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Classification of Chordates
Chordate
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ee Phylum Chordata pase | \1ded into two groups: Nonchordata (or Invertebrata) and Chordata, on the \! notochord (Gr., novon = back; L. chorda = a cord). The nonchordates lack ed into 26 phyla. Their major phyla are Porifera, Cnidaria, Platyhelminthes, opoda, Mollusca, Echinodermata, Lophophorate phyla and Hemichordata. tochord and form a single phylum Chordata. est phylum of deuterostome and includes a majority of living and extinct lancelets, lampreys, fishes, amphibians, reptiles, birds and mammals. ¢s may be either protostomia (in which blastopore becomes mouth) or astopore becomes anus), chordate: s deuterostomia. Lastly, ar, most adaptable, most successful and most widely distributed animals, orm, habitat and habits to an amazing degree. Only the arthropods among the |, phylum Chordata marks the climax of evolution. © from Cambrian to recent period. There are more than 52022 extant species of chordates. |.1_, GENERAL CHARACTERS OF CHORDATES £5 -y y: Segmented body; three germ-layers (triploblastic); well-developed coelom Notochord or skeletal rod present at some stage in the life cycle. Single, dorsal, tubular nerve cord; anterior end of the cord usually enlarged to form brain Phary ngeal pouches present at some stage in the life cycle; in aquatic chordates these develop Postanal tail, usually projecting beyond the anus at some stage but may or may not persist, Segmented muscles in an unsegmented trunk 7. Ventral heart, with dorsal and ventral blood vesse! lar system). iplete digestive system (having mouth and anus). ‘laginous or bony endoskeleton present in the majority of higher chordates losed blood system (or circulatory system FOUR CHORDATE HALLMARKS
Subphylum 2. Cephalochordata J. Vertebrata Divisions/Sections (2) f bas mare 1, Agnatha 2. Gnathostomata Classes @2){ ani? wand 1, Ostracodermi 2, Cyelostomata Superclasses QQ) me 1. Pisces 2. Tetrapoda Classes Q) Classes (4) 1, Placodermi 1. Amphibia 2. Chondrichthyes 2, Reptilia 3, Ostoichthyes 3, Aves 4. MammaliagemesTen Ill ¢ STUDENTS gpomantt ST aryngen! ow 8H dorwal NON re / rely above the caccum “Meuie) dorsal strip mouth pharynx unk — hepatic na mich (Fig. 1.2). 1dr i) mesoderm that a nrchentti oe ‘ ore acomotory suppor It grammatic $i rat andl 10 IM acs Dla re Srwiesscuaaearases Fi 11+ Ctl four chorale Nelms the body of Lr Mtochord serves 8 2008). The note ‘ primitive inter em and the segmental muscles al nervous § oC EEO, Eke} ee! notochord é fibrous sheath elastic sheath f the 4 d and its surrounding sheaths. Cells of the n , dre thick walled, pressed together closely, and filled with semifluid. Stiffness is y ding connective tissue mainly by turgidity of fluid-filled cells and surroun pimive "ype of endoskeleton provides longitudinally stiffening of the main body, pice for trunk muscle, and an axis around which the vertebral column develo Hickman et al,, 1996). Fig, 1.2. Structure of the notochort In Cephalochordata (Branchiostoma) the notochord extends from one end of the other and persists throughout life, In Urochordata (Herdmania) the notochord is res caudal region and is present only in larval stage. Cyclostomata (lampreys and hagfishes) p turgid and rigid notochord persisting throughout life. In Scoliodon, the notochord is constri chain of vertebrae formed of cartilage. It is only represented by a notochordal filament in In vertebrates the notochord is completely replaced by the vertebral column. Notochord is one of key synapomorphies defining the chordates. Synapomorphy is derived or new character or evolutionary novelty (Brusca and Brusca 2003). \ 2. Dorsal tubular nerve cord, The nervous system of chordates is in the form ofa which is situated mid-dorsally above the notochord and immediately beneath the body wall. Itdé pee eee of the neural ectoderm in the mid-dorsal line, so that a medull mb ensiceies aie ale ot (folds) of the groove fuse together to form the Serica low the surface and forms the neural tube. Neural tube devel Pedncs, oe nerve cord is called neurocoel. The nerve cord persists es, becoming differentiated into an anterior short, wide brain and ap long, slender spinal ie, pinal cord. In some lower chordates (urochordates), nerve cord 3. Branchial or Pharyn ; geal clefts, inal the chordates at some ‘stage of their li a The branchial or pharyngeal clefts are fo life cycle. These