are larger still, reaching over a meter in length. The body is often ornamented with ridges, rings, bristles, or other distinctive structures. The head of a nematode is relatively distinct. Whereas the rest of the body is bilaterally symmetrical, the head is radially symmetrical, with sensory bristles and, in many cases, solid 'head-shields' radiating outwards around the mouth. The mouth has either three or Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology six lips, which often bear a series of teeth on their inner edges. An adhesive 'caudal gland' is often found at the tip of the tail. The body features of nematode are shown in Fig.31. Nematodes have successfully adapted to nearly every ecosystem from marine to fresh water, to soils, and from the Polar Regions to the tropics, as well as the highest to the lowest of elevations. They are ubiquitous in freshwater, marine, and terrestrial environments, where they often outnumber other animals in both individual and species counts, and are found in locations as diverse as mountains, deserts and oceanic trenches. They are free-living or parasitic in nature. They are found in every part of the earth's lithosphere. They represent, for example, 90% of all life forms on the ocean floor.Their numerical dominance, often exceeding a million individuals per square meter and accounting for about 80% of all individual animals on earth, their diversity of life cycles, and their presence at various trophic levels point at an important role in many ecosystems. Their many parasitic forms include pathogens in most plants and animals (including humans). Some nematodes can undergo cryptobiosis. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology One group of carnivorous fungi, the nematophagous fungi, is predators of soil nematodes. They set enticements for the nematodes in the form of lassos or adhesive structures.Nematodes have even been found at great depth (0.9–3.6 km) below the surface of the Earth in gold mines in South Africa. The different species of free-living nematodes usually consists of four moults of the cuticle during growth and feed on materials as varied as algae, fungi, small animals, faecal matter, dead organisms and living tissues. Free-living marine nematodes are important and abundant members of the meiobenthos. They play an important role in the decomposition process, aid in recycling of nutrients in marine environments, and are sensitive to changes in the environment caused by pollution. One roundworm of note, Caenorhabditis elegans, lives in the soil and has found much use as a model organism. C. elegans has had its entire genome sequenced, as well as the developmental fate of every cell determined, and every neuron mapped. Nematodes commonly parasitic on humans include ascarids (Ascaris), filarias, hookworms, pinworms (Enterobius) and whipworms (Trichuris trichiura). The species Trichinella spiralis, commonly known as the 'trichina worm', occurs in rats, pigs, and humans, and is responsible for the disease trichinosis. Baylisascaris usually infests wild animals, but can be deadly to humans, as well. Dirofilaria immitisheartworms are known for causing heartworm disease by inhabiting the hearts, arteries, and lungs of dogs and some cats. Haemonchus contortus is one of the most abundant infectious agents in sheep around the world, causing great economic damage to sheep. In contrast, entomopathogenic nematodes parasitize insects and are considered by humans to be beneficial.Some common parasitic nematodes are Ascaris, Trichinella, Wuchereria, Ancylostoma, Enterobius,etc. Depending on the species, a nematode may be beneficial or detrimental to plant health. From agricultural and horticulture perspectives, the two categories of nematodes are the predatory ones, which will kill garden pests like cutworms, and the pest nematodes (Fig.), like the root-knot nematode, which attack plants, and those that act as vectors spreading plant viruses between crop plants. Predatory nematodes can be bred by soaking a specific recipe of leaves and other detritus in water, in a dark, cool place, and can even be purchased as an organic form of pest control Fig.31. Low-temperature scanning electron micrograph ofsoybean cyst nematode and its egg. (Magnified 1,000 times). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Nematomorpha Nematomorpha (sometimes called Gordiacea, and commonly known as horsehair worms or Gordian worms) are a phylum of parasitoidanimals superficially morphologically similar to nematode worms, hence the name. They range in size in most species from 50 to 100 centimetres (20 to 39 in) long,threadlikeand can reach in extreme cases up to 2 metres, and 1 to 3 millimetres (0.039 to 0.12 in) in diameter (Fig.32 and 33). Fig.32. Horsehair Worm (Nematomorpha) and Pterostichus tareumiutbeetleparasitized by horse hairworm (Nematomorpha)parasite. Fig.33. FemaleGordionussp. emerging from anAmara alpine(Carabidae: Coleoptera)and a laboratory over infected variable field cricket,Gryllus lineaticeps withParagordius varius. Horsehair worms can be discovered in damp areas such as watering troughs, swimming pools, streams, puddles, and cisterns. The adult worms are free living, but the larvae are parasitic on beetles, cockroaches, orthopterans, and crustaceans. About 351 freshwater species are known and a conservative estimate suggests that there may be about 2000 freshwater species worldwide. The name "Gordian" stems from the legendary Gordian knot. This relates to the fact that Nematomorpha often tie themselves in knots. Examples:Paragordius, Gordius, Nectonema etc. Nematomorphs possess an external cuticle without cilia. Internally, they have only longitudinal muscle and a non-functional gut, with no excretory, respiratory or circulatory systems. The nervous system consists of a nerve ring near the anterior end of the animal, and a ventral nerve cord running along the body (Fig.34). