You are on page 1of 956
















C. V.


LOUIS. MO. and





will Your opinion of this book appreciated when your review of


has been completed.







Professor of Zoology, Agricultural and Mechanical College of Texas, Formerly Professor of Zoology, Baylor University


With 445 Text





xin^'L-v ^
Iffai:i'Tfii., ^F^^'




C. V.


Copyright, 1938, 1947,

By The

C. V.

Mosby Company

(All rights reserved)

Printed in U.



Press of


C. V.

Mosby Company

St. Loviis


PROFESSOR FRANK A friend and an




to the student


The present edition represents a revision
of certain parts of several chapters, such as those dealing with Annelida, Genetics, Eugenics, Internal Regulation and Endocrines, Physiology, and Phylogenetic A brief section on Mammalian Development Relations of Animals. has been added. Numerous minor corrections or improvements have


made throughout.

Several illustrations have been added and

others improved.

In addition to the acknowledgements included in the preface to the first edition, the author wishes to acknowledge the help of Dr. Kelshaw Bonahm of the Fish and Game Department, Agricultural and Mechanical College of Texas, on the chapter dealing with Pisces. At this point recognition is made of assistance given by Mr. Gordon Gunter in revising the list of animals of the Texas Gulf Coast in the chapter on ]\Iarine Zoology. The author is also indebted to Dr. Fred L. Kohlruss, Biology Department, University of Portland (Oregon), Many valued suggestions have for numerous useful suggestions. been received from individuals in a number of other institutions where
the first edition has been in use.

The author's indebtedness and


expressed to Mr. Phil T. Williams who has furnished a number of the new illustrations. Finally, appreciation is expressed to Agricultural and Mechanical College of Texas for cois


operation in numerous ways to assist in making this revision.

George E. Potter.
College Station, Texas

The important problems of life are common to all animals (including man) as well as to plants. It should be the purpose of a textbook in general zoology to present the animal kingdom in a logical and natural way and at the same time carry the interpretation of the facts
in terms of the principles involved. strike the ideal balance
It is exceedingly difficult to

between the necessity of presenting

"type" material
knowledge of

in order that the student will have the


classification, structure, function,


and organography

to appreciate the discussion of principles,

and the

opposite temptation to go into endless discussions of theories and

comprehension of which is unquestionably beyond the who has not become grounded in fundamentals of animal make-up. Of course it is usually hoped that the laboratory division of the course wiU supply this needed foundation. It seems reasonable that the ultimate aim of the teacher of introductory zoology should be to bring the student to a fundamental and well-grounded understanding of the principles involved in all of the living processes. It is extremely difficult to skim this information from the top of the entire body of zoological knoAvledge, as one can skim cream from a crock of milk, and hand-feed it to the waiting student mind. Apparently there must be a certain amount of personal acquisition of the principles of the subject through attaining a clear-cut knowledge of tlie complete biology of a series of representative animals. Each of these representatives, since it is a living organism, demonstrates certain of these principles. In order to bring this out there must be a rather close coordination between the studies of the laboratory and

capacity of the student

the presentation of principles

by the textbook. Based on a recognition of the above-mentioned situation and


on the realization that the majority of students taking elementary zoology plan to go no further in the field, the author has attempted to strike a workable combination of the two schools of teaching and still cover the fundamental knowledge of the subject. There has been
a definite effort to lead the student to think of biology as related to humankind and to himself. It is hoped that the book will overlap
the laboratory studies just far enough to

the student out of the

laboratory into his o^vn correct interpretation of the facts discovered

It is of course

assumed that the teacher

will naturally elabo-


in the course. The upon particular phases of the topics anticipation of this and limitations of space have reduced the volume

taken up

of detailed information included.


animals from west of the Mississippi River are featured in book. There has been no attempt to limit the scope of the work

to this region,

but since many southwestern and western forms are available and serve as very good illustrative material, they have been It is hoped this wiU make the book more useful and meanutilized.
ingful to students in these regions, as well as



The introduction

of chapters on



of Radiation,

Wildlife Conservation

a slight

Animal Anomalies, Animal ReMarine Zoologij, and departure from the usual textbook

but each of these seems to the author to have enough of special value and current interest to warrant presentation. The chapIts ters on Regulatory Glands, Animal Distribution, The Animal and Environment, Animal Parasitism, Comparative Emhrijology, Animal
Behavior, and Paleontology are also presented with the feeling that they are of exceptional general interest to all students, as well as

being thoroughly zoological.

The arrangement of the chapters on animal groups has been somewhat in the order of complexity and systematic relationships. The
chapters are written in such a way, however, that this order may be modified in any manner to suit the teacher. The chapters dealing with typical Protozoa, Hydra, Planaria, Annelida, Arthropoda, and Amphibia are somewhat amplified and include more detail because

they are so often chosen as typical groups for study. Throughout many the book the genus and species names have been italicized, and


of structures

and functions have


been italicized the


time they occur.

The author


of several teachers

indebted and extremely grateful for the cooperation and specialists who have contributed manuscript

For this service acknowledgment is made Teague Self, University of Oklahoma, Annelida; Elmer P. to: J. Cheatum, Southern Methodist University, Mollusca, and assisted with Marine Zoology; Vasco M. Tanner, Brigham Young University, Arthropoda; Mary Fielding, Public Schools, Waco, Texas, collaborafor chapters in their fields.

Newman, Baylor University, collaboraOttys Sanders, Southwestern Biological Supply Co., tion on Pisces;
tion on Elasmohranchii; Rose




Amphibia, and assisted with Marine Zoology; Leo T. Murray, Baylor University, assisted by James E. Blaylock, Ranger Junior College, Reptilia; Helen Joe Talley, University of Oklahoma, collaboration on Regulatory Glands; T. C. Byerly, United States Bureau of Animal Industries, Ayiimal Regeneration; Titus C. Evans, University of Iowa, Medical College, Biological Effects of Radiation; Willis Hewatt, Texas Christian University, A^iimal Distrilution, and assisted with

Marine Zoology; A. 0. Weese, University of Oklahoma, The Animal and Its Environment ; Sewell H. Hopkins, Texas Agricultural and Mechanical College, Animal Parasitism ; J. G. Burr, Texas Game, Fish, and Oyster Commission, marine data; Walter P. Taylor, Texas Cooperative Wildlife Service and United States Bureau of Biological

Conservation; A. Richards, University of Okla-

homa, Comparative Embryology ; Frank G. Brooks, Cornell College, Genetics and Eugenics; Iva Cox Gardner, Baylor University, Animal Behavior; W. M. Winton, Texas Christian University, Paleontology

To Mr. Ivan Summers goes immeasurable

credit for the excellent

work he has put

into this edition.

Dr. Titus Evans, of the

University of Iowa, Medical College, has also been of great service Mrs. Ruth M. with his excellent talent in creating illustrations.
Sanders, Miss Joanne Moore, and Mr.

Edward O'Malley have


The drawings used in XLII on Genetics and Eugenics were made by Miss Betty Chapter R. Smith of Cornell College. The author is grateful to all of these
by contributing certain illustrations.
individuals for their valuable services.

The author wishes


acknowledge also the friendly and helpful

advice which has been offered by Professors D. B. Casteel, T. S. Painter, and E. J. Lund of the University of Texas, and Professor

Asa Chandler of Rice Institute. Finally, appreciation is expressed to Baylor University for the cooperation which has made the writing of this book possible. George E. Potter.
Waco, Texas.


Introduction The Biological Point of View, 17; Science and the Scientific Method, 18; Zoologj-, a Biological Science, 19; The Subdivisions of Zoologj-, 19; Classification of the Animal Kingdom, 25; Vital Eelations of Animals and Plants, 27; Attributes of Life, 30; Balance in Nature, 31; Zoology as Belated to Man, 33; Agriculture and Zoology, 34; Fisheries and the Application of Zoologj-, 34.

CHAPTEE ___________________


History op Zoology





Protoplasm and the Cell


Living Matter, or Protoplasm, 49; The Cell Principle, 49; General Characteristics of Protoplasm and the Material of the Cell, 53;

Fundamental Properties or Activities of Protoplasm, 54; Physical Nature of Protoplasm, 55; Chemical Nature of Protoplasm, 56; Structure of a Typical Animal Cell, 58; Cell Division, 61.

Phylum Protozoa




Characteristics, 65; Classification, 65; Colonial Protozoa, 75; Tropisms

and Animal Reaction, 77; Economic Relations of Protozoa,

Euglena op Class Mastigophora



Habitat and Characteristics, 81; Structure, 81* Food and Assimilation, 81; Respiration and Excretion, 83; Reproduction and Life Cycle, 83; Behavior, 84; Locomotion and Flagellar Movement, 84.

Amoeba op

Ck<vss Sarcodina




and Habitat, 85; Structure, 86; Metabolism, 86; Reproduction and Life Cycle, 89; Behavior, 91; Amoeboid Movement and Locomotion, 91.

Paramecium of Class Inpusoria



and Habitat, 93 Structure, 93 Metabolism, 95 Reproduction and Life History, 96; Behavior, 100; Locomotion, 102.





Metazoan Organization

__________----CHAPTER IX




General Characteristics, 103; Cellular Differentiation, 104; Cellular Organization, 105; Development of Sexual Reproduction, 111; Meta-

zoan and Ontogeny, 113.



_____-___-------Protozoa, 129.


Sponges, 119; Classification, 120; Fresh-Water Sponges, 121; The Simple Sponge, 122; Habitat and Behavior, 122; External Anatomy, 123; Internal Anatomy, 124; Metabolism, 127; Reproduction and Life History, 127; Economic Relations, 129; Phylogenetic Advances
of Sponges

When Compared With

Phylum Coelenterata
Classification of the




Phylum, 131 Hydra, 144 Habitat and Behavior, 144; External Anatomy, 146; Internal Anatomy, 148; Metabolism, 151; The Nervous System and Nervous Conduction, 153; Reproduction and Life Cycle, 153; Regeneration, 156; Economic Relations of the Phylum, 156 Phylogenetic Advances of Coelenterates, 156.

Phylum Ctenophora

______---------____---------CHAPTER XII



Habitat and Behavior, 157; Anatomy, 157.

Phylum Platyhelminthes


160; Planaria, 163; Habitat and Behavior, 163; External Anatomy, 165; Internal Anatomy, 165; Metabolism, 170; Reproduction and Life History, 170; Regeneration, 173; Economic Relations of the Phylum, 174; Phylogenetic Advances of Platyhelminthes,







Classification, 175; Asearis,

A Representative Roundworm, 179; Habiand Behavior, 179; External Anatomy, 179; Internal Anatomy, 181; Reproduction and the Life Cycle, 183; Relations to Man, 183.

Molluscoida, Trochelminthes, and Chaetoonatha



Molluscoida, 184; Bugula, 184; Trochelminthes, 188.

Phylum Annelida (By

J. Teague Self) Earthworm, 199; Internal Anatomy, 201; Reproductive Organs, 202;



Digestive System, 203; Circulatory System, 205; Respiratory System,



206; Excretory System, 206; The Nervous System, 208; Eeproduction, 209; Eegeneration, 212; Importance of Annelids to Man and Other Animals, 215; Phylogenetic Advances of Annelida, 216.


Phylum Echinodermata




Habitat and Behavior, 226; External Anatomy, 226; Internal Anatomy, 227; Reproduction and Life Cycle, 234; Regeneration and Autotomy, 234;
Classification, 217; Starfish of Class Asteroidea, 226;

Economic Relations, 235.

Phylum Mollusca (By Elmer

P. Cheatum)



General Characters, 236; The Snail, 237; Habitat and Behavior, 237; External Anatomy, 239; Internal Morphology, 243; Respiration, 244; Circulation, 244 Nervous System, 245 ; Excretory, 245 Reproduction and Life Cycle, 245; Fresh-Water Clams, 248; Habitat and Behavior, 248; External Features, 249; Internal Anatomy, 250;

Digestion, 250;

Respiration, 251; Circulation, 252; Nervous System and Sense Organs, 252 Excretion, 253 Reproduction and Life Cycle, 253; Economic Relations of the Phylum, 255; Classification, 256.

Phylum Arthropoda

; ;



Crayfish of Class Crustacea, 266 Habitat and 263 Behavior, 267; External Structure, 268; Internal Structure, 271; Metabolism, 278; Reproduction, 278; Regeneration and Autotomy,



Economic Relations, 281

Characterization of Other Crustacea,


Recapitulation Theory, 284;

Phylogenetic Advances of Arthro-

poda, 286.

Phylum Arthropoda


(By Vasco M. Tanner)



Onychophora and Myriapoda,



Phylum Arthropoda
Arachnida, 292;

(By Vasco M. Tanner)



Spiders, 292;

of the Arachnida, 295.

Phylum Arthropoda

(By Vasco M. Taimer)



Head, 301; Thorax, 305; Abdomen, 307; Body Wall, 308; Metamorphosis, 308; Classification, 309; Hemimetabolous Insects With Incomplete Metamorphosis, 320; Holometabolous Insects With Complete Metamorphosis, 321; Other Orders, 333 Social Life Among the Insects, 334 Guests, 339 Economic Relations, 340; Useful Insects, 341.
Class Insecta, 300;
Insect Characteristics, 301;



Representative Insects (By Vasco M. Tanner) The Locust, 343; The June Bug, 354; The Honey Bee, 357.




Phylum Chordata

_______-____-_--CHAPTER XXIV



Phylogenetic Advances Classification, 361; 360; of Chordata, 362; Protochordata (Lower Chordates), 362; Subphylum Hemichordata, 362; Subphylum Urochorda, Molgula, 365; Subphylum
Cephalochorda, Amphioxus, 368.

The Vertebrate Animal:
Classification, 410.

Subphylum Vertebrata




; ;



Economic Relations of the Class, 413 The Lamprey, 413; Habitat, 413; Habits and Behavior, 415; External Structure,

415; Internal Structure, 415.





Economic Relations of the Class, 425; The Spiny Dogfish, 426; External Features, 426; Muscular System, 427; Skeletal
Classification, 422;

tory System, 435;
Squalus, 439.

System, 427; Digestive System, 430; Circulatory System, 431; RespiraNervous System, 435; Urinogenital System, 437; The Eonnethead Shark, Reniceps (Sphyrna) Tiburo Compared to


True Fish





Bony Fish

445; Economic Relations of the Class, 455; Typical Yellow Bullhead and Some Comparisons With Yellow

Perch, 457; External Features, 457; Digestive System and Digestion, 458; Circulatory System and Circulation, 459; Respiratory System,

463; Excretory Organs, 464; Skeletal System, 464; Muscular System and Locomotion, 467 Nervous System, 469 Reproduction and the


Life History, 470.

Class Amphibia

(By Ottys Sanders)

in the


United (Tailed Amphibians), 484; States, 484; Order Caudata (Urodela) Order Salientia (Anura) (Tailless Amphibians), 485; Economic Importance, 486; Necturus Maculosus, the Mud Puppy, 487; Food and Digestive System, 489; Circulatory System, 490; Respiratory System and Breathing, 491; Urinogenital System, 492; Skeletal System,
Classification, 482;


List of Families of the

494; Muscular System, 496; The Nervous System and Sense Organs, 497; The Bullfrog, 497; Habitat, 497; External Structure, 497; Digestive System and Digestion, 499; Circulatory System, 502; Respiratory Organs and Eespiration, 514; Excretory System and Excretion,




System, 517;

Muscular System, 523; Nervous

The Sense Organs, 528; Reproductive Organs, 531; Embryology, 532; The Toad, 538; Habitat, 538; External Features,


Internal Structure, 541; Respiratory and Digestive Organs, Urinogenital Organs, 541 ; Blood Vascular System, 542 ; Skeleton EmNervous System and Sense Organs, 543 Muscles, 543


bryology, 544.

Reptilia (By Leo T. Murray and James E. Blaylock) Fossil Reptiles, 546; Classification of Living Reptiles, 547; Class Reptilia, 547; Order Testudinata (Chelonia), 548; Order Squamata, 551; Order Rhincocephalia, 560; Order Crocodilia, 560; The Horned Lizard, 561; Habits and Behavior, 561; External Structure, 562;
Digestive System, 563; Respiratory System, 566; The Circulatory Sys-



tem, 566; The Urinogenital System, 571; The Nervous System, 573; The Skeletal System, 574; Muscular System, 576; The Turtle, 577; Habits and Behavior, 577; External Structure, 578; Digestive System,

578; Respiratory System, 579; Circulatory System, 580; Urinogenital System, 581; The Nervous System, 581; The Skeleton, 581; The

Muscular System, 581.


Classification, 584;



Economic Relations, 596; Domestic Chicken, 598;

Habits and Behavior, 598; External Structure, 599; Digestive System, 601; Respiratory System, 603; Circulatory System, 604; Excretory System, 606; Nervous System, 607; Skeletal System, 609; Muscular System, 612; Reproduction and Life History, 613.


Classification, 616;

Economic Relations, 637; The Cat,
Structure, 639;




Mammal, 639; External

Skeleton, 641;

The Digestive System, 646;




Respiratory System, 649; Nervous System, System, 650; Reproduction and Life History, 652.



Animal Anomalies




Harelip and Cleft Palate, 656; Diaphragmatic Hernia (Open Diaphragm), 657; Polydactylism (Extra Digits), 659; Conjoined Twins, 659; Hermaphroditism, 663; Cardiac Anomalies, 664; Abnormalities of Brain and Sense Organs, 664.



The Endocrine Glands and Their Functions


„.„„ PAGE

The Thyroid Gland, 667; The Parathyroid Glands, 670; The Suprarenal Bodies, 671; The Pituitary Gland, 672; The Thymus Gland, 675; The Gonads and Sex Hormones, 675; The Pancreas, 677.

Regeneration (By

T. C. Byerly)



Introduction, 681; Regenerative Capacity, 681; Protozoa, 681; Porif-










688; 692;












Mammalia, 690;



Adaptability and



Biological Effects of Radiations

(By Titus C. Evans) The Structure of the Atom, 697; Biological Effects of Sunlight, 700; Infrared Radiation, 700 High Frequency Oscillations, 701 Effects of Roentgen Radiation, 702 Ultraviolet Radiation, 701 The Funda;














Radium, 708;


Other Radiations, 709;


Animal Distribution (By



Life Regions and Zones of the Earth, 711; Migration of Animals, 716; Means of Dispersal and Barriers, 717; Effects of Man Upon
Distribution, 718.

The Animal and
Its Environment (By A. 0. Weese) The Principal Biotic Formations, 724; Adaptation, 727; Succession, 727; Animal Populations, 730; Seasonal Changes, 733; Summary, 733.



Animal Parasitism (By
of Parasitism, 736;

Sewell H. Hopkins)



Social Relations of Animals, 735; Origin of Parasitism, 736; Degrees

The Successful Parasite, 737; Means of Infecand Transmission, 740; Parasitism and Host Specificity, 741; Parasites and the Groups in the Animal Kingdom, 742; Some Repre-

sentative Parasites, 747.

Marine Zoology







Wildlife Conservation The Abundance of Animals, 789; The fensibles, 792; The

(By Walter

P. Taylor)



Wild Animals, 784; The Natural Range of Wild Coming of Civilization and a Declaration of IndeProblem of Restoration, 794.

Comparative Embryology (By A. Richards)



MAMMALLA.N DEVELOPMENT Organs and Systems, 817.

__________--_-CHAPTER XLIII



Genetics and Eugenics (By Frank G. Brooks)



a Great Discovery, 821; Mendel's Law, 821; Derivatives of Mendel's Law, 823; The Physical Basis, 824; Plotting Crosses, 825; Complications of Mendelian Inheritance, 827; Inheritance of Sex, 831; Linkage, 832; Sex Linkage, 832; Crossing Over, 834;




Mutations, 836;

Human Heredity, 836; Matings Among Defectives, 839; The Differential Birth Rate, 839; Family Size in Eugenic Groups, 841; Family Size in Dysgenic Groups, 842; What Can Be

Done? 844; Some Eugenic Measures,

Animal Behavior (By Ina Cox Gardner)



Reflex Tropistic Behavior, 849 846 Chain Reflex Behavior, 851; Habitual Behavior, 852.



Paleontology (By W. M. Winton)



Phylogenetic Relations op Animal Groups and the Theory of Evolution Colony Formation in Certain Protozoa, 864; Development of the Gastrula, 865; Trochophore Larva, 865; Peripatus and the Wormlike Ancestry of Arthropoda, 865; Eehinoderms and Their Larval Relations, 866; Ancestry of the Vertebrates, 866; Basis for the Theory of Evolution, 870; Darwin and Studies of Evolution, 885,




but she hides the truth. life. such a study deals with the forms and phenomena exhibited by both. on land. he soon realizes the certain fundamental features He common to the structure and function of all living forms. as one striving to get a glimpse the mysteries of life. like the chemist or physicist. largelj^ into tAvo general categories.TEXTBOOK OF ZOOLOGY CHAPTER I INTRODUCTION the air. discourse. A biological concept may rest upon observations. soon recognizes the oneness of all life. her forces are in constant The biologist looks in upon himself into as a seeker after truth. a logos. or in the sea. and himself as a part of one great organic system. but the biologist. which may be changed from day to day by the discovery of new facts. hios. As he succeeds existence of obtaining these glimpses. is justified in holding to a theory or hypothesis as long as for further investigation. In whichever direction we turn or wherever we go. 17 it provides a true working basis . The Biological Point Nature is of . ask questions. What is the many different kinds? things? What is life? Such they come? man to answer these questions and solve the problems of the origin of life has given us the field of study is and nature Biology known as biology. whether in we are surrounded by living creatures. Their very presence presents problems and fills us with curiosity. and and is the name universally applied to the study Since living things fall of living organisms and life processes. plants and animals. We From whence do source of their energy? Why are there so What is our relation to other living questions and endless numbers of similar ones kindle the interest The constant endeavor on the part of of every thinking person. View ever inviting investigation operation about us. word derived from two Greek words. each unit of which has some relation to the whole.

to conclude. These fundamental facts are then classified with respect to previously established facts. the qualities of the thing observed are compared and the essential or fundamental ones are separated from the nonessential. and the scientist respects truth while to little him tradi- tion or mere opinion counts for is as such. and. When the conclusions have been further verified by repeated examination. Facts are indispensable building stones of science. The method is simplicity itself. After the facts are thus established. Facts must be gleaned from careful observations and experiments which have been rigidly checked and will yield identi- cal results Science lays with frequent repetition and by numerous observers. The scientist is not a mato coordinate. That was his way of saying that scientific knowledge is simply an extension and organization of the knowledge based upon common observation and experiment conscience given cerning the facts of nature. Our powers of observation have been increased by the development of the microscope and many other instruments. to experiment. Complete and accurate observations of the objects or phenomena under investigation followed by honest interpretation. It is deduction may be made of the principles involved. rei^resent truth. Experiments are devised to bring to light the features not readily revealed by steps are often tedious direct observation. although the and much involved. by this systematic method of investigation that science has been established. to observe. to deduce.18 TEXTBOOK OF ZOOLOGY Science and the Scientific Method "Trained and organized common sense" was the definition of by Thomas H. are the aims of the scientist. and experimentation. from thin air as Thurston gician Avho can draw his conclusions seemed to produce almost anything imaginable. as it first develops from preliminary observation and experiment. Huxley. on the basis of the relationships of qualities. the hypothesis becomes a theory. . its foundation on accurate observations and depends Established facts on the ability of the senses to reveal the truth. This requires accurate logic and keen judgment. is known as a hypothesis. There nothing mysterious in the scientific method. An idea. to identify by comparison. an eminent English biologist who lived from 1825 to 1895. observation.

which are concerned with living organisms. The expression constitute the science of biology. but up to the present time our scientific However. discourse on. Science is. and a conclusion once In fixed and may not be withdrawn for any reason. A person who study of zoology is known as a zoologist. Scientific hypotheses are frequently shown to be untenable. always subject to modification or even science conclusions are abandonment as investigation continues. increasing body of knowledge which is ever becoming more thoroughly established. is it is of Zoology only one of the divisions of the general field such a broad field in itself that it is neces- sary to subdivide It it into several divisions for convenience in study. The Subdivisions Although zoology of biological science. and logos. not relative. zoology. Zoology. a Biological Science The name. are conveniently divided into two groups: the physical sciences. and astronomy. evidence is produced to show the absolute fallacy of a so-called principle. refers to the study or science of The natural sciences. and the biological sciences. theories are occasionally found fallacious and discarded. There erroneous popular impression that zoologists were was at one time an simply "bug-hunters. geology. which deal with nonliving bodies. and related subjects were studied under the head of natu- . sciences. at any time sufficient principles have remained valid. the scientist will put aside sentiment and prejudice and accept the results of repeated investigation. a chajiging. as distinguished from the social animals. physics. botany together animal hiologij is often used as a synonym for zoology." This conception of the field has been greatly expanded until now it is considered one of the valuable and serious specializes in the fields of science. animal. theory and gives restilts of many drawn people truth is numerous investigators over a long period of time. which is derived from the Greek words zoos. therefore.INTRODUCTION There is 19 always a considerable volume of evidence which supports the Finally. the theory advances to a principle or law after it has been so thoroughly and critically tried as to be generally accepted and assumed This process requires the accumulation of the combined as a truth. all indication that it is a true statement. To is absolute. has been a relatively short time since all of the known biology. such as botany and Zoology and zoology. such as chemistry.

Its organization as a special field of Fig. The latter study comprises a compara- . further divided into several branches.20 ral history. includes all may be studied of form and structure of bodies by dissecting them. Gross anatomy. to such TEXTBOOK OF ZOOLOGY But now the subject matter of zoology alone has grown magnitude that it has become necessary to divide it into numerous special fields. which literally means cutting up. is usually separated from comparative anatomy. —Divisions of study in the field of biology. that It is now A. 1.) curred at about the beginning of the nineteenth century. These subdivisions may be summarized as follows 1. : Morphology is the study of the form and structure of the study oc- bodies of animals. which is one of the fundamental subjects of study in the preparation of the medical student. (Modified from several authors. Human anatomy.

B. there are large numbers of It can readily be seen that a system for putting extinct forms. as usually studied. is a study of the microscopic structure of the various parts of the animal body. Vesalius. we will learn. the store is divided into general departments. 1769-1832. finally. Besides these. Instead of this. and relationship of parts of the digestive system of the cat would be a good example of anatomical study. are the units of structure of all living matter. The dissection. and men's underwear for the toothbrush the its floors at customer desired. Histology or Microscopic Anatomy. size. children's toys. and . and the goods are completely classified within these departments. and study of the parts.000. includes not only the morphology of the cell but a great deal of the physiology in addition. This field of study has yielded many fundamental concepts of the factors involved in the living process.000 to well over a million. observation. The number of described species of animals as given by diftion or orderly ferent authorities ranges from 840. 131-201.D. on random. Cytology. these large numbers of different kinds of animals into a known order is one of the first prerequisites for dealing with them. go out and buy the thousands of different kinds of articles that are handled by a department store. the department store of the same reasons. 2. On a much is systematized for some One can see that it would be next to impossible to do business if a company were to provide a large floor space. To get the toothbrush. Few customers would return a second time if they had to wait hours while the clerk hunted among ladies' shoes. color. One well-known writer says there are probably no less than 2. C. with the human anatomy. Cytology is the The histologist studies the relationship and arrangement of the cells as they cooper- study of the minute structure of the cells which. the customer can be directed to the proper department and counter. 1514-1564. Cuvier. ate to comprise the substance of the organism. and just throw all of them smaller scale. Taxonomy is the subdivision which deals with the classifica- arrangement of organisms according to their natuThis field is often spoken of as systematic zoolral relationships. where kind. ogy.INTRODUCTION tive 21 in turn are study of the form and structure of the other animals and these compared.000 species of living animals. A. Galen. form.

a consideration of metabolism. sensitivity. Each of these sions is known as a phylum and is divided into classes. Such a name is composed of the genus name and species name. Taking the classification of man as an example we have: Phylum rChordata Subphylum :Vertebrata Class rMammalia Order :Primates Family :Hominidae Genus :Homo Species :sapiens The scientific name of man is written. from embryological similarities. each family into genera. is the same in because Latin a constant Russian as 3. functions.22 TEXTBOOK OF ZOOLOGY In this price are all orderly arranged. and followed by the name of the person who wrote the first authoritative description of the particular species. reproduction. The names are is common vernacular and almost universal language. each is class divided into orders. from facts of distribution. This always gives a it double name to a kind of animal. it is in English. excretion. muscular contraction. way the large unwieldy number of different kinds of articles become simply managed. glandular secretion. A group in which the members are very closely related is likely to be comparatively small. is Physiology the study of the functions of the various parts It involves of the organism as well as its living process as a whole. and each genus into species. and adaptation. These groups are ranked together according to evident relationships. nervous activity. many special such digestion. growth. and . and for that reason in Latin instead of is the hinomial system of nomenclature. In this field as is included the study of circulation. respiration. This system was originated by Linnaeus. Zoologists recognize a number of large divisions of the animal kingdivi- dom based on certain general characteristics. The relationships of animals are discovered from similarity of structure. but the Latinized form Homo sapiens Linn. Homo sapiens Linnaeus. and many other comparisons. each order into families. The common names would be almost certain to vary with each different language.

4. and anatomv on the other. through the numerous cell divisions. organization. others are physiological.INTRODUCTION others. physiology. Some of the factors which control this distribution of characteristics are morphological in their nature. is a study of the origin and development of the usually involves the changes occurring in the organism from the time of fertilization by the union of two cells. This field of study could not develop until the sciences of physics and chemistry came forward during the nineteenth century. like morphology. Pathology is diseases in the different groups of animals. is the division which deals with the study of variaresemblances. Genetics independently of one another. This field has advanced rapidly during the last seventy years. however. The development of the individual may be referred to as ontogeny. E. and their inheritance from one generation to tions. changes. and a study of pathol- ogy is likely to involve certain forms from all groups of animals. the next or from parent to offspring. genetics. Phylogeny is a study of the origin of organisms. 5. It is really the science of disease There is a fundamental similarity in in all of its manifestations. zoogeography. . growth. von Baer before the middle of the nineteenth century. and relationships It is of the based on the results of studies of morphology. Physiology depends upon an understanding of physics and chemistry on one hand. Embryology It mdividual. Physiology. The beginning of this study dates back to the work of K. In recent years the field of experimental embryology has developed rapidly. and differentiation leading to the adult conThis process includes both morphological and physiological dition. to as Much of the present study referred is cytology is physiological. 7. the study of the abnormal structures and abnormal functioning of life processes. remained in a crude state long after morphology was fairly well developed. Fairly definite laws governing this inheritance of qualities have been established by the geneticists. one derived from each parent. bringing about a variety of combinations in the progeny. old branch of zoology. 23 in the developing Many of the processes which occur eman bryo are also included here. and different groups and races paleontology. The characteristic features of an animal or plant may be transmitted to the offspring somewhat 6. embryology.

Oceans. or the remoteness of the place of origin of the species. The point of origin of the group may be cut off from other favorable regions by unsurmountable obstacles. Zoogeography or geographical distribution of animals is condis- cerned with the extent of the regions over Avhich species are tributed and the association of species in individual regions. forests. all become a part of an ecological study. found The English sparrow. This branch of the field has become prominent in comparatively recent of the environment. to mating. perhaps because of the topography of the region.24 8. to light. geological history. In con- some respects this field is closely related to ecology. to enemies. which originated in Europe. and many other facThis study usually tors. in 10. have been discovered by studies of the concerning the facts and the fate of extinct species has been learned through this field of study. TEXTBOOK OF ZOOLOGY Ecology is a study of the relation of the organism to its ento bring vironment. lines of descent of animals fossils. to food. It is affecting their distribution. and land are barriers to different types of animals. and in relatively few years it became a dominant bird. medium 3"ears. was not America until after it was introduced by man. Parts of many of the ancient animals are embedded and preserved in the sedimentary rocks. Paleozoology is a study of the animals of the past as they are presented by their fossil remains. draws somewhat upon a knowledge of all branches of biology. The failure of a species to it occupy its a suitable region usually means that has been unable to reach that region. to competition. mountains. cerned with the regions in which species exist and with the factors The regional distribution of an animal group is limited in part by the extent and relations of favorable environmental conditions. The relative age of the fossils is determined from the depth of Many of the probable the rock strata in which they are found. Conditions which prevent dispersal of animals from one area to another are deserts known all as barriers. Many made by animals ism to the adjustments in structure and function have been them into harmony with the conditions in Such conditions as the relation of the organwhich it lives to temperature. 9. Even a slight difference in the salinity or acidity of the water becomes a barrier to many aquatic animals. Paleozoology is ordi- Much . but no species occupies all of the regions where environment would permit.

cells. 2. kinds has been devised. the many-celled animals.000 species have been to 10.000 To date. divided into two subkingdoms: Protozoa. The principal groups subordinate to the class are order. and the geologist uses it in determining the relative ages of the rock strata which compose the crust of the earth.000.000. The secondary groups are phyla. The estimated number of kinds is all the way from 1. and they in turn are divided into classes. scribed briefly: Phylum Protozoa. —Individuals consist either of a single of cell or of aggregates of by each which are performed all the essential . Classification of the Animal Kingdom Very few people realize how many different kinds of animals there are and how greatly they vary in size.000. different forms of animals may a scheme of grouping related The entire kingdom or all is single-celled animals. approximately 840. In order that be known and definitely recognized.INTRODUCTION 25 narily studied under the head of geology. and Metazoa. — Phylum relations in the animal kingdom. ajid habits of life. The principal phyla are listed and defamily. genus. and species. named and described. Cbordata (Frog) /^rthropoda (Crayfbh) MoUusca (Snail) Annelida {Lumbricus) Echinodermata {Starfish) Bryo^oa (Bugulaj Rotifera NemathMnthes {Ascahs) Platyhelminthes (Tapeworm) Ctenophcra (Be roe) sea walnut Coelenterafca Fori f era (Hydra) (5cypha) Protozoa (Paramecium) ANIMAL KINGDOM Fig. structure.

Phylum Porifera (Sponges). symmetrical. There are about 4. The hookworm. Less than one hundred species are known. ascaris. Phylum Ctenophora (Sea "Walnuts delicate. TEXTBOOK OF ZOOLOGY aquatic in habit. nemato cysts. and the body wall is per- forated by species. Phylum Annelida (Jointed worms). They live in marine waters.y. and tentacles provided with stinging bodies.000 are known. triploblastic Phylum Nemathelminthes (Threadworms or Roundworms). and the "horsehair worm" are com- mon representatives.500. This group is characterized by segmented body. unseg- "Flame worms. elongated worms which possess both a mouth and an anus. man and other animals.000 known (Jellyfish). cells" are characteristic excretory structures. liver flukes. They are triploblastic and hermaphroditic. others as parasites in in soil water. About 3. All are aquatic and most of They possess radial symmetry. sea urchins. Phylum Platyhelminthes (Flatworms). many There are approximately 3. Phylum Echinodermata. or Comb Jellies).26 functions of life. They are mostly microscopic in size and largely Some live in the ocean. and the freeliving. species. The earthworm and leech are well-known examples of the phylum. and sea lilies. calcareous. others are parasitic. bilaterally symmetrical. fresh water. sea cucumbers. pores. Phylum Coelenterata — them are marine. a single gastrovascular cavit. — ming.500 species are known. There are at least 4. mented. some in fresh water. and still others About 15. Some are free-living. tached. fibrous. and twenty-one of these are American. bilaterally — These are flat. Approximately 6. well-developed body cavity. and have tube feet as organs of locoThe common representatives are starfishes. Free swimmarine animals that possess biradial symmetry.500 known living radially symmetrical They are motion. The described species number at least 4. Tapeworms. —Aquatic Most of them are marine. These animals may be free-living or parasitic.500 species have been described. aquatic Planaria are commonly known.500 known species. —Un- segmented. — . Marine animals which have — a spiny skin and the body wall usually supported with calcareous plates. or siliceous metazoans which live atThe body is supported by spicules. and nephridia as tubular excretory structures. and in the soil.

carnivorous worms of which Sagitta is an example. frequently similar Nemertinea. Phylum Chordata. Many authors have dignified each of these as a phylum. scorpions. In addition to the above generally recognized phyla. etc. or in snail shells. transparent.000 species belong to it. — Unsegmeuted animals that are usually en- The single muscular "foot" is a charCommon forms include clams. snakes. butterflies. proximately 40. Vital Relations of Animals and Plants There are certain single-celled organisms that are claimed as animals by zoologists and as plants by botanists. of which there are about 1. rats. living either free. —nearly unsegmented. contractile. frogs. and all insects. flies. A number of these are sometimes described under the phylum name sessile — — MoUuscoida.000 species have been described in the group. to certain larval stages of annelids and molluscs. Anyone . snails. This is by far the largest single phylum. some stage. Bryozoa colonial. Of course. Many of our best known animals belong here the phylum includes lampreys. and octopuses. sheep. eral — Segmentally constructed animals with bilat- symmetry and an endoskeletal axis or notochord at . acteristic structure. the majority of which are fossil. As a matter of fact. The group includes crayfishes. horses. or fresh-water forms.750 known species Brachiopoda marine animals enclosed in a bivalve shell. cows. closed in a calcareous shell. Sipunculoidea unsegmented. Most of these groups have certain of the wormlike characteristics. and they have segmented appendages. it is not easy to draw an absolutely clear-cut line of distinction. salamanders. Trochelminthes— unsegmented and — . marine. Some authors believe as many as 675. Phylum Mollusca. rotifers being typical. crabs.: INTRODUCTION 27 Phylum Arthropoda. elongated marine worms. centipedes. monkeys. aquatic or terrestrial. bony fish. there are several other more or less independent smaller but distinct groups. and men. Chaetognatha marine. birds. sharks. slugs. in tubes. They are wormlike forms. it is easy to recognize the extremes. Phoronidea — — marine worms living in chitinous tubes in shallow water. beetles. lobsters.000 species have been recognized. alligators. such as bugs. About 78. — The representatives of this group may be Their bodies are segmented. Apturtles.

published by McGraw-Hill Book i .— The metabolic processes of plants and animals ^s (Redrawn by permission f om Wolcott._The carbon and the fundamental living process. 4.28 TEXTBOOK OF ZOOLOGY Chiorophyll Sunlight \nt(zrmedigte decomposition products Respiration / formoldehyde K ^ fats \ Bacteriai decomposition Living ^/Carbohydrates .food a AnimQ/ Sl l /~^trf i-. animals \ (and proteins) / Dead organisms cycle Pig.t of Oa well as the food Fig. Inyalra -^rnn fO-) Hrnnnir. 3. Inc. Company. manufacturing process (photosynthesis).) Animal Biology.

Si m o ttl 8 fe o >> o C <D ho O C 0) bo 8100H ia KOIidHOSaV existence. All organic animal food is derived either directly or indirectly from the process of photosynthesis carried on is plants to manufacture starch which stored in by green plant tissue. Green in sunlight plants by effect of their chlorophyll (green pigment) . animal forms depend on green plants for their distinctly typical a d a S o O >.INTRODUCTION 29 holding a sunflower in one hand and a frog in the other has no The difficulty in determining which is animal and which is plant.

already described. it. Plants are devoured by animals. the greatest riddle in the biological world. it is said to be adaptive. of plants as well as of animals. The size of living organisms varies within definite limits. What is Life?. Living mate- has the ability to carry on active chemical reactions without losing its body form. by means of which the organiit : zation tends to adjust itself to a constantly varying environment. and chlorophyll. Attributes of Life Most of us think we know what is life is. Much more will be said of living material in the following chapter. a gas of the and water is to unite in the formation of a simple carbohydrate. and the by-products excreted are again incorporated in the earth to be available to other plants. In general. Numerous rial attributes of living material may be given. because they extract simple substances from the earth and unite them into complex foodstufl^s. It is responsive to changes in the environmental conditions therefore. by utilizing the radiant energy of the sun. . we find ourselves confronted by an almost hopeless The ques- tion. Living material is able to sustain and reproduce itself under favorable conditions. as a catalyst in causing the combination of water and carbon dioxide. such as the starches and proteins. form the potential material for the animals. . cause carbon dioxide. and some animals are in turn devoured by others. The term life is an abstraction with no objective reality except as is a phenomenon related to the activities of living units. This the basic food material During daylight hours while photo- oxygen is discharged into the air as a product of the process. plants. to produce protein. The complex substances are then broken down with liberation of energy. but if asked to define task. The excretory products of animals contain nitrogen which is easily transformed into a soluble form and absorbed by plants to be combined with the simple carbohydrate. synthesis is in progress. This oxygen adds to the atmospheric supply and is used by animals in respiration.30 TEXTBOOK OF ZOOLOGY air. The following statement has been given and is probably as nearly a definition as can be found Life is a continuous series of reactions in a complexly organized substance. The carbon dioxide discharged by respiration of plants and animals is made use of by plants in this synthesis of material.

Also. has a large permanent population. When our investigators went to Japan to study the enemies of the beetle in an effort to find a means of control. be balanced. It was hoped and intended by English immigrants there. Every city in the United States. So impressed are some biologists berepression. The story of the rabbit in Australia is likewise an interesting example of the effect of balance or lack of it. as well as many in Canada and Mexico. To the surprise and dismay of these people the rabbits flourished until now they are jeopardizing the enterprises of man. Quite clearly most animals live in a state of repression because relatively few of them become pests and overrun the country. Balanced animal communities can be found the world over. a few pairs were landed in Brooklyn. Its nesting and perching habits in the heart of great cities are a source of great annoyance and expense to building owners. that even when some irritating pest is under discussion. Again. that a few imported pairs of rabbits would increase sufficiently so that the old English sport of riding to the hounds might be developed in Australia. and we are only beginning to get a notion of the extensive ramifications of the forces concerned in maintaining that balance. . In the short years that have elapsed. they consume enormous quantities of the farmers' grain. of we have an example of the effect of the natural agents The Japanese beetle which was recently introduced in the United States hy accident has ravaged the vegetation in several eastern states and threatens other areas. this sparrow has proved so hardy and free of enemies here that it is now our dominant bird. coming with the potential danger of interfering with the natural balance. Consequently. Many men are kept employed full time doing nothing but hunting rabbits. About eighty-five years ago someone who had admired the remarkable spirit of the English sparrow in its native European home thought this hardy little bird would be a cheerful addition on this side of the Atlantic. Not many j-ears ago Australia did not have a rabbit within its boundaries. In an established animal community which might be said to all groups are held in boimds by their enemies. they had to search for weeks to find a seriously infested area.INTRODUCTION Balance in Nature 31 The influence exerted by one animal or one group of animals on another can hardly be estimated until one of them leaves the picture.

32 TEXTBOOK OF ZOOLOGY a 0] C bo u o <u bo a o S d m S o 0) o o 0) Pi s 0) d ft >> o C M d bo •i-t .

frog. our government and many others have placed a restriction on importation of plants or animals. or to the health and life Most of the predaceous animals today are not a menace but they do destroy It is likely man directly. the comprehension of social welfare problems. these fields are in themselves applied of inheritance of Most of the great discoveries as to the nature of disease. the formulation of our conception of religion. The study of philosophy. Zoology as Related to Man The values of the study of zoology may be placed in two classes: and practical. and control and the knowledge fundamental physiological processes occurring in our own bodies have been attained by studies on other animals. rabbit. or guinea pig. by man. lobsters. particularly frogs. and many other similar intellectual and social accomplishments are greatly facilitated by a knowledge and recognition of biological principles. From the purely practical or economic side. One of the most obvious as a source of food supply. fish. many domesticated as well as useful wild animals. Because of the danger of interfering with the normal balance or equilibrium in nature. clams. that the parasites which live on and in the bodies of men. crabs. biology. .INTRODUCTION whose extermination is 33 it easily possible. the manner of human characteristics. of course. To wipe out this form might remove the cheek on others that are still more obnoxious. and their related sciences have cultural profited enormously. One must have permission to bring either into this country. There is hardly a field of endeavor in the realm of human activities which is not greatly influenced by zoology and biology generally. In fact. rat. oysters. they will advise against until all phases of the animal's existence are thoroughly investigated. All of the phyla and classes of larger animals furnish at least a places on our menu cards. The loss of their lives is con- stantly saving millions of uses of other animals is human lives. medicine. usually has its application to man. and on domesticated plants and animals have been much more costly than the depredations of the more conspicuous predators. birds. and even snails. The lives of these laboratory animals have made untold and inestimable contribution to the welfare and comfort of man. monkey. What is found to be true in a dog. few species that find mammals. agriculture. turtles. Many of to animals are important because of their destructive tend- encies in regard to articles valued man.

Drosophila. Injurious insects alone cause an annual loss in the United States of more than one and one-half billion dollars worth of products if they could be sold at the price the remaining portion brings. protozoans. and is insects. of such information and how it lays its eggs . can be made in one man's lifetime than was previously possible through ages. application of parasitology. and yet. The United States Fish and Life Service does an extensive and remarkable work in the study.000. It is and mated. .000. had in the fishing industry. the natural fish life does not flourish as it might. the discovery may influence the activities of our entire cotton A recent instance of the economic importance of zooknowledge is found in the saving of the entire citrus industry in Florida from the Mediterranean fruit fly. lobsters. worms. The fishing industry cultures. shrimp.I 34 TEXTBOOK OF ZOOLOGY Agriculture and Zoology It may frequently bring a smile to the lips of an onlooker to see a full-grown and perhaps intelligent zoologist enthusiastically attempting to learn what. Fisheries and the Application of Zoology A very practical and profitable application of zoology has been made The annual salmon catch alone on the Pacific coast has been known to be worth $25. a field of zoology. when. where. and even sponges. The knowledge and would avoid Agriculture has benefited greatly from the application of the and animal breeding. logical ' least half of this loss could be prevented. and markets not only fish of many kinds but also oysters. Losses almost as impor- tant are caused each year by the parasitism of our domestic animals by bacteria. Much fundamental knowledge has come from the extensive studies on the principles of heredity to plant genetics and breeding of the easily kept in the laboratory common fruit fly. It produces a new generation about once every nine days. Departments. and how much a little boll weevil eats or when. propagation. and care of this natural zoological reEven with this work and that of all the State Fisheries source. crabs. More improvement of strains of animals and plants too. With proper knowledge of animal life and application of this knowledge it is likely that at industry. which this loss. clams. collects. The United States Department of Agriculture and the United States Department of Interior have taken the lead in much of this type of zoology.

and freedom from chemical or oil pollution. it is probably true that nothing has influenced the thinking of the world more than the ideas. In recent times. gives one insight to the solution of many of the problems of life. understand something of that her operations are in accord with definite principles. The strictly intellectual has given man are no less valuable than the tangible gifts. and cultural endowments which zoology To the orderly conduct of Nature and to see . salt balance. and knowledge growing out of biological study. vegetation. principles.INTRODUCTION 35 live. our public more appreciation of conditions necessary for a fish to A fish needs suitable water conditions including proper gas content. Many of the superstitious dreads of unseen monsters have been eliminated by the knowledge of the fundamental principles of life processes. nesting places.

not an attempt to give a complete history of the subject. later from the Greeks indicate that there had early Egyptians and been some concern for the problems of life as well as medicine a number of centuries before living Christ.C. including zoology.. However. Early Roman Scholars. totle The other Greeks who followed Arisadded very little of importance. scholars believed that the ocean supported of the original Hippocrates. Aristotle (384-322 B. Alexander had been one of Aristotle's pupils and had become interested in the development of scientific endeavor. Thus even in those times by Aristotle in his investiendowments were being set up for the support of research. as credit for establishing the scientific method direct observations well as his introduction of animal classification. 23-79). sidered There were individual persons interested in and studying natural The works history long before there was any orgajiized field of study recog- nized under the name of natural history or the more limited diviSome of the translations from the sions of it.000 or more) for use gations. He made a grant of 800 talents ($200. compiled a 37volume work in which much of the scientific knowledge of the time — 36 . a Greek from 460 to 370 B. was the first to think of medicine on a scientific basis. and many other animals. a Roman general.C. To him goes the of study which is based and drawing conclusions on gathering facts from from a study of these facts. His observations on the structure and development of embryo sharks. Pliny (a. From shortly before the time of Christ and extending for about sixteen centuries was a period of "dark ages" in scholarly endeavor. of numerous pioneers in special fields are being conthroughout the text rather than in a given chapter.d. a few contributions of note were made. Some all of the early Greek life. He had the assist- ance of the armies of Alexander the Great in collecting materials.) was an outstanding Greek philosopher and scholar.CHAPTEE II HISTORY OF ZOOLOGY This brief chapter is works and This is lives of a selected organized to afford a slight preview of the few of the historic pioneers of zoology. are contributions which caused him to be called a biologist. chicks.

Barbary apes. coming in the midst of the "dark ages" as he does. His anatomical studies were made principally from direct observations on elephants. should be particularly credited for the contributions he made. His textbook on anatomy became the authority for the next eleven or twelve centuries. father collection of Fulvius tlie of naturalists.). (a. published by Henry Holt and Company.) Inc. Galeai did not take advantage of the work of certain of his predecessors who had been privileged to study human bodies. of Biology. and Fig-. Iconographic bas-relief grecque.) (From Locy. Unfortunately. Growth swine.d. Andreas Vesalius (1514-1564). He was of Greek ancestry but moved to Rome early and became a successful physician. During his time it was strictly against the law to make was not allowed this privilege. It does reflect the tendency of the time in that scientific observation had given way to speculation. The return of interest in zoology came about through the medical schools. From a (Visconti. Ursinus. dissections of the so he human body His conviction in the matter of direct observation as a basis of study handicapped him in this respect. and because of the indiscriminate mixing of fact and fancy it is not scientifically valuable. Vesalius was aji active — . Galen 131-201).HISTORY OF ZOOLOGY 37 and traditional superstitions are woven together. found In —Aristotle (384-322 B. His work was limited to compilations.C. 7.

Holt and Company. Inc.) 5.D. 1719. Acta — GalenLocy. publishedMedicorum (From by Henry of Berolinensium. Vol. 8. (A.38 TEXTBOOK OF ZOOLOGY Fig. . Groivth Biology. 131-200). anatomist.

HISTORY OF ZOOLOGY 39 young student and was not Galen's textbook. Frontispiere. anatomist. Nortliwestern University Library. he transferred to Padua where human dissection was Iislied Fig. Vesahus (1514-1564). the first. Grotvth of Biology. to prove that direct . facsimile edition of 1728. He was and Galen. He later since the time of Aristotle became professor of surgery there.) — then allowed. 9. satisfied to accept the authority of Therefore. (From Locy. after beginning his medical education at Brussels. pubby Henry Holt and Company. Inc.

through a closed system of in 1628.40 TEXTBOOK OF ZOOLOGY observation is the only true criterion of knowledge. 10. of Medicine. and his work on the structure and metamorphosis of the silkworm take rank with outstanding con- . an Englishman." and his teaching is really responsible for the rapid development of biology and medicine following his time. studies on glands.) suspected a circuit of the blood from the heart to the lungs and return. and the This arrive at an idea of a complete circulation of all of the blood vessels. Vesalius. new idea was presented He also did notable work in embryology. William Harvey (1578-1657). B. Galen. "William Harvey. but first to Harvey was the first to demonstrate circulation. and embryologist. Following closely upon the epochmaking work of Vesalius and inspired by several of his pertinent observatiojis on the anatomy of the circulatory system. —History William Harvey (1578-1657). Marcello Malpighi (1628-1694) was a famous Spanish anatomist. (From Garrison. published by W. Vesalius is thought of as the "father of modern anatomy. histologist. Saunders Company. began experiments on the movement of blood in the vessels. father of physiology. His observation of blood corpuscles in capillaries. and three or four others had — Fig.

1685. extended observations on bacteria and microscopic animals. and his . Growth of Biology. pioneer micromotist.HISTORY OF ZOOLOGY tributions to zoological knowledge. Antonj van Leeuwenhoek (1632-1723) lived almost contemporaneously with Malpighi and like him made many contributions to the this scientist of his time. He is said to have possessed a total Further study on of 419 lenses. (From a painting by Fig. first descriptions of spermatozoa. capillary blood circulation. published by Henry Holt and Company. Reprinted by permission from Locy.) — development of the microscope. Veekolje. most of which he had ground. 41 of the Numerous organs human body are named for early microscopists. 11. Inc. he renowned had to build his own microscope. Like other Leeuwenhoek (1632-1723).

(Reprinted by permission from Locy. Inc. published by Henry Holt and Company. Carolus Linnaeus (1707-1778) was a very eminent Swedish biolowho. like many early students of this subject. an outstanding Swedish biologist of his time. gist.) — of a species and introduced the use of anatomical features in distinguishing the larger groups. (1628-1705). and numerous genera and species. Linnaeus (1707-1778). was educated as a physician. His was the . 32 sub-classes. Linnaeus believed in a rigidly fixed species and had divided the animals into six classes. clear. 12.42 TEXTBOOK OF ZOOLOGY valuable contributions to the development of the microscope are the enviable accomplishments of this man. Growth of Biology. He followed somewhat in the footsteps of Ray who had paved the way by fixing a definite conception Fig. In spite of his idea of the invariability of species his classification system was so simple. and flexible that it has persisted to the present time.

is Louis Agassiz (1807-1873) commonly regarded as the father of American zoology and a renowned student of comparative anatomy. the cell theory (principle). Almost immediately following Linnaeus came the Frenchman. Lamarck (1744-1829). Ernst Haeckel organized the modification of this system in modern times. is referred to as the founder of comparative physiology and the first to apply the facts of physics and chemistry to living protoplasm. hence the two names. who among other important things is credited with being first to realize that there are different lines of descent in 1866. His work was a great impetus to modern physiology. and it is known as the Binomial System of Nomenclature. Johannes Miiller (1801-1858).HISTORY OF ZOOLOGY first 43 natural system of classification. Linnaeus is said to have classified and listed 4. He was a recognized paleontologist as well as zoologist. is one who embryology as a field of study. came as a result work and thought. is Georges Cuvier (1769-1832) of comparative anatomj^ credited with establishing the field of He was tlie French ancestry and largely self-educated by his studies at cal structures bear his seashore. Matthias Schleiden (1804-1881) and Theodor Schwann (1810-1882) are the two Germans who in 1838-1839 arrived at one of the most important generalizations of biology. . Avhich of his is explained elsewhere. He is responsible for one of our first and oldest Marine Biological Laboratories.*' thus explaining the unfolding and differentiation of the various organs of the developing animal. a Russian biologist. really established Karl Ernst von Baer (1792-1876). His great inspiration has permeated through his students to nearly every institution in the land. but it is specifically known by the genus and species names used together. Much later. a German scientist. The recapitulation theory. A number of anatomi- name. His notable paper on the development of the chick was published in 1832.378 species of plants and animals. He estab- lished the "germ layer theory. Each individual not only fits into larger general groups by this sj'stem. as used and that no living species is absolutely fixed. This is to be discussed further in the following chapter.

published by Henry Holt and Company. Inc. studies on the problem of the species of organisms arise. made extensive manner and means by which new effectively developed the thesis He very that they originate by a process of natural selection. Those become extinct. Agassiz (1807-1873). with those whose natural variations happen to fit them best to the changing features limitations determining the development of of the environment persisting as crowded out.) — new species. an Englishman. least fitted to the dominant species and others being environment would naturally Darwin did not claim originality in his idea. Biology and Its Makers. Lamarck. that new varia- or groups best suited to their persist and produce progeny. Buffon. This was based on the idea that no two individuals are exactly tions are constantly appearing.44 Charles TEXTBOOK OF ZOOLOGY Darwin (1809-1882). 13. and Erasmus Darwin. grandfather of Charles. and finally that those individuals environment would be the ones to His conception of the factors and P^ig. (From Locy. had presented similar . pictures a constant struggle for existence among organisms. great American pioneer zoologist. alike.

the author of Origin of Species. he derived the original laws of heredity. His results were first published in an obscure Swiss paper in 1866 and were not really discovered and appreciated until 1900. B. He was the founder of genetics.HISTORY OF ZOOLOGY ideas before him. The facts which he established are now known of Heredity. and the remainder resembled the other. From his work there. on the theory of natural selection. History of Medicine. published by W. He crossed different kinds of peas and found that the offspring in the first generation all resembled one parent. Garrison. a contemporary who had reached the same conclusion. In 1858 he read a joint paper with Alfred Russel AVallace. From these facts he referred to characteristics of the former group as dominant and those of the latter as recessive. Saunders Company. 45 It was the vast accumulation of facts covering a period of twenty years which commanded the attention of scientists as well as the public generally. Charles Darwin (1809-1882).:" Fig. (From 14. as Mendel's Laws .) — Gregor Mendel (1822-1884) was an Austrian monk who carried on experiments with the breeding of garden peas in the cloister garden. ' >T. When these oft'spring were interbred he found that three-fourths of their progeny resembled one grandparent. That same year Darwin published his book Origin of Species which is a classic in its field and familiar to all scholars.

46 TEXTBOOK OF ZOOLOGY . taken from an autobiography. the Austrian monk who pioneered In Plaque by Theodor Charlemont based on "Fuschia picture" 1864. Greg-or studies of heredity. C^^f^e^a^-^ Fig.(S. 15. The signature is his.) . — made between 1861 and Mendel (1822-1S84). (Courtesy of the Journal of Heredity.

He rescued the silk industry of southern Europe by discovering the organism which killed the insects. He showed that living organisms cause fermentation and demonstrated that these organisms and others could be killed by heating them to a certain temperature. Comparative anatomy and paleontology were greatly advanced through his influence. He was . Thomas Henry Huxley (1825-1895) Avas one of the most popular English scientists of his day. published by W. In 1861 he put an end to the controversy regarding spontaneous generation of living organisms and established the idea that all life in present times comes from life. and he also discovered an immunization process and treatment for hydrophobia. He showed that materials thus heated and then sealed would not ferment until after they were exposed to the organisms in the air.HISTORY OF ZOOLOGY Louis Pasteur (1822-1895) was a French scientist 47 who had been trained in chemistry but became one of the outstanding pioneers in applied biology and medicine. (From Garrison. B.) process grew out of these experiments. History of Medicine.— Louis Pasteur (1822-1895). 16. He was one of the principal champions of Darwin's ideas and theories. August Weismann (1834-1914) was a German zoologist who started out as a physician after having been trained in that field. The pasteurization preexisting Fig. one of the benefactors of mankind. Saunders Company.

The work of all those mentioned and hundreds of others has given us the background for our present knowledge and grasp of zoology and medicine. is a small book in itself. This book is extremely interesting to . Lincecum. which is important to all modern biological conceptions. particularly E. Geiser of Southern Methodist University. He is widely known for his experimental studies in plant breeding and with evening primrose. D. S. Walker. genetics. is continuity of germ (1848-1935).48 TEXTBOOK OP ZOOLOGY fields of an outstanding scholar in the is for his theory. particularly portant contributions. best known heredity and embryology. and others have been described in the recent book by Dr. transformations. It is an The printed program in the realm of the experimental endeavors. American Zoological Society. W. "Webb. entitled Naturalists of the Frontier. a Dutch botanist. brought out the mutor tion theory. omists of America. Vliet. that there He plasm from generation to generation. Gustaf W. Cope (1840-1897) was one of the greatest comparative anatHe dealt not only with living forms but with fossil materials as well. The works and lives of such prominent pioneer zoologists of the Southwest as Jacob Boll. History is being made so rapidly in these fields durdifficult even to catalogue the imextremely active field. though small. which for the annual meeting of the is made up largely of titles and abstracts of new papers to be pre- ing the current years that it is sented. His idea was that species have not arisen through gradual selection requiring thousands of years for each but by jumps through sud- Hugo DeVries den. read. Belfrage.

This name later came to be applied to the real cells.Matter. During these years it had gradually dawned on biologists that this matter is found in all livthe of Dujardin. characteristics. and plasma. gummy substance and called it sarcode. which means "flesh. for cells do not have a hollow structure but are typically semisolid bodies. as it is more and more being learned about its is nature. an Englishman. Hooke. Purkinje. but the origin of living matter. Malpighi had described 49 . Leeuwejihoek saw spermatozoa and bacteria and included them with single-celled animals as "little beasties". In 1840. in 1835. saw the spaces in it and called them cells because they reminded him of prison cells. or Little is Protoplasm known concerning called. a German botanist." He was able to test its solubility and its behavior with alcohol and acids sufficiently to satisfy himself that it differed from ordinary gelatin or albumin. which comes from the Greek protos. realized that Frenchman by name ing things. The Cells Cell Principle superficially described during the had been seen and even latter part of the seventeenth century the eighteenth century.CHAPTER III PROTOPLASM AND THE CELL Living. a Bohemian biologist. and activities. and called it protoplasm. Living matter ahvays active in some degree. A some of the simple microscopic animals he was studying were composed of a soft. with Avhich it might be confused. but serious study of the material was not begun until approximately one hundred years ago. or protois plasm. In 1846 von Mohl. gave living matter the name protoplasm. in 1665 in observing cork with the microscope he had made. structure. viscous substance similar to that already seen in animals'. He was instrumental in bringing this name into common use. but their significance and numerous times during was not realized. and this activity attracted the attention of scholars at a rather early date. It was an unfortunate term. saw in plants a granular. anything formed or molded. first.

(From Locy. Schleiden (1804-1881). and in 1831 Kobert Brown had discovered the nucleus of the cell. . Inc. but not until the work of Schleiden in 1838 and Schwann in 1839 was the cell theory formally enunciated.50 TEXTBOOK OF ZOOLOGY the nature and appearance microscopically of several organs of the body. each working independently. published by Henry — Holt and Company.) the latter a zoologist. Growth of Biology. Matthias J. Grew had made rather extensive microscopic studies of plants. as they gave it. The former a botanist and Fig. was in substance. noted German botanist who helped establish the cell theory. All living things (plants and animals) are composed of cells. 17. This theory. came to the same conclusion and in 1839 collaborated their ideas.

It was soon found that some tissues are made up not only of cellular structures but included also certain noncellular materials produced by the cells. (From Garrison. the German zoologist who. This matrix is not strictly living matter since it is inactive and passive as far as life processes are concerned. B. Saunders Company. the part. Fig. yet the cell theory proved to be such a unifying generalization and inspiring stimulus to investigation that it became the turning point in the development of biological study. cells since practically cell wall was thought to be the essential though now we know it is not a universal structure of all no animal cells have a cell wall. in 1838 and 1839. Theodor Schwann (1810-1881^). 18. Since living bodies are composed of such an abundance of this noncellular . History of Medicine. The matrix. so abundant between the cells of cartilage.PROTOPLASM AND THE CELL It is 51 that they and no discredit to this theory or these men many dis- other biologists of the time had erroneous ideas concerning the essential features of the cell. was soon found to be noncellular and to be produced by the cartilage cells which became embedded in it. Connective tissue fibers fall in the same category. The notions of the origin of cells and the functional significance were almost wholly fantastic. published by \Y.) — The bare statement that living beings are composed of cells soon became inadequate as studies of cells progressed. Although Brown had recently covered the nucleus. collaborated with Schleiden in formulating the cell theory.

the With the years. can hardly as well as its effect be over-estimated. It was the first great generalization in biology. Through the rather rigid and constant set of developmental changes for which the cells are responsible. cell theory on biological thinking and progress on fundamental thinking generally. and the principles of heredity are some of the results of this conception of life new embraced in the cell theorj^ .52 TEXTBOOK OF ZOOLOGY produced by the : m aterial cells. chromosome rela- and modifications have come to be recognized as being brought about in or by the cells. Many new problems arose with this new knowledge of plants and animals. the cell prmciple soon came to be things are composed of cells and cell products. The cell is now regarded cell as a physiological unit as well as a structural one. The conception of this idea was one of the great landmarks in development of biological ajid scientific thinking. and as almost a corollary to the original statement of the principle. that the activities of the organism equal the cells. there is developed a new individual which usually resembles tionships its parents quite closely. tissue formaof heredity and development. Deliberation and meditation on this fundamental idea seemed to prepare biologists for other great generalizations which followed quite rapidly. the transformation of energy by chemical cells. It is comparable in The influence of the the field of biology to Newton's law of gravitation in the field of physics. reaction within the roles of electrolytes in living substance. sum of the activities of its the embracing of the functional activity of the cell as a part of the principle underlying living processes comes also the inclusion Cell division. Comparative morphology was extensively investigated. namely. Up until this time there had been no single fundamental idea applied to living material that was recognized as being univer- This conception focused the thinking of all biologists sally true. and physiology now has become physiology of cells as a result of this impetus. stated thus conceptions of the nature of the nucleus. formation of cell products. and the composition of protoplasm itself have all added their contributions to the present understanding of the meaning and application of the cell principle. the all living membranes. growth. in the same direction and therefore it had a great unifying influence. of cellular An understanding of the permeability membranes. migration of cells. With tion.

^. or colloidal emulsion.. A colloid a substance of gelatinous nature.. persed particles of the colloid become congested by loss of general fluidity. 19.. This condition is increased water in the dispersion known as the gel state. but they are too small to be seen with the ordinary microscope. — structure of The is living protoplasm as seen in tlie starfish. the colloidal state tends to be- . As a constituent of protoplasm there is always a considerable percentage of water. it may be said that this substance has a variable degree of fluidity under different conditions. however.' S • . : i->. nearly colorand more or less translucent. WT. published by The Macmillan Company. -v •"•<. Km •• •" / *' '''. which conditions the degree of gelatinous in consistency.J^"^-?'^•%^. It is more or less granular. i- --•. it is never perfectly transparent. In the emulsoid. When there is medium and the particles move with greater ease in the more fluid medium.*. Fig.PROTOPLASM AND THE CELL 53 General Characteristics of Protoplasm and the Material of the Cell To begin variation with.•. are larger than molecules. and this is With loss of water from the dispersion medium the disreversible.-.) Protoplasm is in a colloidal state of the emulsoid type. Cell. -- f * ' . viscid and less. A colloid is identified by the presence of particles which are groups of molecules dispersed through a more fluid or watery phase.. '-^J'. It is possible for water and substances in solution to enter protoplasm from without. permeable by crystalloid solutions.. '" ._.^ * - . of course.'. The range It is of this may be from semisolid to semiliquid. >v^^. These particles.. viscosity. the substances are distributed through the more watery or dispersion medium.' J'y- ' . (From Wilson. The trauslucency causes a mass of it to have a lustrous gray appearance. and diffusing not at all or very slowly through animal or vegetable membranes.

materials there are being oxidized to liberate kinetic energy. and produce waste by-products. so this simply mentioned here as another prop- erty which is often listed. All protoplasm has this capacity. such as utilization of food. which refers to the awareness of one's ence. is which is the power of contraction and relaxation that 4. All living organisms are capable of this by some means. be due to chemical changes in dispersion medium of the colloid. own exist- It is certain that some protoplasm possesses consciousness.54 TEXTBOOK OF ZOOLOGY sol. at the same time. the process of continual exchange of food and fuel materials being built into the protoplasm. while. When the rate of the building side of metabolism exceeds the oxidation rate in the protoplasm. there may be mentioned and all 1. Contractility. to change come This transfer of water may from sol to gel state and back to sol repeatedly is the basis of many of the vital activities. Reproduction is the capacity for producing new individuals of the same kind. disposal of waste. 2. because of its colloidal nature. such as heat and movement. all Spontaneity is also considered a prop- erty of protoplasm source of by some. described briefly a number of important activities common to protoplasm. Conductivity refers to the fact that the impulses produced by cell stimuli or irritations at one point in protoplasm are conducted to other parts of not only a single 3. Growth is and hence growth. is frequently given as a property of protoplasm. which refers to the capacity present in all protoplasm for responding to changes in environmental conditions. These properties are: Irritability. or external stimuli. . To be certain that the activity and reaction comes from within is likewise rather difficult is of definite proof. common to the substance of every cell. and movement. Metaholism. Simple cell division is the most primitive process of reproduction among animals. but also to adjoining cells. recognized as any increase in volume. Fundamental Properties or Activities of Protoplasm In addition to the general characteristics. 6. but evidence of this quality is rather intangible. there is storage of materials in the mass of the protoplasm 5. Consciousness. the dispersed particles or in the The ability of protoplasm.

while the particles or molecular aggregations constitute the dispersed phase. This increase in surface is one of the many important significant effects of colloidal organization of substances. only three or four times that of water. one centimeter its total surface will be is 12. because reactions occur at these surfaces. known as disperse systems of the emulsoid type. It changes fluid or sol state to a more solid or gel state and may return in the other direction. that of water. Its physical constitution is similar to glue or gelatin. These up an expression of energy in that they move against each other as though they were dancing in a limited space. This activity can be seen only with a special optical arrangement known as the ultramicroscope and the phenomenon is known as Brownian movement from a (characteristic of colloidal substances). such as sugar or ordinary table salt (sodium chloride). if the same volume of material in colloidal particles of the average size given above.6 square Now. it is a colloidal system of Colloidal systems a number of chemical compounds existing together. it is while with the dispersed particles included only eight or ten The viscosity of the nuclear fluid is only twice Since glycerin has a viscosity about a thousand times as great as water.000001 of a millimeter in diameter. it will be realized that fluid in its active state. Ordinarily the viscosity of the continuous is phase or supporting liquid times that of water.000 square meters.PROTOPLASM AND THE CELL Physical Nature of Protoplasm 55 Protoplasm is a semifluid material which is heavier than water and somewhat more refractive to light. essential to most protoplasm is quite Changes in viscosity accompany and are the activity and functioning of it. in the Instead of being form of a true solution like salt in water. An important consequence of the colloidal systems in protoplasm is the enormous surface of particles exposed to the continuous phase.0001 and 0. it consists of sus- pensions of relatively large molecular aggregations varying roughly particles keep between 0. The more watery or continuous part of the system is loiown as the dispersion medium. If a sphere of material has a radius of centimeters. the total surface of these will be approximately 7. rather than to crystalloids. By the presence of salt ions in the continuous phase and these becoming adsorbed . fuses slowly or not at all Protoplasm difthrough animal membranes. is Protoplasm are not a single compound.

carbon. bromine. When a molecule of glucose is burned. The organic compounds include carhohydratcs. nitrogen 3 per cent. hydrogen. protoplasm has eluded complete and Nevertheless the compounds of living matter are composed of several elements. sodium. and frequently others like silicon. hydrogen. The proportion of the hydrogen to oxygen in the molecule is the same as found in water. sulphur. iodine. two to one. The carbohydrates. its colloidal nature. are compounds of carbon. Fats.'ater dioxide (CO2). chlorine. glucose. calcium. aluminum. for storage in the various animal tissues. phosphorus. povmds consist of several inorganic salts and water. and also enzymes. hydrogen 10 per cent. many of them the most ordinary and abundant in the world. or simple sugar. are composed of carbon. and hydrogen. part of the The most abundant of these are found named in the list. fats. they acquire an electric Protoplasm exhibits these several phenomena because of upon the surfaces charge. A few of them are usually given as constitut: ing approximately the following percentages of protoplasm oxycarbon 18 per cent. phosphorus 1 per cent. The principal carbohydrate found in protoplasm is the monosaccharid. like carbohydrates. magnesium. copper. proteins. calcium 2 per cent. and first fluorine. oxygen. the potential energy is released as kinetic or mechanical energy. Chemical Nature of Protoplasm Up to exact chemical analysis. There is much more carbon and hydrogen with less oxygen. As a rule the elements found in protoplasm are oxygen. Glucose is of energy. and oxygen but in more complex molecular arrangement. the present time. and all others makijig up the remaining 1 per cent. potassium. nitrogen.56 TEXTBOOK OF ZOOLOGY of the colloidal particles. and there are formed (H2O) and six molecules of carbon converted to a starchlike substance. manganese. which include starches and sugars. This substance must be reconverted to glucose before it is available for production six molecules of v. glycogen. These elements are found combined to form compounds. which allows the fats . iron. whose formula is CeHioOeThis is actually built into some parts of the cell. but its chief function is to furnish the most available source of energy by its ready oxidation. The inorganic com- gen 65 per cent. The list of elements necessary to make human protoplasm could be gathered in almost any locality on the face of the earth.

The general function of the catalyzer or catalytic agent is that of facilitating and speeding up certain chemical exchanges without the agent itself The well-known example of catalysis amount of platinum in increasing the rate at which hydrogen and oxygen combine to form water. Egg albumen. under favorable conditions. A particular enzyme is usually specific for one kind of reaction. and lean meat are common examples of proteins. gelatin. posed of thousands of atoms teins rent. Since many enzymes influence only one specific type of chemical reaction and since there are numerous entering into the reaction. Proteins constitute the bulk of the foundation or framework of the and are the most abundant organic constituents. Enzymes taken from one species will usually facilitate the same kind of specific reaction in other species. alcohol. Such common substances as lard. with the frequent addition of traces of sulphur. phosphorus. . high resistance to electric curand usually coagulate upon heating or upon addition of acids. Enzymes are substances whose exact chemical nature is not yet known. : of the structure of the cell and. Chemically and physically they seem to be more like proteins than anything else. These substances are found not only in the cells. tallow. the effect of a small is . and iron. magnesium. or salts to form a clot. and cottonseed oil are good examples. whether it is in the stomach of a frog or of a man. Fats serve a double function in protoplasm constitution of a part more energy. whale blubber. pepsin will bring about the same general reaction. since Aveight by weight it contains more potential energy than any of the organic group. All of the proteins have large molecules. as an illustration. secondly. but they are also secreted by cells into the digestive tract and into the blood stream. take hemoglobin of the red blood corpuscles with its formula C7i2Hii3oN2i40245FeS2.PROTOPLASM AND THE CELL to 57 combine with more oxygen in oxidation and therefore release Fat is extremely well adapted as a form of material for storage. but whose importance to protoplasm is probably unequaled. each being comcellular structure They are composed of carbon. The digestive enzymes may be thought of as an example. Of these. They are split into numerous amino acids which serve as the building stones of the stable portions of protoplasm. hydrogen. the storage of food. but not for the species of animal in which it will function. where they act as organic catalysts. and nitrogen. oxygen. butter. Pro- have a slow rate of diffusion.

and when active it is not the same for any two consecutive moments. and sulphates of sodium. changes in this balance have regulatory effects on metabolism. that there must be is seen numerous enzymes present in the cells of every organism. are stances. potassium. Young cells contain more water than old ones. . and because of its high specific heat. but it is important to protoplasm because of its comparatively high surface tension. and slight salts are The inorganic relatively small amount. magnesium.58 TEXTBOOK OF ZOOLOGY it types of reactions going forward in active protoplasm. which are able to combine with all the other substances in protoplasm. as a part of protoplasm. its more complex than any other sub- is quite unstable in that it changes composition in response to every change in the environment. and iron are important salts of living cells. The chlorides. Even the proteins. The majority of cells. it is On account of the extreme complexity of protoplasm all not sur- prising that the chemistry of pletely understood. The relative proportion of these salts is kept at a fairly constant level. of its activities is not yet com- Structure of a Typical Animal Cell The quantity of protoplasm comprising a single cell varies within wide limits therefore cells vary greatly in size. The relative amounts of water in relation to other materials of the protoplasm vary in different cells and in different species. From the chemical standpoint. young organisms likewise contain more than old ones. carbonates. phosphates. but not all of them. protoplasm all of the necessary adjustments of living matter to its environment. because its presence gives the protoplasm a consistency compatible with the range of variation necesThis sary for metabolism. Water is not only a very solvent. Water constitutes 60 to 90 per cent of protoplasm and maintains efficient many substances in solution. The exceeding variability of protoplasm chemically. and therefore split up in aqueous solution into ions. living protoplasm is considered the most complex of all systems of compounds. iodides. latter point is important in maintaining protection against sudden and extreme temperature changes in the living organism. require considerable magnification to be seen. Cer- . calcium. present in considerable numbers but in They are electrolytes. makes possible In a sense.

which lives in the intestine of the lobster. Egg cells. polygonal. Plasma /Atmirane Eciop/asm Chondriosoma En</op/asm Lin in Chromaf/n Mucf^us Nucteof Sap Vcict/o/e A/</c/zar Membrane Plastid Fig. all cells are far from this shape. Some of the nerve cells. specialized functions. —Diagram showing a typical animal is cell. Porospora gigantea. 20. Muscle cells are relatively also. and may reach from one-half to two-thirds of an inch in length. and spindleshaped forms. long may exceed this may be several size.PROTOPLASM AND THE CELL tain of the single-celled blood parasites are about as small as cells 59 known. any They are barely seen with our highest magnifications. . but due to the effects and unequal growth They vary greatly in shape and include platelike. columnar. feet in length. though of less mass. cubical. The particular form of any cell is not a haphazard matter but strictly controlled by morphological and functional of mechanical pressure. The shape almost of the typical cell spherical. necessities. At the other extreme of size we may refer to another parasitic single-celled animal. including the yolk.

which have a specific arrangement of grouping. The cytoplasm usually includes the larger part of the substance of the cell. the chromomeres. may may be stored as gi-anules in the cytoplasm. c to li are other groupings of chromomeres along the chromonemata of the chromosome. or oils. The more nearly fluid. Genetics 22: 616. and the interspersed fibrillar substance known Within the cytoplasm. showing. the protoplasm having more density here to form the plasma membrane. 21. Threadlike Golgi elements or apparatus may be observed in the cytoplasm. particularly near the nucleus. lA is a semidiagrammatic representation of the types of chromomeres and the ways in which they are connected. At a in the main figure there are two rows of dotlike chromomeres which tend to associate in pairs. The nucleus.the details which appear on the upper 5?urface of a fully developed salivary gland chromosome (large vesicle type) — from Simulium fly larvae. This membrane. The band labeled h is composed of 15 or 16 vesiculated chromomeres closely pressed together. lying near the nucleus. structureless fluid. most animal cells is the centrosome. Secretions produced in the cell cavities filled with water. It may be subdivided into the more nearly clear. (From Painter and Griffen Chromosomes of Simulium. or This cell membrane. consists of a part of the protoplasm whose density is somewhat greater than the adjacent portions.) : Scattered through the cytoplasm also are numerous rod-shaped bodies known as mitochondria. In some types of cells two separate membranes may be distinguished.60 TEXTBOOK OF ZOOLOGY A cell consists of a mass of jellylike cytoplasm surrounding a The outer surface of the cytoplasm is modified. transparent . Its substance is known as kinoplasm and is made up of two parts. A camera lucida drawing. the larger ce7itrosphere. In plant cells the plasma membrane is covered by a cellulose cell wall. 1937. gases. hyaloplasm. enclosing a (two if divided) centriole. is surrounded by the cytoplasm and separated from it by the nuclear membrane. in as spongioplasm. which is usually round and centrally located. like the plasma membrane. also certain inclusions be seen here. which is the outer covering of the animal cell. Vacuoles are often present as small lis I h ' Fig. and these consist of a linear series of granules. The protoplasm which constitutes the nucleus is usually known as karyoplasm. nucleus. The longitudinal threadlike bands are called chromonemata. membrane is living and semipermeable.

His idea was that the nucleolus split first. and this may be expressed in a ratio. while the meshwork it is called linin net. which is thought to be the center of functional activities of the nucleus. It is not entirely possible to define the part each plays in the metabolism of the whole. although catabolism goes on until the cytoplasm is depleted. and finally the cytoplasm divided. There is a definite relation between volume and surface in any mass of material. Cells that are deprived of their nuclei are unable to carry on assimilation. With variation of the size of the mass. The cell is often spoken of as the unit of fine fibers extending through and function in living material. The threads of chromatin are called cliromonemata. Mitochondria. When the limit of growth is reached the cell divides. or plasmosome.PROTOPLASM AND THE CELL portion of this is 61 haryolymph. was called method of amitosis. This of size is fixed primarily by the physiological necessities which are transmitted through the surface of the cell. or nuclear sap. are also found in the nucleus. the volume varies according to the cube of the diameter while the surface area varies according to the square of the diameter. Supported on this net is a dark-staining granular or fibrillar substance known as chromatin. The usual There mitotic cell division is far more complex and less direct. most nuclei have anotlier body composed of material thought to be temporary storage products of nuclear metabolism. This is . was the first to describe cell division. It is generally thought that in these bodies are located the units of material (genes) which function in the transmission of hereditary characteristics from one generation to the next. Chambers. in 1855. are several preliminary changes or phases which must occur before the actual cleavage of the cell into two new ones. placing each porThis direct method of division tion with its share of the nucleus. Then besides these. the nucleolus. Both nucleus and cytoplasm are necessary for its normal activities. of structure Cell Division The limit cell is limited in its size. and this restores the proportion of the surface area to volume that will again permit growth. it is possible to dissect the nucleus of a cell. There are usually one or two knots of more dense chromatin in the nucleus which are called karyosomes. then the entire nucleus. Its actual occurrence is quite rare. Remak. similar to those of the cytoplasm. Since the development of the microdissector by Dr. as is the complete organism. the chromosomes. During division of the cell this granular material becomes arranged into definite bodies.

62 cell TEXTBOOK OF ZOOLOGY division. DAUGHTER mitosis : CELLS 22. (indirect cell division) as . phase. late anaphase.shown in in order resting cell. an'^ (Drawn by Titus Evans. more briefly mitosis. MEfv»BRANE CENTROSPHERE tvlUCUEAR CENTBIOLE MEMBRANE CHROMATIN NUC!_EOI_US CYTOPLASM EARL-Y SPINDL. and the two pieces have moved considerably apart. late prophase. daughter cells. telophase. or indirect cell division.) in stages typical They follow each other early metaphase. The . and the early prophase condition is seen. In this stage the centriole has divided. This method of division was first described by Fleming in 1878. Following the resting cell condition come the first changes. is Although the process of mitosis for convenience in stud3% these CEL-l- a continuous series of changes. changes will be set out as six phases. early proanaphase. though Schneider in 1873 described much of the complicated process.E prophase: EQUATOR P1_ATE r Al_ ASTRAU RAYS C H ROMOSO M E ASTER LATE PROPHASE ADJACENT CEI_i_ MEXAPHASE Fl INTER ZONAU BE RS EARLY ANAPHASE CL-EAVAQE LATE ANAPHASE FURROW TELOPHASE Figr. —The fertilized Ascaris eggs.

The chromosomes. in certain cells at least. the nuclear membrane begins the chromosomes become arranged on the spindle fibers in an orderly fashion midway between the centrioles to form the equatorial plate.PROTOPLASM AND THE CELL 63 surrounding protoplasm has produced some rays radiating from each centriole. These two bodies are now known as asters because of their starlike appearance. are involved in the phenomenon.) linear arrangement (Some hold that from one cell genera- Each consists of a double linear series of chromatin bound closely together to form the thread. each of the chromatin threads now shortens and condenses to become a chromosome. bodies like two chains A single one of these . The nucleolus disappears and the chromatin which ap- parently up until this time has been somewhat dispersed through the nuclear substance in the form of granules. The spindle extends between the two asters and to degenerate. arranged in the equatorial plate. chromatin its fibers or threads. and they carry the chromatin material of the new cells which result from the ensuing division. with which they are associated. There is a constant number of these in the cells of each species. These " half -chromosomes " each soon come to have two chromonemata. The two asters taken together are called the aniphiaster. the centrioles have reached the polar positions on opposite sides of the nucleus. becomes organized into long. Following this stage each "of the new chromosomes. or pole. changes. resulting from this splitting or division. It has been reported that. placing one chromonema in each part. According to modern explanation.. . two series in the thread is known as a chromonema (pi. During the above In the late prophase. although there is general belief that the spindle fibers. the prophase stage al- requires about eight minutes. The characteristic feature of the metaphase is this splitting. This period is the anaphase. In the middle prophase the centrioles have migrated still farther from each other and the splindle fibers between the centrioles as well as the astral rays around them have become well established. The explanation of the movement of these chromosomes from the equatorial plate out to the poles is not entirely forthcoming. possibly tangled. still The metaphase follows with no interruption. 21). now each split longitudinally. the chromatin retains tion to the next. migrates along the spindle fibers toward its respective centriole. The nuclear membrane now has most completely disappeared. chromonemata) The chromatin bodies comprising the chromonemata are often called chromomeres (Fig.

Then after a further period of from one to twelve hours. it is stated that this requires less than two hours. "continuity of protoplasm. At this time a constriction of the cytoplasm begins in the plane of the equatorial plate. so it takes its normal position just outside the nuclear membrane in the cytoplasm. declared the doctrine that all cells must be derived from previously existing cells. In 1855 identical hereditary qualities. and immediately the chromosomes begin to separate from the group. "omnis cellula e cellula." This supposes that in the first living material created were inherent all of the possibilities which have been realized in all living things that have existed since. A nuclear membrane forms to enclose each chromosome gi'oup. growth is rapid until they reach their typical limit of size. These two new cells resulting from the division are spoken of as daughter cells. mation of the new nuclear membrane excludes the centrosome. These cells have each received the same quantity and quality of chromatin material. of this process in all types of organisms. nized as the material which makes possible the inheritance of qualities from cell to cell and. in case of sex cells. another mitotic division may take place. both qualitatively and quantitatively. which are in the resting stage as far as division is concerned.64 TEXTBOOK OP ZOOLOGY the chromosomes approach the poles of the spindle they crowd very close to each other. The fortelophase stage. One or more nucleoli soon become evident. The nucleus resumes its granular appearance of the resting cell. in his statement. both plant The universality and animal. Virchow. This is the beginning of As the The cytoplasm perfects its constriction and divides into two parts. The chromosomes progressively lose their identity and their staining qualities." and that present chromatin comes from pre-existing chromatin. For most average cells under optimum conditions. . which the chromatin is divided between the two cells seems to Chromatin is recogindicate that this is a most significant step. a German pathologist. from generation to generation. The purpose of the splitting of the chromosomes in the metaphase stage seems to be to provide each daughter cell with suggest that This equal division of chromatin. and the regularity of the occurrence of the phases of the process The great precision with it is of vital significajice. At about this time the centriole divides into two. has given rise to the thought expressed in the phrase. although certain ones still clump together. Following the organization of these daughter cells.

Most them live in water while a few live in the body fluids of other animals. Certain types are found living rather abundantly in the soil water. and total lack of such features. changing shapes others are nearly spherical. spindle-shaped. that modern Protozoa have descended. . A few are encased in hard coverings or shells which are made up of a secretion from the cell alone. therefore. as in one of the 65 classes. Being single-celled. families. or of a combination of of . cylindrical. they are considered the oldest. from primitive organisms that were also the ancestors of Metazoa. systematically classified and divided into classes. beginning with the simplest. Protozoa are univer- when animal groups are placed in the order of complexity. In spite of the exceedingly large of species and microscopic size. They are found in almost all conceivable shapes. although many of them are perhaps more complicated than numerous many-celled or metazoan forms because of the extensive modifications of the one sally placed first cell. such a secretion with a foreign material like sand. the phylum has been quite is of the animal kingdom. Characteristics The great majority of Protozoa are microscopic creatures. The phylum is usually divided into four classes. without changing their single-celled condition. "With the excep- tion of one class the Protozoa have characteristic locomotor organs. Most Protozoa exist singly as an independent cell. orders.CHAPTER IV PHYLUM PEOTOZOA first to IN GENERAL The animals included in this group are usually said to be the have existed on earth and. each characterized by a distinctive locomotor structure or by the species. ^to' Classification This group is often spoken of as a subkingdom as well as the first phylum number genera. Some have irregular. oval. but some are organized into groups called colonies. and vase-shaped. they are usually referred to as the simplest known animals. It has been supposed and with reasons to support the supposition.

TEXTBOOK OF ZOOLOGY Class Mastigophora (mas ti gof '6 ra) which means whip bear- ers. The flagellum is a contractile There are some species in which exist both flagellate structure. and amoeboid stages. — Group of representative Curtis and Guthrie. and means of locomotion. material. as in the intestine or the blood stream. published by John Wiley Inc. This class also has a close relationship of its representatives possess chlorophyll. saprophytic.) (Figure of Chilomonas modified. Phaous Trachelmonas Peranema Fig.) These forms are frequently classified as plants class b}^ botanists. and (b) those more plantlike forms which may be holophytic. In some species they serve the organism in feeding. includes forms that possess one or more whiplike extensions of The number of flagella is limited. saprophytic. Entozoic is a name applied to forms which live within the bodies of other animals. This seems to show a rather close relation the cytoplasm. Textbook 23. Maatigamoeba from and Sons. . with plants in that many 111 _ \>asis Monosiga Cercomonas Cht lononas Codonosiga. Mastigophora.66 1. Holozoic refers to forms which ingest and digest food Saprophytic refers to the habit of absorbing nonliving organic matter in solution directly through the surface of the body. The Mastigophora is divided into two groups: (a) the animal-like forms which may be holozoic. or flagella. (Reprinted by permission of General Zoology. or entozoic. or entozoic. they serve the animal as its of this class to the next.

