CHAPTER XVIII.
PEDIGREE AND HISTORY OF THE ANIMAL KINGDOM.
I. ANIMAL-PLANTS AND WORMS.
The Natural System of the Animal Kingdom.—Linnæus and Lamarck’s Systems.—The Four Types of Bär and Cuvier.—Their Increase to Seven Types.—Genealogical Importance of the Seven Types as Independent Tribes of the Animal Kingdom.—Derivation of Zoophytes and Worms from Primæval Animals.—Monophyletic and Polyphyletic Hypothesis of the Descent of the Animal Kingdom.—Common Origin of the Four Higher Animal Tribes out of the Worm Tribe.—Division of the Seven Animal Tribes into Sixteen Main Classes, and Thirty-eight Classes.—Primæval Animals (Monera, Amœbæ, Synamœbæ), Gregarines, Infusoria, Planæades, and Gastræades (Planula and Gastrula).—Tribe of Zoophytes.—Spongiæ (Mucous Sponges, Fibrous Sponges, Calcareous Sponges).—Sea Nettles, or Acalephæ (Corals, Hood-jellies, Comb-jellies).—Tribe of Worms.
The natural system of organisms which we must employ in the animal as well as in the vegetable kingdom, as a guide in our genealogical investigations, is in both cases of but recent origin, and essentially determined by the progress of comparative anatomy and ontogeny (the history of individual development) during the present century. Almost all the attempts at classification made in the last century followed the path of the artificial system, which was first established in a consistent manner by Charles Linnæus. The artificial system differs essentially from the natural one, in the fact that it does not make the whole organization and the internal structure (depending upon the blood relationship) the basis of classification, but only employs individual, and for the most part external, characteristics, which readily strike the eye. Thus Linnæus distinguished his twenty-four classes of the vegetable kingdom principally by the number, formation, and combination of the stamens. In like manner he distinguished six classes in the animal kingdom principally by the nature of the heart and blood. These six classes were: (1) Mammals; (2) Birds; (3) Amphibious Animals; (4) Fishes; (5) Insects; and (6) Worms.
But these six animal classes of Linnæus are by no means of equal value, and it was an important advance when, at the end of the last century, Lamarck comprised the first four classes as vertebrate animals (Vertebrata), and put them in contrast with the remaining animals (the insects and worms of Linnæus), of which he made a second main division—the invertebrate animals (Invertebrata). In reality Lamarck thus agreed with Aristotle, the father of Natural History, who had distinguished these two main groups, and called the former _blood-bearing animals_, the latter _bloodless animals_.
The next important progress towards a natural system of the animal kingdom was made some decades later by two most illustrious zoologists, Carl Ernst Bär and George Cuvier. As has already been remarked, they established, almost simultaneously and independently of one another, the proposition that it was necessary to distinguish several completely distinct main groups in the animal kingdom, each of which possessed an entirely peculiar type or structure (compare above, vol. i. p. 53). In each of these main divisions there is a tree-shaped and branching gradation from most simple and imperfect forms to those which are exceedingly composite and highly developed. The _degree of development_ within each type is quite independent of the peculiar _plan of structure_, which forms the basis of the type and gives it a special characteristic. The “type” is determined by the peculiar relations in position of the most important parts of the body, and the manner in which the organs are connected. The degree of development, however, is dependent upon the greater or less division of labour among organs, and on the differentiation of the plastids and organs. This extremely important and fruitful idea was established by Bär, who relied more distinctly and thoroughly upon the history of individual development than did Cuvier. Cuvier based his argument upon the results of comparative anatomy. But neither of them recognized the true cause of the remarkable relationships pointed out by them, which is first revealed to us by the Theory of Descent. It shows us that the common _type_ or plan of structure is determined by _inheritance_, and the degree of development or differentiation by _adaptation_. (Gen. Morph. ii. 10).
Both Bär and Cuvier distinguished four different types in the animal kingdom, and divided it accordingly into four great main divisions (branches or circles). The first of these is formed by the vertebrate animals (Vertebrata), and comprises Linnæus’ first four classes—mammals, birds, amphibious animals, and fishes. The second type is formed by the articulated animals (Articulata), containing Linnæus’ insects, consequently the six-legged insects, and also the myriopods, spiders, and crustacea, but besides these, a large number of the worms, especially the ringed worms. The third main division comprises the molluscous animals (Mollusca)—slugs, snails, mussels, and some kindred groups. Finally, the fourth and last circle of the animal kingdom comprises the various radiated animals (Radiata), which at first sight differ from the three preceding types by their radiated, flower-like form of body. For while the bodies of molluscs, articulated animals, and vertebrated animals consist of two symmetrical lateral halves—of two counterparts or antimera, of which the one is the mirror of the other—the bodies of the so-called radiated animals are composed of more than two, generally of four, five, or six counterparts grouped round a common central axis, as in the case of a flower. However striking this difference may seem at first, it is, in reality, a very subordinate one, and the radial form has by no means the same importance in all “radiated animals.”
The establishment of these natural main groups or types of the animal kingdom by Bär and Cuvier was the greatest advance in the classification of animals since the time of Linnæus. The three groups of vertebrated animals, articulated animals, and molluscs are so much in accordance with nature that they are retained, even at the present day, little altered in extent. But a more accurate knowledge soon showed the utterly unnatural character of the group of the radiated animals. Leuckart, in 1848, first pointed out that two perfectly distinct types were confounded under the name, namely, the _Star-fishes_ (Echinoderma)—the sea-stars, lily encrinites, sea-urchins, and sea-cucumbers; and, on the other hand, the _Animal-plants_, or _Zoophytes_ (Cœlenterata or Zoophyta)—the sponges, corals, hood-jellies, and comb-jellies. At the same time, Siebold united the Infusoria with the Rhizopoda, under the name of Protozoa (lowest animals), into a special main division of the animal kingdom. By this the number of animal types was increased to six. It was finally increased to seven by the fact that modern zoologists separated the main division of the articulated animals into two groups: (_a_) those possessing _articulated feet_ (Arthropoda), corresponding to Linnæus’ Insects, namely, the Flies (with six legs), Myriopods, Spiders, and Crustacea; and (_b_) the footless _Worms_ (Vermes), or those possessing non-articulated feet. These latter comprise only the real or genuine Worms (ring-worms, round worms, planarian worms, etc.), and therefore in no way correspond with the Worms of Linnæus, who had included the molluscs, the radiates, and many other lower animals under this name.
Thus, according to the views of modern zoologists, which are given in all recent manuals and treatises on zoology, the animal kingdom is composed of seven completely distinct main divisions or types, each of which is distinguished by a characteristic plan of structure peculiar to it, and perfectly distinct from every one of the others. In the natural system of the animal kingdom—which I shall now proceed to explain as its probable pedigree—I shall on the whole agree with this usual division, but not without some modifications, which I consider very important in connection with genealogy, and which are rendered absolutely necessary in consequence of our view as to the history of the development of animals.
