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THE HISTORY OF CREATION.

Hypothetical Sketch of the Monophyletic Origin of Man

THE
HISTORY OF CREATION:
OR THE DEVELOPMENT OF THE EARTH AND ITS
INHABITANTS BY THE ACTION OF NATURAL CAUSES

A POPULAR EXPOSITION OF
THE DOCTRINE OF EVOLUTION IN GENERAL, AND OF THAT OF
DARWIN, GOETHE, AND LAMARCK IN PARTICULAR.

FROM THE GERMAN OF
ERNST HAECKEL,
PROFESSOR IN THE UNIVERSITY OF JENA.

THE TRANSLATION REVISED BY
E. RAY LANKESTER, M.A., F.R.S.,
FELLOW OF EXETER COLLEGE, OXFORD.

IN TWO VOLUMES.
VOL. II.

NEW YORK:
D. APPLETON AND COMPANY,
1, 3, AND 5 BOND STREET.
1880.

A sense sublime Of something far more deeply interfused, Whose dwelling is the light of setting suns, And the round ocean, and the living air, And the blue sky, and in the mind of man; A motion and a spirit that impels All thinking things, all objects of all thought, And rolls through all things.

In all things, in all natures, in the stars Of azure heaven, the unenduring clouds, In flower and tree, in every pebbly stone That paves the brooks, the stationary rocks, The moving waters and the invisible air. Wordsworth.

CONTENTS OF VOL. II.

LIST OF ILLUSTRATIONS.

PLATES.

FIGURES.

THE HISTORY OF CREATION.

CHAPTER XV.

PERIODS OF CREATION AND RECORDS OF CREATION.

Reform of Systems by the Theory of Descent.—The Natural System as a Pedigree.—Palæontological Records of the Pedigree.—Petrifactions as Records of Creation.—Deposits of the Neptunic Strata and the Enclosure of Organic Remains.—Division of the Organic History of the Earth into Five Main Periods: Period of the Tangle Forests, Fern Forests, Pine Forests, Foliaceous Forests, and of Cultivation.—The Series of Neptunic Strata.—Immeasurable Duration of the Periods which have elapsed during their Formation.—Deposits of Strata only during the Sinking, not during the Elevation of the Ground.—Other Gaps in the Records of Creation.—Metamorphic Condition of the most Ancient Neptunic Strata.—Small Extent of Palæontological Experience.—Small proportion of Organisms and of Parts of Organisms Capable of Petrifying.—Rarity of many Petrified Species.—Want of Fossilised Intermediate Forms.—Records of the Creation in Ontogeny and in Comparative Anatomy.

The revolutionary influence which the Theory of Descent must exercise upon all sciences, will in all probability affect no branch of science, excepting Anthropology, so much as the descriptive portion of natural history, that which is known as systematic Zoology and Botany. Most naturalists who have hitherto occupied themselves with arranging the different systems of animals and plants, have collected, named, and arranged the different species of these natural bodies with much the same interest as antiquarians and ethnographers collect the weapons and utensils of different nations. Many have not even risen above the degree of intelligence with which people usually collect, label, and arrange crests, stamps, and similar curiosities. In the same manner as some collectors find their pleasure in the similarity of forms, the beauty or rarity of the crests or stamps, and admire in them the inventive art of man, so many naturalists take a delight in the manifold forms of animals and plants, and marvel at the rich imagination of the Creator, at His unwearied creative activity, and at His curious fancy for forming, by the side of so many beautiful and useful organisms, also a number of ugly and useless ones.

This childlike treatment of systematic Zoology and Botany is completely annihilated by the Theory of Descent. In the place of the superficial and playful interest with which most naturalists have hitherto regarded organic structures, we now have the much higher interest of the intelligent understanding which detects in the related forms of organisms their true blood relationships. The Natural System of animals and plants, which was formerly valued either only as a registry of names, to facilitate the survey of the different forms, or as a table of contents for the short expression of their degrees of similarity, receives from the Theory of Descent the incomparably higher value of a true pedigree of organisms. This pedigree is to disclose to us the genealogical connection of the smaller and larger groups. It has to show us in what way the different classes, orders, families, genera, and species of the animal and vegetable kingdoms correspond with the different branches, twigs, and groups of twigs of the pedigree. Every wider and higher category or stage of the system (for example a class, or an order) comprises a number of larger and stronger branches of the pedigree; every narrower and lower category (for example a genus, or a species) only a smaller and thinner group of twigs. It is only when we thus view the natural system as a pedigree that we perceive its true value. (Gen. Morph. ii. Plate XVII. p. 397.)

Since we hold fast this genealogical conception of the Organic System, to which alone undoubtedly the future of classificatory Zoology and Botany belongs, we should now turn our attention to one of the most essential, but also one of the most difficult, tasks of the “non-miraculous history of creation,” namely, to the actual construction of the Organic Pedigree. Let us see how far we are already able to point out all the different organic forms as the divergent descendants of a single or of some few common original forms. But how can we construct the actual pedigree of the animal and vegetable group of forms from our knowledge of them, at present so scanty and fragmentary? The answer to this question lies in what we have already remarked of the parallelism of the three series of development—in the important causal relation which connects the palæontological development of all organic tribes with the embryological development of individuals, and with the systematic development of groups.

In order to accomplish our task we shall first have to direct our attention to palæontology, or the science of petrifactions. For if the Theory of Descent is really true, if the petrified remains of formerly living animals and plants really proceed from the extinct primæval ancestors and progenitors of the present organisms, then, without anything else, the knowledge and comparison of petrifactions ought to disclose to us the pedigree of organisms. However simple and clear this may seem in theory, the task becomes extremely hard and complicated when it is actually taken in hand. Its practical solution would be very difficult even if the petrifactions were to any extent completely preserved. But this is by no means the case. The obvious records of creation which lie buried in petrifactions are imperfect beyond all measure. Hence it is necessary critically to examine these records, and to determine the value which petrifactions possess for the history of the development of organic tribes. As I have previously discussed the general importance of petrifactions as the records of creation, when we were considering Cuvier’s merits in the science of fossils, we may now at once examine the conditions and circumstances under which the remains of organic bodies became petrified and preserved in a more or less recognizable form.

As a rule we find petrifactions or fossils enclosed only in those stones which have been deposited in layers as mud by water, and which are on that account called neptunic, stratified, or sedimentary rocks. The deposition of such strata could of course only commence after the condensation of watery vapour into liquid water had taken place in the course of the earth’s history. After that period, which we considered in our last chapter, not only did life begin on the earth, but also an uninterrupted and exceedingly important transformation of the rigid inorganic crust of the earth. The water began that extremely important mechanical action by which the surface of the earth is perpetually, though slowly, transformed. I may surely presume that it is generally known what an extremely important influence, in this respect, is even yet exercised by water at every moment. As it falls down as rain, trickling through the upper strata of the earth’s crust, and flowing down from heights into hollows, it chemically dissolves different mineral parts of the ground, and mechanically washes away the loose particles. In flowing down from mountains water carries their debris into the plains, or deposits it as mud in stagnant lakes. Thus it continually works at lowering mountains and filling up valleys. In like manner the breakers of the sea work uninterruptedly at the destruction of the coasts and at filling up the bottom of the sea with the debris they wash down. The action of water alone, if it were not counteracted by other circumstances, would in time level the whole earth. There can be no doubt that the mountain masses—which are annually carried down as mud into the sea, and deposited on its floor—are so great that in the course of a longer or shorter period, say a few millions of years, the surface of the earth would be completely levelled and become enclosed by a continuous sheet of water. That this does not happen is owing to the perpetual volcanic action of the fiery-fluid centre of the earth. The surging of the melted nucleus against the firm crust necessitates continual alternations of elevation and depression on the different parts of the earth’s surface. These elevations and depressions for the most part take place very slowly; but, as they continue for thousands of years, by the combined effect of small, interrupted movements, they produce results no less grand than does the counteracting and levelling action of water.

