AN INTRODUCTION TO ANIMAL MORPHOLOGY AN INTRODUCTION TO ANIMAL MORPHOLOGY \M> Systematic Zoology. BY ALEXANDER MACALISTER, M. B. DUEL., "1- Co.Ml'AKATIYi: ANATOMY A NI) /( H )L( )( ; Y, \' Nl Y V. KS 1 IT t i F DTBI.IN. PART [.— INVERTEBRATA. LONDON: \ (, M A \ S, GREEN, AND CO ". IHT.LIN : prinlrb at the eluibcr:;itn PREFACE. THE study of Biology in the University of Dublin has been within the last few years greatly en- couraged by the introduction of Natural Science into the Undergraduate Course, and by the exten- sion of the means of practical instruction in Zoology and Botany. In teaching Zoology and Comparative Anatomy, I have found that Students desire to have a text- book in their hands to enable them to learn the terminology of the science, and by giving them a connected view of the varieties of animal forms, to assist them in remembering the practical instruc- tion of the class-room. There are in English many good works not intended to supplant such books, but to lit Students tor the profitable perusal of such works lose of Rolleston, IIuxl"-.-, l-'lou-er, To reduce my lecture notes to a suitable size, I have had to condense many parts to the smallest bulk, and to leave out many matters of detail, such as many of the recent embryological researches of Lankester, MetschnikofF, Kowalewsky, and others. I regret that, owing to the long time that this work (written in 1873) has been going through the press, I have not been able to introduce into it references to recent discoveries, such as that of the unisexuality and tracheal system of Peripatus, &c. For the sake of Junior Students, I have printed the chief paragraphs in larger type, the details in smaller, so that the general principles can be more easily grasped. I make no claim to originality, but have bor- rowed largely from Gegenbaur, Carus, Haeckel, Huxley, Lankester, Van Beneden, Schmarda, and others. I have endeavoured to avoid the errors of second-hand quotations, and of making extracts from other Manuals, which are easily accessible to Stu- dents; and, having been for fifteen years engaged in the practical study of Comparative Anatomy, I have been enabled to verify very many of the state- ments herein made. I return my best thanks to Mr. H. W. Mackintosh for his valuable assistance in correcting the press, and to Professor Reynolds for some suggestions in Chapter I. t'acc. vii I regret that some errors have escaped notice in correcting the proofs, the most serious of which is that of Pig. 4, p. 59, which should read Heliophrys variabilis, Grccf, instead of Heliosphaera radiata, Sch. CONTENTS. CHAPTER I. PAGE . Protoplasm.— Its properties and derivatives. Plastides, I CHAPTER II. General Morphology. — Organs, and their groupings, .... 8 CHAPTER III. Histology. — Epithelium and connective tissues, 12 CHAPTER IV. logy. — Muscular and nervous tissues, 20 CHAPTER V. U'y.— Individuality and the formation of 01 -an-. . . . 2.\ CHAPT1.K VI. iuction. — S; I u'lmvth, j>.ir.i>i- •Icgradation, 3-4 \ Contents. CHAPTER VII. PACK. Distribution of Animals. — Primary classification, . . . .42 CHAPTER VIII. Sub-kingdom I. — Protozoa. — Class i,Monera; Order I, Gymno- monera; Order 2, Leponumera. Class 2, Labyrinthuleae. Class 3, Rhizopoda ; Order I, Imperforata ; Order 2. Perforata. Class 4, Flugcllata ; Order I, Monadina ; Order 2, Noctthu ' 5, I'mtoplasta; Order i, Catallacta ; Order 2, Amu-bina ; Order 3, Heliozoa, . . 49 CHAPTER IX. Class <>, K.uliolun.t; Order I, Moiiucytt.irui ; Oidu 2. I'oly- ryttaiia. ( 'la-^ 7. < nr-.irinina. (>r> CHAPTER X. 7, InliiMtria ; Order I, Opalines; ('idci 2. iViuii: ( )idci v Acinetina ; Order 4, Stomatoda, .... (^> CHAITKR XI. Sub-kingdom II. i'nly^tnnuta or I'orifera ; Order I, Jlalisarcia ; Onler j icr 3, Gumminea2 ; Order 4, Halichondrue ; order 5. ( 'mtic.u.r order o, Acantho- spongia ; Order 7, Potamospongia ; Order 8, Cakispongia, . 72 CHAPTER XII. Sub-kingdom III. — Oeleiiterata. < 'la--- i , 1 lydru/o.i ; Sub-cl Hydroida; Order i, Eleutheroblastea, 79 CHAPTER XIII. Order 2, Gymnoblastea, CHAPTER XIV. Order 3, Calyptoblastea ; Order 4, Monopsca 05 n tents. xi CHAITKR XV. PAGB. Sub-class 2, Siphonophora ; Order i, Calycophoridae ; Order 2, Physophorida.'. Sub-class 3, Calvco/.oa ; Order I, Lucer- naridce, ........... 