The aniable to accomplish locomotion by extending the protoplasm into these pseudopodia. Retortamonas in the and Enteronomas are all genera with representatives occurring digestive tract of man. some bind in sand or other solid substances with one The class is commonly divided into five orders. Euglena is a very common!}^ studied fresh-water an interesting marine form which is pelagic (lives at the surface) in its habits and appears as a thick. Chilomastix. fleshy) or Rhizopoda (rizop'Oda. ponds. 386). and others are covered by a shell. Inc. square rods. oily odor and unpleasant taste in the Giardia (Fig.. cellulose. — ^A distinctive feature of nearly all species of this class is the capacity to form protoplasmic processes called pseudopodia (false which are temporary structures and can be withdrawn. Trichomonas. The representatives of this class include many freeliving forms as well as numerous parasitic ones. 24. Textbook of General Zoology. silicon. published by John Wiley and Sons. — Group of typical Sarcodina. creamy scum. Class Sarcodina (sarkodi'na. which secretes its shell. most common parasitic form. which constructs its . and Difflugia. A number of the repfeet) mal is resentatives of class Sarcodina secrete an external shell of lime. (From Curtis and Guthrie. Arcella Actinophrya Fig. root foot). lakes.) 2. Endamoeba histolytica is the Arcella. Some of the species are naked. number of Mastigophora live in quiet streams. studied. PHYLUM PROTOZOA IN GENERAL 67 A larg-e form. is a fresh-water form which is often found in water supply basins and causes a pungent. This soupy mass of organisms may cover an area of hundreds of Noctiluca is and in the ocean. (a) Amoebina are irregularly-shaped forms with lobelike pseudopodia. Amoeba proteus is the free-living naked form which is commonly chitin. When stimulated. these animals are luminescent and at night frequently give up an attractive greenish or bluish white light Uroglena water. or of the secretions.

shell-bearing pseudopodia are very slender a^d order of shelled forms whose the through small pores The pseudopodia are extended m sheU Only a very few of this group live m fresh water.M iU- e crw^^A^ Parker and Foraminifera PoZysfomeZZa Fig.D. Zoology.68 Shell of TEXTBOOK OF ZOOLOGY secretion.nmcrosphericin^^^^ dividual developing from amoeboid Kearawn gametes escaping from it F.-Life cycle of one of the from it ^ youn^^^ spheric individual. .f.) . commonly observed reticnlar. Haswell. published by The Macmillan Co. B amoeboid 9^11^ .^^P^^^ cell . union of gametes. 25. are two of the most sand cemented together by a (b) Foramimera is an forms. The vast ^^iijA^l^ "s :.

and Glohigerina IN GENERAIv is 69 a typical example. are character- ized as being able to produce enormous plasmodia containing hun- dreds of nuclei and contractile vacuoles. (d) Heliozoa a group with thin. Badiolaria a marine group with ray- . threadlike. B. The Invertebrata. as well as having ability to Shells of different Foraminifera. C. . (c) Mycetozoa. The dis- integrating calcareous shells of this organism constitute a great mass of material on the bottom of the ocean which is known as globigerina ooze and from which chalk is formed. Nodosaria hispida (X18). of is They live quite commonly in masses decaying vegetable material upon which they feed. — reproduce by spore formation. Rhahdamina abyssorum (X4. published by The Macmillan Company. Actinophrys (e) sol is a common one found is in fresh-water fine. streams and ponds. (From Borradaile and Potts. 26. after various authors. Globigerina buUoides (X55). A. radially arranged.) PHYLUM PROTOZOA majority are marine.5) Fig. unbranched pseudopodia.

There are a few parasitic forms. Paramecium. Class Infusoria (infuso'ria. crowded). waters as free-swimming organisms. The pseudo- podia extend through the relatively large apertures in the shell. notably Balantidium coli.— This those single-celled animals covered with small hairlike. cytoplasmic They occur in both fresh and marine processes known as cilia. and Vorticella are -ZT^ffflara 0i^» — .70 like TEXTBOOK OF ZOOLOGY pseudopodia and a shell composed largely of silica. Stentor. group includes 3.

These forms are typically attached by stalks. (d) Peritrichida is an order composed of sedentary ciliates with a whorl of oral cilia continued into a depression in which are located the oral spot and aperture of the contractile vacuole.water genera while In some forms this oral region Stentor. —Representatives from Textbook class Infusoria. of (Reprinted by permission from General Zoology.) most of the genera use creeping as their means of locomotion. There are no body cilia in certain phases of the life history. Vor'ticeJla is probably . 28. Curtis and Gutlirie. fresh. Stylony cilia. a parasite in the intestine of man and some (c) Tlypotrichida possess . possesses membranelles. Kerona is a parasitic form and is often found creeping over the ex- ternal surface of fresh-water Hydra. and Bursaria are common is Balantidnim (Fig. trichida possess a well-developed undulating membrane pharynx. John Wiley and Sons. 389) other mammals. formed by fusion of cilia these are found prinThe cell is flattened dorsoventrally and Prorodon From ton I a iupiom StyhnyoMa Laorymaria i^m lionotw Fig.) (Figure of Frontonia modified. The body cilia are small or partially absent. but the cilia of the oral region are well developed. Halteria.PHYLUM PROTOZOA sitic IN GENERAL 71 (b) Heteroin the cyto- genus which inhabits the large intestine of the frog. At the base of this depression is located the mouth. cirri or structures cipally on the ventral side. Inc. Oxytricha and Euplotes are common fresh-water genera.

72 the most TEXTBOOK OF ZOOLOGY common living genus. . The animals of this class are entirely parasitic ^^(^^''f^^T^ Ep . :.. includes animals that are not ciliated. it is encysted. except during a free- swimming stage which may occur following division or encystment. as is sometimes called. The cyst is dissolved upon entrance to a host and liberates the organisms. The spores are produced by the parent animal stage. i th e Hum i<m^l . — These protozoans there in their early stages are often amoeboid. but in the completed life history locomotor structures are wanting. Class Sporozoa (sporozo'a.Intermediate sta^e -natarc 5ta^e Fig. its — Gregarina attached an epithelial development are shown within adjacent to cell of host's cells. hfH-Early sta§e . are protective and enable them to withstand adverse conditions. multiple fission is followed by gamete formation. is During the life cycle a spore stage. and the gametes fuse to form a zygote. but Podophrya is an example of a fresh-water genus. Intestine. Suctoria or TentacuUfera. stalks while Epistylis is Vorticella and Carchesium have contractile attached by noncontractile branching stalks. 4. dividing into fragments while This class of Protozoa is among the most widely distributed of the animal parasites. it The second subclass. seed animal). stages of 29. Most of them are marine. All Sporozoa reproduce by sporulation in which asexual. These little cysts. which are secreted by the protoplasm of the animal. Other and they are usually transmitted to other animals in the spore They often pass from one host in its feces and enter another in contaminated food or drink or they are drawn from one host by a blood-sucking animal and transmitted to the blood of ajiother. These are attached forms with protoplasmic projections which are used in the capture of food. and their life cycles are often quite compli- . Epistylis and Carchesium are well- known colonial genera.

The life history involves a period of asexual reproduction (schizogony) and a period of so-called sexual reproduction which ends in spore formation (sporogony). but the saclike. The and urinary bladder are usual seats of infection for the free forms. (b) Haplosporidia are single cells. Myxidium and myxoholus are gall-bladder. the The third subclass Acnidosporidia includes forms which produce Again there are two orders: (a) Sarcosporidia. There are two orders (a) Myxosporidia are found chiefly as : fish parasites. have in each spore a single polar capsule. and occasionally vertebrates (b) their early stages. This group parasitic chiefly in arthropods.PHYLUM PROTOZOA cated. simple spores. including man. commonly of earthworms. leeches. cockin roaches. The malaria parasite and the causal agent of Haemosporidia. The first subclass is Telosporidia in which the spores produced have neither a polar capsule nor polar filament. and vertebrates. The encysted forms attain a length of several millimeters. other insects. characteristic genera. Texas fever in cattle are the most important forms. (b) Microsporidia is but occur occasionally in reptiles and amphibia. and they have a relatively simple structure. the spores of which contain from one to four polar capsules each with a coiled polar filament. The complete life cycle is is known not known. while the gills and muscles of the fishes are choice tissues for the cysts. IN GENERAL 73 There are three subclasses of the class. Coccidia are minute monocysted forms which are permanent intracellular parasites of molluscs. fish and amphibia. Cnidosporidia is the name of a second subclass. . but later they may become free in the cavities of the host. and ticks. and occasionally in other invertebrates. and ultimately each becomes a mass of sickle-shaped spores. encysted form in muscles of mammals as Miescher's tube or sac. The former occurs in the blood of the vertebrate and the latter takes place in such hosts as insects. uriniferous tubes. and each of these is divided into some orders. (a) called gregarines. inhabit the cells (cystozoic) Gregarinida (Fig. arthropods. As name infers these occur in muscles of several mammals. In this group are three orders. notably the cockroach. They may attain considerable size. 29). each with a single nucleus. red blood corpuscles of vertebrates. This order parasitizes fishes and certain insects. (c) The representatives of this order live chiefly in the Again the life cycle involves both schizogony and sporogony.

A union of the flagellate microgametes with the egglike macrogametes takes place in the stomach of the mosquito. The new individuals (merozoites) are freed by destruction of the corpuscle and almost immediately enter new corpuscles where repetition of events occurs. and others become micro- If a female gametes which develop from the male gametocyte. The union is commonly called fertilization. forming sporohlasts. caus- ing estivo-autumnal or subtertian fever. These enlarge and motile. The parasite. which causes tertian causing organisms: (b) Plasfever. and is now called an oocyst which grows at the expense of the adjacent tissue. spindle-shaped sporozoites develop within. Inside of the oocyst the nucleus divides repeatedly. it requires twelve days or more for . and there may be 500 capsules in one mosquito. this person. while slender. is characterized by an attack each forty-eight hours. the mosAnopheles mosquito bites and sucks blood from quito becomes infected with gametocytes of Plasmodium. invades the red corpuscles. in which there are numerous nuclear divisions before the mass of cytoplasm divides. This example will illustrate also the relationship of insects to diseaseproducing organisms. and its life cycle will be given to illustrate the intricate life history Life cycles in which there are primary of certain of these forms. The parasite (see Fig. the malaria parasite. the attacks of which recur each day.000 or more. This ookinete enters the wall of the mosquito's stomach where it encysts in the form of a ball with a shell. Some of these merozoites become sexual cells (gametocytes). known as an ookinete. or there may be a somewhat constant fever. coalesce. is characterized by an attack every seventy-two hours. 393) generations may live in the blood of man by which may continue throughout the a series of asexual life of the person. which causes quartan fever. is one of the Haemosporidia. There are three species of human malaria(a) Plasmodium vivax. spoken of as female. Part of the gametocytes develop into macrogametes. This cyst protrudes like a little wart on the outside of the wall of the stomach. modium malariae. each with a chromatin dot as a nucleus.74 TEXTBOOK OF ZOOLOGY Plasmodium. is The capsule of this oocyst crowded full of these sporozoites which may number 10. where it reproduces by a sort of multiple division called sporulation. Depending somewhat on the temperature. while in the spore stage. and a fused cell or zygote thus formed soon becomes a wormlike form. (c) Plasmodium falciparum. and intermediate hosts are quite common among parasites.

of these forms only two cells adhere. In. ' process of asexual multiplication begins over again in the red corpuscles of this person as a new host. Codonosiga. In some exist in Codonosi^a Carchejium Joanne Moore. Eudorina. sporozoites make their way to the salivary where they may remain for weeks. a linear colony Carchesium. giving a branching colony. In certain species the cells are stalked. This formation frequently results from incomplete separation of the cells following division. a simple colony. spherical envelope inside of which the colony of cells remains. Colonial Protozoa There are some species of Protozoa in which the individual cells groups called colonies. These types of colonies are known as spheroid and arhoroid or dendritic respectively. Different types of colonial Protozoa. and the ' ' ' These little parasitic gland of the mosquito mosquito bites a man. a stalked flagellate colony. while Epistylis and Carchesium are typical examples of the latter. with the result that thousands of cells are built into the group. Pandorina and Eudorina are typical examples of the former. and the new cells remain attached to the stalk. some species there is a jellylike. When tliis some of the saliva with sporozoites flows into the wound. within gelatinous envelope Ceratium. (Drawn by . but in others the cells may remain attached after many divisions. Pandorina. Colonies like that of Ceratium with individuals ar- . stalked infusorian colony.PHYLUM PROTOZOA this IN GENERAL 75 development to go on in the mosquito. — . 30.) Fig.

) numbers of cells. (From White. Mosby Company. General Biology j The C. and colonies of irregular arrangement are spoken of as gregaloid. Tlie difference between these colonial Protozoa and simple Metazoa is a difference in the relationship of single cells to the group as a whole and not a simple difference B Fig.several daughter colonies. B. V. In the structure of Metazoa the cells are specialized and in .— 76 TEXTBOOK OF ZOOLOGY ranged in a line form a linear one. Formation of daughter colony by development of a parthenogonidium. A. Volvox. Mature colony containing. In the colony each cell is an independent or almost independent individual so far as the functions of living are concerned. 31.

cells They are and reproductive (germ) classified into general cells. If the animal attracted to the source of the stimulation and turns is toward repelled it. such as Volvox. temperature. f. Taxis may also be used here if the response involves the movement of the organism as a rents. Both are spherical organizations of cells. has not been thoroughly determined in a certain way. Phototropism. although he does draw on it indirectly by a food . so that certain groups carry out a definite portion of the entire metabolism. Thigmotropism. response to gravity Rheotropism. response to light d. response to chemical changes Thermotropism. Tropisms are named with respect to the stimulating agent. body (somatic) Certain of the spheroid proto- zoan colonies. which means turning. and electric currents. either positive or negative.: PHYLUM PROTOZOA IN GENERAL 77 distributed. The simpler ani- mals under a given set of conditions respond to these stimuli in a certain way not because of power of choice. b. response to mechanical currents Electrotropism or galvanotropism. response to temperature Geotropism. e. but because they cannot behave in any other way. The Protozoa are controlled in their behavior largely by tropisms. the response negative. have a rather striking resemblance to the blastula stage in the early development of metazoans. whether plant or animal. chemical change. of all degrees of complexity respond to various kinds of stimuli. Tropisms and Animal Reaction Organisms. The important stimuli which call out immediate or direct response by the animal are light. c. response to electric currents is g. and the common ones usually recognized are a. mechanical cur- The response to a stimulus may be Tropism. whole. is If the It organism is by the stimulus. response to contact Chemotropism. gravity. Economic Relations of Protozoa Man has not yet found a way or need to eat Protozoa directly as food material. The sary to get a response why an animal responds to a specific stimulus minimum strength of stimulus which is necesis known as the threshold. bodily contact. refers to the reaction of an organism to a stimulus. the response said to be positive.

and it can be rather successfully eliminated from human beings by use of such drugs as emetine.78 TEXTBOOK OF ZOOLOGY chain including water fleas. by man for example. in human beings. much of northern South America and Central America was. and fish. at one time. There are other Protozoa that render fertilize the soil. Foraminifera which is an order in class Sarcodina has some economic importance because of the limestone which is formed by the concentration of the material of the dead tests or shells. About 10 per cent of our population are said to be carriers of these organisms. It is about the size of a pinhead. EndEndolimax nana. larger crustaceans. will be carried to the liver where The infection is usually obtained serious abscesses are formed. contaminated with the encysted organisms from fecal matter. so far as amoeba coli. intestine of this. and chiniofon. Most of the diseases of this origin lion. other Amoebae have been found known. and Endamoeba gingivalis are such Some examples. Endamoeba histolytica (see Fig. they are not pathogenic. it sinks to the bottom of the ocean where the mass forms the globigerina ooze which hardens into solid chalk. water unfit for drinking or help Amoebic Dysentery. administered by a physician. Each of its repretheir skeletal sentatives has a complicated skeleton of silica. and as it dies. the protozoans are not classed as predators on man as would be the but many of them are parasites. directly through drinking water or eating food which has been tion. From this is formed quartz or flint. The causal agent is one of the Amoebae. Too. ruled by yellow fever and malaria. are more prevalent in the tropical and subtropical regions of the earth. 391). Technically the disease — . Such diseases may attain sufiScient importance to render . carbarsone. Badiolaria is another order in the same class. Many diseases of man and animals are caused by Protozoa. From remains comes an ooze on the sea floor sometimes hundreds of feet deep. but. man on the inside of the walls of the There results from are caused by this disease. disease of African Sleeping Sickness. and the same applies for large portions of continents uninhabitable sleeping sickness in Africa. A genus by the name of Globigerina is one of the best known members of the group. if From the intestine the infec- allowed to continue. —Ulcers severe diarrhea and dysentery. This malady is the most important man caused by flagellate Protozoa.

388). arsphenamine. extended coma. the chain . and the disease is limited to that area in Africa where this fly is found. Trypanosoma cruzi. and spleen. severe emaciation. It was the first disease proved to be directly caused by a protozoan parasite. Malaria. was able to prove that the female Anopheles mosquito is responsible for the transmission of malaria. one of the true bugs which is closely to our common blood-sucking form. the sporozoan which causes this disease. As early as 1718 a worker by the name of Lancisi ventured the statement that mosquitoes or gnats might transmit malaria however. as well as man. The organisms (Fig. armadillos. it was not until about the opening of the present centuiy that this relationship was understood. cattle. or it may be extended Such animals as antelope. or if aU malaria patients or carriers are thoroughly screened in. lutely fatal. Chag'as' Disease. and some wild game are susceptible to the disease plicates the control of it. has been tried with partial success. This comThe disease has been considered absobut recently a drug. death usually within three or four months. Laveran found a curious parasite in the blood of malaria paSeveral years later Laveran and IManson independently sug- gested that the organism might be transmitted by some bloodsucking insect. and may serve as carriers. has already been discussed under the general topic of Sporozoa. A closely related flagellate. The life history of Plasmodium. After several years more of investigation. anemia. Glossina palpalis (Fig-. monkeys. and disturbance of the nervous system. genus name of the animal that Trypanosoma gamhiense or Trypanosoma rhodesiense. In 1881. If houses are screened to keep out mosquitoes at all times. These organisms are transmitted by the tsetse fly. liver. guinea pigs. The sjonptoms are continued fever. It is transmitted — through the bite of Triatoma. Major Ronald Ross. tributed . swollen lymph related glands. and other organs. The disease is one of the oldest and most widely dis- — among men. Dr. liver. 387) live free in the blood and collect in the lymph glands. the "kissing bug. an Englishman. an arsenic com- pound. The infection will bring about loss of appetite. tients. which ends in causes into years. In final phases it collects and attacks the brain. spleen." Chagas' disease affects dogs.PHYLUM PROTOZOA is IN GENERAL 79 called trypanosomiasis for the it. causes this disease in Central and South America. or if all mosquitoes and mosquito breeding places are destroyed.

and suffering is almost inestimable. The red cormay be reduced from an average of 7. and trachoma should be considered with this possibility. transmitted by the Rocky Mountain spotted fever tick and and unmosquito Aedes comfortable disease. There are many other diseases that are rather similar to the above which may be caused by Protozoa. In some parts of the world they have considerable economic importance. Another means by which the chain may be broken is to cure the carriers by killing all of the Plasmodia in their blood by use of quinine. or by covering the water with a film of oil which keeps out air and smothers the larvae as well as discourages females from laying eggs in such water. fatal to Such diseases as Rocky Mountain spotted fever.80 TEXTBOOK OF ZOOLOGY Mos- of necessary relations for production of the disease is broken. A good protozoologist is one of our most valuable economic assets. a sexual disease of the horse. The disease is transmitted from cow to cow by the cattle tick and its young. ml. . disease in cattle puscle count of the host Nagana. Quinine is a specific drug for this disease. properly administered under a physician 's direction. and surra. quitoes are destroyed serve as breeding places. by draining swamps which by placing mosquito fish (top minnows) in the pools to eat the larvae. — The small sporozoan. Babesia higemina. a very unpleasant smallpox.000 per cu. although the organisms have not been specifically isolated. to less than 1. The cost of the above-mentioned and other Protozoa to man throughout the world in money. scarlet fever.000. Texas Fever. dourine.000 per eu. as well as perhaps rabies. are diseases of domesticated animals and are caused by trypanosomes. . ml. loss of time. causes this by destroying red blood corpuscles. transmitted by the yellow fever (Stegomyia) . similar to African sleeping sickness in all man.000. typhus fever. man dengue or breakbone fever.

and lakes. Food and Assimilation The food problem among Euglenae the biological standpoint. These bodies are called pyrenoid dodies. This an extension of the cytoplasm. sluggish creeks. The mouth of the cell is near the anterior end. Euglenae are sometimes classified as plants by botanists because of the presence of chlorophyll. This genus is also quite well represented among marine animals many Euglenae possess chloroplastids which give them the possibility of photosynthesis. and extending inward from it is the gullet or cytopharynx. the Most of the euglenoid forms have spiral markings (striations) on the surface of the body. Small contractile vacuoles empty from the endoplasm into the reservoir. Bodies of collected protein material may be seen in eoiuiection with most of the chloroplasts which are distributed through the cytoplasm. almost transparent. It is a form which illustrates certain plant characteristics and animal characteristics in the same organism. At is the anterior end. Just anterior to this is the stigma. They are usually found living in the surface waters of ponds. It is usually obliterated from view by the abundant chloroplasts. to It as a group is interesting from seems that some Euglenae are able ingest other small organisms through the mouth and cytopharynx 81 . whiplike filament. which is red in many individuals of E. . bears a very slender. Within the inner portion of the cell or endosarc (endoplasm) is located the nucleus. The superficial layer of the cell or covered by an extremely thin portion.CHAPTER V EUGLENA OF CLASS MASTIGOPHORA Habitat and Characteristics The most common species are Euglena viridis and Euglena gracilis which are found abundantly in fresh water. attached near the mouth. Structure The microscopic. single-celled body has about the shape and proit portions of a cigar with a blunt anterior and a sharp posterior end. viridis. the flagellum. Beside the cytopharynx is the reservoir or large vesicle. (ectoplasm) is ectosarc cuticle.

gracilis. dissolved mineral and with the aid of light and chlorophyll builds up organic food substances. 32. ^^ Ctirom atopt) ore A\^ 1^5 -Nucleus Fig. by absorption through the general cell In fact. are able to surface (saprophytic nutrition). '§ ©. assimilate dissolved nutriment Others. carbon dioxide. The final stage of the carbohydrates formed by this . E. this has been called holozoic nutrition as typical of animals. like E. Those forms plant. Vacuole ^Reservoir )<.— 82 to TEXTBOOK OF ZOOLOGY be digested in a vacuole within the endoplasm. viridis probably does not possess this possibility. E. Euglena like viridis Ehrenberg. salts. A chlorophyll-bearing flagellate. viridis that are abundantly endowed with chloro- phyll obtain their food largely by photosynthesis as does the green This process utilizes water. -Fhgellum 'Mouth -Stigma Contractile :0'». this species has been maintained for more than two years in a nutrient solution in darkness.

such as drought or increased chemical concentration. E. . a permanent with the exterior. cyst. There may be some utilization of the carbon dioxide produced in the metabolic activity by the process of photosynthesis in forms where it exists. in a quiet but not encysted condition in other species. but all are represented in closely related species of these flagellates. Grains of this substance may be observed throughout the endoplasm of these Euglenae when living in favorable conditions. Some observers have reported as many as thirty-two young flagellated individuals coming from a single cell. while a new flagellum is developed by the other. It is not likely that all three of these fundamental types of nutrition are found in any one species of Euglena. favorable conditions these cells emerge from the cyst and assume the active phase. some of the excess oxygen produced by photosynthesis may be used in metabolism. division takes place. On side rare occasions two individual Euglenae come together side by and fuse permanently into a single This is somewhat similar to the zygote formation in sexual reproduction. Upon the return of normal. This division occurs only in the motile state (or active phase) in some species. but there is also some rather authentic work which shows that the old flagellum entirely disappears during division. Likewise. viridis may divide by longitudinal binary fission in either the motile or encysted condition. and secretes a thick gelatinous envelope about itself. vesicle communicating Reproduction and Life Cycle Binary longitudinal fission is the common means of reproduction. According to some authors the original flagellum is retained by one-half. During adverse conditions. In this condition it becomes spherical in shape.EUGLENA OP CLASS MASTIGOPHORA process is 83 paramylum. fission occurs only while encysted (encysted phase). There may be a single division or there may be several. a granular substance much like starch. Euglena becomes encysted. and in a few others. "Water and waste products collect in the several small contractile vacuoles which empty into the reservoir. During the encysted phase. *o^ Respiration and Excretion Respiration is carried through the general surface of the cell membrane. nonmotile. and a new one is developed in each daughter cell.

through the water. When it encounters such a condition in the medium. This animal displays positive phototropism and is easily stimulated by changes in intensity of light. These movements resemble waves of contraction (peristaltic contraction) passing over the cell. reactions this cell exhibits the irritability that is characteristic of all protoplasm. This activity is known as euglenoid movement. and when in the active phase swims about. it stops and turns sharply in another direction and attempts to move out of danger. There is naturally an attraction to light in those forms which utilize Euglena usually lives it in the manufacture of food by photosynthesis. is injurious to them. This is known as the avoiding reaction. Some move about in a crawling fashion by taking advantage of this movement. intense When Euglena swims however. The flagellum . spiral path is followed due to the continuous turning of the body. In these and other sunlight. of the larger species is made up of an elastic outer sheath which encloses an axial filament composed of one or more contractile fibrils. its anterior end with the flagellum goes foremost and is first to reach any injurious or distasteful environment. will be a negative response. Direct. The chief method of locomotion is swimming by means A of the whiplike movements of the flagellum through the water. there A medium light is optimum for it. If the light is too intense.84 TEXTBOOK OF ZOOLOGY Behavior near the surface of the water if the light there is not too intense. Locomotion and Flag'ellar Movement are not actively Contractions and expansions take place in Euglenae when they moving about.

such as debris from watering troughs. the outline is con- stantly changing. in streams where the water runs over rocky ledges. Recently Chaos chaos Schaeffer has been rediscovered. The many kinds soil. and wherever is face of submerged lily pads. lustrous. If amoebae are present. mass of gelatinlike substance with slowly-mov- ing. It is often found on the surA mass of pond weed may be brought into the laboratory in some of the pond water and allowed to stand in the container a few days. they will likely be in the brown scum which forms. Amoeda proteus may be collected in a variety of places where conditions of water. bottoms of ponds. drain ditches. temperature. Amoeba is recognized a simple and low form of life. there spring pools. some simpler and some more complex than Amoeha proteus. fine particles within. It is enormous in size and can be seen with the unaided eye. Chaos diffluens is a very desirable species for study." The pedigree of probably as long as that of any of the animals we know and involves hundreds of times as many generations as many of common animals . Even the writers of fiction speak of the range of the span of complexity of Amoeba the as extending "from Amoeba to Man. and organic food are favorable. or as parasites in the fluids of the visceral organs of higher types of animals. irregular The general appearance of this animal is that of a slate-colored. abandoned tajining pits. or in the sediment at the bottom. .CHAPTER VI AMOEBA OF CLASS SABCODINA It is likely that no microscopic organism has attracted so much as attention and popular interest as by the public generally animal life is Amoeba. Little or nothing is known about the real ancestry of Amoeba. abundant aquatic vegetation. yet Amoeba remains in a relatively primitive and simple state. Characteristics and Habitat marine water. There are many kinds or species of Amoeba. of Amoeba live in fresh water. When S5 it is active.

the plasmagel. The firmer.). endosarc (endoplasm). and crystals. PSEUDOPODIUM ECTOPL. is about M. the inner more fluid. These . passive portion. is much more granular and contains the cytosome. of a living Amoeba inclusions as well as the nucleus. Under favorable conditions the protoplasm can be differentiated two portions. plasmasol in which the streaming movements take place and.ASM ^i^ Fig. The nucleus usually appears somewhat dense and granular.00 iJ^ch (0. vacuoles containing watery and varying in size. Metabolism This refers to the constant building up (anabolism) of living protoplasm and its concurrent oxidation (eatabolism). surrounding this a more viscous. and is located in the portion away from the end which is advancing in a moving specimen. various granules. the ectosarc (ectoplasm). and they are constantly changing in shape in the active animal by the flowing of the protoplasm. somewhat tougher outer portion. It includes all activities necessary for maintenance of itself and its race.25 mm. —Drawing to show the appearance and structure proteus. while its extreme diameter is Y^o inch or barely visible as little specks to the unaided human eye. mitochondria. is nearly homogeneous and includes the plasma membrane (or plasmalemma) the more fluid inner portion. 33. fluid single. water vacuoles. shiny.'©**« V-''^^S? -^ ^U". fat globules.86 TEXTBOOK OF ZOOLOGY Structure Amoeba Its proteus is one of the largest of the fresh-water forms. The larger bodies contractile in the cytosome are food vacuoles. Some authors distinguish two types of protoplasm in the endosarc. The animal owes its irregular shape to the fact that protrusions of its own substance are formed at its surface. cell into : 'fj VACU01_E FOOD I ^^ 1 " '^T^'- I Z-^Z-Z^^^> ^ '^ VACUOLE NUCl_EUS >-* f '. average diameter These are known as pseudopodia.

(Redrawn by permission from Wolcott. A circulatory system is not necessary since the . Here we may study the metabolic cycle in progress within the confines of a single entire Its cell.) — Digestion. Bacteria may Amoeba has no definite mouth but the food is taken body by engulfing it at any point that comes in contact with it. phases are as follows: Food.AMOEBA OF CLASS SARCODINA 87 of life phenomena are the same as those found in the highest forms but reduced to very simple terms. — into the E^ejfion xcretion Fig. and portions of filamen- be used to some extent and rotifers (small Metazoa) are sometimes devoured. 34. is The function of digestion complex materials into a soluble. These vacuoles move about in the endoplasm. A pseudopodium is formed at this point. since enzymes serve this purpose in larger animals where exact study can be made to convert is on the process. Diagram to show the phases of the metabolic process as it occurs in amoeba. A droplet of water is included with the food to form what is called a food vacuole. — Its prey consists chiefly of smaller Protozoa. absorbable form. tous algae. It assumed that the surrounding cytoplasm secretes enzymes into the food vacuoles of Amoeba to perform this function. Ing-estion. published by McGraw-Hill Book Company. small single- celled plants. such as diatoms and desmids. and the end of it flows around the food particle until the particle is entirely enclosed. Anional Biology. — The food gradually disintegrates and much of it goes into solution in the fluid of the vacuole.

The chemical union of the oxygen with the organic substance of the protoplasm liberates kinetic energy and heat. most of the protoplasm of the cell is in rather close contact with the dissolved food. may The contractile vacuole likely assists in discharging CO2. are discharged from the body of Amoeba by way of the conThe contractile tractile vacuole along with some carbon dioxide. In Amoeba this exchange is carried on primarily through the general — body surface.88 TEXTBOOK OF ZOOLOGY vacuoles with the food in the process of digestion circulate so widely in the endoplasm that all parts of the cell may receive nourishment by direct absorption. — This is a process whereby the gas. which are protein by-products. Catabolism or Dissimilation. (CO2). Amoebae. In Amoeba the digested food material is absorbed directly from the food vacuoles by the surrounding cytoplasm. Urea and uric acid. The water in which the animal lives must contain go on. Water. It fills out with liquid which is forced out through the membrane as the vacuole disappears. and salts. leaving the protoplasm. carbon dioxide Respiration. —Indigestible material or is debris that has been ingested with the food carried to the surface of the cell egested by simply being left behind as and cast out or the animal moves away. must be disposed They cannot be allowed to accumulate beyond certain limits in the living organism if life is to continue. The contractile vacuole is absent in some forms. is exchanged for oxygen (O2) entering it. Such a process is essential to all living protoplasm. Its location apparently is not fixed in the cell but is often near the nucleus. Since the vacuoles move rather generally through the endosarc. This is the process of transforming the digested Assimilation. food material into protoplasm. urea. and in such cases. however. This is known as oxidation and is a burning process which goes on within the protoplasm. are able to and do live in rather foul water where the oxygen content is rather low and the carbon dioxide high because Amoebae may live several of the decaying vegetation present. ex- . Egestion. vacuole is formed by the union of small droplets of liquid under the plasma membrane. some mineral matter. — and carbon dioxide are residual products of Excretion. hours in water from which the oxygen is removed before asphyxiadissolved oxygen in order that this diffusion tion occurs. —These by-products of metabolism in the form of waste of. liquids this process. excess water.

growth occurs when the rate of anabolism exceeds the rate of catabolism in the organism. Reproduction and Life Cycle The adult. material In all living organisms growth is is accom- by addition to the protoplasm. (Drawn by Joanne Moore. and likewise all waste substances must be discharged in a similar manner. some excretion by this means in all Amoebae. 35. A. We do know of certain relationships it.) is difficult to maintain cultures in perfectly normal conditions for Ordinarily. attains a cer- when this limit of size has been reached growth ceases. There is Growth. Beginning of the process. is added to the protoplasm than In food plentiful. animal grows when conditions are favorable until tain size. It will be recalled that all materials used by a must pass through the cell membrane. or a man take on the proportions of an elephant? We have not been able to put our fingers on any one factor that completely controls growth. Mathematics states that the volume of a cell increases according to the cube of that influence cell . plished — If there is increase in the volume If is of a body. Why does the cell cease to grow? Why should it not attain the size of a man? Or why should a tree not continue to grow until it reaches the sky. through In Amoeba the cycle is likely only partly embryo state. this is spoken of as growth. -Diagram fission to show fission in amoeba. B. nearing completion.AMOEBA OF CLASS SARCODINA cretion occurs only likely 89 by diffusion through the cell surface. other words. because it Nucleus Contractile iracaole Fig. to known. life history of the niajiy-celled animals to be studied later includes a series of changes from egg. the it sufficiently long periods to get this complete story. more used up in the oxidation which produces active energy.

cells /Nuclear fragment of sporu- —Diagram to show amoeba encysted and undergoing the process lation. fission is repeated. then. states has been definitely established. and out of a cell. fission into two daughters.) that Amoeba starts out as a tiny pseudopodiospore It which has only It one pseudopodium. to assume that a the surface area will not be sufficient may be reached when for the passage of necessary materials into is. curs. an authority on Protozoa. a division oc- Binary by which two new individuals are produced. Occasionally the fission seems to be an amitotic one. 36. size of the When Amoeba fission. the amount of material in cell increases surface needed to surround approximately twice as fast as the plane it. (Drawn by Joanne Moore. and some other methods of reproduction have been presented. while its its surface increases only according to the square of a growing point diameter. then passes through a growth period and it increases in complexity until reaches the full-grown condition. When grown to nearly twice its original size. reaches the limit of size. metabolism certainly would be factors affecting the organism. considerable variation in the size of cells. It is logical.90 its TEXTBOOK OF ZOOLOGY diameter. It is felt that the complete details of the life cycle of many common Sarcodina are not yet available. hence seems there must be other factors besides volume and surface relaModified surface and difference in the rate of tion in operation. During the en- then divides by binary ter has each daugh- cystment the protoplasm undergoes several divisions to produce the several pseudopodiospores which later break from the cyst as infant Amoebae. there is a period of encystment and subsequent sporulation. . Environmental conditions and the variety of Amoebae determine the number of times this phase is repeated. Calkins. New Pig. In other words. There it however. At the close of the fission phase.

It is so named from the perfect exemplification of such activity by Amoeba. Amoeboid Movement and Locomotion The flowing or streaming of the protoplasm and extending the cell some direction by the formation of pseudopodia is usually called amoehoid movement. these activities are "comparable to the habits. Amoeba reacts negatively to concentrated salt. and the other simple sensations. An environmental change to which an animal reacts is known as a stimulus. while the reaction of the animal is called the response. and join so as to enclose the object. ingestion of food. and others. and the process of their formation in most Amoebae." The activities under discussion include the formation of pseudopodia. Amoebae have an optimum temperature range between 15° and 25° C. desire. The edges of the pocket fold in. Temperatures approaching the freezing point inactivate the animal. acetic acid. (86° F. pleasure.) also retard their activities and may soon become fatal. Locomotion is accomplished by the pseudopodia. reflexes. Amoeba exhibits all of the tropisms discussed in Chapter contact. while temperatures above 30° C. According to Jennings. locomotion. This attraction to food is likely a positive chemotropism. The movements made by an animal in response to stimuli are called tropisms. meet. A and automatic activities of higher animals. negative. gentle To physical . hunger." He also feels that Amoeba probably experiences pain. cane sugar.AMOEBA OF CLASS SARCODINA Behavior 91 All of the activities of an animal which come in response to in- make up the "behavior. responds positively is if the impact is otherwise the response its strong light and finds responds negatively to optimum in a moderately reduced light. It When some part of the body surface of this animal comes quietly into contact with food. weak electric current has aai effect on the physical condition of the protoplasm on the side nearest the cathode. in . This part of the protoplasm stops flowing while other parts continue. The tendency is toward the sol state here. hence the animal turns toward the cathode. who has done extensive research on behavior of Protozoa. there is a characteristic response. it IV. thus form- ing a pocket around the particle of food. Amoeba proteus ternal or external stimuli of the animal exhibits either positive or negative reactions to various stimuli. and many other chemicals which have been tried.

the cell membrane (plasmalemma) moves upward and over the upper side of the body it continues to move forward to the tip of the Successive pseudopodia are formed in the it goes in a given direction. Most zoologists explain this movement as being due to the contraction of the more viscous ectoplasm. mounted some of the animals on the edge of a slide in by the projecting edges of two cover glasses and observed their movement from side view by tilting the microscope to a horizontal position. pushing the advancing At the opposite side. The new pseudopodia are formed at the advancing margin of the cell. the more fluid eudoplasm (plasmasol) which causes an outflow at points where the ectoplasm is thinnest. . Dillinger a groove formed .Particle -"" Pseudopodium i Fig. As this plasmasol approaches the advancing tip of the pseudopodium. .) — pseudopodium where it dips down and is laid on the substratum over which the animal is moving and becomes a part of the stationary portion. 1906. one or more may be developing while others are receding. Successive positions in the movements of an amoeba viewed from the side. the back through the centers of the pseudopodia toward the main mass. is In the latter. Temperature and other environmental facflow of plasmasol tors affect the rate of locomotion. Journal of Experimental Zoology. (Modified from photographs by Bellinger.92 TEXTBOOK OF ZOOLOGY moving Amoeba proteus as The pseudopodia are temporary locomotor structures. He describes their movement as a sort of walking on the progressively forming pseudopodia. If the specimen has several pseudopodia. it turns to the sides and changes to more solid endoplasm (plasmagel). gel continues to become plasmasol to provide for fluent material. 37. or where surface tension is lessened.. At the side of the animal away from the advancing pseudopodium. Notice the formation of new pseudopodia and tlae engulfing of the particle on the surface. the plasmatip farther and farther forward. particularly in This brings about a forward movement in the "posterior" region. This process continues.

The groove usually extends obliquely from right to left in P. Paramecia are easily cultured by collecting some submerged pond weeds and allowing them to stand in a jar of the pond water for several days. The is blunt but generally narrower. Or some natural creek or pond water may be placed in a jar with some old dry grass and allowed to stand about ten days. It has a definite axis and permanent anterior and posterior ends. which leads to the gullet. anterior portion.2 and 0. Occasionally cultures are found in which the majority of the individuals show the groove extending from left to 93 . represents the sole portion. It is broad and shallow anteriorly but it becomes narrow and deeper as it ends in a mouth.CHAPTER This animal has been the subject of Yll PARAMECIUM OF CLASS INFUSORIA much study and the victim Paramecium caudatum is probably the species most commonly studied. just about large enough to appear as small white specks in the water. ranging between 0. which passes diagonally from the anterior end to about the middle of the body. caudatum as the animal is viewed from the oral side. the oral groove. Structure Paramecium the heel part . They tend to congregate at the surface and particularly in contact with floating objects. cigar-shaped animal. or ponds polluted by sewage. of considerable experimentation. which sometimes described as being slipper-shaped. which is generally broader is but pointed. but it is asymmetrical in shape. These animals occur abundantly in any water which contains considerable decaying organic matter.3 mm. in length. This animal is a great favorite in zoology laboratories. where they frequently form a white scum. They thrive in all streams. At one side is a depression. It is easily available and is large in size. Characteristics and Habitat Paramecium is an active. creeks. represents while the posterior portion.

tough. The body is covered with fine hairlike cilia which are of even length except in the oral groove and at the posThe cilia within terior extremity. are filled with a semifluid substance and each opens to the outside through the pellicle. The endosarc. structures. 38. There are a great many spindle-shaped cavities located in the These ectosarc with their long axes perpendicular to the surface. nongranular ectosarc composed of ectoplasm. the gullet are fused together into a sheet. composed of The . where they are noticeably longer. CON-r RACTIL. trichocysts. C.94 right TEXTBOOK OF ZOOLOGY from this view. much enlarged. elastic cuticle or pellicle which is marked in hexagonal areas by the distribution of the cilia. forming the undulating membrane. —Diagram cell is showing the structure of Paramecium. by T. The cilia are direct outgrowths of the ectosarc.US CONTRACTILE VACUOLE TRICHOCYSTS PEI_l_ICl_E Fig. Evans. The outer surface of it is a thin.) (Drawn which is divided into the outer.E VACUO l_E ORAl_ GROOVE MACRONUCI_E.

Food. much smaller. — This animal hunts food. bacteria. and particles of debris conon the menu for Paramecia. The vacuoles circulate through the endoplasm in a rather definite course. differing only in certain details. 39. aurelia. The micronucleus is located in the curved surface of the macronucleus and is called cydosis. These same vital functions must take place in all living things (organisms). food vacuoles are formed. it settles down and becomes relatively . This activity is two contractile The numerous the inner end of the gullet The contractile vacuoles are located near each end Each vacuole has several radiating canals entering it. Metabolism The same general activities as described in Amoeba and others occur. Ingestion. The macronucleus is located slightly posterior to the center and somewhat beside the mouth.PARAMECIUM OF CLASS INFUSORIA endoplasm. It is relatively large and rather bean-shaped. Oral qroove J^l^ Nucleus ^ % — Mouth Gullet if M Anus -Food vacuole Fig. These vacuoles expand and contract alternately. vacuoles. a process similar to that described in Amoeba. macronucleus. one at a time. stitute the principal items — Smaller protozoans. another species. It contains food vacuoles. —Cydosis in Paramecium. at by a mass of food material coming in with a droplet of water. and other granular masses. and when locates a its it re- gion where food is abundant. ordinarily has two micro- nuclei instead of one. showing the course of the food vacuoles through the endoplasm while digestion is in progress. is 95 within. P. of the animal.

These food vacuoles move in a definite course through the endoplasm. Finally it passes by means of the action of the undulating membrane into the endoplasm in the form of one food vacuole after another. TEXTBOOK OF ZOOLOGY swept through the oral groove by the beating and carried back through the mouth into the gullet. occurs in the cytosome. caudatum through a long series of generations and observed that conjugation occurs at intervals of approxi- . caudatum. the circula- The food is action of the cilia. twice a day. both the macronucleus and the micronucleus divide. Dig-estion. During this division process in P. Under favorable conditions the storage of nutrient materials. Respiration. Nutrition in this animal is holozoic. and one after another. aurelia and others seem not to conjugate. caudatum is a conjugating form of Paramecium. and its living process is essentially like that of all higher forms of animal life. Since this course is in the form of a cycle. or it may occur to some extent by diffusion through the entire cell membrane.000 individual paramecia. Excretion of the waste products of metabolism in solution of the alternate filling means Growth occurs as it does in Amoeba and in all other organisms.96 quiet. It has been estimated that if all survived and reproduced at a normal rate. depending on environmental conditions. Assimilation. but the division of the macronucleus is not distinctly so. is by and expelling of fluid by the two contractile vacuoles. Division is repeated at least once each twenty-four hours and under especially favorable conditions. Reproduction and Life History itself is asexual. the descendants of one individual over a month's time would number 265. The time required for the completion of a division ranges between thirty minutes and two hours. tion is known as cyclosis. the old gullet divides into two. while P.000. The micronucleus divides by mitosis. Calkins carried some cultures of P. this is repeated for long series of generations. and two new contractile vacuoles are formed by division of the old ones. Egestion occurs at a definite anus. like starch and fats. Conjugation is a temporary union of two individuals with exchange of nuclear material. The actual reproduction is by transverse binary fission which in The cell divides transversely into individuals. P. and Catabolism or Dissimi- lation all occur in a manner very similar to that described for Amoeba.

This union resembles a sexual act and has recently been described as such. fission. The degeneration of the three micronuclei is compared with reduction division in maturation. they come in contact. with their oral surfaces together. Following this comes the long series of generations formed. At least it is known that four of the eight nuclei enlarge and become macronuclei. by transverse binary . These daughter cells then divide to produce a total of four Paramecia which have the typical number of one micronucleus and one macronucleus of the active phase. The two conjugants now separate. This micronucleus divides. and these four divide to form eight. three of the others degenerate. The descriptions of the subsequent events vary somewhat. three of which disintegrate. At regular intervals of about every forty or fifty generations. the macronucleus A . It occurs in a single individual. as the "female. forming two micronuclei. phenomenon. one after the other. and the fusion of the small "male" micronucleus with the larger "female" micronucleus of the other conjugant is compared to fertilization. while the macronucleus undergoes disintegration and final disappearance. The micronucleus enlarges and divides. rather unhealthy appearing individuals. A protoplasmic bridge is formed between the two individuals. and one remains as a micronucleus. forming one large and one small micronucleus. usually small. The conjugants are Shortly after the adherence of the conjugants the nuclei of each undergo changes. forming four nuclei in each animal. but the fourth divides again. When two paramecia are ready to conjugate. has been found occurring in P. Each of these two new micronuclei again divides to form four. known as endomixis. This species has two micronuclei and one macronucleus. The whole series of changes involved in conjugation has been compared to maturation of germ cells and fertilization in sexually reproducing metazoans. and almost immediately the entire animal divides by binary fission with two macronuclei and one micronucleus going to each cell. aurelia by Woodruff. Each individual now has a fusion nucleus. and very shortly the fusion nucleus of each divides by mitotic division each of these divides. Sometimes the smaller of these nuclei is spoken of as the "male" nucleus and the larger.'' In each animal the smaller nucleus moves across the protoplasmic connection to the other animal and fuses with the larger nucleus there.PARAMECIUM OF CLASS INFUSORIA 97 mately every two hundred generations. and adhere in this position.

— Conjugation and subsequent divisions in Paramecium. 40. of outside. The divisions for micronuclei actually occur (From "White. . showing activi- ties of the micronucleus.98 TEXTBOOK OF ZOOLOGY Fig. The shaded ones have been resorbed. as figured for convenience. within the cells instead treneral Biology.) Circles are micronuclei and crescents are macronuclei.

one of the remaining micronuclei goes to each. Endomixis may occur in P. and then the cell divides. . caudatum also. and two remain as micronuclei. Endo- mixis seems to have about the same effect as conjugation. This nucleus then undergoes two divisions. each with two micronuclei and one macronucleus.PARAMECIUM OF CLASS INFUSORIA 99 disintegrates. the typical condition for this species. and the micronuclei undergo two divisions which produce a total of eight. Six of these disappear. The micronuclei then divide again as the entire cell divides to form daughters. Two of these four become macronuclei.

The re- positive. placed in moving water. a.V.) — to sodium chloride.^-_f . and sponse to contact is to electric current.". ^ - s 1 ^ / /^ a ' ^ 9 " JS' rj. positive to weak acetic acid. uniformly through the water. and positive to the negative pole of a weak.-' Be- ss- Reactions of paramecia to temperature. trough with temperature at 19° C. however. are somewhat similar to those described for change in temperature. contact and other minor reactions. In & the temperature is held at 26" C. change of chemical constitution.-". In c. the animals will swim upstream. The animals are generally scattered. and the Paramecia are congregated in the region of higher temperature. It reacts either positively or negatively to contact. Paramecia are in a Fig.Y=iV. galvanic electric current. negative to ultraviolet light. swing on posterior end to a slightly different . at the other. responses to food spiral course in locomotion.-?-. published by The Columbia University Press.). The optimum temperature for Paramecium ranges between 24° and 28° C. This animal to material. 42. . it back away. The animals are collected in the end of lower temperature. it seems not to be affected by ordinary light. negative ——7 = 9 ^ ^ :. Gravity causes the anterior end to point upward. at the left end and 38° C. and when If Parain contact mecium comes will with a solid object when its it is moving. at one end and 10° C.-_. 19- pSn^-". (From Jennings. (71° P. at the other. Its reactions to stimuli Amoeba.' . to gravity. Hehavior of the Lower Organisms. response avoiding reactions."-:-t.100 TEXTBOOK OP ZOOLOGY Behavior is an active swimmer and necessarily shows ready Its behavior consists of its environmental factors. the temperature is 25° C.

i Fig. published by The Columbia University Press. This may be repeated in all directions. Behavior of the Lower Organisins. The same type of persistence is practiced in attempting to surmount a solid barrier.) — . This the "avoiding reaction/' may be repeated.PARAMECIUM OF CLASS INFUSORIA direction 101 and try again. (From Jennings. Diagram of the course and movement of Paramecium through the water. These animals are constantly sampling the water and avoiding the conditions which are least favorable. and is known as Such a reaction really involves simply one or more negative responses. 43. Notice the spiral path.

is will entangle many of the aquatic enemies of these animals. In an effort to defend itself when severely irritated. The reverse stroke is much more passive. substance on one side. . Paramecium will discharge the contents of the trichocysts. Each has an Contraction of the protoplasmic elastic sheath and a fibrillar core.102 TEXTBOOK OP ZOOLOGY Such successive attempts to gain the result desired constitute what known as the "trial and error'' mode of behavior. The cilia are contractile outgrowths of the ectosarc. Paramecium may reverse the direction of the stroke of the cilia and thus move backward just as a car can be thrown in reverse. which harden on conThese threads tact with the water and form a mass of fine threads. Locomotion The beating action of the cilia against the water serves as the means of locomotion. The total effect of these activities causes the course followed through the water to be that of a spiral. bends the cilium in that direction. The movement of one tier of principal cilia seems to stimulate the adjacent ones to bring about coordi- nated. rhythmic ciliary activity and movement. The stroke of the cilia is rather oblique and this coupled with the increased length of the cilia along the oral groove causes the body to turn on its long axis while swimming.

its In return. In the most complicated protozoan colonies di\^sions. such as Gonium. There are several genera of animals which are intermediate between Protozoa and Metazoa. and Volvox. Eudorina. may remain attached after many colonies may range from two to two cells.CHAPTER VIII METAZOAN ORGANIZATION All animals whose bodies consist of few or as a unit are called metazoans. certain individual cells become more proficient in doing certain kinds of work. like primitive man. is largely independent of fellows. doing for itself all that is neces- sary to carry on living processes. Pandorina. for other functions. society this becomes In human more and more complicated as 10. Since Metazoa are more or less like compound Protozoa with some degree of intercellular differentiation. In the many-celled animal. it is thought by many authorities that they arose through organization of single-celled organisms. other special groups care the products of In this way each exchanges labor for the products of the labors of the other groups. only two cells adhere together after cell division. Among the single-celled animals each its cell. a special group able to care for a particular function necessary to the life of the entire organism. but a series of this type shows the possibility of the relative complexity of different colonial forms. as in a highly developed society of men. of many cells functioning In most respects the vital activities Metazoa are similar to those of Protozoa. The organized and classified morphologically as well as physiologically. there may be several different types of The representatives of class Mastigophora are the most likely ancestral forerunners Metazoa. Pleodorma. but in others the cells The size of different thousand similar cells. is and as a result. but for the most part the two groups are fairly distinct.1 civiliza- . General Characteristics This group includes cells are definitely all of the strictly many-celled animals. of In some forms compound or colonial Protozoa. are rather plantlike in characterof istics. The colonial forms. There is a well-regulated division of labor.

When the germ cells reach maturity. ovum of the female. They develop in the reproductive glands or The protoplasm of these cells is known as germ plasm. —Typical germ cells. and so on. so zoans. The female germ cells are eggs or ova. TEXTBOOK OP ZOOLOGY it is Avith development of complexity in metaAnother characteristic of Metazoa is the presence of a definite center of control localized in a particular group of cells which becomes the nervous system in higher forms. The germ cells are set aside early in the life of the individual for reproductive purposes. which cells. Some become protective surface cells.104 tion advances. making B Fig. A. still others specialize in excretion. Certain groups of cells The entire metazoan body is usually divided into germ cells cells. perform a and for other parts to perform other functions. . 44. are specialized for reproduction. The complexity of Metazoa is not the result of great complexity of the individual cells. become specialists in a particular phase of the metabolic activity. but it is due to the special differences between them. The presence of a variety of cells within one body is spoken The modification of metabolic of as intercellular differentiation. they become separated from the body and may give rise to a new generation. spermatozoa of the male. is it seen fair development of intracellular difpossible for one part of a cell to particular function. others secrete special enzymes. gonads of the two sexes. and somatic or body which compose the remainder of the body and are grouped in layers. and those of the male are spermatozoa. Cellular Differentiation In Protozoa there ferentiation. B. activity is the basic factor in the development of all differentiation.

The entire living mass of the metazoan animal body may be classified under five fundamental (four by some authors) kinds of tissues. known and when the is as somatoplasm. connective or supporting. Cellular Organization The simpler Metazoa are composed of only two kinds of somatic cells. . A tissue is an organization of similar cells into a group or layer for the performance of a specific function. protective or covering. this protoplasm is spoken of as being immortal. Potentially. These classes of tissues are: epithelial. this type of tissue can be classified as protective epithelium. circulatory. . —A sheet of cells that covers external or in- is known as an epithelium. glandular epithelium. . a rather wide variety of cells has been produced. The epithelium which covers external surface of an organism usually de- ternal surfaces of the body velops various protective structures in the different groups of animals the hard. These cells are grouped according to kind in two layers. immortal. The glands of the body are developed from epithelium. intercellular substance is characteristic of A certain amount of most tissues and enhances their usefulness. gives rise not only to the protoplasm of the germ cells of the new individual but to the somatic In Protozoa the entire material of the individual is passed on to the two offspring and. according to this idea. muscular. and sensory epithelium. The epidermis or outer layer of the skin and the layer of column-shaped cells lining the inside of the intestine are good examples. This all individual dies. The germ plasm. irritable or conductive. homy chitin of insects scales of fish homy plates : . the In generation. Epithelial Tissue. With advanced differentiation. somatoplasm serves as a means of conveyance for the germ plasm through the current final analysis. nervous. The protoplasm is of the somatic cells rebuilt with each generation. Secretions . for this reason. there is nothing left. sus- tentative. germ plasm is likewise cells as well. and scales of reptiles feathers of birds hair ajid nails of mammals. vascular. and when it is so distributed.METAZOAN ORGANIZATION 105 About forty years ago Weismann presented the idea of the continuity of heredity from generation to generation by way of the germ plasm. contractile. of the somatoplasm perishes. According to function.

and much of the dermis of the skin are composed of connective tissue. semisolid matrix or intercellular substances. contain enyzmes for digestion of food. highly modified in arrangement. The outer membrane of a muscle cell is the sarcolemma. The smooth involuntary muscle is composed of individual. supply regulatory substances directly to the blood. except that the matrix has been replaced by a heavy deposit of calcium phosphate and calcium carbonate. as found in the wall of the heart. in composed of slender cells with is an abundance of intercellular material. spindle-shaped (fusiform) cells. Connective tissue is. are In crayfishes and grasshoppers the supCartilage is porting tissue is chitin instead of bone or cartilage. up the framework of the body called supporting tissues. all tissues whose func- tion is to bind together or support the various parts of the body. voluntary muscle is made up of large multinucleate (many nuclei) fibers. each composed of many fibrils (myofibrils) along which are evenly distributed dense and light areas. There is a single oval nucleus. This is distinctive because of contract and in that way produce movements. . Bone is some- what similar. voluntary. as found in the muscle of the arm and striated. com- posed of scattered cells interspersed with abundant. The cardiac involuntary muscle is said to be made up of individual cells. centrally located. Sustentative Tissue. of which the "hamstring" is a good example. —This type comprises most cases. Cells adapted to this function are more or less elongated and fiberlike. as found in the wall of the intestine striated. two cells. serve as poison to other animals. Tendons. inSkeletal. The scattered its ability to cells are present as bone Muscular or Contractile Tissue. . and some are repellent to enemies. Bone and cartilage.106 TEXTBOOK OF ZOOLOGY from these various glands lubricate the surfaces. solid salts. The definition of cells in : — . voluntary. There are three types of muscular tissue smooth. homogeneous. granular. cardiac. the tough cords that connect muscles to bones. involuntary. skeletal. which make and support the other tissues. which give the general appearance of stripes across the cell. This tissue almost uni- versally present in the various organs throughout the body. and nonstriated. because the dense areas on the adjacent fibrils come at the same level. the cytoplasm of which is largely myofibrils but without striations and therefore smooth.

sec. branched and arborlike.) . . pro- ducing a condition of netlike branching known as anastomosis. vertebrate animals. The cells branch laterally to join each other quite frequently. longitudinal view of smooth (involuntary) muscle cells.g. canaliculi. Nervous Tissue. usually a single unbranched fiber except for infrequent collateral branches. showing process. one extending from the spinal cord to the hand or foot. but the fibers are faintly 107 segmented by thin intercalary disks which define areas each with a single nucleus.. and (b) dendrites.METAZOAN ORGANIZATION this tissue is rather difficult. A nerve cell body together with its processes is called a neuron. Squamous of bone showing Haversian canal and matrix. 8. Typical cells and tissues from epithelial cells Z. The neurons approach each other and pass impulses from one to the other at the synapses. 7^ motor nerve cell. (Drawn by Titus Evans. e. frequently much QO Fig. bone cells. striated (voluntary) muscle. where . An axone may be several feet long. human red blood (nonnucleated) corpuscles and human white (nucleated) corpuscles. — This is specialized to receive stimuli and trans- mit impulses which have been set up by some stimulating agent in some part of the body. 45. lacunae. the dendrites The impulses are conducted toward the cell body over and away over the axone. section of hyaline lacunae. 1. — cartilage showing cartilage cells in tion of tendon composed of white fibrous connective tissue 5. 3. Dendrites may be lacking. and matrix between lacunae i. Two kinds of processes are recognizable : (a) the axo7ie. The structural features consist of nerve cell bodies and their processes. 6. section through a portion (in center).

The Skeletal System composes the supporting framework of the The bony and cartilaginous tissues make up the material of this system. skull. and birds these is cells are nucleated. An organ is an arrangement of two or more tissues as a part of the body which performs some specific function or functions.. such as hair. The Integumentary System Its principal is composed of the skin and its out- growths. and regulation of body temperature. an aggregation of organs properly associated and some general function of life. cavity. primarily. b. proper relation. . ribs. Vascular Tissue. Lymph has no red corpuscles. . The vertebral column. nails. branch through the wall nervous tissue reaches all parts of the organ to receive stimuli and distribute impulses. The cells are the red corpuscles (erythrocytes) and white corpuscles (leucocytes). purposes are protection. horns. with some degree of excretion and respiration. while the intercellular substance. Blood and lymph In the common vascular tissues. sternum. The chief function of this tissue the transportation of digested food and oxygen to the cells of the body and the removal of waste by- products of metabolism from them. and they serve for the support of the body as a whole and for the probody. 108 TEXTBOOK OF ZOOLOGY the brushlike ending of the axone of one comes into close proximity with a dendrite of another. while in frogs. and connective tissue serves to bind all the others in . scales. and similar structures. — This is fluid tissue consisting of cells known as corpuscles in a fluid the plasma or fluid are the two medium called plasma. and bones of the limbs are the general parts of the vertebrate skeletoji. Some organs are made up of scribed. There are ten different systems usually recognized: system is A related to perform a. of the nervous tissue In this way an impulse can The be trans- mitted from one part of the body to other parts. hoofs. tection of the internal. some absorption. vital organs. and lymph spaces bearing lymph. turtles. It is all of the different types of tissues just de- For example the stomach is an organ with an internal covered and lined with epithelium. the wall contains two strong layers of muscular tissue blood vessels carrying blood. blood of mammals the red corpuscles are without nuclei fish. is to relate is chief function the organism to its environment.

it is The general form of the system is that of a tube.METAZOAN ORGANIZATION c. striated group moves skeletal parts and accomplishes locomotion the nonstriated. esophagus. The functions of . small intestine.. stomach. Precaval V. —Ventral view of human maniltin showing parts of the principal systems.1 Ascandinc^ Colon Fig. and the cardiac muscle produces the . 109 The Muscular System consists of muscles.Diaphracfm Liver . large intestine. .) d. {JuqularVdn Carotid Artiry Trachea Sabclavian V. and in- frequently called the alimentary canal. (Drawn by Edward O'Malley.Stomach Gall Bladder Jro-nswrseColon D<?5c ending Colon _ . the voluntary. The Digestive System of the higher animals includes the mouth. heart action.Left Auricle LeftVentrick Xu. and ac- cessory glands. Liuodenam '- -_ . involuntary g-roup is concerned with the movements of the internal organs (viscera). pharynx. Dorsoil Aorta Pulmonary A. .

and other foreign nodes. such as flame tubules. absorption. known as hormones. bacteria. and the islands of Langerhans of the pancreas are They go under the names of typical examples of these organs. capillaries. The kidneys withdraw liquid waste products of metabolism from the blood and deliver them to the outside of the body. lymph The general functions are: (1) to distribute blood carrying food. urea and uric acid dissolved in water. and ganglia. and coordinate the activities of the parts of the body. trachea. and lungs in the terrestrial vertebrate forms accessory to nasal passages. (2) collect and transport to the point of exit carbon dioxide. egestion. spinal cord. and hormones from glands of internal secretion to the tissues. pharynx. Malpighian In the mammals. h. gills of various modifications in many aquatic the lungs are the bronchi. is made up of tubular struc- tures and accessory parts. The Respiratory System consists of structures capable of delivering oxygen to the body and eliminating carbon dioxide. oxygen. liquid wastes. larynx. The Endocrine System includes a number of different glands located in various parts of the body. matter. adrenals located near the kidneys. directly into the blood. ductless glands and organs of internal secretion also. and urethra are accessory to the kidneys. but in all higher forms there are special structures for this function. the anirelates itself to it. The nitrogenous substances. the ureters. In some forms the general surface of the body serves the purpose. peripheral nerves. g. The Circulatory or Vascular System and lymphoid glands. are the principal prod- ucts discharged. lymph spaces. is a very extensive one con- sisting of the heart. veins. hormones cooperate to regulate the metabolic activity of the entire body. carry sensations. These glands discharge chem- The ical substances. By the function of the sensory portion of the system. The thyroid gland of the neck region. . Metazoa. puts the food it may be absorbed by the blood. by this system. Tracheae are found in insects. arteries. little ex- cretion are performed in solution so that e. cells. The Excretory or Urinary System and kidneys. correlate them. mal becomes aware of the environment and In vertebrates the principal parts of the system include the brain. and . digestion.110 TEXTBOOK OF ZOOLOGY and very it gestion. secretion. The Nervous System is an organization of the nerve cell bodies and their processes in such a way as to receive stimuli. green glands. In general. sense organs. urinaiy bladder. i. f. nephridia. autonomic nerves.

which are formed in the sexual phase or generation. produce a number of spores which deform a single zygote. and sj^stems to systems to form the metazoan organism. they break through the wall to the outside of the body. among the simpler metazoans a single individual produces both male and female germ cells. in general. Such an organism is said to be . but. organs to organs to form systems. one male and one female. This is one produced by each parent. tissues to tissues to form organs. The body might be thought of as being constructed by relating cells form tissues. simple fundamental process of reproduction by cell The or division binary fission has been studied already. Conjugation of Paramecium is also looked upon as a forerunner of sexual reproduction.: METAZOAN ORGANIZATION 111 j. egglike. More discussion of this system is found below. this type of animal begins life as a single cell resulting from the fusion of two sex cells. and sexual. The individuals in a colony by peculiarities in cell division become differentiated into two types: reproduction (a) the ordinary. In simple Metazoa there are likewise two forms of reproduction asexual (without sex). These will all be studied in more detail in connection with to cells to the study of specific animals. which involves the union of two germ (sex) cells. Development of Sexual Reproduction Keproduction makes great advances among the metazoans. cells unite to In reproduction these two types of from which a new colony arises by number of the Sporozoa. and accessory structures which function in the reproduction of the species. ducts.). not possible for most metazoan animals. Again. and the smaller. there seems to be the beginning of sexual reproduction. These become nutritive trophozoites and are capable of production of another generation of gametes. When the germ cells are mature. The zygotes. The Reproductive System is an organization of glands. motile microgametes. In simple forms like sponges and jellyfish the germ cells arise from general formative interstitial cells between the two primitive germ layers to form temporary gonads. repeated divisions. as well as possibly in Paramecium. In some of the colonial Protozoa and also in Sporozoa. both sexual and In a asexual generations occur. is fission : whose means of and (b) reproductive individuals or gametes inactive macrogametes of two forms the large. velop sporozoites (already studied under Plasmodium. nutritive individuals. including buddijig and fission.

The sexes are separate under such conditions and are said dioecious. of a number of animals. in which it is possible for the unfertilized q^§ cell to develop without union with another germ cell." The eggs sea urchins. No males are produced until the last generation of the season. This individual feeds on the sap of the particular plant on which she nary aphids or plant ample. even in man. the females lay fertilized eggs which pass through the winter and hatch as the first generation next spring. These mate. Another generation of female aphids arises from these eggs which in turn reproduce in a similar manner. and this time there are both males and females. particularly insects. up to and including the worms. these becoming queens if fed abundantly on proper food or workers if fed otherwise. Some authors speak of this process as "virgin birth. only females are produced. he never has a son. artificially worms. phylogenetic scale. Instead of mating (there are no males in her generation) she produces a series of eggs (macrogametes) which continue to develop without union with a sperm (male germ cell).112 TEXTBOOK OF ZOOLOGY Most of the types of animals in the hermaphroditic or monoecious. known to every gardener. and others have been stimulated to continue . nor did he himself have a father. that though a drone bee may become the father of thousands of daughters. In higher forms the usual method of reproduction involves germ cells produced by two individuals. lives and grows to maturity. If her eggs are not fertilized. molluscs. as parthenogenesis. such as frogs. to be the gonads of the other sex having degenerated in that individual. The case of the ordi- is a good exIn the spring an egg which was fertilized and laid the previous fall hatches to produce an individual known as a stem-mother. "So it comes about. These individuals are the stem-mothers for the new season. as most of them are. are normally Infrequent examples of hermaphrodites occur either normally or occasionally abnormally here and there among the higher groups of metazoans." The honey bee queen can control her offspring to some degree. A series of female generations appears in succession during the summer. This is known lice. There are some forms. the offspring are all males (drones). Each cell is either male or female. In regard to this state of affairs Lane puts it this way. hermaphroditic. If the eggs are fertilized.

They are in a very imand are called pri- mordial germ of the germ cells within the gonads. and the development of the female germ cell is oogenesis. This process is an alternation of production of sexual individuals in one generation and asexual in the next. In brief maturation is the preparation of germ cells for fertilization which may follow. Many of the sexually reproducing plants have a similar alternation of sexual and asexual generations. Ohelia. a coelenterate related to Hydra (to be is a plantlike. life Metagenesis a phenomenon occurring in the history of a number of scattered species of Metazoa. in each case differs The offspring from its parents. are first differentiated. The medusae produce eggs and sperms which unite in the water and develop into asexual colonies. agents. colonial form. studied shortly). Metazoan and OntogenyOntogeny refers to the development and life history of the individual organism. or 113 mechanical This goes under the is name of artificial parthenogenesis. Embryological development is an expression referring to the processes which occur during the earlier portion of the life of the individual. including the coelenterate. which gives rise to sexual. except for the latter part of the process in ova which reaches completion after the cells leave the ovary. the tunicate (a chordate animal).METAZOAN ORGANIZATION development by application of chemical. but possibly it insures better and more complete distribution of individuals than could be secured by only the budding colony. electrical. there In Ohelia. and Salpa. produced sexually from the union of germ cells or gametes. 59). . asexual. The male and female germ mature state cells or gametes are produced in their respective gonads as previously described. This process is quite generally similar wherever it occurs. Metagenesis really involves two methods of repro- duction in successive generations of the same species. The significance is somewhat uncertain. It consists of a series of mitotic cell divisions which is The maturation (gametogenesis) or development still occurs while they are modified at one point to bring about a fusion and subsequent reduction in the number of chromosomes in the cells. when they cells. free-swimming medusae (Fig. differing only in detail. two or three marine worms. This is spoken of as alternation of generation. The development of the male germ cell is known as spermatogenesis.

47. each Each of these chromosome divides with the division of the stitutes the multiplication period of the Oogenesis Primordial _ This series of divisions conmaturation process. cell. The primordial cell divides by is mitosis to form two oogonia. Maturation of the germ cells. Next.114 TEXTBOOK OF ZOOLOGY Oogenesis begins with the primordial germ cell within the ovary. — instances each of these cells divides once more. At the close of this growth each of these cells is called a of these oocytes divides as by meiosis. nearly all of it goes to one of the . though they were single ones in primary oocyte. This division. results in cells with half the somatic (diploid) number of chromosomes and is spoken of as the reduction division. divides similarly. For purposes of illustration the process will be described for a form whose diploid number of chromosomes is eight. and spermatofjenesis is a similar process of division in the development of mature male germ cells or spermatozoa. As typical of mitotic division. Oogenesis includes the maturation divisions of the female germ cells or ova. This fusion is spoken of as synapsis of chromosomes. The cytoplasm does not divide equally. polar body^--'' rertili5ed ovum (Zygote) Fig. cells of having the normal number of chromosomes for the somatic the species. therefore. In some Spermatoqemsis Primordial qermcell f^ WC qerm cell J'permafco- gonia --Zbromosome Primary oocyte Secondary oocyte f. These cells are typically spherical or oval with a prominent nucleus. i -(fi\\ \'^(/ v Primary ipermatocybe _ Secondary "\ / /(• spermatocyte Nature ovum [it. During this time the chromosomes in each unite in pairs and fuse together. //T { jSpermatid fj polar body Mature spermatozoon Zr>d. This number of chromosomes is known as the diploid number. Each the fused chromosomes dividing normal division. each of these oogonia passes through a growth period without division.

METAZOAN ORGANIZATION cells in 1]5 each case. They now degenerate. but none of them have any further significance after cariying half of the chromosomes. Fertilization involves the union of a mature ovum and mature spermatozoon to produce a fertilized ovum or zygote. is mary spermatocytes which have developed during When and the primary spermatocytes divide. The change from spermatid to spermatozoa does not involve a cell division but simply rearrangement. Each species has a definite and constant number of chromosomes. a series of mitotic cell divisions. thereby making possible for the germ unite without doubling the typical number of chromosomes in each new generation. their protoplasm is reabsorbed and Spermatogenesis like oogenesis. The spermatozoon SAvims to the q^q and enters it by penetrating the outer membrane which is called the vitelline membrane. the number of chromosomes it is reduced to half in each cells to mature germ cell. The ovum containing the haploid number of chromosomes is now prepared to unite with a mature spermatozoon in fertilization. Each chromosomes. For most animals. the division an equal one all of the resulting cells are typical secondary spermatocytes with the haploid number of chromosomes. four chromosomes. as soon as one sperm enters an egg. Occasionally the first ovum and another away polar body divides. and this process it continues just as does in oogenesis. It is now able to swim in fluid and prepared to unite with a mature ovum. Each spermatid then undergoes a change of shape or transformation to form the mature spermatozoa. The primordial divide by mitosis to form spermatogonia. is there is very significant for at least two imporduring the fusion and subsequent divisions of given opportunity for variation of the genetic com- position. The series of divisions and changes following the primary oocyte stage constitute the maturatio7i period of the process. These cells divide to form spermatids. in this ease. middle piece. is germ cells is completed within the tubules of the testis. each with its half or. until the division of the prithe growth period. the mature Following this the secondary oocyte divides to form polar body. The spermatozoon is a slender. by the surrounding tissue. the chemical nature of the vitelline . and tail. the cells. number The maturation process tant reasons. motile cell composed of head. is This large first cell is called the secondary oocyte and of these cells has four the small one the polar body. First. Secondly. and.

) . . early gastrula stage (infolding of cell layer at one side) 9. two-celled embryo following first cleavage division. and the fertilization is complete. C. The male and female pronuclei finally fuse The to form the fusion nucleus. the ensuing cleavage . . later stage of gastrulation. Evans. the sixteen-cell stage stage (solid) 7. The egg nucleus is known as the female pronucleus. 1. . yolk material is In eggs where the scant and evenly distributed. Secondly. — . Cleavage is a series of mitotic cell divisions beginning in the zygote immediately following its formation. it is the genesis replaces this function in some cases. These divisions occur in rapid order with but very little intervening growth. the four-cell stage the eight-cell stage //. The infolded layer is the endoderm. morula 5. significance of fertilization is largely centered around two important impetus for the development of an embryo from the egg under most normal circumstances. Fertilized ess (zygote) 2. the of means for inheritance of ancestry. . The head of the sperm carries the nucleus and soon takes the form of a rounded male pronucleus inside the cytoplasm of the egg. . parthenofunctions. First. Diagrams showing cleavage in the young embryo of Asterias. and the resulting cells adhere to each other in a body. blastula stage (hollow) 8. S.116 TEXTBOOK OF ZOOLOGY membrane changes and prevents entrance of others. 48. however. 6. (Drawn by T. characteristics from two different the diploid lines This union also restores number of chromosomes. it brings about Fig.

formed. the wall of which is now a single layer. marks the end number As of cleavage. These pouches push into the remains of the blastocoele. the divisions of the developing divisions all of the cells of the rate that it appears as if embryo are unequal. This gradually crowds out the cavity and forms a wall of two layers of cells. In higher forms. the one on each side in the gastrula. is organized from cells usually contributed by one or the other or both of the other germ layers. immediately following gastrulation. As cell divisions continue in the blastula. the two margins of the infolded part come nearer and nearer each other. which usually comes at the sixty-four cell stage or later. The outer layer is known as the ectoderm (outer skin) and represents the portion of the wall of the blastula which has not folded in. In some cases it arises as two saclike outgrowths from the endoderm. This is the beginning of the two primitive germ layers. In sponges and coelenterates development stops here. crease in the and growth of the embryo which is necessary before any special parts can be formed. a third germ mesoderm (middle skin). is called endoderm (inner skin). or from an undifferentiated . During the early body divide at so nearly the same the zygote were being cut with a knife or This process provides for the rapid in- cleaver into smaller parts. Cleavage will be described more fully in a later chapter under the discussion of the development of the frog. finally meeting the wall of cells from the other side. or that resulting from the infolding of the wall of the blastula. forming nearly equal cells. The inner layer. In other cases separate cells are shed from ectoderm or endoderm or both. of cells divisions proceed.METAZOAN ORGANIZATION divisions extend completely 117 through the zygote. This gradually encloses an outside space which is lined by the endoderm and represents the primitive digestive tract or archenteron. The formation of the blastula. a hlastula is formed by the development of a cavity (blastocoele) within the spherical mass of cells. a gastrula is finally The blastula does not simply increase in circumference. but there comes a time when the wall on one side pushes in (invaginates). As division of cells in this wall proceeds and the infolding continues. layer. ectoderm and endoderm. The blastula stage of an animal like a starfish or a frog resembles somewhat a hollow rubber ball whose wall is made up of a large number of pieces cemented together. If the yolk is concentrated in one end of the egg.

and reproductive systems besides nearly all connective tissue and the endoderm produces the internal linings of the digestive tract. outpushings. Sooner or later a space forms within the mesoderm. and extensions which finally bring about the formation of all parts of the mature individual. of the germ layers. The division proceeds at different rates in different regions and at different times resulting in various infoldings. the mesoderm furnishes the muscles. the circulatory system.118 TEXTBOOK OF ZOOLOGY portion to orgajiize as a distinct layer between the other two. The position of the mesoderm is external to the endoderm and internal to the ectoderm. This cavity the coelom or future particular parts of the is body cavity. outgrowths as the liver and pancreas. The fate of the germ layers is determined as cell division and development continue. excretory. skeleton. . respiratory tract. It nearly encircles the endoderm. and such . From each body are derived. blood. The ectoderm gives rise to the external surface cells or epidermis of the skin and to nervous system. causing the outer limb of it to join the ectoderm and the inner to join the endoderm.

Proterospongia. The organization of the sponges is a loose one.CHAPTEK IX PHYLUM PORIFERA SPONGES The name of this this. This group is thought to be sort of an aberrant type with peculiar relations. of locomotion. sessile in habit. There are only a few small fresh-Avater forms. that they were fully acknowledged as animals. The body is in the form of a hollow sac with many canals life. An animal with hundreds of mouths cannot be very highly organized. is shells. phylum. notwithstanding the presence of a simple mesoderm which is lacking in Coelenterata. calcareous. The germ layers are not well-established the layer which seems to begin a common . Some authorities show a rather close comparison between sponges and colonial Protozoa. pilings. and others have a flexible fiberlike piercing the walls and cavity and the outside. 119 The so-called ecto- . for There entire lack making connection between the internal The pores of these canals are essentially mouths. Most sponges. There are workers who hold that sponges may have arisen from ancestor with the choanoflagellate type of colonial Protozoa. like endoderm develops into the external layer. means ''pore- bearers. only a little over ninety years ago. material as a skeleton. live in the sea. The sponges do not have a distinct enteron or digestive cavity. For a long time sponges were thought to be plants. etc. All sponges have some type of skeletal structure. and the interdependence of part upon part is not great. but digestion is entirely intracellular (within cells). being fastened to piers. and it was not until 1857. They are rocks." and these animals certainly are.. Porifera (p6 rif'er a). For a time sponges themselves were considered colonial Protozoa. There is only one general exit from the cavity. The sponges possess collar cells or choanocytes which are similar to the cells of the colonial mastigophoran. but the group is often considered the simplest and lowest type of Metazoa. some possess hard. bath sponges included. They have tissues but are without organs. or siliceous spicules.

If the type can be classified according to germ layers. Classification Class Calcispongiae. acterized — Single. it might be considered a modiBecause of these pecufied diploblastic (two germ layers) form.. is probably the most beautiful of the sponges. possessing a very thin body wall cells. char- by calcareous spicules. with pores as perforations in individual lined with choanocytes. is —Simplest type. derm comes lating the water. .) — Order Homocoela. shallow-water.120 TEXTBOOK OF ZOOLOGY to line the internal cavities and its function is circuThe middle layer is very poorly differentiated. (Courtesy of General Biological Supply House. and is hardly recognizable as the mesoderm of the typical triploblastic animal. Fig. The internal cavity Leucosolenia. 49. Glass sponge or Venus's flower basket. liarities. some authors have called sponges Mesozoa or Parazoa. There are two orders. marine forms. Euplectella sp. being hardly more than a matrix.

fragilis. Venus's flower 'basket. or finger sponge. Class Demospongiae. which possess with three axes and six rays or a multiple of six.{•• •'I '**-••/•* J ftatocyiea cong-etfaie to jorm gemmates inWrgzd^zrnmuka «. —Forms Order Tetraxonida. Asteromeyenia plumosa.) (Courtesy General Biological Supply Fresh-Water Sponges In the southwestern part of the United States. Usually they are not over 1/4 of an inch in height.— PHYLUM PORIFERA Order Heterocoela. A few fresh-water forms are known. Haliclona. Often called glass sponges because of this skeleton.•'*•/. marine forms and one family of fresh-water sponges (Spongillidae). Spongilla fragilis seems to be the most abundant in this area. —Moderately — Sponges complex Choanoeytes in siliceous spicules ScypJia (Grantia). Class Hyalospongiae. averaging approximately as 6 inches by 2^ inch in diameter. 121 wall. Spicules are white and like spun glass. There are less than two dozen fresh-water sponges known in this country. —These are ordinarily attached — to the bottom in Order Monaxonida. but some are cushion-shaped and a few are branched. which have either nontriaxial siliceous spicules or spongin or no skeleton.'^-*i- K"'-'^ Mi- Fig. but there are some as large inches. Most of the large colonies of sponges in this region are dark grey or chocolate brown in color and are found % . and Ephy- Texas. ^•faiocytes— •^ . Most of the colonies are irregular in shape. House. deep water. and Cliona. radial canals. at least in central there are four species of fresh-water sponges: Spongilla TrocJwspongilla horrida. Most of the colonies of this of an species are irregular in shape. Includes shallow-water. or boring sponge. Thenea.. 50. Of the four. datia crater if ormis. They have complicated canal systems and are often quite large and brightly colored. Spongillaj showing reproduction. Spongilla.

"W. drawn in through the pores or ostia on the sides of the body. 85. The animal is is is attached by the basal or proximal end. de benfels. skeletons of spongin waters. sponge and apparently gemmule formation is a perennial process. Pasadena. 1937. It is not as simple. is also This It is available and comparatively simple in structure.122 TEXTBOOK OF ZOOLOGY on logs either floating in the water or submerged. a commonly studied representative of the phylum. This water is forced up through the cavity and out at the osculum or exit opening at the top." the European form. but in the region referred to they horrida and Ephydatia crater if ormis) (particularly Trochospongilla grow abundantly in muddy ponds and in muddy streams whose turbidity equals 110 parts of In this region again the growth of the solid matter per million. Calif. p. sometimes covering several square inches. 199. The species are usually identified by means of microscopic differences in the gemmule spicules as seen when crushed. posited in a pavementlike layer on the object to which the sponge is attached. however. —A group of important sponges of which man uses The representatives of this order have and are found in subtropical and tropical marine at least a dozen different ones. the bath sponge. as Leucosolenia. by most textbooks for years. even following periods of low These gemmules are ordinarily detemperature in the spring. The maximum production of gemmules seems to be in the late autumn and throughout the winter. Habitat and Behavior This type lives attached to rocks in relatively shallow marine water. 10. Water •A Case of Incorrect Identification American genus is Scypha. M. Feb. —These sponges are entirely devoid of skele- Haliscara. then by way of the canals into the internal cavity.. 2199. In some parts of the country there is the idea that sponges require clear water. Lau- . No. Order Myxospongida. The osculum and ostia maj^ be closed and there opposite end free or distal. Order Ceratosa. the A colony may be formed by budding. THE SIMPLE SPONGE Scypha coronata* is (Ellis and Solander) has been mistakenly called "grantia. Science Vol. Euspongia. ton.

It is reported that an average sponge will pump approximately forty-five gallons of water through his body in forty-eight hours. because they are sessile. at the top.— PHYLUM P0R1FEE.. A row of picketlike spicules or spines encircles the osculum. and is called neuroid transmission. . On the contrary. rather goblet-shaped with the excurrent opening.-:^.-^. These reactions may involve the entire body. and impulses are conducted simply from cell to This results in very slow transmission cell in a primitive fashion. or they may be local. of impulses External Anatomy The average length of Scypha It is is about three-fourths of an inch. Activities and coordination in Scypha and sponges generally are quite limited by lack of a nervous system.^. showing habit of life.i. Scypha coronata (Grantia).A 123 may be contractions of the entire body. Individual cells respond directly to stimuli. • -oiisS^Wy. these ajiimals work day and night to keep a continuous current of water to supply their vital needs.. These movements are accomplished by individual contractile cells.. 51. Fig.LV. inactive forms.-. osculum.. Laymen and many zoologists think of sponges as sluggish..

shown of different types of Laboratories. They join the cavity through small pores called apopyles and extend nearly to the outer surface of the wall where they end blindly. : : S .bcr DiAfftftm of Akod Sponge FlaftlUitd Oitmbc (Radul Canal) \ S-' . each one opening distally by an osculum. of rSagon (Icucon) type of canal 3truc:ure such «9 occurs Sycon Sponge in the Demospongiae Fig. They are quite evenly distributed over the wall.124 TEXTBOOK OF ZOOLOGY and other less conspicuous spicules are distributed over the body.. Caiiral Co. —Structure (Courtesy Pacific Biologicalsponges. This canal system not only serves to . They connect with the radial canals by rather numerous apertures called Stereogram to illuatntc ample Leucen Sponge Dermal Oatta ^Subdarmal Cavity Ch^lcnf Cnal FUstlUied Cham^r Diagram StetcogrKiT) to illustiate L.ihcl.) diagrammaCcally. Communicating with and radiating from this cavity is a set of radial canals. Lying between these and extending inward from the ostia are the incurrent canals. cu. The ostia are the incurrent pores through which water is taken into the body. Internal Internally there is Anatomy is a large central or gastral cavity which is simply a water cavity and not comparable to a stomach or enteron. In 10% 4^ I Dctfnal EpiiKctium . InkaltAI Canal •1 k«l>M«n "« 11. 52..- Charr. more complex sponges there may be several or even many such cavities. A dermal epithelium covers the outer surface of the animal. S.

simplest. there are three types of canal systems. In sponges generally. contractile cells (myocytes).raxon Fis 53. is and still Spongin of the ordinary bath sponge. Some have a skeleton of calcareous spicules. wtich secrete the substance for anchorage. In Order Ceratosa. A great many of the cells in this layer do not have distinct boundaries. The monaxon type . flat. The character of the skeleton is a diagnostic feature in the classification of sponges. the triaxon type possesses six rays lying in three axes. others have no skeleton. The histology of Scypha presents a peculiar arrangement of a number of different types of cells. consists of simple straight spines. dermal layer is composed of simple. The honaxon Triaxon . The outer. the triradiate type consists of those that have three rays joining each other in one plane the tetraxon type has four rays radiating from a common point in four different planes .PHYLUM carry the water. and the poly axon type has numerous rays. and rliagon types of which the first is the and the third.) Jriradiatc found in different sponges. formed by some special cells called spongioblasts. the gland cells . which silk simply the skeleton of one of these animals. others of siliceous spicules. sycon. identified as the ascon. it PORIFERA 125 substantially increases the surface area of the This seems to be a definite provision to allow increase in volume by keeping the ratio of surface to volume. —Types of calcareous skeletal spicules (Drawn spicules are of several tj^pes with a number of modifications of each. The canal system of Scypha is of the sycon type. others of the fibrous spongin. The cells which produce spicules are known as sclerohlasts. making it a syncytium. resembles It is chemi- cally. the second intermediate. the most complex. but animal. Hon ax on by Joanne Moore. as well as the sclerohlasts. epithelial cells.

Co/lencyte -flaciei lum DertnalceU — collar Choanoo/ie -I (Oslhrceli) Porocyte s^. The radial canals and the internal wall of the central cavity have similar histological structure since the former are The cells here are primarily special outpouchings of the latter. The incurrent At each prosopyle there canals are is a single large dermal cell. Ovum '4^~ Fig. drive the suspended food particles into the funnellike the collar cell where a food vacuole fashion. interspersed with scatshaped Jpicule Sckroblast . is formed in the cell in mouths of amoeboid Of course the spicules appear in a histological section. after Minchin. lined with flat pavement cells.) — Each choanocyte has at its free margin a funnel or collar opening to the central cavity and a flagellum or whip extending from the funnel. The flagella agitate the water and tered. Treatise on Zoology. flat. The entire arrangement is quite similar to a large colony of semiindependent cells which do not function as integral parts of a tissue as do the cells of higher animals. It has been found that indi- . Dermal cells AirchaeccYbe. ehoanocytes. published by The Macmillan Company. (Redrawn and modified from Lankester. In the middle layer are found the reproductive cells and some amoeboid wandering cells. 54. peculiar to sponges. Histology of wall of a simple sponge in longitudinal section. a porocyte. epithelial cells.— 126 TEXTBOOK OF ZOOLOGY spongioblasts are located in this layer. which surrounds the aperture.

Budding involves the branching of new individuals from the external surface of an old one. is made with the surrounding water. and the exchange of gases O2 and CO2. Excretion is largely by general diffusion through the surfaces. a From these new sponge will reorganize. Reproduction and Life History Scypha is able to reproduce both asexually and sexually. Egestion is probably accomplished much as it is in Amoeba. This may is be augmented by the carried on Respiration by diffusion through the general surfaces. The former may be by budding or by the formation of gemmules. found particularly in fresh-water sponges. Catabolism. which become infiltrated with siliceous materials. Digestion is intracellular (within cells) in the food vacuoles and the process is much the same as has been described in Protozoa. The digested material is assimilated some authors. The current is produced by the flagella of the choanocytes and contractility of the walls. according to A At any rate the food particles are taken into the cytoplasm of certain of the cells. These new individuals finally become free from the parent. by diffusion from cell to amoeboid wandering cells. As the current sweeps the potential food particles into the collar cells they are seized and ingested by pseudopodia. goes on in the cells in some degree as long as they are alive. Sometimes a colony is formed by the buds remaining attached to the parent. if kept in favorable conditions. They are usually formed during adverse conditions and can withstand desiccation and other severe . Groups of cells become separated from the surrounding deep tissue by limiting membranes. through the canals and central cavity. Gemmule formation or internal budding is another type of reproduction. cell. Metabolism sponge obtains food from the water which is continually passway of ostia. Microorganisms and other particles of organic matter are drawn in with the water.PHYLUM PORIFERA 127 vidual cells can be separated from each other by squeezing some types of sponges through the meshes of a silk cloth. It is ing by controlled by the contractility of the cells surrounding the ostia. and out the osculum. living cells. or the union of oxygen with the fuel substance of the cell to liberate energy. perhaps assisted by the wandering cells.

55. the parent individual then dies. These later settle down. ciliated larvae. and are modified to form adult. — Methods of reproduction in Scypha. of the male usually mature before those of the female. The reproductive cells are produced in the jellylike middle layer. become attached. TEXTBOOK OF ZOOLOGY In fresh-water forms these gemmules are formed in the middle layer of cells. . Sponges are usually hermaphroditic. sessile sponges. but the germ cells Sexual I^eprodaction Fig. and the following spring new individuals emerge from the gemmules. Fertilization takes place here and cleavage division progresses. 56. (Courtesy General Biological Supply House. At about the blastula stage the embryos are liberated through the wall of the body as free-swimming. Osculum breaks thru here -Dermal epithelium Qranular cells (Dermal epithelium) Segmentation cavity Ragellafced cells Gasbral epithelium Gastral cavity (future cloaca) {Qastral epithelium) --Dloitopore Typical f ree-5W/mming Typical /Amphiblaifcu/a Awphiblastula Fig.) 128 circumstances. at time of attachment sections of —Diagrammatic Scypha larvae. Sexual Reproduction occurs here for the first time in our discussions.