Since the elevations and depressions of the different parts of the earth alternate with one another in the course of millions of years, first this and then that part of the earth’s surface is above or below the level of the sea. I have already given examples of this in the preceding chapter (vol. i. p. [361]). Hence, in all probability, there is no part of the outer crust of the earth which has not been repeatedly above and also below the level of the sea. This repeated change explains the variety and the different composition of the numerous neptunic strata of rocks, which in most places have been deposited one above another in considerable thickness. In the different periods of the earth’s history during which these deposits took place there lived various and different populations of animals and plants. When their dead bodies sank to the bottom of the waters, the forms of the bodies impressed themselves upon the soft mud, and imperishable parts, such as hard bones, teeth, shells, etc., became enclosed in it uninjured. These were preserved in the mud, which condensed them into neptunic rock, and as petrifactions they now serve to characterise the respective strata. By a careful comparison of the different strata lying one above another, and the petrifactions preserved in them, it has become possible to decide the relative age of the strata and groups of strata, and to establish, by direct observation, the principal eras of phylogeny, that is to say, the stages in history of the development of animal and vegetable tribes.

The different strata of neptunic rocks deposited one above another, which are composed in very various ways of limestone, clay, and sand, geologists have grouped together into an ideal System or Series, which corresponds with the whole course of the organic history of the earth, or with that portion of the earth’s history during which organic life existed. Just as so-called “universal history” falls into larger and smaller periods, which are characterized by the conditions of development of the most important nations at the respective epochs, and are separated from one another by great events, so we also divide the infinitely longer organic history of the earth into a series of greater and less periods. Each of these periods is distinguished by a characteristic flora and fauna, and by the specially strong development of certain vegetable or animal groups, and each is separated from the preceding and succeeding period by a striking change in the character of its animal and vegetable inhabitants.

In relation to the following survey of the historical course of development which the large animal and vegetable tribes have passed through, it will be desirable to say a few words first as to the systematic classification of the neptunic groups of strata, and the larger and smaller periods corresponding to them. As will be seen directly, we are able to divide the whole of the sedimentary rocks lying one above another into five main groups or periods, each period into several subordinate groups of strata or systems, and each system of strata again into still smaller groups or formations; finally, each formation can again be divided into stages or sub-formations, and each of these again into still smaller layers or beds. Each of the five great rock-groups was deposited during a great division of the earth’s history, during a long era or epoch; each system during a shorter period; each formation during a still shorter period. In thus reducing the periods of the organic history of the earth, and the neptunic strata containing petrifactions deposited during those periods into a connected system, we proceed exactly like the historian who divides the history of nations into the three main divisions of Antiquity, the Middle Ages, and Modern Times, and each of those sections again into subordinate periods and epochs. But the historian by this sharp systematic division, and by fixing the boundary of the periods by particular dates, only seeks to facilitate his survey, and in no way means to deny the uninterrupted connection of events and the development of nations. Exactly the same qualification applies to our systematic division, specification, or classification of the organic history of the earth. Here, too, a continuous thread runs through the series of events unbroken. We must therefore distinctly protest against the idea that by sharply bounding the larger and smaller groups of strata, and the the periods corresponding with them, we in any way wish to adopt Cuvier’s doctrine of terrestrial revolutions, and of repeated new creations of organic populations. That this erroneous doctrine has long since been completely refuted by Lyell, I have already mentioned. (Compare vol. i. p. [127.])

The five great main divisions of the organic history of the earth, or the palæontological history of development, we call the primordial, primary, secondary, tertiary, and quaternary epochs. Each is distinctly characterized by the predominating development of certain animal and vegetable groups in it, and we might accordingly symbolically designate the five epochs, on the one hand by the names of the groups of the vegetable kingdom, and on the other hand by those of the different classes of vertebrate animals. In this case the first, or primordial epoch, would be the era of the Tangles (Algæ) and skull-less Vertebrates; the second, or primary epoch, that of the Ferns and Fishes; the third, or secondary epoch, that of Pine Forests and Reptiles; the fourth, or tertiary epoch, that of Foliaceous Forests and of Mammals; finally, the fifth, or quaternary epoch, the era of Man, and his Civilization. The divisions or periods which we distinguish in each of the five long eras (p. [14]) are determined by the different systems of strata into which each of the five great rock-groups is divided (p. [15]). We shall now take a cursory glance at the series of these systems, and at the same time at the populations of the five great epochs.

The first and longest division of the organic history of the earth is formed by the primordial epoch, or the era of the Tangle Forests. It comprises the immense period from the first spontaneous generation, from the origin of the first terrestrial organism, to the end of the Silurian system of deposits. During this immeasurable space of time, which in all probability was much longer than all the other four epochs taken together, the three most extensive of all the neptunic systems of strata were deposited, namely, the Laurentian, upon that the Cambrian, and upon that the Silurian system. The approximate thickness or size of these three systems together amounts to 70,000 feet. Of these about 30,000 belong to the Laurentian, 18,000 to the Cambrian, and 22,000 to the Silurian system. The average thickness of all the four other rock groups, the primary, secondary, tertiary, and quaternary, taken together, may amount at most to 60,000 feet; and from this fact alone, apart from many other reasons, it is evident that the duration of the primordial period was probably much longer than the duration of all the subsequent periods down to the present day. Many thousands of millions of years were required to deposit such masses of strata. Unfortunately, by far the largest portion of the primordial group of strata is in the metamorphic state (which we shall directly explain), and consequently the petrifactions contained in them—the most ancient and most important of all—have, to a great extent, been destroyed and become unrecognisable. Only in one portion of the Cambrian strata have petrifactions been preserved in a recognizable condition and in large quantities. The most ancient of all distinctly preserved petrifactions has been found in the lowest Laurentian strata (in the Ottawa formation), which I shall afterwards have to speak of as the “Canadian Life’s-dawn” (Eozoon canadense).

Although only by far the smaller portion of the primordial or archilithic petrifactions are preserved to us in a recognizable condition, still they possess the value of inestimable documents of the most ancient and obscure times of the organic history of the earth. What seems to be shown by them, in the first place, is that during the whole of this immense period there existed only inhabitants of the waters. As yet, at any rate, among all archilithic petrifactions, not a single one has been found which can with certainty be regarded as an organism which has lived on land. All the vegetable remains we possess of the primordial period belong to the lowest of all groups of plants, to the class of Tangles or Algæ, living in water. In the warm primæval sea, these constituted the forests of the period, of the richness of which in forms and density we may form an approximate idea from their present descendants, the tangle forests of the Atlantic Sargasso sea. The colossal tangle forests of the archilithic period supplied the place of the forest vegetation of the mainland, which was then utterly wanting. All the animals, also, whose remains have been found in archilithic strata, like the plants, lived in water. Only crustacea are met with among the animals with articulated feet, as yet no spiders and no insects. Of vertebrate animals, only a very few remains of fishes are known as having been found in the most recent of all primordial strata, in the upper Silurian. But the headless vertebrate animals, which we call skull-less, or Acrania, and out of which fishes must have been developed, we suppose to have lived in great numbers during the primordial epoch. Hence we may call it after the Acrania as well as after the Tangles.

The primary epoch, or the era of Fern Forests, the second main division of the organic history of the earth, which is also called the palæolithic or palæozoic period, lasted from the end of the Silurian formation of strata to the end of the Permian formation. This epoch was also of very long duration, and again falls into three shorter periods, during which three great systems of strata were deposited, namely, first, the Devonian system, or the old red sandstone; upon that, the Carboniferous, or coal system; and upon this, the Permian system. The average thickness of these three systems taken together may amount to about 42,000 feet, from which we may infer the immense length of time requisite for their formation.

