The second class of plants with enclosed seeds, the two seed-lobed (Dicotylæ, or Dicotyledons, also called Exogenæ) presents much greater historical and anatomical interest in the development of its subordinate groups. The flowering plants of this class generally possess, as their name indicates, two seed lobes or germ leaves (cotyledons). The number of leaves composing its blossom is generally not three, as in most Monocotyledons, but four, five, or a multiple of those numbers. Their leaves, moreover, are generally more highly differentiated and more composite than those of the Monocotyledons; they are traversed by crooked, branching bunches of vessels or “veins.” To this class belong most of the leafed trees, and as they predominate in the tertiary period as well as, at present, over the Gymnosperms and Ferns, we may call the cænolithic period that of leafed forests.
Although the majority of Dicotyledons belong to the most highly developed and most perfect plants, still the lowest division of them is directly allied to the Gymnosperms, and particularly to the Gnetaceæ. In the lower Dicotyledons, as in the case of the Monocotyledons, calyx and corolla are as yet not differentiated. Hence they are called Apetalous (Monochlamydeæ, or Apetalæ). This sub-class must therefore doubtless be looked upon as the original group of the Angiosperms, and existed probably even during the Trias and Jura periods. Among them are most of the leafed trees bearing catkins—birches and alders, willows and poplars, beeches and oaks; further, the plants of the nettle kind—nettles, hemp, and hops, figs, mulberries, and elms; finally, plants like the spurges, laurels, and amaranth.
It was not until the chalk period that the second and more perfect class of the Dicotyledons appeared, namely, the group with corollas (Dichlamydeæ, or Corollifloræ). These arose out of the Apetalæ from the simple cover of the blossoms of the latter becoming differentiated into calyx and corolla. The sub-class of the Corollifloræ is again divided into two large main divisions or legions, each of which contains a large number of different orders, families, genera, and species. The first legion bears the name of star-flowers, or Diapetalæ, the second that of the bell-flowers, or Gamopetalæ.
The lower and less perfect of the two legions of the Corollifloræ are the star-flowers (also called Diapetalæ or Dialypetalæ). To them belong the extensive families of the Umbelliferæ, or umbrella-worts (wild carrot, etc.), the Cruciferæ, or cruciform blossoms (cabbage, etc.); further, the Ranunculaceæ (buttercups) and Crassulaceæ, the Mallows and Geraniums, and, besides many others, the large group of Roses (which comprise, besides roses, most of our fruit trees), and the Pea-blossoms (containing, among others, beans, clover, genista, acacia, and mimosa). In all these Diapetalæ the blossom-leaves remain separate, and never grow together, as is the case in the Gamopetalæ. These latter developed first in the tertiary period out of the Diapetalæ, whereas the Diapetalæ appeared in the chalk period together with the Apetalæ.
The highest and most perfect group of the vegetable kingdom is formed by the second division of the Corollifloræ, namely, the legion of bell-flowers (Gamopetalæ, also called Monopetalæ or Sympetalæ). In this group the blossom-leaves, which in other plants generally remain separate, grow regularly together into a more or less bell-like, funnel-shaped, or tubular flower. To them belong, among others, the Bell-flowers and Convolvulus, Primroses and Heaths, Gentian and Honeysuckle, further the family of the Olives (olive trees, privet, elder, and ash), and finally, besides many other families, the extensive division of the Lip-blossoms (Labiatæ) and the Composites. In these last the differentiation and perfection of the Phanerogamic blossoms attain their highest stage of development, and we must therefore place them at the head of the vegetable kingdom, as the most perfect of all plants. In accordance with this, the legion of the Gamopetalæ appear in the organic history of the earth later than all the main groups of the vegetable kingdom—in fact, not until the cænolithic or tertiary epoch. In the earliest tertiary period the legion is still very rare, but it gradually increases in the mid-tertiary, and attains its full development only in the latest tertiary and the quaternary period.