Phylog-enetic Advances of Sponges Wlien Compared With Protozoa flagellata of the This group appears to be somewhat advanced over the ChoanoGenus Proterospongia in Class Mastigophora. sorted. and Central America. Next they are trimmed.PHYLUM PORIFERA Economic Relations 129 Many sponges are beneficial to man. Dredges which are sometimes used for this purpose are not very satisfactory. here in a simple way.000.000. and the pieces fastened to hooks or wire on a weighted frame. lesser items include the large flint de- some species and those used for The chief importance lies in the use of the spongin skeletons of certain groups for bath and surgical sponges. Red Sea. destroyed by certain sponges which attach themselves to the mollusc's body or by others which bore through its shell and thus kill it. . the Bahamas. and sometimes bleached. coasts of Florida. If everything has gone as siliceous spicules of from ornaments. Selected sponges of good quality are cut up. because they drag down and kill so many young sponges. expected. The value of the crop each year is at least $5. like plants. They are allowed to rot for a day or two. Most of the commercial sponges come from the Mediterrajiean Sea. the slips will then be marketable. The "seeded" frame is sunk in the ocean and left for a year or two for the sponges to grow. and there are a few wliich Oysters and some other Mollusca are injured or are detrimental. Of posits positive importance. else organized into layers and are differSexual reproduction is developed The group has advanced so little that little can be said. dried. With rapid industrial development sponges have also become useful as a fabric material. and placed on the market. Sponges die quickly when taken. washed. then beaten and squeezed under water. The demand has brought about the establishment of sponge farms where they are raised from fragments about an inch square or slips. In cells are partially sponges the entiated for separate functions. Sponges are usually harvested by diving or hooking between May and October. the West Indies.

Excretion and respiration are carried on tion is by the general surfaces of the body. this cavity. and possess tentacles with "sting bodies" or nematocysts. digested food have been engulfed This is called intracellular digestion. Sexual reproduction. A The radial symmetry is correlated with an attached habit of life. The group is considered among the simplest of metazoans and shows. good many of the attached forms look much like plants and were so described for a long time. Most of the The species are marine. Coelenterata (sel eu ter a'ta). Most of the representatives do not develop skeletal structures. calcareous cases around themselves. Hydra will be studied in detail. being accom- plished by enzymes which are secreted by special cells of the endoderm into the internal or gastrov oscular cavity. They are all modified gastrulas. at least a Most coelenterates are attached or very sedentary for part of the life span. digestion." and all of the representatives bear this out by possessing a single large cavity in the body. and for that reason they are said to be diploMastic. but coral polyps produce hard.CHAPTER X PHYLUM COELENTERATA The phylum name. but there are a few fresh-water forms. In several species there is the typical alternation of generations of attached and free-living forms. takes place within the ticles of partially A limited amount by these of the endoderm cells after parcells. because it is 130 . There is a single opening to anus. involving production of ova and spermatozoa and their union in fertilization. and it functions as both . have radial symmetry. means "hollow intestine. typical features of this great division of the animal kingdom. These two layers are the outer ectoderm and inner endoderm. Asexual reproduc- accomplished by budding and fission. occurs here too. in a simple way. The digestive process is principally extracellular. mouth and There are two general types of coelenterates the polyp form and the jellyfish form. however. body wall is composed of two layers of cells.

lithocyst g. or hydroids. Tubularia. . will be discussed as a general representative of the phylum. floating colonial hydroids. easily collected and handled. . some smaller jellyfishes. of this order. . ve.) . . is divided into three classes. — and the fresh-water polyps. Classification of the Phylum The phylum orders. . —mn. m. velum. radial canal re. Fig. These are typical polyp forms. General Biology. ring canal st.. The group includes marine. The first one is a fresh-water polyp form and is the best known of the group. tentacle. (From "White. t. . gonads nematocyst ra. manubrium Structure of Gonionemus. Gonionemus. The last one listed is a fresh-water form with a small polyp stage lacking tentacles but with a disclike medusa possessing many tentacles. and of multicellular animals of simple formation. many of which produce medusae forms by budding. each with three or four Class Hydrozoa. stage. n. PHYLUM COELENTERATA readily available. . mouth stomach. colonial polyps. one special gro"up of corals. Ohelia. is 131 representative The study of Hydra as a simple metazoan will go far in giving insight into the much more complex make-up of the body and life of man. Canipanularia. but since Gonionemus and Ohelia are common marine forms. Hydra. such as Portuguese man-of-war. ad. Adhesive pad li. Order Leptolina a group which has a sedentary or sessile polyp Such examples as Hydra. and Craspedacusta are well-known forms. 57. a brief description of them may be included — here.

Also as branches of the stem. at the end of each is a mouth and tentacles. This stemlike part gives off lateral branches. leaflike folds hanging in the subumbrella and radiating from manubrium to margin are the gonads. covers the colony. The colony begins as a single individual which buds. with a large amount of jellylike mcsoglea between these two genn layers. From a few to more than eighty almost solid tentacles hang down from the margin of the subumbrella. colonial type resembling a branched plant in appearance. bordered by four oral lobes. is the third type of individual connected with an Obelia perisarc. The animal is able to swim about by drawing water into the partially enclosed cavity of the subumbrella and expelling it through the aperture formed by the velum with enough force to move the animal in the opposite direction. and is found in the pelagic waters. A planula-like hydroid form develops from the egg. the hydrocaidus. hydranths. end is the mouth. They are distributed in the Atlantic Ocean and Gulf of Mexico out to forty fathoms in depth. Obelia colony. The exumhrella is is the convex upper. A short stalklike part.132 • TEXTBOOK OF ZOOLOGY Gonionemus is a small jellyfish form. measuring about a centimeter across. is a marine. The medusa colony. Wa\y. At The mouth the aperture leading into the internal or gastrovascular cavity which has four radial branches or canals. lower. This may continue for several generations. The individuals are attached it is to each other in the and fastened to a rock or other substratum by a root- like hydrorhiza. These are feeding polyps. along our eastern shores. The cell structure of this animal is made up of an outer ectoderm and an inner endoderm. but they do not separate from the preceding or parent generation. nonfeeding polyps capable of producing medusae. From the hydrorhiza there is an upright stem. or aboral side while the suhumhrella its distal is the concave. The pressure is developed by contraction of the body. is The which is In some parts this ringed. These radial canals join a circumferential or marginal or ring canal. A circular ledge or fold of tissue which extends inward from the margin of the subumbrella and partially encloses this saucer-shaped cavity. oral side. is called the velum (craspedon). and it expands at the base of the . there are the hlastostyles which are modified. the manuhiHiim hangs down from the center of the subumbrella. Its shape reminds one somewhat of an umbrella with a fancy fringe but with practically no handle and made of clear cellophane. composed of chitin.

58. Obelia Obelia. structure.) . showing: asexual generation. (Courtesy of General Biological Supply House.— - PHYLUM COELENTERATA 133 Obelia habit Mouth tb/drothecaCoelenteron ? '' hntoderm tctoderm Qonotheca MedU5Q-bud B\a5bo5t\/ie - Radiol canal ^^ Repiroductive Moubh Jcatocyjfc— Tenbacles- 7^^-tirf^ Kedusa Fig. and habit of life. hydrozoan colonial coelenterate. sexual generation (medusa).

The blastostyle with this coverFibrous processes connect the ing is often called the gonangium. the ectoderm. published by McGraw-Hill Book — Company. Animal Biology. Inc. The hydranth captures and ingests small aquatic organisms as food by the aid of stinging bodies (nematocysts) produced in certain ectoderm cells of the distal portions of the tentacles. Another modification is the taller. Germ cells produced by the medusae complete the cycle. (Redrawn and modified from Wolcott. Blastula and planula are free-swimming. ^"^ \ *.^^g ^ cell stage. a part of the gastrovascular cavity. and an inner endoderm layer lining the cavity. There are between twenty and thirty solid tentacles attached around the basal margin of the hypostome.) The coenosarc is made up of an outer layer of cells.Ferbilucd eqq \*" "^ Mature. more enclosed case.> Cleavacfe BJastula. hydranth perisarc to the soft. qonancjlcim. Life cycle of Ohelia. ^ \arva ^ mature Position of colony Fig. . 59. The mouth of the hydranth is located in a domelike hypostome at the free end. Adult hydroid colony with mature gonangium gives rise to sexual medusa which is produced in the gonangium and set free in the water. of the there- hydranth is continuous with that of the hydrocaulus. The cavity is. illustrating polymorphism and metagenesis. gonotheca. Medusa JJ J e x Sperm -from another medusa ^__ . and fore. The digestion of this food is accomplished just beneath the perisarc.. inner parts (coenosarc).134 TEXTBOOK OF ZOOLOGY to form a bowllike case or hydrotheca which supports it. which nearly encloses the Mastostyle.

— . to distinguish The medusae them from of this sort are spoken of as hydromedusae the scyphomedusae or jellyfishes of Class Scyphozoa. C. after Claus. the medusa or jellj'fish form. . : . and others. some produce eggs. The zygote develops into the free-swimming. . sporosac F. Obelia presents a very good example of metagenesis as represented in animals. After producing a generation of medusae. Diagram of a siphonophore colony (Physophorida) A. The medusae arise as buds from the special individuals. published by McGraw-Hill Book Company. protective zooid E. .) . dactylozooids H. ciliated planula stage which soon attaches and develops into a polyp from which a new colony arises. blastostyles. The reproductive cycle is both sexual and asexual. feeding polyps (gastrozooids) (From Van Cleave. is described as meta- genesis in Chapter VIII. alternating between the sexually produced polyp or hydroid generation and the asexually produced sexual generation. 60. Pneuniatophore B.PHYLUM COELENTERATA in the internal cavity. and after producing germ cells. swimming bells D. Fig. which are discharged into the water at maturity and unite to form zygotes. nettling cells. esses. escape through the distal pores. spermatozoa. and develop to sexual maturity as free-swimming individuals. The sexes of these are separate. Zoology. . G. the medusae die. to be studied soon. a single 135 With the exception of reproductive procwill be hydranth of Obelia found similar to an entire hydra. involving alternation of generation. This process. this colony disintegrates. Invertebrate /.

as it is called. A common tube of the coenosarc unites the five kinds of individuals of the colony. the staghorn or stinging coral. branched hydrorhiza and powerful nemato cysts (stinging 'bodies). Fig. Order Hydrocorallina. 61. Rudimentary — — medusalike bodies develop on the coenosarcal canals. Physalia. —This is a pelagic order of colonial coelen- terates with extreme polymorphism.directly with no polyp stage. This group resembles the corals by producing strong calcareous skeletons. Millepora.— 136 TEXTBOOK OF ZOOLOGY Order Trachylina. This order includes two suborders of hydromedusae which come from the egg. the Portuguese man-of-war a floating colonial coelenterate. College Zoology. published by The Macmillan Company. They have extensive. continuous from one individual to another. is a good example.) Order Siphonophora. coenosarcal tube is and this cavity is The blind end of the an air-filled. Campanella and Liriope are generic examples. bladderlike float (pneumatopJiore) . (From Hegner.

There are usually eight or more tentaculocysts perradially and interradially distributed on the margin — . if it exists. The coelenterates belonging here are large having an alternation of generation in which the medusa form is dominant. suffer severe pain. such that the entire colony appears as a single individual. crest. These jellyfish have a complex system of branched radial canals and abundant marginal tentacles as well as oral tentacles. Swimoccur just below the pneumatophore. and almost transparent. — They occur in the open sea. Conical or vase-shaped medusae which usumarginal sense bodies (tentaculocysts). it Order Stauromedusae. saucer-shaped. Pericolpa and Periphylla. 137 into the water beneath The polyps hang down of polyps include : The types gastrozooids (nutritive or feeding). four perradial tentaculocysts. Its sting quite poisonous bathers coming in contact with the trailing tenis tacles. Some of them are more than seven feet in diameter. The scyphomedusa has an eight-notched margin. interradial tentacles. to ming bells (nectocalyces) often Most of the individuals are specialized care for only limited functions. and has gonads connected with the endoderm.PHYLUM COELENTERATA with a superior this float. called taeniolae. but may be that the polyp form has not been discovered yet. Order Cubomedusac. dactylozooids -with nests of nematocysts and having long tentacles (tactile ductive zooids. These are cup-shaped. The tentacles are adradial and perradial. The tentacles are distributed perradially and interradially. and possessing tentacles one hundred feet in length. Lucernaria and Haliclystus ally lack — are usually cited as examples. and protective). which bear batteries of nematocysts. Forms which have rather cubical shape. free-swimming forms with four interradial tentaculocysts. Scyphozoa. — Charyhdea is an example. such a degree that they This specialization and diversity of forms is Physalia. Scyphozoa whose medusae are dominant. The polyps have four longitudinal endodermal folds. reproand others which produce ova-bearing medusae. Order Discomedusae. gonozooids which are male. which form gastral tentacles or filaments in the medusa. the Portuguese man-of-war. Order Peromedusae. Class jellyfishes — Several of the representatives of this class are thought by some zoologists to exist generation after generation only as medusae. . There are records of individuals of this group twelve feet in diameter. lacks the velum (therefore acraspedote). is a typical example. and are chiefly tropical.

Stomolophus meleagris. The generic name flrst . When they are dried. there joining a circumferential •This spelling is according to Mayer's monograph.-Sub-umbrella space — Grtutor muscle Manubrium (cut surface) Central mouth Oral tentacles Fig. This is a the Gulf of Mexico. TEXTBOOK OF ZOOLOGY Tentacles are usually present also on the margin of is the most numerous and extensively distributed the bell.- -Cabbage-head jellyfish. This group of Scyphozoa. Cut surface of to body wall Stomach Qonad ^ Jub-gsnital pit Upper portion of -^r — manubrium Lateral mouth \ Radial canal . The animal has no velum as do the hydromedusae. the typical example. proposed by Peronas Le Sueur was so spelled. but there is a square mouth on the subumbrellar side with wing-shaped. A large number of branching radial canals extend from the gastrovascular cavity out to the margin of the bell. and. like most jellyfishes. A suhgenital pit lies in each quadrant of this The mouth leads through a short passageway There is into the angular gastrovascular cavity which in turn has four lateral gastric pouches containing the fringelike gonads. in the Gulf of Mexico. side of the animal. Aurellia and Stomolophus are common examples. a very common form Bisected to show internal structure. only a thin film Aurellia* is remains. common one and ranges from New England It may reach a foot in diameter. 62. liplike oral lohes or arms.138 of the bell. is composed largely of water. also a row of small gastric filaments here carrying nematocysts.

-iy :T*L^5=f'^^^"""V C Admdial canal vV-i^^:?-----^'"'?^"^' 'S^_]_.PHYLUM COELENTERATA -Chrysaora) 139 (in Lonq tentacles Chryiaoro) Larqe subqenibal pit (as in TentaculocyJt Admdial canal Perradial canal Jhort5f'mp/e oral - arm (Aurelia) . -Aurellia and comparative structure in Pacific of jellyflshes. (Modified from figure Biological Laboratories' catalogue. i^-:- : lIfJ&. cells to the . • . showing staj^es from germ ephyra stage which precedes the adult condition.) Stages in the development of the scyphis Stages in the -development ^of the jbrobila toma Planula larva MM ^^ Sperm from separate adult Zygote Fig.'iv. ' ^ j ) V^ _ _ Aon(7 ribboh-like oral arm (in Chrysaom) "^ - /^C"""^ 'Small 6iibgenibal p/fc (aj /n Aurelia) ^ '^ Qastric pouch Short tentacles (Aurelia) '^ Marqinal lobes (as in Chrysaora) Fig. -^^ >-:. — Life cycle of Aurellia aurita. 64.^-#• . 63. - •*. -Intenadial canal .

Sagartia. even in tide pools. Reproduction involves both sexual and asexual generations. — Metridium usually lives attached to rocks or to solid bodies in They average about three and two or two and a half inches in diameter. The polyp with all of these constrictions is known as a strohila. points ou the margin.e. Subclass Zoantharia. The mouth is located in the center of the crown. The mesogloea is quite abundant and contains a good many cells that resemble primitive connective tissue cells. —All ganization and may be colonial or solitary. It includes sea — This typically occurring in multiples of six.140 canal. constrictions are formed around the body making it resemble a stack of saucers the upper one periodically frees itself and swims away. Near it is the olfactory pit. Then the polyps form medusae by strohilization. group has numerous paired septa. and it can change its location by "seooching" on its lasal disc (attachM end). and plain tubular tentacles. bands are found in the septa. The free end of the jar-shaped body is covered with tentacles which are provided with nematocysts. and Bletridmm are common examples. . or four inches in height . The pigment spot over each is likely sensitive to light. and they pass out through the mouth with the water. TEXTBOOK OF ZOOLOGY The eight tentaculocysts are symmetrically located at eight The tentaculocysts are sense organs of equilibrium. Many of these animals produce a calcareous external skeleton called coral. Order Actinaria. Both sexual and asexual reproductissue tion are common. and the egg develops into a free-swimming plajiula which after attachment becomes a tubelike polyp that reproduces by budding most of the season. Cerianthus. Muscular Class Anthozoa. . anemones and corals. The entire body can be expanded and contracted. Fertilization takes place. Germ cells are produced by the pinkish gonads in the gastric pouches. each between marginal lappets. animals in this class conform to polyp orThey have an ectodermal esophagus and longitudinal partitions called septa (mesenteries) incompletely dividing the gastrovascular cavity. and food is forced into it and on through the gullet (stomodeum) bj^ action of cilia on the tentacles and part of the lining of the mouth. and the new medusa is called an epliyra. At each side of the gullet is usually a the water near shore. i. These anemones are usually solitary polyps they have many complete septa and numerous tentacles but no skeleton..

141 is through which water constantly carried into the gastrovascular cavity for respiratory purposes. A quarternary set is represented by mere ridges on the inner surface of the wall and interspersed among the others. 65. There is a band of muscle running vertically on the face of each septum next to the muscle on the adjacent septa of the same rank. Below the gullet the mesentery has secretory filaments which in turn bear long. Between these and the primaries are some tertiary septa which are still shorter ajid also attached to the inner surface of the body wall. At the basal end these cavities are continuous Fig. showing external features.PHYLUM COELENTERATA ciliated groove. Between the primary septa are secondaries which do not quite reach the wall of the gullet. The gastrovascular cavity is divided into radially arranged compartments by the primary septa or mesenteries which extend from the wall of the gullet to the inside of the body wall. with the main central cavity. the siphonoglyphe. hence their medial ends are free in the cavity. threadlike acontia. — Sea anemone. The primary septa in the axis of and extending parallel with the siphonoglyphes are called directives. Metridium inarginatum. is .

with Aconlium protruding SlereogTam of Anthozoan Polyp MeKtilcrie fiUment. The gonads develop on the edges of the lower part of the septa to provide for sexual reproduction. The development includes cleavage and planula stages. Madrepora. cKamter between .) (Courtesy Mature ova and spermia are discharged into tlie water of tlie cavity and escape through the mouth to unite in fertilization outside. — Diagrams to show the structure of the anthozoan. of Anthozoan Polyp fi\ level A-A Otagrammatic T S. Asexual reproduction by budding from the margin of the basal Occasional longitudinal fission may disc is practiced by this animal. Cinclade. The representatives of this order secrete an The indiexternal limestone skeleton most of them are colonial. occur. Otherwise they are similar to anemones. Astrangia.tfsl ^' Directive* Siphonogtyph SpSincler (Ventral bet of Primary Meccnteriea) Diagrammatic T. and Oculina are examples. — . cKsmber between pair* ^ Hollcw Tentacle o( meicnteriei ' ^ Vf)i. Order Madreporaria. The sexes are distinct. and they are supplied with nematocysts and secretory cells. 66. before the new individual attaches and changes form. of Anthozoan Polyp at level B-B Fig.142 TEXTBOOK OF ZOOLOGY These protrude through pores (cinclides) in the body wall to the outside. Metridium. EnJoceet. S. of Pacific Biological Laboratories. . They serve as defensive as well as offensive structures. two metenteriei the of ume pair "-^^ Cxaeoel. viduals of colonies communicate with each other by coenosarcal connections.

Order Antipathidea. fied so that — —Another is colonial one portion submerged in the substratum. Order Pennatidacea. The common sea fan. One such reef is over 1. — srri.. feathered tentacles. An order composed of branching colonies whose individuals are joined by a branched tubular axis which is covered by an epidermal layer. eight hollow. form whose body is modiThe colony . A. Gorgonia. eight mesenteries.- ^»* A Fig. Body walls of individuals fuse together as one.^^. and one siphonoglyphe. Subclass Alcyonaria. Coral poljqDS vary in size from one-sixteenth of an inch to several inches in diameter. Colonial and pol^'morphic forms are not Order Alcyonacea. — Order Oorgonacea. Cirripathes and Antipathes are typical examples. — Common when cleaned Astrangia danae. as well as the precious Corallium rubrum are well known examples. The colonies are bilaterally symmetrical. Organ pipe coral belongs in this order. A colonial group which has calcareous spicules but lacks an axial rod.PHYLUM COELENTERATA Astrangia secretion is 143 it is the common coral polyp. polyps in natural habitat. — The features of this division include uncommon. Stone produced by the animals B. Many corals are of beautiful colors.100 miles and from ten to twenty-five fathoms deep. Alcyoninm is the type example. "^^^ coral. and quite similar to a small sea anemone to which calcium carbonate has been added by from the ectoderm cells as well as having budded to form In time a colony of numerous individuals. .^ •Trv^. continually growing colonies of these animals can produce enormous stony barriers (reefs) in the in length sea. This is another colonial coral which is sessile and has a calcareous axial rod. 67.

Chlorella vulgaris. . in endoderm cells. sea pens is and sea feathers. There may be dimorphism of zooids within the colony. debris. and the need of food. carnea. and the hydra benefits by receiving oxygen from the photosynthesis of the alga. TEXTBOOK OF ZOOLOGY stem which and the individuals are born on a disc or axial supported by a hard skeleton. Hydra will eat only when it is hungry and will not react to food at other times. such as H. Hyman. and Pelmatohydra oligactis. and 50. HYDRA Habitat and Behavior Hydra (Chlorohydra) of the active viridissima is is likely the most common hydra is Southwest. On the other hand.144 takes a bilateral form. Frequently hydra is able to stretch its body over articles of food which are actually larger than the hydra usually is in normal condition. moving about from place to place. Renilla and Pennatula. Vols. It keeps the tentacles extended. attached to the surface of plant leaves. 49. H. This species has the green color because of the presence of a unicellular alga. is a sedentary kind of animal if and may remain stationary for a considerable period of time living conditions are uniformly the environmental conditions are changing. are sluggish and have longer tentacles than the green ones. in cool fresh water. Most of the hj^dras are found of the water. it becomes quite active. or even the surface film The brown hydras. americana.* Hydra good. published in tlie Transactions of the Society. ready to grasp any food which may come into its reach. The plant uses some of the by-products of metabolism of the hydra. are typical examples. 48. The prey is then carried to the mouth and tucked into it by the tentacles. it will likely be paralyzed by the toxin which is injected by the nematocysts. It the small green hydra which very and has short tentacles. smooth sticks. It is more sensible than many people in this respect. it has been authentically reported that a hungry hydra will perform the characteristic feed- When is animal in found American Microscopical Recent taxonomic information concerning Hydras of the United States may be in the papers of Libbie H. Nematocysts or sting bodies are discharged when the tentacle comes in contact with potential food. This kind of a relationship is the called symbiosis. and if it chances to be a small animal.

The movements of the animal are performed by contraction and relaxation of the contractile fibers connected with certain of the cells. If the stimulus is of a general nature and of considerable ditions. the animal will contract in the affected area. Locomotion in hydra. it These animals show response to a number of environmental conAny sudden change is likely to bring about a negative response. responds to a chemical stimulus alone. (From Jennings. the also. but it will not respond a mechanical stimulus only. . published by The Columbia University Press. by the withdrawal of one tentacle. The activities come in response intensity.PHYLUM COELENTERATA ing movements 145 Thus to when only beef extract is in solution in tlie water. animal will contract all of the tentacles If the stimulus is restricted to one locality to internal as well as external stimuli. Behavior of the Loiuer Organisms. Fig-. Successive positions taken when progressing by somersaults. 68.) — and the body and is not too strong.

holds by the tentacles. they simply become less and less active and finally cease to move as the freezing point is approached. then they attempt to find lower temperature. They likewise possess an optimum for temperature and prefer relatively cool water. Attached around the free end of the column is a circlet of from four to seven Tentacles may fingerlike tentacles. are They respond to light and will find an optimum Green hydras intensity which varies with the different species. Or the animal may bend over and cling to the substratum by the tentacles. where the basal disc is reattached. but in H. In some forms the distal (free. or posterior) end. or anterior) end of the column is much stouter than the proximal (attached.146 TEXTBOOK OF ZOOLOGY present in hydras. cylindrical trunk which ordinarily stands in a vertical position. aboral. being able to extend from a contracted length two or three millimeters to a length of eighteen or twenty millimeters. viridissima there is only a slight tapering toward the basal end. The common tropisms. This process is consecutively repeated and is called "loopOccasionally the animal bends over. which extends free in the water. Locoynotion is accomplished in at least four ways. As the temperature is lowered on them. It remains attached in contact with some solid body most of the time. hence they are adapted to the Southwest. react positively to sunlight and withstand moderate temperature. then draw the body toward this point." bubble of gas at the basal disc and floating back to the top on that. Sudden mechanical stimulation like stirring the water or jarring the attachment of the animal will cause it to contract vigorously. then turns a "handspring" or "somersault" to attach the basal disc on the substratum beyond this point. Contact stimuli are of considerable significance in a sedentary animal like this. The fourth means by which locomotion is effected is by dropping to the bottom. oral. Its body quite contractile. thigmotropism are concerned in food-taking. External Anatomy Hydra is is a macroscopic animal. As pointed out previously. then secreting a ing. The column or body is a tubular. release the basal disc. which have been described previously. Gliding from one point to another by partially releasing the basal disc and slipping it to a new location is common. but it is relatively small. of . both chemotropisms and . They seem not to become particularly uncomfortable until the temperature gets up to 31° C.

PHYLUM COELENTERATA 147 Stretch out to be slender threads five to seven centimeters in length. ally bear buds before the first is separated from the original parent. and these occasionthe side it From conical end of the hypostome. appears simply as an indentation or notch in the The proximal or attached end terminates in a 'basal disc or foot. Mouth Tentacle Hypostome Battery of Nematocysts Bud Basal Disc Fig. This conical elevation between the bases of the tentacles in which the mouth is located is called the hypostome. viridissima. either singly or as a They are very useful delivering food to the mouth. From one to several luds are often found on the sides of the trunk. group in capturing and The mouth is located in the center of the distal end of the column and is surrounded by the tentacles. — Hydra showing external features. Buds are lateral outgrowths of the column and are found when the animal has favorable living conditions. The mouth when closed and viewed from the top looks something like an asterisk. Occasionally there may be observed rounded projections on the side of the column which . Budding usually occurs at about the middle of the body in H. which secretes an adhesive substance which helps the animal in attaching to objects. 69.

148 TEXTBOOK OF ZOOLOGY Both ovaries (female gonads) and are seasonal reproductive organs. Since the large barbed type is the most conspicuous. they are appropriately called epitheliomuscular cells. Internal Anatomy Another feature of the organization of this animal is the diploblastic structure which consists of two layers of cells or the germ layers surrounding an internal space. but it is arranged with an axis of polarity from basal disc to hypostome. tubercles in the ectoderm. testes be formed on a single individual. fibrils which extend in a direction parallel to the length of the animal. sac of fluid within which In the cnidoblast the nematocyst appears as a is inverted a barbed stalk with a coiled thread attached. the gastrov oscular cavity or enteron. but they are usually seen on separate individuals. These cells contract to shorten the length of the animal. it will be described here. Each of these cells consists of a polyhedral outer or epithelial portion and a basal portion which is drawn into one or two long. If these projections are conical and located nearer the tentacles. develop the neniato cysts. slender. These are studied on stained sections. which is essentially equivalent to what is called the ventroAll of the metazoans have a dorsal axis of more advanced forms. Each tubercle contains a large barbed one and several of a smaller variety. Four different kinds have been described. maries. which when chemically stimulated causes the cnidoblast to discharge the nemato- . This animal possesses radial symmetry. The most numerous ones are typically cuboidal in shape and serve both as contractile units and as the general external surface of the body. Sedentary and sessile animals very commonly have radial symmetry. they are ovaries. while the motile or free-living organisms tend toward bilateral symmetry. or nettle Interspersed occasionally tributed over among all these cells are the larger cnidohlasts in which cells cells. The outer one is the ectoderm. but they are much more numerous near the distal part of the column and on the tenThe nematocysts are usually contained in little raised tacles. cnidoblast is Projecting out of the superficial surface of the a triggerlike process called the cnidocil. if primary axis. stinging the These are dis- body except the basal disc. they are testes or sper- (male gonads) may they are more nearly knoblike and are located nearer the base. which is thinner and is composed of four types of cells.

and the thread it becomes coiled around the object against which charged. when added Contact will not. 70. after Toppe. the entire cnido- replaced by an interstitial or spent cell of the cell migrating into position.PHYLUM COELENTERATA cyst. Inc. there are the scattered. insect larva attacked by hydra. Stalk-- Bag- Remains of Cnidoblasb Barbless nemafcocyifc Fig. . Cnidoblast containing an undischarged nematocyst. They crowd in between the other cells.) The cnidoblasts are produced by a third type of cell. C. the stalk is barbless. Nematocysts and their function. after Jennings. after Schneider. This type of nematocyst produces a hypnoit is toxin which anesthetizes the animals into which is discharged. probably by de- velopment of pressure. nematocyst everted and extended but still held in the cnidoblast. In is small. near their bases. will bring this about. McGraw-Hill Book Company. Animal Biology. (Redrawn and modified from . leg of small aquatic insect with barbless nematocysts on its spines. after Jennings E. to the water. . D. A ir- damaged body may be replaced from the inter- stitial cell. or methyl ^een. In this reaction the stalk and thread are everted. another form the sac elastic . especially cell. — Wolcott. the interstitial which is small and rounded. Besides these three types. blast is As a nematocyst is discharged. acetic acid. A._:il1 6orb__. portion of tentacle. it is dis- Cnidocil Filament Nemaboo/st Nucleus a. after Schneider B. such as weak iodine. These are formative cells about the size of the nuclei of the epitheliomuscular cells and quite densely granular in nature. 149 Chemicals. and thus impedes locomotion of the victim.

A 150 TEXTBOOK OF ZOOLOGY TEMTACLE — MOUTH •TESTI S GASXROVASCUl_AR CAVI TY ECTODERM MESOSLOE EfslDODERM OVARY BASAU Fig. DISC — Diagrammatic longitudinal layers.) is the . —gastrovascular through orthe column (Drawn by Cross-section enteron. showing mature gonads and (Drawn by Titus C. typical cell section of hydra. cavity The central space Titus C.) ECTODERM EPITHEl_IO- MUSCUL. AR CELL. Evans. NJTERSTI Tl Al_ CEI_U I NEMATOCYST C N DOBL. Evans. AST I MESOGUOEA DIC3ESTIVE CEUUS G l_ A IM D CEI_U Fig. of hydra. 71. 72.

neuroepithelial cells 151 by intercellular processes. These cells fit which are joined into a net between the others and cells are either sensory or motor in function. Ingestion of the food has been described already. The digestion which occurs here is spoken of as intercellular digestion and is brought about by enzymes produced by the secreting cells of the endoderm. The inner. entering the mouth the morsel of food is moved some distance in the cavity by successive wavelike contractions of the column progressing from distal to proximal.PHYLUM COELENTERATA regular. they wedge is themselves between the nutritive-muscular cells and secrete what probably a digestive fluid into the gastrovascular cavity. These the free margin and at times engulf particles of partially digested food like an amoeba. of organic matter in the water. Here in the upper half of the Upon down enteron digestion takes place. thicker cell layer of the wall is the endoderm which lumen of the gastrovascular cavity. It is produced by the cell layers and serves as attachment for them. thus receiving external stimuli and also causing contraction of the contractile at proper times. . The disgland cells in the endoderm. minute worms. In some of the lines cells other coelenterates. By contraction these cells in- crease the length of the animal cells often possess flagella at by reducing its circumference. Metabolism The food small bits hydra consists of small insect larvae. Glandular cells are also present in this layer. Being slender. The general morphology of the adult animal is very similar to the gastrula stage of the developing embryo of more complex metazoans. The wall possesses many more of the and the food material disintegrates into smaller particles here in this region. and other of small Crustacea. Beneath the ectoderm and embedding the bases of the cells is a very thin layer of noncellular substance called mesogloea. water fleas. The most conspicuous cells here are the nutritive-muscular tures attached to which are long. that they serve both as muscles and as digestive cells. vacuolated structhe mesogloea by fibrils which extend in it parallel to the circumference of the animal. slender. Neuroepithelial and interstitial cells are also interspersed among the other cells of this layer. this layer the is exceedingly thick and heavy. particularly for the fibrils of the epitheliomuscular cells. It is seen then. Such serial contractions are usually called 'peristaltic contractions.

Hydra with of meat. which extends into the tentacles and buds. 73. and by diffusion the nutrient solution reaches effect of the contractions of the body. and the digestion many is of the nutritive-muscular cells by virtue of their amoeboid activity.) As into remembered. There is no separate system of transportation will be This dissolved material reaches the remote parts of the protoplasm by diffusion through the membranes and protoplasm generally. (From Curtis — body turned inside out in attempting: to Ingest a piece and Guthrie. This illustrates something of the primitive organization of hydra as a metazoan. Small particles of the partially di- gested substance are engulfed by the free ends of particles are taken in food vacuoles. Inc. The food is distributed to all parts of the enteron. there just as in These completed it is an amoeba or Paramecium. published by John Wiley and Sons. the ectoderm cells just outside. function of digestion and circulation.152 solution of the food TEXTBOOK OF ZOOLOGY by the enzymes is augmented by the churning The flagella present on the nutritive-muscular cells create currents of water which also hurry The dissolved material is presumably absorhed by the the process. by the action of the flagella and by bodily contractions. cells of the endoderm. this process of converting the digested food an integral part of the protoplasm is known as assimilation. . The gastrovascular cavity has the dual or circulation of nutriment. Textbook of General Zoology. Fig.

Nutrient mate- is . When one sensory cell is stimulated. and water which must be expelled from the body. The Nervous System and Nervous Conduction The neuroepithelial cells are distributed among the other cells of germ layers. Catabolism or dissimilation takes place in the protoplasm and involves the union of oxygen with the substance of the protoplasm transform potential energy there to kinetic energy and heat. uric acid. this oxidation there are produced some waste byproducts in solution including urea. There is a greater abundance of them on the hypostome. which makes this region in a sense comparable to a primitive brain. all of the sensory cells seem to be stimulated in some degree. because the protoplasmic requirements are the same in all to Accompanying animals. There is some indication that there may be accumulation of waste products in endoderm cells as cytoplasmic granules. which finally escape through the gastrovascular cavity and mouth. A sufficiently strong stimulus affecting any sensitive point will stimulate the entire body. This process is essentially reproduction by somatic cell division. very similar to the comparable processes in Protozoa and the same similarity will be noticed when they are compared later with the higher forms of animals. in general. and tentacles than along the length of the column. In hydra this excretion is accomplished by diffusion through the general surface of the body. This is a definite the organized type of nervous system but not a very efficient one. used as food have hard skeletal parts that These indigestible portions are ejected from the cavity through the mouth by reverse peristalsis. basal disc. Asexual reproduction accomplished very efiiciently and quite rapidly.PHYLUM COELENTERATA 153 Many of the animals will not digest. The greatest concentration of these cells is in the hypostome around the mouth. Reproduction and Life Cycle Reproduction is both asexual and sexual. It will be noticed that these phases of metabolism are. Eespiration furnishes the necessary exchange of oxygen and carbon dioxide by diffusion through the plasma memThe dissolved oxygen in the water in which the animal lives the source of this element. and the process is known branes. These cells all over the body are in contact with each other by means of their processes forming what is called a nerve net. is as egestion.

l tiydra. and in the terminal position a mouth is developed. hydra reproduces sexually. The cell division at this point is very rapid.. involving considerable activity in interstitial cells. maturation.154 rial TEXTBOOK OF ZOOLOGY accumulates at some point near the middle of the column. where they multiply by division to form oogonia in . After the bud has attained some size. — but it has been reported that individuals of separate sex (dioecious) have been found. An extension of the eiateron extends into the bud. — Methods of reproduction in hydra. Embrvo ?6md. these animals are hermaphroditic or monoecious as suggested before. This enlargement rapidly increases in size to form a stalk. Tentacles appear as outpushings of ectoderm and endoderm. The germ cells or gametes develop from interstitial cells which accumulate at a certain place between the ectoderm and endoderm. This involves the production. As a rule. 74. which is essentially an outgrowth of the body wall. The first appears as a slight superficial bulge. Testes may appear first and ovaries later on the same individual or both gonads may be present at the same time in which case self-fertilization is possible. During the summer and fall particularly. (Courtesy General Biological Supply House.<§exua. a constriction occurs between it and the parent. This closes the enteron between bud and parent. and union of germ cells.l l^production Fig.) Sexual Reproduction. and the bud finally separates to bud become a free individual.

which when mature emerge periodically from an opening in the tip of the testis and are discharged free in the water. — . cleavage cavity (blastocoele) m. 75. The process continues until a hollow llastula of many cells is formed. In the ovary a single egg develops at the expense of the other oogonia. polar bodies. and when Fig. cyst. Fertilization is accomplished by the en- trance of spermatozoa through a rupture in the overlying ectoderm and cross-fertilization usually prevails. Development of hydra. blastula stage. 14.PHYLUM COELENTERATA the female gonad 155 All phases of and spermatogonia in the male. Biological Bulletin. S. while the original . Fertilized ovum.) . These new cells on the inside become organized as an endoderm layer. i gastrula. Vol. The testis produces large numbers of motile spermatozoa. p. mature it fills the ovary. (After Tanreuther.. and this zygote A single sperm unites with undergoes the total and equal divisions of cleavage here in place. which are engulfed bodily and used for food. Then follows the formation of the gastrula by a shedding of cells into the cavity (blastocoele) from the inside of the original layer of cells. showing ectocferm (ec) and endoderm (en) cc. Z. the mature ovum. This one cell grows rapidly.b. maturation (gametogenesis) may be obsei-^^ed in the testis and ovary. 1. two-cell stage.

The corals are of importance both positively and negatively. A large coral reefs are very costly to navigation of marine waters. presence of tentacles with nematocysts or sting-bodies (5) continuance of sexual reproduction.156 outer layer is TEXTBOOK OF ZOOLOGY now known as ectoderm. If conditions are favorable for development. the embryo does not emerge from the cyst until spring. Hydra is able to replace mutilated parts or an entire animal from a be formed from very small This process is known as regeneration. (4) . it is of great advantage to the animal. (2) well(3) defined gastrovascular cavity with one opening. it increases in length within the cyst. Further changes involve the secretion of the thin mesogloea which seals the two layers together. . nomenon was first discovered 1744 by Trembly. Many of them are valuable as ornaments. pieces Complete animals may (% mm. This individual steadily grows and soon attains adult condition. the mouth. (7) a nerve net. in animals from studies on Hydra in Economic Relations of the Phylum is not worth much in dollars and cents to man number of different ones are used as food by some of the useful fish. when it has attained some size it breaks out. while the enteron develops within the endoderm. while the The entire group directly. This pheportion of one. Many corals are quarried for building stone. and in some instances they protect the shore from being washed by the waves. In the meanwhile a shell is produced about the outer surface of the embryo. Regeneration As is the case in many invertebrate and a few vertebrate animals. and while it is not normally a method of reproduction or multiplication. in diameter) of a hydra. (6) distinct radial symmetry and. after which tentacles and a mouth appear at one end. When the zygote is formed in the fall. and this encysted body falls from the parent to the bottom. Phylogenetic Advances of Coelenterates (1) Definite organization of diploblastic condition.

are often classified as There are only twenty-one Amer- ican species representing this phylum. phylum: (1) Tentac^data. sea walnuts or comb jellies. The symmetry said to be biradial since there are eight rows of radially arranged paddles or plates which are equally distributed on each side of the 157 median . bandlike. transparent. Venus 's girdle. with a faint tint of pink. There are two classes in the blue. lives There is a limited number that (te and moves about on the bottom. conical in shape. is an rows of fused cilia. Ctenophora nof '6 ra comb-bearing). about 10 to 12 cm. (2) Nuda. and green colors. oval-shaped body. These animals and transparent. Habitat and Behavior These are primarily surface-living forms with rather wide distriThey move about very slowly through the water with the oral end forward and the two long tentacles trailing if tentacles are present. and Cestus veneris. and the shape may be spherical. ribbonlike or cylindrical. a class in this phylum. may be four feet long and only two inches in width. most of which are pelagic (float near the surface). with an iridescence showing violet. Mnemiopsis leidyi is a luminescent. The tentacles have adhesive or glue cells (colloblasts) which produce a secretion. with a pair of tentacles present either in the larva stage or throughout life. Anatomy The size of different individuals of this group ranges from five millimeters to four feet in length. or They are often phosphorescent. is pear-shaped. transparent form Pleurobrachia hachei has long tentacles on a relatively short. example. and they are commonly called Most of them swim by means of eight called swimming plates or combs. blue. Beroe ovata. purple.— CHAPTER XI PHYLUM CTENOPHORA This is a group of exclusively marine animals. bution but most abundant in tropical seas. because of their similarity to coelenterates. with no tentacles at any stage. and rather common. are quite clear . and with these they capture any small organisms making contact with them. in length.

with endoderm.r .) angles.'/ . The mouth is in the oral end of the body and leads into the stomachlike stomodeum which is connected with a series of canals running through the body. called excretory canals. (Courtesy of Pacific Biological Laboratories. Bilobfff Stomo<faeuin InfunJibuTum Plate Row with ' ^ ^ Slomodaeuni branch of infundibular canal ayilem undetnealh Mouth Oral End Perspective! Drawing a}ong infundibulai plane Pcispcctivt Drawing along sagittal or 3tomodaeum pUne Yig. showing external features and structure. 76.. ' T. This cavity is lined egested through the mouth..'••'""" —~ - Plate row with Bilobed Stomodaeum Infundibular Canal underneath Diagram looking down on aboral pole Pleurobrachia bachei Aboral End 'C^ Radial \ Canal Inlerradial Canal o. When seen from the side.158 line. undigested food is The six canals mentioned above are There are two blind canals extending from the infundibulum beside and parallel to the stomodeum. these are called paragastric canals.. the paddles resemble a comb. There are two blind .. TEXTBOOK OF ZOOLOGY These paddles are tlie locomotor organs. This stomodeum is lined with ectoderm and leads to an infundibulum or gastTOvascular cavity proper which joins the stomodeum at right iflgitial or StomacS Plane Right CastrovBicular Canal 51 Left CAitrovfticular Cansl lUdial CanaU ^^ Intcrradial Cant.'.••... —Pacific comb jelly.:.^ 'Yvi^^—--''' '• . Tentaeulftt 'NviVrC^j Sheath ""~-.i i^j. The tentacular canals lead to the meridio- nal canals lying beneath the ciliated plates. »j^r) .-• "^ Aboral 5cn«e Orgon . i — • Tranaverse or Infundibular Plane A Tentacle -S '-i /.

The fertilized ovum develops and finally metamorphoses to the adult stage. There is no alternation of generation. which serve as organs of equilibrium by stimulating. in addition. — Sea walnut (CourtesyjeUy. The animals are triplohlastic instead of diploblastic as are Hydras. differentiated muscle fibers lying between the ectoderm and endoderm. from Atlantic Ocean and Gulf of Supply House. instead of a simple mesogloea. of General Biological Beroe ovata. Around the aboral surface of the body- is a collection of sense structures or statocysts. . The mature germ cells rupture into the infundibulum and pass through the stomodeum to the exterior. or comb Mexico. Otherwise they have no economic importance. These animals have no nematocysts and therefore do not irritate bathers as do jellyfish. there are.PHYLUM CTENOPHORA tentacular pouches connected with the outside of the infundibulum. but they do serve as food for a large number of valuable fish. Fig. These animals are monoecious (hermaphroditic) the ova are formed in ovaries along one side of each meridional canal and spermatozoa ternal calcareous ball (statolith) rolls as the level of the . This is a morphological advance when compared to the coelenterates. because.the cilia of the bands on the side against which the inbody changes. . 77. 159 one lies on each side The solid prehensile tentacles emerge from these sacs.) along the other.

there is group of soft-bodied. branch tubules. and a plentiful supply of secreting cells in this layer. and along with it have developed some systems of organs. Tricladida." and centralization is developed.'' The gonads are within the body and are connected with the exterior by accessory organs. Rhabdocoelida. 160 . quite The representatives of the two parasitic classes have. The nervous system in the free-living forms is of the "ladder-type. They possess a fairly well-differentiated mesoderm. for the most complex life histories and special adaptations. . Planaria and Stenostomum are examples. part. They are very important economically because of their injury to man and the domesticated animals. There are definite muscle cells. The common fresh-water planaria is an example of the free-living type while the parasitic flatworms are known as flukes or trematodes and tapeworms or cestodes. The excretory system is composed of a pair of longitudinal tubules. is digestive tube single. The The mouth is located ventrally. three-branched. broad worm) are usually called flatworms and in many ways show considerable advance over the coelenterates. and Polycladida. Four orders are known: Acoela. Class Turbellaria (turbela'ria sists — little stirring).CHAPTER The representatives of XII PHYLUM PLATYHELMINTHES Phylum Platyhelminthes (plat i hel min' thez. and the remainder of them are free-living. and excretory and reproductive systems composed of the new mesoderm layer. — This class con- of a forms. There are both land and water forms. elongate and usually free-living The surface layer or epidermis is ciliated in patches. Classification There are four recognized classes in the group. and" flame cells. The alimentary cavity functions as a gastrovascular cavity and has only one opening to the exterior. All of these worms are bilaterally symmetrical and triploblastic. Some of the species are parasitic. or many-branched.

is also char- by a heavy cuticular and a long. Cyclophyllidea. the neioly developed blood vascular system. The two muscular layers of the body are so effective that an extended worm of fifteen feet is Locomotion the body. have no epidermis but a thick nonciliated cuticle. It seems difficult to know where to classify group since some systematists give it the rank of phylum while others give it lower ranking. These animals. and excretory system. two apertures. are unsegmented "band worms. Ilymenolepis are examples. and cilia over the body are all characteristic of this type. may contract to less than two feet in length. but there seems to be no coelom. accomplished by the cilia and the contractility of The proboscis is a very characteristic organ which is . and the group is parasitic. commonly called flukes. There is present a mesoderm. ribbonlike body divided into sections called proglottides. Tetraphyllidea. They usually live in burrows in sand or mud or beneath solid objects.PHYLUM PLATYHELMINTHES Class 161 Trematoda (tremato'da having pores)." Most — this of them are free living and marine. Diphyllohothrium. A long proloscis. Trypanorhyncha. Fasciola (Figs. Paragonimus. The animals are frequently brightly colored. The animals feed on the bodies of other animals and on certain types of general organic matter. and 402). This group is divided into only two subclasses Monogenea and Digenea with orders Gasterostomata and Prosostomata. 400. Class Nemertina. 397). orders: The class includes five Pseudophyllidea. These tapeworms each have a knoblike "head" or scolex on the This structure is supplied with suckers for attachment and sometimes has hooks. The larger ones reach a length of ninety feet. — This group cover. the alimentary canal. There are numerous mucus-secreting glands in the skin which may produce a tubelike dwelling for the worm. 401. Clonorchis (Fig. A developmental stage of the life history is the bladder worm or cysticercus which lives embedded in the musanterior proglottid. and Heterophyllidea. Taenia (Figs. This entire class is parasitic. There is no alimentary tract. The body is either leaf-shaped or elongate and has from one to many ventral suckers. cular tissue of several different animals. 398 and 399) are genera representing the class. The Nemertinea (nem er tin'e a— unerring) as individuals. : — — Class Cestoda (ses to'da acterized — girdle form). nervous system. and the immature stages frequently make use of snails and crabs as hosts for a phase of their life history.

Lineus about 15 cm. Fig.162 in the TEXTBOOK OF ZOOLOGY form of a hollow tube turned back through the body inside of a cavity called the proboscis sheath. The . the principal nerve center of the animal in this stage.-J^loric caecum .) Fig.Qonad -MetractorM. with the body coiled. socialise length published by The Macmillan Company. Anus Fig. — Both eggs and sperm are discharged from respective individuals through a dorsal pore and fertilization occurs in the surrounding water. of the body plate is Cilia develop on the lappets at the lower margins This and on a patch at the opposite pole or apical plate. Structure of the nemertine worm. sheath.) by —A nemertine worm. College Zoology..^-Lateral \ mrve ffhynchocoe/ /nfccjfcinc Proboscis \\\ vSfcylefc _. the proboscis By contracting the saclike may be everted and extended from the anterior part of the body. Fig. after Coe. published The Macmillan Company. after Coe. Following cleavage there is a helmet-shaped larva pouches. Prostoma rubrum. (Redrawn and modified 78. as it appears when flattened. The sexes are ordinarily separate and each individual possesses gonads which are located laterally and between the intestinal _ Rhynchodeum > « . College Zoology. Natural from Hegner._ -Ocellus qanqlia . (Redrawn and modified from Hegner. called pilidium. 79. 79. 78.

fresh-water. PLANARIA Habitat ajid Behavior This free-living. or debris at the brooks and pools. Inc. Either one pore or several communicate with the exterior.fctodermaf invagination EsophacjUi Ventrolateral lobe Fig. Proscords passing through the body. Georaqe cavity L'lji^^i^— -Mesenchymal cell Stomach Lctodermal invagination / . cool water. The vascular fluid is usually colorless. flatworm thrives beneath the rocks. leaves.) — Structure of pilidium larva of the nemertlne worm in partial section. published by McGraw-Hill Book Company. These . Cerebratulus. logs. Animal Biology. The excretory system includes the usual longitudinal tubules and flame cells characteristic of the phylum. Other tactile organs and eyes are usually developed. ing larva 163 In some forms there is a creepThe vascular system is composed of longitudinal vessels connected by transverse loops. bottom of shallow spring-fed clear. 80. Tetrastemma are representatives.PHYLUM PLATYHELMINTHES adult appears after metamorphosis. A pair of grooves with cilia along each side of the cephalic portion are sensory and are called cerebral organs. algae. The central nervous system consists of two ganglia and three longitudinal known as Desor's larva. (Redrawn and modifled from Wolcott. toma. They must have pure.

They usually feed upon minute plants and animals. published by The Macmillan Company. They possess negative phototropism and thermotropism (as regards high . Planaria partially encompasses the food with the body. The planarians respond to several tropisms. and the pharynx is everted through it as a prohoscis which is used to draw food within. The ability to of slippery move along mucus which way is essentially lays a The behavior of this animal is of a reflex or automatic type. tracted to those points.164 TEXTBOOK OF ZOOLOGY animals are rather gregarious and when at rest will group together beneath objects where the light tively to bright light. Eye Genital pore Side of head PharjTix sheath Proboscis in Fig. If tiny scraps of meat are placed in a dish with hungry planarians. the planarians are atthe surface of fresh meat. 81. It is interesting to watch these animals passing the proboscis about over apparently sucking up the nourishing very minute quantities of meat juice are If fluids from the meat. Hegner. (From College Zoology. after Shipley and is The locomotion accomplished in an easy gliding fashion by the in this action of the beating cilia and muscular contractions of the body. It glides over a surface. receiving or receptor sensory cell transfers the impulse produced The by a stimulus to a ganglion cell or adjustor in the central nervous system which in turn transmits an impulse to an efferent cell carrying it to a muscle or gland. even the under side of the surface film of water. The mouth is located at the middle of the ventral side of the body. and adjusts itself easily to any irregularities because of the soft.) —Entire planaria with pharynx extended position for feeding. They respond negais not intense. while the pharynx is protruded to eat it. enhanced by the secretion smooth track for the moving animal. dead animal bodies. The ciliary action and muscular contractions are both rhythmic and progress in waves from anterior to posterior. flexible nature of the body. they will form a wad of living protoplasm about it. liberated in the water at specific points. such as small arthropods and molluscs. and living forms. McBride.

excretory. and reproductive systems. The common species are Plaiiaria niaculata. Many of the structures of the animal. while in Dendrocoelum lacteum there is much less. acids. The average length maculata is about three-fourths of an inch. alcohol. These cilia along with muscular contractions accomof fully developed active P. The pharynx may be protruded through the mouth in the form of a long. while the mesoderm constitutes the muscular. The undifferentiated mesoderm lying outside the intestine is composed of a meshwork of large cells and is called mesenchyme or parenchyma." called an auricle. strong salts. plish locomotion. The symmetry is distinctly hilateral. . They react positively to contact (thigmotropism) and water currents (rheotropism). The digestive system is composed of a mouth in the midventral position. the geniial pore. At each side of the "head" region is a pointed. External Anatomy The body is elongated. sensitive. The responses to chemicals are positive in case of food juices and the like. broadly wedge-shaped at the anterior and tapering to a point at the posterior end. It is triplohlastic since the ectoderm. P. dorotocephala. scarlike aperture. flat. extension of the epidermis in the These are sensitive to sound. and P. touch and chemical stimulations but not to The mouth is located in the midventral portion of the body. Externally the epidermal cells are soft and the general surface is nearly covered with patches of cilia which are cytoplasmic extensions of these cells.PHYLUM PLATYHELMINTHES 165 temperatures). while they are negative to alkalies. which have been observed in none of the forms previously studied. form of a lappet or "ear. endoderm. In Planaria maculata there is considerable pigment in the skin. and mesoderm are all differentiated and present in a clear-cut fashion for the first time in our studies so far. constricted. trunklike prohoscis which is used in feeding. Internal Anatomy The ectoderm covers the outer surfaces of the body and composes nervous system the endoderm lines the intestine and its branches. agilis. On the dorsal side of the anterior region are two pigment bodies called eyespots which are sensitive to light. Posterior to the mouth is a small. have come into existence with the development of mesoderm. a prehensile pharynx held in the pharyngeal chamber or the . etc.


which receive long cilia that are continually beating in a direction toward the tubule. that of the entire body. . longitudinal. a three-branched enteron or in- which branches immediately from the anterior end of the pharynx into an anterior trunk. and by several other pores along the length. coiled tubules. Cilia Excrelioiy tubule Fig. 83. The excretory system is new to our study and is composed of a set of tubules which relate themselves to all parts of the body. it forms a proboscis whose length may be as great as. Each of the numerous smaller branch-tubules has at its blind end a flame cell which is hollow and contains a mass of of the bod}'. it 167 nearly fills. The whole arrangement represents a complicated gastrovascular cavity whose project wall is endodermal. The trunks of the enteron have ticula many lateral.PHYLUM PLATYHELMINTHES buccal cavity which testine. one along each side of the pharynx. the movements appearing something like a flickering flame. It is attached only at its proximal or anterior end and is perfectly free otherwise. The pharynx is in the form of a cylindrical fold projecting through the full length of the pharyngeal chamber. and two lateral trunks that turn posteriorly. and extend nearly to the posterior end. one along each side many small branches and open by minute pores located just posterior to the eyespots. —Flame cell of planaria. or greater than. When it is extended or protruded through the mouth opening which it fills. blind extensions or pockets called diver- which greatly increase the surface exposure of the organ and among most of the other tissues of the body. There are two principal.

and sensory end areas of the epidermis. entirely is by asexual fission. female organs in the same animal consist of two ovaries located well toward the anterior. regular. It is com- posed of an outer circular layer just under the epidermis. This opens into the cloaca. transverse commissures. a small ganglion composed of a few nerve to the The branch nerves extend is surrounding tissue from The reproductive system reproduction is fairly well developed in most species Its except P. At each point where a is longitudinal cord. internal. besides joining The transverse commissures connect transverse commissure meets a cell the two longitudinal cords at from 15 to 20 points like the rungs of a ladder. auricles. common cavity called the genital atrium or geiiital which opens externally at the genital pore. these points. branch nerves. some authors object to calling arrangement a system. Some authors The describe glands which pour a seminal fluid into the system. These in turn lead posteriorly. longitudinal ducts. dorotocephala which rarely develops sexual organs. longitudinal layer just medial to a circular layer. Another advanced development is the "ladderlike" nervous system which consists of two contiguous lobes of nerve cells just ventral to the eyespots.168 TEXTBOOK OF ZOOLOGY cellular walls of the tubules as well as the flame cells arise in The this the mesoderm. oblique bundles of fibers and at the medial margin of the mesoderm is another ir. the copulatory organ. which rather characteristic of sedentary animals. is Another mesoderm organization the muscular system. Vasa efferentia are slender. and converge to form the penis or cirrus. bodies. a tubular oviduct leading posteriorly from each to join the genital atrium at a common point near its posterior end I . two ventrolateral longitudinal nerve cords. the longitudinal nerve cords. and gives branches to sensory areas of the head. enlarge to become seminal vesicles. The sexual reproduction is of other planarians hermaphroditic. By the alternate activity of these layers the animal is capable of great extension and contraction.. Under strict definition. thin-walled ducts leading from the testes to two larger. The male organs consist of numerous globular testes located in the parenchyma through most of the length of the body. the vasa deferentia. known as the cephalic ganglion etc. an outer longitudinal layer just medial to the circular layer. The double ganglion It is at the anterior is the central portion of the system.

— CerebraL (jancjlion Auricle LorKjitudinaL nerve cord Testis / Vasa efferenbia Lateral nerve lumen of pharynx Intestine Vos deferens l_( K-/rax^ •H : Oviduct -— ^ / Pharyngeal Mouth Seminal vesicle \_^^ chamber Penis 5eminal receptacle Genital pore Fig.PHYLUM PLATYHELMINTHES by way of the vagina. The system is . in the form of a blind tube with an inflated end. —Reproductive system of the planarian worm. 84. oviduct along its 169 There are numerous yolk glands joining each length a glandular structure of questionable func. Male organs shown on one tion. is connected with the genital atrium. side only. It has been suggested that the fertilized eggs accumulate and are retained here for a time.

There is no anus. and the diffusion of materials supplies all no circulatory system is necessaiy to transport nutriment. The flame cells absorb these wastes from the surrounding tissues and force the fluid into the tubules by the action of the cilia. being both intercellular and intracellular that is. It is significant that the reproductive system upon which the continuance of the race depends is highly specialized. The spermatozoa germ cells mature in the testes. Reproduction and Life History Sexually the individuals are hermaphroditic. process is similar to that of Porifera and Coelenterata. and this followed by the nervous system which relates the organism to its surroundings. Catabolism or dissimilation takes place in the cells by union of the oxygen with the organic components of the protoplasm. Here the process of digestion occurs in the cavity of the enteron. is performed by the branched enteron. no respiratory system (breathe through the skin). Since the diverticula of this system pene- trate all parts of the body. . while other food particles are engulfed by pseudopodia extended from cells lining the cavity and are digested in food vacuoles inside the cells. epithelium. Absorption and assimilation take place through the plasma memThe food is principally animal tissue with some plant matter. to the seminal vesicles where they . branes of adjacent other cells. The planarian worms and the representatives sess of this phylum pos- no skeletal system.170 TEXTBOOK OF ZOOLOGY notably quite elaborate. and no circulatory system. this succeeded by the digestive system responsible for nourishment of the individual. part of the food in the intestinal cavity is digested by secretions from cells in their walls. and it is found generally that the flatworms have a highly specialized reproductive system. no coelom or body cavity. Metabolism and The food may be parThe principal tially digested by a fluid produced in the pharynx. however. and oxygen is distributed by diffusion through the protoplasm and fluid-filled spaces of the parenchyma. Excretion or elimination of nitrogenous waste liquids is cared for by the flame cells and system of tubules. then pass through the vasa or male efferentia and vasa deferentia. so all indigestible material is egested by way of the Respiration is accomplished through the general surface mouth. ingestion takes place through the proboscis. this function. cells.

) PHYLUM PLATYHELMINTHES are stored in advance of copulation. published by The Macmillan Company. pass 171 Here they become organized the known as spermatophores. endoderm. Cross-fertilization is practiced by these Planarians have been observed to copulate with an apparent exchange of spermatozoa in the form of spermatophores. . animals. — Planarian Wandering cells " Ectoderm Wandering cells Endodenn Provisional Endoderm -Primitive gut Wandering pharynx — cells ceUs Vitelline Mouth —Development of Planaria later blastomeres from segmented egg Fig. stage E!gg surrounded by yolk. 2. cellular differentiation more advanced. and wandering cells. . S. 5. . (From Hegner. The ova mature in down the oviducts where yolk cells or nurse cells are added by the yolk glands. after blastomeres if. and probably from here to the uterus or seminal receptacle where Young planana batchinq tq(j capsule or cocoon cocoons and the young hatching. 85. a provisional pharynx. College Zoology. 1. embryo becomes flattened and ovoid. In copula- they are thought to be stored. lactea. into pockets ovaries. through the vagina to the genital atrium. 6. 86. four still later. have differentiated into ectoderm. after Lankester after Hallez. Vitelline cells Fig.

secreted by the genital atrium and known is as a cocoon. so fertilization likely occurs somewhere along this tube. blastula stage. many thousands of yolk to several zygotes. From one cells. each to is At breeding time zygotes are found surrounded by a large number of yolk cell in the atrium. —Fission as it occurs in Planaria dorotocephala. 87. this way spermatozoa may be other. Each yolk which contributes its store of nourishment to the egg cell it is attached. In the cocoon the embryo passes through cleavage divisions. These are expelled from the atrium and each sides of attached by a stalk to the under submerged stones or vegetation in the water. The individual constricts and then divides into anterior and posterior portions each of which forms the missing parts by rapid cell division. Spermatozoa have been found along the oviduct as far as the anterior portion. The axial . surrounded by become enclosed in a capsule-like shell Fig.172 TEXTBOOK OF ZOOLOGY is tion the cirrus or penis protruded from the genital pore to enter In transferred from each animal to the the genital pore and extend into the uterus of the other copulant. i Asexual reproduction by transverse fission occurs quite frequently when the mature animals become slowed down. cells and (nurse cells). gastrulation and even later stages before the cocoon ruptures and the small wormlike planarians escape into the water.

Regeneration This group shows remarkable powers of replacing lost or mutilated parts of the body. When the animal is young. i.PHYLUM PLATYHELMINTHES orientation of the tissue is 173 retained . ginning at the anterior there a sudden increase occurs. This process not funda- mentally different from budding in Hydra or strobilization in the Scyphomedusae.. will be A found in a later chapter more complete discussion of this phenomenon on Animal Regeneration. A piece from the middle of the animal will regenerhead portion at the anterior margin and a tail portion at the posterior margin. it is relatively short and the entire body.e. It can be cut into several pieces. is a gradient of decreasing metabolic activity until a level just posterior to the mouth is reached. longer. anterior end of the second individual. sesses a well-defined axial organization in The animal poswhich the "head" porBe- tion as usual has the highest metabolic activity of the body. Child. Such zooids are the result of successive functional isolations of the basal structure accompanying growth in length. A new center of dominance and increased metabolism is established just posterior to the point where this "head" dominajice fades out. it becomes and the entire metabolic rate decreases. Child of Chicago University. As the animal dominance over the grows older. and each piece will replace the missing parts about as the process is carried out in ate a fission. . but particularly the ''head." carries on a high rate The head at this time holds a length of the organism. an anterior portion develops in the position of the original anterior portion. older individuals there may be other such points of increased metabolism posterior to this first one. This means that the head loses its dominance over the entire length. and a posterior poris tion at the original posterior position. The retention of the axial orientation during fission has been explained by Dr. of metabolism. This graduation of the rate of metabolism along the principal axis of an axiate animal has been called an axial gradient by Dr. The level where the metabolic rate suddenly rises represents the point of fission or the This seems to indicate a kind of zooid organization in the animal. and here Posterior to this the decreasing gradient again folloAvs to the posterior tip of the body. In larger.

. and (7) permanent sexual reproductive organs. but the phylum includes a large number of forms. sheep tapeworm. dwarf tapeworm. hydatid worm. (5) central nervous system extending with the axis (1) distinct third of the body. (6) specialized gastrovascular cavity. principally Trematodes and Cestodes. which are parasitic Such groups as the in higher vertebrate animals.174 TEXTBOOK OF ZOOLOGY Economic Relations of the Phylum The planarians and other free-living flatworms are of practically no economic importance. liver flukes. tapeworm of horse. chicken tapeworm. margined tapeworm of dog. lung flukes. the mesoderm. (2) bilateral symmetry. pork tapeworm. (4) an excretory system of flame cells. each year. and fish tapeworm They cost many thousands of dollars and are all serious parasites. including man. blood flukes. common tapeworm of dog. intestinal flukes. A more complete discussion of this topic much debility will be found in the chapter on Animal Parasitism. (3) a germ layer. Phylogenetic Advances of Platyhelminthes Anteroposterior principal axis. gid worm. beef tapeworm.

no segmentation is present. there is a dorsal as well as a ventral nerve cord. to four feet. whipworm. Gordiacea. although some authors prefer to use only two. and several others will be considered in the chapter on Animal Parasitism. Class Nematoda (nem a to'da threadworm) is a group occupying almost every possible habitat capable of supporting life. is shed periodically like the molting of arthropods. and there also lack respiratory definite They is a total absence of cilia. and soil-inhabiting speand large numbers of parasitic forms living at the expense of other animals and plants. to more than a meter in length. The former will be discussed in some detail in this chapter. the flatworm not only in shape. They differ from This group is known as the . Trichinella. The three classes are Nematoda. Filaria. pinworm. In size they range from %o mm. Locomotor organs are found in a few forms. Chemical sense organs called amphids are nearly universal. some are found living in plant tissues and others live in animal tissues as parasites. and circulatory systems. There free-living. This is a very important class parasitically.CHAPTER XIII PHYLUM NEMATHELMINTHES unsegmented roundworms or threadworms. "horsehair snake. and there is no true coelom. and is deserving of considerable known forms are Ascaris (pigworm Some of the better or eelworm). but also in that the intestine has two openings. marine. fresh water. These worms are long. and salt water. fresh water. The majority of them are microscopic. slender animals whose bodies are more or less cylindrical but tapering toward each end. The skeletonlike cuticle. and Acanthocephala. Some of the Nemathelmiuthes (nem a thel min'thez. and locomotor organs." hookworm. but a few are macroscopic in size. threadworms) are free-living in soil. and eye worm. Classification Three classes are usually recognized. they are mostly dioecious. while eyes and tactile organs are common in the free-living forms. Guinea worm. The nervous — are many cies. true coelom. 175 . The group is very widely distributed attention. common to all. The range of length is from i/4 mm.

such as freezing. in the kidneys. loides Ascaroidea. high temperatures. — Order Trichinelloidea. and transmitted by certain insects. cells. the chicken worm. The genus Dioctophyme includes the largest roundworms. These organisms are transmitted by mosquitoes. The common known examples. the human pinworm. — This is a completely parasitic order. is structure of the free-living forms that of the parasitic forms. the common intestinal worm. droughts. This is an entirely parasitic group. The club-shaped esophagus is without a posterior bulb. Ascaridia lineata. habitats generally more complex than to They are adapted a wide variety of and can withstand many rigors of natural adversity. and other unfavorable conditions. elephantiasis (Fig. Order 8trongyloidea. Large numbers of free-living forms have not been named and described. Two distinctive characteristics are: (1) lack of bulb on esophagus. 390) through occlusion of blood and lymph This disease results in enormous swelling of the ai^ected parts. blood. Representatives of this class have an intestine but no proboscis. Order Dioctophymoidea. some reaching more than three feet in length. — This parasitic group has a peculiar is cuticle lining the esophagus. the Strongylus — roundworms of horses. Strongystercoralis. and muscle of chorda te animals.176 TEXTBOOK OF ZOOLOGY is system extend posteriorly. The hookworms of man. connective tissue. The males have caudal bursae with rays. Several Filaroidea are parasites of horses and dogs. and alimentary canal of mammals and birds. modified for living in such tissues as lymph. Enterohius vermicularis. and Syngamus trachea which causes gapes all in birds by obstructing the windpipe are common representatives. Some species cause vessels. composed of a circumpharyngeal ring from which cords The It is a sensory-neuro-muscular system. (2) lateral paired lips or entire absence of lips. and Toxocara canis of dogs are other familiar examples. outside of which a single layer of epithelial well- trichina (Fig. 396) and the whipworm are . Order forms. is the most abundant. Guinea worm. This is another parasitic group living body cavity. — It includes both parasitic and free-living Ascaris (Fig. Ascaris lunibricoides will be discussed later as a typical example of Nemathelminthes. eye worm. and Filaria are the common humaai parasites. 90). Order Filaroidea. another parasite of man.

They leave this host and take up abode in a terrestrial form like that of grasshoppers or beetles. The body is cylindrical and without lateral lines. In the adult worm the intestine is a straight tube. often without a mouth. it is likely proper to give them the rank of They are free-living as adults but as larvae are parasitic on May flies and other insects.PHYLUM NEMATHELMINTHES 177 Class Gordiacea (gor di a'she a. a knot). Gordius. There are four longitudinal spaces or sinuses . but the fundamental structure is quite — and therefore. After complete development the adult "hair snakes'' escape into different. puddle. the water of some stream. —Hair snake. These females again lay eggs in the water in long strings. class. Superficially these animals resemble the nematodes. a Fig. or watering trough. excretory organs. or circulatory system. The outer surface of the body is covered by a cuticle. but opening at the posterior end by an anus. 88. an aquatic roundworm. Some have no intestine at all.

in the adult female the lined with peritoneum. Structure of Acanthocrphala. second impression. published by McGraw-Hill Fig.) Sexually the group is dioecious with the gonads opening into the Fertilization takes place within posterior end of the digestive tract. 89.178 in the TEXTBOOK OF ZOOLOGY parenchyma. Neoechinorhynchus emydis. are the Gordius aquaticus and Paragordius varius of the group. Inc. rudimentary eyespots and scattered sensory over the body. Invertebrate Zoology^ first edition. the body of the female. common examples . The nervous system two lateral spaces are is composed of a midThere are cells ventral cord with a nerve ring at the anterior end. (From Van Cleave. Internal structure of the genital organs a young female nematode. — of Book Company.

The sexes are separate. Finally the embryos are discharged by way of the uterus through the genital pore which is located posteriorly and is the only external aperture of the body. A single ganglionic mass constitutes the central nervous system. females commonly reaching from eight to fourteen inches and males averaging six . but mice. rats. is There The male reproductive organs are the glands joining the cirrus which is testes and a set of cement held in the copulatory bursa at The bursa is capable of eversion. fish." which is absolutely parasitic in its habits. spines. A REPRESENTATIVE ROUNDWORM Habitat and Behavior The animal which is frequently studied as a representative of this phylum is Ascaris lumbricoides which frequents the digestive tract ing suitable food and environment. receptacle or sheath. bears numerous recurved hooks or and in many species capable of receding into a proboscis no digestive tract in this parasite. It is entirely External Anatomy This is a length of one of the largest nematodes. and its food is absorbed through the surface of the body even though it is covered with a cuticle. and in the posterior portion of the body. The adults are from a few millimeters to fifty millimeters in length and have an elongated. flattened body when found in group.PHYLUM NEMATHELMINTHES Class Acanthocephala (a kan th6 sef 'a la. but the female has no permanent gonads. ASCARIS. turtles. 179 thorn head) includes a "spmy-headed worms. Not only man. of men and hogs. as known the intestine of a vertebrate but become distended to a cylindrical shape when removed to some solution outside the body. dependent on its host for furnishThe only time this organism is at the mercy of the elements of nature is during the egg stage when it may remain potent for months or even years if it falls in an environment unsuitable for development. Egg masses develop early and completely rupture to produce a considerable number of embryos in the body cavity. sible proioscis is The protru- a peculiar and characteristic structure located at the It it is anterior end of the body. fact all classes of vertebrates serve as hosts for these animals. pigs.

by three lips. two in lateroventral positions and one dorsal. four longitudinal lines. 90. and one ventral. The genital pore in the female is located on the ventral midline approximate!}^ one-third of the length of the body from the anterior . two on the dorsal and Fig. These lips have small papillae on their surfaces. two lateral.— Male and female Ascaris or eel-worm. one on each of the ventral. TEXTBOOK OP ZOOLOGY Males are always more slender and have a curled The mouth is guarded tail instead of the blunt tail of the female. one dorsal. The shape of the body is generally The smooth surface is marked by cylindrical with tapering ends.180 to twelve inches.

modified from Leuckart. E. cross-section of tlie midregion of the body of female D. /?70uth circumesopfiagea/ There aterc/s ov/c^ucts I \ oyan'es pseudoooe/ ^ excretort/ p/)ari/r}jc ^er7ti. whose medial margins are rather baggy. longitudinal section of posterior portion of male female. are thickenings of the epidermis in the positions of the longitudinal lines.. Diagram of lateral view Fig. The excretory tubes follow the is lateral lines. is The body cavity lined externally of this animal a primitive or false coelom which by the mesoderm of the body wall and internally by the endoderm . and in the male the reproductive aperture and two penial setae or spicules are located just within this opening. Textbook of General Zoology. The body wall is composed of the smooth cuticle. whose cells run together. reproductive system of C.PHYLUM NEMATHELMINTHES extremity. 91.) . (From Curtis and Guthrie. Internal Anatomy thin.a//x>re /r?i>€sC>/r?6 i)oc/i/ yya// Ct/7a5 tiactas deferens se/7?//7a/ ves/ch • e/acL'/atory afc/ct pseadocoeJ psei/cfocoe/ cut/c/e 1 intestine ep/c/er/T}/5\ body ivcr// ^em/'na/ ves/c/e musc/e cutic/e ) seta/ soi inteet/'ne recta/r? ej(cretoru cana/ pe/ifcfl_ o//dact setae: a/7i/: '/Vi/sc/e oyan/ at/c/e ner/e oord ep/der/nh 3 Internal anatomy of Ascaris Iwmbricoides. the epidermis. . published by John Wiley and Sons. A. 181 The anus is located near the posterior tip of the body. Inc. and a thick layer of longitudi- nal muscle fibers. — . reproductive system of male. of dissection of female B. outer.

A contractile pharynx. This tube enlarges posteriorly to become the vas deferens which in turn enlarges still more before reaching the aperture to become the ejaculatory duct. These tubes join in the vagina. lined both laterally is This the simplest type of animal in which the is and medially with mesodermic peritoneum. Inc. There is a nerve ring gives off a large dorsal longitudinal nerve around the pharynx which and a large ventral testis is longitudinal nerve. like structure which is much coiled in the cavity. lies in In higher forms the outer coat of the intestine is The alimentary canal is quite straight and simple and There is no need for the dorsal part of the body cavity. which acts as a pump. is TEXTBOOK OF ZOOLOGY Ordinarily the coelom. longitudinal duets open externally by a single pore near the anterior end of the body. 4. 92. The narrowed posterior portion is the rectum and leads to the anus fertilization /Tjemi^ranC' Sfye// ze/^ade Fig. is great specialization of the digestive system since the food taken from the digested material in the intestine of the host. (From Curtis — ovum. published by- John Wiley & Sons. at the posterior portion of the body. . when fully developed.) 182 of the intestinal wall. B. of Ascaris lumbriand Guthrie. body cavity or coelom mesodermic. found. draws fluid into the long epithelial intestine from which it is absorbed by the other tissues. which is a short tube leading to the genital pore. Textbook of General Zoology. and amoeboid spermatozoa.. There are usually four other smaller longitudiIn the males the a thread- nal nerves and some connectives. In the female the threadlike ovaries join the coiled oviducts which lead forward and join the two uteri. The two laterally located. after Leuckart.. Fertilized coides.

long.000 eggs. The life history is completed only in case the eggs are swallowed by a susceptible host. grow to maturity in two to two and one-half months.000 eggs. the larval worms. Relations to Man is Heavy infestation in is man may cause severe hemorrhages and set up pneumonia that often fatal. Drugs like chenopouium. and other similar symptoms. these larvae and back stomach and intestine. bring on coma. in certain cases the organisms may even tangle in masses and block the intestine until surgical operation is necessary The toxic substances from these parasites may to remove them. make their way to the throat. Anemia often the result of such infection. 2 to 3 mm. the daily production is computed to be something like 200. and at this time the spermatozoa are introduced into the vagina of the female to fertilize the mature ova in the oviducts.000. After reaching the intestine. These eggs are so resistant that they can be successfully cultured in 1 to 2 per cent formalin. They hatch in the small intestine of the host and then go on a ten-day journey by way of the blood stream to the By burrowing liver. and thence to the lungs. and hexylresorcinol have been used successfully under physicians' directions as a cure. convulsions. They likely live a little less than a year in to the the host. delirium. A mature female may contain as many as 27. Effective sanitary disposal of fecal material is the most successful preventive. santonin. and they may be stored successfully for four years in a refrigerator. Based on this figure. nervousness. Some workers have reported that each female worm in an infected host may produce a crop of eggs in excess of two thousand per gram of feces. thence to the heart. esophagus.PHYLUM NEMATHELMINTHES Reproduction and the Life Cycle 183 The animals copulate. . These eggs pass from the host with the feces. out from here.

These individuals are called zooids of which the as polypide. soft parts are known it They are within the primitive coelomic muscles make The presence of retractor withdrawn into the There are some smaller vaselike part of the chitinous skeleton. the individuals of which are associated in a treelike colony that lives attached to some object in the water. which are similar to colonial hydroids in their manner of growth and forms. but many phylum rank. was a long time after their existence was laiown that they were separated from that group. small groups of the unsegmented worm type. are marine. although there are a few fresh water forms. TROCHELMINTHES. AND CHAETOGNATHA chapter. the wall of which is the zooecium. In ex- ternal appearance they resemble certain of the corals and hydroids. subclass. The second subclass Endoprocta is characterized by the circlet of oral tentacles which also encloses the anus. enclosed in this area. usually considered a authors prefer to give each of the classes fication of the latter plan phylum name. animals) includes a group of co- Class Bryozoa (bri 6 zo'a —moss lonial animals often called Polyzoa. individuals whose shape is similar to that of a bird's head and whose bodies are smaller than the other zooids. because they are These groups are rather conveniently considered in the same more or less isolated. Bugula Bugula is a common marine genus. The subclass Ectoprocta includes forms in which the mouth is surrounded by tentacles and the anus is not It Bugula is an example of a treelike type of this Another type is one that grows as an incrusting organism. MOLLUSCOIDA This is the name It is of a group composed of two classes. These are called avicularia. The justi- may be questionable. as they are treated here. possible for each zooid to be 184 . It is true that their Nearly all bryozoans structure distinguishes them very readily. Their cavity.CHAPTER XIV MOLLUSCOIDA. and they are found on the surface of the colony.

TROCHELMINTHES. Yihramdaria (vibracula). . They are thought to be variations of the avicularian stitute modification. AND CHAETOGNATHA 185 movable jaws seem to serve as grappling hooks Avhich operate to keep the colony free from other small organisms and debris which may be present in the habitat. which con- Esophaqus. another modified type of zooid. Bugula. showing the structure and habit of life of two zooids from a colony. whiplike appendages.— MOLLUSCOIDA. are filamentous. 93. -Jntestine Avicularium / jaws open Muscle to body wa// ' Muscle _run/cu/uj (Mesentery) Ooecium Fig. a marine bryozoan.

nates at the anus. The body triploblastic and there- . regular zooid is located at the free end and is surrounded by a tuft of ciliated tentacles.) is for each. 94. The tis- held in place by strands of mesenchymatous tissue Special strands of this is extending from the wall of the coelom.186 TEXTBOOK OF ZOOLOGY The mouth of the larger. This arrangement is known as the lophophore and has the shape of a horseshoe when expanded. sue are termed the funiculus. the digestive tube is U-shaped and termi- 3^ liaublt K rXUAVO'ELLA. Kabi-1- " CKI^TATELLA life — Three forms of fresh-water Bryozoa showing the habit of (Courtesy of General Biological Supply House. which digestive tract is located just outside the lophophore. f- PECTIMATELLA Fig. Within.

They withstand the rigors of winter and are stimulated by it. the zooid of which will form a neAv colony by budding. These fresh water types may be developed from winter eggs. They are produced in autumn and may either float on the water or sink to the bottom. which in the is supported by a secretion of is calcium carbonate. the lophophore is a conspicuous and characteristic structure of this type of animal. however. and the slimy Pectinatella. Class Brachiopoda (brak i op'o da arm and foot) is a group of marine forms. gullet. The brachiopods. stomach. while the molluscan valves are lateral. So far as is known this group has little if any economic value. nerves extend to the tentacles. or new individuals may be produced as internal buds.MOLLUSCOIDA. For this reason they are sometimes confused with the clamlike molluscs. enclosed in shells. and ventrally directed intestine. the individuals of which possess bivalve shells. called the broad-pouch or ooecium. or in the lining of the coelom The branching Plumatella. The tip of the beaklike valves is penetrated by a foramen which serves as an opening for the peduncle. it is a free-swimming. lophophore. The nervous system is centered in a ganglion or mass of nerve cells located in the region of the mouth and from it. This larva becomes attached and transforms into a parent individual. TROCHELMINTHES. whose skeleton form of a gelatinous mass. — The digestive tract is U-shaped and is composed of the mouth. This . are the two forms most frequently found in fresh Avater. The shell is secreted by a mantle which lines the valves. AND CHAETOGNATHA fore composed of ectoderm. This organ is composed of two coiled appendages which bear numerous ciliated tentacles. These buds are called statoMasts. When the embryo escapes. eudoderm. This fleshy organ makes permanent attachment to some object in the water. have dorsoventral valves. Its form resembles certain adult Rotifera. Reproduction is accomplished either by budding or sexually. ciliated larva which is similar to the trochophore larva found as a developmental stage of certain Annelida and Mollusca. The cilia produce water currents in the longitudinal groove and carry food particles to the mouth. Ovaries and testes cavity. make their appearance either in the funiculus and fertilization occurs in the body The early development goes on in a modified region of the zooecium. Internally. 187 and mesoderm.

About two pairs serve in excretion. this Gastrotricha constitutes another small division of group but will not be discussed in detail here. . of little economic significance. lenticularis are commonly studied They are entirely marine and represent an old line of aniis group There are relatively few modern forms in existence. mals. true coelom present. 95. times they were called "wheel animalcules. thence to nephridia and outside. and a free-swimming. The sexes are distinctly separate and mature germ cells are discharged into the coelom. Company. College Zoology. Mouth to —Diagram of a sagittal section of a brachiopod show (From Hegner. Magellania flavescens and M." There is very little difference between the trochophore larva of this group and the adult animal. but the septa are a little bit difficult to distinguish. Extensions of the coelom enter the arms and mantle of this type of animal. and produce the gonads also. The entire tube is lined is internally with ciliated epithelium. Fertilization takes place in the water.) forms. A segmented. published by The Macmillan internal organs.188 tract ends blindly in TEXTBOOK OF ZOOLOGY many bracliiopods. ^ophophore Digestive gland Adductor muscle I / i Stomach Heart Dorsal valve Dorsal mantle ' Intestine Nephridium Muscle Pig. ciliated larva of nephridia are connected with the coelom The coelomic cavities hatches from the egg. The TROCHELMINTHES The rotifers (Rotifera) are common examples el of this group known In early as the Trochelminthes (trok mm'thez —wheel worm).

and a few of the sea. too. and extending posteriorly is the smaller cylindrical intestine which leads by way of the cloaca to ticates the particles of food. trunk. 189 them inhabit They are microscopic in size. tion. The internal organs include several systems which lie in the rather extensive body cavity or false coelom. TROCHELMINTHES. and two winding nephridial ducts which lead posteriorly to a contractile bladder. A short tubular esophagus leads to the pouchlike stomach. the anus. Nearly the entire internal surface of the alimentary is which aid the movement of the food material through it.) The flame cells are distributed in the body wall from the anterior. This bladder is pouchlike and empties into the cloaca. mouth which receives other small organisms as It is a cavity leading to the pharynx. It and also receives products of the urino- opens externally by way of the anus. wheels. composed of chitinous jaws. are located some cement or adhesive glands which assist the animal in adhering to most surfaces.MOLLUSCOIDA. The foot as a whole serves in locomotion. The body of a rotifer is bilaterally symmetrical and can be divided into head. At the posterior end is the tail or foot Avhich is forked or toelike in many species. canal lined with cilia The excretory system of flame cells. often creating two sets of water currents so as to resemble two rotating They are responsible for obtaining food and for locomoThe mouth is located in an anteroventral position. and foot. and they are often associated They are very resistant to adverse conditions produced by drought and may be distributed in dry form. in many forms. The digestive system begins anteriorly at the food. AND CHAETOGNATHA Rotifers are plentiful in fresh water. Inside the pharynx is a mill-like organ or mastax. It is covered externally by a cuticle. The so-called head is rather largely a troclial disc comwith Protozoa. (The name cloaca is applied to any cavity which serves as the posterior portion of the alimentary canal genital system. posteriorly. which mas- The movements of these jaws may be observed in certain rotifers when alive. pushing the animal along. posed of various modifications of two bands of cilia over the anterior end and around the mouth. The stomach and intestine are lined internally with endoderm. Some authors believe that the bladder functions also to assist in . is well developed and consists of a number similar to those of flatworms. The trunk tapers toward the posterior and contains numerous organs. Here. These cilia are in active motion.

b.) — stitute the nervous system. A common fresh-water rotifer. located in a dorsoanterior position and several nerves extending to sense organs and muscles con- . The oxygen is received into the body with water which diffuses through the body wall. A large ganglion.Brain — Pharynx —Eye Tooth "Mastax Salivary gland Salivary gland Salivary gland Stomachintestine - Vitellarium -Egg Flame Blasendarm" _ cell Excretory canal Contractile bladder -Rectum — Anus - Foot-gland. College Zoology. Philodina roseola. dorsal view. published by The Macmillan Company. . 96.Tactile organ -r\ . cuticle over a thin layer of ectoderm. ventral view. (From Hegner. a. dermal tissue The body wall is composed of an outer Under this layer is the mesowhich includes mesenchymatous cells and muscle fibers. after Hickernell. Fig. showing internal structure.190 respiration TEXTBOOK OF ZOOLOGY by collecting the excess water and carbon dioxide.

AND CHAETOGNATHA This group of animals is 191 and dimorphism (striking difThe males are usually much smaller and may even live as a parasite on the female. The less highly developed males possess a single testis in which spermatozoa are produced. or even a Rotifers may be oviparous (lay eggs). In the female of most species there is one ovary which produces the eggs. i —Life cycle of the rotifer. is ferences in form of the two sexes) present. produce females only. Connected with this gonad is a yolk gland or vitellarium. the fertilized eggs are usually carried in the body for a time and then discharged by way . or the young if born alive. 97. Hydatina. few are viviparous. In a few forms there are two ovaries with no distinct bisexual. The males lack a well-developed digestive system and are therefore very short lived. and develop parthenogenetically. At this time the eggs of the female are fertilized.MOLLUSCOIDA. The smaller type produce males. and require The eggs when mature. of two sizes. TROCHELMINTHES. zation. In some there is a peculiar type of copulation during which the special copulatory organ composed of a protrusible cirrus seems to perforate the body wall of the female. ovo viviparous. In oviparous forms. The eggs produced during the fertili- winter are thick-shelled. yolk gland. are carried by the tubular oviduct to the cloaca and are discharged to the exterior through the anus. Female Large Egg - Small Egg / / Females Males Sperm Late SuKiner Winter Eggs (Fertilized) Pass Winter in thick shell Females Females / Eggs'" / Females I ^ Eggs ^ k f Parthenogenesis Parthenogenesis Many Generations Manv Generations Females Small Egg i Svunner season "i ^ Large Eggs Fig. The eggs produced during the summer are thin-shelled.

There is considerable similarity between the adult rotifer and the trochophore larva of some annelids. drawn to show internal organs. an arrowworm. They then lie dormant and inactive in the water for a period before hatching. Nemertinea and others.— TEXTBOOK OF ZOOLOGY 192 of the oviduct.) mollusks. This resemblance has prompted the theory that the above groups are rather closely related to the . published by The Macmillan Company. Sagitta hexaptera. College Zoology. 98. Spines -Hooks -4 . rotifers.Oviduct -Fin Genital -pore Anus Vas deferens —Seminal vesicle • Testis Fig. Brain "Mouth Alimentary canal (_ _ Ventral ganglion — Ovary . (From Hegner.

two eyespots. rapidly. The body is divided into three divisions head. The fertilized ova become small adults without a typical ciliated larval stage. trunk. The ovaries the are located in the posterior portion of the body cavity and the mature ova are carried to the exterior by an oviduct on each side. jaw). TROCHELMINTHES. and other sensory organs. The nervous system consists largely of a supraesophageal ganglion or brain. or excretory structure. which enlarge to become seminal vesicles near the aperture.MOLLUSCOIDA. fins supit port the animal at the surface and also possible for to The prehensile mouth with its bristles have given the animal the name of "bristle jaws" in addition to other names. These are separated by septa and the coelomic cavity is separated into right and left cavities by a longitudinal mesentery. that is. Internally is a tubelike intestine which extends from the mouth move about : at the anterior. These animals are lacking in circulatory Each individual tozoa. Sagitta is the best known genus of the group. and tail. AND CHAETOGNATHA Class Chaetognatha (ke tog'na tha 193 —horse's make small marine worms are often called arrowworms. The spermatozoa are discharged into this cavity and delivered to the exterior by a pair of slender vasa deferentia or sperm ducts. . ventral ganglion. is capable of producing both ova and spermahermaphroditic condition prevails. branch nerves. to the anus located near the base of the caudal fin or tail. These and they are well Horizontal it — adapted to livmg at the surface of the ocean. The testes are located in the cavity of the tail portion. mane.

the most classes: — This class includes commonly known forms of the phylum. or living in especially prepared tubes. The Phylum Annelida may be divided into four Chaetopoda I. Order 1. (2) metameres or segments. such as those mentioned in the beginning of this chapter. type of free-living habitat where moisture These worms are found in almost every There are is present. Gephyrea Order 1. chitinous appendages in most cases. Echiuroidea Order 2. hair and foot). (4) segmented. (3) a ventrally located segmental nervous system. Even then. either swimming freely. (5) an excretory system composed of nephridia.CHAPTER XV PHYLUM ANNELIDA (By J. are common to the entire phylum. Teague Self. Freshwater streams and ponds are inhabited by numerous forms of annelids. Oligochaeta Class II. and (6) a nonchitinous cuticle covering the body. There are marine. Polychaeta Order 2. From this it is evident that the phylum as a group has become adapted to many varied living conditions and comprises one of the large groups of the animal kingdom from the standpoint of numbers. 194 . these distinguishing features are sometimes modified until only an expert can recognize them. and moist soil is usually alive with terrestrial earthworms. In the process of adaptation the annelids have become diversified in their anatomical features until only a very few characters. University of Oklahoma) The Phylum Annelida (a nel'i da. Sipunculoidea Class Class Chaetopoda (ke top'O da. Hirudinea Class IV. non jointed. bur- rowing in the sand. many forms which live in the ocean. Archiannelida Class III. form of a little ring) comprises an extremely large group of worms characterized by (1) the presence of a coelom surrounded by two layers of muscle.