The Devonian and Permian formations are especially rich in remains of fishes, of primæval fish as well as enamelled fish (Ganoids), but the bony fish (Teleostei) are absent from the strata of the primary epoch. In coal are found the most ancient remains of animals living on land, both of articulated animals (spiders and insects) as well as of vertebrate animals (amphibious animals, like newts and frogs). In the Permian system there occur, in addition to the amphibious animals, the more highly-developed reptiles, and, indeed, forms nearly related to our lizards (Proterosaurus, etc.). But, nevertheless, we may call the primary epoch that of Fishes, because these few amphibious animals and reptiles are insignificant in comparison with the immense mass of palæozoic fishes. Just as Fishes predominate over the other vertebrate animals, so Ferns, or Filices, predominate among the plants of this epoch, and, in fact, real ferns and tree ferns (leafed ferns, or Phylopteridæ), as well as bamboo ferns (Calamophytæ) and scaled ferns (Lepidophytæ). These ferns, which grew on land, formed the chief part of the dense palæolithic island forests, the fossil remains of which are preserved to us in the enormously large strata of coal of the Carboniferous system, and in the smaller strata of coal of the Devonian and Permian systems. We are thus justified in calling the primary epoch either the era of Ferns or that of Fishes.

The third great division of the palæontological history of development is formed by the secondary epoch, or the era of Pine Forests, which is also called the mesolithic or mesozoic epoch. It extends from the end of the Permian system to the end of the Chalk formation, and is again divided into three great periods. The stratified systems deposited during this period are, first and lowest, the Triassic system, in the middle the Jura system, and at the top the Cretaceous system. The average thickness of these three systems taken together is much less than that of the primary group, and amounts as a whole only to about 15,000 feet. The secondary epoch can accordingly in all probability not have been half so long as the primary epoch.

Just as Fishes prevailed in the primary epoch, Reptiles predominated in the secondary epoch over all other vertebrate animals. It is true that during this period the first birds and mammals originated; at that time, also, there existed important amphibious animals, especially the gigantic Labyrinthodonts, in the sea the wonderful sea-dragons, or Halisaurii, swam about, and the first fish with bones were associated with the many primæval fishes (Sharks) and enamelled fish (Ganoids) of the earlier times; but the very variously developed kinds of reptiles formed the predominating and characteristic class of vertebrate animals of the secondary epoch. Besides those reptiles which were very nearly related to the present living lizards, crocodiles, and turtles, there were, during the mesolithic period, swarms of grotesquely shaped dragons. The remarkable flying lizards, or Pterosaurii, and the colossal land-dragons, or Dinosaurii, of the secondary epoch, are peculiar, as they occur neither in the preceding nor in the succeeding epochs. The secondary epoch may be called the era of Reptiles; but on the other hand, it may also be called the era of Pine Forests, or more accurately, of the Gymnosperms, that is, the epoch of plants having naked seeds. For this group of plants, especially as represented by the two important classes—the pines, or Coniferæ, and the palm-ferns, or Cycadeæ—during the secondary epoch constituted a predominant part of the forests. But towards the end of the epoch (in the Chalk period) the plants of the pine tribe gave place to the leaf-bearing forests which then developed for the first time.

STRATA CONTAINING PETRIFICATIONS.

The fourth main division of the organic history of the earth, the tertiary epoch, or era of Leafed Forests, is much shorter and less peculiar than the three first epochs. This epoch, which is also called the cænolithic or cænozoic epoch, extended from the end of the cretaceous system to the end of the pliocene system. The strata deposited during it amount only to a thickness of about 3,000 feet, and consequently are much inferior to the three first great groups. The three systems also into which the tertiary period is subdivided are very difficult to distinguish from one another. The oldest of them is called eocene, or old tertiary; the newer miocene, or mid tertiary; and the last is the pliocene, or later tertiary system.

The whole population of the tertiary epoch approaches much nearer, on the whole as well as in detail, to that of the present time than is the case in the preceding epochs. From this time the class of Mammals greatly predominates over all other vertebrate animals. In like manner, in the vegetable kingdom, the group—so rich in forms—of the Angiosperms, or plants with covered seeds, predominates, and its leafy forests constitute the characteristic feature of the tertiary epoch. The group of the Angiosperms consists of the two classes of single-seed-lobed plants, or Monocotyledons, and the double-seed-lobed plants, or Dicotyledons. The Angiosperms of both classes had, it is true, made their appearance in the Cretaceous period, and mammals had already occurred in the Jurassic period, and even in the Triassic period; but both groups, the mammals and the plants with enclosed seeds, did not attain their peculiar development and supremacy until the tertiary epoch, so that it may justly be called after them.

The fifth and last main division of the organic history of the earth is the quaternary epoch, or era of Civilization, which in comparison with the length of the four other epochs almost vanishes into nothing, though with a comical conceit we usually call its record the “history of the world.” As the period is characterized by the development of Man and his Culture, which has influenced the organic world more powerfully and with greater transforming effect than have all previous conditions, it may also be called the era of Man, the anthropolithic or anthropozoic period. It might also be called the era of Cultivated Forests, or Gardens, because even at the lowest stage of human civilization man’s influence is already perceptible in the utilization of forests and their products, and therefore also in the physiognomy of the landscape. The commencement of this era, which extends down to the present time, is geologically bounded by the end of the pliocene stratification.

The neptunic strata which have been deposited during the comparatively short quaternary epoch are very different in different parts of the earth, but they are mostly of very slight thickness. They are reduced to two “systems,” the older of which is designated the diluvial, or pleistocene, and the later the alluvial, or recent. The diluvial system is again divided into two “formations,” the older glacial and the more recent post glacial formations. For during the older diluvial period there occurred that extremely remarkable decrease of the temperature of the earth which led to an extensive glaciation of the temperate zones. The great importance which this “ice” or “glacial period” has exercised on the geographical and topographical distribution of organisms has already been explained in the preceding chapter (vol. i. p. [365]). But the post glacial period, or the more recent diluvial period, during which the temperature again increased and the ice retreated towards the poles, was also highly important in regard to the present state of chorological relations.

The biological characteristic of the quaternary epoch lies essentially in the development and dispersion of the human organism and his culture. Man has acted with a greater transforming, destructive, and modifying influence upon the animal and vegetable population of the earth than any other organism. For this reason, and not because we assign to man a privileged exceptional position in nature in other matters, we may with full justice designate the development of man and his civilization as the beginning of a special and last main division of the organic history of the earth. It is probable indeed that the corporeal development of primæval man out of man-like apes took place as far back as the earlier pliocene period, perhaps even in the miocene tertiary period. But the actual development of human speech, which we look upon as the most powerful agency in the development of the peculiar characteristics of man and his dominion over other organisms, probably belongs to that period which on geological grounds is distinguished from the preceding pliocene period as the pleistocene or diluvial. In fact the time which has elapsed from the development of human speech down to the present day, though it may comprise many thousands and perhaps hundreds of thousands of years, almost vanishes into nothing as compared with the immeasurable length of the periods which have passed from the beginning of organic life on the earth down to the origin of the human race.

The tabular view given on page 15 shows the succession of the palæontological rock-groups, systems, and formations, that is, the larger and smaller neptunic groups of strata, which contain petrifactions, from the uppermost, or Alluvial, down to the lowest, or Laurentian, deposits. The table on page 14 presents the historical division of the corresponding eras of the larger and smaller palæontological periods, and in a reversed succession, from the most ancient Laurentian up to the most recent Quaternary period.