We evidently obtain the greatest amount of information concerning the _pedigree of the animal kingdom_ (as well as concerning that of the vegetable kingdom) from comparative anatomy and ontogeny. Besides these, palæontology also throws much valuable light upon the historical succession of many of the groups. From numerous facts in comparative anatomy, we may, in the first place, infer the _common origin of all those animals which belong to one of the seven “types.”_ For in spite of all the variety in the external form developed within each of these types, the essential relative position of the parts of the body which determines the type, is so constant, and agrees so completely in all the members of every type, that on account of their relations of form alone we are obliged to unite them, in the natural system, into a single main group. But we must certainly conclude, moreover, that this conjunction also has its expression in the pedigree of the animal kingdom. For the true cause of the intimate agreement in structure can only be the actual blood relationship. Hence we may, without further discussion, lay down the important proposition that all animals belonging to one and the same circle or type must be descended from one and the same original primary form. In other words, the idea of the circle or type, as it is employed in zoology since Bär and Cuvier’s time to designate the few principal main groups or “sub-kingdoms” of the animal kingdoms, coincides with the idea of “tribe” or “phylum,” as employed by the Theory of Descent.
If, then, we can trace all the varieties of animal forms to these seven fundamental forms, the following question next presents itself to us as a second phylogenetic problem—Where do these seven animal tribes come from? Are they seven original primary forms of an entirely independent origin, or are they also distantly related by blood to one another?
At first we might be inclined to answer this question in a _polyphyletic_ sense, by saying that we must assume, for each of the seven great animal tribes, at least one independent primary form completely distinct from the others. On further considering this difficult problem, we arrive in the end at the notion of a _monophyletic_ origin of the animal kingdom, viz., that these seven primary forms are connected at their lowest roots, and that they are derived from a single, common primæval form. _In the animal as well as in the vegetable kingdom, when closely and accurately considered, the monophyletic hypothesis of descent is found to be more satisfactory than the polyphyletic hypothesis._
It is _comparative ontogeny_ (embryology) which first and foremost leads to the assumption of the monophyletic origin of the whole animal kingdom (the Protista excepted of course). The zoologist who has thoughtfully compared the history of the individual development of various animals, and has understood the importance of the biogenetic principle (p. 33), cannot but be convinced that a common root must be assumed for the seven different animal tribes, and that all animals, including man, are derived from a single, common primary form. The result of the consideration of the facts of embryology, or ontogeny, is the following genealogical or phylogenetic hypothesis, which I have put forward and explained in detail in my “Philosophy of Calcareous Sponges” (Monograph of the Calcareous Sponges, vol. i. pp. 464, 465, etc.,—“the Theory of the Layers of the Embryo, and the Pedigree of Animals”).
The first stage of organic life in the Animal kingdom (as in the Vegetable and Protista kingdoms) was formed by perfectly simple _Monera_, originating by spontaneous generation. The former existence of this simplest animal form is, even at present, attested by the fact that the egg-cell of many animals loses its kernel directly after becoming fructified, and thus relapses to the lower stage of development of a cytod without a kernel, like a Moneron. This remarkable occurrence I have interpreted, according to the law of latent inheritance (vol. i. p. 205), as a phylogenetic _relapse_ of the cellular form into the original form of a cytod. The _Monerula_, as we may call this egg-cytod without a kernel, repeats then, according to the biogenetic principle (vol. ii. p. 33), the most ancient of all animal forms, the common primary form of the animal kingdom, namely, the Moneron.
The second ontogenetic process consists in a new kernel being formed in the Monerula, or egg-cytod, which thus returns again to the value of a true _egg-cell_. According to this, we must look upon the simple animal cell, containing a kernel, or the single-celled primæval animal—which may still be seen in a living state in the _Amœbæ_ of the present day—as the _second_ step in the series of phylogenetic forms of the animal kingdom. Like the still living simple Amœbæ, and like the naked egg-cells of many lower animals (for example, of Sponges and Medusæ, etc.), which cannot be distinguished from them, the remote phyletic primary Amœbæ also were perfectly simple naked-cells, which moved about in the Laurentian primæval ocean, creeping by means of the ever-changing processes of their body-substance, and nourishing and propagating themselves in the same way as the Amœbæ of the present day. (Compare vol. i. p. 188, and vol. ii. p. 54.) The existence of this Amœba-like, _single-celled primary form_ of the whole animal kingdom is unmistakably indicated by the exceedingly important fact that the egg of all animals, from those of sponges and worms up to those of the ant and man, is a simple cell.
Thirdly, from the “single-cell” state arose the _simplest multicellular state_, namely, a heap or a small community of simple, equi-formal, and equivalent cells. Even at the present day, in the ontogenetic development of every animal egg-cell, there first arises a globular heap of equi-formal naked cells, by the repeated self-division of the primary cell. (Compare vol. i. p. 190 and the Frontispiece, Fig. 3.) We called this accumulation of cells the _mulberry state_ (Morula), because it resembles a mulberry or blackberry. This Morula-body occurs in the same simple form in all the different tribes of animals, and on account of this most important circumstance we may infer—according to the biogenetic principle—that the _most ancient, many-celled, primary form of the animal kingdom_ resembled a Morula like this, and was in fact a simple heap of Amœba-like primæval cells, one similar to the other. We shall call this most ancient community of Amœbæ—this most simple accumulation of animal cells—which is recapitulated in individual development by the Morula—the _Synamœba_.
Out of the Synamœbæ, in the early Laurentian period, there afterwards developed a fourth primary form of the animal kingdom, which we shall call the ciliated germ (Planæa). This arose out of the Synamœba by the outer cells on the surface of the cellular community beginning to extend vibrating fringes called cilia, and becoming “ciliated cells,” and thus differentiating from the inner and unchanged cells. The Synamœbæ consisted of completely equi-formed and naked cells, and crept about slowly, at the bottom of the Laurentian primæval ocean, by means of movements like those of an Amœba. The Planæa, on the other hand, consisted of two kinds of different cells—inner ones like the Amœbæ, and external “ciliated cells.” By the vibrating movements of the cilia the entire multicellular body acquired a more rapid and stronger motion, and passed over from the creeping to the swimming mode of locomotion. In exactly the same manner the _Morula_, in the ontogenesis of lower animals, still changes into a ciliated form of larva, which has been known, since the year 1847, under the name of _Planula_. This Planula is sometimes a globular, sometimes an oval body, which swims about in the water by means of a vibrating movement; the fringed (ciliated) and smaller cells of the surface differ from the larger inner cells, which are unfringed. (Fig. 4 of the Frontispiece.)
Out of this Planula, or fringed larva, there then develops, in animals of all tribes, an exceedingly important and interesting animal form, which, in my Monograph of the Calcareous Sponges, I have named _Gastrula_ (that is, larva with a stomach or intestine). (Frontispiece, Fig. 5, 6). This Gastrula externally resembles the Planula, but differs essentially from it in the fact that it encloses a cavity which opens to the outside by a mouth. The cavity is the “_primary intestine_,” or “primary stomach,” the _progaster_, the first beginning of the alimentary canal; its opening is the “_primary mouth_” (prostoma). The wall of the progaster consists of two layers of cells: an outer layer of smaller ciliated cells (outer skin, or ectoderm), and of an inner layer of larger non-ciliated cells (inner skin, or entoderm). This exceedingly important larval form, the “Gastrula,” makes its appearance in the ontogenesis of all tribes of animals—in Sponges, Medusæ, Corals, Worms, Sea-squirts, Radiated animals, Molluscs, and even in the lowest Vertebrata (Amphioxus: compare p. 200, Plate XII., Fig. _B_ 4; see also in the same place the Ascidian, Fig. _A_ 4).