. Typically. . marine annelid with branching gills Hirudo medicinalis large medicinal leech (upper center) Aphrodita ornata. . Arenicola cristata. and they all possess setae (chaetae). From left to right. the nephrostome. which surrounds the straight digestive in locomotion. tract. each coelomic space possesses a pair of nephridial tubules which communicate with the coelom at one end by means of a ciliated. The setae are or bristlelike appendages on the body segments. PHYLUM ANNELIDA fresh-water. funnellike opening. sand worm or clam worm Lumb7-icus terrestris earthworm or angleworm. (Courtesy of Denoj-er-Geppert Company. chitinous and are embedded make them movable and therefore useful muscle attachments which The coelom. sea mouse (lower center) Nereis vii-ens. 195 and terrestrial forms. — . The nephridia remove nitrogenous waste materials from the coelomic cavities and from the blood.. . The other end opens to the outside by means of a nephridiopore.) worm Fig. is divided between the segments by partitions known as septae. . . 99. Representative annelids. They bear in pits of the integument. lug Amphitrite ornata.

Reactions which require immediate coordination of the whole body are controlled by three giant nerve fibers which run through the entire length of the nerve chain. which may be studied as a representative form. or aciculum. which also pulsate. The principal vessels of the circulatory system are a dorsal one. One of the most widely known forms of this group is Nereis virens or the clamworm. These are connected in the an- body by a varying number of paired. segmental The dorsal vessel exhibits wavelike contractile movements (peristaltic contractions) which force the blood anteriorly. and each surrounds a large seta. and a ventral one through which the blood moves posteriorly. so that the stimulation in one segment automatically stimulates the adjoining ones. a green pigment known as chlorocruorin is found in still others no known blood pigment occurs. The class Chaetopoda may be divided into two orders. however. fleshy parapodia. has little to do with the coordination of different parts of interseg- mental and intrasegmental reflexes. intestine. namely. (1) the Polychaeta and (2) the Oligochaeta. . Hemoglobin is suspended in the blood plasma of some Chaetopoda in others. through which the blood moves forward. and other organs. The parapodium is divided into a dorsal notopodium and a ventral neuropodium. tudinal layer and an outer circular layer of muscle.196 TEXTBOOK OF ZOOLOGY of each segment is The inner body wall made up of an inner longi- Segmental nerves which are derived from segmental nerve ganglia innervate the metameres and coordinate the movements of the body. then backward through the ventral vessel to the skin. stitute the It — possesses many setae (chaetae) located in In this case the parapodia with their setae con- segmental appendages. At the anterior end is the brain. The primary function of the suprapharyngeal and subpharyngeal ganglia is to relay sensory impulses. The segmental ganglia communicate with each other through connections extending from one segment to the other. . which serves as a point of attachment for . The polychaetes (majiy bristles) are typically marine Chaetopoda. which is composed of a suprapharyngeal and a subpharyngeal ganglion joined together by a pair of commissures. Order Polychaeta. The latter passes through the hearts. The principal vessels and hearts have valves on their inner surfaces which prevent the blood from flowing in the wrong direction. The brain. terior region of the hearts or connectives.

. which may be everted by use of protractor muscles to form a sort of prohoscis. anal cirrus.) when the pharynx is extended. Fig. present. anterior end and posterior end. peristomium 7. anus. 10. (Courtesy of General Biological Supply House. 17. 2. . XI. 197 A dorsal and a ventral cirrus are usuallyThe notopodium and the neuropodium each have a large group of setae. The suc- except the posterior one which bears a pair of ventral ciVrt extending posteriorly. 100. 15. The pharynx is equipped with muscles by which it can be everted. The peristomium of tentacles. 13. neuropodium. 16. which bears a pair of eyes. gill plate. ventral cirrus. A. aciculum. The pharynx leads into the relatively narrow esophagus . notopodium. The head of Nereis seems to have resulted from the fusion and It is composed of a prnstoand two pairs constitutes the first segment and bears four paii*s of cirri or tentacles. dorsal cirrus. 5. 1. palp. lateral tentacles 6. eye 5. The jaws serve is to in capturing small organisms and crushing anything which ceeding segments are all alike be swallowed. 12. and a pair of chitinous jaws which protrude specialization of the anterior segments. prostomium 4. . . segment 8. terminal tentacle. The mouth opens directly into the muscular protrusible pharynx. The parapodia are used principally as locomotor and respiratory organs. — .PHYLUM ANNELIDA the parapodial muscles. parapodium 9. . B^ parapodium (enlarged). The digestive tract consists of an essentially straight tube. mium. External anatomy of Nereis virens and parapodium. . a pair of palps. setae (chaetae) Ui.

College Zoology. The remainder of the digestive tract is a straight intestine which continues to the last segment. Internal anatomy of Nereis virens.198 TEXTBOOK OF ZOOLOGY which extends through about six segments and which has a digestive gland opening into it from both sides. 101. is composed principally of a dorsal and a ventral blood vessel joined in each segment by a pair of connecting The blood is forced anteriorly through the dorsal vessel and .) — The circulatory system vessels. Prostomlal Prostomlum tentacles Feristomlal tentacles Parapodia Pharynx CEsophageal glands (Esophagus Intestine Nephrldla Dorsal vessel Ventral vessels Nerye-cord' Fig-. where it opens to the outside. published by The Macmillan Company. (From Hegner. after Parker and Haswell.

The nephridium consists of a ciliated funnel. the common earthworm. body except the peristomium has two nephridia from the coelom to the outside. nephrostome. extends posteriorly on the ventral side of the the anal segment. and vary in number up to 175. Two is eyes receive nervous connections from the brain and the animal detect apparently able to moving objects. Order Oligochaeta. which metamorphoses into the adult animal. Its 199 movement is effected by wavelike contractions in the walls of the dorsal vessel. or metalinear arrangement. in length meres. features represent so well those common EARTHWORM The body of Lumbricus terrestris varies from six to fourteen inches and gives the appearance of a number of rings joined in a The rings are the body segments. In the central nervous system there are two suprapharyngeal ganThese are connected by means of comglia dorsal to the pharynx. It reaches the parapodia and digestive tract through lateral branches of the ventral vessel and is then returned to the dorsal one by parietal branches. the nephridiopore.PHYLUM ANNELIDA passes posteriorly through the ventral one. In the adult the number of segments from the anterior end to the posterior end of the clitellum . walls of the coelom and there are gonads in all the segments except The sex cells arise from the and when ripe pass to the outside. composed of segmental ganglia joined by intersegmental connections. (Dipocardia) but is — used here as an example because its it is so well to known and because the entire order. Lumbricus is not as common in the Southwest as are other large forms of earthworms. which is used almost universally as a laboratory specimen. A nerve chain. fertilization The fertilized egg develops into a trochophore larva. The nephridia serve to convey the excretory and reproductive products to the outside. The best known example of the order Oligochaeta is Lumhricus terrestris. body to Lateral nerves from the ventral nerve chain in- nervate the various organs of the worm. taking place in the water. and a coiled tubule which ends in its external opening. missures to the suhpharyngeal ganglion ventral to the pharynx. The Each segment of the opening directly sexes are separate those in the anterior end of the body.

In studying the earthworm it is customary to number the segments with Roman numerals beginning at the anterior end. The prostomium mere. (Prom Wolcott. It is projecting out over the mouth. not considered a true meta- The first segment is incomplete due to the opening of the Pros'f'omi'u/n Openjngf ofoviducf ^Open!n0 of "^Mpis deferens ^Sem/'ncfl % groove XXVIxxxrr-i zm . This simplifies the study because external as well as internal structures are definitely related to specific seg- . ventral view. — External anatomy of earthworm. Company. is a sort of knoblike lobe at the anterior end. C//fe//u/7i xxxvir- ^Sefae A'lus Fig.200 TEXTBOOK OF ZOOLOGY is remains constant. Animal Biology. segments in published by McGraw-Hill • roman Book Inc. This because growth is accomplished by the addition of segments poste- rior to the clitellum. while the number posterior to this varies.) mouth through its ventral side. numerals. 102.

It contains numerous minute pores through which secretions of the unicellular glands beneath are poured and through which gaseous exchanges between the blood and moist soil can take place. and XVII and XVIII. The body of the earthworm is covered by a thin. The openings of the vasa deferentia (sperm ducts). minute pores. It serves also as a protection against physical and chemical injury to the animal's body. the digestive tube being the inner one and the body wall the outer one. the function of which is to secrete the cocoon in which eggs are deposited during reproduction. are located. one on each side. segments XXX. The constricted regions dividing the segments on the outside correspond to the positions of the septae which divide the coelom into separate segmental compartments. in the XV. They are moved by protractor and retractor muscles and serve to help the worm move through the soil.PHYLUM ANNELIDA ments. . These coelomic divisions communicate with each other of pores in the septae so that the clear fluid by means which fills The septae are absent between segments I and II and incomplete between segments III and IV. They are fine. if cut gives the general appearance of a tube within a tube. In sexually mature worms. transparent cuticle which is secreted by the epidermal cells just beneath it. The space between them is the coelom. and one between X and XI. or XXXII to segment XXXVII are swollen to form anterior part of segment the clitellum. one pair in the groove between segments IX and X. The walls of the coelom are lined by a the coelom can circulate freely. 201 The openings of the oviducts through -which the eggs pass XIV. thin layer of cells known as peritoneum (mesothelium). Each segment except the first and last bears four pairs of chitinous setae. stiff bristles which may be located by passing the hand lightly over the worm. XXXI. one on each side of segment The pores of the seminal receptacles occur in pairs. of the earthworm. A pair of nepliridiopores (the external openings of nephridia) is situated on the posterior ventral side of each seg- ment except the first two or three. a sort of saddle-shaped structure. which to the outside are seen as convey sperms to the outside. Internal Anatomy open along the mid-dorsal The body line.

Mosby Company. Diagram showing reproductive system and nervous system in segments VIII to XV of an eartliworm. (From White. and XII. General Biology. The union of the vasa effer- . testis. 103. V. XI. — duct (vas deferens) opening in the fifteenth somite. The seminal vesicles have been cut away in somites X and XI to disclose the testes and sperm funnels. uals they The seminal vesicles are three pairs of light-colored bodies located in segments IX. published by The C.) are seen to contain the various stages of developing spermatozoa coming from the sperm mother cells. ovaries are female organs. nerve cord ov. septum between two somites sp. oviduct. and and the testes and seminal vesicles are male In sexually mature individ- organs. they Fig. . ovary sf. es. seminal receptacle. seminal vesicle. sperm . sr. oviducts. If their contents are examined with a microscope. sv. t.202 TEXTBOOK OF ZOOLOGY Reproductive Organs The earthworm is hermaphroditic. may extend back through the septae as far as the fifteenth segment. the organs of both sexes being present in every animal. . The testes are the two pairs of very minute bodies projecting into the seminal vesicles in segments X and XI and cannot be seen without first removing the dorsal part of the seminal vesicles. egg sac nc. . The seminal receptacles. seminal funnel sm. . vd.

. General Biology. In segments X to XII three pairs of yellow lateral pouches open into it. pharynx pr. 104. vessel eo. . dorsal blood . . cr. I to XXI. peristomium (From White. . . somites pJi. the animal. prostomium. of segments dv. . nephridium sb. subpharyngeal ganglion st. the The esophagus is a straight narrow tube extending from pharynx through the fourteenth segment. These are the . into the bulbous. . gizzard . stomach-intestine pe. . n. Fig. The ovaries are two minute bodies located one on each side of segment XIII. calciferous an earthwormcrop region dissection of buccal cavity glands in be. esophagus . of small white bodies located in segments Digestive System The mouth cavity extends through segments I to III and leads muscular pharynx which extends through segment The pharynx plays the part of a sucker in securing food for V. . The seminal receptacles are pairs IX and X. — Diagram of dorsaleg.) g. II-XXI.PHYLUM ANNELIDA 203 entia coming from the vesicles on each side forms a single pair of vasa deferentia in segment XII.

chloragogue cells cir. Cross section of the earthworm through a posterior segment. . (Prom White. intestine branch of dorsal vessel loti. the secretions of whicli help to neutralize the acid organic matter taken as food. to play a part in the excretion of nitrogenous wastes. which extends through segment XVI.204 TEXTBOOK OF ZOOLOGY calciferous glands. nerve cord sb. ventral blood vessel. ty. . any kind of organic The animals remain in the during the daytime and work their way through I I I I I .) . typhlosole vv. coelom cti. . . epidermis int. known as chioragogen cells. . intestine is not a simple tube but has a large fold. . lateral blood vessel la. dorsal Id. thin-walled structure. The sole. circular muscle fibers coe. . — . . . cuticle dv. They are generally believed. whose function is doubtful. The food of the earthworm consists of almost its matter which may soil pass through digestive tract. . the intestine is covered with a layer of brown cells. •<lv The coelomic side of Fig. longitudinal muscle fibers n. 105. lateral neural vessel ep. which opens to the outside through the last segment. nephridium nc. . . ch. I . General Biology. a larger. however. subneural blood vessel se seta. The esophagus opens into the crop. the typhloits protruding into lumen from the dorsal side giving it more absorptive surface for the assimilation of food. A thin-walled intestine extends to the anus. This is followed by the muscular gizzard in segments XVII and XVIII.

hence the nervous system receives the most highly oxygenated blood. where it flows posteriorly. (6) the typhlosolar vessel. It is then passed into the crop and is stored there long enough for the secretions of the calciferous glands to neutralize the organic acids which may be present in the food. An- . as seen in a living specimen. At night they come to the surface of the ground. (4) the subneural trunk. Food is drawn into the mouth by suction pro- duced by the muscular pharynx. usually remaining partly within or very close to the burrow. In the intestine food up . secretions Circulatory System The blood of the earthworm consists of a clear liquid plasma in which there are numerous colorless cells. where it receives the secretions from the calciferous glands. the typhlosole into and then returns to the dorsal vessel by way of The blood flows from the intestine through the dorsal vessel by dorso-intestinul vessels. The principal ones are: (1) the dorsal blood vessel. made with the water in the moist and in the nephridia the nitrogenous wastes are removed. and feed on dead organic matter. From the lateral vessels it passes into the subneural. A complicated system of blood vessels makes up the circulatory path of the blood. In the pharynx it receives the from the pharyngeal glands and is then passed on through the esophagus. such as leaves. (3) the ventral blood vessel.PHYLUM ANNELIDA it 205 by passing it continually through the digestive tract. is due to a pigment known as Jiemoglohin suspended in the plasma and not in the corpuscles as is the case in many animals. the uephridia. The red color of the blood. and the intestine. the parietal connectives. The paired which distributes it hearts receive the blood from the dorsal vessel and by pulsating move- ments force into the ventral vessel to the is body taken wall. It is then passed into the gizzard. it integumentary teriorly and forces it The dorsal vessel conveys the blood analong by waveiike contractions. and (7) the intestinovessels. The lateral neural vessels receive freshly oxygenated blood from the skin. in the skin gaseous exchanges are soil . (5) the parietal vessels. From the gizzard the food is passed into the intestine where digestion is completed and the absorption of digested materials is accomplished. (2) the paired hearts (usually five) in segments VII to XI. This process is aided by sand grains which are swallowed along with the food. where it is ground by contractions of the muscular walls of that organ.

which are found in each segment except the by the paired first two and The cilia of A . published by The Macmillan Company. the nephrostome create a current which takes the fluid containing . Dorsal vessel Typhlosolar vessel Ventral vessel Sub-neural vessel Parietal vessel Fig. and B. IX. 106. of excretion is cared for principally nephridia.206 TEXTBOOK OF ZOOLOGY terior to the hearts the dorsal vessel carries the blood posteriorly and the ventral is vessel carries it anteriorly. Since the animal always lives is in a moist environment. after Benham. after Bourne. — Sub-neural vessel Septa Dorsal vessel Septa jl IX Septa VIII Intestlnotegtimentary vessel Dorsal vessel CEsopbagus Typhlosolar vessel Ventral vessel Kephridium Ventral vessel Sub-neural vessel £ Lateral-neural vessel Sub-neural vessel Afferent intestinal vessel Efferent Intestinal vessel. and a nephridiopore. transverse section of the same region. (From Hegner. . Circulatory system of the earthworm.) — X . Dorsal vessel -— vessel . longitudinal view of vessels In somites VIII. single nephridium consists of a ciliated funnel (the nephrostome) a thin coiled tube.Ventral vessel fieart Intostlno-tegxuueniai. College Zoology.. A. The circulatory system equipped with numerous valves which keep the blood from flow- ing in the wrong direction. . this type of respiration possible. Excretory System The function the last one. Respiratory System Respiration in the earthworm is carried on through the skin which is well supplied with blood. longitudinal view of the intestinal region D. transverse section of same region C.

nst. and nonciliated portions of the intracellular canal . after published h^ r^^t'^^^"" Meisenheinmer. nephrostonie^ (internal openfneT ?../**in^ter. Also the wastes in the blood are excreted by way of the nephridial tubules.PHYLUM ANNELIDA 207 nitrogenous wastes from the coelom into the tubule where it can pass to the outside through the nephridiopore. ciliated 107. The nephro- stome is located in the posterior part of the segment and leads into the tubule of the segment just posterior to it. ext. (From Parker and Haswell Textbook of Zoology. : . after MazE^irski.'S-. The nephridium coils two or three times before reaching the nephridiopore.^iii T^'' ^anal.^. Tl^t a..) ^. ampulla between the 6il. mjWikherf by The MacmiUan Company. external opening ^nephridiopore) Zc^ nn^Mr'^f'^^^'^ ^''T' septum between somites.— structure of a nephridium from earthworm. ciliated canal coe coelomic epithelium.

208 TEXTBOOK OP ZOOLOGY The Nervous System The "brain" ganglion. Nerves from these two ganglia innervate the first three segments and the prostomium. and the suhpharyngeal The ventral nerve cord extends posteriorly the length of the body with a ganglion and three pairs of nerves in each segment. —Anterior is portion of the nervous system of an earthworm from dorsal view. The suprasegment pharyngeal ganglion lies dorsal to the pharynx in the third and the suhpharyngeal ganglion lies ventral to the pharynx in the fourth segment. . 108. Each ganglion dition really the fusion of two. of the earthworm consists of the suprapharyngeal ganglion. two circumpharyngeal connectives. a deviation from the con- found in many annelids and arthropods where there are two is ganglia in each segment and the nerve cord double. -prosfomium -SvLpharyn^Zo/ Gang/fo/7 Posterior <Sejfmeti^a/ A/erve Bo</y IVa// Sefi/um ^et-^e Core/ Fig.

grooves . the Self-fertilization does not occur. 109. tions and the ventral surfaces Apertures of of their bodies in close contact from 5eminal receptacles Aperture of Vas deferens C/it el lam 5eminal droove Pore of oviduct Dorsal blood vessel Intestine 5eminal groove body wall Band of mucus secreted elite II am Fig. two worms enclosed in bands of mucus B. transverse section showing the seminal . Reproduction earthworm is hermaphroditic. The stimulus is modified in the ventral efferent neurons. this arrangement a stimulus applied to any part of cause responses to occur in a wavelike manner in both directions from the point of stimulation.PHYLUM ANNELIDA 209 Stimuli are received by sensory cells and are passed into the ventral nerve ganglia by the afferent nerves. A. cocoon. however. O. . In reproduction two animals come together with their anterior ends pointing in opposite direc- As has already been described. ganglia and sent to the responding organs by Nerve impulses then have the nature of a simple reflex except that the ventral ganglia are connected neurons which conduct stimuli from one to the other. By this means the worm can contract its entire body at one time. Located in the dorsal part of the by association Because of the body will nerve cord are three giant fibers which serve as the sole means of conducting an impulse directly from one end of the body to the other.— Reproduction in earthworm showing copulation and the cocoon. each egg being fertilized by a sperm from another individual.

G and D. 110. same as I but in cross section K. . Journal of Morphology. brane B. vesicles and vasa deferentia to the closed passage and move through it blastocoef t^eso blast cell Endoderm Coelomk cavities Hesodermic tissue Ectoderm Development of the earthworm. . . A. (After W^ilson. Embryology of the Earthworm.) . . blastula. . cross section of later stage. longitudinal section of a young worm after formation of the mouth and anus J. . gastrxila in archenteron G. 1889. . — . mesoderm layers on each side of them and the H. later stage showing cavities in the mesoderm cross section /.210 TEXTBOOK OF ZOOLOGY the anterior end to the clitellum. side view of gastrula showing invagination F. section of a blastula showing blastocoele and one of the mesoblast cells cell) of mesoderm layer. surrounded by a memFig. closed passage is With their bodies in close contact a formed between the genital openings of the two inSperms pass from the testes out through the seminal dividuals. section of gastrula along a line to show polar cells. showing stages in the beginnmg of (primary gastrulation E.

is set off. 111. tail regenerated from an anterior piece of a worm union of three pieces to make a long worm E. a certain cell. At this stage the animal constitutes a tube within a tube and from it the organs of the adult develop. A. . Soon after the segmentation cavity is formed. and C represent regenerated tissue. College Zooloc/y. move into the cleavage cavity. they separate by working themselves through the bands secreted by the clitella. As the mesoblast move into the cleavage cavity. the clitellum of each individual secretes a band which binds them together at these two points.PHYLUM ANNELIDA to the seminal receptacles of the mate. B. forming a cocoon and development take place. known as the mesohlast cell. . . in which fertiliza- Regeneration and grafting in earthworm. cells and the cells resulting from its divisions and will form the mesoderm.) . published by The Macmillan Company. grafting of two pieces to form a pieces grafted together to form a double bodied worm F. after Morgan. Cleavage in earthworms is of the unequal holoblastic type. This leaves each animal with a band which is gradually worked off toward the anterior end. The gastrula elongates and the archenteron opens at both ends to form the mouth and anus. and as it passes the openings of the seminal receptacles sperms which came from the reproductive mate are released. portion of a worm C. The stippled areas in A. After each has received sperms from the other. regenerated from posterior part of a worm B. anterior five segments Fig. (From Hegner. tion Both ends of the band close. between the ectoderm and endoderm develop segmental cavities which are the coeloms of the metameres. 211 stored. where they are In the meantime. The mesodermal cells which fill the space. — . As the band passes over the openings of the oviducts eggs are released into it. . tail regenerated from the posterior D. gastrulation occurs by invagination to form the endoderm and ectoderm. anterior and posterior new worm.

In this way numerous unusual forms worms have been produced. Each segment shows externally a variable number of annuli or rings. . but instead another tail will develop from the cut surface. The best known example of this group is Polygordius. the lost parts sults. midgut or crop. which has a long cylindrical segmented body with a pair of tentacles on the prostomium. Leeches are commonly parasitic and live by sucking blood from other animals. in front of the eighteenth segment. Class Hirudinea (hlr u din'e leech). have characteristically thirty-two segments and possess no external appendages. grafted to the cut surface to it Almost any part of an individual properly located) of another will fuse of earth- (if and grow. there is a muscular pharynx. — This class includes numerous small marine forms which resemble Chaetopoda in a number of ways. Two ciliated pits are present as a retenis tion of juvenile characters. common to the entire group. This produces an animal with two tails and no head. and the posterior one comes from the last seven. — These animals are com- monly known They are usually flattened dorsoventrally. setae They are very small and lack both and parapodia. If the cut is made posterior to segment eighteen a new anterior end will not regenerate on the tail half. Internally they are very similar to the earthworm. and death from starvation reWhen any part of the tail region is cut off. It is now believed that they have been de- rived from that group by changes usually involving the reduction or loss of certain structures. It has been demonstrated that when the anterior end is cut off.212 TEXTBOOK OF ZOOLOGY Regeneration Earthworms have been used extensively in regeneration experiments because they possess the ability to regenerate lost parts. of which Hirudo medicinalis is a good example. making the animal appear to possess more segments than are really present. the segments from one to five will be regenerated. possess both an anterior sucker and a posterior sucker. Numerous grafting experiments have also been performed on earthworms. The anterior sucker is formed from the prostomium and first two segments. a short esophagus. readily regenerate. first Annelida). Class Archiannelida (ar ki a nel'i da. The troehophore larva a. as the leeches. In a typical leech.

showing faint external segmentation. B. and. segmenting larva G. Development stages of Polygordius. 112. PHYLUM ANNELIDA intestine. It is then stored in The animal is capable of ing'esting three own weight in blood. gastrula . Textbook of General Zoology. which prevents times before its it the diverticulae of the crop. it Blood which sucked from an- other animal receives a ferment from the salivary glarids of the pharynx. since several months may elapse is all digested. advanced larval condition in which the trochophore larva is seen with head. after Whitman. The Fig. blastula C.) — . head kidney and preoral band of cilia E. . from coa^lating. adult worm.. published by The Macmillan Company. mouth. 213 is and ectodermal rectum. early trochophore larva D. . H. and tentacles. . . (From Newman. frequent feedings are not necessary. trochophore larva with posterior growth region F. A. . eyespots. optical section of trochophore larva showing apical plate with eyespot. an archiannelld.

ductus an. . I to XXVII. crop or stomach oe. and such other places as will afford protec- In this class. . po ?. dt. intestine ifflv. male aperture. and live in shells. (Prom Ward and Whipple. testes.. te. ej. and Fig. Sperms are work Development takes place in a cocoon produced by the clitellum. Class Gephyrea (je fi re'a. where fertilization occurs. bridge). and They are usually comparatively large possess a trochophore larva. . somites ejactulatorius ga and in. 113. — This class is a group of marine annelids which are nonsegmented. In Bonellia. and they apparently into the ovaries. . female aperture. pro. crevices. Each animal hermaphroditic. the female is the normal individual. placed on the skin of another leech. Fresh-water Biology.. Two nephridia are The nervous system is typical of the annelids. the representatives of the order Echiuroidea have tion. as modified from Castle.) . Glossiphonia fusca. . anus 2-70. have no appendages. po d. proboscis. showing annulation. a well-developed prostomium. Inc. esophagus ov. used in capturing prey and in locomotion. Bulletin of Museum of Comparative Zoology. a leech. segmentation. published by John Wiley & Sons. while the . .— 214 coelom is TEXTBOOK OP ZOOLOGY very much reduced due to the excessive is development of the mesodermal through present. ovary. annuli internal organs. it tissue.

(From Hegner. . 215 segmental organ of and lives in the Representatives of the order Sipunculoidea have no in the adult. This in- dicates without doubt that these animals are of great value.— PHYLUM ANNELIDA male has no proboscis. prostomium Nephrostomo Male genital pore Alimentary canal Alimentary canal Anus male Bonellia viridis. College Zoology. Importance of Annelids to Man and Other Animals Even though no casual observer would consider that annelids have any important relationship to other living organisms. known as castings. is ciliated. causing soil also Their continuous burrowing through the a necessary condition for plant growth. . birds. they have been found to be of great importance in a number of ways. Earthworms have sites also less desirable qualities. Darwin concluded from some forty years of observation that the earthworms in an acre of ground could bring to the surface in one year as many as eighteen tons of feces. Female (above) has bifurcated proboscis (below) is ciliated over the surface and much smaller. 114. secondary hosts for parasites of several animals. it porous. serious problem in some of the irrigation districts of the Southwest by burrowing through levees until they are too porous to hold water.) Fig. but with the presence of water they have become very abundant. published by The Macmillan Company. because in stirring the soil they cover up objects. the female. them makes to decay. pigs. Before irrigation was started they They have created a did not appear to be at all numerous. They serve as Most of the para- having the earthworm as a secondary host live as adults in and other animals which use the worms as food.

biting off that part of the worms protruding from the mud or sand. They are not at all averse to attacking human beings when the opportunity presents itself. The fish simply swim along. they cause no great injuries and are important only as pests and as secondary hosts for some and other forms parasites. however. (8) respiratory system (gills on parapodia of Nereis). (6) muscular system. Phylogenetic Advances of Annelida Segmentation. (4) closed circulatory system. Instead of dying the injured worms regenerate one or more new heads and go about their business. (5) excretory system of nephridia. and tentacles of Nereis). thus spreading certain diseases. (3) alimentary canal with defined parts. (7) concentrated mid-ventral nerve (1) cord connected to a dorsal pair of suprapharyngeal ganglia. . as parasites on turtles Various forms of leeches live of animal life in the water. (2) coelom. and (9) improved sense organs (eyes.216 TEXTBOOK OF ZOOLOGY in the bleeding of individuals The medicinal leech was once used as a treatment for various ailments. Marine annelids are important only as food for larger forms. palps. In many regions the burrowing forms along the tide levels literally form a good grazing ground for fishes.

This is indicated quite definitely by the fact that the larvae of echinoderms have a typical bilateral symmetry. — The central disc usually with five general features of the body include a arms or rays radiating from it. The modern echinoderms. at an interradius between arms in such a position that a line drawn through it and on through the radius of the opposite ray. as the group is commonly called. pedicellariae . It is located on the dorsal or aboral side. this entire group was included along with coelenterates in a group called Radiata. There will be a half ray and two complete rays in each half of a five-rayed animal divided in this way. sea urchins. The change which occurs seems to be an adaptation to a sedentary habit. : skin cov- ered with spines lack of segmentation . possess several distinctive characa rather teristics. CHAPTER XVI PHYLUM ECHINODERMATA The Echinodermata (e ki no dur'ma ta hedgehog skin) constitute backward phylum of animals which are thought to have undergone a certain amount of retrogression in structural features. they seem to have a lower level of organization than that possessed by some of their ancestors. — Some of these characteristics are as follows . It was later discovered that the coelenterates have a typical primitive radial symmetry while the Echinodermata have only a secondary radial symmetry which is derived from a bilateral condition. triploblastic radial sym- metry. The basis for this was the apparent similarity of radial symmetry. brittle stars. will divide the body into two similar halves. Classification In earlier classifications as in the case of Cuvier. water vascular or ambulacral system and tube feet circumoral nerve ring ajid radial nerves. animals as starfishes. This phylum is usually divided into five classes of modern forms including such common bers. There are some species which do not adhere to this pentamerous condition 217 .. The external opening into the water vascular system is called the madreporite. and sea Class Asteroidea. and a coelom. sea cucumlilies. That is. a calcareous skeleton composed of plates. subduing a primitive bilateral symmetry.

. Solaster. Fig.218 TEXTBOOK OF ZOOLOGY stricted and have up to forty rays. Asterias. and the radiating parts of the vascular. Oreaster and Echinaster are representative genera. organs called pedicellariae. around the mouth. orally along the ambulacral groove of each arm. Surrounding the spines on the skin are distributed some pincherlike calcareous ^^«SlK*n. 115. species. The rays are not usually sharply confrom the central disc. The anus and madreporic plate are located is The sys- mouth tems located on the oral or ventral side of the central disc. These vary considerably in different aborally. Astropecten. This groove is formed by a particular arrangement of Skeletal plates also support the aboral side of the central disc as well as the region skeletal plates. —A brittle star. An ambulacral groove is present in each ray. ambulacral. lie and nervous. Pisaster. oral view.

Class Ophiuroidea.




and serpent




small central disc with five long, slender rays which are clearly

from the disc. The rays are lacking in ambulacral The tube feet do not serve in locomotor functions but are grooves. Both the madreporite and mouth are located ventactile only.





the oral side of the central disc, five interradial sets of

toward the mouth in the formation of jaws which are operated by muscles in chewing food. The anus is not The brittle present. The viscera are confined to the central disc. serpent stars are quite active and live in the shore waters. stars and At low tide they may be found under rocks and debris, but they move about and feed during high tide. The ability of autotomy
skeletal plates project

Fig. 116.

—Oral view ofofa Generalstar belonging class Ophiuroidea. basket Biological Supply House.)

(By courtesy

(self -mutilation)


so well developed here that



become de-

tached by merely grasping them.
entire animals alive.

This makes it difficult to collect Ophioderma, Ophiura, Ophiothrix and Gorgono-

cephalus are


Atlantic and Gulf Coast genera.

Class Echinoidea.
tives of this


urchins and sand dollars are representalost the

group the members of which have

rays but

retain the pentamerous (five division) condition of the body.

The sea urchins are globular or hemispherical, while the sand dollars are disc-shaped. The skeleton or test is composed of five rows of closely fitting plates which are usually arranged into five pairs of inter-ambulacral rows. The position and condition of these rows

of plates can be



until the tips all

a starfish with its arms turned up over touch each other. The surface of the

skeleton bears processes which support movable spines.



be thrust out through perforations in the plates of the ambuThese rows correspond in position to the ambulacral lacral rows. grooves of the starfish. The plates of the inter-ambulacral rows








Fig. 117. Dried test of the sea urchin, Arhac'xa. A, shows arrangement of the (From plates on the aboral side; B, oral view showing mouth and perioral area. Wolcott, Animal Biology, published by McGraw-Hill Book Company, Inc.)

The mouth of this type of animal is located ventrally (orally), and it is guarded by five projecting skeletal processes called teeth. These converge over the aperture and are set in a skeletal case which is composed of many hard ossicles and contains the muscles for moving the teeth. The teeth are used in
are not perforated.



Aristotle's lantern,

removing algae from rocks for food. This arrangement constitutes and the esophagus leads internally from its
aboral part.

Aborally, each ambulacral area ends in a single ocular

plate at the periproct.


series of

genital plates interspace the

ocular plates of the periproct, and one of these genital plates has

become the madreporite.
center of the periproct.

The anus


a small pore located near the



Fig. 118.


portion of the test of the sea urchin has been removed to internal organs.

show the



view, a dissected sea urchin seems to be almost entirely

gonads, internally.

In each pentagonal section there


a mass of

the gonadal structure which

held in place by a band of tissue


from the aboral wall. Small from each gonad through a pore in its adjacent genital plate. The general arrangement internally includes a water vascular system composed of a stone canal leading from the madreporite to a circumoral or circumesophageal canal which encircles the mouth. A radial vessel extends from the circumoral canal through
as the genital rachis extending

genital ducts lead

the length of each ambulacral arch.

In each ambulacral plate are

found two pores for a tube foot which

a lateral branch of the

circumoral canal.


Each foot is connected with a bulblike ampulla. These tube feet along- with some mobile spmes constitute the locomotor system. Five interradial pouches or branchiae or "Polian vesicles" are in communication with the circumoral canal. The esophagus leads from the aboral part of Aristotle's lantern into the

dilated stomach.

The stomach extends almost around the


nal wall of the body.



the intestine leads in an opposite


in sea urchins finally

opens externally by the median

aboral anus.

In the sand dollar the intestine passes along the posteto the anal

ambulacral plate

of the disc-shaped body.


opening which is near the margin branch from the esophagus runs parit.

the stomach and finally joins

This tube





— Thyone,



sea cucumber.

ciliated internally



as the siphon.

Its function is con-

jectured to be either respiratory or a means of washing refuse from

The principal organs of respiration are the interradial pouches and the tube feet. The nervous system is composed
the intestine.
of a circumoral ring with radial cords extending into the ambulacral areas.

Strongylocentrotus, Arbacia,



and Echinarachnius (sand

dollar) are representative genera of the

Class Holothurioidea. The eehinoderms of this class have only an incomplete skeleton, the body is elongated, the mouth surrounded by tentacles is located at one end and the anus is at the other. These animals are called sea cucumbers because of their shape and



some remnant of the pentamerous condition in that there are five double rows of tube feet extending lengthwise on The five sides of the body of some forms, others have less or none.

expanded body of a holothurian is soft like a bladder partly filled with liquid and the body wall is very muscular. The madreporite The row^s of tube feet serve as structures of is located internally.




cr— -\V^/r?^esf/'^e


b/ooc/ yessef

bfood vessel

/nusc/e ba/7cf







Diagram showing the internal anatomy of a sea cucumber. (From Wolcott, Animal Biology, published by McGraw-Hill Book Company.)

locomotion and for clinging.
assist the tentacles in


of them, next to the mouth,

procuring food. Within the cloaca are the openings of two long tubular respiratory trees which receive water to assist in respiration. The tube feet, tentacles, cloaca, and other

organs serve in respiration. These respiratory trees function also as excretory organs. The madreporite draws water from the inside
of the body cavity.




The digestive system
esophagus which

most sea cucumbers consists of a short

supported by a skeletal structure at the point

body cavity. This structure serves as attachment for the tentacles and retractor muscles. Following the esophawhere

enters the


a short but rather inflated stomach which leads to the long, This tube

by a mesentery which is attached to the midventral line of the body wall. The intestine is thickened in its posterior portion to become the muscular cloaca which contains the openings of the two respiratory trees. In the coelom are fine longitudinal muscles that lie in the ambulacral areas. The gonad and genital duct are in ambulacral areas. They are found free at one side of the esophagus and stomach. This duct opens exteriorly by a pore beside the mouth. The food of the sea cucumbers is largely the organic material derived from mud which is ingested. This class of animals possesses a striking power of autotomy and subsequent regeneration. When they are irritated or disturbed, the muscles of the body cavity contract and produce internal pressure sufficient to cause either the body wall to split near the anus where the viscera are ejected or the viscera are forced out the mouth. Other animals, attempting to attack the sea cucumber, are rendered helpless by becoming entajigled in the visceral mass. The sea cucumbers can then regenerate the lost viscera in a short time. This power to eviscerate itself is a unique charactercoiled intestine.

partially supported


of the group.

Representative genera of this class include

Thyone, Holothuria, Cucumaria, Leptosynapta, Aphelodactyla and
Class Crinoidea.


of the sea lilies live attached

by long


At the free end many-branched arms which make up the calyx. The branches In most forms some lateral proof the anus are called pi^inules.
but a few are free.
of the stalk are located

jections, called cirri, are distributed at regular intervals

along the

The mouth


located in the uppermost center of the calyx



surrounded by the anus.

The anus


also to be

found on the

oral side of the calyx within the enclosure

made by

the arms.


the oral surfaces of the

arms are

amhidacral grooves which

serve to transport food to the mouth.

Modified tube feet are present,

but they serve more as tentacles than as

They lack ampullae


and are



slightly respiratory in function.

Gonads are

borne on the arms.

The skeleton


well developed in all crinoids

and for that reason many

of the ancient forms are preserved as fos-


-Distal portion

of a

stalked crinoid.

(Courtesy of General Biological

Supply House.)

in widely distributed limestone layers of the earth's surface.


nervous and circulatory structures parallel the ambulacral grooves





Neocomatella, Pentacrimis, Rhizocrinns,

Metacrinus and Antedon are representative genera.



Habitat and Behavior


starfish lives

along the shores and in the shore waters (to a

depth of over 125 feet) of our stony coasts of the Atlantic and

with scattered ones occurring in the Gulf of Mexico.



scattered individuals


be found on


or sandy shores, but
to pilings,

they are quite scarce.
old boats,

They are often found clinging
in the water.

and other objects
tide they


action of the tube

feet they are able to cling very tenaciously to almost



At low


be found under the rocks, out of the


where they are protected from the heat and drying. Due to a food relationship they are usually found in the same area with marine clams, oysters, and rock barnacles. During the day they are rather inactive, but at night they are much more active and respond to such stimuli as light, temperature, contact, and chemicals. It has been demonstrated experimentally that starfishes may form They ordinarily live and move about with the oral side habits. next to the substratum, and if turned over, will right themselves in the same way time after time. If the arms which are habitually
used for this are incapacitated, they will acquire the habit of using another combination of rays in this act.

External Anatomy
of a central disc and some (usually five) The mouth is located in the center of the under or oral surface while the upper or aboral surface is covered with spines of various lengths. On the arms these spines are arranged somewhat in rows. Between the spines the exposed skin

The body


radiating arms or rays.


extended into projections known as papula or dermal dranchiae. There are some small pincherlike structures, called pedicellariae, arranged around the bases of the spines, which serve to keep the surface of the exposed papulae clear of debris and foreign material. The pedicellariae are composed of two jaws or Mades and a basal There are large and small plate with which the jaws articulate. pedicellariae. In an eccentric position on the aboral side of the central disc is found the calcareous, sievelike madreporite. The portion of the central disc and two rays adjacent to the madreporite

constitute the hivium.

their adjacent por-

The other three arms and
compose the trivium.
rows of tube

tions of the central disc


the oral side sur-

rounding the mouth


a perioral membrane or peristome.
feet, radiates

An amhuthis along

lacral groove, containing


the oral side of each arm.


arm is called an eye. mouth and along the margins

reddish pigment spot in the end of The spines are longer and stronger around
of the ambulacral grooves than



Fig. 122. Pacific coast


published by The Macmillan Company.)

ochre starfish, Pisaster ocJiraceus, an abundant form along- the (XVi). (Johnson and Snook, Seashore Animals of the Pacific Coast,



fectly rigid.

and hard without being perdue to the presence of the calcareous skeletal plates throughout, which are bound together by connective tissue and muscular fibers. These plates are often called ossicles.

The body wall

relatively strong

This condition

They lie in a The skeleton


position in the aboral portions of the

of the ambulacral grooves consists of four

body wall. rows of

oblong ossicles in each arm.

These ossicles are ar-



ranged with the flat sides together, like cards in a filing ease. The two middle rows of ossicles are called amhulacral plates. Amhulacral pores, through which the tube feet project, are located between these The outer rows of plates, forming the margin of the groove, plates.

Dissection of the starfish, Asterias. The aboral wall has been reFig. 123. the trivium and a portion of the central disc. One ray of the biviuni has been turned to expose the oral surface and tube feet. The organs have been removed from one ray of the trivium to expose the skeleton. Am., ambulacra! groove; C.S., cardiac stomach; D.B., dermal branchiae; E, eyespot Cr, gonads; M, madreporite Os., ossicle; P, pelicellariae P.C., ploric caeca; Py., pyloric sac; ic, T.F., tube feet T.F.2, arrangement of tube feet in skeletal ray. rectal gland (From White, General Biology, published by The C. V. Mosby Company.)

moved from







are shorter and are




adamhulacral plates.


flat oral

surround the mouth. Within the body wall and extending into the arms is a large eoelom which is lined by a pcrito7ieum and filled with coelomic fluid. In The this cavity are located the organs of most of the systems. vertically from the digestive system is a modified tube extending mouth on the oral side to the minute arms at the aboral surface. From the mouth a short esophagus leads to the double-pouched stomach. The larger cardiac portion (or pouch) receives the esophagus and is separated aborally from the pyloric portion by a marked constriction. A large pair of branched glandular structures, known

Fig. 124. Diagram of a cross section through a ray of a starfish, avi, ampulla hca, db, dermal branchia amb, ambulacral ossicle coe, perivisceral coolom hepatic caeca musj muscle oSj ossicle pd, pedicellaria ph, perihemal space ra, Sp, septum m sp, spine rv, radial blood vessel radial canal radial nerve rn, radial radial blood vessel (rv) tf, tube feet; v, valve between tube foot and
; ; ; ;










(From White, General


as hepatic or pyloric caeca,

joins the pyloric

is located in each arm, and each pair pouch by a duct which seems to be a continuation of this pouch. These glands and possibly the pyloric pouch produce digestive enzymes in solution. The fluid secreted by the wall

of the cardiac portion probably does not contain enzymes.



rectum or intestine leads aborally from the pyloric pouch to the poreTwo brown, like anus at the exterior surface of the central disc. branched pouches arise from the rectum. These are known as rectal caeca or glands and probably have excretory function. In feeding, the starfish catches its bivalve prey in the five arms and humps



The tube

feet are attached to the shells, and,

by cooperais

tive activity,

an enormous pull

exerted on the valves of the

After the shell

open, the stomach of the starfish


mouth and

are digested in situ.


spread over the tissues of the prey which abundance of digestive fluid secreted over

the food causes the mollusk to be digested in






then taken into the stomach of the starfish. It is reported that between four and five dozen clams may be eaten by a single starIt has also been shown that a starfish may survive fish in a week. After feeding, the stomach is withdrawn after months of fasting. into the body cavity by five pairs of retractor muscles, one pair exis

tending from the pyloric portion to the ambulacral skeleton of each arm. The branched, treelike gonads fill the remaining space in each

arm and the external tween adjacent arms.


from them are located

in the crevice be-

Fig. 125. Longitudinal section through the central disc and one ray of a stara, anus am, ampulla car, cardiac stomach coe, perivisceral coelom ey, intestine eyespot hca, hepatic caeca m, mouth tnp. madreporic plate nr. i, ra, radial nerve ring oe, esopliagus os, ambulacral ossicle py, pyloric sac canal re, ring canal rca, lectal caeca sc, stone canal sp, spine tf, tube feet. (From White, General Biology.)
; ; ;
















composed of the madreporite, stone Tiedemann's bodies, Water is taken in through lateral canals, ampullae, and tube feet. the sievelike madreporite on the aboral side of the central disc and

The water-vascular system


canal, circumoral or ring canal, radial canals,


conducted by the S-shaped, calcareous stone canal (hydrophoric

canal) to the ring canal, which encircles the mouth.
of the water through the madreporite

The movement

and stone canal


by the action







the medial surface of

the ring canal are nine small Tiedemann's

(racemose) bodies, the

stone canal joining the ring canal where the tenth might be expected.

The five radial canals extend cells. one in the roof of the ambulacral groove of each ray. Numerous paired lateral canals arise along the length of each radial canal. Each ends shortly by connecting with its ampulla and tube
These bodies produce amoeboid



The ampulla is bulblike ajid is located above the roof of the It is connected through its ambulacral groove in the coelom. ambulacral pore with the contractile tube foot which hangs down
into the ambulacral groove.


distal or free

has a slightly inverted, suckerlike shape.

end of the foot The proximal pair of


Radial canal


Stone canal

Tiedemanri's body

Fig. 126.



of water-vascular system of the starfish.

Fig. 127.


"walking" on glass. Notice the extended tube of General Biology Supply House.)



ampullae in each arm of some starfish lack the tube feet and are sometimes erroneously called Polian vesicles. Alternate tube feet are farther from the radial canal than the others on each side. The ampullae and tube feet function effectively in locomotion, the ampullae contracting to force water into their respective tube feet




Fig. 128. Development and metamorphosis of the starfish. A, Dorsal view of early ciliated larva showing ciliated bands, and left and right coelomic pouches. B, Ventral view of bipinnaria larva showing the extension of the left and right coelomic pouches. C, Dorsal view of the same larva showing the left madreponc pore and water tube, and the fusion of the left and right coelomic pouches to form an anterior coelom. D, Dorsal view of an older larva showing the budding of the five water tubes from the left coelom. E, Left side view of a still older larva showing the water vascular system developing from the water tubes, and the rays of the adult starfish developing on the dorsal side. F, Brachiolaria larva in process of metamorphosis. The larva has settled on the preoral region which is greatly shortened. Q, Aboral view of a young starfish showing the developing spines. (Legend continued on opposite page.)



ampullae and tube feet are muscular. In large starfish the tube feet may be extended an inch or two. The sucker ends of these tube feet work like a vacuum cup and will adhere effectively to surfaces over which the animal is
extend them.

The walls

of both


When the pressure is released by the ampulla, the itself. tube foot contracts and draws the animal forward. When water is again forced into the tube, it releases its grip and is again extended. By alternation of the activity of tube feet in different parts of the body the animal is able to move itself from one place
to another.


entire water vascular system


a modified part

of the coelom.

thin-walled system of vessels running parallel to the water vascular is the circulatory system. It is enclosed in a perihemal



In addition to this the coelomic fluid, which occupies the all of the organs, serves as a circulatory medium This fluid in that it absorbs the digested food and distributes it. bears amoebocytes which are cells capable of picking up particles

coelom and bathes

of waste material

they pass through the

dermal branchiae, where These dermal branchiae are pouches of the coelomic wall which extend outward between the skeletal plates and have the additional function of When these pouches are completely extended, they respiration. nearly cover the exterior surface of the animal, and thus expose

and carrying them

to the



the exterior.

an enormous area

to the

water for respiration.

Excretion is carried out in part by the amoebocytes which have been produced by the Tiedemanu's bodies and have migrated to
the coelomic cavity.


rectal caeca serve in respiration to


There is a certain amount of diffusion of dissolved wastes through the dermal branchiae and the walls of the tube feet.
extent also.

The nervous system is radially arranged about the oral ring which From the oral encircles the mouth just orally to the ring canal. length of each arm and ends in the ring, a radial nerve extends the
pigmented eyespot.

These nerves


in the roof of the ambulacral

The aboral surface

supplied by a less conspicuous aboral




adoral ciliated band

m, mouth Ic, left coelomic pouch lateral arm I, mp, madreporic pore and water tube pad, posterodorsal arm band pr, preoral ciliated band re, right coelomic pouch sp, (Modified spines; st, stomach; w. five water tubes of the water vascular system. from Wilson and McBrlde. By permission, The Macmillan Co.)
; ;

b, brachiolar ad, anterodorsal arm anterior coelom es, esophagus dr, dorsal surface developing rays
; ;





point of




median dorsal arm


po, postoral ciliated





Branches of these

nerve which extends from an anal nerve ring.

nerves extend to the numerous nerve

cells distributed in the epi-

The pigment eyespots at the tips The pediof the arms are photosensitive and sensitive to touch. There is little cellaria and tube feet are also sensitive to touch. centralization except in the oral ring and radial cords, still there
dermis above the nerve cords.
is sufficient

centralization for the necessary coordination exercised

by the animal.
Reproduction and Life Cycle

The repro-


starfish is dioecious




the sexes are separate.

ductive systems of the two are similar and each consists of five paired gonads lying in the cavity of the rays beside the pyloric

They open to the exterior by pores in the angles between Mature eggs produced in ovaries of females and mature arms. spermatozoa discharged from testes of males are freed in the ocean


water where they unite in fertilization. Total, equal cleavage is the type of division which follows fertilization, and this finally gives rise to the many-celled, free-swimming, ciliated hlastula. The wall of this infolds to form a gastrula. Following this the rounded body becomes

somewhat elongated and lobed. Ciliated bands develop over its surface and it is known as hipinnaria. This larval stage has bilateral symmetry, and the larva swims about near the surface for weeks by the aid of its ciliated bands. A later modification of the hipinnaria in which there are several extended symmetrical processes,


as the hrachiolarian stage (Fig. 128).

Following this condi-

and symmetry superimposes the bilateral. The presence of the bilateral symmetry in these larval stages seems to indicate that the ancestors of echinoderms were likely animals with this type of

a metamorphosis during which


processes are formed,

the radial

Regeneration and Autotomy




applied to the power some animals have

to replace mutilated or lost parts.


starfish has this

or all of the

quite well developed with regard to





of the starfish

(Fig. 367.)

may be lost and the missing parts regenerated. An arm with a small portion of the central disc will

regenerate the missing parts under favorable conditions. A mutilated arm or one caught in the grip of some enemy may be cast off

by breaking


loose at the constricted point


joins the central

This ability of self -mutilation


as autotomy.


ing autotomy there

regeneration of a

Economic Relations

Compared with many other animals the echinoderms
unimportant economically.

are relatively

The sea cucumbers of several different as food by the Chinese and other oriental people. species are used The larger animals, some of them two feet long, are eviscerated, boiled, soaked in fresh water, dried or smoked and sold under the


of heche-de-mer or trepang.
It is

This dried product



and gelatinous.

quite expensive and

usually served as a

very palatable soup. The chief fisheries are found along the shores of China, the East Indies, Australia, and the Philippines; some, however, are taken in California, Hawaii, and the "West Indies.

Sea urchins of several kinds furnish a sort of caviar known as "sea eggs." The egg masses are taken from the sexually mature females and are eaten either raw or cooked. Each specimen contains




quantity of roe at the season just before Production of "sea eggs" has become quite an industry

in the Orient, Italy,

and the West


ticularly noted for their production of this

The Barbados are parcommodity.

Perhaps the

starfish is the

most important of the group, but its almost entirely of negative importance. It is one

and snails. The starfish around the oyster beds of the Atlantic, grows in enormous numbers attacks the oysters, and feeds on them, leaving only the empty shells. A single starfish may eat as many as two dozen oysters in a day. Oyster hunters formerly attempted to protect the oysters and clams by dragging "tangles" made of frayed rope over the beds, catching large numbers of starfish, breaking them in two, and dumping the scraps back into the water. The fallacy of this was realized when their power of regeneration was learned, so at present they are usually dropped into boiling water or thrown on the bank
of the worst enemies of clams, oysters,
to dry.

delicious food

Salted or smoked starfish roe by many people.


are considered a


brittle stars

cal indices

and crinoids have little value except as geologiand biological specimens. Their skeletal parts contribute

to the formation of limestone.


(By Elmer P. Cheatum, Southern Methodist University)

clams, oysters,

The phylum Mollusca includes such familiar animals as the snails, and cuttlefish. Even though they appear different



unsegmented, usually bilaterally
present which

symmetrical, and most of them produce a shell composed principally^ of

calcium carbonate.
in the


muscular foot



be modified for different functions.

In the snail


used for

clam for plowing through the substrate, and, in the Covering at least a portion of the body is a mantle or dermal fold, the outer surface of which secretes the shell in most species. Between the mantle and main body is a mantle cavity which is usually either provided with gills or modified into a primitive pulmonary sac for use in respiration. Jaws are present in the snails, slugs and cephalopods. Within the mouth cavity of many species is the radula, which is an organ composed of fine chitinous teeth arranged in rows and used in

nautilus or squid for seizing and holding prey.

rasping food.

Approximately 78,000 species of mollusks have been described,
hence they constitute one of the largest groups of animal

With very few exceptions they
ajid in the sea.

are sluggish animals and occupy a

diversity of habitats, occurring abundantly on land, in fresh water,

Although most of the species

live in

moist sur-

roundings, a few inhabit arid regions.
cuttlefish, are strictly



such as the



of the snails are herbivo-


and others feed as scavengers.

The oyster and other species

that are attached during adulthood feed on the floating organisms
in the sea.

From the standpoint of their ancestry, the veliger larva of various marine forms bears close resemblances to the trochophore larva Whether or not they are direct descendants of the of the annelids.



a matter for conjecture since some morpboJogists regard this similarity in larval forms as an example of adaptive parallelism Certainly morphological eviin a similar type of environment.

dence shows a close relationship.

(Detailed description based on Helix)

Habitat and Behavior
Snails occupy a variety of habitats. fresh water, salt water, brackish water,
live in the arid sections of

They occur abundantly in and thermal springs; they the country and occur abundantly in the


where certain arboreal forms are found. Some species belonging to the genera Caecilianella and Helix live underground, feeding
Apical or^an

[Mesodermal band

Apical or^an



Embryonic mMcle


ciliated ring









Rnal vesicle

A, Trochophore larva of Eupomatus (a polychaete annelid), side Fig. 129. (After Shearer.) B, Veliger larva of Patella (a marine snail) frontal secview. (After Patten. (Drawn by Joanne Moore.) tion.

embedded in moist and Helix aspersa, humus. Certain species, such as Helix hortensis excavate holes in rocks and live in them. Although most snails are
on roots of plants


other species live deeply

not tolerant to extremes of cold, Vitrina glacialis lives in the Alps above the timberline where the rocks are covered with snow most of the year; even some of our fresh-water snails in this country, such



Physa gyrina and Helisoma



frozen gradually, can live at least several weeks in solid cakes of
snails are


active either during a light rain or


ately following.

In heavily shaded woodlands where surface moisture prevails, snails are active during the day as well as at night. The same species of snail that exhibits both diurnal and nocturnal activity in the

woodland may show only nocturnal

activity in an open,

such as snails most land Lymnaiza Lymnoea buUmoides techelia palusbris Lymnaea stagnalis Ferrissia (Lymnaeidae) excenthca (Ancylidaej Physa bumerosa Physa anatina ( Fhysidoe) f HelisoiriQ Tropicorbis liebmanni Fig-. — Some common fresh water pulmonate snails. rocks.. . crannies in or under appear to be coPreceding prolonged move to protected places. Movements of incident with moisture rather than darkness. dense mats of humus. During this condition of torpidity the body of the snail may be well protected by one or several thin parchmentlike membranes called epiphragmas which are stretched across the shell aperture. periods of cold. Menetus dllatatus tnvolvis lentum (Planorbidac) t 130. the membranes are broken and the snail resumes its activities. When warm weather arrives. and there begin their period of hibernation. 238 TEXTBOOK OF ZOOLOGY exposed habitat. land snails may beneath dead logs.

and in this able sometimes to survive long periods of drouth. tapering spire (Lymnaea stagnalis) . these structures probably funcsnails embed themselves in moss and mud. or a long. and mollusks are parasitized by secretions produced various species of mollusks. by the latter. tion in retarding water loss. Rurnina roemen Fig. or Relisoma may be observed through the During periods of dry weather when ponds and creeks dry up.PHYLUM MOLLUSCA Water snails are active all four seasons. some of the aquatic genera. then return by sunrise the next ' ' ' ' mornmg. decoUata —Common vary considerably in snails. . but when the pond or stream is frozen over. Physa. broad. manner are During this condition epiphragms may be formed in certain species {Lymnaea palustris). 239 provided open water is available. External Anatomy Shell. and Polygyra roemeri have all been observ^ed to occupy as "home" a definite place and go out from this home on nocturnal foraging trips. humboldbiono chisosensis Polyqyra bulimulus dealbatus liquabilis terrestrial snails. the same as in land snails. such as Lymnaea and Helisoma may live two to four j^ears whereas some species of Helix may live to be six or eight The life span seems to years old. H. Helix aspersa. pomatia. ice. — The shell of the snail may be in the form of a low. or flattened spiral (Humholdtiana chisosensis and Polygyra roemeri). Parasitism and commensalism are both exemplified by certain species of snails. 131. the movements of Lymnaea. squamosum and the crustacean Gehia stellata. A few species echinoderms. At least a few species of land snails possess a homing instinct. Naturally their movements are slowed down in the winter due to cold. annelids. on the other hand. A commensalistic relationship exists between the rare mollusk Lepton The former feeds on of sponges.

.. is may have long peculiarly curved spines extending out from the main shell body that give In the sea and land slugs the shell either rudimentary. The thickness salts in the of the layers dependent on the richness of lime snails living in an acid environment.240 TEXTBOOK OF ZOOLOGY shells are some or shaped like house roofs (Patella that lives in the sea Ferrissia. of i is The shell which is largely composed of carbonate lime secreted by the mantle and usually consists of three layers. to the shell a grotesque appearance.. dextral. Zonitoides orboreus Euconulus chersinus trochulus Strobilops lobyrinthica (qostrocopta armifera texasiana Succinea Qvam Zuqfandina sincjleyana Fig. bog have thin transparent shells. whereas the same species inhabiting an area rich in lime salts have thicker. is a fresh-water form). The worm shell (Vermetus spiratus) so loosely coiled that it superficially resembles a worm. . aperture the shell is is on the the shell said to be sinistral. Polyqyra texasiana ^ Polyqyra dorfeuiHiana Helicina orb'iculata f?etinella tropica indentata paudiirata _ . Emis bedded within the latter may be pigments that give the occasional brilliant colors to certain species. but occa- sionally a reversal occurs which has been found to be inherited. Phiiornycu^ carolinensis — Common is If the shell is held with the aperture toward the observer and the left. such as those belonging to the genus Murex. Most species are normally dextral. Some shells. 132. or absent. internal. thus. Pupo'ides marq'inatus terrestrial snails. if on the right.

(sea slug) Fie:. such as Polygyra roemeri. different light true tentacles which are probably sensitive to contact and smell. Helisoma) have their eyes situated at the . — Limacina ausMb (Ptefopod) Vemetus {Worm spirgtus shelf) Muxextenuispina [Venus's comb) . Certain species. "eyes" which can possibly detect but are not sight organs. and a pair of stalked intensities. alholahris if 241 shells. Physa. . (Ship worn) —Marine mollusks. neck. Teredo navalis. The body of the snail consists of a head. and visceral hump. texasiana are capable of repairing broken shells damage is not too severe. and P. foot. The head of a land snail (Helix) has one pair of the Body. Aeolis Urosalpinx {Oyster drill) . Our common genera of water snails (Lymnaea.PHYLUM MOLLUSCA perhaps opaque P. 133.

highway of mucus over which the Respiratory aperturTz Velum i I Mouth A Genital aperture ' ^V^. respiratory. snail. fresh-water the gliding movements are scarcely The visceral perceptible. Located on the side of the head is the genital pore. Humboldtiana. the latter facili- hump. In some marine snails the surface of the foot is covered with cilia. is Just ventral to the mouth the opening of the pedal gland which snail usually glides. and land snails Lymnaea. A thick tating movement. The broad muscular foot deposits a covered with a mucus-secreting integument. B. land with bodies expanded. is protected by the shell which is lined with the mantle. and reproductive systems. — Fresh-water snail. which encloses the digestive.242 base of the tentacles. Tentack Respiratory aperture Jtaihed eye Edqe of wantle Foot ~1' Genital aperture Fig. excretory. 134. A. TEXTBOOK OF ZOOLOGY Just in front of and below the tentacles is is a mouth. circulatory. .

'Semma/ Receptacle '//nfesfine // Albumen Gland ^. 135. ry Qucr Fig. minute recurved teeth.Hearf /VasOeferens . —Internal anatomy On of Helix. —Just within the mouth of a snail The latter is known as the buccal mass.Orotesfis / . —Arrangement of / '' teeth in the radula of a snail. produced where the mantle joins the this mantle-collar is the respiratory aperture is and just beneath back of the latter the anal opening. the radula.Of/djcf 'Dart Sac ytlcfcovs Gland 0/ane/ / / VoQina / /Solvarv /Penis /Crop '' ^-Tentacle enilal Pore Anierior ''Tentacle -Pharynx 'Mouth ^\ ^Cerebral Ganglion •.bra I Canal.PHYLUM MOLLtJSCA collar is 243 foot. It is then rasped by mixed with saliva which flows lie from salivary glands that on each side of . 136. is a rounded organ composed of a ribbon of Fig. Shell removed. . the roof of the moved by conmouth is a horny jaw which pulls food into the the radula into fine particles and into the buccal cavity mouth cavity. supported and nective tissues and muscles. yry ^Sa/ifo. Internal Morphology Digestion. ^Hermaphroditic Duct _.

ably made possible through contraction of the mantle walls. primitive lung. forming the crop. thus decreasing or increasing the volume of air. have external feather-like gills. whereas the branchiate snails breathe by true In all probability pulmonate snails that inhabit the deep water of lakes use the chiates. blood of the snail consists of a plasma which usually color- but in Helisoma. Enzymes produced by this gland convert starches into glucose. in the case of Helix. Feces are discharged to the outside through the may anus. and. float the colorless corpuscles. and gasses may be carried from one part of the body to another. thus releas- be utilized. Most of the marine species are gill breathers. such as the sea slugs. it is not necessary for aquatic pul- monate snails to make periodic trips to the surface in order to re- new but when the water becomes sufficiently warm. the ferment is powerful enough to dissolve the cellulose of plant ing the protoplasm so that it cells. lies in the pericardial cavity. and some. The pulmonary sac their supply. thus giving it a red color. of aquatic pulmonates not only serves in the capacity of a gill or lung but also may serve as a hydrostatic organ. The heart. which widens.244 TEXTBOOK OF ZOOLOGY The masticated food is then passed into the esophagus the crop. secretions. From the crop. hemocyanin. The blood serves as a transporting medium whereby digested food. Respiration Land and most fresh-water pulmonate snails breathe by a fold of the richly vascularized mantle which has been modified into a gills. Blood is pumped from the . pulmonary sac the water is as a gill and breathe like the bran- When air cold. cutaneous respiration alone is inadequate and the snail must come to the surface to get additional oxj^gen. Here the food may be mixed with a brown liquid produced by the digestive gland which occupies most of the visceral hump. excretions. Circulation I is The less. In the plasma. and in Lymnaea and some species of Helix the blood has a bluish tinge due to the presence of a copper-containing pigment. which consists of an auricle and a ventricle. thus enabling snails Such movements are probto ascend to the surface by flotation. food enters the stomach and is passed on into the intestine where absorption takes place. hemoglobin is dissolved in the plasma.

pedal.PHYLUM MOLLUSCA ventricle through a 245 common aorta which divides into two branches. and visceral ganglia lie below the esophagus and are connected to the cerebral ganglia by commissures. Ova are passed from the ovotestis into the hermaphroditic duct. are hermaphroditic. The reproductive system Goniohasis. parallels the rectum and opens near the anus. of a unisexual snail is relatively simple but is exceedingly complex in the hermaphroditic species. first as an ovary and later as a testis others are protandrous. In bisexual occurs. situated three pairs of ganglionic swellings: above the esophagus. the pleural. By means of the latter organ sperm are transferred into the seminal receptacle of another snail. Veins carry the blood from the hemocoele to the mantle walls where it is purified and then passed through the pulmonary vein to the single auricle and on into the ventricle. supply nerves to the anterior regions of the body. and enter the vas deferens which terminates in a muscular penis. and from there into the oviduct . followed by the production of ova. Nervous System Encircling the esophagus is a ring of nerve tissue which includes the cerebral ganglia. as well as spermatozoa. are produced by the Some snails are protogynous. (hermaphroditic) snails cross-fertilization ordinarily ovo- The ova. Its ureter. The kidney is Reproduction and Life Cycle Most fresh-water and terrestrial sea slugs. one of which supplies the head and foot. Spermatozoa pass from the hermaphroditic duct into the sperm duct. such as Pleurocera. testis. nerves extend out to the visceral hump and basal parts The arrangement of ganglia and their connectives is of taxonomic importance. since male gametes are first formed. a thin-walled tube. since the ovotestis functions . From of the body. pulmonate The majority snails. as well as the of the marine shelled gastropods and our fresh-water branchiates. and the other carries blood The terminal branches of these arteries comto the visceral hump. municate with a hemocoele or series of sinuses. and Amnicola are unisexual. Excretory a yellow gland situated near the heart. them.



which terminates in a thick-walled muscular vagina. During this journey the ova are fertilized by sperm from the seminal receptacle and coated with albumin from the albumin glands. Both the penis and vagina have a common genital opening to the exterior.
Album in qland




^Hermaphroditic qland





Oort sac



Fig. 137.

(Modified, after Cooke, Genitalia of Helix aspersa; act of union. bridge Natural History. By permission of The Macmillan Company.)



of the fresh-water snails deposit eggs in clear gelatinous

masses on submerged objects, such as twigs and rocks. The land snails usually deposit their eggs singly or in clusters in well-protected places, such as in rotten wood or beds of humus. The eggs



3. '©


"Egg masses of common snails. A, Lymnaea (fresh-water; gelatinous mass); B, Heliosoma (fresh-water; gelatinous mass); C, Physa (fresh-water; gelatinous mass) D, Pleurocera (fresh-water branchiate; tough gelatinous mass) E, Polygyra texasiana (terrestrial; eggs in cluster) F, egg capsules of Busy con.
Fig. 138.
; ;

Qoniobasis comolenfis (Pleuroceridae)

Coivpeloma decisum (Vwiparldoiz)

Amnicola comolensis
Fig. 139.

CochliopQ texana (Amnicolidae)
fresh-water branchiate snails.

— Some common



are covered with thin shells which prevent undue water loss.

some marine


such as Busycon, eggs are deposited in disc-

shaped capsules which are spaced equally apart and held together

by a tough band. Some snails, such as the fresh-water Campeloma, have a brood pouch in which eggs are deposited and the young are born alive. The latter is ovoviviparous reproduction in contrast to
oviparous reproduction, as described above.

(Detailed description based on Lampsilis)

Habitat and Behavior Mussels or clams are usually found partly buried in the mud, sand or gravel of ponds, lakes, or streams. By means of the muscular foot which is protruded from between the two valves at the






Fig. 140.




— Some


fresh-water bivalves.

anterior end of the shell they plow their


slowly through the

stream or pond bed, feeding on the microscopic organisms in the


the posterior end of the shell are two openings


ventral siphon which pulls in food and water, and the dorsal siphon

through which wastes and deoxygenated water are eliminated.

sels are




the pelecypods.

may move

rapidly by suddenly contracting the valves, thus ejecting a jet of

Oysters are motile in their larval stages but in the adult

stage are attached to rocks and other objects.

Many marine mus-

attached to objects on the bottom or along the shore. Attachment is made possible by the dissolution of a part of the under valve and adherence of a portion of the body thus exposed.

The life span of clams may be relatively long. It has been estimated that Anodonta, one of our common genera of fresh-water
clams, attains its

maximum growth

in twelve to fourteen years.

External Features

Unlike the snail whose shell is of one piece, the clam shell composed of two parts called valves (hence, bivalves) which are

attached together at the dorsal surface by a hingelike ligament.



Liqamentous hinqe


Ventral siphon Dorsal siphc









Anterior protractor retractor muscle

Pallial \inz

shell features of

Fig. 141.


(A) and Internal (B)

Lampsilis anodontoides.

The oldest part of the shell is the umho which protuberance near the top of the valves and



usually a rounded frequently eroded

due to carbonic acid in the water. Extending out from the umbo on each valve in a concentric manner are the growth lines of the shell, evidenced as slight, medium, or heavy ridges.


shell is covered


by a horny, pigmented periostracum.


lying this

the prismatic layer composed of carbonate of lime.

The inner mother-of-pearl or nacreous layer consists of many thin, usually smooth plates, that in reflected light produce an iridescence





The valves are held together by two powerful transverse muscles, the anterior and posterior adductors. Upon cutting these muscles the shells gape open, exposing the underlying organs. The valves are lined with a mantle which secretes the shell. On the inner surPerkaniial cavity

Poit retractor
Post, adductor

Manble cut free




Ant retractor

Ex siphon











removed and

-Lampsilis anodontoides with the left mantle partially turned back to expose the underlying organs.

face of each shell


be seen the curved pallial line which extends

between the two adductor muscles and indicates the partial attachment of the mantle. Teeth which strengthen the closure of the shell may be present where the two valves come together. Between the two walls of the mantle is the mantle cavity which contains the leaflike gills, the foot, and visceral mass.

During the activity

of the clam a constant current of water


maintained in the mantle cavity. Food material is circulated forward to the mouth which lies between ciliated labial palps. Upon

entering the mouth, food
the saclike stomach.


passed through a short esophagus into

comes in contact with a digestive ferproduced by the digestive gland which is discharged into ment each side of the stomach through ducts. The crystalline style, a diverticulum of the intestine, and found only in mollusks, produces an enzyme mixed with the stomach content which undoubtedly The food, having been facilitates the digestion of carbohydrates. mostly digested and partly absorbed in the stomach, is passed on


ryjfcalline .style

1-* t^ucous qiands

Fig. 143.

— Cross

section through the style sac and intestine of Lampsilis anodontoides. (Modified after Nelson.)

into the intestine

which makes one or more loops in the foot, passes through the pericardium and terminates in the anus near the dorsal
Respiration is carried on through two pairs of vascularized gills which hang down into the mantle cavity on each side of the foot. Oxygenated water drawn in through the ventral siphon is passed through a rather complicated series of water tubes in the gills.




absorbed by the capillaries and carbon dioxide passed

into the water




discharged to the outside through the

dorsal siphon.



composed of a ventricle and two auricles lies The ventricle, a muscular organ, surrounds in the pericardium. drives blood forward through the anterior aorta the rectum and and backward through the posterior aorta. Both aortae give off arteries which ramify all parts of the body. Most of the returning blood is carried to the kidneys by means of the vena caval vein. Within the latter, nitrogenous wastes are removed, and the blood then flows to the gills through afferent hranchial veins; after puri-

The heart which


fication in the gills



efferent hranchial veins.

returned to the auricles by way of the The blood is colorless and contains several

types of white corpuscles.

Nervous System and Sense Organs
Situated on each side of the esophagus is a cerehropleural ganglion, the


two ganglia being connected by means of a cerebral comPerlcardial wall



Reno 'pericardial pore Excretory pore



odductor M.


water tubes
txhalQnt Siphon




Liver Ant.aorta -Stomach Cerebral commisjure
i j

Ant .adductor







\ \

Visceral Q.






Pedal q.

Cerebro pleural &.

Fig. 144.


organs of Lampsilis anodontoides.

missure which passes above the esophagus. Each ganglion gives off two nerve cords, one of which passes ventrally and posteriorly
to the pedal ganglion situated at the junction of the visceral




with the foot. The other nerve cord extends backward, terminating in a visceral ganglion which is usually located just ventral to The visceral as well as the pedal the posterior adductor muscle.
ganglia are united.

The sensory organs of the clam are primitive.
visceral ganglion

Covering each

a patch of sensory epithelium called the osphradium, the function of which may be to test the purity of the short distance water brought in through the respiratory system.


a statocyst which functions in equilibrium. It is composed of a small calcareous concretion, the In addition to statolith, which is surrounded by sensitive cells. the sensory organs named, there are many sensory cells distributed

back of each pedal ganglion


along the mantle edges and elsewhere which probably react to light

and touch.
Paired kidneys lie on each side of the body just below the periEach consists of a glandular portion which excretes cardium. waste, and a thin-walled bladder that is connected with an excretory pore through which wastes are discharged to the outside.

Reproduction and Life Cycle

The small bivalves belonging to the family Sphaeriidae (Sphaerium) are hermaphrodites, but in the larger ones the sexes are usually separate. The paired gonads are situated in the foot; the testis is usually whitish in color and the ovary reddish. A short duct leads from the gonad and opens just in front of the excretory pore. Sperm are passed to the outside through the dorsal siphon and
enter the female clam through the ventral siphon.

The ova, having

been discharged through the genital apertures, become lodged in various parts of the gills, depending upon the species. Within the Thus, the gills serve as brood pouches gills the eggs are fertilized. or marsupia and may become greatly distended due to the tremendous number (as many as three million) of developing embryos.

The small bivalve


which ranges in


from about 0.05


called a glochidium and has a single 0.5 millimeter in diameter, adductor muscle for closing the valves which may or may not be hooked. Extending out from the center of the larva is a long secre-

tory thread, the hyssus.
to the outside

In most clams the glochidia are discharged through the dorsal siphon. They fall to the floor of


lie with their jaws agape, or snap jaws on any object. If the soft filament of a fish's gill or a of the fish comes in contact with the glochidium, it will close

the river, pond, or lake, and




group up around and eventually cover the parasites. Thus a cyst is produced about the glochidium and within this structure the
larval clam undergoes metamorphosis.
its host,

It shortly

breaks loose from

drops to the stream or pond bed, and leads an independent
in a river system can be

The rapid dissemination of mussels accounted for by the movements of their

Economic Relations of the Phylum
Mollusks have been used as food by

man from

the beginning of

Oysters, clams, scallops, snails, and the


of cuttleIt

are found in the


of peoples all over the world.


been estimated that the oyster industry along the Atlantic Seaboard approximates 40,000,000 dollars annually. Along the Texas
coast alone. Federal statistics

show that 51,719 barrels

of oysters

were sold in 1932. Buttons are made from the shells of the large heavy river clams and along the Ohio, Missouri, and Mississippi rivers the button industry amounted to 5,000,000 dollars in 1931.

Within some of the clams are found pearls which are formed by some irritating particle, such as a parasite or sand grain that becomes lodged between the mantle and the shell. Iridescent protective layers of mother-of-pearl are deposited around the foreign parThe Japanese have been successticle, thus producing the pearl. ful in artificially stimulating pearl production by planting small objects, such as pieces of mother-of-pearl, between the mantle and
shell of pearl-oysters.

Pulverized clam shells are also being used as a calcium supple-


to chicken feed.

Shells have also been used as a
of the eastern coast of






North America
brightly colored

consisted of strings of cylindrical beads

made from



Shells have always been

are used for orna-



shells are

used in road construction.

Some mollusks

are injurious to





might be mentioned the marine snail, Urosalpinx cinerea, which drills into and feeds on oysters and other pelecypods; the common shipworm, Teredo navalis, attacks the wood of ships and pilings, making extensive excavations. Certain species of snails serve as the intermediate host of parasitic flatworms or flukes. The liver fluke



(Fasciola hepatica) whose intermediate host

the small fresh-water


huUnioides, causes the disease, liver rot in livestock,

particularly in the sheep of the Southwest.

Since shells are easily fossilized they serve as excellent guides to
the geologists in determining the type of rock formation



age of the strata,

Classification of this



based on the nature of the foot,

and respiratory organs; shape and structure of the shell; arrangement and structure of the nervous and reproductive systems.

Includes the Chitons, which are found abundantly on rocks between

marks along the Atlantic and Pacific Coasts.

This class apits

pears to be the most primitive in the phylum, and







if present,

symmetrical body; tentaculess head, eyes absent;
consists of eight overlapping plates.

foot but other species

Most species have a flattened are slender and wormlike Ischnochiton con-



Includes the bivalve moUusks, such as the oysters, clams, scallops,



More than ten thousand

species have been described,

of which approximately four-fifths live in the ocean.
the class into orders

Division of

based on giU characters.




Marine species;

gills consist

of short, flattened leaflets; dis-

tribution along the Atlantic and Pacific Coasts.

Marine species; gills composed of long filaments which hang down into the mantle cavity. The edible scallops and the sea
mussel, Mytilus, exemplify this order.




Fresh-water and marine species; with two platelike gills which hang down into the mantle cavity on each side of the



Unionidae Fresh-water clams or mussels; shell large or relatively
large; valves equal and


anterior to center.




Fresh-water species.

Shell small;

umbo median

or pos-

terior to middle of shell.




Marine species;
trate this order.


plaited into vertical folds;



quently inequivalve.

The oyster (Ostrea) and Pecten


Dental ium

Loliqo brevipennis

Polypus bimaculatus (Octopus


Class Amphineura, Repre-sentatives of three classes of mollusks. Fig. 146. Ischnochiton; Class Scaphopoda, Dentalium ; Class Cephalopoda, Loligo brevipennis (squid) and Polypus bimaculatus (octopus).
Class III. Gastropoda

Includes the snails and


species have been discovered

Approximately fifty-five thousand and described. Shell, if present, uni-



Mostly marine, but fresh-water and land forms are represented.




implies, the gills are situated in the

mantle cavity anterior to the heart.

This order embraces

such animals as the limpets, abalones, and periwinkles,
as Goniohasis,

of which live in the sea; also a few fresh- water genera, such

Campeloma and Pleurococera; Helicina


a terrestrial southern species which

frequently arboreal

in habit, comes under this order.



Strictly marine.


present, are situated jjosterior

to the heart; shell, if present, small.

Includes the sea slugs.

In the sea butterflies (pteropods), the foot may be modified Some of the into two fins which are used in swimming. heavier types have broad cephalic discs, adapted for burrowing in the sand. Many are found in coral beds and in seaweeds, their vivid colors harmonizing with the background.


Mostly terrestrial and fresh-water snails. Gills are absent, the mantle cavity serves as a pulmonary sac; shell usually present, sometimes rudimentary or absent. Suborder 1. Basommatophora Fresh-water species; eyes located at base of tentacles;








naeidae, Physidae, Planorbidae and Ancylidae.




and slugs; stalked
retractile tentacles;

shell in


and one pair of
or absent. Class IV.

form of

elevated or depressed spire, rudimentary and concealed,


Scaphopoda Mantle edges grown together along ventral side forming tube, with a shell of same shape and open at both ends. Commonly
as tooth shells.


Approximately 300

kno-u-n living





The most highly organized of the mollusks. A definitely formed head is present which bears a pair of eyes that superficially resemble the eyes of vertebrates. The foot is modified into arms or tentacles. They are carnivorous animals and many of them are used as food by man. (Nautilus, Loligo, Polypus.)




The chambered nautilus (Nautilus) is a representative of this order. The animal inhabits the last chamber of a flattened spiral calcareous shell. As the name Tetrabranchiata
implies there are four gills; also four primitive kidneys and



Siph uncle

1 __ - ' ^ Jepta

Fig. 147.


view of internal structure of Nautilus.




Fig. 148.

of the cephalopods.



four auricles;
ink sac

This suborder



peak of development in the Silurian and Devonian periods and is one of the most clear-cut examples of evolutionary
development in the invertebrates.

During the Ordovician period the cephalopods constituted one of the chief groups of marine animals. Even though at that time cephalopods with coiled shells existed, the predominant ones were the orthocones (those with straight This latter group in all probability gave rise to the entire conical shells).
series of

coiled shells, culminating in Nautilus.



nautiloids a series of


termed septa, extend the

full length of the shell.

The point of union

with the septa and sides of the shell may appear as a straight, curved, angulate This line is called the suture and in fossil shells whose or highly complex line.
outer shell coating


lost, it

stands out rather conspicuously.

The suture


used as a taxonomic character for the group.




Octopods and squids are representative types.
or absent;

Shell internal




and two primitive kidneys; ink sac pressurrounded by 8 to 10 tentacles which are

furnished with suckers.

This order includes the largest of

moUusks, the giant squid (Architeuthis princepsj which
attain a total length, including arms, of over fifty feet.



The squids and octopods are noted for their ability to change color by the rapid contraction or expansion of chromatophores Their juovements are rapid and are produced in their skin.
by expelling water from the mantle cavity through the muscular siphon with such force that the animal

jerked backof


In the







facilitate locomotion.

Loligo hreviyennis


the small squid found along the Gulf coast.
it is


taken out of the water

usually a mottled red or tan.

mass and mantle cavity are enclosed by a thick muscular mantle. Beneath the skin along the back is a primitive endoskeleton in the form of a feather-shaped shell. The squid is predaWithin the tory, feeding on almost any animal it can capture. The large jaws moved by powerful muscles. pharynx are two terminates in a pharynx connects with an esophagus which in turn muscular stomach. Digestive juices from the liver and pancreas

and after the food is partially dipassed into a thin-walled cecum where digestion is completed and absorption takes place. Wastes are discharged through the anus which opens near the base of the siphonal fold. The blood
are emptied into the stomach,
it is






Hectocotylhed arm.





Anb. aorta

5yitemic heart—




post cava

Spermabophoric sac






-Cub edge of body wall

Fig. 149.


of squid to sliow internal anatomy.

system, which


is composed of arteries, veins, and two Blood is oxygenated in two feathery gills which project into the mantle cavity. The two light-colored triangular kidneys are situated anterior to the branchial hearts and discharge their contents through small papillae, one located on each side of the intestine. In squids the sexes are separate. The male reproductive system is composed of a testis, vas deferens, spermatophoric sac, and penis the female system consists of an ovary, oviduct, ovidueal gland, and nidamental gland.

branchial hearts.




Lens Ciliary M.



Optic (ganglion

Fig. 150.- -Longitudinal section through eye of squid. (Redrawn and modified after Borradaile and Potts by permission of The Macmillan Co.)

The nervous system of cephalopods shows a high degree of spewhen compared with the nervous system of other mollusks. The "brain" is composed of a close association of ganglia around the esophagus and is protected by a capsule of tough tissue resembling cartilage. Nerves radiate out from the central nerve mass to the various parts of the body some of the nerves terminate in large ganglia, such as the stellate ganglia in the mantle. The eyes of the squid are supported by pieces of "cartilage" and are relatively complicated. Statocysts, which are similar but more

complicated than those described for the clam, are situated near
the brain mass.
in function

Ciliated pits which are supposed to be olfactory open in the form of a slit just back of each eye.




Arthropoda (ar throp'O da, joint foot) is the name of the largest known group of animals. As the name implies, all representatives of the phylum have paired, jointed appendages and a definite tendency toward specialization of them. Their bodies are triploblastic, segmented, bilateral, and covered by a chitinous exoskeleThe coelom is modified by a marked reduction as a result of ton. The segmentation or metamerism of specialized vascular spaces. the body is expressed in a high degree in this phylum and there The segments is a definite relation of appendages to segments.
have undergone greater specialization and greater regional differentiation than was the case in annelids. In forms where there is little or no differentiation of segments, the condition is referred to as homonomous, while a highly differentiated condition of segments This as found in most arthropods is spoken of as heteronomous. distinct head, thorax, and abdomen. The appendgroup has fairly ages on various segments are typically homologous with each other.


are modified as sense organs, others as


parts, others

for walking,

swimming, and reproduction. The skeleton is entirely exoskeletal, composed of chitin, and fits exactly the shape and contour of the body. Since it is fairly unyielding to growth, it becomes necessary for the arthropod to shed the skeleton periodically during its growing periods. This molting
or ecdysis, as
sions of this
it is

called, is quite characteristic of


of the divi-


The circulatory system is of the ojyen type, since there are large sinuses or spaces surrounding most of the organs instead of a continuous circuit of blood vessels. The nervous system is of a modified
ladder type with a ventrally located cord. The digestive system shows specialization in that it is divided into distinct regions as an
adaptation to special types of food which require mastication.

This phylum is divided into two sections and at least five classes some authors recognize as many as eight. The sections are deter-

mined according

to the


of respiration.



Section I. Branchiata (brankia'ta, forms for the most part.

gill-breathing, aquatic

Crustacea, craj^fish, crab,


bug, barnacle, water



o m
oi a>

> u









P O ?>> > m




centipedes and millepedes (thousand legs) having one or two pairs of appendages on each segment. centipedes. grasshoppers. as subclasses. Sacculma (Fig. Order Cirripedia. crickets. Order Isopoda. the wormlike arthropod. Tracheata (tra ke a'ta. book lungs or book gills. and shrimps. Strepsiptera. Division B. springtails. cyclops. Order Amphipoda. all with three pairs of thoracic appendages and most of them with wings. pill bugs and sow bugs. Onychophora. Order Diplopoda. flies. Coleoptera. Emhiidina. Division A. sand fleas and beach fleas. Copepoda. etc. beetles. This section is divided into three divisions depending on the primitiveness of the characteristics. locusts. Isoptera. Class IV. such as nephridia. (Texas. Ostracoda. fish louse (Argulus). More highly specialized forms with one Class II. Order Chilopoda. Order Odonata. embicls.) Orthoptera." Dermaptera. stylopids (parasites in insects). Prototracheata. 404). Order Decapoda. goose barnacle (Lepas). lobsters. sow bugs.) pill Subclass Malacostraca. Insecta. rough) both terrestrial and aquatic arthropods which breathe by tracheae. Section II. millepedes. crayfish. sand fleas. dragonflies and damsel Order Plecoptera.PHYLUM ARTHROPODA — CLASS CRUSTACEA 265 Order Copepoda. roaches. and crabs. weevils and beetles. Thysanoptera. Order Thysanura. bugs. Antennata. California.. thrips. lobsters. Order Ephemerida. silver moth. Myriapoda. crabs. Order Collemhola. stone Order Order Order Order Order Order Order flies. thereby dispensing with Entomostraca. and Cirripedia craj^fish. pair of antennae. The primitive form with some arthropod characteristics and certain annelid features. mayflies. (Some authors prefer to rank Branchiopoda. . rock barnacle (Balanus). Class III. termites or "white ants. bees. Florida. Peripatus. earwigs.

it serves ideally as a representative more detailed study. scorpion butterflies and moths. Order Mecoptera. body lice ("cooties"). scale insects. one Texas species. king crab. Arachnida. bird Order Anoplura. Order Xiphosura. true bugs. A without antennae but Avith tracheae. CRAYFISH OF CLASS CRUSTACEA Since this animal represents a relatively simple type of arthropod and is so generally well known. Order Araneida. ant lions. Order Hymenoptera. lions. mosquitoes. ticks and mites. book Order Mallophaga. The number of species described under the phylum is approximately one-half million. caddis Order Lepidopiera. Order Trichoptera. vinegarroon and tarantula. . Order Pseudoscorpionida. aphis lice. spiderlike). scorpions. book lungs or book gills. Order Dipt era. Order Scorpionida. TEXTBOOK OF ZOOLOGY lice. book scorpion. mite. Order Corrodentia. plant Order Neuroptcra. group and four pairs of thoracic appendages. Class V. lice. and there are large numbers still undescribed and early in the chapter in order that the student nitude of unnamed. king crab or horseshoe crab. spiders. flies. true flies.266 . crab Order Hemiptera. spider. Arachnoidea (ar ak noi'de a. Division C. as squash bug. cicadas. Order Phalangida. scorpion. This summary its size of the classification of the phylum has been placed may realize the magand the great variety of animals included. The former is distributed east of the Rocky Mountains and species for a the latter on the Pacific slope. Order Acarina. The genera Canibarus and Potamdhius or Astacus are commonly found in the streams of North America. wasps. bees. Order Siphonaptera. Order Pedipalpi. louse. etc. ants. flies. fleas. daddy longlegs or harvestmen. Order Romoptera. Order Palpigradi.