96 CHAPTER XVI. Sub-class 4, Discophora; Order I, Rhizostomida ; Order 2, Pelagiada ........... 102 CHAPTER XVII. Sub-class 5, Ctenophora; Order I, Eurystomata ; Order 2, Saccatae; Order 3, Taeniataj ; Order 4, Lobatze, . . . lob CHAPTER XVIII. Class 2, Actino/oa; Sub-class i, Zoantharia; Order I, Malaco- dermata: Order 2, Scleroderniata ; Order 3, Antipatharia, ........... 1 09 CHAPTER XIX. . '< yonaria ;' Order i. Alcyonaceoe ; Order 2, Tubipo- raceae ; Order 3, Pennatulacea? ; Order 4, Gorgon aceas ; Order 5, Isidaceae ; Order 6, Briarcacese ; Order 7,J Coral - ........ . . i i S CHAPTER XX. - .in-doiii IV. Mchinodt-nnata. ("la^ I, Stellerida ; Order I, HI.; ; < )idn ^ ( inioi.U'.i ; ( hder 4, Ophiuroidca ; ( )rdrr 5, l'.ii>in-..i«K .1 ; < >i>K r (>, A^teioidra, . \2\ < HAITI k XXI, • i.>: xii Contents. CHAPTER XXII. Class 3, Holothuroidea ; Order I, Apneumona Order 2, Tetra- pneumona ; Order 3, Dipneumona, . . . . . .148 CHAPTER XXIII. Sub-kingdom V. — Vermes. Province I, Archaeostomata. Cl. Turbellaria; Order I, Dendrocoela ; Order 2, Rhabdoccela ; Order 3, Nemertidea, ........ I ^ I CHAPTKR XXIV. Class 2, Cotylidea; On! lea, ...... i' i CHAPTER XXV. Order 2, Trematoda, ......... . 172 CHAPTER XXVI. ria-> 3, Xcmatelmia Order I, Gordiacere ; Order 2, Xematoda, . 477 CHAPTER XXVII. Class 4, Acanthocephal hyrea ; Order i, Inermia : Order 2, Armata, ......... 187 CHAPTER XXVIII. Class 6, Rotatoria ; Order i, Gasterodela; Order 2, Holotrocha ; Order 3, Schizotrocha ; Order 4, Zygotrocha, . . . .103 CHAPTER XXIX. Class 7, Hirudinea. Class 8, Onychophora, ..... iqS CHAPTER XXX. Province 2, Deuterostomata, Enteroccela. Class 9, Chaetognatha. Class 10, Enteropneusta, ........ 206 Contents. xiii CHAPTER XXXI. PAGE. Deuterostomata Schizocoela. Class 1 1, Chaetopoda, .... 209 CHAPTER XXXII. Deuterostomata Epiccela. Class 12, Bryozoa; Order I, Gymnolse- raata ; Order 2, Phylactolxmata, 227 CHAPTER XXXIII. Class 13, Tunicata, 231 CHAPTER XXXIV. Sub-kingdom VI. — Mollusca. Division, and Class i, Brachiopoda : Order i , Ecardines ; Order 2, Testicardines, . . . .241 CHAPTER XXX Y. -ion 2, Otocardia. Class i, Lamellibranchiata ; Sub-class r, Endocardines ; Sub-class 2, Exocardines; Order I, Mono- mya; Order 2, Heteromya ; Order 3, Isomya, .... 255 CHAPTER XXXVI. I 2, Cephalophora ; Sub-class i, Scaphopoda ; Sub-class 2, Pteropoda ; Order i, Thecosomata ; Order 2, Gymnosomata, . 272 CHAPTER XXXVII. Suh- poda; order i. 1'i.inchiata ; Order 2, I'ul- monata 280 MIAI'TKK XXXVIII. . Dihran- rhiata, CHAPTER XXXIX. l.iopo»l.j. xiv Contents. CHAPTER XL. PACI Sub-class i, Cirripedia ; Order I, Thoracica ; Order 2, Abdo- minalia ; Order 3, Apoda ; Order 4, Rhizocephala, . , . 343 CHAPTER XLI. Sub-class 2, Copepoda; Order i, Gnnthostomata ; Order 2, Sipho- nostomata. Sub-class 3, O>tracnda. Sub-class 4, Bran- chiopoda; Order I, Cladocera ; Order 2, I'hyllopoda. Sub- class 5, Trilobita.% 34^ CHAPTER XLI I. Sub-class 6, Poecilopoda. Sub-class 7, Podophthalmia ; Order i, Decapoda ; Order 2, Stomapoda, CHAITKR XLI 1 1. Sub-class 8. Kdriophthalmia; Order I, Cumatca- ; Order 2, Laemo- dipoda ; Order 3, Amphipoda ; Order 4, Nopoda. . CIIAPTKK XLIV. I >nisi,,n 2, I'raclKMjiiiM;.. Class 2, Arachnoidca. Sub-cl.. I'seudarachna ; Order i, I'antopoda: Order 2, Tardigrada. Sub-cla>s 2, Autarachna ; Oricr i, Acarina ; Order 2. neina ; Order 3, Arthrogai .... CHAPTER XLV. Class 3, Myriopoda; Order i. Chilognatlia . Order 3, Pauropoda CHAPTKR XLVI. ("l.is> 4. Insecta. Contents. x\ CHAPTER XLVII. PAUK. Sub-class I, Ametabolica; Order i, Mallophaga ; Order 2, Rhynchota ; Order 3, Thysanura ; Order 4, Thysanoptera ; Order 5, Euplexoptera ; 'Order 6, Orthoptera, ... CHAPTER XLVIII. Sub-class 2, Holometabola ; Order 7, Neuroptera; Order 8, Trichoptera ; Order 9, Strepsiptera ; Order 10, Aphaniptera : Order 1 1, Diptera ; Order 12, Lepidoptera ; Order 13, Coleop- tera ; Order 14, Hymenoptera, 405 INTRODUCTION TO ANIMAL MORPHOLOGY CHAPTER I. THE group of colloids called Albuminoids is the most complex and unstable series of bodies known to the chemist, and, as we are as yet ignorant of the nature of their apparently definite compounds, wre know nothing of their molecular constitution.