Many attempts have been made to make an approximate calculation of the number of thousands of years constituting these periods. The thickness of the strata has been compared, which, according to experience, is deposited during a century, and which amounts only to some few lines or inches, with the whole thickness of the stratified masses of rock, the succession of which we have just surveyed. This thickness, on the whole, may on an average amount to about 130,000 feet; of these 70,000 belong to the primordial, or archilithic; 42,000 to the primary, or palæolithic; 15,000 to the secondary, or mesolithic; and finally only 3,000 to the tertiary, or cænolithic group. The very small and scarcely appreciable thickness of the quaternary, or anthropolithic deposit cannot here come into consideration at all. On an average, it may at most be computed as from 500 to 700 feet. But it is self evident that all these measurements have only an average and approximate value, and are meant to give only a rough survey of the relative proportion of the systems of strata and of the spaces of time corresponding with them.

Now, if we divide the whole period of the organic history of the earth—that is, from the beginning of life on the earth down to the present day—into a hundred equal parts, and if then, corresponding to the thickness of the systems of strata, we calculate the relative duration of the time of the five main divisions or periods according to percentages, we obtain the following result:—

According to this, the length of the archilithic period, during which no land-living animals or plants as yet existed, amounts to more than one half, more than 53 per cent.; on the other hand the length of the anthropolithic era, during which man has existed, amounts to scarcely one-half per cent. of the whole length of the organic history of the earth. It is, however, quite impossible to calculate the length of these periods, even approximately, by years.

The thickness of the strata of mud at present deposited during a century, and which has been used as a basis for this calculation, is of course quite different in different parts of the earth under the different conditions in which these deposits take place. It is very slight at the bottom of the deep sea, in the beds of broad rivers with a short course, and in inland seas which receive very scanty supplies of water. It is comparatively great on the sea-shores exposed to strong breakers, at the estuaries of large rivers with long courses, and in inland seas with copious supplies of water. At the mouth of the Mississippi, which carries with it a considerable amount of mud, in the course of 100,000 years about 600 feet would be deposited. At the bottom of the open sea, far away from the coasts, during this long period only some few feet of mud would be deposited. Even on the sea-shores where a comparatively large quantity of mud is deposited the thickness of the strata formed during the course of a century may after all amount to no more than a few inches or lines when condensed into solid stone. In any case, however, all calculations based upon these comparisons are very unsafe, and we cannot even approximately conceive the enormous length of the periods which were requisite for the formation of the systems of neptunic strata. Here we can apply only relative, not absolute, measurements of time.

Moreover, we should entirely err were we to consider the size of these systems of strata alone as the measure of the actual space of time which has elapsed during the earth’s history. For the elevations and depressions of the earth’s crust have perpetually alternated with one another, and the mineralogical and palæontological difference—which is perceived between each two succeeding systems of strata, and between each two of their formations at any particular spot—corresponds in all probability with a considerable intermediate space of many thousands of years, during which that particular part of the earth’s crust was raised above the water. It was only after the lapse of this intermediate period, when a new depression again laid the part in question under water, that there occurred a new deposit of earth. As, in the mean time, the inorganic and organic conditions on this part had undergone a considerable transformation, the newly-formed layer of mud was necessarily composed of different earthy constituents and enclosed different petrifactions.

The striking differences which so frequently occur between the petrifactions of two strata, lying one above another, are to be explained in a simple and easy manner by the supposition that the same part of the earth’s surface has been exposed to repeated depressions and elevations. Such alternating elevations and depressions take place even now extensively, and are ascribed to the heaving of the fiery fluid nucleus against the rigid crust. Thus, for example, the coast of Sweden and a portion of the west coast of South America are constantly though slowly rising, while the coast of Holland and a portion of the east coast of South America are gradually sinking. The rising as well as the sinking takes place very slowly, and in the course of a century sometimes only amounts to some few lines, sometimes to a few inches, or at most a few feet. But if this action continues uninterruptedly throughout hundreds of thousands of years it is capable of forming the highest mountains.

It is evident that elevations and depressions, such as now can be measured in these places, have uninterruptedly alternated one with another in different places during the whole course of the organic history of the earth. This may be inferred with certainty from the geographical distribution of organisms. (Compare vol. i. p. [350.]) But to form a judgment of our palæontological records of creation it is extremely important to show that permanent strata can only be deposited during a slow sinking of the ground under water, but not during its continued rising. When the ground slowly sinks more and more below the level of the sea, the deposited layers of mud get into continually deeper and quieter water, where they can become condensed into stone undisturbed. But when, on the other hand, the ground slowly rises, the newly-deposited layers of mud, which enclose the remains of plants and animals, again immediately come within the reach of the play of the waves, and are soon worn away by the force of the breakers, together with the organic remains which they on close. For this simple but very important reason, therefore, abundant layers, in which organic remains are preserved, can only be deposited during a continuous sinking of the ground. When any two different formations or strata, lying one above the other, correspond with two different periods of depression, we must assume a long period of rising between them, of which period we know nothing, because no fossil remains of the then living animals and plants could be preserved. It is evident, however, that those periods of elevation, which have passed without leaving any trace behind them, deserve a no less careful consideration than the greater or less alternating periods of depression, of whose organic population we can form an approximate idea from the strata containing petrifactions. Probably the former were not of shorter duration than the latter.

From this alone it is apparent how imperfect our records must necessarily be, and all the more so since it can be theoretically proved that the variety of animal and vegetable life must have increased greatly during those very periods of elevation. For as new tracts of land are raised above the water, new islands are formed. Every new island, however, is a new centre of creation, because the animals and plants accidentally cast ashore there, find in the new territory, in the struggle for life, abundant opportunity of developing themselves peculiarly, and of forming new species. The formation of new species has evidently taken place pre-eminently during these intermediate periods, of which, unfortunately, no petrifactions could be preserved, whereas, on the contrary, during the slow sinking of the ground there was more chance of numerous species dying out, and of a retrogression into fewer specific forms. The intermediate forms between the old and the newly-forming species must also have lived during the periods of elevation, and consequently could likewise leave no fossil remains.

In addition to the great and deplorable gaps in the palæontological records of creation—which are caused by the periods of elevation—there are, unfortunately, many other circumstances which immensely diminish their value. I must mention here especially the metamorphic state of the most ancient formations, of those strata which contain the remains of the most ancient flora and fauna, the original forms of all subsequent organisms, and which, therefore, would be of especial interest. It is just these rocks—and, indeed, the greater part of the primordial, or archilithic strata, almost the whole of the Laurentian, and a large part of the Cambrian systems—which no longer contain any recognizable remains, and for the simple reason that these strata have been subsequently changed or metamorphosed by the influence of the fiery fluid interior of the earth. These deepest neptunic strata of the crust have been completely changed from their original condition by the heat of the glowing nucleus of the earth, and have assumed a crystalline state. In this process, however, the form of the organic remains enclosed in them has been entirely destroyed. It has been preserved only here and there by a happy chance, as in the case of the most ancient petrifactions known, the Eozoon canadense, from the lowest Laurentian strata. However, from the layers of crystalline charcoal (graphite) and crystalline limestone (marble), which are found deposited in the metamorphic rocks, we may with certainty conclude that petrified animal and vegetable remains existed in them in earlier times.