Now if, having reached our own time, we look back upon the whole history of the development of the vegetable kingdom, we cannot but perceive in it a grand confirmation of the Theory of Descent. The two great principles of organic development which have been pointed out as the necessary results of natural selection in the Struggle for Life, namely, the laws of differentiation and perfecting, manifest themselves everywhere in the development of the larger and smaller groups of the natural system of plants. In each larger or smaller period of the organic history of the earth, the vegetable kingdom increases both in variety and perfection, as a glance at Plate IV. will clearly show. During the whole of the long primordial period there existed only the lowest and most imperfect group, that of the Algæ. To these are added, in the primary period, the higher and more perfect Cryptogamia, especially the main-class of Ferns. During the coal period the Phanerogamia begin to develop out of the latter; at first, however, they are represented only by the lower main-class, that of Gymnosperms. It was not until the secondary period that the higher main-class, that of Angiosperms, arose out of them. Of these also there existed at first only the lower groups without distinct corollas, the Monocotyledons and the Apetalæ. It was not until the chalk period that the higher Corollifloræ developed out of the latter. But even this most highly developed group is represented, in the chalk period, only by the lower stage of Star-flowers, or Diapetalæ, and only at quite a late date, in the tertiary period, did the more highly developed Bell-blossoms, Gamopetalæ, arise out of them, which at the same time are the most perfect of all flowering plants. Thus, in each succeeding later division of the organic history of the earth the vegetable kingdom gradually rose to a higher degree of perfection and variety.
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?
| Haeckel-History of Creation | Pl. VI. |
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, equiformal, and equivalent cells. Even at the present day, in the ontogenetic development of every animal egg-cell, there first arises a globular heap of equiformal 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).
PARALLELISM OF ONTOGENY AND PHYLOGENY
| Definition of the forms of the five first stages of the development of the animal body. | Ontogenesis. | Phylogenesis. |
| The five first stages The five first stages of the individual development. | The five first stages of the phyletic or historical development. | |
| First Stage of Development. | 1. Monerula. |
1. Moneron. |
| A simple cytod (a plastid without a kernel.) | Animal egg without a kernel (when the egg-kernel has disappeared, after being fructified). | Most ancient animal Monera, originating by spontaneous generation. |
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| Second Stage of Development. | 2. Ovulum. |
2. Amœbæ. |
| A simple cell (a plastid containing a kernel.) | Animal egg with kernel (a simple egg-cell). | Animal Amœbæ. |
| │ │ |
│ │ |
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| Third Stage of Development. | 3. Morula. (Mulberry form.) |
3. Synamœba. |
| A community (an aggregation of identical simple cells). | Globular heap of homogeneous “cleavage spheres.” | An aggregation of Amœbæ. |
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│ │ |
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| Fourth Stage of Development. | 4. Planula. (Ciliated larva.) |
4. Planæa. |
| A solid or bladder-shaped, globular, or oval body, composed of two kinds of different cells: externally ciliated, internally non-ciliated cells. | Many-celled larva without mouth, composed of different cells. | Many-celled primæval animal without mouth, composed of two kinds of different cells. |
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| Fifth Stage of Development. | 5. Gastrula. (Larva with mouth.) |
5. Gastræa. |
| A globular or oval body with simple intestinal cavity and mouth-opening. Body wall composed of two layers; an externally ciliated ectoderm (dermal layer), an internally non-ciliated entoderm (gastral layer). | Many-celled with intestines and mouth; intestinal wall with two layers. | Many-celled primæval animal with intestine and mouth; intestinal wall with two layers. (Primary form of zoophytes and worms.) |
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 anima 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 of the Animal Kingdom. |