It is possible for them to walk about on the bottom of the stream or pond. swamp crayfish. They are much more active at night than during the day. small fish. as a result of the strong downward stroke of the tail. crawclads.) swimming habits are rather peculiar in that they dart backward through the water.PHYLUM ARTHROPODA — CLASS CRUSTACEA 267 Habitat and Behavior For the most part crayfishes (crawfishes. touch. Crayfishes may at times desert their aquatic habitat and go foraging out over build their swampy burrows down land. In some localities certain species to the subterranean out in the fields and become important pests. fresh-water lobsters) are inhabitants of fresh-water streams and ponds where there calcium carbonate in solution for purposes of skeleton formation. the the Southern States. Their species carry air tunnels vertically Fig. is sufficient Some from the original horizontal burrow to the surface of the earth and deposit mud around the opening of a tunnel. by waiting in hiding and suddenly seizing . chemoreception are important senses in this animal. or they may be in hiding under some stone or log. a very common species in the (Courtesy of Southern Biological Supply Co. such as tadpoles. or they may be in the mouth of a burrow beneath the water's edge. water table right Sight. and The crayfish captures other animals. These animals may be found moving about on the bottom.— Cavibarus ol: clarkii. moving the body in almost any direction. One stroke of the tail will carry the animal a yard and The daytime is this is commonly sufficient to avoid the enemy. and aquatic insects. usually spent in hiding under objects or in the mouth of the burrow. swamps 152.

to the thorax.268 them. due to its protective which matches the background. The portion anterior to the cervical groove is the head or cephalic portion. the pleuron. The mouth is located on the ventral side of the head portion and not at the tip of the rostrum where most people look for it. is The portion posterior "tail" by fishermen. lateral plate. salamanders. which has no appendages but is often called the seventh abdominal segment. while the portion posterior to the grooves is the thorax. fish (such as bass and gars). . its chitinous skeletal covering. and raccoons in particular. abdomen. turtles. and by their use as food for man. Many have been exterminated by the drainage of swamps. color TEXTBOOK OF ZOOLOGY The crayfish is quite well protected. External Structure The chitin-covered body is divided into cephalothorax. and appendages. The cephalothorax is a compound division of the body including the thirteen most anterior segments and is divisible The boundary between these is marked by into head and thorax. Their ventral edges are free. like covering whose lateral edges are free. the posterior end of this. overhanging and the slender ventral sternum in the form of a narrow bar extending from side to side. The anus is found on the ventral side of this part. The skeletal part of the abdominal seg- ment consists of: the dorsally arched tergum. really the which is frequently called aldomen. frogs. On the ventral side of the thorax between the twelfth and thirteenth segments (about the level of the fourth walking leg) of the female is a cuplike pouch called the annulus or seminal receptacle. The lateral portions of the carapace are known as hranchial areas or hrancliiostegites. alligators. gills. drawn out to almost a point. and the tail proper is at is The abdomen divided into six typical segments and the terminal telson. they are captured by water snakes. A thin arthropodial membrane extends between successive sterna and allows for movement of the segments upon one another. The shellthe oblique cervical groove on each side of the region. In spite of this. receipt It serves in reproduction for the and storage of spermatozoa. a thin. herons. and its pinchers. is known as the carapace. and they cover the anterior end of the cephalothorax is The and this portion is called the rostrum.

. maxilliped Mp. Protopodite Epipodite- Endopodite Protopodite k .3 Eindopodite.. ^. second maxilla. 2. (From Newman. 2. 2. third maxilhped Mx. M. Antennule fir.PHYLUM ARTHROPODA — CLASS .2 Exopodite Endopodite Protopodite.3 CRUSTACEA 269 Protopodite A2 Endopodite A. Mp. ..ChitinouB thrrada Epipodite Examples of cephalic and thoracic appendages of the crayfish. Mx. Outlines of General Zoology. 3.. . 1.2 •Epipodite Mp... Mp.. ventral Fig. A. fourth walking leg view. antenna L. second maxilliped Mp. : — .) . 153. first maxilla. i.Endopodite Mx.l Endopodite Exopodite Exopodite Endopodite Mx.:.l Exopodite Protopodite. 1.l Epipodite Mv. mandible A. ." Endopodite External ope'ning of nephtidium. M. published by The Macmillan Company. after Kerr.

ing the coxopodiie (first segment of protopodite) gill a sheetlike struc- ture which supports a anterior walking legs and some chitinous threads. There are nineteen such pairs. platelike exopodite and endopodite. the exopodite or lateral branch and the The first two endopod'ite or medial branch each have many joints. but in the male the protopodite for at- and endopodite are fused and extended to serve as an organ The posterior pair of swimmerets. Each is is composed Join- of the two joints of the protopodite and five of the endopodite. | head are attached five pairs of appendages. tached to the sixth abdominal segment. Just posterior to the mouth and immediately in front of the first maxilliped are two pairs of maxillae. transfer of spermatozoa. oval. The posterior five thoracic appendages are the walking legs or pereiopods. veloped on the same plan from the typical biramous (two branched) appendage. The three possess pinchers or chela which are formed by and defense. exopodite of the second are fused to form a bladelike hailer or scaphognathite which fits over the gills and by its movement helps circulate the water for respiration. The walking maxillipeds. but the protopodite is broad and foliate. legs are used in locomotion. joining the body and the hasipodite. The first maxilla is reduced The jawlike manto a leaflike protopodite and small endopodite. quite typical of the primitive are much reduced in the female. except the first Each has bear gills. Its endopodite is slender. The epipodite and the first. the second of which overlies They are both leaflike and modified. are broadened into fanlike They are known as uropods and have structures for swimming.270 TEXTBOOK OF ZOOLOGY cal The appendages are paired. with one pair attached to each typiThey are all desegment. dible at each side of the mouth is composed of hard protopodite To the segments of the . ofi'ense. three anterior segments of the thorax bear three pairs of biramous The parts are quite typical in most respects. The five anterior pairs of abdominal appendages are form except for the modification of the first two in connection with reproduction. This group is known as smmmerets or pleopods and all have the fundamental parts consisting of a basal protopodite composed of coxopodiie. an epipodite joining the basipodite and all These appendages are used in getting food to the mouth. The the terminal segment being set on the side of the second segment. These are uniramous due to the complete reduction of the exopodite.





with teeth ajid a fingerlike endopoclite, which is tucked under the anterior edge of the former. This appendage is used for chewing. In front of these are the antennae which are biramous and are sometimes called "feelers."


consist of the protopodite of


a long many-jointed, filamentous endopodite and a


short, fan-shaped exopodite.

which are biramous and
are similar in these.


Anterior to these are the antenmiles The exopodite and endopodite

The principle of homology is excellently illustrated by the appendages of the crayfish. In general, homologous structures are those which have similar structure and similar origin but may have
similar or different functions.

By way

of contrast, analogous struc-

tures are those which,

when compared, show

different structure


origin but similar function.

During early development appendages of the crayfish is similar to all others. Some become modiOther illustrations of homologous structures fied with development. are the human arm and the bird 's wing. In organisms like crayfish
where the appendages of successive segments are homologous to each Homologous other, the condition is spoken of as serial homology. stiiTctures are found in many animal groups and are used in establishing relationships. It ha.s been suggested that the parapodia of Nereis
represent possible forerunners of crustacean legs. They are both typically biramous and both take about the same position on the body, as well as having a similar segmental distribution. There is also considerable similarity in their structure.

each of the

Internal Structure

Beneath the

shell-like, chitinous

exoskeleton there


a very rep-

resentative set of systems.

is retained in a degree in the circulatory system. Earlier in the chapter it was pointed out that the coelom is modified as a provision for increased blood sinuses which have occupied much of the space.

most higher animals the segmenthe muscular system, nervous system, and to



the branchial areas of the carapace may be found the paired, feathery gills held in the gill cavity or There are three types of gills present here: branchial chamber.

Respiratory System.^


pleurohranchiae, attached to the sides of the thorax; podohranchiae,

from the epipodites of the thoracic appendages; and arthro-

branchiae, which arise



from the coxopodites of the thoracic appendSeveral of the segments have lost the pleurobranchiae. The scaphognathite moves in such a way over the external surface of
the gills as to

move the water in an anterior direction. The water brought under the free edge of the branchiostegite or branchial area of the carapace and moved forward to be discharged by an anterior aperture. An almost constant stream of water is pumped over the gills to facilitate the exchange of oxygen and carbon dioxide between the blood in the capillaries of the gills and the surrounding water. The aerated blood is then carried to all of the
tissues of the body.
Pericardial sinus

Carapace removed

Heart Ostium



Efferent vessel




Ventral thoracic

Nerve cord


Wntral sinus




of cross section throug-h the posterior thoracic crayfish. Arrows indicate flow of blood.

region of a

The digestive system is in the form of a modified canal and is composed of mouth, esophagus, stomach, and intestine. The mouth
opens between the mandibles on the ventral side of the third segment. From this the short, tubular esophagus leads dorsally and joins the ventral side of the stomach almost directly above the

mouth. This larger anterior portion of the stomach is the cardiac chamber. Within its wall are a number of hard chitinous bars, known as ossicles, which bear teeth capable of mastication of food when moved over each other by the muscular activity of the wall. This grinding apparatus is known as the gastric mill. Between the cardiac chamber and the posterior or pyloric chamber is an arrange-





of bristles which serve as a strainer that allows only properly

masticated food to pass through. ably smaller and curves

The pyloric chamber




continue posteriorly as the

tubular intestine which extends almost directly posteriorly through
the center of the abdomen to the anus in the last segment.
digestive glands



lead into the pyloric chamber

through hepatic ducts.


secretion of these glands contains diges-

consists of a heart, the

The vascular system

pumping organ;


arteries, definite vessels; the sinuses,


series of blood spaces;

Nood which

6 7 g

It consists of the fluid

and plasma containing



Lateral view of a dissection of the crayfish to show many of the Fig. 155. S, internal organs. 2, circumesophageal connective 1, supraesopliageal ganglion ophthalmic artery J/, stomach, cardiac portion 5, lateral teeth 6, median teeth antennary artery; 8, testis; Oj, hepatic artery; 10, ostiuin 11, heart; 12, dorsal 7, abdominal ganglion; 28, ventral abdominal artery; 29, nerve cord; SO, rectum; gland; 17, esophagus; 18, mouth; 19, subesopliageal ganglion; SO, stomach, pyloric portion; 21, opening of hepatic duct; 23, digestive gland; 23, ventral thoracic artery; 2i, sternal artery; 25, opening of vas deferens; 26, thoracic ganglion; 27. abdominal arterj' IS, vas deferens; li, intestine: 15, renal opening; 16, green (Modified from Turtox Key Card of Crayfish. Courtesy General Bio31, anu9. logical Supply House.)








white corpuscles but without red ones.
sorbs oxygen

The hemocyanin which abFresh blood

dissolved in the plasma.

almost clear




takes a blue color after standing in the air for a

The heart is somewhat flattened and angular in outline, and has a muscular wall which is perforated with three pairs of slitlike ostia. When the muscular wall of the heart is relaxed, the slits open,
short time.



and blood is drawn in from the surrounding pericardial sinus in which the aerated blood accumulates. When the heart contracts, blood is forced into the anterior region of the body through the single anterior median artery, paired antennary, and paired hepatic arteries all of which arise from the anterior end of the heart. The large dorsal ahdominal artery extends from the posterior tip of the heart posIt supteriorly through the abdomen just dorsal to the intestine. of the body wall. The sternal artery plies the intestine and muscles is a large branch arising from the dorsal abdominal artery just after It passes ventrally through the nerve cord and it leaves the heart. divides into a posterior, ventral ahdominal artery and an anterior,
ventral thoracic artery.
portions of the body.

These branches carry blood to the ventral Besides the pericardial sinus already men-


tioned, there are others returning the blood to this one.









it is

located beneath the thorax.


several branches lead into the

This provides for a course

through the

From them


by branchio-cardiac

canals and delivered to the pericardial sinus.


perivisceral sinus

surrounds most of the alimentary canal and collects the venous blood



This kind of system


called the

open type because of the

large irregular spaces or sinuses instead of an evenly constructed set
of veins which


a complete circuit of the course.

The excretory system







bodies located in the ventrolateral portion of the head.
richly supplied with blood

These are

and draw the nitrogenous wastes and

excess water from the blood to deliver them externally through
excretory pores located in the coxopodites of the antennae.

The nervous system is of the same structural plan as that of the earthworm, which is a modified "ladder type.'' The two longitudinal cords have come together in the ventral line and run the entire length of the body to form a ventral nerve cord with ganglia. This arrangement constitutes the central nervous system. The ganglia of the anterior three segments are fused into the "brain" or supraesophageal ganglion which is located anterior to the esophagus and is joined to the cord by two circumesophageal commissures or connecone passing on each side of the esophagus. From this dorsal ganglionic mass, nerves pass to the eyes, antennae, and antennules.

The most anterior portion of the ventral cord

receives these com-





This portion, which consists of the fused ganglia from segments three to seven, is known as the suh esophageal ganglia. Ner\'es go from it to the mouth parts, first and second maxillipeds,

green glands, esophagus, and muscles of the thorax. Each segment posterior to the subesophageal ganglia possesses a segmental ganglion
with branches to its respective appendages and muscles. The sense organs include antennae, antennules, sensorj' hairs, statocysts, and

^ _ _ Supraesoplmqeal qanqlion
- -Orcumesophaqeal connective

'Subesophageal qanqlion
Thoracic ganglion


Ring for sternal artery


abdominal ganglion

Lateral nerve

JVentral nerve cord

Tegmental division

Terminal cjanqlion




view of nervous system of crayfish. Notice merging of anterior thoracic ganglia with subesophageal ganglion.

organs (sensitive to touch), the endopodite of which is a relatively long jointed filament. The exopodite The basipodite and coxopodite are is much shorter and fan-shaped.

The antennae are

closely fused to the ventral side of the cephalic region.



pore opens to the exterior through the coxopodite of each antenna. The hairlike processes along the edge of the carapace, on the legs,

and other parts of the body are

also sensitive to touch.

The anten-



nules are tactile and each has two slender filamentous processes, the

exopodite and endopodite.
esses each

antennule has a saclike statocyst in

In addition to these slender jointed procThis its coxopodite.


an infolding from the outside and

lined with exo-

and sensory

Inside of each are small particles of

solid material, such as grains of sand,

which are called


As the animal changes its position the statoliths move about inside of the statocyst and stimulate the sensory hairs. From these stimulations the crayfish is able to determine its orientation in space,
i.e., it

normal walking position, on its back, or standing on its head. These organs serve for equilibrium. When the crayfish molts, the statocysts are temporarily lost and new ones form as the new skeleton develops. If there are no solid objects in the water in which a crayfish lives during molting, there will be no statoliths in the statocysts and the ajiimal has an im-

knows whether

it is


paired sense of equilibrium.

Experimenters have placed only iron filings in the water at such a time and the animals present have used them for statoliths. By bringing a magnet near the crayfish in this condition the statoliths are moved and the animal goes
through numerous peculiar contortions in attempting to respond to these stimulations of orientation. Besides the above functions the
antennules provide the chemical senses of smell and taste.


mounted on movable They are described as compound because each one is composed of a large number of inEach of dividual sight units, each of which is essentially an eye. these units is called an ommatidium, and the crayfish has about 2,500 in its eyes. A single one is rather spike-shaped, tapering from the

which are

of the


type, are

one on each side of the head region.

broader superficial end to the rather pointed internal extremity. A single ommatidium has an outer cornea which is transparent and

supported by some corneagen cells on the vitrella. Beneath this is the rather long crystalline cone beneath which is the rhabdom, an-

Surrounding the latter are sensory cells making up the retinula. The wall of the ommatidium possesses pigment cells along the sides of the crystalline cone and in the retinula. The distribution of the pigment varies with the intensity of the light. The stronger the light the more these cells are expanded and the more direct must be the ray of light to reach the retinula, because
other lenslike structure.






the possibility of reflection within the



light the pigment is partly toward the basal portion of the ommatidia which allows more refraction of rays by the crystalline cones and a combination of


concentrated partly toward the outer and

images in several adjacent units.

In brighter light only the ray from

Cornea - Corneagen




Distal retinal






Proximal retinal pigment cells



Basement •membrane'

Nerve fibers

Fig. 157.

— Longitudinal

section of

pigment when

light is present

in the latter the distal


is in the


concentrated inwardly.

ommatidia from eye of crayfish, a, position position of pigment when in the dark. Notice outward position and the proximal pig(From Hegner, College Zoology j published by The

Macmillan Company, after Bernhards.

directly in front of the cornea will reach the retinula

and stimulate

the nerve cells there.

These cells are connected internally with
of vision

the optic nerve.

The type


in the

compound eye



"mosaic" in that there is registered only a single image by the Each ommatidium which is in focus on the object registers eye. an image of that part. As the object moves, new ommatidia are stimulated and movement is indicated by the rate of stimulation of successive ommatidia. The farther the object is from the eye, the The crayfish eye is often fewer ommatidia will be stimulated. termed a modified appendage because an antennalike structure will
regenerate in case an eye



The crayfish ingests principally flesh from bodies of fish, snails, tadpoles, insects, and other animals, some caught alive and others found dead. The maxillae and maxillipeds hold the morsels while they are crushed by the mandibles. Mastication continues in the cardiac chamber of the stomach and chemical digestion begins in The digestive juices possess enzymes which the pyloric portion. convert the food into soluble form, and as it passes along the intestine, it is absorbed by the blood and distributed to the tissues
over the body.

This conversion of food material into protoplasm



external phase of respiration has put oxygen in

the blood, and

distributed throughout the protoplasm of the

to kinetic

The energy stored in the food material is released or converted form by union Avith the oxygen (oxidation) in the proto-




union there


excess heat produced.


and chemical


the result of the harnessing of this energy.


a by-product of this cataholism, excretory materials, such as excess water, urea, uric acid, and other substances are formed in solution

and are
of these

The green glands relieve the blood collected by the blood. and deliver them to the exterior. Of course growth results


excess food materials are built into the cells at times


the rate of anabolism exceeds that of catabolism.


and the mating takes place either in the spring or fall or perhaps both. The spring hatch become well developed before winter. The eggs produced
These animals are dioecious
(sexes separate)
in the fall


not be laid before spring.






158. Development of the crayfish. A, Toun^ crayfish clinging to swimmerets of mother. B, Second larval stage (2) attached by its chelipeds to hairs (Pl.H.) on a swimmeret (PI.) of the parent. The molted shell of the first larval stage {l) is clinging by chelipeds. portion of the egg-membrane (wi) and shell (Sh.) are still attached to the swimmeret by a stalk (St.). When the first larva hatches it remains attached to the shell by a filament (A/.). By means of these filaments the young remain fastened to the mother during development. C, First larva hatcliing through shell. D, Tlie second larva. (Reprinted by permission after Andrews, 1916, Smithsonian Contributions, Vol. 35.)





In the case of Cayiibarus clarkii the adults retire to holes or bur-

rows at the water's edge during the summer.

It is

here that the eggs

are laid and carried by the female until after hatching; then the

young cling to her swimmerets. In late summer or fall, soon after the young hatch, the adults become very migratory at night, particularly in rainy weather. In this way they help to distribute the young

new water


The female reproductive organs are composed of a bi-lobed ovary and beneath the pericardial sinus. During development the eggs appear in the ovary. Two oviducts lead, one from each side of the ovary, to a genital pore
located beside the pyloric chamber of the stomach
in the coxopodite of the third walking leg (pereiopod) of each respective side.

The ova develop in

follicles in the ovary.

The maturation
the eggs


take place here and,

when mature,

break into the central cavity of the ovary, from which at the time of
laying, they pass out through the oviducts.

The male reproductive

organs are composed of the bi-lobed testis located dorsal to the pyloric

stomach and ventral

to the heart.

Spermatogenesis takes place here

and mature spermatozoa are shed. The tubular vasa deferentia extend posteriorly and ventrally to open externally on the coxopodite During copulation (mating) the sperm of each fifth walking leg. cells are transferred by the two pairs of anterior swimmerets (pleopods) of the male from the apertures of the vasa deferentia to the
annulus (seminal receptacle) on the ventral side of the thorax of the female. Later, vv'hen the mature eggs are laid, they are likely
fertilized as they pass posteriorly in the groove

between the legs on

The fertilized eggs are fastened to the the two and appear much as small bunches of swimmerets by a secretion The later development continues shot-sized grapes hanging there. here, and they are aerated by movements of the swimmerets through
sides of the body.

the water.

Cleavage divisions follow over the surface of the eg^ and the em bryo develops on one side of the mass. The body form with segments and limb buds appears, and hatching occurs in from five weeks to two

The larvae grasp the swimmerets with their chela and remain with the mother for about a month. Two or three days after

hatching they pass through the


molt or ecdysis; that



shed the outer






repeated seven or eight times during

the first season to allow for growth.

The average


span of the

crayfish that reaches maturity


about four years.

Regeneration and Autotomy
This power

limited to the appendages and eyes in this animal,


it is

quite well developed in these parts.




rate of regeneration are greater in younger animals.
lost legs or

Mutilated or

mouth parts

are readily restored.
ability to allow a
if it is

The genus Cambarus has the
off at

walking leg to

a certain line or joint

caught or injured.



leg will develop




This phenomenon
to help in this

called autot-


There are special muscles

and a membranous

valve stops the passage of blood through the leg, thus preventing excessive bleeding.

Bleeding will stop more quickly

the break oc-

curs at such a point than

would otherwise.

Autotomy often



possible for the animal to sacrifice a leg to save its

Economic Relations
Crayfish and the entire class Crustacea are of considerable im-

portance to man.


crayfish, lobster, crab, shrimp,

and others

are used directly as food to the extent that it is an industry valued The at several million dollars annually in the United States. numerous smaller genera, like Daphnia, Cyclops, Cypris, Gammarus, Asellus, and Euhranchipus, comprise a large part of the food


of our food fish either directly or indirectly.

human consumption.

minute ones also feed many clams and oysters and The shrimp and crab fisheries


The more end in are the most

important of the Crustacea on the Texas coast of the Gulf of Mexico.
In the Mississippi valley and on the Pacific Coast the crayfish

becomes a serious pest in the cotton and corn fields of Louisiana, East Texas, Mississippi, and Alabama, They fill the swampy land with their burrows where they come up to the surface and eat the young plants. Frequently their burrows do serious damage to irrigation ditches and earthen dams. Crayfish also capture numerous small fish which are either immature food
used extensively as a food.
It fish or potential

food of such



Characterization of Other Crustacea

Besides crayfish the order Decapoda includes lobster, shrimp, and

They all have ten walking legs for which they are named. The crayfish and lobster are verj^ similar except in size. The shrimps and prawns are marine and resemble the crayfish except that they do not have the great pinchers (chela) and the abdomen is bent sharply downward. The crabs are quite different in shape
in that the



broader than



long, the

to sev-

poorly developed, and folded sharply beneath the thorax.

of different kinds vary in diameter
eral inches.



few millimeters

There are four species of swimming crabs
the most important and best known.

in the


of Mexico, of which the hlue or edihle crab

{Callinectes sapidus,

Fig. 408)


The lady crab and

calico crab are also interesting species.


the blue crab


tured at molting time
it is



called the soft-shelled crab.

At other

the hard-shelled crab.

They maj^ be caught

in baited nets

or on pieces of meat on a line with which they are brought to the sur-

The hermit crab (genus Pagurus, Fig. 408) is smaller and lives in empty gastropod shells by backing into the shell and carrying it around. Due to the cramping and inThe activity the abdomen has become soft and partly degenerate. fiddler crab (genus Uca, Fig. 408) is another very abundant form found on the coast of the Gulf of Mexico and elsewhere. These are small semiterrestrial crabs which burrow in tunnels, and may thus honeycomb large areas of salt marshes. They can run quite rapidly, often moving sidewise, and thej^ are peculiar in that one pincher
lifted out in a dip-net.


of the male, usually the right,




This gives the

appearance of a fiddle and the other, reduced pincher resembles the bow. The large pincher is used in a nuptial dance, and occasionally a large number of these little crabs will be seen raising and lowering these enormous pinchers in concert.
Asellus communis is a common fresh-water form found in streams and pools. A salt-water genus, Idotea, is found in the ocean. The pill bug (Armadillidum) and the sow bug (Oniscus asellus or Porcellio sp.) are terrestrial, living in damp places under logs, stones, or heavy vegetation, and in cellars or greenhouses. Their legs are arranged Respiration is in two groups, which point in opposite directions. carried on by gills on the ventral side of the body and for this





reason they must live in moist plac6s.

a garden pest in There are a number of aequatic forms which are parasitic on fish and others, such as the gribble (Limnoria), which tunnel in submerged wood. The amphipods are sand and beach dwellers which may be found burrowing or jumping around on the seashore or walking on the bot-

They are

that they eat leaves of delicate plajits.

tom The

in fresh water.



the best


fresh-water form.

legs of representatives of this order are divided into

two groups,
These are

with the legs of each group pointing toward each other.
of particular value as fresh-water fish food.

Entomostraca as a group is composed of many smaller crustaceans occurring in great numbers in both marine and fresh waters. The fairy shrimps (EuhrancJiipus) are delicate, transparent and feathery


are about three-fourths of an inch in length.


Fig. 159.

Asellus, a



fresh-water crustacean. Supply House.)

(Courtesy of General Bio-

up and their long, leaf-like appendages the body; these appendages serve also as respiratory organs. The^^ live in cool streams during the spring ajid fall. The summer is passed in the egg, which can withstand complete dryness. Many of them are parthenogenetic, hence, males are rare. The common marine form is Artemia, often called brine shrimp. The water fleas including Daphnia of order Branchiopoda, Cyclops and Diaptomus of order Copepoda and other small Crustacea constitute an important common group. Daphnia is one that is enclosed in a delicate bivalve shell. The second pair of antennae are very large and are used in swimming. The shell is beautifully marked and terminates in a caudal spine. They are only about

swim with hang from

the ventral side

one-tenth of an inch in length. Cyclops is another common freshwater form with the antennae shorter than the cephalothorax whose



body length is also about one-tenth inch. It has a single median eye, and the females frequently are seen with a pair of egg sacs attached at the base of the abdomen. Diaptomus, another Copepod, is a common form of about the same form and size as Cyclops, except that the antennae are nearly as long as the body. Argulws is a genus of Copepods which is parasitic on fish, and the individuals are called fish lice or carp lice. They are flat creatures and are found running around over the scales of their hosts. Some of the other forms are parasitic on the gills and fins of fish and their bodies become greatly modified. The ostracods are small, swimming, bivalve forms that are sometimes called swimming clams. This group has beautifully marked valves; in fact, these animals are the most beautiful found in the

Adult barnacles of order Cirripedia bear so


resemblance to

other Crustacea that they are usually overlooked as such by the

They are completely encased in a thick shell of several sections and have the general appearance of an oyster or clam. They are sessile in habit as adults, though free-swimming in the
larval stage.

Their entire

life is

spent in marine waters.


are several characteristic barnacles, rock barnacles on rocks, whale
barnacles from ships and pilings, and gooseneck barnacles of the
stalked type.
like bristles

After attachment, the legs become modified into featherwhich are used in gathering food. Sacculina (Fig. 404) is a genus related to true barnacles which has gone parasitic on crabs and has lost all resemblance to animal form. It settles on the body of a crab, makes its way to the interior and there becomes a branched mass of tissue which penetrates by roots to all parts of the body of the crab. After a time a baglike portion forms and projects externally on the ventral side of the abdomen of the crab.
Recapitulation Theory

statement of this idea, which was developed by von Baer, Haeckel, and others, and is so well illustrated by the comparison
of the phylogenic


and embryonic stages of certain Crustacea, may


This theory maintains that certain developmental stages or structures of the individual are related to ancestral conditions. That is, the individual in its development tends to repeat in an abbreviated fashion the history of the development
at this point.


of the race.
is still




Briefly stated ontogeny recapitulates pliylogeny.


some doubt as

to the validity of this generalization in direct




example which


frequently cited


that of the devellarva,


of the shrimp, Penaeus,

which hatches out as a nauplius

Nftuplius «t*c{<«

Fig. 160.


stage of the barnacle, Balanus. logical Supply House.)

(Courtesy of General Bio-

Fig. 161,

Fig. 162.

—Zoea and Megalops stages of developing Crustacea. Supply House.) Crabs include these stages their development. (Courtesy of General Biological Fig. 162. — Schizopod or mysis stage through which the shrimp and lobster pass. (Courtesy of General Biological Supply House.)
having a single median eye and only three pairs of appendages. Following the molt, this nauplius changes to become the Prozoea stage, possessing six pairs of appendages. The next molt brings on segmentation and some change in form. This stage is called the Zoea



and resembles very closely the adult Cyclops of modern Copepoda. The Zoea transfonns during further molts and growth to a stage with thirteen segments and a distinct cephalothorax which resembles the Gammarus is adult Mysis and therefore is called the Mysis stage.
also in about this category of phylogenetical development.


ing the next molt the mysis stage becomes a juvenile shrimp with nineteen segments. The life history of the barnacles and Sacculina

has illustrated quite forcibly the possibility of such a relationship. There are extinct forms also whose adult condition was that of one This idea generally has served as of these developmental stages.
a great stimulus to the study of embryology and the theory of evolution as well as serving to establish natural relationship of

animal groups.
Phylogenetic Advances of Arthropoda

Greater specialization of segments,


paired, jointed ap-

pendages, (3) chitinous exoskeleton, (4) gill and tracheal respiration, (5) dioecious reproduction, (6) development of eyes and other sense organs, (7) green glands and malpighian tubules (insects) as excretory organs, (8) organization of social life.




(By Vasco M. Tanner, Brigham Young University)



interesting group of arthropods,


considered as the class




the more tropical

and semitropical
These primitive

regions of the earth south of the Tropic of Cancer.

nocturnal forms, according to Austin H. Clark, are found in areas that vary in annual temperature from 50° to 80° P.; in fact, most of
the species are confined to habitats in which the temperature does not vary beyond the limits 60° to 70° F.

The onychophores are characterized

as soft-bodied, wormlike, ter-

forms with internally segmented bodies. The body may be divided into a head and abdomen. On the head is one pair of annulate antennae and a pair of jaws. The body bears many pairs of legs which are not distinctly jointed, but are provided with transverse pads

and apical




by means

of tracheae

which communicate with spiracles that are, in some species, arranged in rows on the body. The excretory system consists of nephridia arranged in pairs in the body segments and opening to the outside by a pore at the base of the fourth and fifth legs. The genital organs
discharge at the posterior end of the body.

The nervous system


consisting of separate longitudinal nerve cords connecting
of ganglia.




pair of eyes
consists of


located at the base of each


The body wall

an unsegmented dermomuscu-

lar covering.

In commencmg on the ancestry of the Onychophora, Prof. J. W. Folsom has the following to say: "Onychophora, as represented by Peripatus, are often spoken of as bridging the gulf that separates the Insecta, Chilopoda and Diplopoda from the Annelida. Peripatus
indeed resembles the chaetopod Annelids in

segmentally arranged

nephridia, dermomuscular tube, coxal glands


soft integument,

and resembles the three other

classes in its tracheae, dorsal vessel with

lacunar circulation, mouth parts, and salivary glands.

These resemblances are by no means

however, and Peripatus



does not form a direct link between the other tracheate arthropods and the annelid stock, but is best regarded as an offshoot from the base of the arthropodan stem."


little is

known about

the habits of the onychophores, except

that they live under stones and the bark of trees, feeding upon small insects and spiders which they capture in a slime produced and
forcefully discharged

from glands which open on the oral
in a season.



of the species are viviparous, a single female producing as


as thirty living


About seventy-three species and fifteen genera are known from the two families Peripatopsidae and Peripatidae. A number of species in the family Peripatidae are found in tropical America; Macroperipatus perrieri (Bouvier) is found at Vera Cruz, Mexico; while
Australia, Tasmania, the species of the family Peripatopsidae are confined to New Guinea, New Zealand, Cape Colony, and Chile.



Diplopoda and Chilopoda are considered by some au-

The more recent students thors as orders of the class Myriapoda. of these groups, however, have adopted the plan of classification followed here.



Peripatus capensis. Entomology.



Redrawn by Nelson

(After Moseley A. Snow.)



The Diplopoda are terrestrial arthropods commonly called millepedes. The body is composed of three regions: the head, thorax or trunk, and the ahdomen. The head bears a pair of short antennae, ocelli, and mouth parts consisting of a pair of mandiUes and a pair
of maxillae.

Just back of the head


a segment with a well-developed
of this group as a seg-

tergite, the collum, considered

by some students

ment which has played an in some of the other groups

important role in the formation of the


of arthropods. A however, to belong to this first segment, as does a single pair of legs These four segments are said to to the three following segments. Ducts from the reproductive organs open at constitute the thorax.

single pair of legs seem,

the base of the second pair of legs on

tlie third body segment. an indefinite number of segments, each The abdomen consists of consisting of a tergum and two sterna. Each sternum bears two pairs






of legs and two pairs of spiracles. The spiracles are closed by a valve and communicate with tracheal pockets and unbranched tracheae.

Embryological evidence supports the belief that the abdominal segments have resulted from the fusion of two segments. The legs are jointed very much as in the insects. The tarsus consists of

the three segments.

The heart

a dorsal structure with side valves and an anterior

The diand intestines. The excretory wastes of the body are removed by two or four pairs of Malpighian tubules which discharge their excretions into the ingestive tract consists of a mouth, esophagus, stomach,

tube to the head similar to the arrangement in the insects.



Fig. 164. Julus terrestris. millepede. Side view of anterior end. a, antenna; ab. abdomen; colj collum (first thoracic segment) e, a group of ocelli go, genital opening m, mandible g, gnathochilarium h, head I, labium th, thorax. (After Borradaile and Potts, Invertebrata. Redrawn by Nelson A.

A common






living plant substance.

The millepedes feed upon vegetable matter, decaying as well as They are slow-moving, wormlike creatures,

living in dark, moist places.


disturbed they usually roll them-

laid in damp earth, and young hatch they are very small, consisting of only a few segments and three pairs of legs. The diplopods are found in most parts of the world. In the United States there are six important families and about 120 species. The family Julidae is widely distributed. The species Julus hortensis Wood; J. virgatus; J. hesperus Chamberlin; and Spiroholus margin-


into a little


The eggs are



L. . Scutigera forceps. The nervous system consists of connected paired ganglia in each leg-bearing somite and a subesophageal and supra-esophageal ganglia which supply nerves to the eyes. 627. of Agriculture. Polydesmus serratus Say is a common species in the United States west to the Mississippi River. also upon spiders and Mollusca. S.— 290 atus are fairly TEXTBOOK OF ZOOLOGY common in many parts of the United States. legs. 165. Marlatt. Dept. the last two segthe other body somites bear a pair of first seven-jointed walldng legs. U. and other . The Chilopoda. consisting of fewer segments which bear but one pair of as a rule. of the parts of the head. A pair of Fig. Members of the class Chilopoda differ from the Diplopoda by their dorsoventrally flattened bodies. 1914. and by their long antennae.) poison claws are located on the ments are legless. antennae. The circulatory system is well developed it extends the full length body and gives off in each segment lateral vessels or arteries. mature The Chilopoda are carnivorous. and each of body segment. (From a paper by C. No. In the family Polydesniidae. preying upon adult as well as imThe mouth parts insects. Farmer's Bulletin. mouth parts. consist of a pair of mandibles and two pairs of maxillae. The long-legged house centipede. move faster than millepedes.

heros Girard and Arthrorhahdmus pygmaeus Pocock of the family Scolopendridae. connected to a pair of spiracles empty into The tracheae are branched. heros is a large dark greenish colored species found in Kansas. Scolopendra. Chamberlin.— PHYLUM AETHROPODA The digestive system is —CLASS ONYCHOPHORA 291 fairly simple. reports the following species of centipedes for the Western United States: Scolopendra morsitins Linn. The species S. being on each body segment. Dr. and intestines. The large centipede of the Southwest. and Texas. 166. R. Fig. S.. stomach. Southern California. consisting of the esophagus. . A pair of ]\Ialpighian tubules the anterior part of the intestines. V. one of the leading students of the millepedes and centipedes. Arizona.

In attempting to give the characteristics of this class we will list only the obvious characters and then remind the reader that the members of this group have more likenesses to the Arachnida than to any other assemblage of animals. the are to be found upon this part of the body. each consisting of seven joints. Brigham Young University) Probably the most heterogeneous class of arthropods is the class Arachnida. The poison serves as an aid to kill insects and other animals used by the spider for food. consisting of a basal part or mandible and a terminal claw. which are remarkable number and arrangement in the various families abdominal appendages which have been modified into spinning organs. They are used in handling the food. pseudoscorpions. four pairs of walking legs. consisting of six joints. The poison glands in the mandibles discharge their poison through the movable sharp-pointed claws. hence. the cephalothorax. the cephalothorax and the abdomen. results from the fusion of the head and thorax. respiratory organs. are located in front of the legs. scorpions. chelicerae. lung books. which is now a pigeon-hole for the spiders. the reason for having this class intact. the chelicerae and pedipalpi. as the Pentastomoids and several other less common orders.CHAPTER XX PHYLUM ARTHROPODA ARACHNIDA (CONT'D) (By Vasco M. harvestmen. bear animalcules. The first division of the body. mouth parts consisting of but two pairs of appendages. no antennae or wings. no doubt. and no metamorphosis. such and Pycnogonids. mites. The pedipalpi. simple eyes. are two-jointed. 292 . Tanner. Six pairs of appendages The first pair. The mandible is stout and covered on the inner surface with small teeth and setae. The spiders and their relatives are characterized by having two main divisions of the body. the development being direct. king crabs. ticks. Spiders The spider's body consists of two divisions: the cephalothorax and abdomen. whip scorpions. that are definite in .

The four following pairs are the legs. clearly shown. Latrodectus mactans. the are used in chewing the food. — CLASS ARACHNIDA 293 and in the males the terminal segments are used as copu- latory organs. . Through its small opening the liquid portion of the prey is sucked up by means of muscles which are attached to the dorsal wall of the cephalothorax . The mouth is located just behind the chelicerae. by permission of the Utah Agricultural Experi- — are the front pair of breathing openings or slits which communicate with the lung books. Each leg is made up coxa. Ventral view of adult female. trochanter. The basal segments. ment Station. femur. namely.PHYLUM ARTHROPODA as feelers.) Fig. 167. of These palpal organs are useful in the classification many of the families of spiders. Near the end of the abdomen are three pairs of spinnerets. of appendages of seven segments. tibia. patella. Near these is the opening of the reproductive which in the females is protected by a plate called the epigynum. metatarsus. and tarsus. or maxillae. organs. black widow spider. The cephalothorax and abdomen are connected by a narrow waist. The digestive system of the spiders is well adapted for its fluid food. just back of the last pair of legs. The eyes are located on the front of the head and usually are eight in number some species have fewer. The hour glass on the ventral part of the abdomen is (From Knowlton. In front of these is an opening to the tracheae and just posterior to them is the anus. hanging from web. but never more than this number. On the under side of the abdomen.

The nervous system is composed of a large ganglion in the cephalothorax which is connected with a dorsal brain by a nerve ring around the esophagus. The hatching takes but a day or two. teriorly into the or heart. If the young spiders do not get out of the cocoon soon enough. In the posterior part of the abdomen is is a pair of branching air tubes or tracheae. The excretory wastes are removed by the Malpighian tubules. arteries. and by vestigial coxal glands. which is called the sucking The posterior portion of the stomach gives off five caeca which are supplied with blood vessels from the anterior aorta. while those laid in the summer develop within a few days. receives its blood It is then forced also pos- anteriorly through the aorta into the cephalothorax and abdomen. The eggs are fertilized within the body of the female and then laid in silken bags or cocoons. the development being direct. the testes in the male. located above the intestine. The young spider is pale and soft bodied. eggs laid in the autumn develop slowly all winter. it molts a number of times. which discharge into the posterior portion of the intestines. As the spider grows. from the tips of the pedipalps. The rate of growth of the eggs varies according to the conditions. The respiratory functions are carried on by the lung books or sacs. which contain a number of thin plates through which the blood passes on its way to the heart. they . The male and female openings are near the lung books. one for the reception of the hepatic ducts from the liver and the other at the posterior end for the formation of a stercoral sac or pocket. The heart from the body by means of three pairs of ostia. For example. The vascular system consists of a muscular tube and veins. one to receive the sperms into the sacs. so that they have their freedom. In many species these egg bags are carried on the abdomen of the female. The reproductive organs consist of the ovaries in the female and The female has two sets of openings. and the other is the exterior opening of the oviducts.294 TEXTBOOK OF ZOOLOGY stomach. Nerves pass from the ganglion to the limbs and the abdomen. This system of breathing found only in the arachnids. The intestine passes through the small waist connection into the abdomen and becomes enlarged in two portions. and the anterior portion of the stomach. but in a few days it molts for the first time and then begins to look like an adult spider.

PHYLUM ARTHROPODA — CLASS ARACHNIDA 295 begin to eat one another. and some them are not found The following eight : families. the five of the common orders of the class Arachnida true spiders. The eyes are eight . A Natural Classification of Spiders. They They possess eight are often highly colored for protection. The species. are common in the United states 1. Latrodectus mactans. the king crabs. under stones. Some spiders live two or three years. Theridiidae. developing from over-wintering eggs in the spring. and at They build rather loose webs from times under boards and rocks. crab spiders. Petrunkevitch. Members of this family have a toothed comb on the tarsi of the fourth pair of legs. but in wait in the flowers for insects that visit them. L. Lycosidae. and the body is rather flat. (4) Phalangida. eyes in two rows. and eight eyes. the mites and ticks. Many spiders live only one year. while other species live many years. According families. frondicola. lie These crablike spiders do not con- struct webs. to a recent study. the comb-footed spiders. and grass. The eyes are arranged in three rows of four. or in silk tubes on shrubs long enough to The drassids live mainly on the ground. This family contains many of the largest native species. growing to maturity during the summer. of food. the two anterior pairs of legs are longer than the other legs . They are found mainly upon the ground running around in search The females carry the cocoons attached to their abdomens. and two. the order Araneida Species of of made up of sixty-two many of these families are rarely encountered. and Allocosa parva are 2. however. Classification of the Arachnida For purposes of this chapter we shall confine our discussion to but : (1) Araneida. The spinnerets are generally extend a little behind the abdomen. (2) Acarina. laying eggs. Thomisidae. the scorpions. the daddy longlegs. common species. A. reported as a poisonous species. Lycosa gulosa. buildings. is a member of this family. The black widow. (3) Scorpionida. two. The hourglass on the ventral part of the 3. L. in America. three claws. Drassidae. and (5) Xiphosura. which they hang back downward. common species in the western states. the wolf spiders. fences. 3Iisumena vatia and Xysticus nervosus are 4. The spiders of this fam- ily are found on low growing vegetation. by is Dr. kochii. and then dying in the fall. abdomen is scarlet in color in the live spider.

They are arranged in three rows. occupying an area on the cephalothorax known as the ocular quadrangle. and beet fields. extensa. Argiopidae. and holes. buildings. The trap-door spiders live in tunnels dug in the ground and provided with a hinged door which closes the entrance to the tunnel so perfectly that it is almost impossible to locate the tunnel. tion by their jumping and bright coloration. the fall of the year in potato. Metargiope trifasciata. Some of the common species of this family are Tetragnatha lahoriosa. and on the ground. Icius similis. Representatives of this family are 7. and Salticus senicus. hairy. 6. Drassus negledus is a representative species. P. This is one of the large families of spiders. . the members of which have eight eyes. common garden spider. and Aranea gemma. plants. The males and This family is females differ considerably in size and structure. Members of this family attract attenThey live on fences.296 in TEXTBOOK OF ZOOLOGY number and arranged in two rows. Attidae. fences. They do not spin webs for the capture of prey but only for their protection and cocoons. Most of the species of Tetragnatlia are found in moist places they Their bodies are round and build their webs over running water. black spiders with eight eyes and chelicerae projecting forward. After building the web they lie in wait for some insect which may serve for food to become entangled in it. which are large nocturnal species. They are confined to suitable places on vegetation. found in Metargiope trifasciata is the long. The tarantulas. Aviculariidae. Some of the common species of our fauna are: Phidippus worhmanii. buildings. the orb-web spiders. the tarantulas. They are confined to the south and southwestern United States. . tomato. jumping spiders. world-wdde in distribution. but in this family there are They differ from the wolf spiders in that the body is much longer than wide and flattened slightly on the back. Neoscona benjamina. 5. where they construct their webs and then remain near by to watch them. This is a large family. large. The spiders of this family build rather large typical webs for the purpose of capturing their prey. It is represented by three hundred and fifty species in America north of Mexico. formosus. It has a silvery white color and is about an inch in length when full grown. live under rocks. The eyes on the front row are the largest. and three claws on the tarsus. T. only two tarsal claws. EuopJirys monadnock. The eyes of this group are most distinctive.

with eight . the pholcids.^^nphs. blister pyri.PHYLUM ARTHROPODA — CLASS ARACHNIDA 297 in holes. All mites lay eggs Avhich hatch into six-legged After some growth and moulting they develop into adults legs. and the legs are one and one-half to two inches species are used to carry the eggs. in Utah. (Greatly enlarged. (After Knowlton. red spider. 168. n. 168. In this species the chelicerae are Fig-. Order Acarina.) (After Sorenson. —An adult mite. They are. webs in dark places. tanneri. by permission of the Utah Agricultural Experiment Station. common tarantulas. living upon various small — . such as cellars and There are six genera in the United is Pholcus phalangioides a large species. —The brown mite. telarius. Fig. are spiders with long legs and six or eight eyes. States. Bryobia praetiosa. 170. some of the aquatic species are able to shift for themselves. hentzi are 8. by Professor John H. and under debris in the daytime.) Fig. Eriophyes (After Sorenson. 169.) Fig. 169. in the main. water animals. Fig. ectoparasites or endoparasites however. by permission of the Utah Agricultural Experiment Station. 170. The abdomen is elongated in form. Comstock. the student should consult the most valuable treatise on this subject. by permission of Experiment Station. long. The Spider Book. They feed upon beetles and other ground-living insects. Eurypelma steindachneri and E. —An the Utah Agricultural Tetranychus adult female. Mites and ticks are mostly small creatures with the cephalothorax and abdomen fused solidly together. Pholcidae.) Fig. For a discussion of the other spider families found in this country. Three found in the genus Physocyclus : two of them are Physocyclus glahosus which is found in Florida and P. They build loose protected corners of buildings.

Eight of the species be- long in the genus Eremohates. called the pectine is ( A peculiar comblike structure. are widely distributed. After taking a meal of blood. Hadrurus hirsutus. Texas cattle fever and Rocky Mountain spotted fever are dreaded diseases that are carried by ticks. Four families are represented in the United States. They feed upon insects and spiders. The clover or brown Bryohia praetiosa. trees. Ticks of the family Ixodidae attach themselves to their host. suck blood. practically all kinds of cultivated plants. with a flattened body and elongated abdomen terminating in a They are nocturnal. ticks are all fairly large parasitic species. Tetranychus telarius. The common red spider. is cosmopolitan in distribution. they leave the animal and go into hiding. and shrubs. It is reported that it attacks over 250 Another common mite pest is the leaf blister mite. They breathe by means of lung books and have a ing direct development like the spiders. which is common shade in this country. Another interesting little group of arachnids is the order Solpugida. This species attacks the leaves of the pear and apple. to Eriophyes pyri. the mother takes care of the young. and common annual and perennial plants. and Vejovis mexicanus of the family Vejovidae are common species. protectthem by carrying them around on her back and by helping them to catch their prey. forest trees. which belongs the family Eriophyidae. garden plants. attacks field plants. At times they are a serious pest to poultry. leaves results in the development of defoliation. and they are found only in the southern and western states. feeding upon mite. . It feeds upon fruit and shade trees. species of plants. found in the ventral part of the second abdominal segment. The damage to the brownish blotches and partial Those belonging Argasidae attack only warm-blooded animals. fruit trees. They are viviparous. known as brown mites and red spiders. found in the same territory as the scorpions and represented by twelve species contained in three genera. and grow to many times their original size. also specialized stinging organ.298 TEXTBOOK OF ZOOLOGY group of mites belonging to the A family Tetranychidae. hiding during the day under rocks and burying themselves in the sand. The to the family The order Scorpionida consists of large-sized arachnids with pedipalpi which resemble the chelipeds or pinchers of the crayfish.

eggs are deposited in the summer in shallow water in small sandy depressions w^here they are then fertilized by the male.PHYLUM ARTHROPODA The Phalangida. ovipositor in the female in the United States. an and a penial organ in the male. There are about seventy species only two eyes on the cephalothorax. The males are The a little smaller than the females. The harvestmen do not have silk glands and therefore do not construct cocoons for the eggs. long legs. but similar in appearance. commonly — CLASS ARACHNIDA 299 called harvestmen and daddy longlegs. representing six families. which they lay under stones and under the bark of trees. The order XipJwsura. which consists mainly of worms. . king primitive genus and five species. The respiratory organs consist of tracheae which open through abdominal spiracles. are located on the obscure division between the cephalothorax and abdomen. and the presence of The reproductive organs. On the abdomen are six pairs of appendages. the abdomen consisting of nine segments. the cephalothorax and the abdomen. The basal parts of the appendages situated around the mouth are used for crushing the food. There are six pairs of appendages on the cephalothorax. Phalangididae are the most common and widely crab. Because of its shape and resemblance to the crabs. is represented by only one living Limulus polyphemus is the American species found along the Atlantic Coast from Maine southward. can be distinguished from other arachnids by their body which is composed of a broadly fused cephalothorax and abdomen. The body consists of two regions. the last five pairs bearing book-gill structures used in respiration. Species of the family distributed. it has been called the horseshoe crab.

to ice cold. of the late Paleozoic era. on and in animals. from the tundra is said to be over 650. our crops and damage our stored foods. is because insects exist in every type of habitat known. The Russian biologist. Brigham Young University) Insects are the most abundant creatures on the earth today. H. which are carriers of disease. in trees. in fresh water that ranges in temperature from 50° C. Various explanations have been advanced to account for the great adaptability^ of insects in filling practically every niche in nature. and a small size new place in the world of living animals. on the vegetation of plain and swamp. Kennedy. unlimited external features. of the north to the tropical as well as man. This means that for probably one hundred million years these arthropods have been adjusting to a changing environmental complex.CHAPTER XXI PHYLUM ARTHROPOD A CLASS INSECTA (CONT'D) (By Vasco M. 300 . in dry desert conditions. and the success with which they have met the challenge is quite evident today. There which have never no doubt. however. This great class Insecta has been upon the earth from the Pennslyvanian times. Tanner. in the soil. is is we may say that One noted entomologist has said that this an age of insects. has pointed out that there are advantages as well He says: as disadvantages to the possession of an exoskeleton. They ravage In short. S. insects are omnipresent.000 living species. to the present. They are found in sea water along the shore. and to this we may add that every man's farm "no man's land" and that the contending forces are insects and man. permitted them to develop strong appendages. This. argues that the chitinous in that it has exoskeleton has been of great value in the evolution of this group. entirely which has opened up an Dr. C. Chetverikov. many of been seen by the great majority of mankind. S. many of pampas.

The paired appendages furnish much evidence that the head has resulted from the fusion of several segments. the abdomen bears a variable number of segments in the various groups of insects. when they are present in the nymphs and adults. successful man will be in his evolution during the next fifty million . There are two kinds of eyes eyes. and compound The simple eye is a small area consisting of a single cornea. . also the genital apertures which are situated near the anus at the posterior end of the body.PHYLUM ARTHROPODA "The exoskeleton has made in fully as — CLASS INSECTA 301 possible very definite advances in the evolution of insects. and labium are considered as developing on distinct somites. the Simple eyes are generally found in varying numbers along with compound eyes in adult insects. the sucThis leads us to wonder how cessful creatures they are today. visible. Head The head consists of a number of immovable plates or sclerites forming the head capsule. ment as the Will he be able to meet the demands of a changing environinsects have? INSECT CHARACTERISTICS body is divided into three The head. : ocelli. and abdomen. mandibles. other distinctive characteristics. which keeps the hemolymph or blood from becoming impure and finally their great variability and power to reproduce. bears a single pair of antennae. maxillae. antennae." Aside from the ehitinous exoskeleton. the eyes. or simple eyes. such as power of flight. no doubt. have made the insects. to which are attached the paired appendages. they are. which is possessed by no other invertebrate animal a tracheal system. The eyes. years. which consists of six segments. the thorax consists of three segments and is the region which bears three pairs of legs and two pairs of wings. six anterior Evidence concerning the fusion of the segments of the body in the formation of the head comes through an embryological study of the insect. Insects are Arthropoda in which the regions. however. the head. In many insects the sutures separating the sclerites are and these plates and depressions have been given definite names. thorax. and the mouth parts. but at the same time has limited their evolution many other ways. usually lack- .

(Photographed 171. female. is shape. Brachypephus magnus. Some students of the insects maintain that the I I A. and they vary in among the various groups of I . TEXTBOOK OF ZOOLOGY The compound eye consists of many facets. B.) by Leo compound eyes function ocelli are mainl}^ in detecting movements. The form found on the plains. which are the hexagonal-shaped corneal ends of structures called ommatidia. while used to detect light intensities. Murray. —Western lubber grasshopper. and B.302 ing in beetles. and position on the head but one pair of antennae or feelers. Fig. There size. T. A. male. It has been shown by experimentation that many insects are able to detect colors. and insects are short-sighted. The facets are convex.

labrum i. Antennae are useful structures in the classification of insects. podical plate. wing. the insects by . coxa. pronotum first abdominal segment. 29. — CLASS INSECTA in 303 insects they The . 25. ocellus 8. metaIJi. . some are tactile or in others they are respiratory or olfactory. femur. . V. 12. . or auditory. . pubMosby Company. femur. tergum. spiracle. feelers function in many ways. 10. lished by The C. mandible 3. 19. ovipositor. tibia. 172. compound eye 7. Neuroptera. mesothoracic leg. . supra-anal plate. clypeus 5. mesothoracic 9. Odonata. antenna. 18. magnified. 26 V.) PHYLUM ARTHROPODA insects. 16. 28. 21. auditory apparatus. —Detail of ommatidia. gena 11. 2i. and suctorial or sucking mouths.' Z7 28 . The external features of a grasshopper. 23. 30. 27. 15. . General Biology. frons 6. . . Fig. (After Turtox key card. tarsus. trochanter. femur. spiracle. . 173. thoracic n.) The mouth parts are chitinous structures and are represented in two distinct types: mandibulate or biting. courtesy General Biological Supply House. maxillary palp. prothoracic leg. (From White. sternum 31. vertex 13. Fig. labial palp. wing. thoracic. 26. Some of the insect orders which possess mandibulate mouth parts are Coleoptera. spiracle. 2. may be used to hold the female during copulation. cercus 20. — . metatlioracic leg. : . 1. 22.

.. In the mandibulate mouth parts there is a labrum or upper lip which is attached on its upper border to the clypeus and extends down over the mandibles. — . stipes. Lac. L. G. The mandibles or jaws are true appendages which move in a trajis- . A labial palp. The Lepidoptera. palpifer. • .. labium. and Orthoptera. M. group of organs within the insect body vary in form as do the torial orders. mouth parts. La.. and Siphonaptera are suc- knowledge of the mouth parts is very useful if No other insects are to be effectively controlled and classified.304 TEXTBOOK OF ZOOLOGY Mallophaga. . Dermaptera. The mandibulate mouth parts are the type from which the suctorial type has been evolved. ilfa.. General Biology. Diptera. Pa. first maxilla. clypeus L-i". Heteroptera... Me.. labium (second maxilla). . ligula galea. Fig.) /St.. laclnia mentum MP. mandible. I/t. Mouth parts of Bhomaelia microptera. submentum . Ga.. Homoptera. maxillary palp. Isoptera. 174. (From White. Sin. cardo Gl.

and palpus. galea. In the mosquito the hypo- pharynx long and slender like the mandibles and maxillae. The dorsal or back surface of .PHYLUM ARTHROPODA verse plane. proximal portion of the beak. attached to the distal portion of the cardo is the stipes. The saliva causes an irritation. In the typical sucking insect the mouth parts consist of two pairs of sicklelike or styletlike structures which are modified mandibles maxillae. of the mouth. and metathorax. The maxillae or second pair of jaws in a similar plane. probably detecting senses similar to our taste own senses of and smell. thick plates of chitin with toothed edges adapted for cutting or crushing food. Thorax The thorax is composed of three segments. Projecting from the mentum on . which bears three sclerites the lacinia. — The lacinia is provided with teeth or spines which aid in holding and chewing the food. — CLASS IXSECTA 305 They are hard. lie under the mandibles and move is much more complex. The The cardo maxillae. Food is taken in a liquid form by being sucked up through the labial sheath. In some beetles the mandibles become greatly enlarged and apparently worthless. the salivary duct extends throughout the entire length of the food channel. as and found in the mandibulate orders discussed above. The hypopharynx or tongue arises from the labium into the cavity and bears the opening of the salivary duct. The palpus is composed of four or five segments and is sensory in function. which consist of sclerites. are the piece which hinges the maxillae to the head. The labium is hinged to the head by the mentum from which extends one or two pairs of lobes. The labium forms a long sheathlike structure in which the styletlike manThe upper lip or labrum covers over the dibles and maxillae lie. A pair of jointed legs is attached to each segment and most adult insects bear a pair of wings on the mesothorax and metathorax. and if the mosquito is inis fected with malarial organisms they are introduced into the blood stream of man. mesothorax. each side is a palpus which consists of one to four segments it functions as a sen- soiy organ. The labium or lower lip is formed by the fusion of what is believed to have been a second pair of maxillae. the ligula. which are called the prothorax.

coxa. the under and each pleurum. They are also modified for the production or reception of sound. General Biology. and are useful in classification. — shape. tibia. the ventral or side. and tibia are large. or thigh. and up The coxa forms the joint by means of which the leg is attarsus.) . In some species they also exhibit secondary sexual char- The wings are thin folds of the skin. running. shaped and strengthened with The presence of wings is one of the most . and jumping. walking about. swimming. trochanter. the sternum. pulvilli Ta.306 TEXTBOOK OF ZOOLOGY is the segments of the thorax surface. forming the greater part of the leg. femur. Jumping leg or third thoracic appendage of Rhomaelia microptera. Fig. The legs are made main segments: the coxa. In insects the tarsal segments may differ in size. tarsus. P. acters. Ti. The trochanter is small. (From White. while the femur. The tarsus or foot is composed of five smaller segments and a pair of of five claws. for the collection of food. C. such as pollen. Tr. tibia. veins in various ways. trochanter. The legs are greatly modified for obtaining food. length. 175. called the tergum. tached to the body. and number. and for copulation. femur. F.

. Within the abdomen are found the respiratory. (From Henderson. genera. On the eighth and ninth segments of the female and the ninth of the male are paired structures forming the genitalia which are the external organs of reproduction. the fore of Utali Agricultural wing. and genital systems.PHYLUM ARTHROPODA — CLASS INSECTA 307 characteristic features of the insects. A. and character of the veins and cells are an aid in grouping insects into The majority of insects possess two families. .) Abdomen The segments of the abdomen are usually simple. and species. digestive. Because of their great variawings are of much value in classification. The wing is com- posed of a network of thickened lines called veins and thin areas between the veins called cells. The number. but the number There are only ten present in many insects. Fig. Experiment Station. yet in the embryo of the insect there are eleven. tion. the hind wing. B. arrangement. 17G. permission —Right wings of a grasshopper. that have but one pair and pairs of wings some groups are wingless. there are some. varies greatly in different insects. however. The jointed appendages have been almost entirely lost in adult insects.

It a most interesting organic substance. All the tubercles. All of the . Hemiptera. and Plecoptera. setae. — — There are four types of development or metamorphosis first. and fourth. The hypodermis has its origin in the ectoderm and is the active growing layer of the body wall. pauroametabolous or development without metaholotis or gradual metamorphosis third. and from pupa to adult constitute metamorphosis. The exoskeleton consists of three layers known as the cuticula or outer layer which is impregnated. and which alters. Metamorphosis Metamorphosis includes the alterations which an insect undergoes after hatching from the egg. In the Odonata. second. more or less. the general form and life of the individual. Chitin is a substance found in many is parts of the insect body.308 TEXTBOOK OF ZOOLOGY Body Wall Another distinctive feature found in the arthropods is the chitinous body wall. not destroyed by caustic potash. Collembola. These structures are of importance in the identification of insects. Isoptera. holometaholous or complete metamorphosis. resembling horn in some physical ways. remaining practically the same form as the adult insect during all the development. which after hatching. there is a type of development in which the nymphs gradually increase in size and the rudimentary wings and genital appendages become adult structures. The ametabolous insects are the Thj-sanura. and scales of the body wall are formed by the cuticula. and Dermaptera. Mallophaga. spines. the newly hatched naiads pass through an incomplete metamorphosis. and Pediculidae. All the changes which are undergone by a butterfly in passing from egg to adult each change from egg to larva. Homoptera. This is development without metamorphosis. from larva to pupa. hemimetaholous or incomplete. Thysanoptera. with calcareous matter. but Chitin is it especially serves to give firm- ness to the cuticula. grow through a number of instars. extensively. which provides the only rigid support for the body. This is known as paurometabolous development. In the following orders: Orthoptera. and the 'basement memhrane. the hypodermis or intermediate layer. : . Ephemerida. metamorphosis.

Lepidoptera. minute size. use of flight in locomotion. and rate of multiplication. CLASSIFICATION Because of the important role insects play in the life of man it is worth while to be able to recognize some of the common orders. elongated rather flattened bodies. the arrangement folthat of discussing them according to their development. and abdominal appendages. —Ametabola are insects without metamor- Order Thysanura. the adult or imago emerges. and the type of development. Coleoptera. goes through a complete metamorphosis. The Holometabola. They are soft-bodied small insects. phosis. and Pterygota. growth period times it comes in The remarkable adaptation of the immature stages of insects to their food supply has undoubtedly had much to do with their great success as a group. Subclass Apterygota. The number of orders recognized in this class varies considerably. depending on the authority followed. gradually takes on the adult form and after a few days or even months. grub. com- — . which includes lowed here is all the other orders of insects. It eats almost constantly since this is the an insect's life history. Siphonaptera. or caterpillar. The characters most used for the separation of the orders of insects are the structures of the wings and mouth parts. After molting several The pupa to rest and prepares for the pupal stage. live — CLASS life INSECTA 309 an aquatic which necessitates changes in the adult aerial ex- and physiological adjustments not required In these orders there are greater changes during developin ment than are found gradual metamorphosis. have made possible the development of this dominant group. in which the larva hatched from the egg bears no resemblance to the adult. Neuroptera. tera.PHYLUM AKTHROPODA hemimetabola naiads istence. Their food habits. along with other distinctive characteristics mentioned above. the two wingless orders Thysanura and Collembola. or metamorphosis. long antennae. It has been divided into the subclasses: Apterygota. DipThe larva is variously called the maggot. The members of this order have retracted mouth parts. and Hymenoptera. The holometabolous insects include the following orders : Tri- choptera. Since practically all the orders fall into the subclass Pterygota.

) Fig. and beneath stones. Machilis orhitalis Packard. Melanoplus feniur-rubrum DeGeer. : — The following are some of the families and species found in the western United States: Family Lepismidae. It is about half an inch long ajid has three bristles extending from the tip of the abdomen. C. and loose and One of the commonest species of this order is the "tish bark. .. the paste of wall paper. Haldeman's locust. warm climates. two striped Mermiria. dry leaves. female coralUpes Hald. male B. 310 moil in TEXTBOOK OF ZOOLOGY They feed largely upon dead plant tissue. Mermiria hivittata Serville E. and starched clothes. The silverfish moth. Conocephalus vicinus Morse. Hippiscus Schistocerca shoshone Thomas. " which attacks book bindings. Utah Agricultural hopper. . Experiment Station. Red-legged 177. female. A. D. Lepisma saccharina Linn. Grasshoppers common to western United States. moth" or " silverfish. Machilidae. western meadow grass(From Henderson. Slioshone grasshopper. are found in the soil i locust. .

Japyx huhhardi Cook and Evalljapyx sonoranus Silvestri. abdomen with six segments.PHYLUM ARTHROPODA — CLASS . Campodea folsomi Silvestri and Entry chocampa wilsoni Silvestri.) with retracted Order Collenihola. 177. . Fig. — Small . aud Japygidae. which often carries three ap- . — (Continued. insects mandibulate mouth parts simple eyes antennae with four segments in most genera. INSECTA 311 Campodeidae.

) Henderson. 1. Tull. Sniinthiirus niger (Lubbock) S. : . 179.) (From Fig. Achorutes species follow Family Entomobryidae Folsomides decemoculatus maturits Fols. pendages modified for jumping. About one thousand species have been described. . permission cricket. totJiorax incestoides Mills. per- JtlilLs . titusi Fols and Tomocerus vulgaris Family Sminthuridae. of Utah Agricultural Experiment Station. All the springtales listed above are found . female. — Sand mission of Utah Agricultural Experiment Station. . eisenii . .312 TEXTBOOK OF ZOOLOGY Tlie body is often cylindrical. Anabrus simplex Haldeman. Isoioma dongaia Mac G. . . (From cricket. Mega. Schott and Papirius maculosus Schott and Family Neelidae. Fig. some of the common Family Poduridae. —Western or Mormon Henderson. Stenopelmatus fasciatus Thomas. 178. Podura aquatica L.

— Praying mantis. however. The mouth parts are of the biting type. Members of this order usually po-ssess two pairs of wings some species. and some do not possess wings at all. cockroaches. have their wings greatly modified or reduced.) to Utah (From Knowlton. katy- and walking sticks. tender plant tissue. Subclass Pterygota. The paurometabolous orders. — Paurometabola following are are insects with gradual metamorphosis. Mills reports that the Collembola are of some economic importance. damaging. in the main. riglit). 181. Agricultural Experiment Station. Stagmomantis sp. Order Orthoptera. . . and Oriental Cockroaches —American German (ventralofview). It is named for its pose. crickets. —This order contains the grasshoppers. The legs are highly developed for use in getting and holding food and dids. some of the more important Fig ISO. permission (left Fig.PHYLUM ARTHROPODA in — CLASS INSECTA 313 many of the states of western America. mantids.

— (Legend on opposite page.) . 182.Fig.

Hippiscus comllipes Hald. 5.) the Shoshone grasshopper.) the German roach. Melanoplus f emur-nibrum Haldeman's locust. Blatta orientalis L. 177) (DeG. . The Mormon cricket. conicaudus Hubbell. Hubbell in his classical study of cave crickets and camel crickets reports more than eighty species. camel crickets. native species of the western Some common Hemiptera. tatus Say 2. — . The katydids. Scudderia furcata Brunner and Microcentrum retinerve (Burmeister) are widely distributed in the western states. About seventeen thousand is species have been described. Tryxalinae. The family Blattidae is represented by such common species as the American cockroach. is also a very interesting member of this family.. 178). (Fig. Blatella germanica (L. (From Knowlton. one of the most destructive insects found in this order. adult box-elder bug. and sand crickets. Locustinae. Analjrus simplex Hald. Giniex lectulanus L. The family Tettigonidae consists of the katydids. female bedbug. generally present. Mermiria hivittata (Serville) . 6. a and Orenivaga erratica (Rehn). agassizi Scudder. first. are of considerable economic importance and are widely distributed throughout the western states. Conocephalus vicinus Morse. the oriental cockroach. family is divided into three subfamilies. had the sea gulls not erected on the temple grounds in Salt Lake City (Fig. the Locustidae. United States (Fig. Leptocoris trivitFig. and Oedipodinae. The following species Ceutliophilus utahensis Thomas. and C. adult Nnhis ferns L. nymph or immature false chmch bug i. meadow grasshopper. —CLASS INSECTA 315 The abdomen is and an ovipositor usually provided with jointed eerei. . C. PHYLUM ARTHROPODA for jumping.) . The sand cricket. . 1. the species of this family are widely distributed and are of economic imThe portance. . It has attracted much attention since the Mormon pioneer days of 1848 when it overran the fields of the pioneers and would have deis stroyed devoured them in great quantities and so reduced their numbers that the growing grain was saved. nodidosus Brunner are fairly common. 182. A sea gull monument commemorating this event has been all the grain crops. permission Utah Agricultural Experiment Station. or grasshoppers. Some of the common families and species found in many of the western states are. The red-legged locust. adult false chinch bug S. 180). doing great damage to crops and forage plants. Schist ocerca sliosJione (Thomas) the twoand the western striped Mermiria." Stenopelmatus fasciatus (Thomas). C. Periplaneta americana (L. cave crickets. male bedbug. or "child of the earth. .) .

.316 TEXTBOOK OF 2oOLOGY Fig. .. Some common Homoptera. 9. pygidium. purple scale. San Jose scale. Pulvinaria vitis L. . Pseudococcus maritimus Elirli. ventral view of body i. pine scale. adult beet leafhiopper S. wingless female black cherry aphid 6. — . . . pygidium 8. . Baker's mealybug. cottony maple scale. . ventral view of body 5. i. Chionaspis pinifoliae Fitch. Spring.migrant of rosy apple aphid 2.. lateral view of adult female potato psyllid 7. Lepidosaphes becki Newman. 183. Aspidiotus perniciosus Comst.

adult of one of the varieties 01 grape leafhopper common in Utah.) . adult wingless female. . Erythronewa siczac Walsh 12. 11. 5. courtesy Utah Academy of Sciences. (Nos. 6. 7. 8. beet aphid.PHYLUM ARTHROPODA — CLASS INSECTA 317 /€iSV /2. 12 from Knowlton. 1. adult male of Emvoasca filamenta De L. 10. 4. 9 from Jorgensen. 10. pygidium. 2. Arts and Letters. 3. 11. Nos. permission Utah Agricultural Experiment Station.

Pemphigus hetae the black cherry aphid.318 TEXTBOOK OF ZOOLOGY species of Pliasmiclae is A common cler) is Parabacillus coloradus (Scud- and of the Mantidae Litanseutria ohscura Scudder (Fig. constitute this order. . Order Homoptera. piercing and sucking mouths the winged species with the front wings leathery and hard near the base and Over twenty thousand species of widely distributed. Leptocoris trivittatus (Schilling). is an example of a bloodsucking species which is world-wide in distribution (Fig. belonging to the family Aphididae. Parafriozoa cockerelli (Sulc). . species. The members of the family Miridae are numerous and widely striders. Nysius ericae common species of the family Lygaeidae. The aphids. the San Jose scale. are probably one of a half dozen species of insects known by all. bugs have been described. Many of the most serious insect pests belong Insects to this order. Pulvinaria vitis L.) typical of the family Nabidae.) is one of the The tarnished plant bug. the back swimmers. and the harlequin bug. and scale insects. .) and the family Corizidae by the box-elder bug. 182). water bug-s. leaf hoppers. Aspidiotus control. is a (Say). Notonectidae and the giant . with membranous wings and sucking mouths. also some species that are beneficial to man. stink bugs. Lygus pratensis commonest mirids in the United States. is The lace bug. They are Family Pentatomidae. 181). — This order includes the true bugs. . such as the cicadas. the cottony maple scale. ferus (L. Aphis roseus (Baker) tive apple aphids in the is — one of the most common and destruc- West. Belostomatidae are familiar to of streams and ponds. are destrucDoane. The family Coccidae is a small of the very obscure group of insects. all who are acquainted The bedbugs belong to the family with the life Cimicidae. The water Gerridae. Chlorochroa sayi Stal. Myzus cerasi (Fab. Anasa tristis (DeG. Pseudococcus maritimus Ehrh. yet they are very destructive and hard to Baker 's mealy bug. The false chinch bug. The beet root aphid. Order Eemiptera. is one tive species. insects with . Murgantia histrionica (Hahn) are common The family Coreidae is represented by the squash bug. plant lice or aphids. and Cimex lecUdarius L. The following represent some of the common families and species: membranous over the outer half. (L. and many are of considerable economic importance. the common damsel bug NaUs distributed. destructive Chermidae. CorytJiucha distincta Osborn and Drake a handsome Tingiis tidae. The rosy apple aphid. and The potato psyllid.).

-Mayflies and dragonflies.'! PHYLUM ARTHROPODA —CLASS INSECTA 319 Sympe^ 1. of 'little ^^'^]^:Cvrr^''VZ'%hvistensen (From Needham and Chnstensen. Kennedy) tn liberate her efcgs . Nymph of Rubicund f^^gonfly Fig." Pteronarcrjs sp. nymph of Lestes uncatus. A side view showing n'a>fl>. 2. on abdomen of smaller "prickleback" 4. 7 permission Utah Agricultural Experiment Station.) gni'^%L™fcciden^aZis/6f nymph .. H. prickleback of the trum ntbicundultim. nymph spines on back. the "trailer ^P^t^Jil'I'ern bunch? wesiern "uncii 5. dorsal view. nymph ot (drawing by C. 3. adult Of the "big curler. ^P n^f Ephemerella grandis. ^. i84. .

large — The dragonflies and damsel flies are insects with compound eyes. Spongovostox apicedentatus (Caudell) are species commonly found in the western United . Chionaspis pinifoliae Fitch are important scale insect pests. insects. The Cicadellidae or : leaf hoppers are represented by the following insect enemies the sugar-beet leaf hopper. Order Isoptera. and the grape leaf hopper. four membranous The wings that are finely veined. —-The earwigs are small terrestrial. The winged species have a short leathery anterior pair of wings which resemble the elytra of some beetles. the purple scale. TEXTBOOK OF ZOOLOGY . Erythroneura comes (Say). plant feeders.320 perniciosus Comst. Hemimetabolous Insects With Incomplete Metamorphosis Order Odonata. 183). not more than They are mostly plant feeders. soft-bodied. mandibulate appendages at the tip of the ab- insects with a pair of forcepslike domen. Empoasca filomenta DeL. often wrongly called white ants.) and the toothed earwig. and a long slender abdomen. sucking the juices from the plants. — Thrips are very small two to three millimeters in length. Iteticulitermes Jiesperus Banks. Thrips idbaci Lindeman are common insect pests. termite. The Nevada termite Termopsis nevadensis (Hagen) . The banded thrip. modinaiads are aquatic and possess a labium which has been highly fied. States. are common dis- More will be said of these insects under the cussion of social insects. (Baker) are . Termites are white. Lepidosaphes becki (Newman) . later in this chapter. The small earwig. About five hundred species of thrips are known.) and the onion thrip. and the western and destructive. Eutettix tenellus . Order Thysanoptera. They feed principally upon wood. It can be greatly extended for the catching and holding of . and the pine scale. — and in the tropics they are one of the most destructive insects known. More than five hundred species of termites. Aeolothrips fasciatus (L. Order Dermaptera. Labia minor (L. have been named. forming large colonies which are used for years and contain as many as five hundred thousand to a million individuals. and they are very numerous over sixteen thou- sand species have been described (Fig. mandibulate mouth parts. all Insects of this order . Termites are social in habits. Delong's leaf hopper. mandibulate insects.

the mouth parts are vestigial." as they Order Trichoptera — . four They have mandibulate mouth wings that are not so thickly netted with veins as are They the Odonata. flies). HOLOMETABOLOUS INSECTS WiTH COMPLETE METAMORPHOSIS The following are some group. Order Plecoptera (Stone flies). gnats. They serve as food for larger insects and fishes. and many other kinds The order is divided into the suborders. Their food consists of mosquitoes. The Mayflies are aquatic insects. Ephemerella grandis Eaton. The life cycle occupies from one to three years. such as May flies. parts.: PHYLUM ARTHROPODA prey. Much has been written on the dragonflies of the United States. This order includes about eighteen hundred species of "case flies" or "rock rollers. are representative species. of the important orders that fall within this (Caddis flies). They require running water that is well aerated. The stone flies are found near — — streams. and the stalked-winged. There are four families the little curler. but with longer antennae than the Odouata. . Pteronarcella hadia Hagen and Perla modesta Banks . Siphlurus occidentalis Eaton are common species. with mandibulate naiads. The prickleback. The adults have well-developed wings and two or three long abdominal cerci. brightly colored. dragonflies Lihellula pulchella Drury and Sympetrum ruMcund^dum United States (Fig. —CLASS INSECTA predaceous 321 in- The adults are swift flying. spot. and the western bunchgill. Hetaerina americana Fabr. Lestes The uncatus Kirby are damsel flies that are widely distributed. Order Ephemerida. scribed. Their food consists largely of aquatic insects. sects. Aeschinidae and Libellulidae. but since the adult stage lasts but a day. damsel flies. flying low over the water. the Agrionidae and CaenagTionidae also two families The beautiful ruby of dragonflies. where they pass their are found on stones along lakes and streams naiad stage. Zygoptera (damsel There are two families of and Anisoptera (dragonflies). are common west of the Mississippi River in the 184). They are sometimes used as bait for trout. About twenty-eight hundred species have been deof flying insects. The food of the naiads consists of small aquatic plants and organic matter which is obtained from the rocks and mud on the bottom of streams and along the shores of lakes where they live.

and Order Coleoptera (Beetles).— Larva (From Needham of net-making caddis worm. . but with vestigial ably take no food. permission of Utah Agricultural Experiment Station. and Platyphylax Pupation takes place in the water. and The larvae feed upon plant tissue and small animals which plants. The lacewing or golden beneficial and widespread eyes.) and Order Neuroptera (Nerve Winged Insects). They have biting mouth parts. calif ornica Coquillett. scalaris nized. they capture in case. designata (Walker) of the family Limnophilidae are common in the western states. and the first pair of wings. snake flies Chrysopidae. the elytra. lacewing flies and the Myrmeleonidae. all the species. have biting mouths and two pairs of net-veined membranous wings. and in variety the water.322 are sometimes called. . are leathery or hard. Cliristensen. lacewings. on land. dobson flies and mantispids. It feeds in the larval stage upon aphids. little nets that are placed near the entrance to their The adults lay their eggs About eighteen families are recogThe species Hydropsyche partita Banks and H. Hydropsyche. however. ally encountered. Fig. is a species. in the water on sticks or stones. doodlebugs or ant lions contain insects most gener- upon other insects. well-developed wings. with mouth parts since they probThe larvae inhabit the bottoms of lakes. shells. This order contains the doodlebugs. They feed on all possible kinds of — . It is — probably the most heterogeneous order of insects. but probably the families Raphidiidae. rivers. and as a means of protecting their soft bodies they build cases or tubes of small rocks. bits of wood. 185. Chrysopa psyllids. snake flies. and feed mainly There are thirteen families. ponds. scale insects. living on plants and animals. TEXTBOOK OF ZOOLOGY The adults are less than an inch long. Hagen of the net-making family Hydropsychidae. The beetles are world-wide in their distribution and contain the largest number of species of any order They are adapted for an almost unlimited in the animal kingdom. The larvae are both terrestrial and aquatic. and creeks. of conditions. thrips.

spotted B.PHYLUM ARTHROPODA — CLASS INSECTA 323 c. common blister beetle. permission Utah Agricultural Ex- . Epicmita jmnctiblister beetle. A. periment Station.) . Three species of Coleoptera. LeptvnoFig. (From Knowlton and Sorenson. adult Colorado potato beetle. 186. Epxcauta maculata Say . — collis Mann. . C. larva or slug of Colorado potato beetle tarsa decimlineata Say D.

Silphidae. larva Alfalfa weevil. In the United States. Curculionidae. Many Some beetles. weevils. Phytonomus porticus. north of Mexico there are one hundred and nine families and Over 200. are. ground and the carrion Anthonomus grandis. Rhynchites hicolor var. lower left. The following families beetles. of the families are the leaf beetles. Other groups of weevils of which the following are typical do considerable damage the billbugs. Calendra mormon Chitt. Elateridae.) . lower right adult. have done millions of dollars' worth of damage.324 food. pupa. cockerelU Pierce and Apion pro: . while in some of the most beneficial insects are beetles. 188). and Cicindelidae . Chrysomelidae . very beneficial: the tiger Carabidae . . permission Utah Agricultural Experiment Station. TEXTBOOK OF ZOOLOGY species do an enormous amount of damage. Coccinellidae beetles. Above. ladybird beetles. the June . which contain the most destructive species the long-horned wood-boring Cerambycidae the click beetles. — and the main. Buprestidae. the metallic wood-boring beetles. the alfalfa weevil. Scarabaeidae . beetles. dive Lee. from all parts of the world have been contrast. . (Fig. Fig. Phytonomus posticus. described.000 species twenty-four thousand species recognized. (From Knowlton and Sorensen. in the . beetles. 187. The cotton boll weevil.

The antennae are shapes and sizes. cockerlli Pierce. . ings by Tanner. the rose weevil.) . of which about eight thousand is the second largest order of insects.PHYLUM ARTHROPODA — CLASS INSECTA 325 Order Lepidoptera (The Butterflies and Moths). 5. 188. and garden. also. the moths. orchard. field. species. stored food and animal products. Rhynchites bicolor var. the bill-bug. which are highly colored in many Common weevils. and Heterocera. while the adults highly specialized suctorial structure. The order is divided into the sub- orders Rhopalocera. tera the larvae have biting —In the Lepidophave a of various mouth parts. butterflies. . Apion Fi^. The two pairs of wings are covered with scales. Calendra mormon Chitt 2. are found in the United States. The larvae or caterpillars are among our most destructive insect They attack the foliage and fruit of the forest. 1. pests. (Drawproclive Lee. — This five Approximately ninetythousand species are recognized.

— Representative of order Lepidoptera. Above.) . (From Knowlton berry leaf roller. Heliothis obsoleta. adult tomato fruitworm moth. and R. Arts and Letters. Sorensen and Knowlton. 190. courtesy of Utah Academy of Sciences. larva of Capitophorus potentillae (Walker) right. strawFig.) .326 TEXTBOOK OF ZOOLOGY At left. fragariae (W. — . Ancylis comptana var.) and Smith. 189. (From corn-ear worm) below. permission Utah Agricultural Experiment Station. tomato fruitworm (or Fig.

Above. cabbage. Some moths. cocoon and empty pupal case borer. and at times alfalfa (Fig. Danails menippe (Hubner).PHYLUM ARTHROPODA — CLASS INSECTA 327 The following are some examples of common species: the monarch butterfly. adult female moth of peach below. and south into the This species is typical of the family Danaidae which is one tropics. strawberries. of the nine families of butterflies in this country. adult exitiosa. the green bolls of cotton. Insects tree borer. parts of Canada. of the most destructive species of this order are (millers) is among the The Noctuidae The corn-ear worm (Fabr. a large family of injurious cotton bollworm. a few of which are tomatoes. Fig.) . corn.). The gooseberry fruitworm. permission Utah AgriStation. 191. (Pi-om Sorensen and Knowlton. is widely distributed through the United States. squash. Heliothis ohsoleta or . species. feeds upon many plants. Aegeria male moth of peach cultural Experiment — of the order Lepidoeptera. Zophodiu grossulariae The clearRiley. 190). center. is a pest belonging to the snout moths or Pyralididae.

a rather distinctive family. —Life historylished by Thebutterfly. pub- leaf roller. blackberries. family it feeds on both wild raspberries and is is an imported species from Europe. are represented by the peach-tree worm. 192. . V. General Biology. of monarch Mosby Company. and cultivated strawberries.328 TEXTBOOK OF ZOOLOGY wing moths. Eucosmidae. The strawberry Fig. Aegeriidae. (Prom White. Aegeria exitiosa Say. a serious enemy of the peach in most parts of the United States (Fig.) C. and found in many parts of the United States. 191).

(slightly magnified) 2. 1. —The Diptera may be char/ acterized as insects with mouth parts specialized for sucking. mosquito larva or wiggler and Community Health. adult (From Turner. Life history of the mosquito. about four hundred wings. Personal . . Mosby Company. 193. laterally flattened bodies. in some species for piercing. species have been described. J.PHYLUM ARTHROPODA — CLASS INSECTA 329 Order Siphonaptera (Fleas). mosquito eggs floating S. V. All of the species in the adult stage They are are external parasites on warm-blooded vertebrates.) Order Dipt era (Flies and Mosquitoes).— Fleas have strong jumping legs. piercing and sucking mouth parts. — m . after Turner and Collins. mosquito pupa or tumbler. published by The C. but no They are world-wide in distribution.. pests on cats and dogs and known to be carriers of bubonic plague. the halters or second pair being vestigial structures. © m^^^- the water Fig. and with only two wings.

A. C. of the (From Knowlton.inn. Syrphidae.— Life history of the housefly. B. Muscidae. by permission of the Utah Agricultural Experiment Station. A. Rowe. Hippoboscidae. The robber flies..) .) Fig. Some larvae are parasitic. or seaven- . others predacious. maand the fruit Trypetidae. eggs.lophaous ovinus L. Utah Agricultural Experiment Station. Tachinidae. contain many species that are Many valuable to mankind. and the taehinids. D. the botflies. pupa inside puparium Madsen. the syrphids. 195. ture larva. The larvae of some families are called maggots. by permission —Adult female sheep tick. Culicidae Fig. the bee flies. On the other hand. Bombyliidae. spread disease and suffering. Asilidae. Rowe. and the sheep tick. adult. damage food and flies. Me. 194. Musca domestica L. 330 TEXTBOOK OF ZOOLOGY of the most useful insects are found in this order. Oestridae. and Madsen. (From Knowlton. the mosquitoes. the houseflies.

sand of which are known to occur in the United States. root aphids. by permission periment Station. Utah Agricultural Ex- parts are both biting and sucking. S. birds. and the females are provided with ovipositors that have become greatly modified.western green-headedMeig. 196. mon In the ichneucomposed of long slender bristlelike structures. causing con- siderable damage. In the winged species there are two pairs of wings. flies. Rowe. the ovipositor is mutillids. (From Knowlton. The mouth are so — Fig. The sheep tick is a fairly common species. Above. horsefly. the second pair being smaller than the first pair. and bees use their The pigeon ovipositors for stinging as well as for horntails bore into trees. "Wasps. and Madsen. and mammals. Order Hymenoptera (Bees. . The ants.) Chloropisca glabra —Below. which are used for drilling through the bark of trees and depositing their eggs upon insect larvae under the bark. depositing eggs. Its of the maggots feed upon beet Tabanits phaenops O. ten thougers.FHYLUM ARTHROPODA — CLASS INSECTA 331 There are over fifty thousand species of Diptera. and Ants). The Hymenoptera named because of their membranous wings. the word hymen means membrane. The suborder Pupipara is a most interesting group. adult Flies. containing the bloodsucking ectoparasites which live upon bats.

(From Knowlton. our garden. Pipunculus subvirescens Loew. The . and forest vegetation. Below. and Letters. thousands of individuals living in a single colony. ichneumon group of parasites. since they feed upon the leaves of Station.) Rowe and Madsen. adult female Simulium vittatum Zett. —Above. and chal- cid flies are examples of this A number of the Hymenoptera are not beneficial.332 TEXTBOOK OF ZOOLOGY of the ! of great value flies. attacking a wide variety of plants. orchard. Arts. Many in biological Hymenoptera live as parasites and are control work. There are many species that are gall makers. Many species are highly developed as far as social organization is concerned. (From courtesy of Utah Academy of Sciences.) Fig 197. female big-headed fly. The braconids. by permission of the Utah Agricultural Experiment Knowlton.

These are in the main. the true lice. Other Orders Other orders than the ones discussed above are included in the notable treatises on entomology. and social wasps are examples. (Prom Sorensen and Knowlton. commonly called scorpion a group of about forty American Brues and flies. the Strepsiptera. a group of small twisted. twentyeight families and about twelve thousand species. . Professor Comstock in his An Introduction to Entomology. a small group of about seventy species found in the warmer parts of the world.winged insects that live as parasites and the Mecoptera. the Embiidina. Vespula pennsylvamca Saussure. insects resembling termites in many respects.PHYLUM ARTHROPODA ants. The common wasp. living under stones and in the detritus of the soil. Melander in their Classification of Insects recognize thirty-four or- . — CLASS INSECTA 333 honey bees. wingless ectoparasites of birds. the lice. an order consisting of sixty-five species of blood-sucking parasites found on the mammals. in addition to the eighteen orders discussed above. The honey bees and silkworms are the only really domesticated insects. Corrodentia. the Anoplura. within the body of other insects species. The Hymenoptera found in this country are divided into three suborders. or yellow-jacket. rare and little known insects. 198. however. recognizes twenty-five orders the Zoraptera. by permission of the Utah Agricultural Experiment — Station. psocids and book the Mallophaga. and consisting of but : Fig-.) six known species in a single genus Zorotypus.

F. Alfalfa-seed chalcis-fly.334 TEXTBOOK OF ZOOLOGY ders. the noted English entomologist. eggs (greatly enlarged) H. has included twenty-three orders in his A General Textbook of Entomology. pupa. larva. 199. Brucophagus funetris How. B. permission Utah Agricultural Experiblack ant. G. worker of the anterior view of right mandible. Fig. (enlarged) (From Sorensen and Knowlton. male antenna. —Hymenoptera. C. . . A. female antenna. female. ment Station. while Imms. with- relationship existing between parents The processes that have ever been operative have emphasized the importance of the individual in the scheme of prog- and offspring. D. . E. In this elementary consideration of insect classification we have tried to include information and illustrations which will be of value in interesting the student in the thousands of insects of our environ- ment.) SOCIAL LIFE The great majority of out any cooperation or filial AMONG THE INSECTS insects live an individual existence.