Our record of creation is also extremely imperfect from the circumstance that only a small portion of the earth’s surface has been accurately investigated by geologists, namely, England, Germany, and France. But we know very little of the other parts of Europe, of Russia, Spain, Italy, and Turkey. In the whole of Europe, only some few parts of the earth’s crust have been laid open, by far the largest portion of it is unknown to us. The same applies to North America and to the East Indies. There some few tracts have been investigated; but of the larger portion of Asia, the most extensive of all continents, we know almost nothing; of Africa nothing, excepting the Cape of Good Hope and the shores of the Mediterranean; of Australia almost nothing; and of South America but very little. It is clear, therefore, that only quite a small portion, perhaps scarcely the thousandth part of the whole surface of the earth, has been palæontologically investigated. We may therefore reasonably hope, when more extensive geological investigations are made, which are greatly assisted by the constructions of railroads and mines, to find a great number of other important petrifactions. A hint that this will be the case is given by the remarkable petrifactions found in those parts of Africa and Asia which have been minutely investigated,—the Cape districts and the Himalaya mountains. A series of entirely new and very peculiar animal forms have become known to us from the rocks of these localities. But we must bear in mind that the vast bottom of the existing oceans is at the present time quite inaccessible to palæontological investigations, and that the greater part of the petrifactions which have lain there from primæval times will either never be known to us, or at best only after the course of many thousands of years, when the present bottom of the ocean shall have become accessible by gradual elevation. If we call to mind the fact that three-fifths of the whole surface of the earth consists of water, and only two-fifths of land, it becomes plain that on this account the palæontological record must always present an immense gap.

But, in addition to these, there exists another series of difficulties in the way of palæontology which arises from the nature of the organisms themselves. In the first place, as a rule only the hard and solid parts of organisms can fall to the bottom of the sea or of fresh waters, and be there enclosed in the mud and petrified. Hence it is only the bones and teeth of vertebrate animals, the calcareous shells of molluscs, the chitinous skeletons of articulated animals, the calcareous skeletons of star-fishes and corals, and the woody and solid parts of plants, that are capable of being petrified. But soft and delicate parts, which constitute by far the greater portion of the bodies of most organisms, are very rarely deposited in the mud under circumstances favourable to their becoming petrified, or distinctly impressing their external form upon the hardening mud. Now, it must be borne in mind that large classes of organisms, as for example the Medusæ, the naked molluscs without shells, a large portion of the articulated animals, almost all worms, and even the lowest vertebrate animals, possess no firm and hard parts capable of being petrified. In like manner the most important parts of plants, such as the flowers, are for the most part so soft and tender that they cannot be preserved in a recognizable form. We therefore cannot expect to find any petrified remains of these important organisms. Moreover, all organisms at an early stage of life are so soft and tender that they are quite incapable of being petrified. Consequently all the petrifactions found in the neptunic stratifications of the earth’s crust comprise altogether but a very few forms, and of these for the most part only isolated fragments.

We must next bear in mind that the dead bodies of the inhabitants of the sea are much more likely to be preserved and petrified in the deposits of mud than those of the inhabitants of fresh water and of the land. Organisms living on land can, as a rule, become petrified only when their corpses fall accidentally into the water and are buried at the bottom in the hardening layers of mud. But this event depends upon very many conditions. We cannot therefore be astonished that by far the majority of petrifactions belong to organisms which have lived in the sea, and that of the inhabitants of the land proportionately only very few are preserved in a fossil state. How many contingencies come into play here we may infer from the single fact that of many fossil mammals, in fact of all the mammals of the secondary, or mesozoic epoch, nothing is known except the lower jawbone. This bone is in the first place comparatively solid, and in the second place very easily separates itself from the dead body, which floats on the water. Whilst the body is driven away and dissolved by the water, the lower jawbone falls down to the bottom of the water and is there enclosed in the mud. This explains the remarkable fact that in a stratum of limestone of the Jurassic system near Oxford, in the slates of Stonesfield, as yet only the lower jawbones of numerous pouched animals (Marsupials) have been found. They are the most ancient mammals known, and of the whole of the rest of their bodies not a single bone exists. The opponents of the theory of development, according to their usual logic, would from this fact be obliged to draw the conclusion that the lower jawbone was the only bone in the body of those animals.

Footprints are very instructive when we attempt to estimate the many accidents which so arbitrarily influence our knowledge of fossils; they are found in great numbers in different extensive layers of sandstone; for example, in the red sandstone of Connecticut, in North America. These footprints were evidently made by vertebrate animals, probably by reptiles, of whose bodies not the slightest trace has been preserved.[1] The impressions which their feet have left on the mud alone betray the former existence of these otherwise unknown animals.

The accidents which, besides these, determine the limits of our palæontological knowledge, may be inferred from the fact that we know of only one or two specimens of very many important petrifactions. It is not ten years since we became acquainted with the imperfect impression of a bird in the Jurassic or Oolitic system, the knowledge of which has been of the very greatest importance for the phylogeny of the whole class of birds. All birds previously known presented a very uniformly organized group, and showed no striking transitional forms to other vertebrate classes, not even to the nearly related reptiles. But that fossil bird from the Jura possessed not an ordinary bird’s tail, but a lizard’s tail, and thus confirmed what had been conjectured upon other grounds, namely, the derivation of birds from lizards. This single fossil has thus essentially extended not only our knowledge of the age of the class of birds, but also of their blood relationship to reptiles. In like manner our knowledge of other animal groups has been often essentially modified by the accidental discovery of a single fossil. The palæontological records must necessarily be exceedingly imperfect, because we know of so very few examples, or only mere fragments of very many important fossils.

Another and very sensible gap in these records is caused by the circumstance that the intermediate forms which connect the different species have, as a rule, not been preserved, and for the simple reason that (according to the principle of divergence of character) they were less favoured in the struggle for life than the most divergent varieties, which had developed out of one and the same original form. The intermediate links have, on the whole, always died out rapidly, and have but rarely been preserved as fossils. On the other hand, the most divergent forms were able to maintain themselves in life for a longer period as independent species, to propagate more numerously, and consequently to be more readily petrified. But this does not exclude the fact that in some cases the connecting intermediate forms of the species have been preserved so perfectly petrified, that even now they cause the greatest perplexity and occasion endless disputes among systematic palæontologists about the arbitrary limits of species.

An excellent example of this is furnished by the celebrated and very variable fresh-water snail from the Stuben Valley, near Steinheim, in Würtemburg, which has been described sometimes as Paludina, sometimes as Valvata, and sometimes as Planorbis multiformis. The snow-white shells of these small snails constitute more than half of the mass of the tertiary limestone hills, and in this one locality show such an astonishing variety of forms, that the most divergent extremes might be referred to at least twenty entirely different species. But all these extreme forms are united by such innumerable intermediate forms, and they lie so regularly above and beside one another, that Hilgendorf was able, in the clearest manner, to unravel the pedigree of the whole group of forms. In like manner, among very many other fossil species (for example, many ammonites, terebratulæ, sea urchins, lily encrinites, etc.) there are such masses of connecting intermediate forms, that they reduce the “dealers in fossil species” to despair.

When we weigh all the circumstances here mentioned, the number of which might easily be increased, it does not appear astonishing that the natural accounts or records of creation formed by petrifactions are extremely defective and incomplete. But nevertheless, the petrifactions actually discovered are of the greatest value. Their significance is of no less importance to the natural history of creation than the celebrated inscription on the Rosetta stone, and the decree of Canopus, are to the history of nations—to archæology and philology. Just as it has become possible by means of these two most ancient inscriptions to reconstruct the history of ancient Egypt, and to decipher all hieroglyphic writings, so in many cases a few bones of an animal, or imperfect impressions of a lower animal or vegetable form, are sufficient for us to gain the most important starting-points in the history of the whole group, and in the search after their pedigree. A couple of small back teeth, which have been found in the Keuper formation of the Trias, have of themselves alone furnished a sure proof that mammals existed even in the Triassic period.