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig.34. Structure of Paragordius, a nematomorph. A, Longitudinal section through the anterior end. B, Transverse section. C, Posterior end of male and female worms. Nematomorphs, or “horsehair worms,” are very long and very thin. Their pharynx is usually a solid cord of cells and is nonfunctional. Paragordius, whose pharynx opens through to the intestine, is unusual in this respect and also in the possession of a photosensory organ (“eye”). (iii)Subdivision: Schizocoelous Coelomata Coelom is a schizocoel which originates as a space by the splitting of the embryonic mesoderm. Phylum: Priapulida Priapulida (priapulid worms or penis worms, from Gr. πριάπος, priāpos 'Priapus' + Lat. -ul-, diminutive) is a phylum of marine worms (Fig.35). The name of the phylum relates to the Greek god of fertility, because their general shape, and their extensible spiny introvert (eversible proboscis) may recall the shape of a penis. Priapulida are found in colder waters at a variety of depths from tidal to abyssal. They live in the mud, which they eat, in comparatively shallow waters up to 90 metres (300 ft). Some species show a remarkable tolerance for hydrogen sulfide and anoxia. Priapulids are cylindrical worm-like animals, ranging from 0.2 - 0.3 to 39 centimetres (0.08 - 0.12 to 15.35 in) in length, with a median anterior mouth quite devoid of any armature or tentacles. The body is divided into a main trunk or abdomen and a somewhat swollen proboscis region ornamented with longitudinal ridges. The body is ringed and body surface is covered with spines and tubercles, which are continued into the slightly protrusible pharynx. Some species may also have a tail or a pair of caudal appendages. The body has a chitinous cuticle that is moulted as the animal grows.Their body cavity has a mesodermal lining, so they can be regarded as coelomate; however the lining is unlike that of other coelomates. Proboscis anterior and peritoneum of coelom greatly reduced. Circular and longitudinal muscles exert the pressure required to evert the head region (Fig. 37). The larvae move through the sediment using their head region as an anchor, but adults are rarely good Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology burrowers. They are predators feeding on slow-moving animals, especially worms. At present only about 16 species of Priapulida have been described, though fossil evidence dates back to the Cambrian period. Example:Pripulus. Fig.36. Priapulid worm Priapulus caudatus in a Petry dish.Fig.37.Priapulus bicaudatus. Phylum: Sipunculida The Sipuncula or Sipunculida (common names sipunculid worms or peanut worms) is a group containing 144-320 species (estimates vary) of bilaterally symmetrical, unsegmentedmarine worms. Traditionally considered a phylum, molecular work suggests that they might be a subgroup of phylum Annelida. Fig.38. Sipunculid or peanut worm. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Sipunculids are all marine and are relatively common, and live in shallow waters, either in burrows or in discarded shells like hermit crabs do. Some bore into solid rocks to make a shelter for themselves. Although typically less than 10 cm long, some sipunculans may reach several times that length. Sipunculans are worm-like animals ranging from 2 to 720 millimetres in length, with most species being less than 10 centimetres. The sipunculan body is elongated and cylindrical with retractile anterior end. Lobes or tentacles are around the mouthand anus is on dorsal side.The body is divided into an unsegmented trunk and a narrower, retractable anterior section, called the "introvert". Sipunculans have a body wall somewhat similar to that of annelids (though unsegmented) in that it consists of a non-ciliated epidermis overlain by a cuticle, an outer layer of circular and an inner layer of longitudinal musculature. The body wall surrounds the coelom that is filled with fluid on which the body wall musculature acts as a hydrostatic skeleton to extend or contract the animal. When threatened, Sipunculids can retract their body into a shape resembling a peanutkernel - a practice that has given rise to the name "peanut worm". The introvert is retractable into the trunk via two pairs of retractor muscles that extend as narrow ribbons from the trunk wall to attachment points in the introvert. The introvert can be protruded from the trunk by contracting the muscles of the trunk wall, thus forcing the fluid in the body cavity forwards.Examples: Sipunculus, Aspidosiphon. Fig.39. A bucket of deliciously-looking purple worms 'Sand worms' from Beihai. Sipunculids are capable of regenerating lost parts of their tentacles, introvert, trunk, or internal digestive system. Some species are able to "clone" themselves by breaking into a Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology large front portion and a smaller back portion, each capable of regrowing their missing parts, and resulting in two separate organisms. To reproduce, sipunculids release their sex cells into the surrounding water to produce free-swimming juveniles. Fig.10. A dish of Sipunculid worm jelly made with Sipunculus nudus. Phylum: Mollusca Mollusca are one of the most diverse groups of animals on the planet, with at least 50,000 living species (and more likely around 200,000). It includes such familiar organisms as snails, octopuses, squid, clams, scallops, oysters, and chitons. Mollusca also includes some lesser known groups like the monoplacophorans, a group once thought to be extinct for millions of years until one was found in 1952 in the deep ocean off the coast of Costa Rica. Molluscs are a clade of organisms that all have soft bodies which typically have a "head" and a "foot" region. Often their bodies are covered by a hard exoskeleton, as in the shells of snails and clams or the plates of chitons. A part of almost every ecosystem in the world, molluscs are extremely important members of many ecological communities. They range in distribution from terrestrial mountain tops to the hot vents and cold seeps of the deep sea, and range in size from 20meter-long giant squid to microscopic aplacophorans, a millimeter or less in length, that live between sand grains. These creatures have been important to humans throughout history as a source of food, jewelry, tools, and even pets. Besides having yummy soft parts, molluscs often have desirable hard parts. The shells of some molluscs are considered quite beautiful and valuable. Molluscs can also be nuisances, such as the common garden snail; and molluscs make up a major component of fouling communities both on docks and on the hulls of ships. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology They also have a very long and rich fossil record going back more than 550 million years, making them one of the most common types of organism used by paleontologists to study the history of life. Morphology Despite their amazing diversity, all molluscs share some unique characteristics that define their body plan. The body has a head, a foot and a visceral mass. This is all covered with a mantle (also known as a pallium) that typically secretes the shell (Fig.40-44). In some groups, like slugs and octopuses, the mantle is secondarily lost, while in others, it is used for other activities, such as respiration. Fig.40. Snail,Pilasp. Fig.41. Molluscan internal body parts. The buccal cavity, at the anterior of the mollusc, contains a radula (lost in bivalves) — a ribbon of teeth supported by an odontophore, a muscular structure (Fig.42). The radula is generally used for feeding. The ventral foot is used in locomotion. This foot propels the mollusc by utilizing muscular waves and/or cilia in combination with mucus(Fig. 43). Fig.42. The freshwater Sinistral Pond Fig. 43. A cuttlefish, a coleoid Snail (Physella sp.) scrapes algae from cephalopod, moves primarily by the glass with its radula, the two undulating its body fins. "toothy" arcs you can see lining the mouth. Typically, at least in the more primitive members of each group, there are one or more pairs of gills (called ctenidia) which lie in a posterior cavity (the pallial cavity) or in a posterolateral groove surrounding the foot. The pallial cavity typically contains a pair of sensory osphradia (for smelling) and is the space into which the kidneys, gonads, and anus open. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Molluscs are coelomate, although the coelom is reduced and represented by the kidneys, gonads, and pericardium, the main body cavity which surrounds the heart. Examples: Chitons, snails (Pila), mussels (Unio), squids (Loligo), etc. Fig.44. On the left is a marine snail, the California Trivia (Trivia californiana). Here the mantle covers much of the shell. Note how a portion of the mantle is rolled into a tube shape to form the siphon just above the head. At the right is a restoration of one of the largest of all molluscs, the Giant Squid (Architeuthis). Phylum: Echiurida The Echiura, or spoon wormsor adder-tailed worms, are a small group of marineanimals(Fig.45). Once treated as a separate phylum, they are now universally considered to represent derived annelid worms. The Echiura fossilise poorly and the earliest known specimen is from the Upper Carboniferous (called the Pennsylvanian in North America). However, U-shaped fossil burrows that could be Echiuran have been found dating back to the Cambrian. Fig.45. (a)Echiurussp. entire animal (ant.set, anterior setae; post.set, posterior setae;prob, proboscis);(b) Echiurus sp. showing an extensible proboscis and posterior end with a set of small hooks; (c)Echiuran long and coiled intestines. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Echiurans have a worm-like body with a large flattened proboscis projecting forward from the head (Fig.45 and 46). The body is typically drab in colour, but bright red and green species are known. Its body is cylindrical with anterior retractile proboscis. Its trunk is with setae. The proboscis is a sheet-like structure, rolled around into a cylindrical tube with an open gutter at the ventral surface. The length of the proboscis varies greatly between species, and in some species is many times longer than the rest of the body. It is probably homologous with the prostomium of other annelids. Fig.46. Echiura Urechis unicinctuson a market in South Korea. Compared with other annelids, echiurans have relatively few setae. In most species, there are just two, located on the underside of the body just behind the proboscis. In others, such as Echiurus, there are also further setae near the posterior end of the animal. Unlike other annelids, adult echiurans have no trace of segmentation. Echiurans are marine worms similar in size and habit to sipunculans. Many genera, such as Echiurus, Urechis, and Ikeda, live in burrows in sand and mud; others live in rock and coral crevices. One species, Thalassema mellita, which lives off the southeastern coast of the US, inhabits the tests (exoskeleton) of dead sand dollars. When the worm is very small, it enters the test and later becomes too large to leave. The majority of echiurans live in shallow water, but there are also deep sea forms. More than 230 species have been described.[ Phylum: Annelida The annelids (also called "ringed worms"), formally called Annelida (from Latinanellus "little ring", are a large phylum of segmented worms including ragworms, earthworms and leeches (Fig.47). They are found in marine environments from tidal zones to hydrothermal vents, in freshwater, and in moist terrestrial environments. They are bilaterally symmetrical, triploblastic, coelomate organisms. They have parapodia for locomotion. There are over 22,000 living annelid species, ranging in size from microscopic to the Australian giant Gippsland earthworm and Amynthas mekongianus (Cognetti), which can both grow up to 3 metres long. The basic annelid form consists of multiple segments, each of which has the same sets of organs and, in most polychaetes, a pair of parapodia that many species use for locomotion. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Septa separate the segments of many species, but are poorly-defined or absent in some, and Echiura and Sipuncula show no obvious signs of segmentation. In species with welldeveloped septa, the blood circulates entirely within blood vessels, and the vessels in segments near the front ends of these species are often built up with muscles to act as hearts. The septa of these species also enable them to change the shapes of individual segments, which facilitates movement by peristalsis ("ripples" that pass along the body) or by undulations that improve the effectiveness of the parapodia. In species with incomplete septa or none, the blood circulates through the main body cavity without any kind of pump, and there is a wide range of locomotory techniques - some burrowing species turn their pharynges inside out to drag themselves through the sediment. Fig.48. Fig.47. Fig.11. Bloodworm, Glycera sp., abounds in salt marsh sediments. Fig.12. Bamboo worm(Capitella capitata F.). Fig.50 Fig.49. Calcareous tubeworm, fan worm, plume worm or red tube worm(Serpula vermicularis L.) Fig.51.Nereis pelagica.Fig.51. Rag worm, clamworm or Nereid worm-Live Nereissp. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig. 52. Fig.52.Hediste diversicolor. Fig.53. Lamellibrachia luymesi Fig.54. Amynthas sp., a common Asian earthworm often cosmopolitan and introduced around the world Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig. 55.Earthworms mating. Fig. 56. Cocoons of earthworms, L. rubellus. Fig. 57. Earthworms’faeces in form of casts. Fig. 58. Earthworm (L. terrestris) - permanent vertical burrow. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Although many species can reproduce asexually and use similar mechanisms to regenerate after severe injuries, sexual reproduction is the normal method in species whose reproduction has been studied. The minority of living polychaetes whose reproduction and lifecycles are known produce trochophore larvae, which live as plankton and then sink and metamorphose into miniature adults. Oligochaetes are full hermaphrodites and produce a ring-like cocoon round their bodies, in which the eggs and hatchlings are nourished until they are ready to emerge. Earthworms support terrestrial food chains both as prey and by aerating and enriching soil (Fig.59). The burrowing of marine polychaetes, which may constitute up to a third of all species in near-shore environments, encourages the development of ecosystems by enabling water and oxygen to penetrate the sea floor. Fig. 59. Staphylinus olens fighting an earthworm (Lumbricus sp.) near Nettersheim, Germany. Fig. 60.A close up of an earthworm in garden soil. In addition to improving soil fertility, annelids serve humans as food and as bait. Scientists observe annelids to monitor the quality of marine and fresh water. Although bloodletting is no longer in favor with doctors, some leech species are regarded as endangered species because they have been over-harvested for this purpose in the last few centuries. Leeches have been used to treat patients for centuries. Leech therapy, which is called in Ayurveda system as Jalauka or Rakta Moksha, is an old form of Ayurvedic detoxification with the help of the creeping insect leech. Ayurvedic scriptures such as Charak Samhita and the Sushrut Samhita give many details of this therapy. The impure blood which is the root cause of most of the human ailments is removed from the body. Leeches have the ability to suck blood clotted around muscles or skin (Fig.61). The anti-blood clotting enzymes in their saliva make the blood circulation normal. In this way, leech therapy does wonders in many diseases connected with blood clotting.There are approximately 600 leech species which have been identified to date but only about 15 are used in medicine.Leech therapy has been used Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology in European countries since 18th and 19th centuries. Today, doctors use leeches for treating abscesses, painful joints, glaucoma, myasthenia, and to heal venous diseases and thrombosis. Medical leeches are used in plastic surgery, for improving brain circulation and for curing infertility. Some even claim that this could be an alternative acne treatment (Fig.65). In Fig.62 the leeches hanging from the chin of a Kashmiri womanare shown, who is undergoing a leech therapy session at a roadside in Srinagar, Kashmir, India during summer. Leeches, which are widely used to promote bloodletting, are being used to heal pain and other ailments in Kashmir.Even with osteoarthritis of the knee joint, a leech therapy may be effective (Fig.63.). Painful, locally jammed and metabolically abnormal processes can be treated this way. Most common use of leech therapy is in local inflammation, arthritis and chronic pain syndromes (Fig. 64 and 65). Potentially, this could be a business opportunity for those who are interested in alternative medicine (Fig.66). Fig.61. Leech Therapy: An alternative therapy for cancer. Fig.63. Leech therapy for osteoarthritis. Fig.62. Leech therapy session in Kashmir, India. Fig.64. Leech therapy session in Kashmir, India. In tropical regions, leech bites on the skin are commonly encountered, especially, when people walk carelessly through bushes and water (Fig.67). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig. 65. Russian woman takes treatment in a laboratory in Moscow. leech Fig.66.Booming leech trade raises concerns. Vaginal leech bites, in children are also commonly heard in rural part of north-eastern India. Leeches have been reported in body cavities open to and nearer to the exterior. We report a 2-year- old girl with intraperitoneal leech. The leech entered her vagina and uterus, perforated the uterus and entered into the peritoneal cavity (Fig.68 and 69). Fig. 67. Leech attacks on the legs of men. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig.68: Leech popping out from the pelvic cavity. Fig.69: Postoperative leech specimen. The marine annelid (Nereis)is used to derive biopesticide which is the only pesticide of animal origin (Fig.70). Cartap Hydrochloride(Imidacloprid) is an analogue or pro-Crop Protection product of the natural toxinNereis. It is a systemic insecticide with stomach and contact action. Fig.70. External body features of ragworm, Nereis sp. (dorsal view). Ragworms' jaws are now being studied by engineers as they offer an exceptional combination of lightness and strength. Examples: Earth worms, Nereis, Leech, etc. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Tardigrada The name water bear comes from the way they walk, reminiscent of a bear's gait. The biggest adults may reach a body length of 1.5 millimetres; the smallest below 0.1 mm. The freshly hatched tardigrades may be smaller than 0.05 mm. About 1,150 species of tardigrades have been described. Tardigrades occur throughout the world, from the Himalayas (above 6,000 metres), to the deep sea (below 4,000 metres and from the polar regions to the equator. Tardigrades (commonly known as water bears or moss piglets) are small, waterdwelling, segmented micro-animals with eight legs.Body segmented with pairs of unsegmented legs terminates in claws.Usually, tardigrades are 1 millimetre long when they are fully grown. They are short and plump with 4 pairs of legs, each with 4-8 claws also known as "disks."The animals live in freshwater, terrestrial and marine ecosystem and are prevalent in moss and lichen and, when collected, may be viewed under a very low-power microscope. The diagram showing internal and external body features and electron micrograph showing features and colour of different species are given in Fig.71-78. Tardigrades are notable for being one of the most complexes of all known polyextremophiles. An extremophile is an organism that can thrive in a physically or geochemically extreme condition that would be detrimental to most life on Earth. For example, tardigrades can withstand temperatures from just above absolute zero to well above the boiling point of water, pressures about 6 times stronger than pressures found in the deepest ocean trenches, ionizing radiation at doses hundreds of times higher than would kill a person, and the vacuum of outer space. They can go without food or water for more than 10 years, drying out to the point where they are 3% or less water, only to rehydrate, forage, and reproduce. Examples: Macrobiotus hufelandi, Echiniscussp., Hyphsibussp., etc. Fig. 71. Macrobiotus hufelandi. I-IV, appendages; b. c, buccal cavity; gl, accessory gland; mal, Malphigian tube; ov, ovary; ph, pharynx; r, rectum; sal.gl, salivary glands; st, stomach; t, teeth. Fig. 72. Echiniscus sp. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig. 75. Echiniscus testudo Doyère, Fig. 73. Water bear (tardigrade), Hypsibius dujardini, scanning electron micrograph. Fig.76.Coloured scanning micrograph of a marine tardigrade (Macrobiotus sp.) showing 4 pairs of stumpy legs terminating in claws for clinging to sand or soil. Fig.77. Coloured scanning electron micrograph of a water bear (Echiniscus testudo) in its crypto-biotic tun, or barrel, state. Fig.74. Giant yellow water bear (Richtersius coronifer); 2. Large carnivorous water bear (Milnesium tardigradum); 3. Tidal water bear (Echiniscoides sigismundi sigismundi); 4. Turtle water bear (Echiniscus testudo); 5. Balloon water bear (Tanarctus bubulubus). Fig.78.Pseudobiotus sp. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Onychophora The velvet worms (Onychophora — literally "claw bearers", also known as Protracheata) are a minor ecdysozoanphylum with ~180 species. These obscurely longsegmented organisms have tiny eyes, a pair of anteriorantennae, multiple pairs of shortlegs ending in clawsand slime glands. Body surface with thin cuticle and live in moist terrestrial. ecosystem. Examples: Peripatopsis, Peripetus.They have variously been compared to worms with legs, caterpillars and slugs. Most common in tropical regions of the Southern Hemisphere, they prey on smaller animals such as insects, which they catch by squirting adhesivemucus. In modern zoology, they are particularly renowned for their curious mating behaviour and for bearing live young. Fig.79. Peripatoides novaezealandiae clearly showing the stub feet. Fig. 80. Velvet Worm (Onychophora) from the Amazon Rain Forest in Peru. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology The two existing families of velvet worms are Peripatidae and Peripatopsidae. They show a peculiar distribution, with the peripatids being predominantly equatorial and tropical, while the peripatopsids are all found in what used to be Gondwana. A B Fig.81.A. Cephalofovea tomahmontis. B. Epiperipatus biolleyi. Phylum: Arthropoda: (Joint- footed animals). This is the largest and most successful of the animal phyla, with over 1 000 000 species identified to date and an estimated 1 000 000 000 000 000 000 (1 quintillion) individuals alive today. Some scientists estimate that there are over 200 million insects for each single person. That's just insects alone, and doesn't include the countless millions of spiders, crustaceans, millipedes, centipedes, etc that make up the phylum Arthropoda. Arthropods are the most easily identifiable of all the animals, as they all share several things in common that no other animals share. First, all arthropods are protected by a hard exoskeleton (outer skeleton) composed of chitin (humans and other chordates have Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology endoskeletons). This exoskeleton protects the organs, gives support for the body, and allows for efficient locomotion. However, because the exoskeleton cannot grow, it must periodically be shed to allow for the organism to grow. 1 2 3 4 5 6 Fig.82. (1) Kolihapeltis, (2) Stylonurus, (3) Scorpion, (4) Crab, (5) Centipede, (6) Butterfly. All arthropods have segmented bodies divided into a head, thorax, and abdomen. In some cases, such as with the lobster, the head and thorax are fused together. This is called a cephalothorax. Arthropods also have jointed appendages. Arthropods have a complex internal design, with ganglia for a brain and an organ that acts as a heart. They have a complete digestive and excretory system. They also have welldeveloped sensory organs, that include antennae, eyes, and in some cases internal ears. Respiration occurs through the body surface, gills, trachea, and/or book lungs. The oxygen is carried in an open circulatory system (humans have a closed circulatory system). Arthropods are very diverse, and therefore can locomote in a wide variety of ways. They can walk, crawl, climb, hop, fly, glide, swim, skate, dive - almost every type of locomotion imaginable (other than drive!). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Arthropods can be found on every continent and in every ocean, from the harsh climate of Antarctica to the dryness of the desert to the dampness of the rainforests, and even to the darkness of the ocean's depths. They can be found in ponds, on plants, in and on other animals, and even in your own house!Examples:Scorpions, prawns, flies, centipedes, etc. Phylum: Pentastomida Pentastomida are an enigmatic group of parasiticinvertebrates commonly known as tongue worms due to the resemblance of the species of the genus Linguatula to a vertebrate tongue.Alternative names for the Pentastomida include Pentastoma (strictly a genus name), Linguatulida, and Acanthotheca Examples: Porocephalus, Cephalobaena etc. There are about 130 extant species of pentastomids; all are obligate parasites with correspondingly degenerate anatomy. Adult tongue worms vary from about 1 to 14 centimetres in length, and parasitize the respiratory tracts of vertebrates. They have five anterior appendages. One is the mouth; the others are two pairs of hooks which they use to attach to the host. This arrangement led to their scientific name, meaning "five openings", but although the appendages are similar in some species, only one is a mouth. Pentastomids are worm-like animals with two anterior appendages terminating in claws and ranging from 2 to 13 centimetres in length. The anterior end of the body bears five protuberances, four of which are clawed legs, while the fifth bears the mouth. The body is segmented and covered in a chitinous cuticle. The digestive tract is simple and tubular, since the animal feeds entirely on blood, although the mouth is somewhat modified as a muscular pump (Fig.83 and 84). These are blood sucking endoparasites of vertebrates. The nervous system is similar to that of other arthropods, including a ventral nerve cord with ganglia in each segment. Although the body contains a haemocoel, there are no circulatory, respiratory, or excretory organs Fig.84. Adult female Linguatula serrata. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Subdivision(iv): Lophophorate Coelomata Coelom develops as schizocoel or enterocoel and with a crown of hollow tentacles (Lophophore) surrounding the mouthy. Head indistinct. Gut U- shaped. Phylum: Phoronida The Phoronida is one of the smallest and least familiar phyla; there are about twenty species in two genera, Phoronis and Phoronopsis. However, phoronids -- or "horseshoe worms," as they are sometimes called -- may be abundant in shallow marine sediments at certain localities. Phoronids are elongated and worm-shaped, but the gut loops and ends close to the mouth, instead of passing straight through the body as in annelids and many other wormlike organisms (Fig.85 and 86).The mouth is surrounded by the ciliated feeding structure known as a lophophore. The lophophore of phoronids is a simple ring in some species, folded into a horseshoe shape in others, as shown in Phoronopsis viridis(Fig.87)(hence the common name "horseshoe worms"), and coiled in other species. Fig.86. Phoronid secreted tubesof agglutinated sedimentas shown around their bodies. Fig.87.The horseshoe shape lophophore of a phoronid (Phoronopsis viridis). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology They are found in all oceans and seas (except the polar seas), and all species have wide geographical ranges.They occur at depths up to 400 metres, but mainly between 0 to 70 metres. The life span is thought to be about a year. The adults are tube worms, and secrete chitinous tubes in which they live. These tubes can be buried in the mud or sand or rest on the surface of a rocky substrate. If on rocks, they may live in colonies with their tubes become twisted around each other for support. Some species can dissolve away holes in rocks such as limestone, calcareous seashells or even cementpiers; they then live in these holes which they line with their secreted tubesof agglutinated sediment, as shown in Fig.86, around their bodies. Though they are normally long, up to 50cm. Phoronids are normally very thin.Examples: Phoronis, Phoronopsis etc. They feed using a lophophore, a ciliated structure that surrounds the mouth. Phoronids are suspension-feeders. They move their lophophores into the prevailing water current. Food particles in the water current are trapped in a stream of mucous that travels along the tentacles until it reaches the oral ring. There it is drawn into the mouth and then on into the digestive tract. Direct uptake of amino acids through the epidermis also occurs. Phylum: Bryozoa or Ectoprocta The Bryozoa, also known as Ectoprocta or commonly as moss animals, are a phylum of aquatic invertebrate animals. Typically about 0.5 millimetres long, they are filter feeders that sieve food particles out of the water using a retractable lophophore, a "crown" of tentacles lined with cilia. Most marine species live in tropical waters, but a few occur in oceanic trenches, and others are found in polar waters. One class lives only in a variety of freshwater environments, and a few members of a mostly marine class prefer brackish water. Over 4,000 living species are known. One genus is solitary and the rest colonial.Examples: Bugula, Plumatella, etc. Fig.88. Bugula flabellate colony hanging from rock race. Fig.89. A marine colony of Bryozoa. Phylum: Brachiopoda The word "brachiopod" is formed from the Ancient Greek words βραχίων ("arm") and πούς ("foot"). They are often known as "lamp shells", since the curved shells of the classTerebratulida look rather like pottery oil-lamps.Brachiopoda (from Latin bracchium, arm + New Latin -poda, foot) is a major invertebrate phylum, whose members, the brachiopods, are sessile, two-shelled, marine animals with an external morphology Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology resembling bivalves (that is, "clams") of phylum Mollusca to which they are not closely related. Brachiopods are found either attached to substrates by a structure called a pedicle or unattached and resting on muddy bottoms. Brachiopods are suspension feeders with a distinctive feeding organ called a lophophore found only in two other suspension-feeding animal phyla, the Phoronida (phoronid worms) and the usually colonial Ectoprocta or Bryozoa. Characterized by some as a "crown" of ciliated tentacles, the lophophore is essentially a tentacle-bearing ribbon or string that is an extension (either horseshoe-shaped or circular) surrounding the mouth.Examples: Lingula, Crania, etc. 1 2. Fig.90. Lamp shells:1. Kraussina mercatori; 2.Pajaudina atlantica. Fig.913. Spiriferina rostrata with visibleskeleton of the lophophore. Division 2: Deuterostomia Mouth arises some distance away from blastopore. Subdivision: Enterocoelous Coelomata Coelom is an enterocoel which originates as pouches of embryonic gut (archenteron). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Chaetognatha (Arrow worms). Chaetognatha, meaning bristle-jaws, and commonly known as arrow worms, are a phylum of predatory marine worms that are a major component of plankton worldwide. About 20% of the known species are benthic, and can attach to algae and rocks. They are found in all marine waters, from surface tropical waters and shallow tide pools to the deep sea and Polar Regions. Most chaetognaths are transparent and are torpedo shaped, but some deep-sea species are orange. They range in size from 2 to 120 millimetres (0.079 to 4.7 in).They have small elongated body bearing fins and anterior end with grasping spines. There are more than 120 modern species assigned to over 20 genera. Despite the limited diversity of species, the number of individuals is large. Examples: Sagitta, Spadella, etc. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Echinodermata Echinodermata (əkī´nōdûr´mətə) [Gr.,=spiny skin], phylum of exclusively marine bottom-dwelling invertebrates having external skeletons of calcareous plates just beneath the skin. The plates may be solidly fused together, as in sea urchins, loosely articulated to facilitate movement, as in sea stars (starfish), or reduced to minute spicules in the skin, as in sea cucumbers. The skin usually has warty projections or spines, or both. Echinoderms display pentamerous radial symmetry, that is, the body can be divided into five more or less similar portions around a central axis. Unlike other radially symmetrical animals, they develop from a bilaterally symmetrical larva and retain some degree of bilateral symmetry as adults. There is no head; the surface containing the mouth (the underside, in sea stars and most others) is called the oral surface, and the opposite side, which usually bears the anus, the aboral surface. There are five living classes of echinoderms. There are a total of about 7,000 extant species of echinoderm as well as about 13,000 extinct species. They are found in habitats ranging from shallow intertidal areas to abyssal depths.Spiny-skinned animals, about 6,000 species, secondarily pentamerous radial symmetry. A part of coelom as water vascular canals. Locomotion by tube feet. Examples: Brittle stars, starfish, sea urchins, sea lilies, etc. 1 2 3 4 Fig.95.1. Sea urchin (Strongylocentrotus purpuratus); 2.Ruby brittle star or brain coral (Ophioderma rubicundum); 3. Echinaster sepositus;4. Sea lilies or feather star (Crinoidea). Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Pogonophora The name Pogonophora is Greek for "beard-bearers," and comes from the fact that many species have from one to many tentacles at the anterior end. These tentacles somewhat resemble the lophophore found in animals like brachiopods and bryozoans, as well as the feeding tentacles of certain chordates. The incompletely known anatomy of pogonophorans was interpreted to show that pogonophorans were chordate relatives. Because pogonophorans live with their lower ends buried in mud, and were broken during the dredging process, it was not until 1964 that a complete pogonophoran was recovered. It turned out that pogonophorans have a segmented posterior end of the body -- the opisthosoma -- that bears setae and resembles an annelid body. The forward part of the body, or prosoma, is unsegmented. Because of the segmented opisthosoma, and because pogonophoran larvae have been found to look very much like annelid larvae, pogonophorans are now considered to be close relatives of the annelids, and are classified with them in a larger group, the Trochozoa. 1 2 Fig.98. Pogonaphora: 1.Lamellibrachia luymesi van der Land and Nørrevang; 2. Riftia pachyptila Jones. About 80 pogonophoran species are known today, with new species still being discovered. One of the most spectacular zoological discoveries of recent years was the Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology finding in 1977 of giant pogonophoran worms, 1.5 meters long, growing in heated, sulfurrich water around warm-water vents in the Pacific Ocean, 2600 meters below the surface (pictured at right). These worms are sometimes placed in their own phylum, the Vestimentifera, but they are similar to pogonophorans in most respects, and the current tendency is to group these rift-dwelling worms together with the rest of the Pogonophora into one phylum. Phylum: Hemichordata Hemichordates ("half (½) chordates") have some features similar to those of chordates: branchial openings that open into the pharynx and look rather like gill slits; stomochords, similar in composition to notochords, but running in a circle round the "collar", which is ahead of the mouth; and a dorsal nerve cord - but also a smaller ventral nerve cord. There are two living groups of hemichordates. The solitary enteropneusts, commonly known as "acorn worms", have long proboscises and worm-like bodies with up to 200 branchial slits, are up to 2.5 metres long, and burrow though seafloor sediments. Pterobranchs are colonial animals, often less than 1 millimetre long individually, whose dwellings are interconnected. Each filter feeds by means of a pair of branched tentacles, and has a short, shield-shaped proboscis. The extinct graptolites, colonial animals whose fossils look like tiny hacksaw blades, lived in tubes similar to those of pterobranchs. Hemichordata is a phylum of marine deuterostomeanimals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and include two main classes: Enteropneusta (acorn worms), and Pterobranchia. A third class, Planctosphaeroidea, is known only from the larva of a single species, Planctosphaera pelagica. The extinct class Graptolithina is closely related to the pterobranchs. Fig.99. Acorn worm, a hemichordate. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Fig.14. Body parts of acorn worm. Acorn worms are solitary worm-shaped organisms. They generally live in burrows (the earliest secreted tubes) and are deposit feeders, but some species are pharyngeal filter feeders. Many are well known for their production and accumulation of various halogenated phenols and pyrroles. Pterobranchs are filter-feeders, mostly colonial, living in a collagenous tubular structure called a coenecium. The body plan of hemichordates is characterized by a tripartite organization. The anteroposterior axis is divided into three parts: the anterior prosome, the intermediate mesosome, and the posterior metasome. The body of acorn worms is worm-shaped and divided into an anterior proboscis, an intermediate collar, and a posterior trunk. The proboscis is a muscular and ciliated organ used in locomotion and in the collection and transport of food particles. The mouth is located between the proboscis and the collar. The trunk is the longest part of the animal. It contains the pharynx, which is perforated with gill slits (or pharyngeal slits), the esophagus, a long intestine, and a terminal anus. It also contains the gonads. The prosome of pterobranchs is specialized into a muscular and ciliated cephalic shield used in locomotion and in secreting the coenecium. The mesosome extends into one pair (in the genus Rhabdopleura) or several pairs (in the genus Cephalodiscus) of tentaculated arms used in filter feeding. The metasome, or trunk, contains a looped digestive tract, gonads, and extends into a contractile stalk that connects individuals to the other members of the colony, produced by asexual budding. In the genus Cephalodiscus, asexually produced individuals stay attached to the contractile stalk of the parent individual until completing their development. In the genus Rhabdopleura, zooids are permanently connected to the rest of the colony via a common stolon system. They have a diverticulum of the foregut called a stomochord, previously thought to be related with the chordate notochord, but this is most likely the result of convergent evolution rather than homology. A hollow neural tube exists among some species (at least in early life), probably a primitive trait they share with the common ancestor of chordata and the rest of the deuterostomes. Some species biomineralize in calcium carbonate. Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology Phylum: Chordata Chordates, members of the phylumChordata, are deuterostomeanimals possessing a notochord, a hollow dorsal nerve cord, pharyngeal slits, an endostyle, and a post-anal tail for at least some period of their life cycles. Taxonomically, the phylum includes the subphyla Vertebrata, including mammals, fish, amphibians, reptiles, birds; Tunicata, including salps and sea squirts; and Cephalochordata, comprising the lancelets. Primitive chordates are known from at least as early as the Cambrian explosion. There are more than 75,000 living species of chordates, about half of which are bony fish of the class osteichthyes. The world's largest animal, the blue whale, and fastest animal, the peregrine falcon, are chordates, as are humans,snakes, birds, Ascidians, Amphioxus, fishes, frogs, rabbit, etc. Subphylum: Urochordata The tunicates are marine, filter-feeding animals (Fig. 102). The most prominent tunicates are the sea squirts (class Ascidiacea), which show affinities to other chordates only in the juvenile stage. Adult sea squirts are sessile (attached), globular or tubular animals, often with prominent incurrent and excurrent siphons; many kinds grow in colonies. Most of the body of the adult is occupied by a very large pharynx with numerous gill slits that act as a sieve for food. Water taken into the incurrent siphon enters the pharynx and passes out through the gill slits, leaving food particles trapped in the pharynx. A groove in the pharynx called the endostyle secretes mucus that traps the particles and conveys them into the digestive tract; the movement of the mucus is caused by the action of cilia. Water leaves the atrium, a sac surrounding the pharynx, by way of the excurrent siphon. Thus the gill slits in tunicates serve a feeding function, not a respiratory function. The sea squirt larva is a free-swimming animal resembling a tadpole. The head, which will become the entire body of the adult, contains a rudimentary brain and sense organs, a small pharynx and digestive tract, and a ventral heart. Incurrent and excurrent openings are Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology located at the top of the head. The tail is a muscular appendage that functions as a swimming organ. It contains a hollow nerve tube (connected to the brain), and a notochord that extends into the head and keeps the animal from telescoping when its muscles contract. When the larva is ready to undergo metamorphosis it attaches to an object head downward. The tail, notochord, and nerve cord degenerate, the pharynx enlarges, and the other organs shift in position; the incurrent and excurrent openings develop siphons. Fig.102. A colony of sea tulips - There are two other classes of tunicates, both consisting of small planktonic animals. The salps (Thaliacea) metamorphose into barrel-shaped adults that swim by muscular contractions. The larvaceans (Larvacea) are neotenous, that is, they achieve sexual maturity and reproduce without losing the larval form. Many zoologists believe that tunicates of the sea squirt type were the first chordates and that the larval tail, with its notochord and nerve chord, was evolved as a means of dispersing their larvae. According to this theory, the later chordates, including the vertebrates, are descended from neotenous tunicates that, like the larvaceans, failed to assume the adult form. Subphylum Cephalochordata This class includes the several species of lancelets (Fig. 103), or amphioxi, small, fishlike, filter-feeding animals found in shallow water. A lancelet has a long body, pointed at both ends, with a large notochord that extends almost from tip to tip and is present throughout life. At one end is a mouth surrounded by prominent bristles and leading into a pharynx. The Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology pharynx has gill slits, an endostyle similar to that of a sea squirt, and an atrium surrounding the pharynx. Water enters the mouth and leaves through the gill slits, and food is trapped in the pharynx. The dorsal, tubular nerve cord is slightly enlarged in the anterior region, forming a rudimentary brain. Nerves extend from the nerve chord to other parts of the body. The muscles, as in fishes, are a series of cone-shaped blocks that fit into each other like stacked paper cups. This is the most primitive occurrence of the segmental body wall structure characteristic of lower vertebrates. The colorless blood moves forward through a ventral vessel and back through a dorsal vessel, in the typical chordate pattern. There is no major heart, although many small enlargements of the vessel serve the function of hearts. There are no blood cells and no respiratory pigments. The excretory system, like that of many invertebrates, consists of segmentally arranged nephridia; there is no kidney. The gonads, unlike those of any other chordate, are numerous and segmentally arranged. Fig.103.ALancelet (or Amphioxus) specimen (Branchiostoma lanceolatum)collected in coarse sand sediments (600 µm) on the Belgian continental shelf. Subphylum: Vertebrata Vertebrates constitute the vast majority of living chordates, and they have evolved an enormous variety of forms (Fig.104). The backbone of vertebrates protects the nerve cord and serves as the axis of the internal skeleton. The skeleton provides strength and rigidity to the body and is an attachment site for muscles. The vertebrae in the middle region of the trunk give rise to pairs of ribs, which surround and protect the internal organs. A cartilaginous or bony case encloses the brain. Bone is a substance unique to vertebrates. It was formerly thought that vertebrates with cartilage skeletons (cyclostomes and shark like fishes) were descended from early vertebrates that had not yet developed bone. However, very primitive fishes with bone skeletons are known from the fossil record, so lack of bone is now believed to be a degenerate rather than a primitive feature. All but the most primitive vertebrates, known as jawless fishes, have jaws and paired appendages. The fishes and, to a lesser extent, the amphibians and reptiles show a segmental arrangement of the muscles of the body wall and of the nerves leading to them. There are eight vertebrate classes. Four are aquatic, and may be grouped together as the superclass Pisces, or fish; four are terrestrial or (in the case of amphibians) semiterrestrial, and may be grouped as the superclass Tetrapoda, or four-footed animals. Fishes breathe water by means of gills located in internal passages, although they may also have lungs as Professor Paras Nath’s Agricultural Biology Lecture Note CAFF, FNU-2013 L 4 ENT 402: Agricultural Biology supplementary air-breathing organs. Most move through the water by weaving movements of the trunk and tail. All have fins, and most have two sets of paired fins (pelvic and pectoral). Tetrapods breath air, usually by means of lungs, and never have gills as adults, although the amphibians go through a gilled, water-breathing stage. Except where the appendages have been lost, as in snakes, all have two pairs of limbs, generally used for locomotion; these are homologous to the pelvic and pectoral fins of fish. 1 2 3 4 5 Fig.104. 1. Fire Salamander (Salamandra salamandra); 2. Saltwater Crocodile(Crocodylus porosus) 3. Southern Cassowary (Casusarius casuarius); 4. Black-and-rufus Giant Elephant Shrew (Rhynchocyon petersi) 5. Ocean Sunfish (Mola mola). XXXXXXXX