Definition of the _forms_ | +Ontogenesis.+ | +Phylogenesis.+ of the five first stages | The five first stages | The five first stages of the development of | of the individual | of the phyletic or the animal body. | development. | historical development. -------------------------------+------------------------------+------------------------ | | _First Stage of Development._ | 1. | 1. | +Monerula.+ | +Moneron.+ A simple cytod (a | | plastid without a kernel.) | Animal egg without a | Most ancient animal | | kernel (when the egg-kernel | Monera, originating by | | has disappeared, | spontaneous generation. | | after being fructified). | | | | | | | | | | | | _Second Stage of Development._ | 2. | 2. | +Ovulum.+ | +Amœbæ.+ A simple cell (a | | plastid containing a | Animal egg with kernel | Animal Amœbæ. kernel.) | (a simple egg-cell). | | | | | | | | | | | | | | | | | _Third Stage of Development._ | 3. | 3. | +Morula.+ | +Synamœba.+ A community (an | (_Mulberry form._) | aggregation of identical | | An aggregation of simple cells). | Globular heap of homogeneous | Amœbæ. | | “cleavage spheres.” | | | | | | | | | | | | _Fourth Stage of Development._ | 4. | 4. | +Planula.+ | +Planæa.+ A solid or bladder-shaped, | (_Ciliated larva_.) | globular, or oval | | Many-celled primæval body, _composed of two | Many-celled larva | animal without kinds of different cells_: | without mouth, composed | mouth, composed of externally ciliated, | of different cells. | two kinds of different internally non-ciliated | | | cells. cells. | | | | | | | | | | | | | | _Fifth Stage of Development._ | 5. | 5. | +Gastrula.+ | +Gastræa.+ A globular or oval | (_Larva with mouth._) | _body with simple intestinal | Many-celled with intestines | Many-celled primæval cavity and mouth-opening. | and mouth; intestinal | animal with intestine Body wall composed | wall with two | and mouth; intestinal of two layers_; an | layers. | wall with two externally ciliated ectoderm | | layers. (Primary form (dermal layer), an | | of zoophytes and internally non-ciliated | | worms.) entoderm (gastral layer). | |
From the ontogenetic occurrence of the Gastrula in the most different animal classes, from Zoophytes up to Vertebrata, we may, according to the biogenetic principle, safely draw the conclusion that during the Laurentian period there existed a common primary form of the six higher animal tribes, which in all essential points was formed like the Gastrula, and which we shall call the Gastræa. This Gastræa possessed a perfectly simple globular or oval body, which enclosed a simple cavity of like form, namely, the progaster; at one of the poles of the longitudinal axis the primary intestine opened by a mouth which served for the reception of nutrition. The body wall (which was also the intestinal wall) consisted of two layers of cells, the unfringed entoderm, or intestinal layer, and the fringed ectoderm, or skin-layer; by the motion of the cilia or fringes of the latter the Gastræa swam about freely in the Laurentian ocean. Even in those higher animals, in the ontogenesis of which the original Gastrula form has disappeared, according to the laws of abbreviated inheritance (vol. i. p. 212), the composition of the Gastræa body has been transmitted to the phase of development which directly arises out of the Morula. This phase is an oval or round disc consisting of two cell-layers or membranes: the outer cell-layer, the _animal or dermal layer_ (ectoblast), corresponds to the ectoderm of the Gastræa; out of it develops the external, loose skin (epidermis), with its glands and appendages, as well as the central nervous system. The inner cell-layer, the _vegetative or intestinal layer_ (hypoblast), is originally the entoderm of the Gastræa; out of it develops the inner membrane (epithelium) of the intestinal canal and its glands. (Compare my Monograph of the Calcareous Sponges, vol. i. p. 466, etc.)
By ontogeny we have already gained five primordial stages of development of the animal kingdom: (1) the Moneron; (2) the Amœba; (3) the Synamœba; (4) the Planæa; and (5) the Gastræa. The former existence of these five oldest primary forms, which succeeded one another, and which must have lived in the Laurentian period, follows as a consequence of the biogenetic principle; that is to say, from the parallelism and the mechanico-causal connection of ontogenesis and phylogenesis. (Compare vol. i. p. 309.) In our genealogical system of the animal kingdom we may class all these animal forms, long since extinct, and, which on account of the soft nature of their bodies could leave no fossil remains, among the tribe of Primæval animals (Protozoa), which also comprises the still living Infusoria and Gregarinæ.
The phyletic development of the six higher animal tribes, which are all derived from the Gastræa, deviated at this point in two directions. In other words, the _Gastræads_ (as we may call the group of forms characterized by the Gastræa-type of structure), divided into two divergent lines or branches; the one branch of Gastræads gave up free locomotion, adhered to the bottom of the sea, and thus, by adopting an adhesive mode of life, gave rise to the _Protascus_, the common primary form of the _Animal-plants_ (Zoophyta). The other branch of the Gastræads retained free locomotion, did not become adherent and later on developed into the _Prothelmis_, the common primary form of _Worms_ (Vermes). (Compare p. 133.)
This latter tribe (as limited by modern zoology) is of the greatest interest in the study of genealogy. For among Worms, as we shall see later, there are, besides very numerous peculiar families, and besides many independent classes, also very remarkable forms, which may be considered as _forms of direct transition_ to the four higher animal tribes. Both comparative anatomy and the ontogeny of these worms enable us to recognize in them the nearest blood relations of those extinct animal forms which were the original primary forms of the four higher animal tribes. Hence these latter, the Molluscs, Star-fishes, Articulated animals, and Vertebrate animals, do not stand in any close blood relationship to one another, but have originated independently in four different places out of the tribe of Worms.
In this way comparative anatomy and phylogeny lead us to the _monophyletic pedigree of the animal kingdom_, the outlines of which are given on p. 133. According to it the seven phyla, or tribes, of the animal kingdom are of different value in regard to genealogy. The original primary group of the whole animal kingdom is formed by the Primæval animals (Protozoa), including the Infusoria and Gastræads. Out of these latter arose the two tribes of Animal-plants (Zoophyta) and Worms as diverging branches. Out of four different groups of the Worm tribe, the four higher tribes of the animal kingdom were developed—the Star-fishes (Echinoderma) and Insects (Arthropoda) on the one hand, and the Molluscs (Mollusca) and Vertebrated animals (Vertebrata) on the other.
Having thus sketched out the monophyletic pedigree of the animal kingdom in its most important features, we must now turn to a closer examination of the historical course of development which the seven tribes of the animal kingdom, and the classes distinguished in them, have passed through (p. 132). There is a much larger number of classes in the animal than in the vegetable kingdom, owing to the simple reason that the animal body, in consequence of its more varied and perfect vital activity, could differentiate and develop in very many more different directions than could the vegetable body. Thus, while we were able to divide the whole vegetable kingdom into six main classes and nineteen classes, we have to distinguish, at least, sixteen main classes and thirty-eight classes in the animal kingdom. These are distributed among the seven different tribes of the animal kingdom in the way shown in the Systematic Survey on pages 132 and 133.