occur.as paired series ofBe ee exer Sn ihe arpa lefts cvery chordstes oo . r oil throughout life i lower chondates Selacrahy, They persis het Oe larval ae poucioes tailless hans te ae cowed races, the Paar atea! fier feeding and to some extent it the gills » 4. Postanal tail. The tail isthe part . br min, nerve cord, blood vessels, nerv visee a! the chordates (even human embryo (fishes). the 2 tc vecomovory organ, while in the Serres) shapes and functions he ea cee cris ch HO of the body behind the cloacal or anal #PeaNtE: not actos, bus lacks. corkons 5 potoch Some Additional but Unique Chordate Characters of the prasy=™ 1. Endestyle. Endostyle is 2 ciliated, glandulss groove in the ¢ hordates and c and in the larva 1n the adult uroc! cephalochordates, Soe bich wraps tiny food partcies Sor SaasPoce “The thyroid functions 25 25 :semester lll ) develop from the ortebrate phyla) s WITH NONCHORDATE PHYLA posterior ends, and nirror images of e 1 (including opti com the epider p BY CHORDATE! ral symmetty ft sides of the Jane divide: 1. Bilateral symm dian lon vertical) P! jority of n° es with the ant ital medi Fall chordates anid ma ; notry of body ves polarity to the animal's body so that it mov ral SYTHE owever, the anterior end of the chordates does not co ; nt do nthe rte echinoderms, lophophe ates and her | called the protoston nofichordates (exce} chorates. In the vverived from the blastopore of the ‘embryo (ie., gastrual is (tig, 14A), In contrast, in chordates > also in enchinoderms, lophophe pore and the mouth develops as an invagination the blast n (Fig. 1.4B). vad also all nonchordates above the coel and endoderm, and are said to be triple (erm arises from the do anus appears ne stopore), called t Joblastic nature. All chords layers: ectoderm, mesoderm different! In chordates, mesod ene of archenteron, whereas in nonchordates it mostly develops & totem and endoderm fig. 1.4A ein echinoderms and chordates, them 3 a chenteron itself (ie., a preformed endoderm), either as a solid ordat he deuterostome conditi lop from three germ | cof mesoderm is, howe m the junction of arises from the wall of the as pouches (Fig. 1.48) ~- ectoderm ~~ so, S= blastocoe! — presumptive: mesoderm archenteron — ‘endoderm ‘mesoderm from derivative es of a me: Brusca and Brusca 2 003).7. Endoskeleton PHYLUM CHORDATA, 7 nonchordate phyla lack metamerism, except ‘Annelida and h present, is of different type. 5. Cephalization seine the formation of head by concentration of some vice organs at the anterior end) The head contains brain, nares or olfactory sense ofgans, eyes, ears, gustatory sense ‘ "Wy and directs its activities, for example, head being at the cata changes during progression. In chordates, the size and sm the cyclostomes to the mammals. The phenomenon onchordates, have only two pairs of paired appendages (fins or limbs). One lost) Mud-eel and whale have lost hindlimbs and snakes have ichardates have more than two pairs of paired appendages, if 6. Appendages. endoskeleton of cartilage or/and bone is characteristic of most oath the animal's growth, for it can increase in size with rest of the shed it, The skeleton of nonchordates, if present, is always non-living, ¢ «or chitinous. The rigid exoskeleton is periodically moulted to allow ¢ body as in arthropods, ‘Coclom. Covlom is the perivisceral space lined by mesodermal epithelium. It is present in s nonchordates. The animals with coelom are described as coelomated. In vim mostly arises by a split in the mesoderm and is said to be schizocoelie cc ample. in schizocoely of annelids, bilaterally paired packets of mesoderm gradually Cventually becoming thin-walled coelomic spaces (Fig. 1.5A, B). The number of ‘ns varies among different animals and is frequently associated with segmentation, d worm (Fig. 1,5C). ¢a—anmea ren tl ENTS say Fon DRANEE ATUDEN coelomic > ypaces mesoderm iy 1.8 Coelom formation by schizocoely (Irontal sections). A—Precoelomic cor paired packets of mesoderm. B—Hollowing of the mesodermal packets to of coelomic spaces, C—Progressive proliferation of serially arranged pairs of spece®. This process occurs in metameric annelids (after Brusca and Brugea In lower chordates (cephalochordates) the coclom develops as linear se es of pai ch terocoelic, The higher chordates woatified form of enteroooelic coelom (Pig. 1.6; Box 1.1), teron and is described asPHYLUM CHORDATA, | enterocoaly. In most direct sort of enteroooely, rm snd the same process (Fig. 1.6A). Its called arc! he walls of these pouches: Chordates have two types and coolom formation are or Pouch or pouets form in the gut as a complete coelomic compartment 1 ‘ BBouch or pouchos jor in some cases the mesoderm arises from the well of Srcie Taz as oer Senet or late that later bocomes Bllayered and hollow (Fig. 1.68). Some authors consider this Procons to boa form of schizocos! (because of the “spliting” of the mesodermal plate), but it is in Process 10 De a foro eri econly (@rusea and Brusca 2003). Enlerooosly frequently resets parte arrangoment of the body recs hich are disignated procoel, mesocoel and metacoe! (Fig. 1.6¢ 9. Digestive tract. Most chordates have a complete digestive tract, Such an alimentary canal A eee erat imesuon of food and an anus or cloacal aperture for egestion of Taeesé Nonchordate phyla above platyhelminthes also possess similar complete digestive tract cal 10. Closed circulatory system, In chordates the blood flows in a continuous system of blood onal vessels (arteries, capillaries and veins), all with definite walls Such @ circulatory system is described Y 4s closed. In this case blood does not come in direct contact with body tissues. Some nonchordates, aan rclida. alco have elosed circulatory system. Many nonchordates (arthropods, molluses), llow however, po: 11 open cireulatory system in which blood flows in sinus . Le., blood channels 38a having novwalls In open eitculatory system, the blood isin direct contact with the tissues of the body. IL. Special respiratory and_exeretory organs. For some primitive chordates, respiration and excietynn by body surface diffusion is adequate, However, in higher chordates, special organs for respiration (ills or lungs) and for excretion (kidneys) have developed. Special respiratory and crctetory orpans also occur in many nonchordates, ¢.g., gills and nephridia in Annelida; tracheae and Malpighian tubules in Insecta; and ctenidia and kidneys in Mollusca. 12. Endocrine glands. The glands that pour their secretions (i.., the hormones) into the blood for transport to the target organs are called endocrine or duetless glands. Such endocrine glands are found in chordates as well as nonchordates, but they are better developed in chordates than in nonchordates. 13. Single pair of gonads with gonoducts. single pair of gonads each of which is provided with a gonoduet for th similar condition is found in many nonchordates. 14. Mesentries. In chordates, the visceral organs are interconnected and suspended from the dorsal body wall by double-fold of peritoneum, called mesenteries. Mesentries are absent in most chordates. However, some nonchordates such as Neries and Asterias, have small mesenteries in nection with the alimentary canal. 15. Energy compound. Chordates and some nonchordates such as echinoderms, use sphocreatine in the energy cycle of muscle contraction. Most nonchordates use phosphoarginine the same purpose. 16 Cleavage. Chordate eggs undergo equipotential and radial cleavage. In equipotential “each early blastomere on separation from others may give rise to a complete embryo +h cleavage is called indeterminate cleavage). Most nonchordates have spiral cleavage which ‘determinate type, i., cach early blastomere has a fixed fate. First two divisions are alike in both radial and spiral cleavages. However, in radial cleavage third subsequent division planes are symmetrical to the axis of polarity. ‘Therefore in radial cleavage, eight cell embryo, cells of each tier lie directly above and in line with those of the lower tier i, 1.7). In case of spiral cleavage, subsequent divisions of 4-cell embryo result in the displacement blastomeres in such a way that they lie in the furrows between one another (Box 1.2). The chordates are generally unisexual and bear @ '¢ discharge of the sex cells. APHYLUM CHORDAT® 14 FERENCE NCES BETWEEN CHORDATES AND NONCHORDATES \ ' ih tying ton bse a ale J exoskeleton) a s, the food- cand epatic portal vein bri ver, where the the alimentary canal is not carried Such hepatic portal system is not found in the no nchordates. ually haemoglobin, which 1s contained in th RECs found dissolved in plasma of Jeated red blood cells veins and is od, ramifies into the heart but in chordates, the respiratory (red blood cells). In nonchordates, the haern the blood. (Note. The blood in phoronids (lophophorates) p that contain haemoglobin for carrying oxygen; Hickman e tal. 7. In nonchordates the blood in the dorsal