Of the incompleteness of the geological accounts of creation, Darwin, agreeing with Lyell, the greatest of all recent geologists, says:—

“I look at the geological record as a history of the world imperfectly kept, and written in a changing dialect; of this history we possess the last volume alone, relating only to two or three countries. Of this volume, only here and there a short chapter has been preserved; and of each page, only here and there a few lines. Each word of the slowly-changing language, more or less different in the successive chapters, may represent the forms of life which are entombed in our consecutive formations, and which falsely appear to us to have been abruptly introduced. On this view, the difficulties above discussed are greatly diminished, or even disappear.”—Origin of Species, 6th Edition, p. 289.

If we bear in mind the exceeding incompleteness of palæontological records, we shall not be surprised that we are still dependent upon so many uncertain hypotheses when actually endeavouring to sketch the pedigree of the different organic groups. However, we fortunately possess, besides fossils, other records of the history of the origin of organisms, which in many cases are of no less value, nay, in several cases are of much greater value, than fossils. By far the most important of these other records of creation is, without doubt, ontogeny, that is, the history of the development of the organic individual (embryology and metamorphology). It briefly repeats in great and marked features the series of forms which the ancestors of the respective individuals have passed through from the beginning of their tribe. We have designated the palæontological history of the development of the ancestors of a living form as the history of a tribe, or phylogeny, and we may therefore thus enunciate this exceedingly important biogenetic fundamental principle: “Ontogeny is a short and quick repetition, or recapitulation, of Phylogeny, determined by the laws of Inheritance and Adaptation.” As every animal and every plant from the beginning of its individual existence passes through a series of different forms, it indicates in rapid succession and in general outlines the long and slowly changing series of states of form which its progenitors have passed through from the most ancient times. (Gen. Morph. ii. 6, 110, 300.)

It is true that the sketch which the ontogeny of organisms gives us of their phylogeny is in most cases more or less obscured, and all the more so the more Adaptation, in the course of time, has predominated over Inheritance, and the more powerfully the law of abbreviated inheritance, and the law of correlative adaptation, have exerted their influence. However, this does not lessen the great value which the actual and faithfully preserved features of that sketch possess. Ontogeny is of the most inestimable value for the knowledge of the earliest palæontological conditions of development, just because no petrified remains of the most ancient conditions of the development of tribes and classes have been preserved. These, indeed, could not have been preserved on account of the soft and tender nature of their bodies. No petrifactions could inform us of the fundamental and important fact which ontogeny reveals to us, that the most ancient common ancestors of all the different animal and vegetable species were quite simple cells like the egg-cell. No petrifaction could prove to us the immensely important fact, established by ontogeny, that the simple increase, the formation of cell-aggregates and the differentiation of those cells, produced the infinitely manifold forms of multicellular organisms. Thus ontogeny helps us over many and large gaps in palæontology.

1. Man, 2. Gorilla, 3. Orang, 4. Dog, 5. Seal, 6. Porpoise, 7. Bat, 8. Mole, 9. Duck-bill.

To the invaluable records of creation furnished by palæontology and ontogeny are added the no less important evidences for the blood relationship of organisms furnished by comparative anatomy. When organisms, externally very different, nearly agree in their internal structure, one may with certainty conclude that the agreement has its foundation in Inheritance, the dissimilarity its foundation in Adaptation. Compare, for example, the hands and fore paws of the nine different animals which are represented on Plate [IV]., in which the bony skeleton in the interior of the hand and of the five fingers is visible. Everywhere we find, though the external forms are most different, the same bones, and among them the same number, position, and connection. It will perhaps appear very natural that the hand of man (Fig. 1) differs very little from that of the gorilla (Fig. 2) and of the orang-outang (Fig. 3), his nearest relations. But it will be more surprising if the fore feet of the dog also (Fig. 4), as well as the breast-fin (the hand) of the seal (Fig. 5), and of the dolphin (Fig. 6), show essentially the same structure. And it will appear still more wonderful that even the wing of the bat (Fig. 7), the shovel-feet of the mole (Fig. 8), and the fore feet of the duck-bill (Ornithorhynchus) (Fig. 9), the most imperfect of all mammals, is composed of entirely the same bones, only their size and form being variously changed. Their number, the manner of their arrangement and connection has remained the same. (Compare also the explanation of Plate [IV]., in the Appendix.) It is quite inconceivable that any other cause, except the common inheritance of the part in question from common ancestors, could have occasioned this wonderful homology or similarity in the essential inner structure with such different external forms. Now, if we go down further in the system below the mammals, and find that even the wings of birds, the fore feet of reptiles and amphibious animals, are composed of essentially the same bones as the arms of man and the fore legs of the other mammals, we can, from this circumstance alone, with perfect certainty, infer the common origin of all these vertebrate animals. Here, as in all other cases, the degree of the internal agreement in the form discloses to us the degree of blood relationship.

CHAPTER XVI.

PEDIGREE AND HISTORY OF THE KINGDOM OF THE PROTISTA.

Special Mode of Carrying out the Theory of Descent in the Natural System of Organisms.—Construction of Pedigrees.—Descent of all Many-Celled from Single-Celled Organisms.—Descent of Cells from Monera.—Meaning of Organic Tribes, or Phyla.—Number of the Tribes in the Animal and Vegetable Kingdoms.—The Monophyletic Hypothesis of Descent, or the Hypothesis of one Common Progenitor, and the Polyphyletic Hypothesis of Descent, or the Hypothesis of Many Progenitors.—The Kingdom of Protista, or Primæval Beings.—Eight Classes of the Protista Kingdom.—Monera, Amœbæ, or Protoplastæ.—Whip-swimmers, or Flagellata.—Ciliated-balls, or Catallacta.—Labyrinth-streamers, or Labyrinthuleæ.—Flint-cells, or Diatomeæ.—Mucous-moulds, or Myxomycetes.—Root-footers (Rhizopoda).—Remarks on the General Natural History of the Protista: Their Vital Phenomena, Chemical Composition, and Formation (Individuality and Fundamental Form).—Phylogeny of the Protista Kingdom

By a careful comparison of the individual and the palæontological development, as also by the comparative anatomy of organisms, by the comparative examination of their fully developed structural characteristics, we arrive at the knowledge of the degrees of their different structural relationships. By this, however, we at the same time obtain an insight into their true blood relationship, which, according to the Theory of Descent, is the real reason of the structural relationship. Hence by collecting, comparing, and employing the empirical results of embryology, palæontology, and anatomy for supplementing each other, we arrive at an approximate knowledge of “the Natural System,” which, according to our views, is the pedigree of organisms. It is true that our human knowledge, in all things fragmentary, is especially so in this case, on account of the extreme incompleteness and defectiveness of the records of creation. However, we must not allow this to discourage us, or to deter us from undertaking this highest problem of biology. Let us rather see how far it may even now be possible, in spite of the imperfect state of our embryological, palæontological, and anatomical knowledge, to establish a probable scheme of the genealogical relationships of organisms.

Darwin in his book gives us no answer to these special questions of the Theory of Descent; at the conclusion he only expresses his conjecture “that animals have descended from at most only four or five progenitors, and plants from an equal or less number.” But as these few aboriginal forms still show traces of relationship, and as the animal and vegetable kingdoms are connected by intermediate transitional forms, he arrives afterwards at the opinion “that probably all the organic beings which have ever lived on the earth have descended from some one primordial form, into which life was first breathed by the Creator.” Like Darwin, all other adherents of the Theory of Descent have only treated it in a general way, and not made the attempt to carry it out specially, and to treat the “Natural System” actually as the pedigree of organisms. If, therefore, we venture upon this difficult undertaking, we must take up independent ground.