* The phenomena of lifef are always associated with and manifested in a viscid, transparent, colourless, unstable albuminoid, whose two special properties are contractility and the power of converting other Carbon, Hydrogen, and Nitrogen compounds, into a material like itself. An ammoniacal solution of carmine stains it crimson. Of • Albumen is slightly acid in reaction, nnd forms compounds with Ba, Ca, I'b, K, Xa, &c., the la>t pair being probably the piotun < t Mulder, irtzcnbach regards its compound with Platino-cyanide of 1'otassium as being definite, but this is not confirmed by Diakonow's analysis. :i-e, the evolution of heat, and the assi- milaiiim of othci i.uw generally regarded as a mode of energy. B 2 Introduction to Animal Morphology. this body (Protoplasm or Bioplasm), white blood corpuscles, the yelk spheres of eggs, the contents of all growing cells, are examples. This substance shows three forms of motion under different conditions : — ist. When diluted with water, internal currents indicated by the circulation of enclosed particles, the outline of the mass being unaltered ; a more definite motion than the Brownian cyclosis of non-living fluids, and con- nected with chemical change. 2nd. Motion attended with change of form, irregular contractions of the whole mass, with temporary protrusions at one part (called pseudopodia),and recessions elsewhere ; thus locomotion can take place.* This form of motion maybe changed into the first by dilution, or the former into it by the action of a weak (2%} solution of NaCl. 3rd. Ciliary motion, f the protoplasm mass having on its surface constant hair-like processes of its own substance* (cilia), rapidly vibrating to and fro, either from within (as in spermatozoa) or from the outer layer of the mass. Long whip-like cilia are called flagclla, if sub-rigid, * Thus Amoebae wander in the waters wherein they live, and white blood corpuscles through the tissues of the bodies of higher animals, as Waller, Recklinghausen, and Cohnheim have described. Such cells are probably used as tissue-pabulum, as Joung noticed in connective tissue, Biesiadecki and Pagenstecher in epithelium, and Strieker, &c., in deve- loping embryos. t Gegeribaur regards ciliary motion as different from protoplasmic. \ Flagella have been seen changing into pseudopodia in Protomyxa (Haeckel) and in Myxomycetes (De Bary). The converse, or cilia changing into pseudopodia, occurs in Magosphaera (Haeckel). Cilia are never out- growths from a cell-wall, nor vacuolated nor granular. The granules observed by Alexander Stuart in Opisthobranchiata have not been con- firmed by others. Stuart has also described fine lines passing from the cilia to the cell nucleus, which by contraction moved the nucleus ! Pro- bably these are of the nature of the longitudinal striae found by Eberth in cells of ciliated epithelium. Introduction to Animal Morphology. 3 sfyh\ A flagellum originates as a process from the protoplasm mass, which first undulates, then becomes globular, and finally elongates (Engelmann}. In the formation of a row of cilia the undulating knob elongates into a ridge, which splits into parallel bands, rapidly changing into cilia. The rate of motion of cilia varies per minute from 720* to 480 (Engelmann}, 190 (Krausc\ 150-100 (Valentin}. The effects of reagents on protoplasmic motions are as follows: — i. Very weak alcohol, moderate heat,f and very diluted alkalies favour motions of the 2nd and 3rd form. 2. Ether, strong alcohol, chloroform, stop motions and render the mass granular. 3. Hydrocyanic acid arrests movements. 4. Vegetable alkaloids, as a rule, affect them but little. 5. Weak acetic acid causes irregular contractions, and finally checks motion. 6. Fresh water makes protoplasm masses spherical and develops the first form of motion.f 7- Carbonic acid de- stroys ciliary action (Kiihne}, but the access of air, if resorted to speedily, restores it. 8. Hydrogen checks ciliary action, but cilia thus stopped, if soon exposed to carbonic anhydride, are at first set in motion, and then finally stop (Engelmann}. 9. Cold checks amoeboid motion and rotation of the yelk- spheres, but (if not excessive) only retards, but does not arrest, growth or life-progress. 