Main Classes, Branches or Clades of the Animal Kingdom. |
Classes of the Animal Kingdom. |
Systematic Name of the Classes. |
||||
| A. Primæval Animals Protozoa |
I. Egg-animals Ovularia |
1. | Archaic animals | 1. | Archezoa | ||
| 2. | Gregarines | 2. | Gregarinæ | ||||
| 3. | Infusoria | 3. | Infusoria | ||||
| II. Mulberry animals Ovularia |
4. | Planæads | 4. | Planæadas | |||
| 5. | Gastræads | 5. | Gastræadas | ||||
| B. Animal Plants Zoophyta |
III. Sponges Spongiæ |
6. | Sponges | 6. | Porifera | ||
| IV. Sea-nettles Acalephæ |
7. | Corals | 7. | Coralla | |||
| 8. | Hood-jellies | 8. | Hydromedusæ | ||||
| 9. | Comb-jellies | 9. | Ctenophora | ||||
| C. Worms Vermes |
V. Bloodless worms Acœlomi |
10. | Planary worms | 10. | Platyhelminthes | ||
| V. Bloodless worms Acœlomi |
11. | Round worms | 11. | Nemathelminthes | |||
| 12. | Moss-polyps | 13. | Bryozoa | ||||
| 13. | Sac-worms | 13. | Tunicata | ||||
| 14. | Proboscideans | 14. | Rhynchocœla | ||||
| 15. | Star-worms | 15. | Gephyrea | ||||
| 16. | Wheel animalcules | 16. | Rotatoria | ||||
| 17. | Ring-worms | 17. | Annelida | ||||
| D. Molluscs Mollusca |
VII. Headless shellfish Acephala |
18. | Lamp-shells | 18. | Spirobranchia | ||
| 19. | Mussels | 19. | Lamellibranchia | ||||
| VIII. Head-bearing Eucephala |
20. | Snails | 20. | Cochlides | |||
| 21. | Cuttles | 21. | Cephalopoda | ||||
| E. Star-fishes Echinoderma |
IX. Ringed-arms Colobrachia |
22. | Sea-stars | 22. | Asterida | ||
| 23. | Lily-stars | 23. | Crinoida | ||||
| X. Armless Lipobrachia |
24. | Sea-urchins | 24. | Echinida | |||
| 25. | Sea-cucumbers | 25. | Holothuriæ | ||||
| F. Articulated Animals Arthropoda |
XI. Gill-breathers Carides |
26. | Crab-fish | 26. | Crustacea | ||
| X. Armless Lipobrachia |
27. | Spiders | 27. | Arachnida | |||
| 28. | Centipedes | 28. | Myriopoda | ||||
| 29. | Flies | 29. | Insecta | ||||
| G. Vertebrate Animals Vertebrata |
XIII. Skull-less Acrania |
30. |
Lancelets |
30. |
Leptocardia |
||
| XIV. Single-nostriled Lipobrachia |
31. |
Lampreys |
31. |
Cyclostoma |
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| XV. Amnion-less Anamnia |
32. | Fishes | 32. | Pisces | |||
| 33. | Mud-fish | 33. | Dipneusta | ||||
| 34. | Sea dragons | 34. | Halisauria | ||||
| 35. |
Amphibians |
35. |
Amphibia |
||||
| XVI. Amnion-bearing Amniota |
36. | Reptiles | 36. | Reptilia | |||
| 37. | Birds | 37. | Aves | ||||
| 38. | Mammals | 38. | Mammalia | ||||
MONOPHYLETIC PEDIGREE OF ANIMALS
| Vertebrata (Vertebrated animals) Craniota |
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| Arthropoda (Articulated Animals) Tracheata |
│ │ │ │ |
Mollusca (Molluscs) Eucephala |
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| Echinoderma (Star-fishes) |
│ │ |
│ │ |
│ │ |
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| Lipobrachia | Crustacea | Acrania | │ |
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| │ |
Annelida | │ |
│ |
Tunicata | Acephala | |||
| Colobrachia │ |
│ │ |
│ │ |
│ │ |
│ │ |
Bryozoa │ |
│ │ |
||
| │ |
Gephyrea | │ |
│ |
Rotatoria | ||||
| │ |
│ |
│ |
│ |
│ |
│ |
│ |
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| Vermes (Worms) |
│ │ |
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| │ |
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| Cœlomati (Worms with a body-cavity) |
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| │ |
Platyhelminthes | |||||||
| │ |
│ |
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| Zoophyta (Animal Plants) |
Acœlomi (Worms without body-cavity) |
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| Spongiæ | Acalephæ | │ |
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| │ |
│ |
│ |
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| │ |
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| Protascus | Prothelmis | |||||||
| │ |
│ |
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| │ |
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| │ │ │ |
Protozoa (Primæval animals) |
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| Gastræa | ||||||||
| │ |
Infusoria | |||||||
| Planæa | │ |
Gregarinæ | ||||||
| │ |
│ |
│ |
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| Synamœbæ | ||||||||
| │ |
│ |
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| │ |
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| Amœbæ | ||||||||
| │ |
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| 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.
| Pl. VII. |