The group of _Primæval animals_ (Protozoa) within the compass which we here assign to this tribe, comprises the most ancient and the simplest primary forms of the animal kingdom; for example, the five oldest phyletic stages of development previously mentioned, and besides these the Infusoria and Gregarinæ, as well as all those imperfect animal forms, for which, on account of their simple and indifferent organization, no place can be found in any of the other six animal tribes. Most zoologists, in addition to these, include among the Protozoa a larger or smaller portion of those lowest organisms, which we mentioned in our neutral kingdom of Protista (in Chapter XVI.). But these Protista, especially the large division of the Rhizopoda, which are so rich in forms, cannot be considered as real animals for reasons previously given. Hence, if we here leave them out of the question, we may accept two main classes or provinces of real Protozoa, namely, _Egg animals_ (Ovularia) and _Germ animals_ (Blastularia). To the former belong the three classes of Archezoa, Gregarinæ, and Infusoria, to the latter the two classes of Planæads and Gastræads.
SYSTEMATIC SURVEY
_Of the 16 Main Classes and 38 Classes of the Animal Kingdom._
------------------+-----------------------+----------------------+----------------- _Tribes or Phyla_ | _Main Classes_, | _Classes_ |_Systematic Name_ _of the_ |_Branches or Clades_ | _of the_ | _of the_ _Animal Kingdom._ | _of the_ | _Animal Kingdom._ | _Classes._ | _Animal Kingdom._ | | ------------------+-----------------------+----------------------+------------------
A. { =Primæval= { I. Egg-animals { 1. Archaic animals 1. Archezoa =Animals= { _Ovularia_ { 2. Gregarines 2. Gregarinæ { { 3. Infusoria 3. Infusoria +Protozoa+ { {II. Mulberry animals { 4. Planæads 4. Planæadas { _Blastularia_ { 5. Gastræads 5. Gastræadas
B. { =Animal= { III. Sponges { 6. Sponges 6. Porifera =Plants= { _Spongiæ_ { { +Zoophyta+ { IV. Sea-nettles { 7. Corals 7. Coralla { _Acalephæ_ { 8. Hood-jellies 8. Hydromedusæ { { 9. Comb-jellies 9. Ctenophora
C. {V. Bloodless worms {10. Planary worms 10. Platyhelminthes =Worms= { _Acœlomi_ { {11. Round worms 11. Nemathelminthes +Vermes+ { VI. Blood-bearing {12. Moss-polyps 12. Bryozoa { worms {13. Sac-worms 13. Tunicata { _Cœlomati_ {14. Proboscideans 14. Rhynchocœla { {15. Star-worms 15. Gephyrea { {16. Wheel animalcules 16. Rotatoria { {17. Ring-worms 17. Annelida
D. {VII. Headless shellfish {18. Lamp-shells 18. Spirobranchia =Molluscs= { _Acephala_ {19. Mussels 19. Lamellibranchia { +Mollusca+ { VIII. Head-bearing {20. Snails 20. Cochlides { _Eucephala_ {21. Cuttles 21. Cephalopoda
E. { IX. Ringed-arms {22. Sea-stars 22. Asterida =Star-fishes= { _Colobrachia_ {23. Lily-stars 23. Crinoida { +Echinoderma+ { X. Armless {24. Sea-urchins 24. Echinida { _Lipobrachia_ {25. Sea-cucumbers 25. Holothuriæ
F. { XI. Gill-breathers {26. Crab-fish 26. Crustacea =Articulated= { _Carides_ { =Animals= { { XII. Tube-breathers {27. Spiders 27. Arachnida +Arthropoda+ { _Tracheata_ {28. Centipedes 28. Myriopoda { {29. Flies 29. Insecta
{ XIII. Skull-less {30. Lancelets 30. Leptocardia { _Acrania_ { G. { =Vertebrate= { XIV. Single-nostriled {31. Lampreys 31. Cyclostoma =Animals= { _Monorrhina_ { { +Vertebrata+ { XV. Amnion-less {32. Fishes 32. Pisces { _Anamnia_ {33. Mud-fish 33. Dipneusta { {34. Sea dragons 34. Halisauria { {35. Amphibians 35. Amphibia { { XVI. Amnion-bearing {36. Reptiles 36. Reptilia { _Amniota_ {37. Birds 37. Aves { {38. Mammals 38. Mammalia
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+Vertebrata+ (_Vertebrated animals_) Craniota | +Arthropoda+ | +Mollusca+ (_Articulated Animals_) | (_Molluscs_) +Echinoderma+ Tracheata | Eucephala (_Star-fishes_) | | | | | | Lipobrachia Crustacea Acrania | | Annelida | | Tunicata Acephala | | | | | Bryozoa | Colobrachia | | | | | | | Gephyrea| | Rotatoria \--v---/ \--v---/ | | | | | | | | | | | | | | \---------v-------------------/ \------v-----/ | | | +Vermes+ | | (_Worms_) | \------------------v-------------------/ | | CŒLOMATI (_Worms with a body-cavity_) | Platyhelminthes | | | | \-----v------/ +Zoophyta+ | (_Animal-Plants_) +ACŒLOMI+ Spongiæ Acalephæ (_Worms without body-cavity_) | | | | | | \------v-------/ | | | Protascus Prothelmis | | | | \-----------v---------------/ | | +Protozoa+ | (_Primæval animals_) | |/-------------^-------------\ +GASTRÆA+ | | Infusoria | | +PLANÆA+ | Gregarinæ | | | | | | +SYNAMŒBÆ+ \----v---/ | | | | \---------v--------/ +AMŒBÆ+ | +MONERA+
The first province of the Protozoa consists of the _Egg animals_ (Ovularia); we include among them all _single-celled animals_, all animals whose body, in the fully developed state, possesses the form-value of a _simple plastid_ (of a cytod or a cell), also those simple animal forms whose body consists of an aggregation of several cells perfectly similar one to another.
The _Archaic animals_ (Archezoa) form the first class in the series of Egg animals. It contains only the most simple and most ancient primary forms of the animal kingdom, whose former existence we have proved by means of the fundamental law of biogenesis; they are, (1) Animal Monera; (2) Animal Amœbæ; (3) Animal Synamœbæ. We may, if we choose, include among them a portion of the still living Monera and Amœbæ, but another portion (according to the discussion in Chapter XVI.) must on account of their neutral nature be considered as Protista, and a third portion, on account of their vegetable nature, must be considered as plants.
A second class of the egg animals consists of the _Gregarines_ (Gregarinæ), which live as parasites in the intestines and body-cavities of many animals. Some of these Gregarines are perfectly simple cells like the Amœbæ; some form chains of two or three identical cells, one lying behind the other. They differ from the naked Amœbæ by possessing a thick, simple membrane, which surrounds their cell-body; they can be considered as animal Amœbæ which have adopted a parasitical mode of life, and in consequence have surrounded themselves with a secreted covering.