Four years ago I set up a number of hypothetical genealogies for the larger groups of organisms in the systematic introduction to my General History of Development (Gen. Morph. vol. ii.), and thereby, in fact, made the first attempt actually to construct the pedigrees of organisms in the manner required by the theory of development. I was quite conscious of the extreme difficulty of the task, and as I undertook it in spite of all discouraging obstacles, I claim no more than the merit of having made the first attempt and given a stimulus for other and better attempts. Probably most zoologists and botanists were but little satisfied with this beginning, and least so in reference to the special domain in which each one is specially at work. However, it is certainly in this case much easier to blame than to produce something better, and what best proves the immense difficulty of this infinitely complicated task is the fact that no naturalist has as yet supplied the place of my pedigrees by better ones. But, like all other scientific hypotheses which serve to explain facts, my genealogical hypotheses may claim to be taken into consideration until they are replaced by better ones.

I hope that this replacement will very soon take place; and I wish for nothing more than that my first attempt may induce very many naturalists to establish more accurate pedigrees for the individual groups, at least in the special domain of the animal and vegetable kingdom which happens to be well known to one or other of them. By numerous attempts of this kind our genealogical knowledge, in the course of time, will slowly advance and approach more towards perfection, although it can with certainty be foreseen that we shall never arrive at a complete pedigree. We lack, and shall ever lack, the indispensable palæontological foundations. The most ancient records will ever remain sealed to us, for reasons which have been previously mentioned. The most ancient organisms which arose by spontaneous generation—the original parents of all subsequent organisms—must necessarily be supposed to have been Monera—simple, soft, albuminous lumps, without structure, without any definite forms, and entirely without any hard and formed parts. They and their next offspring were consequently not in any way capable of being preserved in a petrified condition. But we also lack, for reasons discussed in detail in the preceding chapter, by far the greater portion of the innumerable palæontological documents, which are really requisite for a safe reconstruction of the history of animal tribes, or phylogeny, and for the true knowledge of the pedigree of organisms. If we, therefore, in spite of this, venture to undertake their hypothetical construction, we must chiefly depend for guidance on the two other series of records which most essentially supplement the palæontological archives. These are ontogeny and comparative anatomy.

If thoughtfully and carefully we consult these most valuable records, we at once perceive what is exceedingly significant, namely, that by far the greater number of organisms, especially all higher animals and plants, are composed of a great number of cells, and that they originate out of an egg, and that this egg, in animals as well as in plants, is a single, perfectly simple cell—a little lump of albuminous constitution, in which another albuminous corpuscle, the cell-kernel, is enclosed. This cell containing its kernel grows and becomes enlarged. By division it forms an accumulation of cells, and out of these, by division of labour (as has previously been described), there arise the numberless different forms which are presented to us in the fully developed animal and vegetable species. This immensely important process—which we may follow step by step, with our own eyes, any day in the embryological development of any animal or vegetable individual, and which as a rule is by no means considered with the reverence it deserves—informs us more surely and completely than all petrifactions could do as to the original palæontological development of all many-celled organisms, that is, of all higher animals and plants. For as ontogeny, or the embryological development of every single individual, is essentially only a recapitulation of phylogeny, or the palæontological development of its chain of ancestors, we may at once, with full assurance, draw the simple and important conclusion, that all many-celled animals and plants were originally derived from single-celled organisms. The primæval ancestors of man, as well as of all other animals, and of all plants composed of many cells, were simple cells living isolated. This invaluable secret of the organic pedigree is revealed to us with infallible certainty by the egg of animals, and by the true egg-cell of plants. When the opponents of the Theory of Descent assert it to be miraculous and inconceivable that an exceedingly complicated many-celled organism could, in the course of time, have proceeded from a simple single-celled organism, we at once reply that we may see this incredible miracle at any moment, and follow it with our own eyes. For the embryology of animals and plants visibly presents to our eyes in the shortest space of time the same process as that which has taken place in the origin of the whole tribe during the course of enormous periods of time.

Upon the ground of embryological records, therefore, we can with full assurance maintain that all many-celled, as well as single-celled, organisms are originally descended from simple cells; connected with this, of course, is the conclusion that the most ancient root of the animal and vegetable kingdom was common to both. For the different primæval “original cells” out of which the few different main groups or tribes have developed, only acquired their differences after a time, and were descended from a common “primæval cell.” But where did those few “original cells,” or the one primæval cell, come from? For the answer to this fundamental genealogical question we must return to the theory of plastids and the hypothesis of spontaneous generation which we have already discussed (vol. i. p. [327]).

As was then shown, we cannot imagine cells to have arisen by spontaneous generation, but only Monera, those primæval creatures of the simplest kind conceivable, like the still living Protamœbæ), Protomyxæ, etc. (vol. i. p. 1[186], Fig. 1). Only such corpuscules of mucus without component parts—whose whole albuminous body is as homogeneous in itself as an inorganic crystal, but which nevertheless fulfills the two organic fundamental functions of nutrition and propagation—could have directly arisen out of inorganic matter by autogeny at the beginning (we may suppose) of the Laurentian period. While some Monera remained at the original simple stage of formation, others gradually developed into cells by the inner kernel of the albuminous mass becoming separated from the external cell-substance. In others, by differentiation of the outermost layer of the cell-substance, an external covering (membrane, or skin) was formed round simple cytods (without kernel), as well as round naked cells (containing a kernel). By these two processes of separation in the simple primæval mucus of the Moneron body, by the formation of a kernel in the interior and a covering on the outer surface of the mass of plasma, there arose out of the original most simple cytods, or Monera, those four different species of plastids, or individuals, of the first order, from which, by differentiation and combination, all other organisms could afterwards develop themselves. (Compare vol. i. p. [347].)

The question now forces itself upon us, Are all organic cytods and cells, and consequently also those “original cells” which we previously considered to be the primary parents of the few great main groups of the animal and vegetable kingdoms, descended from a single original form of Moneron, or were there several different organic primary forms, each traceable to a peculiar independent species of Moneron which originated by spontaneous generation? In other words, Is the whole organic world of a common origin, or does it owe its origin to several acts of spontaneous generation? This fundamental question of genealogy seems at first sight to be of exceeding importance. But on a more accurate examination, we shall soon see that this is not the case, and that it is in reality a matter of very subordinate importance.

Let us now pass on to examine and clearly limit our conception of an organic tribe. By tribe, or phylum, we understand all those organisms of whose blood relationship and descent from a common primary form there can be no doubt, or whose relationship, at least, is most probable from anatomical reasons, as well as from reasons founded on historical development. Our tribes, or phyla, according to this idea, essentially coincide with those few “great classes,” or “main classes,” of which Darwin also thinks that each contains only organisms related by blood, and of which, both in the animal and in the vegetable kingdoms, he only assumes either four or five. In the animal kingdom these tribes would essentially coincide with those four, five, or six main divisions which zoologists, since Bär and Cuvier, have distinguished as “main forms, general plans, branches, or sub-kingdoms” of the animal kingdom. (Compare vol. i. p. [53.]) Bär and Cuvier distinguished only four of them, namely:—1. The vertebrate animals (Vertebrata); 2. The articulated animals (Articulata); 3. The molluscous animals (Mollusca); and 4. The radiated animals (Radiata). At present six are generally distinguished, since the tribe of the articulated animals is divided into two tribes, those possessing articulated feet (Arthropoda), and the worms (Vermes); and in like manner the tribe of radiated animals is subdivided into the two tribes of the star animals (Echinodermata) and the animal-plants (Zoophyta). Within each of these six tribes, all the included animals, in spite of great variety in external form and inner structure, nevertheless possess such numerous and important characteristics in common, that there can be no doubt of their blood relationship. The same applies also to the six great main classes which modern botany distinguishes in the vegetable kingdom, namely:—1. Flowering plants (Phanerogamia); 2. Ferns (Filicinæ); 3. Mosses (Muscinæ); 4. Lichens (Lichenes); 5. Fungi (Fungi); and 6. Water-weeds (Algæ). The last three groups, again, show such close relations to one another, that by the name of “Thallus plants” they may be contrasted with the three first main classes, and consequently the number of phyla, or main groups, of the vegetable kingdom may be reduced to the number of four. Mosses and ferns may likewise be comprised as “Prothallus plants” (Prothallophyta), and thereby the number of plant tribes reduced to three—Flowering plants, Prothallus plants, and Thallus plants.