10. Weak induction currents accelerate amoeboid motions or produce contraction (Kiihne, Golubew} ; they also accelerate ciliary action (Kistiakewsky). Every variation of intensity of the current acts as an excitant, producing alternate contractions and relaxations. The closure of a constant current is a stronger stimulus than the opening. Strong currents stop all motions (Engelmann}. • I observed this rate in Planaria. t M .t accelerates the motions of cilia (TTWvr), and the loco- motion of corpuscle, which is greatest at ioo13 F. (Schultzc). Yelk masses contract and dilate at 90° 1 . chko). 1 13° F. coagulates • t The spherical state may be strong contraction (fCuhnc) • 4 Introduction to Ammal Morphology. Protoplasmic motion is sometimes attended with the evolution of gas.* In an atmosphere of Hydrogen and in an oxygenated solution of Haemoglobin, cilia move until the free oxygen is gone (Kiihne}, but under some conditions vibra- tions continue for a time without oxygen and without food, as if the mass had in itself a store of energy. The first stage of decomposition in protoplasm is indicated by the formation of granules, and the disen- gagement of fluid or gas making little clear spaces (vacuoles). On account of the constantly occurring partial contractions, homogeneous protoplasm masses are rarely of large size, except where feebly acted on by external stimuli ; usually they split into smaller particles. The contact of air, of oxygenated water, or other stimuli, seems to cause the surface to undergo chemical change ; it ceases to be protoplasm, and be- comes a limiting membrane,t through deficiencies in which protoplasmic threads may protrude (cilia r) As this material is constantly decomposing, new protoplasm needs to be as constantly forming, but its production is often irregular, and may proceed more actively in one spot than elsewhere. Such a spot is usually denser, and takes a deeper carmine dye than the rest of the mass, and is often sharply defined. It is called the nucleus, but it is not essential to growth,:}: « As in Arcella (Engeltnann). Protoplasm is normally neutral or slightly alkaline, but after exhaustion of motion it is faintly acid, and some- times weak alkalies revive for a time suspended motions. Possibly the contraction may be the result of the force evolved by oxidation. The contractions in a mass of protoplasm are always partial ; one side of a cilium, or one part of a mass contracts, and drags the rest after it. The evolution of light in animals is sometimes attended with the decomposition of proto- plasm into urate of ammonia. t Often porous. \ Many cells have no nuclei, as all Monera, Hydrodictyon (Alga), the segmentation spheres of ova, &c. Introduction to Animal Morphology. 5 nor has it any properties other than those of any other particle of vigorous protoplasm.* It may be granular, sometimes vacuolated (Rollett\ and often has a denser spot within it called the nucleolus. Each independent protoplasm mass is named a plastide ; a plastide with no nucleus is a cytode ; a nucleated plastide is a cell ; a naked cytode is called a gymnocytode ; one invested by a membrane is a lepocytode ; a naked cell is a gymnocyte (monoplast Sharpcy] ; one with a cell- membrane is a lepocyte (H&ckel}. The largest single cells are the myeloplaxes of growing bones and the muscle cells of Nematodes. Plastides multiply by gemmation, a bud arising as a small process from a parent cell, then enlarging and becoming detached ; by fission, each splitting into two or more nearly equal parts — in this process the nucleus may or may not take partf ; — by free cell formation, nuclei arising in a mass of protoplasm, and around each an area of protoplasm becoming isolated by the differentiation of cell-membranes — this occurs in developing embryos ; — by endogenous formation, as shown by Wcissmann in the eggs of Diptera, and by E. Van Bencden in the formation of the nucleus in Pseudofilaria of Gregarina gigantea.J It occurs pa- thologically in tumours. • Gegenbaur states that the nucleus differs from the surrounding pro- toplasm by being non-contractile, but Bottcher has demonstrated that a free nucleus or a nucleus removed from a cell contracts actively. Neumann describes the nucleus as showing motions in