The following excerpt from a study of the nesting habits of Odynerus dorsalis Fabr. in colony life. ground which provisioned and then an egg is sealed in the No other atten- tion is given to the developing young and the new generation never knows the old. up small spheres of fresh cow manure. In the Hymenoptera are found varying stages of social the solitary wasps. '' Odynerus dorsalis is a solitary wasp in that each female builds a separate nest. the female digs a In is burrow in the cell. Edwin Vest gives a good picture of the activities of this solitary wasp. in the family Passalidae. — CLASS INSECTA 335 Despite this. neath the letting it gradually into a hole in the ground. rolls A common down species. pointed out that at least twenty-four different times communism or societies have appeared in the class Arthropoda. Canthon sim- plex var. yet there are often several nesting individuals in the same vicinity forming a kind of community. and cover over the sphere of manure upon which the female has deposited an egg. The male helps to dig No The beetles are probably the least social of all the orders listed. The developing larvae feed upon wood that has been prepared by the adult beetles. Another beetle. and in the Dermaptera. The French naturalist and entomologist. In the beetle family. Fabre. Let us look at some of the ways social life has manifested itself. which the writer has ofttimes observed. Scarabaeidae. made by Mr. The labor of dig- . and then excavates beroll. fifteen families of Hymenoptera. Each species of beetle grows a species of fungus which is fed to the developing larvae by the adult beetles. and the males take but little part life. J. Wheeler. castes have been developed. the great authority on insect societies. The colony is kept together by audible noises made by the mature beetles.PHYLUM ARTHROPODA ress. lives in rotten logs. The ambrosia beetles of the families Scolytidae and Platypodidae form colonies by making their burrows into the wood of both living and dead trees. Embiidina. corvinus Harold. reported many interesting observations relating to the preparation of manure pellets for the deposition of eggs by several different scarabaeids. H. He reports that social life occurs in six families of Coleoptera. and Isoptera. Passalus cornutus. we find a number of species in which there is a cooperation between the male and female for the perpetuation of their offspring.

They often alight on a female as she is working about the nest or returning to the nest with food and knock her to the ground. The bottom of the hole is enlarged The slightly into a cell and is made very smooth on the inside. cell might be lined with a secretion from the body which forms a cementlike protection to the larva during the winter. Desiring to learn how Odynerus handled the larvae before putting them in the nest. In general they are ovoid-elliptical in shape. and composed principally of clay. by grasping them with her mandibles just back of the head and supporting them somewhat with her two front legs. another male attacked with such vigor that the female flew other male in pursuit. away with still an- "The ground where the nests are made is hard. "Each The wasp cell is provisioned with from five to twelve Pieridae larvae. in diameter to 6. In order to penetrate it the female fills a thin pouchlike sac. After the nesting is begun the females spend the night in the burrows with the head uppermost. the depth varying from 48 to 110 mm. They fly rapidly back and forth over the community usually from six to eight inches above the ground.336 TEXTBOOK OF ZOOLOGY ging the hole for the nest and gathering the provisions is apparently done entirely by the female. Favorable results were obtained in two cases. with water and uses this to moisten the ground. "Their attempts at copulation are very amusing as well as inBeginning about one or two o'clock in the afternoon the males become very active. located within the second segment of the abdomen. When the wasp found the worm she applied her mandibles to various places on the body but spent most of her time biting just back of the head as if carries these larvae . are usually one or two. distance away from the hole.8 mm. cells constructed in the tunnel for the deposition of eggs. dry. while the males roost upon nearby herbs or shrubs. The average size of the cells is 23 by 14 mm. This work is continued until the hole There is as deep as desired. teresting. the writer attempted to induce several wasps to pick up worms that were dropped on the ground about the nests. rarely three. At no time was the male seen to engage in any part of this work. One female was resting on the ground when a male flew down and alighted on her back as if attempting to copulate. With her mandibles she digs the dirt out in small pellets. varying in size from These pellets are carried a short 2 mm.

in length with the point of attachment to the cell wall concave and about 2 mm. During the warm weather they were kept moistened by placing a few drops of water on blotting paper covering the cells. primarily to two-celled nests. while with the two-celled nests the larva in the lower cell always developed into a female and the upper individual into a male. in diameter. The above condition applies it might be equally true of evident from this study that the eggs laid in July and August hatch and remain in a late larval instar throughout the winter.PHYLUM ARTHROPODA trying to cut it off. that the male develops more rapidly than the female. On August 2 a number of larvae were collected and placed "It is in glass vials. After that time they began to decompose rapidly. About the middle of September they were placed in a north room of the writer's home where they were left throughout the . It was noted that the wasp in the lower cell did not emerge until three days after the top cell had been vacated. although the three-celled types. with single-celled nests ranking second and three-celled nests third in frequency. — CLASS INSECTA 337 This is of paralyzing the victim.8 mm. since the egg in the lower cell is laid before that in the upper cell. No successful observations were made on the The facts of the case would seem to indicate three-celled nests. Of those individuals successfully reared in the laboratory it was found that in the case of the one-celled nests the individual invariably developed into a female. Only one egg is deposited in each cell. In order to observe the writer used a small pocket mirror to reflect the this process. This did not seem to interfere with the apparently air- activity of the wasp. light down into the hole. The writer was not successful in hatching out all the "Most individuals of any three-celled nest dug from the ground but those containing one or two cells were often hatched successfully. The cell is then sealed over by wetting the soil at the surface and then carrying it down to be moulded into an and water-tight compartment. "After the cell is provisioned with the Pieridae larvae the female attaches the egg to the upper part of the cell by a short hairlike process 1. probably a part at least of the process These paralyzed Pieridae larvae have been kept in the laboratory in bottles for two weeks in warm weather before there began to be any change in their appearance. of the nests observed in this study consisted of two cells.

One female was kept in a breeding cage and fed on a syrup of cane sugar and times going slightly below the freezing point of water. workers. the queen feeds the larvae until they are completely deThese workers then come to the aid of the exhausted veloped. distilled water. After the nests are built and the eggs begin hatching. It seems evident that "It is somewhat these early wasps build their nests in the spring and that young emerge during the same season. The adults emerged fully developed about the middle of July. thought that under natural conditions the insects emerge earlier in the summer than was indicated by the artificially reared specimens since they have been observed to be very active even during the early part of May. Only the individuals their nest- ing in the late summer spend the winter in the larval stage. to carry on the life cycle. Polistes. The queen's only duty now It will be noted that the Vespidae attend their young is to lay eggs. The social wasps. the queen. The workers assist in building and feeding the colony. and males die. the bumblebees also start a colony in the spring by overwintering queens seeking out an unoccupied mouse The queen gathers hole or some other suitable hole in the ground. which he chooses to call "trophallaxis. by gathering food and feeding them. belonging in the genera Vespula. the temperature some- About the last of May the specimens were removed to the Brigham Young University where they were kept on the writer's desk. All this is very similar to the life habits of the social wasps. also that in turn the adults may feed upon the saliva of the larvae. The room in which they were kept was cold. She then deposits an egg in each cell and waits for them to hatch and develop into workers. Some of the larvae spun their cocoons in the vials while others had already done this before being removed from the ground. start new colonies each spring from overwintering queens. Wheeler believes that the exchange of food in many of the social insects." and Polyhia. pollen and nectar with which she fills a few cells. which developed late in the summer and which hibernate. . When the winter comes on.338 TEXTBOOK OF ZOOLOGY winter. In the family Bremidae." has been the source of the social habit. of the family Vespidae. leaving only the females. founder of the colony by taking over the enlarging of the nest and the feeding of the larvae and the queen.

There is also an interesting symbolic relationship existing between numerous intestinal protozoa and the termites. a worker caste. Ants and termites have many different forms of individuals in each species. males and females are found in the various castes of termites. these guests are called they are never found outside of the colonies. ants. caste system. a less fully developed sexual caste. and workers. social life is carried to its is highest state of perfection. and finally a caste known as nasuti. In a termite colony there are many The principal kinds are perfect males and females. Wheeler reports that fully two thousand species of myrmecophiles and one thousand termitophiles have been described. wingless individuals. building up large colonies consisting of Other queens take up the job fifty to eighty thousand individuals. also polymorphism. of continuing the colony. A well-developed social insects. The Formicidae have a highly developed caste system and usually the workers and at times the males and queens are polymorphic. Wheeler believed that ants are the most highly developed as well as the dominant group of social insects. is found in these In a swarm of bees there are three kinds of individThe workers are females that are uals. tive tracts. or the royal stock. Many of the guests have become so dependent upon living with ants or termites that social insects. since about four thousand species are known today. undeveloped sexually. morphologically and which are small Both individuals with the head produced into a kind of snout. The wood eaten by the termites is made soluble by the infusoria found in their digesdistinct from other individuals because of their large heads strong jaws. sterile. perpetuated for hundreds of years. Aphids and mealy . and termitophiles when with the termites. Ants are world-wide in their distribution they are also very numerous as individuals and species. with rudimentary wings. In these groups the colony probably Some ant and termite queens live from ten to fifteen years.PHYLUM ARTHROPODA In the honey bees. males. castes. of fairly small. females. —CLASS INSECTA 339 and termites. Guests There are many species of insects that live in the nests of the myrmecophiles when found with ants. the fecund pair of the colony. a soldier caste. .

Some Diptera are also guests in ant colonies. thenogenetically. niger var. C. zelus Fall are fairly common in ant nests in the states west of the Rocky Mountains. which are gathered and stored for the winter by the ants. which they greedily feed upon. After mating. Aphis maidi-radids Forbes and the brown ant. are found in ant hills in the United States and Mexico. Reichenl)achia. Many insects. In the spring before the com commences to grow. members of the family Staphylinidae. are found in the colonies of several of A number of species of Batrisodes the mound ants. and KnocJii LeC. S. Dr. The ants are repaid for the care they bestow on the aphids by receiving a honeydew given off by the aphids. are placed upon the roots of smartweed and some of the grasses. A. and The deto and their products valuable to man amounts over a billion dollars annually. ECONOMIC RELATIONS Insects attack all kinds of growing crops struction of plants V plants. Some of the . As soon as the corn has started to grow the agamic female aphids are transHere many generations are produced parferred onto the roots. eggs are laid. but species live as endoparasites or ectoparasites on animals and man. Animals and man suffer greatly from the attacks of insects. americanus Emery. most dreaded diseases known to man are carried by insects. pselaphids. Many atus of the insect guests are beetles. and and Cremastocheilus angularis LeC." This great loss goes on because unabated and persistent struggle of the insects to maintain Plants are not only eaten and damaged by many plant diseases are spread by them. Because of this there has recently developed a new branch of entomology known as "medical entomology." Some of the most notable progress during and in so doing also spread disease. Then in later September or October wingless males and females are produced. The two histerids. and Scarabaeidae. Histeridae. Hetaerms tristri- Horn and H. after hatching. Forbes has reported most interestingly upon the activitiv'^s of the cornroot aphid. Several species of Xenodusa. Lasius in October The little ants gather the aphid eggs and take care of them during the winter. scarabaeids. the aphids.340 TEXTBOOK OF ZOOLOGY bugs are kept as guests for the droplets of lioneydew which they excrete when stroked by the antennae of the symbiont. Staphylinidae. of the their "place in the sun. Pselaphidae.

In India a small scale insect. as well Everyone knows that honey is produced by the honey bee and by the silk moth. tion. granary weevil. the pigment cochineal. bean weevil. The pea and confused flour beetle feed and seeds and their upon and damage products. or store Many insects his upholstered furniture. of dwellings. Much damage is also done to the same products by such species as the Mediterranean flour moth and the Indian meal moth. even furniture Termites also attack the wooden parts and books. Insects belonging to the orders Coleoptera and Lepidoptera are the main offenders. The tobacco beetles and home furniture. tularemia. shellac. clothing. Tachardia lacca Kerr. After man has produced his crops and harvested them he finds many insects ready to take their toll from these trated products. and many drugs.PHYLUM ARTHROPODA — CLASS INSECTA 341 the past thirty or forty years has been made in the field of medical entomology. but there are many people who do not know that certain insects produce shellac. lives on trees and produces a secretion that forms a layer over the branches. and silk beeswax. tannic acid. Many species are as in man in the struggle many other ways. drugstore beetles live upon tobacco products. against the injurious insects. Rocky Mountain spotted fever. This substance. due to clothes moths. Much remains to be done in this new entomological field. Diseases such as malaria." inks and dyes. cantharidin or "Spanish fly. for use. bubonic plague. typhus fever. Great losses are suffered annually by the producers of clothing. practically all kinds of grains Practically all pests of stored foods are world-wide in their distribu- which makes it difficult to ship food products long distances them for future use without running the hazard of insect damage. and elephantiasis are now known to be insect borne. furs. as well as in the homes. living upon and rugs. African sleeping sickness. yellow fever. Useful Insects Fortunately not allies of all insects are our enemies. weevil. have taken up their abode with man. concen- The "board bill'* of the insect pests of stored foods annually amounts to about four times the cost of all higher institutions of learning in this country. formic acid. is removed by the natives in .

seeds. is used for making varnishes and polishes. and mammals. many fishes.342 TEXTBOOK OF ZOOLOGY Shellac various ways. including man. and orna- mental plants to develop. amphibians. . as an electrical insulating material. in airplane construction. birds. many the blossoms. It is important that insects be recogWithout the nized as playing a major role in this connection. Finally. reptiles. insects the food habits of many of the vertebrates would be entirely changed. vegetables. and Insects serve as food for many other ways. plants depend upon insects to assist in pollenizing Only as the insect helps in transferring the pollen from plant plant is it to plant or from the stamens to the pistil of the same possible for some fruits. millions of pounds being sold throughout the world.

in certain regions of the body. between the eyes. Again. Eegardless of the amount of variawe find that the segments are always grouped into three regions. thorax. forming a boxlike structure known as the epicranium. Smith reported the damaging activities of Melanoplus atlanis in the New England states. and along with the beetles and bees. and down the sides of the head to the base of the mouth parts. The head is made up of a number of segments. known as the head. which are fused parts of the segments may be together. In the United States many destructive species are found. very few boys and girls grow up without having some experience with a grasshopper. to the transverse suture. As early as 1743 The locust or grasshopper is known to practically all people. while the front of the head between the eyes is called the frons. or cheeks. They are widely distributed throughout the world.CHAPTER XXII REPRESENTATIVE INSECTS (By Vasco M. or segments. Brigham Young Universitt) THE LOCUST being one of the most common insects. and the depression between the plates is called a suture. The grasshopper is a typical insect. and abdomen. The sides of the epicranium below the compound eyes are known as the genae. The insect body the segments are is divided into a series of rings. to be discussed later in this chapter. lost. The hardness of the plates is due to the deposition of a horny substance called ckitin. gro^\Ti together. living on grass and low-growing plants of the fields and open country. tion in this respect. Tanner. or are fused. may serve to illus- trate the general structure of the class Insecta. This boxlike piece which surrounds the eyes and forms the basis of attachment for the movable parts of the head extends down the front of the head. 343 . because Mr. In many places two or more of these rings have made up as sclerites. and from 1855 to 1877 many outbreaks of grasshoppers were reported in the western United States. and known These plates are of hardened plates. Even today the national government is expending large sums annually to keep down the activities of the many destructive species.

.g. maxillary palpus O. . ver.-.g.) . .e.. .. . tempora.. crest of the metazone c. by permission of the Utah Agricultural Experi. temporal foveola. ant. . a. The external structure of the grasshopper. median carina of the fastigium in. .e. Antennal groove.. Dissosteira spurcata. 200.. .. . Front view of the head of the grasshopper.e.e. lateral carina of the metazone m. . t.-cj. I U i. . gena inan. crest of the prozone g. ocellus. Hind angle of lateral lobe cm. 1.p. compound eye c. labrum carina of the fastigium I. genal groove I.o.e. Fig. L 1J^V' n^. al.f. lateral carina c.p. gena g. 2.— r C.. central foveola e.. vertex..'p. transverse incision.... mandible in.. fastigium of the vertex f. c-. c.e. antenna. ..C..L.. s.p.c.... .. . .i. clypeus c.. ment Station. frontal costa la. . ... . lateral l.. .. (From Henderson. .w A ^' r. central ocellus fas.f. sulcation of the frontal costa.p.e. Dissosteira spurcata..g. labial palpus g. carina of the antennal groove cl. maxillary palpus t.

the dorsal part known as the pronotum and the ventral portion known as the sternum. just beloAV the margin of the impression which contains the bases of the antennae. which is attached by its upper edge to the epicranium. The simple eyes or ocelli consist of One of the ocelli is three small.REPRESENTATIVE INSECTS 345 The grasshopper has both compound and simple eyes. situated on the front of the head. oval areas. Each maxilla is composed of a number of parts. . and the maxillary palpus. The caudal part of the mouth parts is the lower lip or laliium. The first noticeable part the Idhrum. On the front of the head there is a short rectangular piece. a sclerite which bears some teeth on its terminal end. It is made up of a fusion of four plates. . called the clypeus. which is composed of the siibmentum which acts as a hinge on the epicranium above. It may be divided into two regions. Each mandible consists of a single piece which is notched on the inner grinding surface to form a number of ridges or teeth. The antennae or feelers are two threadlike processes situated median to the compound eyes. and two large outer flaps. the surface will be seen to be made up of a number of hexagonal areas. and are large. the lacinia. highly polished surfaces. the maxillae. The compound eyes are situated upon the upper portion of the sides of the head. which are indicated by the transverse sutures. Each consists of about twenty-six segments. 200). The free edge is deeply notched on the median Just beneath the labrum are the mandibles. may be exposed by the removal of the mandibles. The anterior sclerite bears a spine on the median line. oval areas with smooth. or sclerites. and in contact with the upper portion of the compound eye. The prothorax is the segment to which the head is attached. and on the lower edge to the labrum. A second pair of jaws. If the eye is examined with a dissecting microscope. a mentum. edge of the clypeus. The pronotum is a saddle or bonnetlike piece extending over the dorsal and lateral regions of the prothorax. the ligulae (Fig. consisting of the cardo or proximal hinge part of the structure the stipes. or upper lip. The sternum or ventral side of the pronotum is also made up of separate plates. a flaplike piece attached to the lower line. almost transparent. the outer lobe or galea. or first pair of jaws. labial palpi. which are known as facets. The mouth parts is consist of a number of separate parts attached to the ventral region of the epicranium.

j2 c o 5 o So-I ^t^CLj '^ -73 I !-o y a> . . a •? o c> 2i S ^. cq .- ll . o3 -^^ .5 ^ to aoK J^ o r s^oj.J +-> « <D oT « t.. bo S ri o o S o c .^!^ Eh c3 !-i O " aj aa r a'O 3 <p Sg - °t?a.^1 fe 0) .346 TEXTBOOK OF ZOOLOGY aO S-S S <D ra tJ3^ cd o) -^ ^1 .

Like the prothorax these segments are made up of separate plates. boxlike structure for the support of the wing and leg muscles. or dorsal region. is attached to the dorsal region of the mesothorax. The second pair of wings is attached to the metathorax. They are leathery in texture and do not fold fanlike over the abdo- men. however. On the sides of the body. and fold fajilike over The metathoracic wings are used the abdomen when not in use. The femur the third and largest segment of the leg. running sutures. the mesothorax and metathorax. The pleura of each of the posterior thoracic segments are again divided by transverse suture. The next segment. and their structure The mesothorax is joined to the prowill be discussed together. The first pair or wing covers. and in the case of the metathoracic leg contains the muscles used in jumping. be divided into three groups: the terguni. A pair of legs arises from the lateral and ventral portions of each Each leg is composed of five parts. On the dorsal and ventral regions of the body the sutures separat- ing the mesothorax from the metathorax are not very distinct. held together by a tough. The segments up of three segments. or ventral region. There are two pairs of wings. Posteriorly the metathorax is joined immovably with the first abdominal segment. the trochanter. They are strengthened by many veins and cross veins. They are membranous. there is a very distinct line. and the pleuron. These plates may. also called tegmina. is slender. piece which is is hard to distinguish except in the first pair of legs. are made up of sclerites that are intimately associated. with many veins to strengthen them. The coxa is the first segment and is attached to the thorax by a tough elastic membrane. so that each pleuron consists of two sclerites. The last division of the leg is the tarsus suckerlike disc known as the pulvillus. which terminate on the last one in a large as the femur. each movable bear a series of pads.REPRESENTATIVE INSECTS 347 The next two segments. the tibia. is a very short of the segments of the thorax. This suture divides the mesothorax from the metathorax. connecting membrajie. . thorax by a membrane which permits of more or less movement. The mesothorax and metathorax form a strong. however. but about the same length which is made with the other. or lateral region. or from the posterior border of the attachment of the second pair of legs toward the dorsal part of the body. in flight. The fourth seg-ment. the sternum.

is The eleventh segment represented only by the tergum.absent. The plates called cerci and podical plates are similar to those in the male. or near the ventral part of the body. except that the sternum is nearly twice as long. pointed tip. which forms the termi- nal. where side pieces. Attached sternum is the subgenital plate which forms the most posterior ventral plate of the body. This piece partially" surrounds the tympanic membrane. platelike piece. or ear. and known as the and eleventh segments are essensegments nine and ten being partially fused. which terminates just above the attachment of the third pair of legs. eighth segments are joins the sternum. which is a large. The ventral pair arises just dorsal to the eighth sternum and resembles the dorsal . to which extends laterally it The pleura. and extending back. tenth. The seven anterior segments are similar in both the male and female. dorsal shield. It is composed of eleven segments. crescent-shaped area covered with a semitransparent brane. In the female the eighth segment resembles the other segments. The dorsal pair lies just ventral to the eleventh tergum and each plate is long. and with a hard. In the male the first abdominal segment is made up of a curved dorsal shield. the terga of prominent. past the end of the eleventh tergum. The cerci constitute a pair of plates attached to the lateral posterior border of the tenth segment. and tergum eleven forming the terminal. and the genital chamber lies to the ninth The anus opens between these directly below them. owing to the large size of the attachment The pleura are entirelj'. all quite similar. and ventral to the eleventh tergum. have been inseparably fused to the tergum. except that the podical plates are much more subgenital plate. is not attached to the tergum. mem- The ventral part of the first segment. the sternum. tially like those of the male. noted in connection In with the thorax. lance-shaped. forming a broad. much modified. The ovipositor consists of three pairs of movable plates. 201). The ninth. the tergum. dorsal. shield-shaped piece (Fig.348 TEXTBOOK OF ZOOLOGY The last main division of the insect body is the abdomen. consisting of a dorsal tergum. The podical plates lie directly beneath the cerci plates. The second to the of the legs. the ninth and tenth segments the terga are partially fused together. the union of the two being indicated by the presence of a transverse The sterna of these two segments are entirely fused and suture.

This is because there are never more than eight pairs of abdominal spiracles Air passes through the present in any fully developed insect. the blood is forced to the anterior part of the body where it flows out into the body cavity and slowly returns to the abdominal region. body. which extends along the length of the median dorsal part of the body. spiracles into the tracheae and is carried to the tissues of the body. forming a sharp organ by male bores the holes in the ground The third set of plates are known as the egg guides. There are ten pairs of spiracles. but this belongs dorsal to the From the second to the eighth to the first abdominal segment. In the abdomen of the fully developed insect this vessel is divided into a number of chambers with side valves. near the There is another spiracle just attachment of the metathoracic leg. or openings in the respiratory system on the body of the grasshopper. It will be noted that the circulatory system has practically nothing to do with the carrying of oxygen to the tissues. The digestive system straight tube extending of the grasshopper consists of a practically from the mouth to the anus through the . 349 When these four pieces are brought together. The circulator}^ system consists of a single dorsal tube. The mesothoracic spiracle is concealed by the posterior edge of the pronotum. The spiracles are one of the most useful sets of structures for determining the segmentation of an adult insect body. means of which the fewhich to deposit her eggs. The metathoracic spiracle is located just dorsal to the mesothoracic leg. except through the vessel toward the head. These are much smaller and are located median to the plates of the true ovipositor. which has been called the heart. Two pairs of these liplike structures are situated on each side of the thorax on the anterior margin of the pleural plates.REPRESENTATIVE INSECTS pair. which allows the blood to enter but not to escape. In this process the tissues are supplied with nourishment from the food materials carried in the blood. Due to the pulsating of this portion of the tube. their points in are in contact. abdominal segments there is one pair of spiracles located on the anterior margin of each segment near the union of the sternum and tergum. or heart. This unique system of breathing enables the insect to keep the body tissues well aerated and the carbon dioxide eliminated from the suture separating the two segments.

. begins. -A\..C. colon Int.X H'lnt.T. From the mouth the food is conveyed through the esopha- — \-/A 's-. Malpighian tubules G. Sal.. A. salivary glands. . anus. Co. General Biology. invertase. O. The food after being ground up by the mouth parts passes into the mouth or pharynx where it is mixed with the salivary mucin and the action of the enzyme.A 202. . V..) Fig./ 5t. The C. Oe. R. — . mouth M.. crop. rectum. -R. intestine M.. Digestive system of Rhomaelia microptera.e. esophagus. n -to. . .. J.. 350 TEXTBOOK OF ZOOLOGY central portion of the body. gastric caeca (From White. Mosby Co.

. (From White.) . . Tiie C. \— Ab Nervous system of Rhomaelia microptera. supraesophageal ganglion ganglion C. Sp. Mosby Co. The gizzard is muscular and lined with chitinous ridges which strain the coarse particles of food and prevent their entering the next division of the system. General Biology.REPRESENTATIVE INSECTS 351 gus to the crop and gizzard which are dilatations of the tract filling a great portion of the thorax.. They secrete a weak acid which helps in the The food is . which are glandular bodies opening into the anterior end of the stomach. acted upon in the stomach by the secretions of the gastric caeca. . the stomach. circumesophageal commissure subesophageal ganglion. first abdominal Fig 203 Su. Ah.. V.

) . which is divided into the ileum. The rectum has thick muscular walls with six-surface rectal glands. — commissures or connecting nerve fibers lying along the ventral body wall. TEXTBOOK OF ZOOLOGY and the conversion is of albuminoids into pep- absorbed into the hemolymph from the stomach. Five ganglia are located in the abdomen. well-developed ganglia are found in the thorax the anterior one is connected with the subesophageal ganglia which in turn are connected with the brain or supraesophageal ganglia by nerve fibers which pass on set of . Since there are at least eleven segments in the abdomen of the adult grasshopper. Anterior aspect of brain (supraesopliageal ganglia) of Rhomaeha microptera. General Biology. (Magnified. it is apparent that the ganglia of some of the segments have fused together. V. 2. The subesophageal ganglia supply the mouth parts with nerves. showing fibers to ommatidia S.) 1. In the larvae of insects there is usually a ganglion to each segment. Between the stomach and the intestines is a pyloric valve which permits the contents of the system to pass in only one direcIn the intestine. Just back of the stomach many threadlike tubes enter the intestine.352 emulsification of fats tones. absorption of food continues. The nervous system connected by a double consists of a series of ganglia or nerve cells Nerves pass from the brain to the eyes. colon. circumesophageal commissure. antennae. Three large. In the vertebrates the . Fig-. known as Malpighian tulules. in their movements by the thoracic ganglia. especially in the ileum. and perform a similar function to the kidneys of higher animals. 6. The C. These tubes are the excretory organs. 204. nerve to unpaired ocellus. 4 and 5. and palpi of the head. nerve to eye. rectum. nerve to paired ocellus. nerves to mouth parts. 7. (From White. The legs and wings are coordinated each side of the esophagus. The feces are expelled from the rectum to the outside of the body through Much of the food the anus. and tion. Mosby Co. nerve to antenna.

) — — which when mature fill the major portion of the abdomen.. which consists of chitinous styles used in copulation with the female. and the foreshadowing of this evolutionary change is initiated in the insects by the development in the cephalic region (Figs. C.. vagina.. The oviducts convey the eggs to the vagina. O.REPRESENTATIVE INSECTS 353 nervous system is dorsal to the digestive tract. (From White. 205. General Biology. the abdominal structures separating the two sexes are distinctive. which discharges the eggs through the opening at the base of the egg guide to the outside of the body. Fig. V.. The external genital structures have been discussed above. (From White. Y. The sperms are borne in ducts which communicate with the penis.) Fig. The grasshopper is dioecious.S. 206. Female reproductive organs of Rhomaelia microptera. The C. testes. 203 and 204). The female is able to dig a hole in the ground with the ovipositor .T.D-. Fig. 206. The male organs consist of testes located dorsal to the intestines. vas deferens. copulatory sac. oviduct. Ov. which is dorsal to the vagina and which is connected by means of a sperm duet. The eggs are fertilized by the sperms from the spermatheca. In the female there are two ovaries. General Biology. a duct made by the union of the two oviducts. ovarian tube with eggs. Mosby Co. Va. Te. Male reproductive organs of Rhomaelia microptera.

the may be seen in the lateral margins of the dorsal surface of the abdomen. More than one hundred and twenty-five species of these beetles have been reported as occurring in the United States and Canada. The mouth parts are of the biting type. from the frost. are covered with a frothy substance which protects The eggs them from in moisture and. An examination of a specimen of the genus Phyllophaga reveals that there are three body regions: the head. cereal. to some extent. The thorax is provided with The abdomen. which spiracles is broadly fused with the metathorax. retracted head bears antennae of nine or ten joints and a club composed of three elongate leaflike joints. destroying the roots of grain. the largest order of arthropods. Because of the general distribution of these beetles. or elytra. leaving the prothorax free and movable. thorax.354 TEXTBOOK OP ZOOLOGY to the and deposit the eggs depth of an inch or more. The larvae or white grubs live underground. the grasshopper is The eggs are laid The development of by gradual metamorphosis. consists of eight external segments. . The metathorax is fused with the first abdominal segment and with the mesothorax. The thorax consists of three segments. The hind wings are membranous and folded under the elytra. The legs are well developed. which cover and protect the back of the thorax and abdomen. yellow setae. Attached to the dorsal portion of the mesothorax are the fore wings that are modified into horny sheaths. as well as great tracts of pasture and grasslands. and garden crops. truck. the prothoracic ones being adapted for digging in the ground. There are no ocelli. The antennae are located just beneath the lateral edge of the prominent clypeus. The adults live upon the leaves of many kinds of trees and shrubs. When the elytra are removed. often completely defoliating the trees. and abdomen. THE JUNE BUG The June bugs or May beetles are members of the family Scara- baeidae. they have been selected as a type to illustrate the characteristics of Coleoptera. The genital organs of both sexes are simple. The rather small. a very large and important family of beetles. The compound eyes are on the sides of the head near the prothorax. similar to those of the grasshopper. the fall and hatch in the spring of the year. the majority of them being considered as pests.


four years. except for a bend in the colon. and some soil. and rectum. many changes in the digestive passes from the larval stages to the imago. which means that the heart serves mainly as an agitator of the body fluids. At the posterior end of the midintestine and in front of the pyloric valve are ten pairs of pyloric caeca. The larvae are commonly known as ''white The life histories of the June bugs vary in length . The tracheal system There are as it is well developed for carrying the air from the spiracles to all parts of the body. arise in the ileum just posterior to the pyloric valve. a brain. There are eight ganglia in the ventral nerve chain. The blood is forced forward through the heart chambers by the pulsations of the heart walls. which consists of roots. In the adult the excretory organs. Two rows of gastric caeca are present on the anterior end of the midintestines. as it is rarely met with in larval stages of other insects. without the two rows of gastric caeca. There are four Malpighian tubules connected to the hind intestine. depending upon a number of ecological factors. system of the June bug The alimentary tract of the larva consists of a straight tube. The food passes from the mouth or buccal cavity into the esophagus and then into the crop. colon. It is much greater in diameter than in the later stages due to the nature of the food. In the pupal stage the gastric caeca have disappeared. humus. There are no arteries and veins.356 TEXTBOOK OF ZOOLOGY carefully removing the By membranous tergites of the abdomen the heart can be seen to consist of a thin-walled dorsal vessel with paired lateral openings into the body cavity. from three to The adult females dig into the ground and deposit from a hundred to two hundred eggs. or supra-esophageal ganglion. the Malpighian tubules. helping to distribute the absorbed food to the tissues. They extend into the body and then end blindly at the junction of the colon and rectum. a nerve ring which connects the brain and the foremost or infra-esophageal ganglion. The nervous system consists of a ventral nerve chain. four in the thorax and four in the abdomen. At this point there is a valve between the crop or gizzard and the mid-intestines. The hind intestine consists of the ileum. and the tract is becoming much elongated and coiled. The large saclike stomach or mid-intestine of the larva is transformed into an elongated coiled stomach in the adult. This is a very unique feature.

known and the large spinelike structure. as the pollen Irush. The worker bee is provided with large compound eyes on the sides of the head and three small ocelli near the median part of the frons. mesotlwrax. Many of the Hymenoptera. The maxillae and labial palps are used in sucking the nectar from the flowers. On the tibia are the curved bristles. live a social life. modified spines and structures on the last pair of legs known as the The modifications found in the pecten. legs of the bee are remarkable adaptations for the specialized life of this insect. developing colonies consisting of three types of bees: a queen. feeding upon the foliage of many plants. and pollen combs. At the end of the abdomen is a highly specialized organ. the sting. and (3) the maxillae. The abdomen is composed of six external segments consisting of a dorsal tergum and a ventral sternum. and metaThe wings are borne thorax. The lahrum is attached to the lower edge of the chjpeus. which secrete a substance composed of an acid and an alkali. such as the honey bee. . The thorax is divided into the prothorax. lateral to the labial palps.: REPRESENTATIVE INSECTS 357 grubs. and the females usually with a stinging organ. well-developed biting or sucking mouth parts. which is associated with the antenna comb. THE HONEY BEE The honey bee belongs to the order Hymenoptera. and workers. auricle. The metathoracic There are also legs have the tibia modified to form a pollen basket. upon the mesothorax and metathorax. Beneath the mandibles is the proboscis made up of several separate structures (1) the glossa or long tongue. dibles are attached to the ends of the labrum and lie over it. (2) the laMal palps. drones. The antennae are attached to the anterior surface of the head. Associated with the sting are the poison glands. the velum. The legs are very well adapted for the work of the hive. Each segment bears a pair of legs. The first pair of legs are provided with hairs adapted for various uses. composed of insects with two pair of membranous wings. A The manlittle organ. is just below the upper lip." and live The adults come forth in great numbers in May or June from one to two weeks. the epipharynx. The mouth parts are adapted for both sucking up nectar and chewing.


and hind intestines. the gathering of use.REPRESENTATIVE INSECTS 359 digestive tract is well adapted for the specialized life of the bee. mid-intestines. its life in the hive. study of the digestive sj'stem of the larva. . by Mr. while the hind intestine may be divided into the ileum. is Odynerus dorsalis. as Hymen- in the June bug. in the lar\^a. rectum. The divisions of fore intestines are buccal cavity. Just how the queen is able to regulate the laying of eggs that are fertilized by the sperms from the seminal receptacle or those that are not fertilized is not fully known. There a marked change in the length of the esophagus during metamorphosis. are fascinating subjects dealt with in the many books devoted exclusively to a study of the honey bee. The queen bee when fully developed mates with a drone on the virgin flight. looped stomach. Edwin Vest reveals that it optera. It is con- nected by a nerve rmg with the subesophageal ganglion. water sac or Jioney The mid-intestine consists of the stomach. pupa. larval stage to the adult. or in the honey bee. The number of Malpighian tubules increases from the anus. The life history of the bee. as well as many other In Odynerus or the honey bee the digestive tract may be divided into the fore intestines. which are conof thoracic spiracles ajid eight pairs on the is is similar to that of the grasshopper. and the pro: the In the larval and early pupal stages the mid-intestine is a thin but in the adult it has developed into a convoluted. The nervous system brain is well filled with tracheae. while there are is also Only four Malpighian tubules are found around one hundred in the adult. as well as nectar and development into honey for table swarming and the rearing of a queen. The a ganglion in the head above the esophagus. and ventriculus. flat tube. The fertilized eggs develop into workers and the unfertilized eggs into drones. esophagus. that of gathering and feeding upon the nectar of flowers. and adult of the The A solitary wasp. very similar to that of the honey bee. rectal glands. the honey stomach. mouth or stomach. In the adult the esophagus extends from the buccal cavity through the thorax into the first abdominal segment where it enters the water sac. The two ganglia of the head are connected with two in the thorax and four in the abdomen. which is in the head but below the esophagus. By means of the copulatory organ the male transfers a supply of sperms to the seminal receptacle of the queen. The body of the bee nected with two pairs abdomen.

The pharyngeal clefts or gills provide a more effective mode of respiration for aquatic animals than that used by most non- chordates because the passes through them. a series of paired slits in the wall of the pharynx and in the body wall of some (3) dorsally located tiibular nerve cord. These three characteristics are: (1) noiochord. then as a groove. gills are thus interposed directly in the course of the circulation. such at some time during the it is every cliordate animal. lying dorsal to the digestive tube and ventral to the nerve cord. and finally a tube which results in the spinal 360 . All individuals classified in the phylum possess three distinctive characteristics that are most conspicuous in certain primitive forms. a flexible rod extending from anterior to posterior in the longitudinal axis of the body. mud-burrowing forms. of the vertebrae. The three features clearly distinguish the phylum from all others and bind together individuals which are widely separated in appearance but characterized by certain traits peculiar to this group alone. as a plate. (2) pharyngeal clefts or gills. small colonial forms. extending the length of the body dorsal to the notochord and other organs. cord) is made up of the group of animals which includes man himself and in general the more con- known animals. CHAPTER Phylum Chordata spicuous. Although the become modified to form other structures in the adult terrestrial chordates including man. In the adult vertebrate replaced by the centra time in the gills The gill clefts are present at some life of all individuals placed in this phylum. It includes minute sessile forms. Characteristics There is a rather wide range of variation as to form and size in the group. they have had rather typical ones as embryos. and the entire blood supply of the body The central nervous system is derived from first the ectoderm along the middorsal line of the embryo.. and on up to the largest and most complex of living animals. and is The notochord life of serves as a stiffening rod It is present as the foundation axis for the endoskeleton. better XXIII PHYLUM CHORDATA (kor da' ta.

ten genera and twenty-eight species. fishlike forms Cephalochorda (sef a of which Aniphioxus (Branchiostoma lanceolatus) is the most common representative. free-swimming. head cord) includes approximately twenty-eight different species of marine. either free-swimming or sessile. These are all small wormlike animals. and more intimate association of ganglionic masses to furnish better coordination. three classes: (1) Larvacea. 16 kor'da. tail cord). eration. Thaliacea. so named because it retains the larva form throughout colonial. 361 In higher forms the anterior end of the tube beexpanded and modified to form the brain. increased accessibility. and order Cephalodiscida with its two genera C ephalodiscus and Bhahdopleura. Genus Appendicidaria is an example. simple or may reproduce sexually or by budding. but it is progressively obscure as one proceeds from simpler to more complex forms. The chordates possess segmentation (metamerism). cludes the tunicates. life. shore-loving. the sea squirt. Cynthia. half cord) or sometimes known as Enteropneusta (en ter op nus'ta) includes order Balanoglossida with its four families. . These are all advances in both structure and function when compared with other groups. (3) Molgula. Urochorda (u r6 kor'da. usually exhibit alternation of gen- Salpa and Doliolum are the most common examples. and Ascidia are common examples. but cal freedom allowed and this is it greatly increases the mechania distinct advantage as well as an advance in structure.000 different species in this phylum which is divided into four established subphyla as follows: Hemichorda (hemikor'da.PHYLUM CHORDATA cord and brain. and allows for an increase in number of nerve cells. all or Tunicata (tunika'ta) in- of which are marine and mostly small. (2) Ascidiacea. compared with the external one of others studied does not give as great a leverage for the muscles. it is Adults show a high degree of degeneration so the larvae only There are that exhibit distinctive characteristics of the phylum. pelagic. The continuous comes tubular nerve cord is at the apex of the development of centralization in the nervous system. There is a tendency toward fusion of metameres and The internal skeleton of this group shifting of superficial muscles. solitary or colonial forms. Classification There are approximately 40.

The proboscis. Others of the subphylum may be as short as one into the mud and body are The three portions of the and a segmented trunk. (4) development of five senses. the former was with Annelida and the latter was independent. inch or still others as long as four feet. Hemichorda and Urochorda were not classified as Chordata. This process. proboscis. The cavity of the proboscis is filled with water which is dra"wn in and expelled through a proboscis pore or vent located on its dorsal Supporting the base of the side and just anterior to the collar. a ringlike collar. (5) pharyngeal gills (1) and lungs for respiration. in a peculiar position. this group is of no economic importance. and coordination of muscles. (3) development of dorsally located nerve cord with anterior brain. With the exception of the value as biological specimens and the use of amphioxus as food by Chinese. jointed) animals with backbone —frog. digestive tube which is characteristic in the embryonic development of the notochord for certain higher chordates. (6) voice production. Phylogenetic Advances of Chordata Notochord and endoskeleton.362 Yertehrata (ver TEXTBOOK OF ZOOLOGY te bra'ta. is usually referred to as the rudimentary notochord. (2) pectoral and pelvic girdles with limbs. These are the larger. They range from 6 to 10 inches in length. called the diverticulum. is hollow and serves as a water chamber. (7) specialization Protochordata (Lower Chordates) Until relatively recent years the two subphyla. will be man. They are wormlike animals which burrow sand along the seashore. it is it very poorly developed and has the relationship to the Nevertheless. more conspicuous animals and discussed at length in later sections of the book. The mouth opens on the ventral side just anterior to the collar and leads into the straight alimentary canal which extends to the posterior end of the body . is a short skeletal process which is stiff and extends anteproboscis riorly from the roof of the mouth region and assists in burrowing. However. SUBPHYLUM HEMICHORDATA One of the species of Balanoglossus or Dolichoglossus koivalevskii will serve as an example. as well as the collar.

There is no differentiation of a distinct pharynx. absorbing the organic matter from it Balanoglossus has numerous paired gill slits. 209. located in the lateral walls of the anterior (supposedly pharyngeal) position of the digestive tube. 210. (From Hegner. water is passed through them for respiratory purposes. glossus. it lives 363 mud in which as nutriment. —External Glomerulus features of DoUchoglossits kowalevskii. and a The nervous system . published by The Macmillan Company. is of these main vessels composed of a dorsal cord which is tubular in the region of the collar and extends the length of the trunk. Where gills are present.) PHYLUM CHORDATA and ends in the anus. oxygen being absorbed and carbon dioxide being discharged from the blood here. a more or less concentrated center of nerve cells in the collar. Pericardium / Collar Nerve cord Dorsal vessel Proboscis coelom Heart / Trunk Proboscis Notochord Mouth Ventral vessel Gill slits Alimentary canal Diagram of a sagittal section through anterior portion of DolichoFig. (Courtesy of Denoyer-Geppert Company. Like the earthworm. this animal utilizes the for food. Sinuslike branches supply various parts of the body. after MacBride. College Zoology.) — The circulatory system is rather rudimentary. At the collar it is joined by lateral connectives vessel extending posteriorly which encircle the body to connect with a ventral below the intestine. It includes a sinuslike heart which is held in a pericardial sac located in the basal part of the proboscis. are much reduced in Fig. In some of the other representatives the gills numbers or are lacking. A dorsal vessel extends posteriorly from the heart to the posterior end of the trunk.

College lished . but the dominance and hollow structure of the anterior portion of the dorsal one. ventral cord certainly The is not a chordate characteristic. cavity (coelom) and pass out the pore with the water as it is exThese animals are dioecious. and the dominance and grooved structure of the dorsal nerve cord. though lacking in complete conformity to chordate characteristics. . Dolichoglossus and its subphylum. pub- larvae. Zoology. which are globular in shape and form a pattern of In this respect and ciliated bands over the body. are fertilized in the water. hatch out Apical plate and become tornaria Proboscis ccelum Mouth Anus Fig. Excretion seems to be accomplished by a mass of vascular tissue (glomerulus?) located in the proboscis just anterior to the heart. theoretical relationship has been proposed. is classified here because of the diverticulum supposedly representing a rudimentary notochord. (From Hegner. with gonads in the form of a genital ridge extending leng-thwise along each side of the anterior portion of the trunk. The group includes Cephalodiscus and Rhaldopleura which are colonial forms living in deep sea. The mature germ cells escape through the body wall. after Metchnikoff larva Hemichorda. 211. represent features which are homologous to the central nervous system of higher chordates.) 364 TEXTBOOK OF ZOOLOGY ventral cord rimning longitudinally on the floor of the trunk. On this basis a Until relatively recent times this larva was mistaken for a form of adult nonchordate animal and went under the genus name of Tornaria. —Tornaria by TheofMacmillan Company. the gill clefts in the alimentary canal. The excreted materials are received by the water in the proboscis pelled. in habit of life these larvae resemble the larvae of the echinoderms.

excurrent siphon. It is saved to the chordates by the presence of all three of the is characteristic features in the larval stage. Clavelma.Digestive glands -. manhatt encidia. The adult is covered externally by a cellulose coat or tunic (test). given particular consideration here. AsThe latter genus represented by M. In saclike this condition it would be an outcast among chordates because as an adult it has no notochord. tubular nerve cord. or sea squirt.. 212. Some are brilliantly tinted with color.End05tyle /Atrium . When the tunic of Molgula is removed. Cynthia. Inside the mantle is the extensive atrial cavity. However. incurrent siphon or mouth) and the other is the atrial siphon (atrial funnel. and Molgula. while the adult globular and sessile common forms. or atriopore).PHYLUM CHORDATA 365 SUBPHYLUM UROCHORDA. MOLGULA Subphylum Urocliorda includes a number of common represen- tative marine forms. such as Salpa.PharynK Fig. On the dorsal (unattached) side of the body are two funnellike siphons. which is secreted by the cells of the underlying mantle.. and averages about one inch in diameter. one may see most of . Diagram of Molgula manhattensis from the left side to show the structure with the courses of water and food through the body indicated by arrows. and no dorsally located. This animal is comsis will be monly known of the adult is The body as sea lemon.Esophagus --Intestim -Stomach — Branchial fold — . The anterior one is the Iranchial siphon (oral funnel. it does present pharyngeal gill slits. The larva is free-swim- ming and shaped in most of the like a tadpole. Ciona. sea peach. Incurrent siphon Excurrent i/phon Mantle Tunic Qanqiion Ana5 ^ Genital duct Testis Ovary r .

Vessels extend in one direction to the pharynx. excretory sac which may be seen from the right side. The censide of the body. one on the left side in the loop of the intestine and the other on the right closed tral tunicates. There is an oblong. is tween the siphons in the dorsal portion. fertilize ova from another however. It ends with the anus which opens into the atrial cavity shortly below the atrial siphon. may water outside the body. the endowhich extends from the branchial siphon along the ventral midline of the pharynx to the esophagus. and finally from here passes through the the surrounding atrial cavity.fertilization is the rule. A current of water carries food into the digestive system and oxygen for respiratory purposes. The heart is a contractile tube which pulsates. mucus secreted by a glandular groove. The larva possesses the typical notochord. For some reason fertilization occurs in the The . there . finally narrowing at its dorsoposterior extremity to become the small tubular esophagus which turns sharply downward and anteriorly to become the stomach. The life of interest. Cross. absorbed by the blood in the walls of The animal's food consists of minute organisms which is Oxygen style. passes into the sievelike pharynx. primarily. as Molgula. which turns ventrally on itself in a U-shape. the atrial siphon. this Upon viewing from the left the large saclike pharynx may be seen continuing ventrally and posteriorly from the branchial siphon. Each has two compound sets of gonads. It lies ventral to the stomach and forces the blood in one direction by a series of contractions and then in the opposite direction by another are entangled in series. reproduce Some of the sessile by budding. The water enters the branchial siphon. gill slits or stigmata in its wall into and finally leaves the body by way of the stigmata. digestive gland. gills. The eggs hatch to produce larvae somewhat similar to amphibian tadpoles which are free-swimming. These animals are hermaphroditic or monoecious. it The esophagus is partially embedded in a dark-colored The stomach continues anteriorly and upward where becomes intestine. This food mass passes into the esophagus and out through the alimentary canal where digestion and absorption occur. spermatozoa from nervous system one individual usually be exceptions to this. that is. and in the opposite direction to other organs and the body wall.366 TEXTBOOK OF ZOOLOGY the internal organs through the transparent mantle. and nerve cord of Chordata. reduced to a nodulelike ganglion located beNerves branch from this to history of the tunicate is one the various parts of the body.

larva ready for fixation. glion. e. ce... mouth The external opening of this cavity is the atrial siphon. tnt. — . epw." It now undergoes regressive saying this changes involving loss of tail. This places The dorsal side shortens while the ventral side leug-thens. d. .s. cerebral vesicle . notochord st.ves.. ganglion fixation papillae cardium est.. endostyle . / . ci. intestine (From Borradaile and Potts. fix. statolith . ad~ga. an intermediate stage of metamorpliosis.. celluThis process of metamorphosis has caused an active lose covering. rudiment of atrium adult ganglion . at.. stomach stat. G.c. The outer wall of this newly formed cavity is the mantle.. nerve cord. A. 213. completion of metamorphosis. The atrial cavity is formed by in- foldings from the exterior on each side which surround the pharynx and meet each other. The Invertebrata.) ber of gill slits increases greatly. int. eye ciliarv funnel d. published by The Macmillan Company... notochord.. Later the tunic is secreted by the mantle to become a protective. .e.. h. B. ganglion. epigill slits . . atrial opening at. dorsal nerve cord g. The anterior portion of the cord becomes a simple ganThe paired eyes and otocysts (ear structures) also disappear.. ga..oj}. .n... . ^'9-3- cie. . . and posterior portion of located in the anteroventral position. trhqa. . .n.. eiric. and bends the alimentary canal into a U-shape. the mouth in a dorsoanterior position. . .c. the anus in the dorsoposterior The numposition... trk. Metamorphosis of an ascidian lari'a. heart m.f.oji Fig. PHYLUM CHORDATA it 367 then settles on the bottom and attaches itself by adhesive papillae Some authors express it by larva settles on its "chin. at. trunk .

B. the tail — The body of this animal is shaped it is like a being the point. lermudae. is is and the next produced by budding (asexually). — It is of the subtropical found in shore water and on the sandy beaches and tropical portions of the world. fishlike. It is a small. and on the southern Pacific Coast. : form it represents clearly the phylum in a simple condition. but comes to rest with the anterior end exposed to the water. at certain points in the Gulf of Mexico. AMPHIOXUS group which There are four species on American shores Bra7ichiostoma virginiae. B. It burrows rapidly. Habitat. They may be found along the shores of the Mediterranean Sea. This alternation of generation SUBPHYLUM CEPHALOCHORDA. small lance. These ani- mals are found along our Atlantic Coast as far north as Chesapeake Bay. Habits and Behavior. and along the southern coasts of China. In its adult listed twenty-eight species in this There are usually are rather locally distributed over the world. Certain of the sessile forms. stationary form which is not much more than a water-bag whose level of development has degen- erated almost to that of a sponge. the European form. floridae. distinctive is characteristics of the a ehordate. This form is one of the few colonial ehordate animals. calif orniense. floridae is more commonly studied in the United States. the animal leaves the burrow and swims about like a fish by means of lateral strokes of the posterior portion of the body. it is likely that B. However. Branchiostoma lanceolatus. the Indian Ocean. particularly at night and during breeding season. Amphioxus or the lancelet. similar to . another retrogressive feature. head first. In a few instances tunicates reproduce one generatioji sexually. — External Structure. marine animal whose average adult length is about two or three inches. It essentials. in the sand by means of a vibratory action of the entire body. At times. virginiae or B. and B. which reproduce also by budding. possessing only rare It is usually referred to as a close ancestral relative of Vertebrata. In general. develop colonies with a common tunic.368 TEXTBOOK OF ZOOLOGY respectable ehordate to become a lazy. is an admirable representative of the subphylum and has become classical in its use.

side .Cerebral vesicle Oral cirri Velum Velar tentacles Spinal cord Atriopore Intestine Ventral fin Anus Caudal fin Fig. —Diagram of Branchiostoma to (Amphioxus) lanceolatus from the right show the structure. 214.

The segmental divisions of the muscles are apparent on the body wall. There are no clearly defined lateral fins. Adjacent ones are separated by a myocomma or myoseptum. is an opening. the metapleural folds. two-thirds of the ventral surface of the body are thought to be their forerunners. The mouth opens on the ventral surface of the anterior portion of the body.370 a small TEXTBOOK OF ZOOLOGY fish. the atriopore. just posterior to the metapleural folds. 215. but alternate with each other. The ventral and dorsal fins are supported by small vertical rodlike fin rays. extending along the anterior Dorsal fin F/n ray Epidermis Neurocoek NotochordoL _ sheath '/ '/^-^ . Fig.Nephridium Atriaicavity ^ Pharynx Gil/rod Gill _ _ Liver bars Qonad Hypobranchial qroove Ventral aorta. This hood is fringed with sensory fingerlike oral tentacles. There is a median fin along the dorsal side. There are from fifty-eight to sixty-four of them on each side in B. On the ventral side. lanceolatus but sixty-nine in B. calif orniense and they are known as myotomes. but it does not have a distinct head. Endostyle Coelom Metapleural fold S.^ Spinal \ ^j nervz Nerve cord Nobochord S Myoto. . The myotomes on the two sides are not paired. but a pair of skin structures. — Cross section of Amphioxus thirough the level of the posterior portion of the pharynx. and beside the ventral margin of the caudal fin is the muscle \ >] Myocomma Dorsal Aorta Coelom Atrial cavity- ^K— = A^-Epibranchial qroove . continuing around the tail as the caudal fin and anteriorly about one-third of the length of the body as the ventral median fin. It is beneath a rostrumlike projection and is nestled well up in an oral hood which is shaped like an inverted funnel.

.HepaticV. -Atriopore -Vcntro.PHYLUM CHORDATA 371 Buccal cirri QUI slit inwall of phorynx Afferent branch- Jal arteries Ventral aorta Dorsal aorta ..Dorso-intestinai A. —Notochord _ -Spinal cord -Distribution throuqh liver ^ . . Subintestinal vein . 216. Anus Caudal vein 1 Caudal artery Fig. —Diagram of the circulatory system of Amphioxus.intestinal V. .

body giving off the flow of the blood is just opposite to that in the dorsal vessel of the from the inand continues anteriorly to the liver as the hepatic portal vein. Here these vessels is branch into capillaries. oral and velar tentacles. which branches to form the afferent branchial arteries to the gills. The circulatory system does not include a heart. The nerve cord gives off nerves to the organs of the body. In small cleared specimens the internal organs are easily observed. There are dorsal sensory nerves going to the skin and ventral motor nerves wise through going to the myotomes. The blood in these ventral veins flows from posterior toward the anterior (Fig. Surrounding the mouth is a membranous velum to which are attached twelve velar tentacles. The posterior direction of sally to join the paired dorsal aortae. subintestinal vein receives the blood testine earthworm. which fold across the mouth and serve as a strainer to hold back the coarser particles. There are smaller pigment bodies distributed along the length of the cord. 216). but the blood moved by the contractions of a ventral aorta. The mouth leads to the large. These cilia form what is called a wheel organ because of their rotary motion. The gill slits are clefts in the lateral walls of the pharynx. The Digestive System. These capillaries converge to form the efferent branchial arteries which lead dorteriorly to the tip of the The dorsal aorta extends posnumerous branches to myotomes and internal organs along the way. but those behind the cerebral vesicle alternate on the two sides. A mass of dark pigment is located at the anterior end which is known as the eyespot. providing aeration for the blood. There are sensory cells in the skin.if 372 TEXTBOOK OF ZOOLOGY Internal Structure and Metabolic Activities. The notochord — extends the length of the body as a slender rod of vacuolated cells which are body. These are thought to be sensitive to light. The hepatic vein collects from the liver and leads forward as the ventral aorta. Immediately dorsal to this rod the nerve cord. which also runs the length of the It has a small central canal or neurocoele it extending length- and is dilated at the anterior end to form the cerebral vesicle or rudimentary brain. barrel-shaped pharynx. A current of water is carried into the mouth by the ciliated bands on the inner surface of the oral hood. The blood in the subintestinal and hepatic portal veins is laden with dissolved nutriment. The number of clefts — . tentacles. The two anterior ones are paired. as well as being sensory. filled with fluid to give is it turgor or stiffness.

373 ranging between fifty and ninety pairs. The gill-'bars. Excretory System and Excretion. In the midline of the roof of the pharynx is an inverted trough. which are capable of secreting mucus. In the floor of the pharjTix Its another ciliated groove. Reproductive System and Life Cycle. contain the blood The gills are on the vessels. The strings of It functions on the same plan here as in tunicates. extends anteriorly from its connection on the anterior part of the intestine to lie on one side of the pharynx. — Ciliated nephridia similar to those of the earthworm lead from the dorsal portion of the coelom to the atrial cavity.PHYLUM CHORDATA varies. Respiratory System and Respiration. the JiypohrancJiial groove. and absorbed from. Behind the atriopore is relatively larger. The food is digested in. — . The intestine extends posteriorly to the anus as a relatively straight tube. the hyperhrancliial groove. Between the posterior end of the pharynx and the atriopore. the water through the gill slits delivers oxygen to the blood in the and absorbs carbon dioxide from it. and are supported by chitinous rods. The cilia here move the mass back to the intestine. constitute the endostyle. The coelomic cavity is reduced in the pharyngeal region to a narrow space surrounding the dorsal aorta above the pharynx and a narrower one around the ventral aorta below. the coelom consists of a narrow space surrounding the intestine with a thin membrane separating it it from the atrial cavity. glandular walls. the liver or hepatic caecum. mucus entangle the food particles and are moved anteriorly. faces of the gill bars and are covered with cilia which help move the water through its course. which is is ciliated. and then by two peribranchial grooves are carried dorsally to the hyperbranchial groove. fingerlike diverticulum of the intestine. This organ is a digestive gland and empties a digestive juice containing enzj^mes into the intestine. which separate the slits. the intestine. calif orniense) nodular gonads embedded in the body wall near the base of the metapleural folds. The blood then distributes the oxygen to all tissues of the body. These open into the atrial cavity which surrounds the pharynx and other visceral organs. in passing capillaries there —As stated above. This animal is dioecious with each mature individual possessing 26 pairs of (31 to 33 pairs in B. A blind. When the germ cells mature. The water then passes back through the atrial cavity and out through the atriopore.

they break through the wall of the gonad into the atrial cavity and pass out through the atriopore with the water. Fertilization occurs Early summer is the breeding season. only to begin bashfully burying itself in the sand. Fol- lowing fertilization comes a series of cleavage divisions which are This is followed by the infolding of one side of the total and equal. in the water. form the gastrula and this in turn becomes a freeswimming larva which reaches adult condition without metamorphosis. and at that time the animals are quite active during the evenings and nights. spherical to body .374 TEXTBOOK OF ZOOLOGY |.

The nech which is a constricted region between trunk and head is conspicuous in terrestrial forms. man himself belongs are found the dis- tinctive chordate characteristics at some time in the life of the indi- In terrestrial forms there are certain modifications to pro- duce other structures. chambers. The sense organs are in an advanced state of development. five-lobed brain located in the more or less distinct head. The appendages are usually arranged with one pair attached to the anterior. fins. excretion. and in terrestrial forms. acteristic The body wall is composed of the skin. such as scales. wings. In the vertebrate body is a well-developed coelom. which usually has chartegumentary outgrowths. respiration. Paired appendages are usually present at some time in the life of the individual. forelegs. trunk. and tail. reproduction. The majority have two pairs of fins or limbs in adult conThere is a ventrally located heart which is divided into dition. feathers. and hair. which encloses advanced systems of organs for digestion. nails. The tail is a posterior prolongation of the body behind the anal opening and is found in some degree in all vertebrates. pelvic region. beneath which is the muscular 375 . universal characteristics symmetry are The segmented vertebral column and other supporting structures form an endoskeleton (internal skeleton) which is the basic support of the body. circulation. This arrangement is less consistent in the aquatic types where the weight of the body is buoyed up by the water and the limbs are used less for support and locomotion. shells. pectoral portion of the trunk and one situated at the posterior. Cephalization is developed in all vertebrates and along with this they possess a hollow. as the outer layer. In Metamerism and bilateral among vertebrates.CHAPTER XXIV THE VERTEBRATE ANIMAL: SUBPHYLUM VERTEBRATA In this group to which vidual. The body is divided into head. The Mood contains hemoglohin hearing red corpuscles and amoeboid white corpuscles. different types of vertebrates there are various modifications of pectoral appendages as arms. and The same is generally true for the pelvic limbs. pectoral flippers.

. the second the lungs. external and internal sheaths (From Maximow and of hair root. hair shaft. stratum . Cor. dermis section of —Diagrammatic gland M. and aldominal cor muscle n. u. . Saunders Co. myelinated nerve fibers P. D. cavities. Histology. . reproductive. 217. corneum . Bloom. papilla of hair Sc. In In all vertebrates. nm'". B. nonmeduUated nerve nm'. and the third the organs of the excretory. and digestive systems. nm". published by W. the coelom consists of only two parts: the pericardial cavity and the general abdominal cavity. nina.. . 376 I'EXTBOOK OF ZOOLOGY is coat and internal to this the membranous peritoneum. except mammals. layer niu. sebaceous gs. . endings of nonmyelinated nerve fibers. . fat tissue v and w. Malpighian . thoracic. mam- mals it is further divided into pericardial. the human skin. and nmb. The first contains the heart.) Fig. x and y.

these proc. It also helps in excretion through the sweat glands. (3) digestion. and assimilation. since it is a study of the functions of living organisms. and the elimination of wastes incidental to it. release of energy through oxidation. feathers. production of heat. (5) circulation. The integument is composed of an outer stratified epithelial epidermis which consists of several layers of cells. movement. blood vessels and lymph spaces. must be concerned with metabolism. The maintenance several functions which any living body requires the cooperation of will attain similar fundamental results wherever in living material they occur. — The collective term metabolism is employed when re- ferring to all of the interactions involved in the living process of protoplasm. therefore. transformation of energy. here are ingestion. few nerves. The processes concerned with the conversion of food material into protoplasm (building up) Included constitute the phase of metabolism known as anaholism. elimination of wastes or. and (9) internal regulation. integumental glands. The oxidation of materials of the protoplasm to liberate energy. nerve endings. egestion. and oil glands as well as Such exoskeletal strucfacilitating temperature regulation in some. in other words. digestion. tures as scales. absorption. mucus glands. hoofs. It includes the processes concerned with conversion of food into protoplasm. stantly being changed from the potential to the kinetic form. claws.THE VERTEBRATE ANIMAL —SUBPHYLUM VERTEBRATA 377 The vertebrate animal is covered by an integument or skin which serves as a protective and sensorj. and reproDuring the execution of these activities energy is conduction. These functions merge into one living process which involves the building up of protoplasm. (7) reproduction. (2) movement and locomotion. respiration. Metabolism. The principal functions performed by the structures in the animal body are: (1) support and protection. nails. which consists of areolar connective tissue. (4) respiration. oxidation. digestion. and elimination. transportation. transporta- tion. It includes all Metabolism is . absorption. The membrane type of bone of is developed in the dermis. one of the fundamental features of all protoplasm. hairs and enamel of teeth are produced by the skin. and the inner fibrous dermis or corium. (6) excretion. nerves. esses are chiefly : Ingestion. and no blood vessels.organ. (8) reception and conduction of stimuli. is known as cataholism or the "breaking down" phase. all physiology.

Cranium Sense capsules Jaw apparatus (Visceral arches) . The exoskeleton is a rather minor part in vertebrates and consists of nails. thoracic vertebrae of the chest. firmness and rigidity. claws. axial and vertebral column. and the caudal vertebrae of the tail Bone is a firm. composed of inorganic salts. hard tissue consisting of abundant matrix. The appendicular the anterior and posterior girdles and two to be called cartilage I ones or they develop in the connective tissue of the dermis. Axial Skeleton (a) Skull 1. ribs. lumbar vertebrae of the small of the back. The skeleton is quite well developed in the vertebrates and serves them quite efficiently for support. The endoskeleton includes the hair. five regions as follows: cervical vertebrae of the neck. scales. portion is pairs of limbs. and is divided into hones. Caustic solutions will destroy the animal matter and make the bone brittle. A acid. The following outline weak presents a summary of the principal parts of the terrestrial verte- brate skeleton. The first is composed of the skull. Divisions of Skeleton of Terrestrial Vertebrate I. composed of In their development bones either replace cartilage and in some a sternum. 3. 2. appendicular portions. sacral. stature. region. protection. to be is known as membrane composed of segmental divisions. The vertebral column the vertebrae. and the bone cells which are held in The outer membranous covering The mineral part of the bone consists of bone is called periosteum. They give it chiefly of calcium phosphate and calcium carbonate. and other outgrowths. It is composed of cartilage entirely in some of the simpler forms and of bone and cartilage in higher types. and muscle attachment. It is divided into an exosJceleton which is superficial and an inner endoskeleton which includes all of the deeper skeletal parts. in which case mineral the bone will lose its rigidity. feathers. vertebrae of the hip region. will dissolve the matter of bone if allowed sufficient time. pocketlike lacunae in the matrix.378 TEXTBOOK OF ZOOLOGY of the chemical changes and transformations by which energy is supplied for the activities of the protoplasm. The animal matter is composed of the bone cells and cartilage which serve to give the bone life and resilience. such as the acetic acid in vinegar.

metacarpals (palm). or horny skeleton extending throughout the body. and ischium Limb: Femur (thigh). animals like the starfish. Appendicular Skeleton (girdles and limbs) (a) Pectoral (anterior) 1. ilium. metatarsals (sole). Sternum (breastbone) II. crayfishes. phalanges (bones of digits) (b) Pelvic (posterior) 1. 2. carpals (wrist). The presence of a cuticle in some and the secretion of a hard shell in others seem to be the particular developments related to these special functions in this group.THE VERTEBRATE ANIMAL (b) Vertebral column 1. tibia and fibula (shank). 5. There are special cells of the epidermis which function primarily The echinoderms. as a rule. beetles and representatives of their respective phyla secrete a well-developed exoskeleton as an external cover over most of the other tissues of The muscles and other tissues are attached within. 3. Such forms as snails. Lumbar Thoracic vertebrae (chest) vertebrae (small of back) Sacral vertebrae (hip) Caudal vertebrae Eibs (paired) (tail) (c) Thoracic basket 1. Girdle: 2. including in production of this skeletal material. possess calcareous skeletal plates which are essentially similar to exokeleton except that they are principally beneath the skin. Girdle: Limb: scapula. pubis. In Protozoa there is The skeleton and integumentary structures serve the Metazoa primarily for a support and protection. clavicle. The corals of the phylum Coelenterata secrete a calcareous or horny skeleton around the exThe sponges. patella (knee cap). Arcella. the body. Difflugia. siliceous (glassy). A number of exoskeletal modifications are used for There is . —SUBPHYLUM VERTEBRATA 379 Cervical vertebrae (neck) 2. 4. the Foraminifera. procoracoid and coracoid Humerus (upper arm). radius and ulna (forearm). no well-developed endoskeletal structure known in nonchordate animals but the endophragmal structures extending into the thorax of some Crustacea are thought to be the forerunner of the endoskeleton. tarsals (ankle). phalanges (bones of toes) no very elaborate adaptation toward a skeleton. and Radiolaria exemplify this adaptation. 2. each ternal surface of the body proper. have a calcareous.

published by the McGraw-Hill Book Company. hair. girdles. Animal Biology. the original endoskeletal structure of are developed the basic structures of Around Cranium Ska// Orbit Mandib/e Ceri//ca/ vertebrcxe C/av/c/e Scapu/a Sternum Thoracic Humerus Rib vertebrae Lumbctr vertebrae Pe/vis Sccrum Radius Utna ^Carpa/s Mefacarpa/s Hc^nd Tibia Metatarsals Fig. is it Primitively the notocJiord the chordate group. nails. and even enamel of teeth are of this type. horns. Such structures as scales. life of vertebrates and the necessity . developed with the terrestrial for locomotion on land.) the vertebral column which functions as the principal axial support The sternum.380 TEXTBOOK OF ZOOLOGY protection and temperature regulation in most of the groups of vertebrates. 218. (From Wolcott.- -Human skeleton. and paired limbs have of all vertebrates. shells. feathers.

cells. they all pass through a free. The muscles are usually attached to the skeleton or occasionally to other muscles by fibrous cords called tendons.. During the shortening of the muscle there a hydrolysis or absorption of water by the protein product. A muscular locomotor system conis opposing muscles. Cardiac muscle is the highly specialized involuntary muscle which makes up the wall of the heart. but more often they cover only areas cilia of free surface of epithelium. are involuntary. corals. Oxidation and heat production are involved in the process. cilia. fundamental basis for all animal movement. to move separate parts in adult Simpler forms of locomotion have already been seen in Protozoa which move from place to place by means of pseudopodia. . creatinine. By-products of the process include carbon dioxide. active larval stage. and others. Most of them retain the power condition. those of the visceral organs. Carbohydrates in the form of glucose are oxidized (burned) in the is reaction. In many Protozoa the entire body is covered with while in Metazoa the entire body may be covered where they are used for locomotion. e. creatine-phosphorie acid. are sessile however. In muscular contraction there a cycle of rapid chemicophysical rearrangement in the cells. particularly the linings of passages. so timed that the beat passes in a wavelike progression from one end of the ciliated area to the other (metachronous rhythm). . and phosphoric acid. of a high degree of contractility in special makes possible muscular movement which the principal kind in higher animals. oysters. urea.g. or flagella. The stroke of a eilium consists of a vigorous bend in one direction and a very deliberate recovery in the other. In ciliary movement the numerous small strands of protoplasm beat rhythmically with a stroke in one direction. Voluntary muscles are usually connected with the skeleton.THE VERTEBRATE ANIMAL SUBPHYLUM VERTEBRATA 381 The muscular system represents a system of cells highly specialized in contractility. almost a characteristic of Contractility as a property of all protoplasm is the animal life. lactic acid. barnacles. is The development such as muscle sists of sets of cells. such as sponges. The adult forms of cer- Independent power of movement is tain animals. intestine. Here they serve to move materials along and keep the surface free of foreign material and excess mucus.

such and respiratory organs. Its internal lining is provided with numerous fine fingerlike projections which The digested increase the inner surface and enhance absorption. Following the mouth is the pharynx or throat region which receives the internal nostrils.382 TEXTBOOK OF ZOOLOGY In animals without a skeleton muscle bands are arranged in both circular and longitudinal directions. In echinoderms with the water vascular system the pressure is exerted on water in a system of tubes which extend is to make contact with the surface over which the animal moving. The Dig"estive System. . This is a convoluted tube in most of the advanced forms of vertebrates and is divided into the anterior duodenum. and the shortening of the longitudinal strands draws the body along. In the higher forms there are many outgrowths. peristaltic contractions continue along the wall of the stomach to help digestion by churning and mixing the food with digestive juices. and receives saliva from salivary glands. it is It is usually coiled. The anterior region of the digestive tube is the mouth cavity which contains teeth on the jaws. middle jejunum. secretes the bile which is stored in the thin-walled gall Madder. At the posterior end a pyloric valve in the form of a sphincter muscle guards the entrance to the small intestine which follows. a tongue. The contraction of the circular group tends to lengthen the body. Its walls : are embedded in the wall just outside of the lining epithelium. The small intestine serves both as a digestive organ and as the principal absorptive organ of the body. — The digestive system is typically a straight tube extending through the length of the trunk of primitive vertebrates. food is taken up by the lymphatic spaces and by blood vessels which posterior ileum. The It liver is the largest organ in the body of most vertebrates. a process known as peristalsis. The esophagus is usually tubular and propels the ''swallows" of food posteriorly by consecutive waves of contraction. the Eustachian tubes from the middle ears and opens into the esophagus posteriorly. and longer than the body and therefore produce digestive enzymes from glands and the liver and it receives digestive juices from two other glands the pancreas. as digestive glands containing enzymes whose walls possess gastric glands for secretion of a digestive fluid The (ferments) and weak hydrochloric acid. It leads to the saclike stomach. The pressure exerted on the coelomic fluid is thought to be a factor in bringing about an even extension of the body by this means.

THE VERTEBRATE ANIMAL which is —SUBPHYLUM The VERTEBRATA 383 attached to one of its lobes. and hydrolysing enzymes by causing a reaction between a substance and water. is and finally glucose. amino and fats an organic substance which by its pres\vill cause or hasten chemical reaction between other substances without itself being consumed. Most of the digestive enzymes fall in this latter class. There are different types of enzymes each capable of producing speIn general. The materials commonly used for foods have large molecules. Digestion then must serve to break up these large molecules into smaller ones. termed activating agent or coenzyme. proteins are converted to soluble to maltose materials in solution. . The large intestine which forms of vertebrates the posterior portion of the large intestine is modified to become a cloaca. lases) which cause clotting or coagulation. coagulating enzymes (coagu. usually called colloidal in nature. The enzymes are formed in the protoplasm of cells and their action is similar to that of a catalyst. the precursor of pepsin is pepsinogen which becomes activated in the presence of dilute (hydrolases) act acid. There are oxidizing enzymes (oxidases) capable of bringing about oxidation reducing enzymes (reductases) which produce reduction in tissues. which receives also the products from tlie urinary and reproductive organs. As an example. starches to and sugars fatty acids and glycerin. than the small intestine possesses no In many receives the fecal matter and delivers it to the anus. may be excreted is shorter from It villi or digestive glands. Most enzymes consist of a parent substance or precursor (zymogen) which becomes active only in the presence of a certain other substance. an enzyme ence under certain conditions cific kinds of reactions. Digestive enzymes are responsible for placing the food So. thus forming solutions of substances in order that they will readily diffuse through membranes. chief function of this entire system is The that of dissolving and converting complex food materials into a form which may be absorbed and assimilated by the protoplasm of cells throughout the body. since they accelerate chemical action. converted into urea and uric acid in order that they the blood in the kidneys. acids. liver also serves to convert ener^ carbohydrates to glycogen (animal starch) and store it for future It is also in the liver that protein wastes are production.

In carnivorous animals it is about one-half the area of the skin while in herbivorous animals it is about twice the area of the digestive tract is skin. tein. Diastases or diastatic (a) Ptyalin in saliva (b) (c) enzymes — split carbohydrates Amylase in pancreatic juice Glycogenases liver and muscles Converts glycogen to glucose — 2. In carnivores (flesh-eaters). In hydrolysis the large molecules of procombine with water and then split into Some foods may require more than one such first . carbohydrate. relative length of this canal varies considerably depending on the habitual diet of the organism. 3. and and general because of the omnivorous food habits. while in herbivorous forms (plant-eaters). such as horses and cows. and (2) The associated glands which discharge digestive juices into it. The length of the it is over twenty times as long as the body. The digestion of all organic food materials is brought about by hydrolysis in the same The process is fairly of digestion in it typical kind of chemical change. Trypsin in pancreatic juice Erepsin in intestinal juice Clotting or coagulating enzyme (a) Eennin in gastric juice —functions In higher Metazoa digestion is accomplished principally extrathrough secretion of enzymes by certain groups of cells. man is quite well understood. The relative proportion of the internal absorptive surface of the alimentary canal to the external surface of the body is significant. it is human approximately ten times the length of the body. or fat simpler products. The action of the several enzymes produced by different glands is a very essential part of the process. Such systems consist of: (1) an alimentary canal proper. Lipase or lipolytic enzyme splits fats (a) Steapsin in pancreatic juice Inverting enzymes convert disaccliarids to the less complex monosaccharids (simple sugars) (a) — — —intestinal juice Maltase (b) Lactase (c) Sucrase (invertase) 4.384 TEXTBOOK OF ZOOLOGY Classes of Digestive Enzymes 1. cats and dogs. such as cellularly from three to five times as long as the body. Proteases or proteolytic enzymes (a) Pepsin in gastric juice (b) (c) — split complex proteins in small intestine 5.

such as cream. steapsin. respectively on proteins This protease is in the form of trypsinogen until it reaches the intestine and is activated by an intestinal enzyme. Be^inin causes the casein in milk This is the first step in its digestion. —When chyme is ejected through the pylorus into the duodenum. the Intestinal Digestion. starches and sugars. The pepsin when present in the acid medium brings about the splitting of complex proteins into intermediate proteoses and peptones. enterokinase. There is some evidence that secretin lates secretion in the liver. Trypsin completes the work begun by the pepsin in that it converts proteoses and peptones into amino acids. but it also digests complex proteins which have escaped the action of pepsin. SUBPHYLUM VERTEBRATA 385 The splitting of the disaccharide. it is discharged through the pylorus. will serve as this process: an example of (Malt sugar) (Water) (Glucose) The two molecules of glucose formed are absorption. known as secretin. In a complex protein like casein. and the lipase. the diastase.ijlopsin. etc. in a form for ready Gastric Digestion. These act and peptones. — The tubular gastric glands located in the secrete the acid gastric juice which is a and two important solution of 0.5 per cent hydrochloric acid enzymes.) a clear. maltose.THE VERTEBRATE ANIMAL splitting. some that emulsified fats. the hydrochloric acid stimulates certain cells of the intestinal lining. into the small intestine by way of the also stimu- pancreatic ducts. and fats. pancreatic juice. . am. pepsin and rennin. tinually churned and mixed by muscular activity of the walls. alkaline solution containing and three enzymes. are regarded as hmlding stones of the protein molecule. It is claimed by to coagulate.2 to 0. It acts more rapidly and There are nineteen amino acids that efSciently than does pepsin. causing them to secrete into the blood a substance of hormone nature. as many as sixteen of these amino acids will trypsin.4 per cent) with acid and has been reduced mucous layer of the stomach to the consistency of soup. are partially digested by a The digesting mass or chyme in the stomach is congastric lipase. Upon reaching the pancreas this secretin stimulates it to secrete the digestive fluid. the protease. "When it becomes saturated (0. Pancreatic juice inorganic salts is (carbonates. watery.

When the chyme reaches the large fermentation. The alkaline salts which are introduced by the bile. Certain bacteria (B. levulose. common This neutralize the acidity of the chyme . ting of fats into glycerin (glycerol) and one or such as stearic acid. Maltase converts maltose and dextrin into Invertase changes sucrose (cane sugar) into dextrose and Lactase converts milk sugar (lactose) into galactose and dextrose. intestine it is about the consistency of thick cream. has been mentioned already. The undigested residue others) attack passes into the large intestine where prob- ably no enzyme digestion occurs. Intestinal secretions or succus entericus which are produced by glands in the mucous membrane of the small intestine include five enzymes. Cholesterin and two pigment materials are excreted in bile duct. and it is able to of bring about hydrolysis of carbohydrates in the alkaline the intestine without activation. coli and any undigested protein and bring about putrefactive Products of this action may be absorbed. It medium (malt sugar). which activates trypsinogen to form trypErepsin. Certain other bacteria here feed upon cellulose and may produce some sugar from it. dextrose. both simple sugars. This with the pancreatic juice brings about the emulsification of fats mentioned above. combine with these fatty acids to form soaps which help in emulsifying the remaining fats. but it becomes more and more solid by absorption of water here until finally only concentrated fecal matter remains. some of them are frequently toxic and must be eliminated in either the urine or the feces. sin. more fatty acids. oleic acid. The pancreatic lipase. Amylopsin (amylase) is the pancreatic diastase.386 be found. discharged into the The secretion of the liver is bile and is duodenum of the small intestine by way of the is — an alkaline solution which serves to help as it comes from the stomach. Enterokinase. brings about the splitetc. the intestinal protease. produces dextrin and maltose steapsin. supplements the activity of trypsin by converting proteoses and peptones into amino acids. thus making them more readily split. butyric acid. Functions of the Liver. TEXTBOOK OF ZOOLOGY The tissues of the animal body must not only have avail- able a wide range of amino acids but must also select in the proper proportion the ones needed to reconstruct their specific protein constituency.

THE VERTEBRATE ANIMAL —SUBPHYLUM VERTEBRATA 387 The Digestive Enzymes and Their Functions ENZYME .

305 Calories Some portion of the dextrose is distributed it and oxidized directly for immediate energy. of proteins in the is collected by the body are to rebuild debiliand energy to the body by oxidation. and water there is another indispensable class of food material. and help supply heat parative heat production values of the three are as follows One gram of protein z= One gram of carbohydrate z= One gram of fat = 4. the vitamins. Car- bohydrates in excess may be converted to fat. hepatic portal vein The blood supplying the intestine and delivered to the liver. but much of is transformed into glycogen This by the enzyme glycogenase tissues as needed. in the muscles to be reconverted into dextrose for oxidation stored as a reserve fuel supply in adipose tissue over the body. Besides proteins. or in the case of fats into the lacteal IjTnphatics and from here into the subclavian vein. In They are recognized usually through the abnormal condition brought on by their deficiency. The final oxidation products of carbohydrates in the body are heat. liter of water one . may be stored here or by the Normally there is a constant supply of dextrose (0. Much of our knowledge concerning the symptoms brought on by lack of different substances has been •A Calorie equals the amount of heat necessary to raise one degree centigrade under standard conditions. and stored. The com: The two functions tated protoplasm. Vitamins and Their Functions.1 to 0. and carbon dioxide. The last two are diskinetic energy. They are natural substances found general. in the liver. Fat is converted to dexIt is usually trose and oxidized to produce heat and kinetic energy. They serve first and best for the purpose mentioned first. inorganic salts. Oxidation of protein requires the disposal of much more waste products. charged from the body as waste products. Carbohydrates and then fats are more economical and efficient as sources of fuel for production of heat and energy. fats.: 388 TEXTBOOK OP ZOOLOGY Absorption and Utilization of Food Materials.100 Calories 9. There is little danger of vitamin deficiency for adults living on a balanced and mixed diet. in relatively small quantities in a regulatory.15 per cent) in the blood and this level must be maintained. water. their function is — number of different foods. — The soluble prod- ucts of digestion are absorbed through the semipermeable epithelial lining of the intestine into the blood of the adjacent capillaries.100 Calories* 4. carbohydrates.

THE VERTEBRATE ANIMAL —SUBPHYLUM VERTEBRATA 389 gained by feeding experiments on different kinds of laboratory animals and results applied to human beings. egg yolk. (c) Effects of Pellagra in primates (man and monkeys). and muscular activity of digestive Cessation of growth. egg yolk. carrots." 1. and milk. green leaves. human (cheilosis). Bj or Thiamin (Ci2Hi. Vitamin A (C20H30O) —antixerophthalmic—fat soluble. . (b) Functions: witli Necessary for growth. The following outline will give much of the essential information concerning vitamins. (a) Sources: carotene (CjoHBe) a yellow pigment in green plant leaves. Nicotinic Acid (CgHsNOz) (a) Sources: —antipellagric. yeast. Meat. (a) Sources: Milk. stimulates appetite. 2. Germ of wheat and other cereal grains. Xerophthalmia (lack of tear secretion and dry "Nutritional" roup in cornea). Polyneuritis develops in birds. (b) Functions: essential for (c) Promotes tone in alimentary tract. May influence oxidation of food. cod. wheat germ. Transformation of this pigment into the vitamin which is especially stored in shark. liver. 4. Dermatitis of turkeys. (b) Functions: (c) Necessary for growth. (a) Sources: Eggs. peanuts. balances deficiency: lular function. and "night blindness" in human. cel- (b) Functions: Produces active "coenzymes" (I and II). milk. loss of tonus tract. and egg yolk. important factor in regeneration of visual purple of retina. Effects of deficiency: still Paralysis in chickens. "Curl-toe" paralysis of chickens. Be or pyridoxine (CgHuOaN). yeast. and such plant tissues. There are other recently discovered fractions of Vitamin B. yeast. Swine pellagra. "Yellow liver" of and inflammation at corners of dogs. green leaves. II. Vitamin B* "Complex. birds. active relation to several enzymes intermediate metabolism of food. The Vitamins and Their Characteristics I. normal growth. halibut or other fish liver oil. liver. (b) Functions: Influences efficiency and acuity of vision. liver. cereals. Beri-beri (neurodigestive disturbance following Effects of deficiency: diet of polished rice). liver. strengthens and pro- motes hardiness in epithelial (c) Effects of Deficiency: tissue. Riboflavin (C^H^oOoN^). of in (c) Effects deficiency: Irritation mouth 3. whose func- tions are specific. Black-tongue in dogs.ON4S) (a) Sources: —Antineuritic. essential for carbohydrate metabolism.

also certain bacteria of the "intestiInfluences the production of prothrombin by the liver is nal flora. Dermatitis in rats and 6. The Respiratory System. Biotin (doHieOsNjS). E or Tocopherol (CjaHjoO. beneath skin and into joints). have developed the trachea (windpipe) and lungs as another outgrowth of the pharynx. Effects of deficiency: Thickening of skin and dermatitis in clucks and III. between the external environmental medium and the blood and internal respiration which is the . A certain amount of respiration takes place through the skin. Sources: Green leaves. brates respiration is is at least in In most aquatic verte- accomplished by drawing water through gill Air-breathing. bones (osteomalacia) especially in V. mammals except pri- Citrus fruits. cod-fish liver oils." (b) Functions: (prothrombin (c) Effects of deficiency: Blood fails to clot. terrestrial forms slits in the wall of the pharynx. oxygen and carbon dioxide. Failure of Death of rat embryos in uterus. (b) Functions: Maintains structure of capillary walls. egg yolk. peanuts. green leaves. The respiratory process is composed of two phases: exterTial respiration which includes the exchange of the gases. alfalfa. Vitamin (a) K (C3. rats. Effects of deficiency: chickens. and differentiation. Vitamin (a) women of the orient. IV. black rats. molasses. (a) Sources: Liver. (b) Functions: (c) Essential for growth. (a) Sources: Tuna and Exposure of skin to ultra- violet radiation. (c) Effects of deficiency: deformed bones in young (rickets). Vitamin in D (C2SH44O) —antirachitic. male fowls and rats.— The respiratory system part an outgrowth of the digestive canal. (b) Functions: (c) Promotes rapid Effects of deficiency: spermatogenesis.)- —antisterility. Soft. tomatoes. yeast. Graying in (b) Functions: (c) Essential for growth. other vegetable fats. (b) Functions: Eegulation of calcium and phosphorus metabolism. cell proliferation Sources: Wheat germ oil. TEXTBOOK OF ZOOLOGY Pantothenic aeid (CgHiTOgN). milk. yeast. (c) Effects of deficiency: Scurvy in human and guinea pig (bleeding mucous membranes. (a) Sources: Egg yolk. turnips (most mates and guinea pig can synthesize this vitamin). .390 5. KeSoft quired for normal growth and mineralization of bone. cereal grains. Vitamin C or Ascorbic Acid (CoHsOe) (a) Sources: —antiscorbutic-water-soluble.H4e02) —antihemorrhagic. molasses. necessary for blood clotting). Sterility in VI.

exposed to surrounding water from which the dissolved oxygen is In many aquatic worms the gill filaments are outgrowths of the sides of the body wall. Also the possibilities of oxygen absorption are greatly increased by the development of respiratory pigments like hemoglobin and hemocyanin. This movement of gas through the cell membranes depends on the partial pressure of the particular gas on the two sides of the membrane. In protozoa and simple metazoa. cells becomes carbonic acid and carbonates which may be transported by the plasma (fluid) of the blood. Respiration has been defined as the process involving the ex- change of gases between the protoplasm of an organism and its environment. All living protoplasm must be provided with a means of receiving oxygen and giving up carbon dioxide. Gas will flow in the direction toward the least pressure. hemoglobin. the oxyhemoglobin releases its oxygen rapidly. Thus the blood is enabled to absorb far more oxygen than an equal quantity of ordinary liquid. coelenterates. Carbon dioxide accumulates in excess in the tissues and diffuses from the cells to the lymph. a number of aquatic insects. such as sponges.THE VERTEBRATE ANIMAL —SUBPHYLUM VERTEBRATA 391 exchange of the gases between the blood and the protoplasm of the Much of the carbon dioxide given up by the cells over the body. round- worms. which are These pigments readily unite with oxygen to form oxyhemoglobin in the case of the former. and even some annelids. . In the larger and more complex animals where the volume of is such that a more active interchange of gases is required than the general body surface will permit. fishes. flatworms. the more or less plumelike In gills of crayfish are pocketlike outpushings of the body wall. special organs or moditissue fications of the surface must be provided. and turtles. thence to the plasma where much of it combines with sodium Small amounts of CO2 combine with the as sodium carbonate. The gills of most aquatic forms are richly supplied with a capil- lary supply of blood and then membranous surfaces are directly absorbed. worms. the rectum serves as an accessory respiratory organ. When the oxygen pressure of the surrounding tissue is sufficiently low. Likewise. this gaseous exchange is made by almost direct diffusion through the cell membranes to the surrounding medium. carried in a blood vascular system all over the body.

There are also nerves from the lungs themselves which extend to this center and contribute to the maintenance of the proper rhythm.392 Aerial respiration is TEXTBOOK OF ZOOLOGY accomplished in terrestrial animals through In insects a special internal surfaces which must be kept moist. The blood circulatory system centers in a contractile heart from which tubular arteries lead out to various organs of the body where they branch into minute vessels or capillaries. The capillaries converge as they carry the blood away from the organs to form the veins which carry the blood back to the heart. The blood is composed of the clear fluid. and it is a specialized surface derived from the anterior or pharyngeal portion of the digestive tube. This is a closed system of vessels. The muscles which control these actions are automatically stimulated through the nervous system to contract when the carbon dioxide level of the blood reaches a certain point. is affected by the carbon dioxide and determines the rate of respiratory movements. The circulatory system is a closed system of vessels supplying all parts of the body with blood and a system of spaces. located in the medulla oblongata. hemoglobin. sinuses. and made spongy by the innumerable small air sacs which provide the enormous respiratory surface necessary. Abundance tory action. The real lung is a development found in the terrestrial vertebrate. the area of it would be more than 100 square yards. as are also the tracheae of insects. such as birds and mammals. It has been estimated snails the "lung*' is simply that if all of the average of these pitlike alveoli of the internal lining of the lungs human being were spread out in an even surface. A respiratory center. In higher vertebrates. The red corpiiscles contain the red pigment matter. In pulmonate an invagination of the skin. which open through spiracles along the sides of the body. and the Uood corpuscles. The Circulatory System. and vessels collecting lymph from the various organs to return it to the blood vessels. they are extensively lobed. of venous blood stimulates an increase of the respira- moisture and give In addition to exchanging gases the lungs also discharge off a certain amount of heat. The mechanism for the accomplishment of breathing in the cat and other mammals by the use of the diaphragm and thoracic wall is described in the chapter on mammals. plasma. system of branched tubes called tracheae. distribute oxygen to and receive carbon dioxide from all of the cells of the body. which — .