vesse! flows from posterior to the anterior direction whereas in chordates blood always flows in anterior direction. . In nonchordates the coelom occurs as a single wide perivisceral cavi organs. In chordates the perivisceral cavity (coelom) is divided into Sev has a specific function. These are dorsal myocoel, middle nephrocoel, later capi In this way, the blood from nent is ossesses nuc! 1996) around the visceral ns and each al and ventral splanchnocoel 9. Inthe nerve cord, the nerve cells are arranged deep around the central canal in chordates and superficially on the periphery in nonchordates a of the nerve cord, whereas 10. In nonchordates the nerves arise mostly from the segmental gangli d ventral nerve roots. in chordates these arise from the dorsal an 11. The paired appendages in nonchordates vary in ‘number and may even correspond to the number | of body segments. In chordates, however, limbs are invariably two pairs (tetrapods) 12. Inhigher chordates (i bral column. Vertebral column is not found CLASSIFICATION form a large and heterogenous group, Phylum Chordata is divided yyla (Brusca and Brusca 2003) , L. Subphylum Urochordata (= Tunicata). The tunicates. Adult body form varies, but usually ing obvious trimeric organizations; body covered by thick « jysaccharide; without bony tissue; notochord restricted to tat! and usually found only in larval {and in adult appendicularians); gut U-shaped, pharynx (branchial chamber) typically a gill slits (stigmata ‘coelom not developed; dorsal nerve cord present in larval stagess . It includes four classes: Class 1. Ascidiacea, idia, Botrylus, Ciona, Molgula, etc. e., vertebrates) the notochord is replaced by the vert in nonchordates and lower v ertebrates. into following three e.g, Herdmania, Asc«), Small (to 7 em), fis it in adults, but wi data (= Acrania) \ eerially arranged, Maing ee 8) an fc in which they burrows g J segment vertebral cola Chordates usual sin case and (exceptifge am Verteb ith paired ape" recognized, although not alligg shi are the hagfishes and lamy Cephalaspidomor? on han fishes); Cl hy: Class Amphibia inglud - rout, tuna, perch) hie Clas tis traditionally included the tu cs, sry s; Class Reptilia oer tc (Class Aves) and reptiles togetig ification ples Opies the mammals (See Buse s Mammalia cc Myxini ANCESTRY AND EVOLUTION known truely vertel freshwater forms abundant during eee are collectively known as Ostracodertig oc n head. These jawless bottom-dwelling orming a shield on head wless bottom Drie ‘feeding (Fig, 1.8). The oldest fossils belonging te filter feeding (Fig. 1.8). The Cambrian and middle Ordovician periods Time of origin. The earl nian periods. T y a bony adapte Fig. 1.8. A0stracoderm (Cephalaspis) in dorsal (A) and lateral (B) view. intl recently, it was thought that the earliest known vertebrates w. fishes". However, it now appears that fossils long known from the called conodonts, belong tothe earliest known vertebrates, Teeth et al., 1992). Conodont ay imal (Fig, 1.9) were ‘small and ‘notochord In these respects, they resemble living hag-fs 2, Place of origin, origin of chordates. It w was also considered as the Protochordates and d marine forms. ere Ordovician jas | Cambrian, tooth-like structtnt — are made of cellular bone (Sams shaped; they lacked vertebrae, but had ‘o which they may be related, American geologist, Chamberlain (1900) Bave the idea of freshwatel 3.Hy developmes period som echinodem Althe group of f (450 mili small non covered13 PHYLUM CHORDATA t animal showing tooth-like structure, stor (Prevertebrate). There are several lines of anatomical, agesting that somewhat later, atthe base of the C ‘ambrian ‘70 million years ago, the frst distinctive chordates appeared, evolved from an early ordate, of a common ancestor of both (Fig, 1.10; Hickman ef al., 1996) ail ike--modern chordates, there is one curious group of fossil echinoderms, the *Caletchordata” which have lived during the Ordovician period 0 million years ago). Caleichordates had gill slits and possibly other chordate attributes. These da raene mmetrieal forms had a head resembling a long-toed medieva) oot, a series of gill slits iu with laps much like the gill openings of sharks, a flexible arm’ that could be interpreted as esi aad structures that are tentatively interpreted as a notochord and muscle blocks. They fs for suspension feeding, as do the primitive chordates today, Although, tthe right chordate characters based on soft anatomy, there