As a third class of egg animals, we adopt the real _Infusoria_ (Infusoria), embracing those forms to which modern zoology almost universally limits this class of animals. The principal portion of them consists of the small _ciliated Infusoria_ (Ciliata), which inhabit all the fresh and salt waters of the earth in great numbers, and which swim about by means of a delicate garb of vibratile fringes. A second and smaller division consists of the adherent _sucking Infusoria_ (Acinetæ), which take their food by means of fine sucking-tubes. Although during the last thirty years numerous and very careful investigations have been made on these small animalcules,—which are mostly invisible to the naked eye,—still we are even now not very sure about their development and form-value. We do not even yet know whether the Infusoria are single or many-celled; but as no investigator has as yet proved their body to be a combination of cells, we are, in the mean time, justified in considering them as single-celled, like the Gregarines and the Amœbæ.
The second main class of primæval animals consists of the _Germ animals_ (Blastularia). This name we give to those extinct Protozoa which correspond to the two ontogenetic embryonic forms of the six higher animal tribes, namely, the Planula and the Gastrula. The body of these Blastularia, in a perfectly developed state, was composed of many cells, and these cells moreover differentiated—in two ways at least—into an external (animal or dermal) and an internal (vegetative or gastral) mass. Whether there still exist representatives of this group is uncertain. Their former existence is undoubtedly proved by the two exceedingly important ontogenetic animal forms which we have already described as Planula and Gastrula, and which still occur as a transient stage of development in the ontogeny of the most different tribes of animals. Corresponding to these, we may, according to the biogenetic principle, assume the former existence of two distinct classes of Blastularia, namely, the _Planæada_ and _Gastræada_. The type of the _Planæada_ is the _Planæa_—long since extinct—but whose historical portrait is still presented to us at the present day in the widely distributed _ciliated larva_ (Planula). (Frontispiece, Fig. 4.) The type of the _Gastræada_ is the _Gastræa_, of whose original nature the mouth-and-stomach larva (Gastrula), which recurs in the most different animal tribes, still gives a faithful representation. (Frontispiece Fig. 5, 6.) Out of the Gastræa, as we have previously mentioned, there were at one time developed two different primary forms, the Protascus and Prothelmis; the former must be looked upon as the primary form of the Zoophytes, the latter as the primary form of Worms. (Compare the enunciation of this hypothesis in my Monograph of the Calcareous Sponges, vol i. p. 464.)
The _Animal-plants_ (Zoophyta, or Cœlenterata) which constitute the second tribe of the animal kingdom, rise considerably above the primitive animals in the characters of their whole organisation, while they remain far below most of the higher animals. For in the latter (with the exception only of the lowest forms) the four distinct functions of nutrition—namely, digestion, circulation of the blood, respiration, and excretion—are universally accomplished by four perfectly different systems of organs: by the intestines, the vascular system, the organs of respiration, and the urinary apparatus. In Zoophytes, however, these functions and their organs are not yet separate, and are all performed by a single system of alimentary canals, by the so-called gastro-vascular system, or the cœlenteric apparatus of the intestinal cavity. The mouth, which is also the anus, leads into a stomach, into which the other cavities of the body also open. In Zoophytes the body-cavity, or “cœloma,” possessed by the four higher tribes of animals is still completely wanting, likewise the vascular system and blood, as also the organs of respiration, etc.
All Zoophytes live in water; most of them in the sea, only a very few in fresh water, such as fresh-water sponges (Spongilla) and some primæval polyps (Hydra, Cordylophora). A specimen of the pretty flower-like forms which are met with in great variety among Zoophytes is given on Plate VII. (Compare its explanation in the Appendix.)
The tribe of animal-plants, or Zoophytes, is divided into two distinct provinces, the _Sponges_, or _Spongiæ_, and the _Sea-nettles_, or _Acalephæ_ (p. 144). The latter are much richer in forms and more highly organized than the former. In all Sponges the entire body, as well as the individual organs, are differentiated and perfected to a much less extent than in Sea-nettles. All Sponges lack the characteristic _nettle-organs_ which all Sea-nettles possess.
The common primary form of all Zoophytes must be looked for in the _Protascus_, an animal form long since extinct, but whose existence is proved according to the biogenetic principle by the Ascula. This Ascula is an ontogenetical development form which, in Sponges as well as in Sea-nettles, proceeds from the Gastrula. (Compare the Ascula of the calcareous sponge on the Frontispiece, Fig. 7, 8.) For after the Gastrula of zoophytes has for a time swum about in the water it sinks to the bottom, and there adheres by that pole of its axis which is opposite to the opening of the mouth. The external cells of the ectoderm draw in their vibrating, ciliary hairs, whereas, on the contrary, the inner cells of the entoderm begin to form them. Thus the Ascula, as we call this changed form of larva, is a simple sack, its cavity (the cavity of the stomach or intestine) opening by a mouth externally, at the upper pole of the longitudinal axis (opposite the basal point of fixture). The entire body is here in a certain sense a mere stomach or intestinal canal, as in the case of the Gastrula. The wall of the sack, which is both body wall and intestinal wall, consists of two layers or coats of cells, a fringed _entoderm_, or gastral layer (corresponding with the inner or vegetative germ-layer of the higher animals), and an unfringed exoderm or dermal layer (corresponding with the external or animal germ-layer of the higher animals). The original _Protascus_, a true likeness of which is still furnished by the Ascula, probably formed egg-cells and sperm-cells out of its gastral layer.
The Protascads—as we will call the most ancient group of vegetable animals, represented by the Protascus-type—divided into two lines or branches, the Spongiæ and the Sea-nettles, or Acalephæ. I have shown in my Monograph of the Calcareous Sponges (vol. i. p. 485) how closely these two main classes of Zoophytes are related, and how they must both be derived, as two diverging forms, from the Protascus-form. The primary form of Spongiæ, which I have there called Archispongia, arose out of the Protascus by the formation of pores through its body-wall; the primary form of Sea-nettles, which I there called Archydra, developed out of the Protascus by the formation of nettle-organs, as also by the formation of feelers or tentacles.
The main-class or branch of the _Sponges_, _Spongiæ_, or _Porifera_, lives in the sea, with the single exception of the green fresh-water Sponge (Spongilla). These animals were long considered as plants, later as Protista; in most Manuals they are still classed among the primæval animals, or Protozoa. But since I have demonstrated their development out of the Gastrula, and the construction of their bodies of two cellular germ-layers (as in all higher animals), their close relationship to Sea-nettles, and especially to the Hydrapolyps, seems finally to be established. The _Olynthus_ especially, which I consider as the common primary form of calcareous sponges, has thrown a complete and unmistakable light upon this point.
The numerous forms comprised in the class of Spongiæ have as yet been but little examined; they may be divided into three legions and eight orders. The first legion consists of the soft, gelatinous _Mucous Sponges_ (Myxospongiæ), which are characterized by the absence of any hard skeleton. Among them are, on the one hand, the long-since-extinct primary forms of the whole class, the type of which I consider to be the Archispongia; on the other hand there are the still living, gelatinous sponges, of which the _Halisarca_ is best known. We can obtain a notion of the Archispongia, the most ancient primæval sponge, if we imagine the Olynthus (see Frontispiece), to be deprived of its radiating calcareous spiculæ.