These cells may make their way among cells of other tissues Veins from upper. Upon exposure to air the dissolved fibrinogen in the blood becomes fibrin and forms a clot which is semisolid and blocks flow of blood from most wounds. The white corpuscles or leucocytes are of several varieties and they are amoeboid in character. Due to this sub- stance. The oxygenated Fig. — Mosby Company. Veins from lowcp papt of Body ArtGplo3 to lowcp par>t of Body Lympfiaticj — Diagram of circulation of the blood in a mammal. The lymphatics are the black (From Pettibone. V. irregular lines. the cells have oxygen-carrying power. part of Bcxjy Arteries to upper* part of Body Lymphatics Thoracic duct ^uporiop vena cava 'PulmonaP3^ artary Pulmonapy vein Ui^ht aupicla — — Left auricle Infcpiop vena cavavcntpiclG . blood is shown in black the venous blood in white. 219. . Lymph . published by The C. Physiological Chemistry.Left ventpjclc fli'^ht Lacteal^ — Hepat ic vain.) where they engulf bacteria and foreign matter.THE VERTEBRATE ANIMAL was mentioned in connection —SUBPHYLUM VERTEBRATA 393 with respiration. The remaining' fluid after the blood clots is called serum.

as cyclosis. which cyanin. always in the heart. the necessity of increased food distribution is cared for by branching of the gastrovascular vision necessary except cavity into diverticula. but the necessary exchange and movement of food materials. such as planaria. and the blood is circulated by the action of a single heart. simple. Here a closed system of vessels forms a complete In circuit to carry a circulating medium to all parts of the body. the blood is very unequally distributed. Circulation. Another fourth is held in the hepatic portal system. and it serves to circulate the food vacuoles. a fluid similar to plasma which has seeped through the walls of it carries amoeboid white Certain of them are produced in the lymph glands. because it bears no red this group the fluid is known as hemolymph The hemoglobin is borne in the fluid. The amount of blood in a mammal is approximately one-twentieth of the body weight. In most Protozoa there is no special arrangement for this function.394 is TEXTBOOK OF ZOOLOGY the capillaries in the various organs. however. The spleen is a lymphoid organ in which debilitated red corpuscles are disintegrated and the products placed in the blood. is carried in the red blood In molluscs and some crustaceans there is a similar corpuscles. The plasma contains enough inorganic salts to taste slightly salty. When the body Its salt content is about equal to that of sea water. saccular forms like hydra there is no proan exchange of the water in the gastrovascular cavity. the . as was studied in the earthworm. large arteries. and corpuscles. — Transportation of materials through the protoplasm and from cell to cell of is of a single cell or a single-celled organism the metazoan a fundamental function among living things. rials Vertebrate blood is largely water carrjdng dissolved mate- and suspended corpuscles. and lungs. The hemoglobin. an iron compound. or in the average man a little more than a gallon. is called hemo- Instead of iron. the liver and its sinuses. corpuscles. respiratory pigment carried in the plasma. copper is the principal constituent of this pigment. In double-walled. transporting materials. is One-fourth is active. of which Paramecium is an example. veins. there This is known is a definite course of movement by the endoplasm. stances. The fluid part is known as plasma. In flatworms. waste sub- and gases is accomplished by simple diffusion of materials. In a few forms. contractile parts is A In sponges the wandering cells assist in distinct system of tubelike vessels with developed in the annelid worms. The vertebrate system is closed.

000.000 to 10. The acrial in the tissues. then.000 red corpuscles (erythrocytes) per cubic normally about millimeter of volume in the male and about 4. antithrombin. others dissolve precipitated. Each erythrocyte is essentially a little capsule filled with hemoglobin which is a compound peculiarly fitted to unite with atmospheric oxygen. Thrombin reacts with fibrinogen to produce fibrin. the bacteria. When united with oxygen it is known as oxyhemoglobin. the blood cells and platelets produce a substance. essential information concerning blood The plasma antibodies. The average person. neutralizes the antithrombin.000. cephalin. One of their chief functions is the destruction of bacteria and other foreign mateThis process is known as phagocytosis. companying table summarizes cells. allowing the formation of thrombin.THE VERTEBRATE ANIMAL —SUBPHYLUM .000 in the female. weighing 150 pounds. Howell 's theory of coagulation of blood holds that there is also an inert substance.000. of the blood contains a group of substances called These have been produced by various tissues of the body upon contact with certain foreign proteins. the solid fibers of the clot. The plasma normally contains a soluble protein. substances bring about the clumping or agglutination of foreign bacteria. and still others cause them to be The chemical nature of these bodies is not yet known. They are amoeboid and therefore not confined to the blood vessels. principally in the plasma and returned to the lungs. The leucocytes or white corpuscles are quite variable in form and number from 6.000. which prevents the activation of the prothrombin of the plasma to become thrombin.000 per cubic millimeter. they stimulate the body tissues to the production of specific protective antibodies and physicians have come to make use of these antibodies in sterile serum for preSome of these antigen vention and treatment of several diseases. in the presence of calcium. which is readily reduced to give up the oxygen to the cells when the blood reaches The carbon dioxide given off by the cells is collected the tissues. Human blood 5. Since bacteria and pathogenic Protozoa react as foreign protein. and any wound is likely to be fatal. The rate . which. would possess ap- proximately 20. called fibrinogen and calcium in solution. VERTEBRATA 395 is skeletal muscles require another fourth and the remaining fourth contains distributed through all of the other organs. When blood is shed and exposed to air.000 (20 trillion) of them. There are individuals known as hemophiliacs or bleeders whose blood will not clot.500.

396 TEXTBOOK OF ZOOLOGY Average Characteristics of Human Blood Cells KINDS OP CELLS AND .

finally. and mammals. followed by the mesonephros which is the dominant func- kidney. and. The flickering movement of the cilia in the cell gives the appearance of a flame and moves the accumulated excretion down the tubule.^A certain olism is result of the oxidation necessary for metab- the production of end-products which are not only of no further use to the protoplasm but may be a distinct menace to the welfare of the organism because of their toxic effects. The quantity of water which passes through the protozoan in twentyfour hours is several times the volume of the animal itself. The life tional excretory organ when in its glory. the sole kidney for a time. birds. is SUBPHYLUM VERTEBRATA 397 The former is seldom functional. This arrangement is sometimes called a protonephridial system. The substances are usually dissolved and removed as a waste liquid or occasionally as crystals by special parts of the body. but the latter up to and including the and salamanders. the develop- of the others. This is an illustration of the Theory of Recapitulation which says that each individual in its development lives through abbreviated stages of ment of the metanephros with retrogression the history of the development of the race. Excretion.THE VERTEBRATE ANIMAL as a mesonephros. excretion is accomplished by a system of canals which begins in numerous capillary-sized tubules whose blind ends are composed of individual cells called flame cells. Among sponges and coelenterates diffusion of liquid wastes through the general surfaces of the body to the surrounding water serves for excretion. The nephridial system is found in Annelida and has been studied in connection with the earthworm. and a series of segmentaUy arranged pairs of coiled tubes or . The main excretory ducts open to the surface of the body by excretory pores. The ureter is the excretory duct which leads from the metanephric the functional organ in vertebrates as in frogs history of these animals as individuals includes successive stages as follows: the pronephros. higher developed kidney as found in reptiles. Here a coelomic cavity is present. planaria. In Protozoa this function is performed by general diffusion through the plasma membrane and in many forms by the contractile vacuoles. The waste liquid of the surrounding tissues diffuses into this cell. In an animal like the flatworm. The metaiiephros is the Amphibia. These flame cells are irregular in shape and each bears a tuft of cilia extending into the end of the tubule.

and finally with their enclosed granules are cast out through the membranes of the respiratory papillae. these tubules open into the coelom as nephrostomes. . substances from essential structures of the kidney for taking waste the blood and delivering it to the exterior of the body are the Malpighian corpuscles. This eoelomic fluid is drawn into the canal of the nephridium by the beating of the cilia and is delivered of the body at the nephridiopore of the next segment. care of In the insects excretion is by the Malpighian tubules. below mammals (excepting some fish). or to a urinary bladder in the mammals. and the coiled uriniferous tubules which discharge the excretion through collecting tubules into the ureter at the pelvis This canal leads to the cloaca in most vertebrates of the kidney. known as the to the outside nephrostome. Soluble materials in solution also diffuse through the memtaken branous walls of these structures. makes it seem possible that in vertebrates as well The as in annelids the coelom was once important in excretion. In known as pericardial glands. The fact that in vertebrate embryos as well as in lower chordates. They are bunched in the posterior part of the body cavity and discharge excretions into the intestine at its junction with the rectum. which are considered modified nephridia. wastes accumulate in the eoelomic cavity and are nephridia through the ciliated funnellike internal The excreted moved into the end. each made up of a glomerulus and a Bow- man*s capsule. — The chief excretory organs of vertebrates are called kidneys. amoeboid cells of the eoelomic fluid. and they are thought by some authors to have developed by modification and condensation from segmentally arranged nephridial tubules. although of unit organs. The green glands of crayfish are much more concentrated. even the frog.398 TEXTBOOK OF ZOOLOGY nephridia extend through the wall to the exterior. they are modified nephridia. They function as a pair each opening by a duct on the basal segments of the antennae. These cells wander out into the cavities of the respiratory organs where they coalesce into large masses. The echino- derms make use of direct diffusion as well as intracellular excretion by which excreted materials are taken up from the eoelomic cavity by the numerous phagocytic. Kidneys. mollusks there are both nephridia. and the special cells formed from the eoelomic epithelium.

THE VERTEBRATE ANIMAL The wall of each SUBPHYLUM VERTEBRATA is 399 very thin and readily permits diffusion of water and dissolved materials from the blood into the cavity of the uriniferous tubule on the opposite side of the mem- Bowman's capsule The glomerulus carries arterial blood from the afferent arThe terial branch and discharges it into the efferent arterial branch. A portion of this latter division. and water is excreted by different parts of the The substances which are excreted by the kidney are not formed there. of the ammonia which results from protein metabolism is converted into urea in the liver and is carried in that form to the kidneys where it is removed from the set of capillaries ramifying over the convoluted tubules by a process of true secretion. the water eliminated by the lungs. other soluble materials. consisting of two longitudinal trunks with ganglia distributed along them. is taken Some monium most of the nitrogenous wastes are excreted in the form of amHowever. Water constitutes the largest volume of materials to be excreted in most animals. The kidneys are the most important organs in the excretion of water. and kidneys. and the sense organs which serve for receiving stimuli are usually grouped together under the name peripheral nervous system. uriniferous tubule. this organ. which has few sweat through panting. but are merely removed from the blood by The Nervous System. lies parallel to the spinal cord. ganglia which are groups of nerve cell bodies outside the central nervous system. Water is eliminated by lungs. is proportionately greater than in man. Each ganglion has a connection with the adjacent spinal nerve or cranial nerve as the case — . skin. nerves extending to all parts of the body. salts and some free or combined amino acids. glands. soon spreads into a capillary network which surrounds the latter convoluted portions of the uriniferous tubule. In the dog. the urine which consists of urea. various salts. except in some desert forms where water is conserved and the excretion is in crystalline form. In man the quantity of sweat dis- two or three liters a day. alimentary canal. brane. and the amount they eliminate is inversely proportional to the amount charged may amount to excreted by the skin. According to this idea. and constitutes the sympathetic system. The nervous system in this type of animal is composed of a hrain and spinal cord forming the central nervous system. Most of the water to be excreted from the blood in Malpighiaji corpuscles.

the relationship of parts in regard to function is similar to what has already been seen in the higher nonchordate animals. — . 9. . (From Zoethout. cut end of spinal nerve root of spinal nerve 8. dorsal . Each spinal nerve has two roots where it A dorsal root receives fibers from sensory end- ings and therefore conducts impulses toward the cord. sliowing a simple reflex circuit. efferent i. might This system controls the involuntary muscles. 1. Textbook of Physiology. joins the spinal cord.400 TEXTBOOK OF ZOOLOGY be. The pe- ripheral system includes ten to twelve pairs of cranial nerves from the brain. In gencells. its 5. located in the spinal ganglion nerve fiber 6. ventral root of spinal nerve dendrites of motor nerve cell body in gray matter of the cord. and ten to thirty-one pairs of spinal nerves in different forms of vertebrates. sensoi-y surface of skin 2. . which is the simplest kind of a nerve conduction circuit. The ventral root of each of these Fig'. after Morat. cell of origin. pass cells in the cord to The impulses. . eral. . V. voluntary muscle 7. published by The C. 220. Cross section of spinal cord and roots of spinal nerves. The reflex arc. This root has a spinal ganglion located on it. is set up by the connectives from the sense organ or receptor to the gray matter of the cord and then the return connection from the motor nerve cells over the ventral root to the muscles. afferent nerve fiber with S.) . Mosby Company. nerves carries fibers extending from the motor the motor end plates on the voluntary muscle therefore. from the spinal cord to the muscles over these roots. .

is a highly developed organ. (From Zoethout. sound waves.-y\/. 221. It is constructed on the plan of a camera with the eyeball forming the light-tight box. Beneath the sclera is a black. The eye. the cornea. It lies as a lining of the inside of the posterior half of the cavity of the eye and is connected directly with the brain by the optic nerve.THE VERTEBRATE ANIMAL SUBPHYLUM VERTEBRATA 401 A ing. The pupil appears black because there is no light behind it.n. chemical changes. pigwhich continues anteriorly the colored part of the eye. high development of sense organs for the senses of sight. transparent lens whose surfaces are curved to bend the rays of light in such a way as to focus them on the sheetlike retina behind. cm. lens. The wall of this is composed of an outer fibrous sclera (white of eye) which continues anteriorly as a mented and vascular as the iris. hearand touch is characteristic of vertebrates. a. ciliary process. chorioid coat. s. transparent front. the chorioid. optic nerve.Vw crjo. c. ciliary muscles. The retina is a lateral extension of the brain and is the sensory part of the eye. v. The iris is like a curtain layer.p.) surrounding a space at the anterior surface of the eye and this space between its medial margins is the pupil.. The . sclerotic coat. r. iris. fovea centralis. i. The organs are receptors and they are stimulated by changes in external environ- mental conditions. vitreous humor. c. cornea. Textbook of Physiology. ch. such as light. which is the organ of sight.L. Behind the pupil is a. published by The C. Diagram of the eyeball. — Mosby Company. I. The general cavity of the eyeball is divided into some chambers. suspensory ligaments.. Fig. taste. C. sc. and contact. retina: /.. op. smell. aqueous humor. V.

. 402 TEXTBOOK OF ZOOLOGY external or aqueous chamber between the cornea and the lens. scala tympani containing the three auditory ossicles Eustachian tube. vertebrates. cochlea. then aqueous humor. The This makes possible an adjust- curvature of the lens can be controlled by the action of the ciliary muscle which encircles its margins.) . they tend to lose this accommodation because of loss of get older The tension on it due to the attachment of elasticity in the lens. external auditory meatus. the inside of the eyeball by the ciliary process tends to hold it out of the eye to near Fig. scala vestibuli. o. Diagram of a section through the right ear. (From Zoethout. B. . is filled with aqueous humor. in a flattened condition. 8. A ray of light enters the eye by passing through the cornea. with the iris extending into is it. Eyeglasses are used by older people to supply this lost phase of accommodation. round window Pt. V. Mosby Company. . lens. pupil. oval window (fenestra ovale) r. Yt. — . This focuses the eyes very well on distant objects but does not provide the necessary curvature of the lens to bring near objects in focus.. vitreous humor and then to the retina where the sensory cells are stimulated and the impulse carried to the brain by the nerve fibers of the optic nerve. membrana (fenestra rotunda) below r is seen the tympani. is This cavity subdivided by the iris. . Textbook of Physiology. after Czermak. published by The C. semicircular canal P. tympanic cavity a. 222. T. Behind the lens the large internal or vitreous chamber which is filled with a jellylike vitreous humor. ment and far objects and particularly so in higher As people This power is known as accommodation.

which serves in catching and directing sound waves within. The external ear is still further limited to reptiles. The sound waves which stimulate the sensory cells of hearing enter the external ear and set up vibrations in the tympanic membrane. incus. and the dorsal portion related to equilibrium. These are in turn transmitted by the ossicles to the fluid endolymph within the labyrinth. thus giving a sense of position. there are two vertical canals. The sense chamber. epiglottis. The middle ear is a space beneath a tympanic memhrane which separates it from the external auditory canal. rial of smell is centralized in the epithelial lining of the nasal Special olfactory cells are stimulated by particles of mateair dissolving cells. and mammals. This organ consists of an external ear. birds. brates while the cochlea The latter are common to all verte- is limited to Amphibia and higher classes. which contains the sensory cochlea with its organ of Corti for hearing. containing ossicles. These cells are connected with the brain by way of the auditory or eighth cranial nerve (Fig. which are concerned with equilibrium rather than hearing. and . join the body of the utriculus in as In the higher forms many different planes as there are canals. one anterior their planes at right angles to each other. and one posterior.UM VERTEBRATA 403 The ear structures provide most classes of vertebrates with facilities for two functions: hearing and equilibrium. from the on this membrane and making contact of taste is with the sensory is The sense similar except that it located in sensory cells in taste buds on the tongue. In this cavity are three bony ossicles. which transmit the sound vibrations from the tympanic membrane to the membrane The memover the fenestra ovalis leading into the internal ear. and the inner ear. 222). in which the sensory cells are supported on the organ of Corti stretched across it. the fluid in the canals changes its level and position to stimulate the sensory hairs. The vibrations of the fluid extend through the cochlea.THE VERTEBRATE ANIMAL —SUBPHYI. the malleus. branous labyrinth inner ear. At one end of each canal there is a bulblike swelling or ampulla which contains sensory hairs. with and one horizontal canal. a middle ear or tympanum. When the position of the head is moved. and the semicircular canals. and stapes. is the name often applied to the chambers of the is Its ventral chamber the sacculus connected with the is organs of hearing. the utricidus which is forms and the three in The two semicircular canals in simpler higher.

I 404 lips TEXTBOOK OF ZOOLOGY (and barbels of some vertebrates). it reaches the taste cells. Irritability and conductivity are fundamental functions of all protoplasm. In Annelida and Arthropoda a modified ladder type in which the two longiganglia have fused along most of the midventral tudinal cords of Toward the anterior end. Such a system was studied the nervous sj'stem is in planaria. The protozoan organism has only neuromotor apparatus and depends largely on the primitive properties of irritability and conductivity to guide its activities. there are scattered nerve cells connected with each other form a nerve net. each cord lying lateral to the digestive tract with transverse connectives and predominant ganglia at the anterior end. ladder. the cords separate at a paired ganline. In the final analysis. Every part of the receptor surface of such an organism is in physiological continuity fibers to cells by with every other part of the body. The neuroepithelial or neurovmiscular which make up this continuous net through the body are the forerunners of the typical neurone and are called protoneurones by Parker. such as the co- elenterates. In the net system there is no central exchange and no specific path of conduction. effects of previ- possesses the power up the ous stimuli. Nervous Function Reception and Conduction. but it — — Living protoplasm to record or store is not only excitable. A protoneurone transmits in every direction while a true neurone transmits in only one. . Next comes the linear type of nervous system in the form of a It is composed of an organization of neurones into a double chain of ganglia. of this type. In the simpler Metazoa. of food The particles come in by way and drink and as the material dissolves. organisms to change of conditions both externally and internally determines their behavior. whether The responsiveness of it be in the body of an Amoeba or a man. and pressure sense organs are located just A few of the pressure sense organs are found in certain of the internal structures of the body. The lateral line system in fishes is sensory to vibrations carried in the water and is quite important to aquatic animals Most of the tactile beneath the skin over different parts of the body. the perceptions and reactions of man are but expressions of these primitive functions in a more specialized organism.

The reflex arc and reflex actions illustrate the simple form of nervis ous conduction circuit. starfish for example. In its simplest form the reflex arc com- . —SUBPHYLUM VERTEBRATA 405 the siil) esophageal ganglion. is The fact that the central nervous system of verte- dorsally located and hollow has been brought out previously. On the other hand. and each of these units must conduct nerve impulses in its normal function. Each neurone is a nerve cell with processes extending from it. the nervous system shows a progressive increase in complexity. At room temperature the sciatic nerve of a frog will transmit a nerve impulse at the rate of about 100 feet per second. but since there it only slight increase in temperature during the change. seems not to be a typical metabolic oxidation process not to fatigue the nerve fiber. . and radial branches extend into each arm. and encircle the alimentary canal to join on the dorsal side as the pair of siipraesophageal ganglia or ''brain. impulse or is a result of it rather than the impulse The speed of different of electrical transmission has been measured in a number animals and nervous transmission is much slower than electrical. but charge follows the wave apparently accompanies the itself. the thorax have undergone considerable fusion.THE VERTEBRATE ANIMAL glionic enlargement. measurements of the rate of conduction in man show a velocity of about 400 feet per second. furthermore. the activity seems An electrical it of activity along the nerve fiber. The exact nature of the nerve impulse is still somewhat of a question. Conduction over nonmedullated fibers of invertebrates is much slower than this. Branches from these communicate with the sensory structures of the skin and tube feet. the central group of ganglia makes up the circumoral nerve ring around the mouth. at least cer- partially a chemical change in which oxygen is necessary tain and a is amount of carbon dioxide is produced." In Arthropoda the ganglia of the In Echinodermata. Even within the the group of vertebrates. Concentration of the tissue of the nervous system into definite organs brates is carried farther in vertebrates than in the less highly or- ganized forms. The neurones have been referred to before as the units of structure and function of the higher type of nervous system. from worms to man. It is thought to be transmitted as a metabolic change This is passing along the nerve fiber (axone or dendrite). The highly developed brain of mammal is the climax of this tendency.

The cereassociations. individual actions included in walk- breathing. be conducted in either direction by the fiber but can cross a synapse only from axone to dendrite. It also regulates digestive secretions. of speech. Functions of the Divisions of the Brain. and others. visceral organs. This is known as a reflex of the first level. Functions of the Spinal Cord. Below and behind the cerebellum is the medulla oblongata which controls breathing and may be an bellum is inhibitor on heart action. — memory The diencephalon serves as a center for sponThe midbrain is one of the centers of coordinated taneous actions. and vasomotor activity of the blood . reflex centers — This organ serves as a system of which control the actions of glands of the trunk. The impulses from the muscles. and skeletal muscles. or shivering or formation of goose flesh. thus serving like a valve in a pipeline. for these come in contact only by a synapse which brings them It has been found experimentally that nervous in close proximity. it is usuallymore complex. tendons. particularly with reference to equilibrium. as with- no protoplasmic union impulses may drawal from unexpected pain. movement which has to do with posture and eye muscles. because it returns the motor impulse over the motor fibers of the same nerves which brought in the sensory impulse.406 TEXTBOOK OF ZOOLOGY posed of one motor and one sensory neurone. Reflex actions may be in the form of motion. another center of coordinated movement. There between the axone of the sensory neurone and the dendrite of the motor. The spinal cord is also a nervous pathway between the brain and numerous organs of the body. movements of digestive organs. and semicircular canals of the ear are coordinated so that in a movement or posture the proper muscles may be contracted to the proper extent at the proper time. Still other reflex actions include secretion by glands. The classical example involves the spinal cord and a spinal nerve. joints. Conscious sensations and intelligence are centered in the gray matter or cortex of the This section controls voluntary actions and provides cerebrum. movements ing. or the contraction of the pupil of the eye with increased light intensity. however. It is said that more than half a million neurones join the cord through the dorsal roots of the spinal cord. The motor axone carrying cell in the impulse from the motor nerve the gray matter usually is j ends in a muscle cell or a gland.

The sexes are almost universally separate. and is later born alive. the coordinator of the The Reproductive System. in those in which the egg is retained in the body and the embryo develops there. Reproductive Function. Therefore. SUBPHYLUM VERTEBRATA 407 As a Avhole. rial — —A living organism is in numerous ways similar to a machine. The distinct gonads develop to produce special germ cells. Our present conception is that the protoplasmic sub- stance of the but a continuation of the specific protoplasm peculiar to an earlier individual or in sexual reproduction to two individuals. the brain serves as the organ of coninuinication is between the sense organs and the body and bodily activities. New organisms all arise from preexisting organisms of the same kind. and here too the young are born alive. The process of cell divi- by existing organisms processes known in sion is the fundamental basis for all reproduction. The female gonads are ovaries. The vertebrate reproductive system shows a fairly high degree of development. and in the forms in which the fertilized ovum is retained in the uterus. The males of some classes possess for use in copulation certain accessory organs which tend to insure fertilization. or from the dead bodies of plants and animals. and they produce spermatozoa which are carried from the gonads by the vasa deferentia. the embryo being nourished by diffusion of nutriments from the blood of the parent. The vertebrates which lay eggs are spoken of as being oviparous. For centuries before the invention of the microscope it was commonly believed that living things arose spontaneously from nonliving material. . Certain old books carry directions for the artificial generation of mice or bees. with the exception of some cyclostomes. They are carried from the body by oviducts. under ordinary circumstances the structural and physiological complexities which arise through individual is new embryonic development must be generally similar to those of the predecessors. The male gonads are testes. Louis Pasteur did as much as anyone to discredit this idea of spontaneous generation.THE VERTEBRATE ANIMAL vessels. the condition is known as ovoviviparous. feeding on the yolk of the egg. the condition is said to be viviparous. and they produce ova or eggs. In vertebrates the possible offspring produced each season by a single individual varies from one to thousands. but reproduction of new units of living mate- is hardly comparable to any mechanical our industries.

Sexual Reproduction. volvox may reproduce for several generations by cells for example. planaria. there are no specially organized gonads for the production of germ cells. the colony asexual division of the individual but sooner or later the cells of the colony become specialized into conjugating individuals. both being formed by aggregations of formative or interstitial cells between the ectoderm and endoderm. but as a rule the germ cells are produced in such organs set apart for this purpose. After the seasonal production of germ cells is completed. Protozoa as independent individuals. a distinct sexuality exists Examples of asexual reproduction by budding and Paramecium. Previous studies made on the reproduction of hydra have brought out that the gonads are temporary. have al- fission ready been pointed out in the studies of reproduction of sponges. and even in tunicates. In the simplest of Metazoa. may reproduce also by sporulation. by which process the forms a protective cyst and by a series of simple divisions (fragmentation) the internal protoplasm breaks into a number of smaller Following this the cyst ruptures and releases these new units units. reproduction among protozoans lished is taken to be asexual. The ovary produces mature or nearly mature ova and the testis produces mature spermatozoa. In such forms it is possible to see foreshadowed sexual reproduction as it is known in Metazoa.408 TEXTBOOK OF ZOOLOGY In most of the single-celled organisms reproduction may occur by such equal division of the protoplasm (binary fission) that the new cell individuals cannot be distinguished as parent and offspring. hydra. In certain is . Both ovaries and testes are present. Even in these true hermaphrodites cross-fertilization insured by copulation or union in such a way that the spermatozoa of one individual fertilize the ova of another. For the most part. a few higher ones. Hermaphroditism is the condition in which the same individual produces both ova and spermatozoa. as in sponges. In flatworms and annelid worms the gonads are permanent structures of the mesoderm. and rarely among normal vertebrates. It occurs principally in the simpler Metazoa. the gonads disappear. In some forms this goes to the extent of certain cells becoming distinct gametes with male and female characteristics. —In certain of the colonial Protozoa. but according to a recently pubin work by Sonneborn.

female. and the ova are fertilized within the genital tract of the In birds and most reptiles after the addition of nutritive and protective coats the egg passes to the outside to develop and hatch (ovipall arous animals) but in the mammals. This act is known as amphiplexus. In animals like the toads and frogs. The majority of bisexual fertili- make a still greater provision to insure zation of the ova by copulation or coitus. vertebrates. At the time of breeding the mature spermatozoa are delivered to the cloaca or vagina of tha female. except monotremes. It will be remembered that the first and second pairs of abdominal appendages of the male crayfish are modified for transferring spermatozoa into the seminal receptacle of the female. In the females of viviparous mammals the posterior portions young The internal wall of the uterus and the external embryonic membranes (serosa and allantois-chorion) cooperate to form a placenta through which food. where gether in that the they remain until the eggs are or dioecious animals laid. Here the sexes are each with functional gonads abd ac- cessory structures capable of producing only one kind of germ cells. In some of the types of animals already studied individuals of the two sexes have simply deposited the mature germ cells in the same Under the sections on revicinity and at about the same time. metabolic wastes. This represents a beginning in the development of a copulatory organ. (fish) such a procedure has production in starfish and the bullhead been described. and young are bom as more or developed individuals (vivipa- rous). less it is retained within the uterus during the period of embryonic development. a special provision is made to bring the individuals of the opposite sexes to- male clasps the female and sheds sperm over the eggs as they are expelled from the cloaca. and respiratory gases diffuse between parental and embryonic blood.THE VERTEBRATE ANIMAL —SUBPHYLUM VERTEBRATA 409 other hermaphroditic forms (as some cyclostomes) the spermatozoa and ova of a particular individual are usually not mature at the same time. is Bisexual reproduction the form of reproduction all common to many groups of the higher invertebrates and nearly distinct. . The blood does not pass from parent to embryo of the two oviducts are modified into a uterus within which the are retained and nourished until ready for birth.

The immediate cause of the development by an egg thus stimulated is not known. has a dual function: (1) that of stimulating the egg to develop. Electric and change of temperature are also used. The female produces ova which as parthenogenesis. bees. provided with protective hard shells. starfish. bile salts. and frogs. are produced by the females of this generation After winter is over such fertile eggs to live through the winter. mechanical pricking. Fatty acids. the second is sometimes found as a subclass under . artificial parthenogenesis. There are indications that this is also the case in stimulus. during which period no males are developed. . develop into fall of the new individuals like herself without fertilization for is a whole season. This oxidation may Classification In most recent classifications this subphylum is divided into seven classes however. toluol. ether. saponin. benzol. molluscs. wasps. thrips. chloroform. a few moths. In normal fertilization of an egg by only one spermatozoon. sexual re- < production may lapse for considerable periods of time. These classes are as follows: fish Cyclostomata (si klo sto'ma ta. scale insects. the third. be the cause of the development in the ovum. This known Usually in the year males are developed. and jawless. Fertilization. Parthenogenesis. and fertile eggs. Round-mouthed with only median fins. circle and mouth). is This process common in many smaller Crustacea. Lampreys and Hagfish. solanin.410 or vice versa but tlie TEXTBOOK OF ZOOLOGY necessary materials are allowed to diffuse dis- through the tissue of the placenta in which both systems are tributed. where it occurs. some ants. Such methods have produced artificial parthenogenesis in eggs of sea urchins. and alcohol are other substances which will induce it. moths. annelids. aphids. —In some species of invertebrates. unsegmented notochord. and rotifers. and (2) that of introducing the properties of the male parent. hatch into parthenogenetic females for the next season. Artificial partheno- genesis may be induced in many mature eggs by change of osmotic pressure due to change of salt content in the surrounding medium. it has been found that the rate of oxidation then increases from 400 to 600 per cent.

metal plate and gills). mammary tebrates with hair and with Cats. paired appendages.THE VERTEBRATE ANIMAL —SUBPHYLUM VERTEBRATA 411 Elasmobranchii (e las mo bran'ki i. both lives). Rays. Whales. Monkeys. Seals. erect forms possessing The forelimbs have become wings. Catfish. . Cold-blooded. and Salamanders. persistent notochord. and Chimaeras. Snakes. nonscaled aquatic and terrestrial vertebrates with five-fingered. fishes). Warm-blooded ver- mammary glands for suckling the young. crawling). Bats. . or breast). birds) Warm-blooded. gill respirawith jaws and paired lateral fins. Most of them breathe by gills in the larval stage and by lungs in the adult. Men. Cold-blooded forms which are fundamentally ing by lungs. usually possessing Turtles. Amphibia (am fib'i a. terrestrial. tion. Aves (a'vez. All birds. feathers. Lizards. Frogs. Mammalia (mama'lia. and plaeoid scales. Pish with jaws. Pisces (Pis'es. True fish with bony skeleton. cartilaginous skeleton. Perch. a scaly skin and breathand Crocodiles. Bass. etc. Reptilia (rep til'i a. Sharks. Toads.

and the lack of the oral funnel or sucker. Bdellostoma and Paramyxine of the Pacific. Chapters XXV. four tentacles on each side of the mouth. nets or those dead from natural causes and devour all of the inter•If the frog or toad Is to be used as the laboratory animal representing the typical vertebrate. This name is in contrast Gnathostomata (jaw mouth) which includes all other vertebrates. The body is slender and eel-like in shape. and suctorial. Classification The group is divided into two subclasses (or orders according to some authors) distinguishable by presence or absence of tentacles around the mouth. There are some exoskeletal teeth located on the roof and floor of the mouth and on the tongue. 412 . They all possess twelve or semicircular canal in the inner ear. These subclasses are Myxinoidea (Hyperotreti) including the hagfishes. all : included in one family Myxinidae Myxine of the Atlantic and Pacific These each Oceans.CHAPTER XXV CYCLOSTOMATA* Because of the absence of jaws this group is sometimes known as Agnathostomata (ag nath o sto' ma ta). jawless. only one and a functional pronephros. smooth skin and has only dorsal and to ventral median fins. The mouth of the cyclostomes is round. It is covered with a slippery. ability to produce enormous quantities of mucus. The development of the hagfish does not include a metamorphosis. more pairs of gills. have but they all agree in having a terminal nostril. and the instructor so desires. They usually live in the mud of the sea bottom except when they are feeding either on the dead body of a fish or attached to a live They frequently enter the mouth or gills of fish caught in one. and Petromyzontia (Hyperoartii) including lamprey (or improperly. XXVI. and the number of semicircular canals. divided into three genera specific characteristics. lamprey Myxinoidea or hagfish are which is eel to some). number of gill slits. and XXVII may be omitted until after the study of Chapter XXVIII and then assigned if time permits.

Economic Relations of the Class In a general all way lampreys are both beneficial and injurious. Petro- myzontidae. tridcntatus ciliatus is the southern form. Both return to the fresh-water streams to spawn a few years later. formed. which follows the type name. Sea lampreys and lake lampreys are both valuable as human food. including Petromyzon. The characteristics of the group will be brought out under the discussion of Lamprey as a typical representative. Entosphenus tridentatus trident atus is the northern form and E. especially just preceding the it spawning season. fish frequently and release itself. Subclass Petromyzontia likewise includes only one family. it does little harm newly matured ones are making is to fresh-water fish except as the their trip to sea. fill. it sucks the One will remain to a single fish for about five days. for the two or three years at the expense of spends in the ocean. . and through the hole thus fish's blood. sea In adult the sea lamprey goes to the open and the lake lamprey goes to the deep water of the lakes. It attaches itself and rasps a hole in the side of a fish about once a month. Ichthyomyzon of the lakes and streams and Entosphenus of the Pacific coast. They serve as excellent fish food and fish bait when they are in the larval stage. THE LAMPREY Habitat All live on or in the muddy bottoms life of fresh-water streams dur- ing larval stages. and they are quite predaceous. Since the sea lamprey does not feed after it starts The up stream. They since they are predaceous and spend their adult lives in the lakes. lives marine fish.CYCLOSTOMATA nal organs 413 living fish and flesh. They frequently attack which have been otherwise injured. its similar except that it spends are very destructive to lake fish The lake lamprey entire life in fresh water. get its dies as a result. the There are several genera common Atlantic form. The lampreys live in both salt and fresh water. attacking fish of considerable size. The sea lamprey. Brook lampreys are classed as wholly beneficial since they feed on microscopic organisms while larvae and do not feed as adults.

—Lateral view of the Pacific lamprey. Entosphenus tridentatus. Evans.) (Drawn . 223. by Titus C.414 TEXTBOOK OF ZOOLOGY SUCTORIAL MOUTH NASAL OPENING \- EYE -GILL CLEFT •DORSAL FIN V -URINOGENITAL PAPILLA CAUDAL FIN Fig.

Along the length of the lateral axis of the body are distributed There are two dorsal fins and a caudal fin. The color is rather variable but might be expressed as being a variegated olive brown. and several bundles of muscular tissue radiate through the wall of the funnel to expand and contract it. Entosphenus trident aUis. and the Pacific form. A large lingual muscle is differentiated for moving the tongue. behind it is the urinogenital opening at the tip of a papilla. Internal Structure The muscular system is quite primitive. plungerlike tongue supported by a cartilage and bearing teeth. The following account will fit them generally.CYCLOSTOMATA Habits and Behavior 415 The animal is a rather inebriate type of swimmer because it is long and slender and does not possess paired fins. In a middorsal position on the head is located the single nostril which leads into an olfactory chamber. They may reach a length of three feet and three inches. holds open. correspond quite closely in structure and make excellent representatives for study of the group. Immediately a poorly developed eye. Petromyzo7i marinus. body wall of the host The annular cartilage supports the margin of the funnel and it Along the margin is a fringe of papillae. The papilla is larger in the male specimens. The anus is located in the midventral line a short distance anterior to the tail. It winds its way through the water and occasionally comes to rest by attaching itself to a rock by means of an oral funnel. This funnel is provided with sensory organs. simply being covered with transparent skin. of zig-zag . but no paired fins. ehitinous teeth used in rasping through the fish. is gill slits along each side of the anterior In front of the gills on each side of the head It has no lid. It is principally a series myotomes along the length of the body very similar to those in Amphioxus. The mouth In the floor of the mouth is a lies at the bottom of the funnel. There are seven uncovered portion of the body. and on ventrally as a pituitary pouch or caecum. At the anterior end of the animal is the mouth with the luccal funnel extending from it. External Structure In most respects the Atlantic lamprey.


dorsoventral bars. The intestine is slender and almost straight. two principal The and anterior cardinal veins located just lateral to the lower side of the notochord collect blood from the body wall and head region. The intestine ends posteriorly at the anus. but it has a slight This is internal fold which extends spirally through its length. lateral bars. Six pairs of afferent branchial arteries carry the blood to the gills where capillaries supply the gill it lamellae. The blood then passes through the sinuauricular valve to the single auricle. and it tends to increase the absorptive surface. and nine much-curved. thence through hulhus arteriosus to the ventral aorta. veins.CYCLOSTOMATA 417 The skeletal system called neural arches. and empty it into the common cardinal vein which extends ventrally to the sinus venosus. The liver is found in the anterior part of the body cavity. and the main aorta passes . except for a transverse bar. The sinus venosus receives also the single inferior jugular and the hepatic vein from the ventral region. not very highly developed because the adult lives entirely on blood and lymph of other fish. supplies the brain region. The blood is passed is The digestive system from the mouth down the esophagus which continues into the intestine at the level of the posterior end of the branchial region. A carotid branch of this artery posteriorly. called a typhlosole or spiral valve. The circulatory system posterior consists of a heart with chambers. The cartilaginous pericardium joins the branchial basket branchial area is at the posterior end. obtained by rasping a hole through the body wall and sucking it out. two pairs of sinuous. and lymphatic spaces. cartilage already There are two auditory capsules near the posterior The buccal funnel is supported by the annular mentioned and three sets of labial cartilages. developed around a nonseg- rnented notochord along each side of which are paired cartilages At the anterior end is a skull whose floor and sides are cartilaginous. They take a meal about once in three or four weeks. part of the skull. capillaries. thence by the auriculoventricular aperture to the single ventricle. arteries. carry dorsally to join the dorsal aorta which made up by their convergence. The supported by the cartilaginous hranchial basket which is composed of a i)air each of dorsal and ventral longitudinal bars. is cartilaginous. while the roof is membranous. The anterior one of these is not in contact with a gill aperture. The efferent branchial arteries is collect this blood.

arteries. R. of lamprey. Evans. RENAL V. 4TH. JUGULAR VEIN V. (Drawn by and veins Titus Arrows of C. —Diagram indicate direction of flowview blood. VENTRICLE SINUS VENOSUS ATRIUM HEPATIC V. RENAL A. 225. EFFERENT BRANCHIAL A. AFFERENT A. — DORSAL AORTA 5TH.R. Fig.) 418 TEXTBOOK OF ZOOLOGY There is giving branches to the viscera and body wall. When the . BRANCHIAL VENTRAL AORTA INFERIOR JUGULAR R. The seven pairs of gills cipal features of the respiratory and respiratory tube constitute the prinsystem of this animal. INTESTINAL V. CAUOAL VEIN CAUDAL ARTERY of oblique ventrolateral of heart. INTESTINAL A. giving one part to each posterior cardinal vein. the caudal vein simply divides. R. POST CARDINAL V. no renal portal system.

as it is not entirely closed over. The roof of the brain is rather membranous. — OLFACTORY SAC OLFACTORY LOBES PINEAL EYE RIGHT CEREBRAL HEMISPHERE OPTIC NERVE RIGHT GANGLION HABENULAE DIENCEPHALON PITUITARY POUCH OCULO-MOTOR PLEXUS OPTIC LOBES N. insignarrow bandlike cerebellum just behind the optic lobes. through them. —Brain of lamprey. under the velum. the water is drawn into the respiratory tube through the gill slits and then discharged apertures. cerebral hemispheres closely fused to preceding. Lateral view Evans. (Drawn by Titus C. The nervous system shows the development of a brain. This continues directly posteriorly as the flattened spinal cord. brain.CYCLOSTOMATA 419 animal is not attached to a host.) . through the respiratory tube. The sense organs include with its nificant . While the lamprey is attached to a host fish. water may be drawn through the mouth. 226. dorsal view. CEREBELLUM FOURTH VENTRICLE AUDITORY SAC AUDITORY NERVE MEDULLA OBLONGATA NOTOCHORD VAGUS N. single dicncephalon dorsal epiphysis. and the medulla just posterior to it. small. MIDDLE CHOROID TRIGEMINUS N. primitive which possesses all five principal divisions of a vertebrate it is From anterior to posterior composed of olfactory lobes. Fig. midbrain with a pair of optic lobes. through the paired gills and to the outside through the seven pairs of external The blood in the gill capillaries is aerated from the oxygen carried in the water as it passes over the gill lamellae.

supported by cranial cartilage. under lip. oral hood. . ANC. it consists simply of a vestibule and two vertical semi- circular canals. eye. anterior end of notochorcl CC. O. velum.. but hermaphroditic con- ditions are occasionally found. ventricle of heart. dorsal aorta F. (Courtesy of Albert E. caudal fin DA. OH OP cc vf a. extending from tip of upper lip to a point slightly anterior to end of notochord. folds in intestinal wall. . to the peritoneal lining of the body cavity. and sight is not used extensively by The audiis tory organ. . VN. OL. VA. upper lip. ear. The rib- one at each side of the notochord and just dorsal SC AU HB OA N PfJ VN OE MM M LP-. . . intestine L. . showing pronephric tubules with their ciliated funnels (nephrostomes) SC.) . fair development. The sense of taste centers in taste buds located in gill slits the respiratory tubes between the and possibly near the inner margin of the buccal funnel. . pineal body. esorhagus OH. . 227. liver LP. only for equilibrium . spinal cord. ventral aorta. M. PC. olfactory organ. OP. This is located just posterior to the rectum externally by the urinogenital papilla just behind the anus. N. oral papillae. MA''. AU. which does not include an organ of hearing. suggesting a possible origin of the spiral valve FB. pericardial cavity PN. pronephros. hind brain I. branauricle of heart. The urinogenital system shows only bonlike kidneys P lie. The eyes of this animal are not it. Fig. 420 TEXTBOOK OF ZOOLOGY is the single nasal chamber which the brain. Entosphentos tridentatus. located immediately anterior to Extending ventrally from the nasal chamber and projectis ing beneath the brain to end blindly just above the esophagus pituitary pouch. A mesonephric duct ex- tends posteriorly along the free edge of each to join the small urinogenital sinus. T. notochord. BA. . Galigher. duct connecting pharynx and thyroid cartilage. hepatic vein. position in which mesonepliros will develop. cranial chial arteries (afferent) G. anus. the As it passes beneath the diencephalon it makes contact with the infundibulum. P. UL. Ammocoetes larva of the lamprey. forebrain G. body muscles (myotomes) MB. Germ cells when mature are discharged from the gonad into the body cavity and go by way of two . . midbrain.jc t c VA V A G L F AN CF A. Inc. and opens The single gonad is rather large and is suspended by a peritoneal fold into the coelom. V. . . . where it divides to form two lateral rods CF. gall bladder HB. AN. OE. — . highly developed. VE. The sexes are presumably separate. thyroid.

The eggs first stick to objects. b. At 20° C. in about six hours when the optimum temperature of 22. c. The adults spawn but once and then die. then out through the papilla to the environmental water where fertilization occurs. e. this division requires nine days. The sea lamprey then migrates to the ocean and the lake lamprey moves down stream to a large fresh-water lake. hatches from the egg and lives from four streams where the eggs are laid.5° C. They both become parasitic on other fish and continue this existence for from one and one-half to three and one-half years. history may be summarized as follows: The eggs which contain considerable yolk in the vegetal portion and are about one millimeter (%5 inch) in diameter are laid in fresh-water streams. to five years in the mud along the five years the ammocoetes undergo meta- or August to They remain under the mud from July February or March while undergoing this transforma- tion to adult condition. A tadpolelike larval form. d. . prevails. life The a. then fall in the sand. when they return to fresh-water streams to breed again.CYCLOSTOMATA 421 genital pores into the iirinogenital sinus. At the end of four or morphosis to become adult. ammocoetes or mud lamprey. Cleavage follows. They are fertilized in the water almost immediately after laying. usually between March and June for all kinds of lampreys.

The Elasmobranchs have no operculum or Classification air The classes. The apertures. possess laterally located gill and are active swimmers. and dwell on the bottom of is This subclass usually divided into two orders. there are first. Subclass Selachii. unpaired or median fins. and the exterior is covered and protected by placoid scales. The males have a modification of each pelvic fin known as a clasper which is circular. cylindrical in shape. except for the or spiracle. but they rarely attack man unless the person is already wounded. and man-eater shark {Carcharodon carcharias [Linn]) are forms com- monly found. class is divided into two rather very easily distinguished subto The is first group is common American shores and the second rarely seen in our waters. are less active. Orders Euselachii and Cyclospondyli. used as a copulatory organ.CHAPTER XXVI ELASMOBRANCHII* Unlike the cyclostomes. •In collaboration with Miss Mary Fickling. are slitlike instead of The gills are supported by gill arches. The skeleton is entirely cartilaginous. and the mouth has an upper and lower jaw. which are dorsoventrally flattened. 422 . the Elasmobranchs are covered with scales and have two body. gill sets of paired fins on the ventrolateral surfaces of the In addition to these. bladder. There is present in the ileum of the small in- a spiral valve which increases the internal surface. rays. The majority of sharks are carnivorous and active. The sharks make up The dogfish sharks (Squalus acanthias and others). and the ventrally located the sea. cub shark {Carcharias platydon [Poey]). — This group includes the sharks which are slits. The mouth testine is not right at the anterior end of the body but is ven- tral or subterminal. — these tiger shark (Galeocerdo arcticus [Faber]). as seen in the lamprey. thus add- ing absorptive area. orders. shovelhead or bonnethead shark {Reniceps tiburo [Linn]. possess gills. there is a partially persistent notochord.

—Southern human sting ray. The former has been considered sufficiently interesting to warrant fureating beings. 228. ther discussion of it as an example of the class. The shape of the head of each is about the shape ascribed to it by the common name. a common form in the Gulf of Mexico. The shovelnose (bonnethead) and hammerhead sharks are very interesting forms. Their natural food consists principally of Crustacea. . squids. Dasyatis americana. and refuse. small fish.ELASMOBRANCHH 423 The average length of most sharks commonly observed ranges between three and six or eight feet. In the Gulf of Mexico and other warm seas the so-called man-eater may occasionally reach a length of thirty feet and is sometimes charged with Fig.

with a saw five feet in length. but they bear their young alive and tend to have a smoother skin. The rays are more numerous in the warmer waters. — This "These is dorsoventrally flattened fishes in ventral side. ackleyi are arous. Pristis pectinatus a sharklike ray with a long tooth-bearing rostral process or snout that resembles a double-edged saw. of skates. ring a doorbell. The saw-fish. Fig. has attracted considerable attention because of its ability to generate and store electrical energy in the muscles of the bases of the broad pectoral fins. The torpedo ray of family Torpedinidae. They have hornlike quadrate in outline and of considerable size. or light a . common ones. a common bottom feeder. B.424 TEXTBOOK OF ZOOLOGY Order Batoidei the broad. — Skates and Rays. These electric organs are capable of discharging suffi- cient current to paralyze other animals. diaphanes. and B. — Butterfly ray. of which Raja erinacea. The skates are distributed along our Atlantic shores and are ovipThe eggs are enclosed in dark brown cases or capsules. There are about six species processes extending from each corner. These animals may reach a length of fifteen or twenty feet. 229. The rays are of similar shape. flat. which the gill a group of depressed or slits are located on the anal fin fish lack and the is caudal is absent or reduced.

The skins of certain sharks and pointed. Shark skins are now being manufactured into leather on a commercial basis. squid. as often the only species taken from the waters of the coasts of is North America. Large quantities of oil are extracted from some of the sharks. as the cub shark for exskates. are quite common in the water of the Gulf of Mexico. Along the They also coasts of California and in the Gulf of Mexico both sharks and rays are a nuisance to the seining fisherman. slender. called) is Psychichthys (or Chimaera affinis. too. is a broad-bodied form with an exceptionally short tail. — This group contains an order with three affinis modern genera. toothlike scales. This oil is used in currying leather in the tanning industry. The average width of those usually seen is from eighteen inches to two feet. shrimp. Bathers particularly dislike "stingaree" infested beaches as well as those infested with the less common torpedo ray. Pteroplatea micrura. and then only rarely.ELASMOBRANCHn flashlight bulb. whiplike tail with a strong spine or sting on the dorsal side of its proximal third. The butterfly ray. Subclass Holocephali. They have a long.000 in Massachusetts alone. They. 425 The sting ray or stingaree is very common in the Gulf of Mexico. gear by tearing through nets. Chimaera monstrosa species which is found in South American waters. "With a sudden swing of the tail one can inflict an ugly and extremely painful wound. It is called butterfly ray because of the manner of flapping the lateral expansions about as a butterfly moves its wings in flight. Some people become severely ill as the result of such a sting. crabs. . Dasyatis sahina and Dasyatis americaiia are two common forms. another Economic Relations of the Class Many of the smaller sharks. like Squaliis acanthias canis are very destructive to lobsters. and Mustelus and valu- able fish which they use for food. The sting is usually obsolete in this form. The sting rays or "stingarees" which are armed with the barbed stinging spine on the proximal portion of the tail are generally common in most of the warmer fishing waters. which have the sharply wood and other materials and is called shagreen. are used as a polisher of ample. in this damage much fishing It is estimated that the damage done way averages $400.

the tail is described as heterocercal. It apparently makes a spring migration northward along the coast and a return movement in the fall. The flesh of sharks and rays is also ground up and used extensively as In some parts of the world the fins of sharks are used fertilizer. many countries." The wingSawfish steaks like fins of skates and rays make delicious steaks. seen as a scavenger in harbors as well as going out to sea for extended periods. Because of the ventral location of the mouth. or tail. which is dorsally located and greatly . The dorsal and ventral lobes of the caudal fin. there are two unpaired. or ventral fins. the anterior pectoral and the posterior pelvic. each with a spine at its anterior margin (hence spiny). Male individuals may be distinguished from females by the fingerlike extensions. dorsal fins. In addition to the paired fins. but dogfish are now being canned and sold under the name of "grayfish. in the manufacture of gelatin. are unequal and based on this. are quite desirable and the saws are preserved as ornaments. The average length of Squalus is between two and one-half and three feet. these fish are com- monly used small sharks and skates are offered for sale right along with other fish in the markets of China. There are two pairs of fins. There are six pairs of uncovered gill clefts in the walls of the pharynx. or claspers on the pelvic fins.426 TEXTBOOK OF ZOOLOGY liver oil is of Shark In high vitamin content and has an important medicinal use. It is said that unfounded prejudice against eating these fish. The anterior one. They are also salted and dried. A good many dogfish and bonnethead sharks are sold for purposes of study in zoology laboratories. External Features The body is generally spindle-shaped (fusiform) tapering at both head and tail. It is a strong swimmer and is frequently Elasmobranch group. median. these fish find it necessary to turn ventral side up to eat morsels of food from the surface of the water. In the United States there is an as food. particularly of the Orient. THE SPINY DOGFISH This shark is the most commonly studied representative of the Squalus acanthias is the scientific name applied to the common form taken along the Atlantic coast and Squalus suckleyi is the name given the similar one of the Pacific coastal waters.

is This structure is considered its surface by enamel. The centrum. and is said to be amphicoelous.ELASMOBRANCHII modified. and. are all examples of special developments. and supplied with sharp located on the teeth on the jaws. The paired nostrils are to be openings on the ventral side of the snout. It conand vertebral column) visceral skeleton . is 427 a rudimentary gill struc- called the spiracle. the lids of which are immovable. connected with the lower jaw. The The anterior exits tension of the skull the rostrum and the depression in dorsal . The vertebral column and skull are gill arches) . Skeletal System The endoskeleton sists of axial of the sharks is skeleton (skull composed of cartilage. The mouth aperture The placoid scales are primitive exoskeletal structures with a hasal plate embedded beneath the skin and a spine projecting on the surThis spine has a pulp cavity. are situated on the sides The cloacal aperture is located between the bases of of the head. plate. The trapezius found above the branchial area. which has replaced a considerable portion of the notochord in each vertebra. is It contains somewhat the shape of an inverted U. Some of the remains of the notochord fills these interstices between much more developed than vertebrae. Muscular System The segmental arrangement eommas. become segmented and partially replaced by cartilage. separated by myo- The complete along both sides of the body. covered on the pelvic fins. The eyes. which face. The skull is laid is on a foundation of the ventral hasal is dorsal side fairly well enclosed with cartilage. (jaw and and fins. The notochord has in the cyclostomes. homologous to the vertebrate tooth. and the adductor mandihularis. surrounded by dentine. anterior to the mouth. in the principal specializations of independent muscles are found form of myotome modification in the region of the mouth gills and paired appendages. is of myotomes. ventral side of the head. appendicular skeleton (pectoral girdle pelvic girdle and fins). fairly the superficial constrictors extending from the head to beyond the gill slits and assisting in their operation. is deeply concave at each end. ture. These teeth are developed by modification of the placoid scales which cover the skin over the body generally.

ceratohyal. known as preorhital process. Introduction to Vertebrate Zoology. The nasal capsules are rounded. laterally located spherical depressions in The orbits are which the eyes are set. V. one at each side of the base of the rostrum. Sqimlus acant. aosTuuM OLFACTORY CAPSULE EPIPHYSIAL FORAMEN LATERAL LINE FOR^M^^»A ENDOLYMPHATIC FORAMEN QU ADR ATE MECKELIAN PTERYOO- DUCT ORblT OPT\C CARTIlAOE CERATOHYOID HYOMANDl&ULAR CERATO&RANCH lALS EPI- . The visceral skeleton consists of the upper jaw (palatopterygoid (Meckel's cartilages). Dorsal view above. supra- and postorbital process respectively. (From Latei'al aspect on the riglit. car- tilage-encased cavities. epibranchial. lower jaw hyoid arch (hyomandibular. branchial arches (each typical one has pharyngobranchial. . The skull and visceral arches of the dogfish shark. and posteriorly by slight extensions of the cartilage orbital crest. cera- and basibranchial cartilages). BBAMCHIALS CAROTID I FORAMEN PHARYNGO-J BRANCH ALS OTIC CAPSULE \. ventral view below.) 428 TEXTBOOK OF ZOOLOGY side is the anterior fontanelle. five or quadratopterygoid cartilages). hypobranchial. Each orbit is guarded anteriorly. Fig. These capsules house the olfactory sense organ in life. The orbits are laterally located. spherical depressions which normally hold the eyes. published by The C. basihyal). dorsally. Mosby Atwood : — Company. and tobranchial.hias. 230.

by The C. (From Atwood: Introduction to Vertebrate Zoology.\C\.ELASMOBR AN C HH 429 NOSTWU OILL CONUS ARTERIOSUS VENTP. male. Squalus Fig.&PLE. V. COELIAC ARTERY PANCREAS . 231.EN MESENTERIC ARTSRV RECTAL GLANO ARTERY RECTAL OLANO SEMINAL VESICLE INTESTINE A60OMIMAU PORE PELVIC FIH CLASPtR Ventral view of the visceral anatomy of the dogfish sharlc. published acanthias.E TESTIS UVER GALL feL^DOER STOMACH B\LE DUCT DORSAL AORTA . Mosby Company.) — .

. spiral valve of Raia. Dissection of the valvular portion of the small intestine. the bonnet-head shark to show the spiral valve. The basals of this fin are fused into one cartilaginous plate. a series of radial cartilaginous rays. and a series of exoskeletal dermal rays..430 TEXTBOOK OF ZOOLOGY girdle is The pectoral composed of the ventral coracoid bar and the dorsolateral scapular process at each end of it. mesopterygimn. (B reproduced by permission. from General Zoology by Wieman. and metapteryginm). B. Inc. The fin consists of three flat basal cartilages (propterygimn. McGraw-Hill Book Fig.) — Digestive System Most of the organs of this system and other viscera lie in the pleuroperitoneal portion of the coelomic cavity. Anterior to this the pericardial portion of the coelom contains the heart. A. The digestive organs are in the form of an alimentary canal with accessory glands. copyrighted 1938 by Co. The pelvic girdle is made of one cartilaginous bar (ischiopubis) with a fin joining at each end. A. 232.

Following this is the short tubular esophagiis. intestine is This section of the small of considerably greater diameter than the is duodenum and contains internally a spiral valve. is rather U-shaped. which is a lymphoid rather than digestive organ. This arrangement serves to slow the passage of food and increases the absorption surface. . which is somewhat broader than the esophagus. is divided into two lobes. in whose lateral walls are the gill slits (and spiracle). The liver is a large. which a spirally arranged in- folding of the mucous lining. a membranous extension of the peritoneum. The principal absorption takes place through this part of the intestine. The mesentery supporting the stomach is the mesogaster. Circulatory System This centers in the heart. pancreas The digestive tract and adjacent organs in both species are suspended from the body wall by mesenteries. lobe. narrow. which leads directly into the pharynx. which controls the passage of food materials into the intestine. which are extensions from the peritoneum. pancreas. The gall bladder stores bile produced by the The liver and delivers it to the duodenum through the hile duct. and the mesorectum supports the large intestine and rectal gland. and rectal gland are accessory glands con- nected with this system. This organ. the pylorus. large intestine. lies around the greater curvature of the stomach. The liver. which empties into the cloaca (Figs. or pyloric. which leads into the cardiac end of the stomach. The duodenum is the short anterior portion of the intestine which follows the pyloric portion of the stomach and leads into the valvular portion of the intestine (ileum). which is located ventrally at about the and consists of two principal chambers and two accessory chambers. The reddish spleen. the one extending between the spleen and stomach is the gastrosplenic. The posterior. 231 and 232). The rectal gland is a spindle-shaped gland leading into the large intestine directly. portion of the stomach is provided with a sphincter muscle. The pericardium. three-lobed organ with the saclike gall bladder located just dorsal to the junction of the right and middle lobes. The valvular portion leads to the short. an oval ventral and a slender dorsal Ducts lead from it to the duodenum. or membranous lining of the coelom. The two level of the posterior pair of gills. encloses the heart.ELASMOBRANCHH 431 The canal begins anteriorly witli the mouth.


Lateral abdominal V. Efferent branch- Ventricle Dorsal aorta V< Subclavian Duct of Cuvier Subclavian A. Rectal qiand Pelvic fin Cloaca Uiac A: Pig.ELASMOBRANCHII 433 Internal carotid Ventral carotid Hyoidcan artery 1st afferent branchial A ZQd qui Ventral aorta slit W. cardinal ial A. Coeliac A.Gonad Post. LPoit. mesenteric A. Ventral pancreas Qastro-splenic A. Caudal A.) . — Diagram of lateral view of the circulatory bonnet-head shark. (From dissections by system and other organs of Mary Fickling. mesenteric A. - Kidney Spleen A. Ant. Sup. cardinal Ventral gastric A.mesenteric A Intestine Inf. 234.

renal arteries to the kidneys. The two testines. and spleen and empties into the liver. inthe sinus. A bulblike enlargement at the base of the ventral aorta. and the ventral. gastrosplenic to stomach and spleen. where it leaves the venThe ventral aorta leads anteriorly tricle. Hepatic veins lead directly from the liver to the sinus venosus. through the ventral pancreas. ward from the spiracle to the internal carotid. The hyoidean extends from the ventral portion of the spiracle first pretrematic to the where it spreads in capillaries. . bring blood to the kidneys from the single caudal renal portal veins This blood spreads through the capillaries of the vein of the tail. superior mesenteric to the valvular intestine to become the posterior mesenteric artery there. teric arise very near each other. The ventral carotid leads forExtending anteriorly from the dorsal part of each first efferent branchial is a common The carotid which supplies arterial blood to the head and brain. muscular ventricle.434 TEXTBOOK OF ZOOLOGY main chambers are the single. which empties into the auricle by way of the sinu-auricular aperture. which receives blood from the veins of the body. soon giving off the subclavian arteries to the pectoral stomach. the inferior mesenteric to the rectal gland and large intesThe subclavian tine. coeliac to . and the iliac to the pelvic fins and cloaca. The pretrematic and postreniatic branches of the efferent branchial arteries which form four efferent branchials leave the gills and join the dorsal aorta. pancreas. coming off ahead of the posterior efferent branchial the coeliac is farther back and sends a pancreaticomesenteric artery above the duodenum. all The systemic veins return venous blood to the sinus venosus. and pancreas. The blood spreads from these by capillaries through the gill lamellae for oxygenation. Leading into the auricle is the sinus venosus. artery leaves the aorta more anteriorly. from the conus and gives off three pairs of afferent branchial arteries which branch to the five pairs of gills. dorsal aorta extends posteriorly from the junction of the efferent brachial arteries. kidneys and is collected into the postcardinals through the renal . liver. more dorsal auricle. and along the valvular intestine a gastric to the stomach and the hepatic artery to the liver the gastrosplenic and superior mesenfins. is the conus arteriosus. The ducts of Cuvier collect blood from the anterior cardinals of the head region and posterior cardinals of the trunk region and empty it into A hepatic portal system collects from the stomach. which is formed by them in the dorsal midline.

III. followed by two cerebral hemispheres. There is a very slight conthis the tnedulla oblongata. IV. These last two supply the gills. a well-developed cerebellum. lateral line. abducens. trochlear from dorsal side of optic lobes between them and cerebellum. VI. On the lateral walls of the medulla are located the acusticolateral areas. It extends the off paired spinal nerves seg- . The cerebellum is divided into quarters by a longitudinal groove and a transverse and behind striction groove. and certain viscera. oculomotor from ventral side of optic lobes or midbrain. 435 spaces.ELASMOBRANCnn branches. audi- IX. Respiratory System The gills in the wall of the pharynx are constantly bathed in water forced through from the mouth. oculomotor. There are ten pairs of cranial nerves which are numbered and named from anterior to posterior: I. The cavity within the medulla. It covers a part of the optic lobes as well as the anterior portion of the medulla oblongata. is length of the vertebral column and gives tubular and somewhat flattened. This is made possible by diffusion of these gases through the membranes gill of the gill lamellae. An exchange of carbon dioxide for oxygen is made by the blood as it passes through the capillaries of the gills. including the earlike auricles. facial. the glossopharyngeal from a more posterior part of the sides of the medulla. . is the fourth ventricle. which opens to the dorsal surface beneath and behind the cerebellum. The spinal cord mentally. optic. trochlearis. There is also a system of lymph which supple- ments the blood circulatory system. trigeminus. VIII. The gills are supported by cartilaginous arches. The olfactory nerves extend from the olfactory lobes optic from diencephalon and optic lobes. The brain includes two large olfactory lobes at its anterior. and auditory all from the anterior portion of the medulla oblongata. olfactory. tory . vagus. VII. II. the trigeminus. . glossopharyngeal X. abducens. a pair of large optic lobes. between olfactory lobes and cerebrum. a depressed diencephalon. V. Nervous System The central nervous system consists as usual in vertebrates of the hrain and spinal cord. facial.

The internal ears are composed of a vestibule . The olfactory OUPACTORV e>ULB OUFACTORV TERMINAL TRACT CEREBRUM OPTIC MAX>LUAR. Fig. Introduction to Vertebrate Zoology.*S OPTIC LO&E TROCHLEAR OPHTHALMICUS SUPERFICIALIS OPHTHALMICUS PROFUNDUS MANDI&ULARIS OCULOMOTOR A60UCENS CORPU& RESTIFORMUS TRIGEMINAL FACIAL PALATINUS AUDITORY MEDULLA HYOMANDIBULARIS OLOSSOPHARYNGEAL VAOUS Dorsal view of the brain and cranial nerves of the dogfish shark. 235. and the lateral line system. described in the general chapter on phylum Chordata. The nasal chambers are blind sacs and lined with a sensory lamellated olfactory membrane in which the olfactory nerve ends. published Sgualus acanthias. (From Atwood by The C. The eyes are in the orbits. internal ear.) : — organ consists of a pair of nasal sacs on the ventral side of the rostrum which open by nostrils.436 TEXTBOOK OF ZOOLOGY The sense organs include the eyes. Mosby Company. one on each They are quite typical of the vertebrate eye side of the cranium. V. the olfactory organ.

A posterior pouch of the sacculus is considered serves the foreranner of the cochlea of higher vertebrates. and the Wolffian duct serves as the urinary duct of the female. which the dorsal exterior into the lower part of the vestibule the sacculus. carried by several vasa efferentia through the mesochorium to the convoluted cranial portion of the Wolffian duct. portion functions in excretion. The posterior.ELASMOBRANCHII 437 and three semicircular canals. An is endolymphatic canal leads from the lagena. A canal extends along the side of the body in the lateral line and forward onto the head. A papilla leads from the sinus to the cloaca in both. In the male spermatozoa are produced in the testes. Urinog-enital System The kidneys are thin. This is the lateral line system. the epidid^Tnis of each side. There is an accessory mesonephric duct embedded in each kidney which carries urine to the urinogenital sinus of the male. or caudal. Endolymphatic duct Anterior semicircular canal Posterior semicircular canal Utriculus Horizontal semicircalar canal Recessus utnculi L^ \^ ^^^ '-'"?^'^ Sacculus Fig. 236. The spermatozoa then pass out through the papilla to the cloaca. — Diagram of lateral view of left internal ear of Reniceps tiburo. thence to the outside by way of body . slender organs extending along the dorsal wall. The ear the sense of equilibration in the fish. one on each side of the vertebral column. emptying into the urinary sinus. which continues posteriorly as the vas deferens. lying just beneath the skin. On the head there are some pores with tubes extending beneath the skin to small bulbs called ampullae of Lorenzini. The function of the lateral line system ajid these ampullae is perception of water pressure and vibrations. This tube enlarges to become the semincH vesicle and continues into the inflated sperm sac which is directly connected with the urinogenital sinus.

—Urogenital systems of Sgualus acanthias. B. The ova of the female are produced in the ovaries which are located one on each side of the median dorsal line in the anterior portion of the coelom. Fig. TEXTBOOK OF ZOOLOGY During copulation they are transferred to the cloaca of They swim by their own motility into the uteri and oviducts. Each gonad is suspended in a mesentery. 237. Mature ova rupture from the ovary into the body cavity and enter . A. the mesovarium. male. female.438 the anus. the female by use of the claspers.

yolk mass of the egg. and the ground shark. they receive a covering which is secreted the shell gland in the wall of the duct. This is skull is shaped considerably different from that brought about by lateral extension. The embryo is nourished by the large THE BONNETHEAD SHARK. which is 439 held by the falciform mesentery at the anterior end of the peritoneal cavity.) The peculiar shovel-shaped head with the eyes out on the lateral margins is one of the striking features of Reniceps by which it differs from the others mentioned. It occurs abundantly along the Louisiana and Texas Gulf coast during May and June. RENICEPS (SPHYRNA) TIBURO COMPARED TO SQUALUS The bonnethead (or shovel-nosed) shark is common in the At- lantic along the coast of the Southern States and in the Gulf of Mexico. leaving only the five pairs of gill slits.-|-°'^°^torycap5a/e ^^^^ Supraorbital process -Preorbital process . Anterior fontanel/e M m^''"m^ . Endolymph i —Dorsal view of the skull of bonnet-head shark. Squalus.ELASMOBRANCHII the funnellike ostium of the oviduct. Carcharhinns.PosbDrbitcil process • Endolymphatic fossa Foramen maqnum' Fig. The other external features are similar to those of Squalus. pass As the ova bj'' down the oviduct. It averages about the same size as In many respects it is similar to the smooth dogfish. Each olfac- .. Mary (From dissections by Fickling. In Reniceps there are no spines in front of the dorsal fins and a single anal fin is present on the ventral side between the anus and tail. Mustelus canis. 238. Fertilization occurs in the is oviduct and the embryo develops in the uterus which the expanded lower portion of the oviduct. The spiracle is absent. Epiphyseal foramen ... The skeleton of the of Squalus..

. Fickllng. The posterior part of the skull is somewhat narrowed and flattened but otherwise similar to that of Squalus. (From bonnet-head by MaryReniceps tiburo. shark. and bladelike postorhital process are also Ventral pancreas Cardiac stomach H\dr)ey SmaW'mtzstme - Recta/ qloDd Larqe intestine Spleen stomach Mesoneplinc duct -- Cloaca Fig. anatomy of —Internalview.440 toiy capule is TEXTBOOK OF ZOOLOGY extended far to the lateral of the base of the rostrum it. preorhital process.) dissections from ventral projected at the terminal portion of this arm of cartilage. 239. The orbit with the modified supraorbital instead of lying beside crest.

lon is entirely hidden from dorsal view by the cerebrum and cerebellum. In the pharynx there are no spiracles. There The diencephais no line of demarcation between the hemispheres. there is a long glandu- which extends from the level of the gonad proper to the region of the cloaca. at the time in Texas Gulf waters in late August and early September. Fairly mature "pups. The shape of the testes in male Reniceps is much longer and more slender than in Squalus. and convoluted. They contain extensive folds or lamellae of the olfactory membrane. somewhat different in the two animals. The medulla has well-developed acousticolateral areas. The latter is large. The spiral folds of the spiral valve are more telescoped into each other than in Squalus.ELASMOBRANCHII 441 There are a few differences in the digestive systems of the two. Copulation in Reniceps probably occurs during May and June in when they are so numerous in the shore waters. the epigonad. Reniceps has five afferent branchial arteries branching from the ventral aorta while Squalus has only The branching of the coeliac artery is three." ^s the developing young are called. The olfactory lobes are broader and almost completely fused to each other. . The brain has the same general parts as it does in Squalus but they are quite modified. have been found in the uteri of specimens collected off shore the Gulf of Mexico. The cerebrum is a somewhat smaller single lobe just posterior and dorsal to the olfactory lobes. The stomach of Reniceps is J-shaped instead of U-shaped and the long slender pyloric portion is armlike. It covers not only the diencephalon but also most of the optic lobes (midbrain) and much of the medulla oblongata. The nasal chambers of Reniceps are quite large and kidney-shaped. There are only two lobes in the bonnethead and the gall bladder is nearly embedded in its In the circulatory system. lar body. In addition to this. two of which branch. liver of tissue. irregularly divided into three lobes.

assists in dividing the water as the fish moves through body. and paired fins as well as median fins to assist in locomotion. all of the original features of The shape it. a pair of pelvic or ventral fins which are quite and in some forms rudimentary. sometimes divided into two a single caudal fin over the tail a ventral median anal fin . The endoskeleton is primarily bony. in most species. but the pelvic girdle is usually small. The fins are supported by fairly well-developed fin rays. The shape of the body varies from that of the long slender eel to that of the globe-shaped box-fish and inflated puffers which can float like balloons. Most forms within the class have scales as an exoskeletal covering of the skin.CHAPTER XXVII PISCES. The structure of the caudal fin and posterior end of the vertebral column is distinctive and has been classified. Still another peculiar adaptation is the flying fish. of speed. ferent forms. . The fins of fish are found singly in the form of a dorsal median fin. 442 . The most primitive type of tail is the diphyceroal in which both the cutaneous and osseous parts are equally divided between dorsal and ventral regions. while in the bullhead catfish they are just anterior to the anus. which is composed entirely of skin and is called an adipose fin. The typical shape of the fish's body is fusiform or spindle-shaped. an adaptation for easy production The sedentary forms of fish usually tend to lose this shape and become flattened or otherwise modified. The heterocercal tail is asymmetrical and the tip of the vertebral column exvariable in position pectoral fins. The majority of families in this class possess a swim bladder. TRUE FISH dif- This important class includes quite an extensive variety of They are aquatic and possess the usual adaptation of gills for respiration. The sea horse is one of a group of very peculiarly shaped forms. Pectoral and pelvic girdles are developed to support the paired fins. The pelvic fins of the perch are located almost immediately ventral to the pectoral fins. and the paired These paired fins are supported by bony girdles. it As the water passes over the thicker part of the rushes in to push forward on the posterior slopes of the This is spindle form of the body. with stream-lining. In this catfish there is a second dorsal.

. 240. published by Henry Holt fish. but the lobes stroke the water with about equal surface and force. (From Krecker. common eel -B. C. — into the dorsal lobe. after Jordan. sea Fig.) . from Gulf of Mexico and Atlantic Ocean. A. The original notochord turns Fig-. It forces the fish through the water in a hori- zontal plane and is correlated with a terminal mouth. General Zoology. Chilomycterus schoepfli. horse. 241.PISCES. tends into the dorsal lobe as has already been seen in the of tail is the liomocercal. — Spiny boxflsh. TRUE FISH 443 dogfish. Company. flying & Diagram showing some peculiar bony fish. which is internally unStill another type balanced but externally symmetrical.

after Her twig. cycloid. and ctenoid. placoid. General Zoology. concentric lines.) The different types of fish scales. (From Krecker. 1. ctenoid. lines formed during nongrowing periods fuse closely together. The bones of the operculum arise as a part of this division. The skeleton includes. 2. shingle roof. . thus indicating seasonal periods on the scale. clams. insect larvae. The third type is similar to the cycloid except that the free edge of the scale bears some spiny projections or cteni. the jaw and five gill arches. which skeleton. These are: ganoid. It passes growths. cycloid. published by Henry Holt and Company. The first the are usually rhombic or oval in shape and are covered by a dentinelike Such fish as gar pikes and bowfins possess Cycloid scales are rather disc-shaped with conspicuous this type. Crustacea. The digestive tract is in the usual form of a canal with outFood ranging from vegetation. ganoid. and snails to small fish and amphibia is utilized. the amphicoelous (concave in both ends) vertebrae and bony cranium. besides the paired iins and girdles already mentioned. Fig.444 TEXTBOOK OF ZOOLOGY There are three principal types of scales which cover and protect body of most true fish (a notable exception is the catfish to be described later). The age of many fish can be The determined by the distribution of the concentric lines on the scales. like a substance called ganoin. i. 242. They are usually imbricated on the skin. 5. — Cycloid scales are found on the carp while the ctenoid are characteristic of the perch and sunfishes. is complete and independent of the visceral This latter portion consists of seven arches. structures.

pharynx. TRUE FISH 445 through the toothed mouth. The system includes the paired anterior and posterior cardinal veins meeting in the duct of Cuvier which joins the hepatic portal vein leading to the liver. Oxygen is absorbed by the blood. and carbon dioxide is discharged to the water. are located on the jaws. pharynx. in general. similar to that described for the lamprey. roof of the mouth. of North America. and walls of the and are used primarily for holding prey. each of which is connected by a mesonephric duct by to a urin<iry sinus or bladder. mouth and forced out The mouth and pharynx form or expired through the a water-tight pumplike arrangement with the help of the lips flaplike oral valves just inside the and the IrancJiiostegal memhrane at the margin of the operculum. the swim bladder in certain fish. in The circulation most fish is. and the various branches. gills. the two-chambered heart with accessory sinus venosus and lulhus arteriosus. The exchange of oxygen and carbon dioxide between the blood in the capillaries of the gills and the water occurs as the water passes over the gill lamellae. but a general scheme of the like 20.000 acceptable plan will be presented. the its vein from liver to sinus venosus. and. the hepatic sinus venosus. There are something Few species of fish described. esophagus. accomplished by a pair of dorsally located mesonephric kidneys. to some Water is drawn in or extent. Teeth ileum.PISCES.300 species occur in There are two subclasses. except for certain specializations and phylogenetic developments. . of their orders which more than 3. Gastric glands in the wall Pyloric caeca of the stomach supply some of the digestive juices. Classification students of the fish are in complete agreement on the classimost fication of all groups of fish. and large intestine during the process of digestion. anterior portion of the duodenum increase the which join the absorptive and digestive surface in many fish. inspired through the gill slits. dorsal aorta. ventral aorta. An abridged summary is and families included here. This bladder opens to the exterior Excretion is an aperture located just posterior to the anus. duodenum. branchial arteries. The respiratory system consists of the mouth. stomach.

There are only three genera — usually described for this country. suborders and families. 243. each including subdivisions of either families or in the larger orders. rows of keeled. Sturgeons. tesy of American Museum of Natural History. Family Acipenseridae. It has a smooth skin and long. spoonbill cat or paddle fish and its associates. ganoid shields The body is covered with five and the tail is distinctly heterocercal . — Fig. Paddlefishes or Spoonbills. A ganoid type of fish with abundant proportion of cartilage in the skeleton and with bony ganoid ula scales. 446 Subclass Teleostomi. Polydon spathula. many In this group are found all of the marine fish which frequent our waters and shores. with a few ganoid scales on the tail. Family Polyodontidae. except lamprey. Polydon spathis found in the Mississippi Valley. In this general division of the group are four orders in the world. an Afrifish using the swim bladder as an accessory respiratory organ. flat paddlelike snout. fresh-water fishes and TEXTBOOK OF ZOOLOGY —True Fishes.— . can Order Crossopterygii (Family Polypteridae) Polypterus. —Paddlefishes — and Sturgeons.) (Cour- Order Chondrostei.

but in a — few they are cycloid and imbricated. fin. it can now be found only occaand. —Bowfin or Fresh-water Dogfish. Family Elopidae. Both families of (caviar) as food for man. Salmon. Fish of this family have greatly compressed bodies which are covered with large. find the cycloid scales and another form capable of accessory respiration by means of swim bladder.PISCES. ctenoid. the scales are of the enamellike ganoid variety. though sionally. TRUE FISH 447 in the mouth. Tarpon. This is a very active game type of fish with large scales It is and an extended filament from the dorsal very abundant throughout the Gulf of Mexico and is a fish all famous game along the Texas-Louisiana coast. The long-nosed gar and alligator gar are the most common southwestern forms. Gar pikes and Bowfins. silvery. Found in warm marine waters. ganoid scales. the only species in existence. This is another ganoid type but with a more complete bony skeleton. Mooneyes. Ten-pounders. or no scales and their type of tail is usually homocercal. The pelvic fin is abdominal and the dorsal fin is — far to the posterior. while the long-nosed and short-nosed are the common species of These are covered with a strong armor of the Middle West. — Tarpons. They inhabit the fresh-water lakes and sluggish streams as far southwest as east Texas. Family Megalopidae. this order furnish both flesh and roe Order Holostei. with the upper lobe quite extended and slender. Some are soft-rayed fishes with open connection between swim bladder and alimentary canal (Physostomi). It is thought to have descended from the ganoid type. Suborder Isospondyli. They have cycloid. Their skeletons are highly ossified. Herring. enamellike. cycloid — . rhombic. — — Flops saurus is the typical representative. Family Hiodontidae. — True Bony Fishes. The snout is conspicuously extended. and the Order Teleostei. and others are spiny-rayed fishes with no air duct from swim bladder (Pkysoclisti). tail is heteroeercal. Gar pikes (garfishes). There are no teeth Acipenser fulvescens is the Mississippi Valley form. we In Amia calva. A group which includes the majority of existing fishes. In most of the representatives of the order. it was once abundant. Family Lepisosteidae. etc. Family Amiidae.

in warm and swim in Anchoviella (Anchovia) mitcMlli one of the most common American water lake species. Thymallus tricolor. large-mouthed fish with fine scales. fresh-water species. They are found many species enter fresh water to and there are very few spawn. large schools. —Whitefishes. This There are about twenty freshan important commercial fish. prolific. compressed bodies. Family Coregonidae. It is a northern fish. is — Gizzard Shads. fishes Family Dorosomidae. is have elongated. Hiodon tergisus is the form found in the Mississippi Family Albulidae. TEXTBOOK OF ZOOLOGY There are three species in our western streams and in the Great Lakes. This is elongate. These differ from Salmonidae chiefly in the presence of a blind saclike stomach. —Anchovies. Valley. with a rounded belly outline. but almost inedible. They lati- are principally northern and abound in clear waters north of tude 40°. but they have a large dorsal fin. This is a scarce. widely fish. waters. Michigan Grayling. Ciscos a group of are included in this group also. is a typical species. ber of species. small. seas is They are abundant forms. moderately compressed. group of fin. Graylings. bluish fish. a large number of pyloric caeca. Family Clupeidae. The body There is quite flat from side to side. Family Salmonidae. This family includes a large numin it is thought that there are more individuals this group than in any other. The esophagus and pylorus join close together. distributed. and the edges are These fish thin. A small group of a few species warm marine Family Dussumuriidae. internally. a lateral line and. —Round Herrings. and —Herrings. but in most seas. — Smelts. These fish resemble the previThe ous group in many ways. They are noted as game and food fish. found in —Ladyfishes. This is an abundant. . — Salmon and Trout. a filament extending from last ray of the dorsal Family Engraulidae. — Family Osmeridae. Family Thymallidae.448 scales. of which Clupea harengus The family includes many valuable food is the most important. small.

scale carp. Fishes with elongated slender body and no pelvic fins. TRUE FISH This is 449 a small Family Argentiuidae. leather carp. — True Eels. A group of marine eel. a. eels.PISCES. Family Ophichthyidae. Snake Eels. dis- tantly related to the Anguillidae.) after Fig. published by Lea and Febiger. European carp. (From — streams of the Southwest. Metcalf. Family Congridae. Carjlina. Family Anguillidae. Cyprinus carpio. the common They are withalso without and the body often entirely black. Suborder Apodes. These eels are Bcales and have the tongue more or less adherent. The scales are embedded in the skin. They are among the most voracious of fishes and are quite hardy. b. Eels come several hundred miles inland from the Gulf of Mexico in the 244. They go out to sea to spawn and the young make their way back up the streams. — . —Argentines. out scales. Textbook of Economic Zoology. Fishes of North. — Conger is Eels. It is thought that the female spawns once and then dies. group of northern or deep-sea species. —Eels. etc. Smith.

fish is — — Family Synodontidae. tough air bladder.450 TEXTBOOK OF ZOOLOGY Family Muraenidae. —Moray eels. These fish are devoid of scales. felis. are very common along the shores of the south- ern Atlantic and the Gulf of Mexico. These. toothless jaws without barbels. As a group. and Minnows. as well as pelvic. The air bladder is large and connected with the alimentary canal. Lizard Fishes. Family Ameiuridae. The bullhead catfish and channel catfish are very well known and almost universally distributed over the United States. and all are tenacious of life. the Gaff-topsail Catfish. too. — Carp. a large mouth with the lower jaw projecting. The skin is thick. Family Umbridae. Fresh-water Catfishes. — — Catfish. and a round sucking mouth. and Suborder Eventognathi. Family Esocidae. Family Catostomidae. lateral line are covered with cycloid scales. east of the Rocky mountains. This is a large and important group in fresh water. Suborder Nematognathi. These carnivorous They have no There are at least three minnows and American have a mud of sluggish streams. scaleless. species. and a large. and Minnows. The head is depressed and the mouth is quite wide with strong teeth. — Family Ariidae. two inches in length. Sea Catfish. Carp. These fish slender body. As in the suckers. — Suckers. There are 60 species in the fresh waters of North America. leathery. —Mud Minnows. and Pickerels. live in the —Mud Minnows. Suborder Haplomi. fins. the jaws of this group are toothless. The body of this type of elongate and covered with cycloid scales. A group of fish whose air bladders are usually large and commonly divided into anterior and posterior lobes. These fish have elongated bodies covered with cycloid scales. and Oaleichthys — common sea cat. Pikes. Suborder Iniomi. they are excellent commercial and food fish. They feed on plant tissue and small animals. —The Pikes or Pickerels. Dace. have barbels on the lips. Family Cyprinidae. to large carp of two feet in length. They range from small minnows. and a . barbels. Some of the most abundant species of general distribution fall in this family. are scaleless and have Bagre marina. In this type there is an absence of pectoral. Suckers.

Fish. top minnow. heteroclitus is a much used a very labo- ratory animal along the Atlantic coast. near the base of the forked lucius. This is is common mosquito-eating fish of Ganibusia a widely distributed Family Amblyopsidae.very peculiarly-shaped this family —Pipefish. Esox vermiculatus. Killifishes. — Cave Fishes. These are large-mouthed fish with numerous teeth and pyloric caeca. Poeciliidae. Codfishes. Family Syngnathidae. Sea Horses. —Sticklebacks. Some of these peculiar fishes are able to leave the water and glide through the air for several The pectoral surfaces. and Cave Family Cyprinodontidae. —Pipefishes fishes and Sea Horses. (Flat-fishes. common pike. by Funduhts heteroclitus. and Tongue Family Gadidae. vorato the warm seas. Family Belonidae. Family genus. This group of minnows. Esox and Esox masquinongy. There are a few species of these small fish living in subterranean streams of the central and Suborder Synentognathi. —Halibut Flounders. and Sticklebacks. —Mosquito the South. the muskellunge. the tail. yards. fins are greatly developed and serve as planing There are about sixty different species inhabiting the tropical seas. Family Gasterosteidae. Most representatives of fin. are found commonly in shallow well represented estuaries or fresh-water streams. Suborder Cyprinodontes.PISCES. — Codfishes. They are food fish of northern seas. — Killifishes. and Esox niger penetrate the Southwest. Fishes. Mosquito Fish. southern United States. Suborder Heterosomata. TRUE FISH 451 dorsal fin far to the posterior. Soles. the mud or grass pickerel.) — . They range from Maine Texas Coast of the Gulf of Mexico. They are principally northern Suborder Anaca7ithini. cious fish of the — Sea These are long-bodied. —Flying Fishes. Family Exocoetidae. A group of whose bodies are quite long and slender. Suborder Thoracostei. or Zygonectes notatus. — F. are familiar northern forms. Fish. Garfishes. the common killifish. have prominent spines just anterior to the dorsal in distribution. — Garfishes and Flying Fishes.

mon forms shores. etc.452 TEXTBOOK OF ZOOLOGY cipally marine Family Hippoglossidae. — . 245. Family Achiridae. monly along the Gulf compressed. published by Lea and Febiger.) Fig. particularly along the coast in the warmer regions. Soles. —Mullets. Textbook of Economic Zoology. flat —Tongue In these the eyes are on body and the entire body resembles a tongue Fig. Textbook of Economic Zoology. —Mullets. (From Metcalf. published by Lea and Febiger. -White mullet. Scomheromorus maculatus. Pompanos. Family Mugilidae. left side. The Halibut family and is used a great deal for food. Mugil curema. They live in schools. Halibuts. lie — laterally —Broad coast. Snappers. 246. Most of these fish are small.) — Suborder PercomorpM. Family Atherinidae. Basses. Like the other flounders they are greatly on the and have both eyes on the Fish. the left side of the in shape. (From Metcalf. These are found* quite comright. Silversides. 245. Family Cynoglossidae. Mackerels. after Smith. Perches. after Smith. coast.) it is There are numerous species of comand river mouths along our southern This is an esteemed food fish along Atlantic but ignored in the Gulf States. — is prin- Family Pleuronectidae. Flounders. compressed and covered with even cycloid scales. Spanish mackerel. These fish have a flattened body and both eyes on the right side of the head. in the estuaries (Fig.