is wrdates (which was calcium carbonate) d skeleton of calcichot dd ofa hydrated complex of calcium and phosphates called hvdroxyapatite). While such speculation bring us closer 10 an understanding of chordate origins, \vc are not yet in a position to know the precise characteristics of the long-sought chordate ancestor. However, we do know a great deal about the two living, protochordate groups that descended from ut, Le, Urochordata (Tunicata) and Cephalochordata (see Hickman et al., 1996). ‘4. Theories of Chordate Ancestry. (i Echinoderm ancestry. On the basis of anatomical, embryological, palaeontological, biochemical and serological evidences, various workers have tried to establish that chordates derm larva. The hemichordate larva probably origi ‘ated from some primitive echinoderm or echino But this view is (tornaria) is strikingly similar to the bipinnaria or dipleurula larva of echinoderms. not accepted these days. (ii) Hemichordate ancestry. Hemichordates are sedentary and have pharyngeal gill slits and a hollow dorsal nerve cord. But the presence ofa true notochord is doubtfull and their adult body plan is different from chordates. Hence they have been put under the separate phylum. Fig. 1.9. A Cambrian conodon! 3. Hypothetical vertebrate am developmental and molecular eviden period some rm, early hemicl Although modem echinoderms look nothing ap ipparently used their gill he calcichordates apparently had some o} ‘nv ineing similarity between the har that of vertebrates (which is composes andmodem ‘ampnibians = teeing TN cori reese primitive sessile fiter-feeder | OSs PRECAMBRIAN: PALEOZOIC ‘Geciogie time (My ago) Fig. 1.10. Phylogenetic tree of the chordates, The #ach group through geological time, bulging and thinning of that group's felative abundance in numbers of s indicated by the fossil record, is sug ine of descent (after Hickman ef ‘Amphioxus) js an interes aS a model Prevertebrate15 ; Bont ; Neuer & : rece Brusen and Brass? QUESTIONS Long Answer Questions 2W dsingushing fe eee De ories of ©avn anne: ATHDENTE ‘ wine ll vata il 41 annwnere wor Oven tan gy etna Faint of li wind ae # A svat ove toe oi lgnnci@ te Ful in the Blanka ' Jot pout Heong HE A vst get we ed waren nya wal jesntinne my 1 Chawla eevee iw jpharyi connotes yeh he eater Toit w part of body 6 Chote eat Ok Jn potion Woapiatiory pment (prevent (the Cleavage In ohana et nd % ave endonkeleton of fiving Uawuen, Multiple Choloe Questions 1, Chovlatoy are different fiom nonehordates in having () #hotwehord th), (0) a ventral solid nerve oor Ww) 2 Which one of the following is found in the ehardaton (9) chitinous exoskeleton () (6) postanal int ) i achizoonel wll the above willy spltweloy 4 How many pins of appendages are presen in vertatnaten (one pair fe) four tains 4 Chordates with backbowe are vatled (2) protoehordaten (h), Wy (6) gnathostomatey Hi y The wroup amnniotn inches (2) ids onl rainy b) (0) epviten nit vamiaaly i Renan OF Verlebratey in modified (0) umbitiog! cord i Iwo pairs Variable Horn wpecion to 9 on andl 1 vd Hoy He vertebral eavein eplavei oF blood in thie ehorditen, but not in nonehorditay brady sini Wile FOP en, Ini) Notoehord SPrNMLLE Gord ond enaee PHYLUM CHORDATA Ww 7, In vertebrates the skin is covered by ' (a) scale (b) hairs (e). feathers (d) any one of the above 8. Vericbrates resp (a) lungs (b) gills (e) skin (d) all of the above 9. The amniote egg first evolved in (a) bony fish (b) amphibian (c) reptiles (d) birds 10, Which group of the invertebrates are the closest relatives of the chordates? (a) Annelida (b) Mollusca ) Arthropoda (d) Echinodermata ANSWERS Very Short Answer Questions 1. Notochord is an elongated ular cord, skeleton of chordate embryos and adult cephalochordates, sence of notochord, (b) presence of dorsal tubular central nerve cnclosed in a sheath, which forms the primitive axial us system and (a) pr (©) presence of branchial clefts 3. Closed circulatory system. 4. Two pairs (hence called tetrapods) 5. Late Cambrian and middle Ordovician. 6. Vertebrates 7. Haemoglobin 8. Chordates, nonchordates except Echinodermata’ and Hemichordata, 9. Annelids. Fill in the Blanks slochordates, 2. Skeletal. 3. Middorsal, 4. Pharynx. 5. Postanal, postcloacae, aperture 1. Cepha 6. Ventral. 7. Erythrocytes. 8 Radial, equipotential. 9, Chordates Multiple Choice Questions 1. (a) 2 © 3. (b) 4. (d) 5. (d) 6. (b) 7 (@) 8. (d) 9% () 10. (d) jb 5
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