The second legion of Spongiæ contains the _Fibrous Sponges_ (Fibrospongiæ), the soft body of which is supported by a firm, fibrous skeleton. This fibrous skeleton often consists merely of so-called “horny fibres,” formed of a very elastic, not readily destructible, organic substance. This is the case for instance in our common bathing Sponge (Euspongia officinalis), the purified skeleton of which we use every morning when washing. Blended with the horny, fibrous skeleton of many of these Sponges, there are numerous flinty spicula; this is the case for example with the fresh-water Sponge (Spongilla). In others the whole skeleton consists of only calcareous or silicious spicula which are frequently interwoven into an extremely beautiful lattice-work, as in the celebrated Venus’ Flower Basket (Euplectella). Three orders of fibrous sponges may be distinguished according to the different formation of the spicula, namely, Chalynthina, Geodina, and Hexactinella. The natural history of the fibrous sponges is of especial interest to the Theory of Descent, as was first shown by Oscar Schmidt, the greatest authority on this group of animals. In no other group, perhaps, can the unlimited pliability of the specific form, and its relation to Adaptation and Inheritance, be so clearly followed step by step; perhaps in no other group is the species so difficult to limit and define.
This proposition, which applies to the great legion of the Fibrous Sponges, applies in a still higher degree to the smaller but exceedingly interesting legion of the calcareous sponges (Calcispongiæ), on which in 1872, after five years’ careful examination, I published a comprehensive Monograph. The sixty plates of figures accompanying this Monograph explain the extreme pliability of these small sponges “good species” of which, in fact, cannot be spoken of in the usual systematic sense. We find among them only varying series of forms, which do not even completely transmit their specific form to their nearest descendants, but by adaptation to subordinate, external conditions of existence, perpetually change. It frequently occurs here, that there arise out of one and the same stock different form-species, which according to the usual system would belong to several quite distinct genera; this is the case, for instance, with the remarkable Ascometra (Frontispiece, Fig. 10.) The entire external bodily form is much more pliable and protean in Calcareous Sponges than in the silicious sponges, which are characterized by possessing silicious spicula, forming a beautiful skeleton. Through the study of the comparative anatomy and ontogeny of calcareous sponges, we can recognise, with the greatest certainty, the common primary form of the whole group, namely, the sack-shaped _Olynthus_, whose development is represented in the Frontispiece (compare its explanation in the Appendix). Out of the Olynthus (Fig. 9 on the Frontispiece), the order of the Ascones was the first to develop, out of which, at a later period, the two other orders of Calcareous Sponges, the _Leucones_ and _Sycones_, arose as diverging branches. Within these orders, the descent of the individual forms can again be followed step by step. Thus the Calcareous Sponges in every respect confirm the proposition which I have elsewhere maintained: that “the natural history of sponges forms a connected and striking argument in favour of Darwin.”
The second main class or branch in the tribe of Zoophytes is formed by the Sea-nettles (Acalephæ, or Cnidæ). This interesting group of animals, so rich in forms, is composed of three different classes, namely, the Hood-jellies (Hydromedusæ), the Comb-jellies (Ctenophora), and the Corals (Coralla). The hypothetical, extinct Archydra must be looked upon as the common primary form of the whole group; it has left two near relations in the still living fresh-water polyps (Hydra and Cordylophora). The Archydra was very closely related to the simplest forms of Spongiæ (Archispongia and Olynthus), and probably differed from them only by possessing nettle organs, and by the absence of cutaneous pores. Out of the Archydra there first developed the different Hydroid polyps, some of which became the primary forms of Corals, others the primary forms of Hydromedusæ. The Ctenophora developed later out of a branch of the latter.
The Sea-nettles differ from the Spongiæ (with which they agree in the characteristic formation of the system of the alimentary canal) principally by the constant possession of nettle organs. These are small bladders filled with poison, large numbers—generally millions—of which are dispersed over the skin of the sea nettles, and which burst and empty their contents when touched. Small animals are killed by this; in larger animals this nettle poison causes a slight inflammation of the skin, just as does the poison of our common nettles. Any one who has often bathed in the sea, will probably have at times come in contact with large Hood-jellies (Jelly-fish), and become acquainted with the unpleasant burning feeling which their nettle organs can produce. The poison in the splendid blue Jelly-fish, Physalia, or Portuguese Man-of-war, acts so powerfully that it may lead to the death of a human being.
The class of Corals (Coralla) lives exclusively in the sea, and is more especially represented in the warm seas by an abundance of beautiful and highly-coloured forms like flowers. Hence they are also called _Flower-animals_ (Anthozoa). Most of them are attached to the bottom of the sea, and contain an internal calcareous skeleton. Many of them by continued growth produce such immense stocks that their calcareous skeletons have formed the foundation of whole islands, as is the case with the celebrated coral reefs and atolls of the South Seas, the remarkable forms of which were first explained by Darwin.(13) In corals the counterparts, or antimera—that is, the corresponding divisions of the body which radiate from and surround the central main axis of the body—exist sometimes to the number of four, sometimes to the number of six or eight. According to this we distinguish three legions, the Fourfold (Tetracoralla), Sixfold (Hexacoralla), and Eightfold corals (Octocoralla). The fourfold corals form the common primary group of the class, out of which the sixfold and eightfold have developed as two diverging branches.