dying cells. Free nuclei may exist in masses of protoplasm. t Remak and // ccn cells divide while the nucleus was ad- herent to one side. The same has been noticed by many other observers. J The multiplication of cells in cartilage, some tumours or ova, is sometimes regarded as of this kind, but in reality it is more likely a con- cealed form of fission. The instance givrn by liuhl of the formation of 6 Introduction to Animal Morphology. In the simplest animals the protoplasm is homo- geneous, but in the more complex the homogeneous yelk protoplasm becomes differentiated in development, and thus we find cells whose contents vary. It has been supposed that protoplasm may be a compound albuminoid resolvable into its elements (myosin, neurin, &c.) in the course of growth ; at least two substances may be identified in it, one of which is active and dilates on dilution with water, while the other is not.* All cells are at first protoplasmic, whatever their ulti- mate contents may be, and protoplasm seems to possess the sum of the properties of the active cells (muscle and nerve) derived therefrom. The chief ma- terials derived from protoplasm in animals I have appended in a note.f To complete our introductory studies, it may be pus cells within epithelial is more probably a case of wandering (Stcudder and Volkmann}. * Heidenhain and Brucke suppose some cells to have a lacunary structure, with a basis of a more solid albuminoid and a more fluid proto- plasm contained therein. f Protoplasm derivates are of two kinds, Nitrogenous and non- Nitrogenous. The first are : — Albumen, CTsHnoNisSO^HgO, soluble in water, not precipitable by alkaline carbonates nor sodium chloride, but is by heat. It varies in pro- perty according to its source. Seralbumen (from blood serum) is not coagulated slowly by ether. Ovalbumen is, and when injected under the skin of an animal, speedily appears in the urine, which the other does not. The ovalbumen of Raptorial birds is with difficulty coagulated by heat or acid (Fr&my and Valenciennes). In some swimming birds and reptiles the ovalbumen, when diluted, is slowly coagulated by heat. Globulin is insoluble in water, soluble in sodium chloride, coagulated by heat, changed into acid albumen or syntonin by HC1 ; varieties of it are Vitcllin, not precipitable by NaCl, when added in substance to saturate its solution ; Myosin is thus precipitable. Paraglobulin or Fibrinoplastin^ when combined with another albuminoid fibrinogen, forms fibrin, as in the coagulation of blood. Introduction to Animal Morphology. 7 observed that the following inorganic materials are found in animal bodies : — Oxygen in all the fluids, Hydrogen in the intestinal gases of vertebrates with H2S, Carbonic acid in all tissues, Calcium Fluoride in bones and corals, Magnesium Fluoride in some corals, Water in all tissues from enamel which contains 0*2 per cent, to the connective tissues of jelly fishes with 99-5 per cent., Sodium Chloride also in all Peptone is a dialysable form of albumen formed in the process of digestion, not precipitable by heat, dilute acids, nor alkalies. Fibrin is insoluble in water or NaCl, swells in dilute acids, and this softened form is coagulated on boiling ; found in blood. Casein is probably a mixture of Alb. with a non-albuminous substance, which is separated from it in digestion as a dyspeptone ; soluble in alkalies, coagulable by mineral acids, precipitable by lime and magnesia salts, in- soluble in water or NaCl. Keratin, the material of the cell-wall in epithelium, resembles casein in many of its properties ; it may be a compound of several albumen- derivates. Gerhardt regards these as similar in composition, but differing in molecular arrangement. Lebonte and others regard them as each com- posed of several radicles united in different proportions. From these are derivable the following nitrogenous non-albuminoids : — Protagon, present in nerve tissue. A mixture of Cerebrin Cn / OiCieHaiO with Lecithin, a body derived from ) OiCisHaaO Glycerin C.,11'"* OiPO /OH \ OC,H4 Leudn CGHn (NH2) O2 present in the tissues of Arthropods, in pancreatic juice, &c. Ty rosin C^HnNOa found with Leucin and in Cochineal. Kreatin CiHaNaOjH^O in smooth muscle, and with Kreatinin (C