Textbook of Economic Zoology. water game fish. pikelike fishes of the There are 15 species. after Smith. This group includes vicious. The tail is usually quite widely forked. seas. published by Lea and Febiger. metallic blue in color and are very important commercial fish of the Atlantic. They are fine food fish and are very abundant in the Gulf of Mexico (Fig. — Large-mouthed black bass. Micropogon undulatus. The dorsal fin projects high above the body. warmer Threadfins. (From Metcalf. Family Istiophoridae.— True Mackerels. They live in great schools. These mackerels have a scaly body. — . voracious. These fish are a pretty.PISCES. after Smith. —Barracudas. This common name comes from These are found among collections from the Gulf of Mexico.) Economic Zoology. 247. Family Cybiidae. (From Metcalf. 246). published by Lea and of Family Scombridae. the pectoral filaments. Textbook —Marine croaker. Among this group are found some of the swiftest of fishes. Spanish Mackerel. Family Poljaiemidae. The prize freshPig-.Febiger. 248. Sailfishes. Micropferus salmoides.) — Fig. TRUE FISH 453 Family Spliyraenidae.

swiftly-moving streams. The bodies of most of the sunfishes are about the shape of a person's hand. This family is another which is related to the mackerels. These Family Serranidae. remora attaches itself to the ventral side of a shark and is carried — — about with it. These fish have rather small fusiform difference between these — — — bodies with ctenoid scales. They are prized commercial fish. Family Echeneidae. Gurnards or Sea Robins. Perches. There are at least 500 species. They reach large size and are quite numerous on the high seas. Family Triglidae. These are tropical fish and several species of them are choice food fish. the gaspergou or drum is found — in fresh water. Gobies. Suborder Gobioidea. Suborder Cataphracti. — Sea Basses. Aplodinotus. They are carnivorous. Family Percidae. Family Gobiidae. important fish. the red snapper. Sunfishes and Fresh-water Basses. A brilliantly colored group of small fish found in clear. and common on sandy shores of warm seas.. the yellow perch of the are several northern food fish states. Sea Robins. This is one of the widely distributed families with numerous species and abundant individuals. — Tunnies — and Bonitos. Family Centrarchidae. — Snappers. They are — — common in all warm seas.Remoras. Family Etheostomidae. — — These small. Remoras or Shark Pilots. Perca flavescens. Family Carangidae. There Family Lutianidae. In this group the air bladder is usually large and constructed to enable the fish to make a grunting or drumming sound. The bodies of members of this family are usually covered with rough scales. particularly Lutianus campechanus. Darters. There is not a wide and the other mackerels. The spinous dorThe sal fin of this fish is modified to form a sucker on top of its head. chiefly of tropical seas. They all build nests and are desirable game fish. Suborder Discocephali. Family Sciaenidae.454 TEXTBOOK OF ZOOLOGY Family Thunnidae. . are abundant in the deep waters off the Gulf coast. carnivorous fish are found creeping about on sandy bottoms along shores and in mouths of rivers. included in this group. Drumfishes. Pompanos. is almost a classical classroom form.

They possess characteristics which apparently place them intermediately between fishes and Amphibia. by two lungs formed from a modified Lepidosiren paradoxa is the single form found in South is America. velvety skin. They feed is the genus found in the marshes of Africa. tion. worms. paired The air bladder is connected able to breathe air. TRUE FISH 455 Suborder Jugulares. There are only three living genera. This group includes a number of species which have large mouths and somewhat the appearance of toads. The fixed spines on the bony plates in the skin are characteristic of the group. There is only one species in this family and it is Neoceratodus fosteri.— Porcupine Fishes. Suborder Pledog^iathL—Trigger^hes. Fishes. Filefishes. aestivation by burrowing into the mud of the marshes during the into dry summer season. — Anglers. slender. —Triggerfishes. and smaller vertebrates. — Family Batrachoididae. Lungfishes of South America and Africa. tip to hang over the mouth as a bait for other fish. with the pharynx and is modified to serve as a lung. found in stagnant waters of Australia. and Porcupine fins of Family Balistidae.— Filefishes. Suborder Pediculati. Economic Relations of the Class Fish have been one of the stable sources of food for man all through history. Family Lepidosirenidae. They go on insects. fish is Family Monacanthidae.—Australian Lungfish. — Here they carried on secrete a cocoon of slime for protec- Respiration air bladder.PISCES. The spines of the dorsal these fish are long and saw-toothed. broad-bodied Family The first spine of the dorsal fin is extended and enlarged at the fish. crustaceans.— Toadfishes. H^idreds of millions of dollars . Proiopterus This species is have long. Family Diodontidae.—Lungfishes. fins — Lophiidae. and it is in this respect that they are most important to man in present times. The fins are poorly developed. Subclass Dipnoi. The body of this type of much compressed and covered with rough. A heavy appearing. Family Ceratodontidae. These fish and a well-developed median fin.

shad. and the pectoral fins of catfish have a barbed poisonous spine. catfish. The sharks and crabs. game fish. Several of the food fish are also game fish. americana. haliver oil. such as bass. Fish and Wildlife Service and most states through their Fish and Game departments are making continuous studies of the fishing industry and are conducting fish culture on a rather large scale. such as malaria. a top minnow of the South. demand as caviar. annually are yielded by the commercial of The annual catch salmon alone is estimated at $17. and pike perch are all important food and commercial flesh. living largely on other fish. extracted from certain fish scales in an known ammonia . Certain rays. Many of the game fish. as pearl essence. The United States government through the U. clothing. Some highly desirable such as the tarpon. as well as the larger ones. destroy many other fish. shark-liver oil. carp. In addition to the food value of the the eggs of several species. A liquid. and dengue is fever.456 TEXTBOOK OF ZOOLOGY fisheries. halibut. Several groups of fish have a distinct negative value. The gar pike is decidedly a predator. is an asset because of its appetite for mosquito larvae. fish. such as sturgeons and paddlefish. cod- fish. Fish scrap. are much used as medicinal products for production of vitamin D which is a preventive for rickets. which is left after the oil is extracted. by destroying mosquitoes. there tackle. by this sport. red snapper. pain- wound which is dreaded by all fishermen. There are still several other valuable relations of fish to the welfare man. ful Either of these fish can inflict an ugly. ciscoes. The hides of sharks and some other fish make fine leather when tanned. Such oils as cod-liver oil. trout. yellow of fever.000. lobsters. have a poisonous spine on the tail. Besides salmon. Ganibusia. inspired is much money spent on the trips. mullet.. trout. are in great greatly prized as crappie. mackerel. is used as a fertilizer and is also put up in the form of fish meal and sold as protein feed for other animals. and the pikes. such as Dasyatis sahina and D. are choice food. especially. are almost worthless as food. herring. called the mosquito fish. S. Besides having recreational value. buffalo fish. The value of the sport of fishing is usually underestimated. injuring or even killing a human being occasionally. etc. It is used to control the spread of mosquito-borne diseases. damage perhaps one- half million dollars' worth of fishing gear per year.000.

It is a External Features The body wide. and lakes. to the maxillary process at each side of the These are distributed. one attached mouth. fin. The tail is homocercal. These fish will be the principal subject of this description. the head is short and broad. Ameiuriis natalis. Perch has none. TRUE FISH 457 "When glass beads are coated with wax and covered with this solution they become artificial pearls. bladelike anal fins. There are two pairs of nostrils on the head. broad. Perca The bullhead inhabits nearly all sluggish streams. colors of the upper parts of the bullhead range from yellowish green to dark brown. Perch lives in clear lakes and ponds. ponds. but during recent years The natural range extends it has been successfully introduced. as well as eight feelerlike harhels. will be some comparisons made with yellow perch. while those ventral to the mouth are pinkish cream.PISCES. It lives along the muddy banks and around submerged rocks and logs very hardy fish and is able to thrive in almost any aquatic condition. in the water. The eyes are On each side of the head is a relatively small and without lids. and a rounded caudal fin forming the Just anterior. Its distribution did not originally reach the Pacific coast. and the ventral side is yellow. The sides are a lighter waxy yellow or yellowish brown. Extending along each side of the body is a lateral line. flaplike operculum. solution. an adipose fin back near the tail. two dorsally. throughout the Middle West. Anterior to this gills is a pair of ventral or pelvic Just posterior to the position are the pectoral fins. and well into the Southwest. is a single. which covers the gills. The dorsal barbels are brown. There is a darker longitudinal band running lengthwise of the anal The •In collaboration with Rose Newman. to the caudal fin on the ventral side of the body tail. is widely distributed through the fresh waters of the United States. and the mouth is There is a relatively small dorsal fin located anteriorly. . but there flavescens. TYPICAL BONY FISH—YELLOW BULLHEAD AND SOME COMPARISONS WITH YELLOW PERCH* The yellow bullhead catfish. is stout. and four pale- colored ones on the skin of the lower jaw. South. The skin of and in a lateral Ameiurus is smooth and without scales whereas the skin of perch is covered with ctenoid scales.

249. esophagus. as well as insects. insect larvae. is similar to that of the yellow perch. The dead bodies of almost any animal Avill be eaten by the bullhead. The digestive system consists of the mouth.-Digestive system and other visceral organs of Ameiurus natalis. the body splits the water as it passes through it. In fact.. ESOPHAGUS UIVER 6AL_U Bl_ADDER DUODENUM 7t'!l\ CABDI AC STOMACH PYLORUS :iT4-FUNDIC STOMACH SPl_EEN I_EUM Al R BUADDER Kr DNEV LARGE INTESTINE TEST I ANUS ^BL-ADDER I —VAS DEFERENS MESONEPHRIC DUCT UROGENI TA1_ PAPIUL.) Digestive System and Digestion Except for some habits of a scavenger. water and other small fish. and the water closes in on the posterior slopes of the spindle shape to help force it forward. the food of the bullhead snails. and anus.S 458 fin. which includes crayfishes.A Fig. The mouth is large and has teeth . stomach. (Drawn by Titus Evans from dissections by Rose Newman. pharynx. intestines. TEXTBOOK OF ZOOLOGY The general shape of the body is fusiform or spindle-shaped and hence offers little resistance to the water.

Circulatory System and Circulation The heart lies almost free in the pericardial space at the extreme anterior end of the body cavity. bullhead. . thence to the ventral aorta which branches into four pairs of afferent branchial arteries into the Here the blood passes through finely branched capillaries gill arches. pyloric valve to the enteries. Following the small intestine the There short broad large intestine leads to the anus where fecal material is discharged. but is cylindrical in perch. It passes auricle to the valve. TRUE FISH 459 by the maxillary jaw above and the mandibular jaw below. Gastric glands in the walls secrete enzjones which start the digestive process. The bones in the roof of the pharynx bear superior tooth pads which are round or oval in shape. diges- which and continues directly to the pylorus in the bullhead. There it. It is composed of two principal is chambers and two accessor}^ ones. The blood enters the saclike sinus venosus from the common cardinal veins and hepatic veins. testine is It receives the hile duct from the liver and possibly some small pancreatic ducts from small masses of pancreatic tissue held in the mesregion. with the pyloric portion extending from the side. through a valve to the auricle.PISCES. is not a distinct pancreas in either the perch or the Digestion continues in the small intestine and most of the absorption of food by the blood occurs through the walls of the posterior portion of the ileum. then with contraction of the muscular ventricle through the auriculoventricular The contraction of the ventricle forces the blood through the semilunar valve into the conus arteriosus. which is supported by the hyoid hone. The esophagus is a straight muscular tube near the posterior end of which enters the ductus pneumaticus from the air bladder. The teeth serve to hold the prey or food in the mouth. tion begins in the stomach As is is generally the case in saclike fish. has a row of papillae running along its midline posteriorly into the pharynx. The small inshorter and less coiled in the bullhead than in the perch. The at the front supported pharynx is rather funnel-shaped and has four gill slits in each lateral wall. a single auricle with the accessory sinus venosus leading into and the single ventricle which leads into the accessory conus arteriosus. The tongue. In perch there is a group of fingerlike pyloric caeca attached to the side of the pyloric The mass of partially digested material passes through the duodenum of the small intestine.


I. — . ^ox. urinary bladder.. ileiim L. 251.. Bl. stomach. testis. large intestine.. T.ii:-. TRUE FISH 461 CAROTID EFFERENT BRAMCHl Al_ SULBUS ARTERIOSUS AFFERENT BRAtslCMI AU DORSAL AORTA SUBCI_AVI AN TO HEAD KIDNEY HEPATIC COEI_l ACOMESENTERIC PNEUMATOCVSTtC GASTRIC MESENTERIC RENAL- NTESTINAU SPLENIC I GENITAU IL-t AC HYPOGASTRIC CAUDAL Arteries and most of the visceral organs of Ameiurus natalis Le i-16. Sto. auricle.Ht. A.PISCES. H. Sueur. il. kidney proper. (Drawn by Titus Evans from dissections by Rose Newman. head kidney .K..) . Fig-... lateral view.

kidney.. L. 252.462 TEXTBOOK OF ZOOLOGY ANXERlOR CARD NA. head Icldney. H.K. POST CARDINAL. lateral view. St.) — . PtslEUMATI C GASTRI C I NTESTINAL. Kid. large intestine.. MESENTERIC INTESTINAL ABDOMI NAt_ PORTAI_ ll_l AC RENAU PORTAl_ I GEN TAl_ CAODAU Veins and viscera of Ameiurus natalis Le Sueur.l_ DUCT OF CU VI ER I SINUS VEISJOSUS HERAT C I HERAT IC PORTAl_ ACCESSORY PORTAL. ileum..I. Fig. II.. (Drawn by Titus Evans from dissections by Rose Newman. stomach..

taking up almost one-half of the space of the abdominal The pneumatic duct extends from its midventral region to the posterior part of the esophagus. The blood consists of the fluid plasma. The branchiostegal membranes relax and when pressure is applied to the water in the mouth. Water is drawn into the mouth by lowering the floor. while subclavian veins return from the pectoral fin region. This is the principal artery of the trunk of the body and gives subclavian branches to the fins. The opening of the mouth is guarded by fleshy flaps or oral valves. the hepatic portal from the visceral organs to the liver. In the perch. it is found that oxygen is secreted into the closed swim bladder during periods of plentiful supply in the water then drawn upon by the blood at times when the environmental supply is scant. TRUE PISH 463 where it is aerated by absorption of oxygen from the water passing gills. corpuscles.PISCES. of air through this There is likely some exchange and the possibility of some degree of diffusion of gases between this and the blood in its walls. over the surrounding This blood continues by convergence of the capillaries into the efferent branchial arteries which lead dorsally and join pectoral in the formation of the dorsal aorta. air bladder likely has sac. and finally ends in the caudal artery supplying the tail. oval nucleated red. Another function of the air bladder is to decrease . a large. is it is forced out over the gills. This process repeated in rhythmic sequence in order that the blood passing gills will through the capillaries of the be constantly aerated. and the hepatic from the liver to the sinus venosus. These capillaries also waste prod- and converge to form the veins which carry the blood back to the heart. When the mouth is filled with water. oxygen are supplied ucts to the tissues The food substances and of the body by means of capillary collect the branches through them. The 'posterior cardinals return from the posterior portion of the trunk. some respiratory function. parietal ar- teries to the body wall. it is closed. and amoeboid white corpuscles. It is The tough cavity. which has no pneumatic duct. and these valves prevent water from escaping through this aperture. Respiratory System The mouth is used in forcing water over the four pairs of gills. renal arteries to the kidneys. coeliaco-mesenteric artery to the viscera. while the branchiostegal membranes at the margins of the opercula are closed down over the gills.

The head kidney — head kidney (pronephros) pneumato— cystic duct l_EFr a r bladder OVARY i RIGHT OVARY KIDNEY (mesonephro^ URINARY BLADDER OVIDUCT MESONEPHRIC DUCT Fig. They are located in the dorsal wall of the abdominal cavity. Excretory Organs The kidneys of Ameiurus are similar to those in the perch. posterior to the air bladder and just outside of the peritoneum. as in perch. 253. the volume is regulated by secretion or absorption of oxygen. The urine is stored in the bladder and finally expelled through the urinogenital (pronephros) pore just posterior to the anus. as the need may be. (Drawn by Titus Evans from dissections by Rose Newman. is a paired mass of lymphoid tissue in front of the air The functional kidneys are composed of numerous uriniferous tubules supplied with blood capillaries which extract urea. . creatinin. Skeletal System of the soft fin rays which support the fish like Since there are no scales on the bullhead.. The ctenoid scales of a the perch are also exoskeletal. exterior the volume of air in In forms where the bladder is open to the it can be regulated by direct exchange and allow the animal effort. and other wastes from the blood.) bladder. the exoskeleton consists fins. A slender mesonephric tube leads from each kidney to the urinary bladder.-Urogenital system of bullhead. lateral view. to take a definite level in the water without In the closed type.464 TEXTBOOK OF ZOOLOGY the specific gravity of the body of the animal in water and to serve as a hydrostatic organ.

quadrates. Each is composed of either four or its five They are parts and has spiny gill rakers at located in the lateral wall of the pharynx and are covered by the four opercular bones of each side which compose the framework of the anterior margin. A neural spine extends from the neural arch. pectoral and The main axis of the skeleton is the vertebral column and skull. alisphenoids. Vertebrae six to fourteen bear ribs from the dorsally transverse processes. referred It supports the tongue. and parasphenoid. supraclavicle. The skull may be divided into cranium and The cranium encloses the brain and is composed visceral skeleton. and to as number two. The adjacent vertebrae articulate at the centra and are held in place by ligaments. ribs. The maxillary arch supplies both upper and lower jaws. The parts of one of the simple vertebrae are the body or centrum just mentioned. hasioccipitals. epiotic. sphenoids. pterygoids and nasals protect and support the auditory and olfactory organs. The premaxillae and mandible both have short spinelike Just behind the mandible is found the hyoid arch. coracoid. exoccipitals. The upper jaw The develops from a cartilaginous pterygoquadrate process into the pair The lower jaw is the of premaxillae and pair of maxillae bones. and the neural arch over the spinal cord which lies in the neural canal. TRUE FISH 465 is The encloskeleton is principally bony and composed of skull. The ethnoids. known as the axial portion. parietals. vertebrae. The skull is very largely bone with some cartilage. operculum. mandible. . or cervical region are fused together. The skeleton of the paired fins is known as the appendicular skeleton. as gill or hranchial arches. teeth.PISCES. floor of the mouth. The first five vertebrae of the neck pelvic girdles. and bony fin supports. 'basisphenoids. There are haemal arches supporting haemal spines on the ventral side of the posterior vertebrae. The bones are arranged bilaterally. of the frontal hones. The pectoral girdle. supraoccipital. visceral skeleton supports the gills and includes the jaws. and the parapophyses extend laterally from the centrum and support the ribs. The next five arches support the four gills and are known operele. but the remainder are separate and are called amphicoelous because each end of the centrum or body is concave. made up of scapula. postfrontals.


TRUE FISH 467 and infraclavicle. It consists Fron tal Supraocopital Postfrontol Parasph enoid Prefrontal |— Paris tal Epiotic — Po St temporal Nasal Pterotic Premaxilfa -Hyomandibuhr -Opercular r-Entopteryyoid L acrimal Suborbital Ectopteryg old Maxil/a- Denfary ^Metop terygoid Quadrate Symplectic A rticular Angular Ceratohyoid Preopercular Subopercular Brancti/osteyal Pay Hypohyal Urohyal Pig. sists of a has a rudimentary pelvic girdle. and distal to this. terminates in a sharp spine. Inf er opercular —Lateral view of tlie sltull of tlie yellow perch. Perca flavescens. but is not the only function of the voluntary muscular it is an important one. a row of radial elements. 255. Posterior to fin. Galloway. are much stronger in the bullhead than in the perch. The anal fin and the dorsal fin are supported by interspinous bones.) PISCES. In addition. supports the pectoral fin which conrow of hasal elements. Muscular System and Locomotion Locomotion system. In addition to this function. . These it has a poisonous secretion with which to inflict wounds. and has the posterior edge serrated. The most anterior radial is completely ossified. (Drawn by B. these in the midline a platelike fusion of the basals of the The radials are all fibrous. The anterior ray of the dorsal is a bony spine. but the perch does not. The bullhead mesoclavicle. of two similar ischiopubic plates united is in the middle.

In connection with the eral distinct muscles. and guide and in guiding the body as it is forced The fins act somewhat as a combination keel "Without them the fish is unable to hold its upright itself through the water. CEREBRUM PINEAU BODY OPTIC LOBE CEREBELLUM AUDITORY N. fsl. M. GLOSSOPHARYNGEAL N. The myotomes are modified in the regions of the paired The action of the fins serves fins to supply certain muscles to them. mouth and feeding process there are sevThe adductor mandihularis elevates the lower jaw while the geniohyoid ancf mylohyoid raise the floor of the mouth and tongue. N. position and rudder. OUFACTORY BULB OI_FACTORY TRACT OPTIC N. POSTERIOR LOBES MEDULLA OBLONGATA VAGUS IM. to help in equilibration through the water. —(Drawn viewTitus Evans from dissections of Ameiurus natalis Le Dorsal of brain and cranial nerves by by Rose Newman. TROCHLEAR OCULOMOTOR PACIAL- TRIGEMIMUS N. since most of the power is delivered by lateral strokes of the tail against the water. 256.468 TEXTBOOK OF ZOOLOGY certain muscles are specialized for feeding respiration.) Sueur. and others to assist in The segmental myotomes divided dorsally and ven- trally by the lateral line are the chief muscles of locomotion. There are eight different sets of muscles connected with the respiratory movements of the opereula and gills. . Fig.

256). 257. three are the large posterior lobes. cerebellum and is quite medulla oblongata lies just posterior to the The central nervous system. of cranial nerves emerging from various strictly motor in of these have strictly sensory function. fingerlike pineal body (epiphysis) which is the vestige of a third or median eye. The bullhead has forty-one pairs of spinal nerves arising segmeutally from . Rose Newman. lamprey or shark. Also partially covered by the cerebellum The is the midbrain which is divided into two rounded optic lobes. Ca3 AND FACIAL N. and four have both sensory and motor function. The diencepMlon. due to the large posterior lobes (tubercula acoustica) on each of its anterolateral positions. composed of the brain it is in the OI_FACTORV TRACT OPTIC NERVE C2) CEREBRUM OCUL. This is the fourth ventricle. On the ventral side of the diencephalon is the optic chiasma where the optic nerves meet. fe) GL_OSSOPHARYNGEA1_ N . Two peculiarities of the brain of Ameim^us are the There are ten pairs large cerebellum and Three levels of the brain. -Ventral view of brain and cranial ner^^es of (Drawn by Titus Evans from dissections by Ameiurus natalis Le Sueur. is a diamond-shaped slit which leads into the cavity of the brain. which is immediately posterior to the cerebrum. Dorsally. TRUE FISH 469 Nervous System and spinal cord. is covered dorsally by the large dome-shaped cerebellum (much smaller in the perch). and behind this are the inferior lobes with the stalklike infundibulum joining the glandular hypophysis. CT) ABDUCENS Nt (6) MEDUUl_A OBUONGATA AUDITORY N. The is a little more developed than cerebral hemispheres are closely fused with the olfactory tracts which extend by long tracts anteriorly to the olfactory bulbs (Fig. function.(9) VAGUS N.OMOTOR HYPOPHYSIS Ni C35 OPTIC UOBES NFERIOR LOBES TROCHL-EAR TRIGEMINIES N. <'lO'> Fig.) prominent. between these lobes. 0*5 N. Extending dorsally from the roof of the diencephalon is a slender.PISCES.

Ameiurus. and guard the The details of the reproduction and nest until the eggs hatch. The sense of taste is highly developed and is centered in the numerous and well-developed taste huds which are distributed on the inside and outside of the lips. even to the tail. lay the eggs in the nest. and is better for detecting motion than for recognizing objects. and other fish have a well-developed pressure and water-vibration sense centered in the lateral line system. breeding are not so well known in Ameiurus natalis as they are in . — the ear structures serve in the sense of equilibrium. House. The eyes are small and without lids but have fair power of vision as this sense goes in fish. perch.) Fig. fry. B. TEXTBOOK OF ZOOLOGY Each has a dorsal raraus. perch.470 the spinal cord. The focal distance is between twelve and eighteen inches. and in gTOups over the external surface of the body. in the lining of the first three gill slits. fry. on the barbels. 258. eggs with embryos. The fish does not have a sense of hearing. is particularly keen in the lips and Reproduction and the Life History The bullhead. sunfish and many other common fish build nests of one sort or another. The sense of touch is dis- tributed over the epidermis but barbels. Eggs of trout with well-developed embryos. and recently hatched (Courtesy of General Biological Supply A. and a ventral one extending out to certain parts of the body in the region. or branch.

the brown bullhead. days. They were 4 mm. D. by the fourteenth day. where they remained for two ton. C. Ninety-five per cent of them hatched in five days with the water at 77° F. The observations were made on a pair in an aquarium in AVashing- A. By the end of the seventh day they were swimming actively and most of them collected in a school just beneath the surface. leaving the slate bottom bare. These operations were continued until the fry (newly hatched fish) swam freely. then they began to swim. It is also reported that they ate finely on the sixth day and had enormous appetites after the eighth day. keeping them agitated constantly by a gentle fanning motion of the ventral fins. The egg masses are also sucked into the mouth and then blown out with some force. Both parents assume responsibility in caring for the eggs. At the age of two months their average length was 50 mm. On July 5 about two thousand eggs were deposited in four masses. afterwards scattering.PISCES. The young remained in masses until six days old. a brief description will be given for the latter. ground liver . long when hatched and had attained a length of 18 mm. nehulosus. They made a nest on July 3 by removing with their mouths more than a gallon of gravel from one end of the tank. TRUE FISH 471 This being the case and since the two are very similar.

There are other exceptions to the general characteristics of this A large group of salamanders. and never have paired fins like fishes. and salamanders usually lay their eggs in water. the plethodontids. was group these together under the classification based on habits rather than on structure. Today the name is restricted to a group of vertebrates which we know as frogs. Except in caecilians. The frogs. Kicord's frog. toads. salamanders. The amphibians are cold-blooded vertebrates. 472 . They are intermediate between fishes and reptiles. They have a moist. These animals and a few others such as the large South American frog ceratopharys. Eleuthero- dactylus ricordii. They have small scales between their transverse body rings. which has dermal bones or ''scales. and as soon as such animals as the seal and crocodile were studied structurally they were removed from the class. These land eggs lack the calcareous shell of reptile and bird eggs. they have paired limbs. naked skin lacking the protective hair of mammals or the feathers of birds. are examples of species that lay their eggs on land. and caecilians.CHAPTER XXVIII CLASS AMPHIBIA (By Ottys Sanders. both of which are richly supplied with blood vessels. Amphibia. in contrast to the warm-blooded mammals and birds. do not have lungs even as adults. These develop into tadpoles or larvae breathing with gills before metamorphosing to become adults which breathe with lungs. are wormlike burrowing creatures of the tropics. toads. none of which has been reported from the United States. usually with fingers and toes. although these are not usually seen unless a dissection is made. and their respiration takes place in the mouth cavity and through the skin. Plethodon glutinosus. The caecilians. of course." are the only ones of the class to have scales. diverse class. and the slimy salamander. it was natural for Linnaeus This. A few species of frogs and salamanders lay their eggs on land and pass their entire development in the egg. Southwestern Biological Supply Company) As class there are many vertebrate animals which lead an amphibious to life.

which had a skull 4 feet long and a total length Among living amphibians. Megalobatrachus japonicus. 259. Florida. Adults measure only % to about 18 inches long and of Africa reaches a % . Bufo marinus. The smallest frog in the United States is the swamp tree frog. which and commonly grows to be Fig. the "hellbender. V. the largest salamanders to a length of 5 feet. 473 —While most modem Amphibia are small creatures. C. which ranges from North Carolina to southern of an inch in body length. is legs are extended. as. and attains a body length of 8% inches. Pseudacris ocularis. and a larva showing external gills on the right. Ichthyophis glutinosus. which attains a length of about 30 inches. grows In the Southwest. are Siren lacertina. may grow to be over 7i/2 inches in body length.) eggs on the left. Mosby Company. adult female. published by The Sarasins. guarding her AmpMuma. the largest of the true toads. paleon- tological species reached large proportions. the Congo eel." Cryptobranclnis. : —The caecilian.CLASS AMPHIBIA Size. with a total length of 16 to 18 inches when the The giant toad or marine toad. the Mastodonsaurus. for example. salamander of Japan and China. the giant of probably 15 or 20 feet. Partly after (From Atwood Introduction to Vertebrate Zoology. The goliath frog body length of nearly a foot. larger than their northern relatives. while southern bullfrogs.

or red results these predominate. pigment distributed m response modified from Noble. temperature. the small grayish canyon tree frog. These color chajiges are .^^^^Ji^ A. in general. reeds and cattails in the toad. species of toads may live about thirty years. are but a few examples of beautiful species. y. (Redrawn and Their different colors are due primarily to various combinations of three kinds of pigment cells in their skin. these pigment cells produce color changes which are initiated by various stimuli. moisture. which makes green tree frog. Green color results from the from guanin granules wherein all the light rays Different arrangements of escape absorption except the green. frogs probably Coloration.474 TEXTBOOK OF ZOOLOGY far as As ten to known.. and the chemical composition of the frog's habitat. bell-like calls The bright from the summer months. when cells. Amphibians possess considerable ability to change color. is longer than the smaller species.:. Yellow from the action of lipophores contained in spherical reflection of light and white from the guanophores. 260. with its red warts. The black melanophores are branching pigment cells which may contract or expand. B. Eyla cinerea. the varicolored common Hyla versicolor. such as light. —Amphibians as a group are very colorful. Bufo pnnctatus.) North America published by McGraw-Hill Book Fig. the skin appears black or brown. seem to live fifty-two years. with its orange groins. —Melanophore from Rana temporia. the length of life of Amphibia ranges from The larger ones. and. pigment contracted. Amphibia of Company. Some less. and many of the tree frogs equal or surpass the chameleon in this respect. to light.

Scaphiopus. . Burrowing frogs and toads. Scaphiopus dartlike spades.CLASS AMPHIBIA directly beneficial to the animal closely its surroundings 475 when they help it to resemble more and thus avoid capture. by Thos. showing the dark-colored. (Photograph coucliii. smooth skins. Amphibians have a soft. The Skin.) Fig. (Photograph by Thos. Aquatic — Fig. such as the spadefoot toad. Scaphiopus coucliii. — Spadefoot tlie toad.) and forest-inhabiting frogs and toads have a smoother skin than species which live in drier places. 262. 261. moist skin which is kept in that condition primarily by a rich supply of mucous glands. Mebane Jones. showing the shape of the pupil of eye. also have thin. Mebane Jones. —Feet of spadefoot toad.

During hibernation. in the male sex. It has patches of vascular villosities on the thighs and sides has been named the ''hairy frog.) North lungs. practically all of the respiration of frogs is taken care of through the skin. and therefore seek moist places where they may to such an extent that it absorb water through their skins. has developed a strange aid to respiration. portant function is its use as a respiratory organ. which." 5^ of (Redrawn and modified from Noble. Since amphibiajis have moist skins. A most imlight but also has other functions. with greatly reduced Pig. publislied by McGraw-Hill Book Company.476 TEXTBOOK Of zoology The skin not only protects the underlying tissues from excessive With its pigment it helps to regulate temperature by transformijig light into heat. Most of them are nocturnal in ." These villosities help provide sufficient oxygen for its increased metabolism during the breeding season. one large group of salamanders. the plethodontids. As previously mentioned. In Africa there is a frog. — "Hairy ^ frog. they are in constant danger of drying out. lack lungs and use the skin and buccal cavity for respiration. 263. AmpMiia America.

frogs and toads will seize a moving object without much examination. dragonfly nj^mphs. spiders and other small invertebrate animals. it be retracted into the head and by this action they help to push food in the mouth cavity toward the esophagus. Nor are amphibians immune to disease and gross infestation by parasites. while the tadpoles eat either animal or plant food. The power of regenerating lost parts Young tadpoles may is one way in which Nature aids the group. birds. and other aquatic insects. or in other situations where they can protect themselves against this constant threat of desiccation. and mammals feed upon the adults and young. of the adults consists primarily of living insects. gators. allilarvae. the toads quite often stalk their prey and inspect it. Amphibians depend to a large extent upon their sight in detecting food. Amphibians can go for a long period of time without food. During the hibernation season and breeding season most salamanders may live for months. — . Many large frogs and a few smaller ones are cannibalistic. and fish appreciate their good flavor. — The enemies of amphibians are many. Powers of Regeneration. If a disagreeable insect. The food worms. although adult The axolotl larva of ently unable to regenerate lost appendages. Man Enemies of Amphibia. snails. Food and Feeding Habits. and frogs do not feed. enjoys the hind legs of frogs. Small crustaceans devour the gills of salamander Snakes. the toad has a wide esophagus. In food for giant their larval or tadpole stages they are a delicate water bugs. While. larvae of water beetles. Their eyes can swallowed. frogs and toads are apparregrow limbs or tails. turtles. such as a stag beetle with strong mandibles. Most of the frogs and toads and many salamanders utilize their eyeballs in swallowing food. in general. and there is an increasing demand Man also destroys amphibians by polluting the streams where they breed.CLASS AMPHIBIA their habits 477 logs. fortunately. —Adult frogs and toads eat animal food. and his automobile kills countless toads and frogs on the highways. in crevices or and therefore can be found during the daytime under burrows in the earth. for these as food. is can be disgorged because. Tadpoles and experiments made on axolotls (larvae of the tiger salamander) have demonstrated that they may live for about a year on the food stored in their own tissues.

The air is forced back and forth between the . of the most protective is weapons that amphibians have. — The amphibians were probably the The calls of first vertebrates to de- modern frogs and toads are very distincMost of the tive. Colorado. and their habit of remaining immobile frequently causes them to be overlooked. Bufo marinus. —Amphibians have few ways of protecting themselves from their enemies. which ranges from Texas to Patagonia. Voice. It is night with their choruses. species practice death feints lungs. and elsewhere. although certain species other times. known as parotoid glands. Their coloration often blends in with their surroundings and camouflages them. in the case of salamanders particularly. One ever. toads. the ability to break off their tails and escape. produces one of the most virulent poisons known among amphibians. many of which have large glands on their shoulders. An animal that has attempted to bite or swallow a toad and felt the effects of the poisonous secretion of the parotoids upon the mouth tissues will not soon forget the experience. during the breeding season that the air resounds at may croak at The croaking of frogs and toads is usually done with the mouth and nostrils closed. their writhing and twisting movements when captured make them hard to hold. The mucous glands of frogs and salamanders make them slippery. The secretions of toads are ordinarily quite harmless to man. The largest known toad of the North American continent. stream. have been used by the Indians of Colombia for poisoning their ari'ows. Means of Defense. howThis is the secretion of their poison glands. and the primary function of these calls seems to be to attract females and other males to the pond or velop a voice. has been used extensively in experiments for studying the nature of this regeneration. croaking is done by the males.478 the tiger salamander. each species having its own particular call. unless they happen to get into his mouth or eyes. A few salamanders have. Dendrohates. which is found in Texas. especially effective in the case of toads. New Mexico. TEXTBOOK OF ZOOLOGY Amhystoma tigrinum. in addition. Many and some swell up by inflating their making themselves more difficult to swallow. There are records of dogs which have been killed by Glandular secretions of certain South American its secretions. however. and.

in tropical climates. however. Vocal sacs. usually lay their eggs in cracks and hollows in its logs. which undergoes its entire development on land and cannot swim when placed in water. salamanders lay on land. which lives in the cold mountain streams of Washington and the northwestern United States. as are the eggs of is its relatives and the tadpole stage their eggs passed in the egg. Many in Mexico. Some salamanders and caecilians. give While the majority of amphibians lay their eggs in water. for its sound would not carry above the noise of the mountain streams. toads. during the rainy season. usually occur during the spring months or. Arthroleptella lightfooti. A few frogs. there are many The eggs of the Texan cliff frog. lying either in the floor or at each corner of the mouth. when present usually call. Salamanders often come to the pools much earlier than do — make the frogs and toads and may also begin their egg-laying earlier. called the breeding season. have given up their voice and reduced their lungs. the on land. are laid birth to living young. causmg them to vibrate. such as Plethodon cinereus. Among salamanders in many species. Apparently voice would not be as useful to this species as to frogs living in quieter places. such as Ascaphus. and young pass through tadpole or larval stages. Breeding and Egg-Laying Habits. and most of them are silent throughout their existence. and their eggs are fertilized by body of the female. none of the salamanders use voice in attracting mates. Some species of Oriental frogs are reported to lay their eggs in trees high out of the water.CLASS AMPHIBIA 479 lungs and mouth over the vocal cords. So far as is known. have the eggs fertilized before they are the male after they leave the laid. Most amphibians are oviparous. Plethodon glutinosus. Salamandra salamandra. puff out to increase the volume of the make resonating chambers which Bullfrogs quite often call while under water. the males deposit spermatophores containing sperm which are picked up by the females and provide internal fertilization. Eleutherodactylus exceptions. lays often in the walls of caves. and salamanders periodic migrations to ponds and streams for the purpose of egg-laying. A few species of salamanders such as the fire salamander of Europe. latrans. These periods. . There is also reported a South African frog. Species in the Southwest. eggs in moist places. The slimy salamander. Frogs.

glandular masses at the base of the tail or elsewhere may the females. two frogs from the northern part of South America and Central America. Hibernation. —Adult Ambystoma tigrinunij tiger salamander. Gastrotheca. carry their eggs in a dorsal sac or brood pouch which is found in the female. Phyllolates and Dendrobates. 264. ^Amphibia are more or less adapted to their environment. tween males and females of a —Secondary sexual characters comspecies. which is larger than that of the female. their tadpoles on the Fig. Familiar secondary sexual char- acters are the nuptial pads of male frogs. the male carries the eggs wrapped around his legs until they hatch. In the case of the obstetrical toad of Europe (Alytes olstetricans). In some salamanders the teeth may elongate. be- These differences may be both structural and physiological. bringing low temperatures and — . (Photograph by Sanders. when winter comes. transport pass the rest of their tadpole stage back of the male to the stream where they and metamorphose. the male has the columella of the ear pushed through the drum to form a noticeable projection. and. the the tympanum.480 TEXTBOOK OF ZOOLOGY of South America. The marsupial frogs The Amazonian frogs Pipa and Protopipa carry their eggs and tadIn poles in individual dermal chambers on the back of the female. is One of the be present in the males and absent in most bizarre secondary sexual characters In this frog found in an African frog (Petropedetes newtoni). pose those differences. in the male bullfrog. their swollen thumbs size of during the breeding season. These sexual characters of the male may be various. and. the ease of a small frog (Bhinoderma) in Chile. eggs are carried in the vocal pouch of the male where they metamorphose and hatch as fully formed young. exclusive of the reproductive organs. in others.) Secondary Sexual Characters.

one of the most common salamanders (Photograph by Sanders.— CLASS AMPHIBIA a scarcity of food. the depth to which they go depending on the type of They may go as far as 18 inches underground in sandy soil.) in Texas. After establishing itself in hibernation quarters the amphibian reduces all vital activities to a the skin. or crawl into cracks and crevices. Frogs crawl into the ponds or other damp spots. 266.) Ambystoma tigrinum. most of of 481 them hibernate. under rocks soil. reducing their activities until the severest weather is over.«».•' T Fig. Salamanders may bury themselves in the mud. Respiration is carried on and the body in its dormant state secures entirely through the slight amount of nutriment needed from the food stored in its In some hot countries during the dry. dig into the ground under logs. graph by Sanders. . in rotting tree stumps or in burrows in the mud in the bottom ground.(*'• "^m^: Ambystoma texanum. ians aestivate in a protected moist place. —^Axolotl larva of the tiger salamander. torrid season amphibtissues. 265. minimum. <«»'. in running streams. '*g! *. (Photo- 'i'-S*. Fig-. Toads burrow into the ground.

— 482 TEXTBOOK OF ZOOLOGY Classification frogs. 4th edition. Thos. In the United States there occur about 79 species of salamanders and about 70 species of frogs and toads.* Many of According to the Check List of North America Amphibia and Reptiles by Stejneger and Barbour. Typhlomolge rathbuni. Mebane Jones. grapii (Photo- by Sanders. and about 60 species Fig.) (Photograph by of caecilians. Pseudacris streckeri. the blind cave salamander of Texas. toads.t ' '^M Fig. Strecker's ornate chorus frog. None of the caecilians have been reported from the United States.900 known species of living and salamanders in the world. 268. 267. There are estimated to be about 1.) ''^^i. .

. : Some or absence of characters used in classifying salamanders are the presence gills. . color markings. presence or absence of parotoid glands and their shape presence or absence of tympanum presence or absence of . presence or absence of lungs presence or . number of digits position of teeth presence or absence of a nasolabial groove plantar tubercles shape of vertebrae form of cranial bones and cartilages. length. The student interested in classification and identification of species should consult appropriate keys for the various groups of Amphibia. Hyla crucifer. shape of head.) nature of skin. (Photograph by Thos. . shape and appearance of body. Fig. . absence of ypsiloid cartilage. 483 The Southwest contains a large proportion of all of these. 269. the shape of the vertebrae. . number of costal grooves. . presence or absence of teeth and their situation. shape of the sacrum and pectoral girdle shape of pupil of the eye presence or absence cranial crests and their shape . . — Tree frog.: CLASS AMPHIBIA these species are subdivided into several subspecies. Some characters used in classifjdng adult frogs and toads are color markings. There is appended at the end of the book a list of references dealing with this class of animals. of adhesive discs at the ends of digits. Mebane Jones. . . either external or internal. length of body and of hind limb.

Bhyacotriton olympicus (1 species). This so-called "hellbender" ranges from the eastern states west to Iowa. Dismal Swamp. Triturus (5 species in the United States). Typhlotriton spelaeus (1 species). Pennsylvania to Louisiana. species in the Southwest are: the Tiger . Gyrinophilus porphyriticus (1 species). Range from New England to Texas. The blind salamander of the caves of Typhlomolge rathbimi (1 species). The blind cave salamander of Texas. salamander has only four toes. Calif. Stereochilus marginatus (1 species). A. Distributed over almost the entire United States. the three-toed congo to eel. meridionalis. Eastern states west to Kentucky. south to Louisiana. Suborder Ambystomoidea Family Ambystomidae Amby stoma (13 species). Southern Canada to the Gulf of Mexico. texanum) Marbled salamander {A. Common . Leurognathus marmorata (1 species). ranges from northern Florida Family Plethodontidae south to Georgia. Manculus quadridigitatus (1 species). salamander {A. tigrinum) the Texan salamander {A. Missouri and Arkansas. North Carolina to Texas. Desmognathus (5 species). Most common species in Southwest is D. west is Triturus viridescens louisianensis. Family Amphiumidae Amphiuma (2 species). The common newt of the SouthThe other species represented is T. and the Dicamptodon ensatus Suborder Salamandroidea Family Salamandridae (1 species). Virginia to Georgia. eastern states westward to Illinois. tridactylum. eastern Texas. Eegion of San Francisco. Common in the Southwest is P. Wash. North Carolina mountains. Brimley's triton. Canada to Louisiana. Hemidactylium scutatum (1 species). brimleyorum. Plethodon (15 species). the slimy salamander. Another four- .484 TEXTBOOK OF ZOOLOGY A List of Families of the cited Amphibia in the United States dis- The ranges below are not exact but give an idea of the tribution of the genera. Fseudotriton (2 species). Order Caudata (Urodela) (Tailed Amphibians) Suborder Crypto'branchoidea Family Cryptobranchidae Cryptohranchus alleganiensis (1 species). This dwarf Eurycea (6 species). glutinosus. toed salamander. opacum). Olympic Mountains.

Syrrhophus (2 species). insidior. Leptodactylus labialis (1 species). widespread throughout eastern and central United States. Species of Bufo are distributed over the entire United Among common species in the Southwest are B. the species which seems to be the most common in southern states is N. woodhousii. B. The worm salamander. and B. describing two new species from Alabama and two new species from Louisiana. fowleri. Family Proteidae (with external gills and 2 pairs of limbs) Necturus. without hind limbs) Suborder Meantes Family Sirenidae (with external Siren (2 species). when Suborder Procoela Family Bufonidae (Toads) Bufo (17 States. gills. Both species limited to Texas. of New Orleans. South Carolina to Florida.CLASS AMPHIBIA Flethopsis wrighti (1 species). S. Washington and a few other points on the Suborder Anomocoela Family Pelobatidae (Spadefoots) Scaphiopus (5 species). 485 Oregon. Percy Viosca. Pacific Coast. lacertina and S. Aneides (4 species). four species in the Southwest. marinus. B. (1 species). On All on the Pacific Coast. Batrachoseps Pacific. ' Pseudobranchus striatus (1 species). species). com- B. one species in Florida. (2 species). Eydromantes platycephalus Suborder Proteida Yosemite salamander. Both S. punctatus. According to a recent revision of the genus by Mr. B. Order Salientia (Anura) (Tailless Amphibians) Suborder Amphicoela Family Liopelmidae Ascaphus truei (1 species). Found only in Texas. Both species on the Ensatina (3 species). B. B. pactilis. Eleutherodactylus (3 species). Family Hylidae (Tree Frogs) Acris gryllus (1 species). The common large Necturus from the Great Lakes region is N. Family Leptodactylidae (Robber Frogs) valliceps. debilis. is increased from three to seven. Eastern Virginia to Texas. cliff One species in Texas (Texas frog). One species the pupils of whose eyes are vertical in the East. . B. The cricket frog. in daylight. one each in Florida and California. which extends into Texas. These are the spadefoot toads. cognatus. heyeri Viosca. intermedia are found in the Southwest. including the Southwest. Pacific Coast and in southeastern states. one species in Arizona. maculosus. the number of species in the U.

In southern Texas. Within recent years businesses have developed which are devoted to supplying bullfrog legs. species in the Southwest in- H. bullfrogs crayfish. B. man of food Dried frogs and toads have been used in China both as a source and for medicinal purposes. The axolotl is used in Mexico as food. The recently described P. Ranges from Virginia to Texas. H. Frogs are used throughout the world as an article of food by as well as by other animals. clamitans. nigrita) are common Southwest. H. such as our Necturus. and water dogs. thus becoming valuable aids to the farmer in controlling noxious insects. flooded rice fields of Louisiana. Family Brevicipitidae (Narrow-mouthed Toads) Bypopachns cuneus (1 species). catesbeiana. bullfrog legs have been a favorite food for years. They are the most colorful of all the frogs. and its high-pitched staccato chirp is one of the earliest to be heard at breeding pools in Texas. Throughout the same regions as Acris. grow fat eating insects. ranging throughout Texas. is a very colorful species. Common crucifer. H. and the demands for these from all parts of the country have been so great that it may become necessary to afford some protection to prevent the rapid depletion of these animals. pipiens. arenicolor. Hyla (12 clude: Various species in all of the United States. cricket frog Various in the subspecies of the swamp (P. cvnerea. are reputed to have a good flavor. In the eastern United States. large quantities of the leopard frog and wood frog are consumed. Com m on species in the Southwest are: B. Various species occur in all parts of the United States. squirella. northward souri and Indiana. Microhyla (3 species). Most of these attempts have been failures because of the high overhead cost. Suiarder Diplasiocoela Family Ranidae (True Frogs) Baria (18 species). Attempts have been made to operate frog farms and raise a supply.486 Fseudacris TEXTBOOK OF ZOOLOGY (6 species). and other small animals. B. In the southern states. B. streckeri Wright. to Mis- Economic Importance The entire group of Amphibia are of considerable economic value In the because they feed to such a large extent on insects. sphenocephala. H. It is reported that toad skins have been used in Japan and elsewhere for making a fine type of . versicolor. species).

crayfish. but swimming and crawling of the salamander division of about at night. Insect larvae. 270. . THE MUD PUPPY Necturus maculosus. Fig. Mebane Jones. spending most of the time in the mud at the bottom. and occasionally fish comprise much of its food. as well as larval stages. the mud puppy or water dog.) NECTURUS MACULOSUS. (Photograph by Thos. 487 Dried salamanders have been used as a vermifuge. It comes ashore only occasionally. frogs.— CLASS AMPHIBIA leather. Adult frogs and salamanders. Bufo valUceps is a common toad. are widely used as laboratory animals. worms. and is the one most commonly used for laboratory study. is a very com- mon example Amphibia found from the Mississippi basin eastward. It lives in ponds and streams.

testis.) . right lung. subclavian vein. pylorus 37. 0. J. vesical vein. . stomach S5. . Fig. H. A. IS. IS.0. 11. 25. 19. 29. 26.. SO. 22. gall bladder. 28. bulbus arteriosus 7. . gill slit. . spleen 36. S2. caudal vein. . 1)2. cloaca. 10. mesonephric duct. kidney. ileum. — . hepatic sinus 5. afferent branchial artery. 271. pulmonary vein Sk. Jil. external jugular vein. pelvic vein. JiS. femoral vein. afferent branchial artery. . large intestine. mesenteric vein. common cardinal vein or duct of Cuvier 3/. liver. . external gills. S9. SO. . internal jugular vein. subclavian 6. left auricle. 16. S. 1. 11. It. 12. ventral abdominal vein. 21. urinary bladder. 15. 27. 23. hepatic portal vein.. dorsal aorta. gastrosplenic vein. I S. renal portal vein. afferent branchial artery. . postcaval vein. duodenum S8. III artery. ventricle i. pancreas. II 2. . postcardinal vein. House. gastric artery. 21/. (Courtesy of General Biological Supply . Diagram of dissection to show principal organs of Necturus. left lunng SS.

possessing only the cardiac porThis tion anteriorly and the posterior narrowed pyloric portion. although not highly functional. the glottis. Also lungs are developed for aerial respiration. The retention of external gills a very marked Food and Digestive System This animal does make use is quite inactive and requires relatively little food. other small crustaceans. larval feature. and occasionally fish eggs for food. vomer. by the newly veins are still partially taken over Necturus remains in a larval condition throughout its life. and the dentary and splenial bones of the lower jaw. becomes sexually mature. two auricles The heart has become three-chambered.CLASS AMPHIBIA 489 is The group and of vertebrates which Necturus represents of par- ticular interest because of its transitional position between aquatic session Necturus is aquatic and fishlike in its posand use of external gills. by lateral terrestrial forms. The body is used like that of a fish in SAvimming that is. strokes of the tail against the water. This portion of this arch known as the duct of Botallus. insect larvae. and reproduces without metamorphosis. More posteriorly. It of several aquatic inhabitants including crayfish. fairly and with a Teeth are located on the premaxillae. but their function developed post cava. The internal nares enter the mouth cavity as a slit on each The mouth is located in the anterior. some minnows. The sixth or last pair of aortic arches of the primitive series gives off a pulmonary artery and still retains the connection is to the dorsal aorta. Such a condition is is referred to as neoteny. . in the lateral walls of the . snails. leeches. and palato-pterygoid bones of the upper jaw. terminal position wide gape. The posterior cardinal is present. The stomach has the typical shape and appearance of this organ in the lower vertebrate groups. pharynx and joins the anterior or cardiac portion of the prominent stomach. two dorsal rows of teeth near their posterior termipharynx are the two pairs of gill slits or pharyngeal clefts. The tongue is broad and only slightly movable. although it has only three arches. side between the nations. It is terrestrial in the de- velopment of pectoral and pelvic girdles and limbs for crawling. Still more posteriorly there is a very small inconspicuous pharjoigeal prominence with a The esophagus leads posteriorly from the tiny slit. there are now instead of only one as in fish.

hepatic and pulmonary veins. renal portal. The bladder duct which is obscured by is and the mesenteric vein. the tine first S-shaped turn of the tube. and the atrium of divided into two parts to keep the systematic and aerated. With certain modification of the situation in fish. The more coiled This empties part of the small intestine following this the ileum. and urinogenital products from the wolffian ducts. pulmonary blood partially separated. somewhat broadened large which opens common receptor of faecal matter from the intes- der and miillerian ducts. . Blood in passing through the heart may be traced by entering the sinus venosus from systemic veins. intestine. The panhile creas is divided into slender lobes and lies in the vicinity of the juncOne lobe extends to the tip of the tion of stomach and duodenum. The gall bladder is a membranous sac attached to the margin of the liver (usually at The liver is connected with the duodenum by a a mass of pancreatic tissue. pass by way of sinu-atrial (sinuauricular) valve to portal. Another slender lobe spleen which lies extends posteriorly in the mesentery that supports the anterior part the right side). The right atrium receives the systemic blood and the left atrium. in that the number of functional aortic arches is reduced. Both of these lobes join the mass of pancreatic tissue around the bile duct and the pancreatic ducts of ileum enter the duodenum at this level. whose lumen it joins. the system consists of heart. There is pulmonary the heart is vessels are added to supply the lungs. right atrium. is duodenum. the pulmonary blood by way of pulmonary veins. an elongated. thence through the atrioventricular valve to the ventricle which it enters simultaneously with aerated blood from left atrium.490 TEXTBOOK OF ZOOLOGY leads into the anterior section of the small intestine or which includes the into the short but into the cloaca. aortic arches. dark-colored and somewhat serrated organ lying in the ventral portion of the body cavity. urinary bladThe urinary Madder is a thin-walled sac hanging at the ventral side of the cloaca. the sinus venosus which joins the right atrium and the conus arteriosus which leads from the ventricle to the ventral aorta. as well as systemic. Circulatory System a partial conversion from the straight branchial type of circulation of the fish to the pulmonary type of the terrestrial vertebrates. The heart consists of right and left atria (auricles) and one ventricle with the two usual accessory chambers. dorsolateral to the stomach. systemic and pulmonary arteries.

performed through gills and pidmonary through the . manders this number may be referred to as four but considerable The first (anterior). and internal carotid arteries. The lateral veins of sharks form a similar cut-off but enter the duct of Cuvier instead of the hepatic portal system.* Respiratory System and Breathing The lungs. and third modification has occurred. As will likely be remembered. usually by reducIn sala- Even teleost fish have only four branchial arches. The two pelvic veins pass ventrally to meet each other at the midventral point of the pelvis and this union forms the ventral abdominal vein which either enters the liver or a branch of the hepatic portal before it enters the liver. of aortic arches in vertebrates vertebrates. Chicago. original arches have been rearranged and combined to form the common external.CLASS AMPHIBIA 491 The blood is expelled from here through the conus arteriosus into the ventral aorta. primitive. second. may the wet skin branchial through the be divided into cutaneous. The special modification of the veins is centered around the development of the post cava which is formed posteriorly by the junction The pair of posterior cardinals. is known as the duct of BotaUus. There are several parts to the latter arrangement. in . characteristic of fish. They usually join the post cava some distance anterior to the kidneys and parallel the aorta to the heart where they enter the ducts of Cuvier. which are of urinogenital veins. is six. from the point where the pulmonary branches off. one on each side. The portion of the sixth aortic arch which continues on dorsally to join the aorta. The sixth arch is modified to supply a large pulmonary artery from each side to the respective lung. respiration .. are retained also but are greatly reduced. and embryonic typical number This number is modified in most adult tion. A pelvic vein branches from the femoral on each side before it joins the renal portal. the theoretical. system of Necturus may be found Denoyer-Geppert Co. The small external nares lead by way of passages to the slitlike internal nares which open into the mouth between the posterior ends of the two Helpful Stuart: Anatomy Illustrations of the circulatoi-y of Necturus maculosus. thus providing a cut-off in the course of the venous circulation in going anteriorly from the posterior limbs. The fourth and fifth supply the external gills with the fourth becoming the systemic arches which meet dorsally to form the dorsal aorta. Another modification is the pelvic-ventral abdominal complex which connects the renal portal arrangement with the hepatic portal.

TEXTBOOK OF ZOOLOGY The mouth is made airtight by the shape and The general portion of this cavity posterior to fitting of the lips. . Upon becoming filled with urine the bladder contracts and forces the urine back into the cloaca and from here it passes to the exterior by way of the anus. After the urine enters the cloaca it collects in the urinary bladder which hangs ventrally and serves as a storage reservoir. The glottis leads into a recess called the larynx and the two smooth-walled. Urinogenital System The following organs constitute this composite system a pair of mesonephric kidneys. Well back in the floor of it. the urinary bladder The kidneys are somewhat elongated and flat but thicker toward the posterior. The air is pumped into and from the lungs by the movements of the floor of the airtight mouth and change of posi- from this. arches 4 and 5 provides most of this blood supply to the gills. only vestigial in male) single and the mesenteries (mesovarium. Inside the kidney the Malpighian corpuscles. the angle of the jaws is the pharynx. suspended in the dorsal peritoneum and lying dorsal The kidney of the female is smaller than to the large intestine.492 dorsal rows of teeth. The Wolffian duct leads from the lateral margin of the kidney in either sex and proceeds directly from the posterior portion of the kidney to make a dorsolateral entrance into the cloaca. one pair of Miillerian ducts or oviducts (in female. is the tiny slitlike glottis in the midst of a slightly thickened laryngeal prominence. Branchial respiration accomplished largely by waving the highly vascularized external The capillary branching of aortic gills back and forth in the water. mesotubarium. that of the male. one pair of Wolffian : or mesonephric ducts (ducts of Leydig in male). tion of visceral organs within the is body cavity. ovaries in female) numerous vasa efferentia from testes. a pair of gonads (testes in male. the opening of which would receive only an object the size of the head of a pin. which in turn join the collecting tubules and they lead to the Wolffian duct. including glomeruli. are connected with the uriniferous tubules. saclike lungs extend posteriorly These saclike lungs have a fairly abundant vascularization (blood supply). and mesorchium). The pulmonary artery supplying the lungs is formed by a large branch from aortic arch number 6. cloaca.

When the ova reach maturity inside the ovary they escape by a rupture Due to the in its wall which frees them in the coelomic cavity. When the spermatozoa reach the cloaca and bladder they clump into bundles called spermatopliores.CLASS AMPHIBIA 493 In the male specimen the yellow or brown-colored cylindrical testes are located one in either side of the dorsal part of the body cavity and each suspended by a fold of the dorsal peritoneum. As these ova pass down the MuUerian tube (duct) they are met by spermatozoa. a mucous substance is added as a cover by the glands in the oviduct. The vasa efferentia. The Wolffian duct carries them to the cloaca. white oviduct or Miillerian cavity at each side of the other organs whose antube in the body terior end is suspended in the anterior portion of the body cavity and spreads into a wide membranous funnel called the ostium. In specimens with immature ovaries the eggs may be about the size of pinheads. medial side of the kidney (except at its anterior) and deliver spermatozoa to a longitudinal Bidder's canal just within. and are stored until breeding time. as does the urine. When fully mature each ovary seems to be a large sac full of large yellow mature eggs about the size of small peas. The varium. logs. fertilization occurs. There is mesentery which supports the oviduct is the jnesotuharium. In the female the pair of ovaries can usually be recognized by the presence of eggs of some stage of development in them. etc. shape of the body cavity and position of visceral organs these eggs to the anterior part of the cavity and the ciliated mouths of the two ostia receive them one at a time in each. This is connected with the medial ends of collecting tubules and through them the spermatozoa reach the Wolffian duct. in the water in small clutches of from 25 to 90 individuals. The embryonic stages are passed here and the larvae hatch out as tiny fishlike organisms. These fertilized cells in a pouchlike posterior part move called the uterus and after a few accumuby passing from the body by way of the cloaca and anus. of each oviduct which is late they are deposited . Each ovary is suspended from the dorsal peritoneum by a mesentery. the mesorchium. the meso- a prominent. and the spermatic blood vessels The vasa efferentia enter the are suspended in this mesentery. which are tiny sperm tubules about the size of very fine threads. coiled. These zygotes (fertilized eggs) are deposited by attachment to the under sides of rocks. A Wolffian duct (mesonephric duct) which serves both for conveying urine as canal well as spermatozoa is called a duct of Leydig.

is located at the mid-posterior position and an occipital condyle is . The axial portion consisting of skull. is the (flat when compared with large pair of ixirietals. Denoyer-Geppert Co. each of which bears a few teeth.* Skeletal System The skeleton is of these animals is classified as a bony skeleton but not completely ossified and a considerable part of it is cartilage. consisting of the two girdles with limbs constitute the essential parts of this system. depositing them here while the female follows and collects them into the cloaca by use of its swollen lips. at the side of the skull. dorsal surface of it is covered by a single. and extending beneath and somewhat lateral to this. The actual breeding and copulation activities (if any) do not seem to be very well understood. . irregular one which lies between the anterior part of the squamosal and the parietal. The braces. which is at the posteroThe foramen viagnum (large opening) lateral corner of the skull. are the palatopterygoid bones. Just posterior to this and somewhat covered by the frontal the vomer. the quadrate cartilage. the opistliotic. and the mucus which is secreted by the cloacal glands that lie at the sides of the cloacal aperture. At is the anterior tip of the frontal are the premaxillae which bears teeth. which appears more dorsally. quadrate hone. The skull is platybasic tive bones and broad) with a marked fusion and loss of primithe teleost fish. a small. The group is represented only by the prootic. the first vertebra. vertebral column and ribs. The act of transferring spermatophores is described as occurring in shallow water or on the muddy margin of the pond or stream by the male. located at each side of it for articulation with atlas. and the appendicular portion. Chicago. The principal part of the In Stuart: floor of the skull consists of the large flat •Helpful Illustrations of the urinog-enital systems of Necturus may be found Anatomy of Necturus maculosus. which articulates with the lower jaw otic and the squamosal. The anterior. posterior to which. There is a prevailing idea that the spermatophores are passed from the male to the female in the autumn and held in the genital tract of the female until the succeeding spring when the eggs mature and pass down the oviducts. which also bears teeth. the papillae there.. fused frontal bone. and another small one. Both the nasals and maxillae are absent.494 TEXTBOOK OF ZOOLOGY At about a year of age they are one and one-half or two inches in length with a stripe down the side which gives them a peculiar appearance.

In the lateral position where the ilium meets the two ventral parts of the girdle is a concave recess into which the head of the femur of the thigh articuThis recess is called the acetahidum. the scapula. ventral part is the puhic plate consisting of cartilage. Each rib has a double head (bicipital) the dorsal head or tuherculum articulating with the transverse process of the vertebra and the ventral head or capitulum articulating with Following the thoracolumbar group is a the side of the centrum. dentary bones. 495 The lower jaw is composed of a pair of each. Necturus usually has forty-six amphicoelous They articulate with each other by anterior and posterior zygapopJiy. free The margin of this is frequently referred to as suprascapula. The pelvic girdle is likewise largely cartilage. splenial bones. which bear teeth. Distal to this are the four elongated metatarsals and beyond each is the digit. six carpals in the wrist.. composed of phalanges. devoid of teeth and articulating with the quadrate of the vertebrae. four metacarpals in the palm. one are about eighteen thoracolumbar vertebrae each of which bears a pair of short Y-shaped ribs. which bear the last few teeth. The pectoral tral portion is girdle principally cartilage in structure. atlas. There is one cervical Posterior to this vertebra. The ven- formed by a posterior coracoid cartilage in the muscles Projecting dorsally of the body wall. Anatomy . Distal to the thigh is the shank with two bones. the tibia and the fihula lying parallel to each other. but it is fused in the midventral line. Chicago. The remainder is of the series. formed at the junction of scapula with the ventral parts into which the arm articulates is called the glenoid fossa. the radius and idna in the forearm (antebrachium).* •Illustrations of the skeleton of Necturus may be found in Stuart: of Nectwrus maculosus. and the angular bones. There are six somewhat fused tarsals in the ankle. single sacral vertebra to which the ilium of each side is attached by way of the sacral rib. posterior to this is the pair of iscMa which are partly ossified. with which the skull articulates.. The most dorsal. The skeleton of the anterior appendage includes the proximal humerus (in the brachium). consists of caudal vertebrae. posterior to this point. lates. and four digits recess each composed of joints or phalanges. Denoyer-Geppert Co. skull. and an anterior as well as the ends of the centra. CLASS AMPHIBIA parasphenoid. and laterally is the third unit of each side. Extending dorsally on each side is a slender ilium which joins the sacral rib and this in turn the sacrum. The anterior.

) — i— ~ ~ rectus abdominis myotomes '• pectineus I p _^ pubofemoralis '^ internus ischiofemoralis— rectus exteraus "" -gracilis • *- - - pubotibialis femorofibulariS:. 1. The Blakiston Company. 271. Comparative Yertebrate Dissection. and 3 are extensors of the foot. and 4 is a flexor. (From Atwood. 271. A horizontal septum sets of super- along the side of the body divides the muscles into a dorsal. Left lateral view of the muscles of the head and shoulder region of the salamander.) — . ventral view on the left. In the dorsal view tlie ilium has been cut from the pelvic girdle and deflected downward. 2. dorsal view on the right. The Blakiston Company.ischiocaudalis'' - gluteeus maxim us _/— . C The muscles of the hind legs of Necturus maculosus . (From Atwood. Nectiirus maculosis.496 TEXTBOOK OF ZOOLOGY Muscular System The muscles of the body are divided into segmental myotomes with intervening connective tissue sheets or myosepta. portion and a ventral hypaxial portion. epaxial The principal Fig. B. Comparative Vertebrate Dissection.semimembranosus pyrifonnis— flexor '' commimis communis extensor Fig.

•Illustrations of this system mav be found in Stuart: maculosusj Denoyer-Geppert Co. These furnish not only a good hiding place but a good hunting ground for the crayfish. In such a situation. they are essentially the same anatomically. the southern bullfrog. which is deit scribed in the next section of the book. pickerel weeds. This filled and may even include younger frogs. species. and other aquatic organisms which make up the bullfrog's diet. insect larvae. Bullfrogs are found in North America east of the Kockies from Canada to Mexico. particularly when alive. and the shore waters are with aquatic plants. They have also been introduced into the western portion of the United States and into various foreign countries. the shore is protected by low willows or other trees. is a solitary animal except during the breeding seaaquatic and does not leave the pools as does the It is strictly leopard frog. and the cypress ponds of swampy regions.CLASS AMPHIBIA ficial 497 muscles or those of the head and gills. snails. diet is quite varied External Structure Bullfrogs obtained in the South and Southwest are usually of two Rana catesheiana Shaw. such as lagoons. on a study of Rana catesheiana. the common bullfrog. it seems unnecessary to here. and this chapter is based usually are Rana catesheiana.. body wall.* THE BULLFROG Habitat The bullfrog son. Individuals of the former species attain larger sizes. and the giant bullfrogs of the southern swamps The two species differ not only in size but also in external appearajice. describe Since this system resembles that of the fish Avhich has been studied and is so closely similar to that of the frog. Anatomy of Necturus . Because of the development of the terrestrial limbs. and floating lily pads. or Bana grylio Stejneger. It prefers bodies of quiet water where there are both shallows and deeper water. water beetles. and the appendages. specific muscles the student will depend on the accompanying illustrations and the laboratory study. The Nervous System and Sense Organs already. small lakes. Chicago. the latter group is much more complicated in this amphibian than For the detailed information concerning the it was in the fishes. However.

while in the males it small fold of skin. The upper eyelid is immovable. runs from the eye around the posterior margin of the tympanum. Underparts are mottled with dark spots on a white background. The head has two prominent eyes which protrude above its surThese can be drawn back into their orbits and forced some- Fig. but as a result of the retraction of the eye into the orbit. The legs are marked with crossbars and other splotches of dark Preserved specimens appear brownish gray with the dark The body of the bullfrog includes the head and trunk. — External features of the common bullfrog. not by independent movement of the eyelid. 272.) (Courtesy what its into the mouth cavity.498 TEXTBOOK OF ZOOLOGY The common bullfrog is ordinarily greenish or olive brown. larger than the eye. color. the tympanic fold. Attached trunk on either side anteriorly are the forelegs and posteriorly mottling lighter in color than on the living specimen. The lower is lid of the frog's eye with attached nictitating membrane drawn up over the eye. the tympanum or eardrum. to the the hindlegs. In the females this is is about the size of the eye. face. and The mouth reaches from one side of . Back of each eye is a circular oval area. near the anterior part of the head. A The two each is nostrils or nares are guarded by a valve. and the upper surfaces may be plain or marked with large dark splotches. Rana catesbeiana. of Southern Biological Supply Company.

trunk. The forelimbs are used not help support the body but also as an aid in pushing food Bullfrogs into the mouth. moult or shed the superficial layer of epidermal at varying intervals. particularly during the breeding season. . and the pyloric or posterior end is narrowed and constricted where it joins the small intestine. At the point where the intestine turns back posteriorly the duodenum becomes the ileum. The latter narrows toward the esophagus. Dig^estive cells of their skin System and Digestion The mouth cavity. or tarsus. or anterior part of the small intestine. Following the tarsus is the foot with five digits (toes). The duodenum. The anus or vent is at the extreme posterior end of the trunk. five ordinarily leap about three feet but can easily cover a distance of The hindlegs are composed of the thigh. which are connected by a web. which is a short gullet leading directly from the pharynx to the stomach. the innermost thumb. wrist or carpus. The hindlimbs are long and have powerful muscles. is composed of two layers. the forearm.CLASS AMPHIBIA the head to the other 499 and has an upper and lower jaw. Its anterior or cardiac end is wide. which joins the trunk the shank. The lining of the esophagus has a number of longitudinal folds and is ciliated. The large intestine or rectum is sharply . an outer epidermis and an inner dermis. digit. The forelimbs are composed of the upper arm. which aid in keeping it moist. Bullfrogs or six feet. is enlarged. with the convex side toward the bullfrog's left. producing a very efficient swimming organ. which composes the remainder of the small intestine and is considerably coiled. and their positions in relation to various bones of the to specific only to hand give rise names for these tubercles. The smooth damp skin. The stomach normally lies on the left side of the body. The skin is pigmented and very rich in mucous glands. It is curved. and the ankle. apparently the same The digits may have tubercles on them. whereas the thumbs of females remain size. runs forward almost parallel with the stomach. or buccal cavity continues directly into the pharynx with no sharp line of bounjiary between them. or In the male. which is soft and loosely attached to the body except in the head region. which joins the and the hand with its four digits.

stomach. The tongue of the bullfrog is is somewhat smaller proit portionally than that of the grass frog. The stomach can be greatly expanded and aid in mixing it tions of these muscles pass the food through the digestive tract with the gastric juice in the stomach. and its posterior end is free. In addition. —Mouth or buccal cavity of the bullfrog. Esophagus. and they tached to the floor of the mouth just back of the tip of the lower jaw. _ lustach'ian tuoe Qlotth C Tonque Fig. Maxillary teeth . or other animal captured for food. in which case the stomach turns inside out and protrudes into the mouth cavity. called vomerine teeth. the cloaca. Its anterior half is atfish. as might be expected. making it bicornute. The tongue is somewhat leaflike in shape and is deeply notched behind. It passes directly into a muscular part.. from the small intestine and is wide and short. as maxillary teeth. for the latter inlajid. The peristaltic contrac- and They may also be used to regurgitate a disagreeable substance swallowed by the frog. which terminates in the anus The buccal cavity has in its roof near the end of the snout two patches of small conical teeth. 273.500 TEXTBOOK OF ZOOLOGY off marked or vent.^^. insect. In order to get the tongue out of the mouth the posterior part has to somersault over the attached anterior part. and intestine have an outer longitudinal and an inner circular layer of smooth muscle. more dependent on this organ when hunts insects Taste buds are present on the tongue and palate. the upper jaw has a single series of small conical teeth on its edge known These teeth serve primarily to help hold the craymay help at times in crushing it. acts as a reservoir for food which may be available only at irregular intervals and the frog has to take advantage of a food supply when . Vomerine teeth InLcmol nares Fbor of orbit M/' Sulojs marqinalis Isophaqus Vocal 5ac M.

the intestinal glands release into the blood stream a substance. the bile itself contains it secretes per- mits the fats to be more easily digested by a lipase from the pan- no digestive enzymes. It is also concerned in the formation of urea blood corpuscles.CLASS AMPHIBIA it is 501 The mucosa of the intestines has a number of longitudinal and transverse folds which produce a great absorptive surface through which the digested food can be taken up by the blood stream and transported to different parts of the body. Between the right and left lobes present. connected by narrow bridges of liver tissue. Here. which converts the proteins to peptones. Peristaltic contractions of the stomach cause a thorough mixing of the gastric juice with the food and then this partly digested food (chyme) is passed posteriorly into the small intestine. Their vomerine and maxillary teeth are too feeble to do more than slightly crush their prey. Bile is carried from the gall bladder to the duodenum by the hile duct. is Although its function in altering fatty substances tance is its ability to store important. for contains three taken from the pancreas by pancreatic ducts which empty into the bile duct that passes through the pancreas before entering the duodenum near its beginning. — Since frogs live primarily on insects. which passes through the pancreas on its way. so digestion begins in the stomach. The mesenteries which hold the body organs in position and the internal surface of the body wall likewise are made up of this peritoneal membrane. irregularly-lobed gland whose alkaline secreit of considerable importance in digestion. is the gall Madder. The liver is from the not primarily a digestive gland. which on reaching the pancreas causes it to pour forth into the duo- . and is other small invertebrate animals. This secretion is Intestines. The liver lies on each side of and behind the heart. Here the gastric glands secrete hydrochloric acid and an enzyme. secretin. It is a long. while the bile creas. It is threelobed. pepsin. and pancreas are covered with peritoneum. for. activated by the acid nature of the food. digestive enzymes. whitish. of prime impor- glycogen and the fat upon which a hiber- nating frog lives. and in the destruction of red lies in The pancreas tion is the loop between the stomach and duodenum. two lobes being on the left and one on the right. crayfish. Digestion. their food very rich in proteins. liver. which receives an alkaline secretion liver known as bile and stores it until needed in the process of digestion.

A thymus gland side. : In addition. system comprises the Mood vascular system and the The two systems are closely interrelated in that they both carry to the tissues of the body nutritive material neces- . which. steapsin. carbon dioxide. converting proteins to amino acids. this pancreatic juice contains three digestive enzymes which continues the digesbegun by pepsin in the stomach. The two thyroid glands are small and lie in front of the glottis under the floor of the mouth. Bile also contributes to the trypsin. which changes starches into sugars. spherical. leaving as by-products urea Food as feces. annjlopsin. causes a splitting of the fats into glycerol and fatty acids. Circulatory System The circulatory^ lymphatic system. tissue or for supplying energy.502 TEXTBOOK OF ZOOLOGY its demim tion highly alkaline secretion. reddish. The secretion and functions are discussed in the chapter on Internal Regulation. and a lipase. the functions of which are but The destroying of red blood corpuscles is an important duty. incompletely known. These foods in solution are taken by the blood stream and lymph vessels to various parts of the body where they are utilized for building and Sugars that are not used are stored as glycogen in the liver and in voluntary muscles. The process of digestion is completed in the intestine and the food products are taken up by absorption in its mucosa layer. Further discussion of will be taken up in the chapter on Internal Regulation. —Attached by a mesentery to the wall of the end of the rectum is intes- tine near the anterior the spleen. alkaline condition here. that is not digested passes to the large intestine where it is retained for a time and then passed to the outside through the anus Other Glands. as possibly also is the formation in its tissues of lymphocytes. an amylase. There is one on each side of the hyoid apparatus. lies on each It is it partly covered with muscle and under the skin behind the tympanic membrane is small. Ijnnphoid organ. In mammals the spleen is also believed to accumulate iron freed by the metabolism of other tissues. The liver also serves to store fats and to secrete urea and sugar directly into the blood stream. It is a small. one type of white blood corpuscle. aided by the bile. This iron is subsequently used in the formation of hemoglobin.

and thus become free in the surrounding tissue to engulf bacteria or other harmful objects. a tendon has few capillaries. in contrast. or foreign substances accidentally introduced. is a closed one. forces the blood to flow to the tissues. they break away from the heart are the arteries. The blood vessels leading When The in these reach the tissues. The capillaries are very small vessels. Blood is comprised of a clear liquid called the 'plasma. the capillaries. up into very small vessels. may contain dissolved nutritive substances from the digestive system. The abundance of the capillaries varies with the activity of the organ the greater the rate of metabolism the greater their abundance. . Other differences will be noted in the discussion. food products brought by the arterial blood pass into the tissues.CLASS AMPHIBIA 503 sary for metabolism aud remove from them to the excretory organs. arteries The vessels leading back to the heart are the and veins are connected by the capillaries. Leucocytes are able to get out of the capillaries. waste products from tissue repair aud destruction. suspended which are blood corpuscles of three kinds. the walls of which are made up of endothelium continued from the linings of arteries and veins. The pump which distribute it to the heart. acting as semipermeable membranes. by its conSince the system tractions. and the spindle cells or thrombocytes. hormones being transported from organs of one part of the body to another. They differ in several respects. and carbon dioxide and waste products are taken up to be conducted into the veins. the white blood corpuscles or leucocytes. the red blood corpuscles In addition. the lymph neither contains red blood corpuscles for transporting oxygen nor moves in a continuous closed vascular circuit as does the blood. veins. but in so doing they branch extensively and anastomose Through their thin to form fine networks in the tissues invaded. oxygen is unloaded from the red blood corpuscles. squeezing their way between the cells of the capillary walls. walls. which. The}^ connect the distal ends of the arteries with the proximal ends of the veins. the blood or erythrocytes. Examples of such are the various glands and the mucous membrane of the digestive tract. . waste products of body activity. The Blood Vascular System. system all — The blood moves through is a closed of tubelike vessels of various sizes parts of the body. the blood eventually returns to the heart.

-j1 \uricL:. . 1 / h Irtternal care ^^ ' I .504 TEXTBOOK OF ZOOLOGY The arteries are large vessels with elastic walls and carry blood from the heart to the capillary networks in the various organs and The arteries arise from the conus arteriosus tissues of the body.5»> .iari '-- \ _ -.

systemic arch. the anterior carotid arch. giving branches to the thyroid. -'h^^^^^emora I h Sciatic Fig. the middle.vesical i . and then extends along the edges of the lower jaw. (Drawn by Ruth M.) view. Systsmic arch Lateralis \^Left qastric PANCREAS ]. muscles of the hyoid — External c ral Auricularis i Occipital Internal carotid Cutaneous Carotid qiand Brachial Vertc bra I Conus arteriosusPulmonary. The Carotid Arch. or external carotid. It has at its base a spongy enlargement known as the carotid gland which by its structure serves to steady the pressure of blood passing into the artery. The internal branch is larger and is called the internal carotid.rnesenteric l_Coeliac Anterior mesenteric Splenic (Irinoaenital Epigastric RECTUM! pogfer/or mescnter/c Viae Recto. and the posterior. This artery follows the side of the neck to the base . The more ventral. pulmo cutaneous arch. passes forward. and tongue. 275. Each carotid arch divides into two branches. —Arteries of the bullfrog from ventral Sanders. Ungual artery. pseudothyroid.CLASS AMPHIBIA 505 each side of the body.Riijht Cjastric Dorsalis Cozliaco.

and the turning posteriorly along the spinal column. The small posterior mesenteric artery is given off near the posterior end of the aorta. passing to a portion of the rectum and. the dorsal side of the skull to the orbit vertebral artery. adjacent body wall and enters the arm The systemic arches from each side. the dorsal aorta. and few small lumbar arteries arise either as branches of these or directly from the aorta and go to the body wall on each side. At or just posterior to this meeting point. It often anastomoses on the rectum with fat bodies. The systemic arch soon after it leaves the truncus supplies a small laryngeal artery to the larynx and muscles of the hyoid. the cerebral carotid which enters the skull and supplies the brain. ImmeIt diately posterior to the occipito vertebral artery the large subclavian artery arises from the systemic arch. in the female. A descending branches of the anterior mesenteric. while the former sends small pancreatic arteries to the pancreas liver. gall bladder. reproductive organs. where it is The anterior mesenteric artery gives off the splenic (lienal) artery to the spleen and then divides into two parallel vessels which send numerous smaller arteries to the small and large distributed. latter runs directly to the dorsal or left side of the off stomach. gives off an occipitovertehral artery which sends a small artery to the dorsal side of the esophagus. . meet near the anterior end of the kidneys and fuse into a single large artery. The urinogenital aorta to right and arteries consist of about four to six small off much- divided arteries which are given left. — then curves It downward and around the esopha- gus on each side. arteries. and and continues to the ventral side of the stomach. then branches at the spinal cord into the occipital artery. the coeliac artery. after curving under the alimentary canal. giving off the palatine artery to the roof of the mouth. to the ovisac. and the ophthalmic artery to the eye. and a posterior branch. branches to the shoulder and as the brachial artery. intestines. The coeliac artery divides into right and left gastric The . from the ventral side of the dorsal supplying the kidneys. which extends posteriorly. there arises from the aorta the large coeliacomesenteric artery which divides into an anterior branch.506 TEXTBOOK OF ZOOLOGY of the skull. It The Systemic Arch. a larger hepatic artery to the pancreas. the anterior mesenteric artery. running anteriorly on and tympanum.

now called the a branch to the right and to the left.OBC . The pulmocutaneous .v^. lumbar- Renal Renal portal. gives off In the upper leg the continuation of the iliac.- CLASS AMPHIBIA 507 body cavity the dorsal aorta divides Each of these gives into two iliac arteries going to the hind legs. sending off several branches at the knee. Vesical External iliac Fe moral Fig. 276. skin. LIVER LOWER L^FT I. bladder. and (2) just below it. As the iliac artery enters the leg.V CutanecusPosterior ^^^\ Hepatic portal Gastric Splenic /.^^0. —Veins of bullfrog from ventral view. a rectovesicular artery is sent Mandibular !!^lnternal jut^ular External juijular . Subscapular Innominate -Pre cava I Brachial. (Drawn by Ruth M. and buccopharyngeal cavity. Sinus venos CardiacHepatic. a femoral artery passing to the body wall. off (1) an epigastric artery suppljang the bladder and dorsal and ventral body walls of the region.Mesenteric SPLEEN vena cava Soermatic Dorso. The pulmo cutaneous arch takes blood to the respiratory organs: the lungs. sciatic.) off to the rectum.>-. supplying the muscles.^\ . and skin on the dorsal surface of the thigh. and proximal muscles of the Near the posterior end of the thigh. and then continues down the leg. Sanders. skin.

thyroid. supplying the skin is and the which distributed to the skin of the The Veins. and the subscapular vein bringing blood from the back of the arm and shoulder. The two precavals empty into the anterior end of the sinus venosus of the heart. circulation. two anterior precavals receives blood from three branches: (1) the external jugular bringing blood from the tongue. which passes outward to the skin. Important branches of the cutaneous are: the auricularis. and of the (3) the subclavian vein. the dorsalis. may be divided into four main the systemic. and pulmonary The systemic veins carry the greatest load of blood to the heart. The larger collecting veins of the system consist of two precavals receiving blood from the anterior parts of the body. tympanum and of the back side. and the large musculocutaneous vein. (2) the innominate vein. . pseudothyroid. adjacent head region lateralis. Pulmonary veins from the lungs carry oxygenated blood. and floor of the mouth. Many flow have semilunar valves on the internal surface of the wall which open in the direction of veins. made up of a fusion of the internal jugular returning blood from the brain and other parts of the head.508 TEXTBOOK OP ZOOLOGY arch on each side divides into a pulmonary artery to the lungs and a large cutaneous artery. renal portal. which differs from the type of blood found or hepatic in the other veins. The venous systems: systems. providing renal filters to eliminate urea and other waste products from the blood or to alter it chemically. hyoid. a fusion of the brachial vein. The walls of the veins are thinner — and not as elastic as those of the arteries. . which forms an ellipse down the side of the body and extends up into the head region. except the lungs. returning blood Each from the forelimb. particularly those of the limbs. hepatic portal. therefore. and prevent the backflow of blood. supplying the . and a single postcaval or posterior vena cava receiving blood from the posterior parts of the body. and the posterior vena cava empties into its posterior end. returning blood from the skin and outer muscles in these regions. These vessels usually parallel the arteries that brought blood to the tissues from which the veins are returning it. the venous system carries some of the blood through the kidneys or through the liver. In returning blood to the heart.

these unions is the hepatic portal vein. from the The gonads by small spermatic or ovarian veins. comprised of two chief veins. a cardiac vein from the heart.CLASS AMPHIBIA 509 large posterior vena cava originates between the kidneys and receives blood from each kidney by five or six renal veins. discharging its blood into sinusoids. after giving off the pelvic vein. — This from the hindlegs. parietal veins from the body wall and. The abdominal vein arises as follows: Two large veins. Near the heart the vena cava receives two large hepatic veins from each side of the liver. diverts blood to a purifying organ instead of carrying it directly to the heart. It passes through the anterior portion of the pancreas and sends a large branch into the lower left lobe of the liver. and from the fat bodies by other small branches. bring blood from the hindlimbs. runs anteriorly and joins the sciatic. system. pancreatic veins from the pancreas. It is returned to the systemic system The Hepatic Portal System. The femoral. to make the renal portal vein. In this ease. gives off the pelvic vein. and enters the right and upper left lobes of the liver by short branches. as it enters the body cavity. the hepatic portal vein and the ventral abdominal vein. and gastric The vessel resulting from veins from both sides of the stomach. the sciatic and femoral. The pelvic veins from each side of the body join in the middle to form the large ventral abdominal vein. at its anteIn the region of the liver rior end. — This system is veins that join the postcaval. It then continues a short distance to join the abdominal vein just below the heart. is joined by the hepatic portal vein. teric vein Veins from the large and small intestines unite to form the mesenwhich is joined as it progresses forward by the splenic vein from the spleen. like the hepatic portal system. As the abdominal vein runs toward the heart along the median portion of the ventral body wall. it receives vesicular veins from the bladder. bring (modified it to the liver to pass through a netAvork of sinusoids through hepatic capillaries). instead of carrying blood directly to the heart. These veins. The outer femoral vein and the medial sciatic vein collect blood The Renal Portal System. it leaves the body wall. the blood is taken to the kidneys. Near the kidney this vein receives the dorsolumbar vein from the body . At about this point it often receives a final gastric branch which has passed on top of the pancreas to join it. The femoral vein.

is On the ventral side a muscular tube. It is the left auricle on its dorsal side. backflow. where they break up into capillaries. which is lined by a transparent tissue. the auriculoventricular aperture. Blood leaves the ventricle and enters the arterial system through The opening into the conus is protected by three pocketlike semilunar valves which open inwardly into the conus when blood is passing out but are tightly closed at other times. several vessels from the ovisacs (uteri). The right and left pulmonary veins unite to form a single vessel which empties into the Other veins which take on oxygen are and buccopharyngeal cavity. which receives venous blood from the systemic The sinus venosus empties auricular aperture. Pulmonary Veins. which is divided by the inter- This aperture has two on each side and two small valves at each end which large valves regulate the discharge of blood into the ventricle and prevent its auricular septum separating the two auricles.510 TEXTBOOK OP ZOOLOGY wall and. The the conus arteriosus. . the C07ms arteriosus. It is conical in shape and in the frog consists of a right and left thin-walled auricle above a single thick-walled ventricle. Both auricles pass blood into the ventricle through a common open- ing. is heart a thin-walled sac. Valves are not necessary at this opening. The heart is enclosed in the pericardial cavity. The Heart. purified of some of its waste products Blood which passes through these capillaries is and then leaves the kidney through the renal veins which empty into and originate the posterior vena cava of the systemic system. triangular in shape. On the dorsal side of the It conducts blood away from the heart. veins. returning the oxygenated blood to the heart. described with the arteries. The renal portal vein follows the dorsolateral margin of the kidney. those coming from the skin — rhythmically contracting organ that circulates the blood. into the right auricle through the sinu- This aperture has liplike valves on each side to prevent the blood from flowing back into the sinus when the auricle The smaller left auricle receives oxygenated blood from contracts. and is separated from the remainder of the body by the transverse septum. the pulmonary vein. the sinus venosus. send- ing numerous transverse branches into the organ. for pressure on the auricular w^alls tends to close the small oblique aperture when the auricle contracts. the pericardium.— These veins run along the inner walls of each lung. in the female.

and the synangkim. dorsal wall of the pylangium and the other edge lying free in the Upon contraction of the conus this structure is brought into contact with the ventral wall and helps direct the flow of blood into the arches. Running through the length of the pylangium is a longitudinal spiral valve.CLASS AMPHIBIA 511 proximal portion of the couus is known as the pylangium. one edge attached to the distal portion as the vessel. one to the right and the other to the left. Just below these valves is an aperture which leads into the trunk formed by the union of the two pulmocutaneous arteries. shown through with the end of the spiral valve. separate the pylangium from the synangium.-/luricufo. 277. there Carotid A is of the spiral valve where it is the a pair of small synangial valves which. Near the anterior free end widest. The synangial chamber is very short and gives off almost immetwo large branches.. All three trunks . together System. R auricle - -Sinu-auricular aperture Spiral valve —Loft auricle Conus orte-_ nosus — Intzrauricular septum --Dmtle inpulmoSemilunar vaUz cutaneous A. They are formed by two longitudinal diately septa dividing the vessel into three compartments. .-Heart of frog with the ventral wall removed and bristles the arteries of the truncus arteriosus.-">:7 Ventricle Fig. In each of these branches originate the three main trunks or arches of the arterial system.ventricular valve .A Pulmocuta- neous Truncus arteriosus '^-Pulmonary aperburz f\.

pylangium pulmonary arch. forcing out the remaining oxygenated blood. As the systemic arteries fill. in a wavelike peristaltic manner. the ventricle immediately forces the blood into the conus through the semilunar valve. The heart beats contracts first. together with that of the synangial valves which are anterior to the common opening of the pulmonary arches. but enters the posterior compartment. and finally the conus. then the ventricle. sufficiently fast The heart must beat and pump a sufficient of blood at each stroke to insure an adequate supply of volume oxygen and food to the body tissues. the systemic arch from the middle compartment. they offer more resistance to the blood. and the pulmocutaneous arch from the posterior com- Blood enters the anterior and middle compartments from the synangium. As the contraction of the ventricle comes to an end. The rate of pulsation is influenced greatly by temperature up to a certain maximum rate. Venous blood from the right auricle is closest to the conus. while resistance in the carotid arteries de- creases due to their emptying into capillaries so the last oxygenated blood from the ventricle passes into the carotids and is conveyed . Since the heart's contractions are wavelike. Since the carotid arteries offer some resistance systemic arteries to blood flow. The blood therefore passes into the synangium and enters the chambers leading to the systemic arteries or the carotid arteries. partment.512 TEXTBOOK OF ZOOLOGY are therefore enclosed in one large vessel for a short distance before breaking up into three separate vessels. as well as to remove waste products as they form. flowing into the closest openThis is the opening in the ing offering the least resistance. the pylangial part of the conus contracts. and reduce mixing. for the activity and metabolism of the bullfrog are considerably affected by temperature. completely shuts off the flow of blood into these arches. T