SYSTEMATIC SURVEY
_Of the 4 Classes and 30 Orders of the Animal Plants, or Zoophytes._
----------------+--------------------+--------------------+--------------- _Class of the_ | _Legions of the_ | _Orders of the_ | _A Genus Name_ _Zoophytes._ | _Zoophytes._ | _Zoophytes._ | _as example._ ----------------+--------------------+--------------------+--------------- I. { I. Myxospongiæ { 1. Archispongina | Archispongia { _Mucous Sponges_ { 2. Halisarcina | Halisarca =Sponges= { | { II. Fibrospongiæ { 3. Chalynthina | Spongilla +Spongiæ+ {_Fibrous Sponges_ { 4. Geodina | Ancorina or { { 5. Hexactinella | Euplectella +Porifera+ { | { III. Calcispongiæ { 6. Ascones | Olynthus {_Calcareous Sponges_ { 7. Leucones | Dyssycus { { 8. Sycones | Sycurus | | II. { IV. Tetracoralla { 9. Rugosa | Cyathophyllum {_Fourfold Corals_ { 10. Paranemeta | Cereanthus =Corals= { | { V. Hexacoralla { 11. Cauliculata | Antipathes +Coralla+ {_Sixfold Corals_ { 12. Madreporaria | Astræa or { { 13. Halirhoda | Actinia +Anthozoa+ { | { VI. Octocoralla { 14. Alcyonida | Lobularia {_Eightfold Corals_ { 15. Gorgonida | Isis { { 16. Pennatulida | Veretillum | | III. { VII. Archydræ } 17. Hydraria | Hydra {_Primæval Polyps_ } | =Jelly-polyps= { | { VIII. Leptomedusæ { 18. Vesiculata | Sertularia +Hydromedusæ+ {_Soft Jelly-fish_ { 19. Ocellata | Tubularia { { 20. Siphonophora | Physophora or { | { IX. Trachymedusæ { 21. Marsiporchida| Trachynema =Hood-jellies= { _Hard Jelly-fish_ { 22. Phyllorchida | Geryonia { { 23. Elasmorchida | Charybdæ +Medusa+ { | { X. Calycozoa } 24. Podactinaria | Lucernaria { _Stalked Jellies_ } | { | { XI. Discomedusæ { 25. Semæostomeæ | Aurelia { _Disc-jellies_ { 26. Rhizostomeæ | Crambessa | | IV. { XII. Eurystoma } 27. Beroida | Beroe { _Wide-mouthed_ } | =Comb-jellies= { | { XIII. Stenostoma { 28. Saccata | Cydippe +Ctenophora+ { _Narrow-mouthed_ { 29. Lobata | Eucharis { { 30. Tæniata | Cestum
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Ctenophora Hydromedusæ Tæniata Lobata Rhizostomeæ | | | | | | \---v----/ Semæostomeæ Saccata DISCOMEDUSÆ STENOSTOMA | | Trachymedusæ | Siphonophora | | | | | | Lucernaria | | EURYSTOMA> | Calycozoa | | | | | | | | | | | | \-------------------------v----------------/ LEPTOMEDUSÆ Coralla | Octocoralla | Hexacoralla | | | | | | | | \----v----/ | Tetracoralla | Spongiæ | | Fibrospongiæ Calcispongiæ | | Chalynthina Leucones Sycones| | | | | | | | | | | | | | | | | Hexactinella|Geodina Dyssycus Sycurus | Hydroida | | | | | | | Cordylophora | | | | | | | | Hydra \------v---/ \-----v--/ | | | | | Ascones | | | | | Myxospongia | | \-----v-------/ | Halisarcina | | HYDROIDA | | | Procorallum | | | | | | CHALYNTHUS | OLYNTHUS | | | | | \-----v--------/ \------v----------/ Hydroida Archispongiæ | | | | Archydra | | \--------------v--------/ Protascus | Gastræa
The second class of Sea-nettles is formed by the _Hood-jellies_ (Medusæ) or _Polyp-jellies_ (Hydromedusæ). While most corals form stocks like plants, and are attached to the bottom of the sea, the Hood-jellies generally swim about freely in the form of gelatinous bells. There are, however, numbers of them, especially the lower forms, which adhere to the bottom of the sea, and resemble pretty little trees. The lowest and simplest members of this class are the little fresh-water polyps (Hydra and Cordylophora). We may look upon them as but little changed descendants of those _Primæval polyps_ (Archydræ), from which, during the primordial period, the whole division of the Sea-nettles originated. Scarcely distinguishable from the Hydra are the adherent Hydroid polyps (Campanularia, Tubularia), which produce freely swimming medusæ by budding, and out of the eggs of these there again arise adherent polyps. These freely swimming Hood-jellies are mostly of the form of a mushroom, or of an umbrella, from the rim of which many long and delicate tentacles hang. They are among the most beautiful and most interesting inhabitants of the sea. The remarkable history of their lives, and especially the complicated alternation of generation of polyps and medusæ, are among the strongest proofs of the truth of the theory of descent. For just as Medusæ still daily arise out of the Hydroids, did the freely swimming medusa-form originally proceed, phylogenetically, out of the adherent polyp-form. Equally important for the theory of descent is the remarkable _division of labour_ of the individuals, which among some of them is developed to an astonishingly high degree, more especially in the splendid _Siphonophora_.(37) (Plate VII. Fig. 13.)
The third class of Sea-nettles—the peculiar division of Comb-jellies (Ctenophora), probably developed out of a branch of the Hood-jellies. The Ctenophora, which are also called Ribbed-jellies, possess a body of the form of a cucumber, which, like the body of most Hood-jellies, is as clear and transparent as crystal or cut glass. Comb or Ribbed-jellies are characterized by their peculiar organs of motion, namely, by eight rows of paddling, ciliated leaflets, which run in the form of eight ribs from one end of the longitudinal axis (from the mouth) to the opposite end. Those with narrow mouths (Stenostoma) probably developed later out of those with wide mouths (Eurystoma). (Compare Plate VII. Fig. 16.)
The third tribe of the animal kingdom, the phylum of _Worms_ or worm-like animals (Vermes, or Helminthes), contains a number of diverging branches. Some of these numerous branches have developed into well-marked and perfectly independent classes of Worms, but others changed long since into the original, radical forms of the four higher tribes of animals. Each of these four higher tribes (and likewise the tribe of Zoophytes) we may picture to ourselves in the form of a lofty tree, whose branches represent the different classes, orders, families, etc. The phylum of Worms, on the other hand, we have to conceive as a low bush or shrub, out of whose root a mass of independent branches shoot up in different directions. From this densely branched shrub, most of the branches of which are dead, there rise four high stems with many branches. These are the four lofty trees just mentioned as representing the higher phyla—the Echinoderma, Articulata, Mollusca, and Vertebrata. These four stems are directly connected with one another at the root only, to wit, by the common primary group of the Worm tribe.
The extraordinary difficulties which the systematic arrangement of Worms presents, for this reason merely, are still more increased by the fact that we do not possess any fossil remains of them. Most of the Worms had and still have such soft bodies that they could not leave any characteristic traces in the neptunic strata of the earth. Hence in this case again we are entirely confined to the records of creation furnished by ontogeny and comparative anatomy. In making then the exceedingly difficult attempt to throw a few hypothetical rays of light upon the obscurity of the pedigree of Worms, I must therefore expressly remark that this sketch, like all similar attempts possesses only a provisional value.
The numerous classes distinguished in the tribe of Worms, and which almost every zoologist groups and defines according to his own personal views, are, in the first place, divided into two essentially different groups or branches, which in my Monograph of the Calcareous Sponges I have termed Acœlomi and Cœlomati. For all the lower Worms which are comprised in the class of Flat-worms (Platyhelminthes), (the Gliding-worms, Sucker-worms, Tape-worms), differ very strikingly from other Worms, in the fact that they possess neither blood nor body-cavity (no cœlome); they are, therefore, called Acœlomi. The true cavity, or cœlome, is completely absent in them as in all the Zoophytes; in this important respect the two groups are directly allied. But _all other Worms_ (like the four higher tribes of animals) possess a genuine body-cavity and a vascular system connected with it, which is filled with blood; hence we class them together as _Cœlomati_.
The main division of _Bloodless Worms_ (Acœlomi) contains, according to our phylogenetic views, besides the still living Flat-worms, the unknown and extinct primary forms of the whole tribe of Worms, which we shall call the Primæval Worms (Archelminthes). The type of these _Primæval Worms_, the ancient Prothelmis, may be directly derived from the Gastræa (p. 133). Even at present the Gastrula-form—the faithful historical portrait of the Gastræa—recurs in the ontogenesis of the most different kinds of worms as a transient larva-form. The ciliated Gliding-worms (Turbellaria), the primary group of the present Planary or Flat-worms (Platyhelminthes), are the nearest akin to the Primæval Worms. The parasitical Sucker-worms (Trematoda) arose out of the Gliding-worms, which live freely in water, by adaptation to a parasitical mode of life; and out of them later on—by an increasing parasitism—arose the Tape-worms (Cestoda).
Out of a branch of the Acœlomi arose the second main division of the Worm tribe, the Worms with blood and body-cavity (Cœlomati): of these there are seven different classes.
The Pedigree on p. 151 shows how the obscure phylogeny of the seven classes of Cœlomati may be supposed to stand. We shall, however, mention these classes here quite briefly, as their relationships and derivation are, at present, still very complicated and obscure. More numerous and more accurate investigations of the ontogeny of the different Cœlomati will at some future time throw light upon their phylogenesis.
The Round Worms (Nemathelminthes) which we mention as the first class of the Cœlomati, and which are characterized by their cylindrical form, consist principally of parasitical Worms which live in the interior of other animals. Of human parasites, the celebrated Trichinæ, the Maw-worms, Whip-worms, etc., for example, belong to them. The Star-worms (Gephyrea) which live exclusively in the sea are allied to round worms, and the comprehensive class of Ring-worms (Annelida) are allied to the former. To the Ring-worms, whose long body is composed of a number of segments, all alike in structure, belong the Leeches (Hirudinea), Earth-worms (Lumbricina), and all the marine bristle-footed Worms (Chætopoda). Nearly akin to them are the Snout-worms (Rhynchocœla), and the small microscopic Wheel-worms (Rotifera). The unknown, extinct, primary forms of the tribe of Sea-stars (Echinoderma), and of the tribe of the articulated animals (Arthropoda), were nearest akin to the Ring-worms. On the other hand, we must probably look for the primary forms of the great tribe of Molluscs in extinct Worms, which were very closely related to the Moss-polyps (Bryozoa) of the present day; and for the primary forms of the Vertebrata in the unknown Cœlomati, whose nearest kin of the present day are the Sea-sacs, especially the Ascidia.
SYSTEMATIC SURVEY
_Of the 8 Classes and 22 Orders of the Worm Tribe._
(Compare Gen. Morph. ii. Plate V. pp. 75-77.)
------------------+-------------------------+----------------------+------------------ _Classes_ | | _Systematic_ | _of the_ | _Orders of the_ | _Name of the_ | _Name of a Genus_ _Worm Tribe._ | _Worm Tribe._ | _Orders of Worms_. | _as example._ | | | ------------------+-------------------------+----------------------+------------------ 1. _Flat_ { 1. Primæval worms | 1. Archelminthes | Prothelmis _Worms_ { 2. Gliding-worms | 2. Turbellaria | Planaria Platyhel- { 3. Sucker-worms | 3. Trematoda | Distoma minthes { 4. Tape-worms | 4. Cestoda | Tænia | | 2. _Round_ { 5. Arrow-worms | 5. Chætognatha | Sagitta _Worms_ { 6. Thread-worms | 6. Nematoda | Trichina Nemathel- { 7. Hook-headed | 7. Acanthocephala | Echinorhynchus minthes { worms | | | | 3. _Moss_ } 8. Horse-shoe-lipped | 8. Lophopoda | Alcyonella _Polyps_ } 9. Circle-lipped | 9. Stelmopoda | Retepora Bryozoa } | | | | 4. _Sea-sacs_ { 10. Sea-squirts | 10. Ascidia | Phallusia Tunicata { 11. Sea-barrels | 11. Thaliacea | Salpa | | 5. _Proboscideans_ } 12. Tongue-worms | 12. Enteropneusta | Balanoglossus Rhynchocœla } 13. Cord-worms | 13. Nemertina | Borlasia | | { 14. Star-worms without | 14. Sipunculida | Sipunculus 6. _Star-Worms_ { bristles | | Gephyrea { 15. Star-worms with | 15. Echiurida | Echiurus { bristles | | | | 7. _Wheel_ } | | _Animalcule_ } 16. Wheel-worms | 16. Rotatoria | Hydatina Rotifera } | | | | { 17. Bear-worms | 17. Arctisca | Macrobiotus { 18. Worms with claws | 18. Onychophora | Peripatus 8. _Ring_ { 19. Leeches | 19. Hirudinea | Hirudo _Worms_ { 20. Land-worms | 20. Drilomorpha | Lumbricus Annelida { 21. Mailed worms | 21. Phracthelminthes | Crossopodia { 22. Bristle-footed | 22. Chætopoda | Aphrodite { worms | |
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Chætopoda | Drilomorpha | | \-------v-------/ | Phracthelminthes | | | Echiurida| | Hirudinea Sipunculida | | | | | | | | | Onychophora \----v---/ | | | | +Gephyrea+ | | | | Arctisca | | | | | | | | | | \--v-—/ Chætognatha | \--v-----/ | | | | | \-----v-----/ Nematoda | | | | | | | \---------v---------/ \--v--/ | +Annelida+ | Acantho- | | | cephala | Stelmopoda | | | | | | Enteropneusta \--v--/ | | | | +Nemathelminthes+ | Lophopoda | Ascidia | | | +Bryozoa+ | Thalicea | | Nemertina | | | | | | | | | | | +Rotifera+ | | | | | | | | | | | | \---v-/ | | | | | | | +Rhynchocœla+ | | | | | \--v---/ | | | | \----v---/ +Tunicata+ | | | | | | | \-----------------------v-----------------------------/ +Cœlomati+ (_worms with body-cavity_) | Cestoda | | | | | Trematoda | | | | | Turbellaria | +Platyhelminthes+ | | | | | \-------v---------/ +Acœlomi+ (_worms without body-cavity_) | | Archelminthes +Prothelmis+ | | +Gastræa+
The class of Sea-sacs (Tunicata) is one of the most remarkable among Worms. They all live in the ocean, where some of the Ascidiæ adhere to the bottom, while others (the sea-barrels, or Thaliacea) swim about freely. In all of them the non-jointed body has the form of a simple barrel-shaped sack, which is surrounded by a thick cartilaginous mantle. This mantle consists of the same non-nitrogenous combination of carbon, which, under the name of cellulose, plays an important part in the Vegetable Kingdom, and forms the largest portion of vegetable cellular membranes, and consequently also the greater part of wood. The barrel-shaped body generally possesses no external appendages. No one would recognise in them a trace of relationship to the highly differentiated vertebrate animals. And yet this can no longer be doubted, since Kowalewsky’s investigations, which in the year 1867 suddenly threw an exceedingly surprising and unmistakable light upon them. From these investigations it has become clear that the individual development of the adherent simple Ascidian Phallusia agrees in most points with that of the lowest vertebrate animal, namely, the Lancelet (Amphioxus lanceolatus). The early stages of the Ascidia possess the beginnings of the _spinal marrow_ and the _spinal column_ (chorda dorsalis) lying beneath it, which are the two most essential and most characteristic organs of the vertebrate animal. Accordingly, of all invertebrate animals known to us, the _Tunicates are without doubt the nearest blood relations of the Vertebrates_, and must be considered as the nearest relations of those Worms out of which the vertebrate tribe has developed. (Compare Plates XII. and XIII.)
While thus different branches of the Cœlomatous group of the Worms furnish us with several genealogical links leading to the four higher tribes of animals, and give us important phylogenetic indications of their origin, the lower group of Acœlomi, on the other hand, show close relationships to the Zoophytes, and to the Primæval animals. The great phylogenetic interest of the Worm tribe rests upon this peculiar intermediate position.