Very important facts in the anatomy and the history of development, both in the animal and vegetable kingdoms, support the supposition that even these few main classes or tribes are connected at their roots, that is, that the lowest and most ancient primary forms of all three are related by blood to one another. Nay, by a further examination we are obliged to go still a step further, and to agree with Darwin’s supposition, that even the two pedigrees of the animal and vegetable kingdom are connected at their lowest roots, and that the lowest and most ancient animals and plants are derived from a single common primary creature. According to our view, this common primæval organism can have been nothing but a Moneron which took its origin by spontaneous generation.

In the mean time we shall at all events be acting cautiously if we avoid this last step, and assume true blood relationship only within each tribe, or phylum, where it has been undeniably and surely established by facts in comparative anatomy, ontogeny, and phylogeny. But we may here point to the fact that two different fundamental forms of genealogical hypothesis are possible, and that all the different investigations of the Theory of Descent in relation to the origin of organic groups of forms will, in future, tend more and more in one or the other of these directions. The unitary, or monophyletic, hypothesis of descent will endeavour to trace the first origin of all individual groups of organisms, as well as their totality, to a single common species of Moneron which originated by spontaneous generation (vol. i. p. [343]). The multiple, or polyphyletic, hypothesis of descent, on the other hand, will assume that several different species of Monera have arisen by spontaneous generation, and that these gave rise to several different main classes (tribes, or phyla) (vol. i. p. [348]). The apparently great contrast between these two hypotheses is in reality of very little importance. For both the monophyletic and the polyphyletic hypothesis of descent must necessarily go back to the Monera as the most ancient root of the one or of the many organic tribes. But as the whole body of a Moneron consists only of a simple, formless mass, without component particles, made up of a single albuminous combination of carbon, it follows that the differences of the different Monera can only be of a chemical nature, and can only consist in a different atomic composition of that mucous albuminous combination. But these subtle and complicated differences of mixture of the infinitely manifold combinations of albumen are not appreciable by the rude and imperfect means of human observation and are, consequently, at present of no further interest to the task we have in hand.

The question of the monophyletic or polyphyletic origin will constantly recur within each individual tribe, where the origin of a smaller or of a larger group is discussed. In the vegetable kingdom, for example, some botanists will be inclined to derive all flowering plants from a single form of fern, while others will prefer the idea that several different groups of Phanerogama have sprung from several different groups of ferns. In like manner, in the animal kingdom, some zoologists will be more in favour of the supposition that all placental animals are derived from a single pouched animal; others will be more in favour of the opposite supposition, that several different groups of placental animals have proceeded from several different pouched animals. In regard to the human race itself, some will prefer to derive it from a single form of ape, while others will be more inclined to the idea that several different races of men have arisen, independently of one another, out of several different species of ape. Without here expressing our opinion in favour of either the one or the other conception, we must, nevertheless, remark that in general the monophyletic hypothesis of descent deserves to be preferred to the polyphyletic hypothesis of descent. In accordance with the chorological proposition of a single “centre of creation” or of a single primæval home for most species (which has already been discussed), we may be permitted to assume that the original form of every larger or smaller natural group only originated once in the course of time, and only in one part of the earth. We may safely assume this simple original root, that is, the monophyletic origin, in the case of all the more highly developed groups of the animal and vegetable kingdoms. (Compare vol. i. p. [353.]) But it is very possible that the more complete Theory of Descent of the future will involve the polyphyletic origin of very many of the low and imperfect groups of the two organic kingdoms.

For these reasons I consider it best, in the mean time, to adopt the monophyletic hypothesis of descent both for the animal and for the vegetable kingdom. Accordingly, the above-mentioned six tribes, or phyla, of the animal kingdom must be connected at their lowest root, and likewise the three or six main classes, or phyla, of the vegetable kingdom must be traced to a common and most ancient original form. How the connection of these tribes is to be conceived I shall explain in the succeeding chapters. But before proceeding to this, we must occupy ourselves with a very remarkable group of organisms, which cannot without artificial constraint be assigned either to the pedigree of the vegetable or to that of the animal kingdom. These interesting and important organisms are the primary creatures, or Protista.

All organisms which we comprise under the name of Protista show in their external form, in their inner structure, and in all their vital phenomena, such a remarkable mixture of animal and vegetable properties, that they cannot with perfect justice be assigned either to the animal or to the vegetable kingdom; and for more than twenty years an endless and fruitless dispute has been carried on as to whether they are to be assigned to this or that kingdom. Most of Protista are so small that they can scarcely, if at all, be perceived with the naked eye. Hence the majority of them have only become known during the last fifty years, since by the help of the improved and general use of the microscope these minute organisms have been more frequently observed and more accurately examined. However, no sooner were they better known than endless disputes arose about their real nature and their position in the natural system of organisms. Many of these doubtful primary creatures botanists defined as animals, and zoologists as plants; neither of the two would own them. Others, again, were declared by botanists to be plants, and by zoologists to be animals; each claimed them. These contradictions are not altogether caused by our imperfect knowledge of the Protista, but in reality by their true nature. Indeed, most Protista present such a confused mixture of several animal and vegetable characteristics, that each investigator may arbitrarily assign them either to the animal or vegetable kingdom. Accordingly as he defines these two kingdoms, and as he looks upon this or that characteristic as determining the animal or vegetable nature, he will assign the individual classes of Protista in one case to the animal and in another to the vegetable kingdom. But this systematic difficulty has become an inextricable knot by the fact that all more recent investigations on the lowest organisms have completely effaced, or at least destroyed, the sharp boundary between the animal and vegetable kingdom which had hitherto existed, and to such a degree that its restoration is possible only by means of a completely artificial definition of the two kingdoms. But this definition could not be made so as to apply to many of the Protista.

For this and other reasons it is, in the mean time, best to exclude the doubtful beings from the animal as well as from the vegetable kingdom, and to comprise them in a third organic kingdom standing midway between the two others. This intermediate kingdom I have established as the Kingdom of the Primary Creatures (Protista), when discussing general anatomy in the first volume of my General Morphology, pp. 191-238. In my Monograph of the Monera,[(15)] I have recently treated of this kingdom, having somewhat changed its limits, and given it a more accurate definition. Of independent classes of the kingdom Protista, we may at present distinguish the following:—

1. The still living Monera; 2. The Amœboidea, or Protoplasts; 3. The Whip-swimmers, or Flagellata; 4. The Flimmer-balls, or Catallacta; 5. The Tram-weavers, or Labyrinthuleæ; 6. The Flint-cells, or Diatomeæ; 7. The Slime-moulds, or Myxomycetes; 8. The Ray-streamers, or Rhizopoda.

The most important groups at present distinguishable in these eight classes of Protista are named in the systematic table on p. [51.] Probably the number of these Protista will be considerably increased in future days by the progressive investigations of the ontogeny of the simplest forms of life, which have only lately been carried on with any great zeal. With most of the classes named we have become intimately acquainted only during the last ten years. The exceedingly interesting Monera and Labyrinthuleæ, as also the Catallacta, were indeed discovered only a few years ago. It is probable also that very numerous groups of Protista have died out in earlier periods, without having left any fossil remains, owing to the very soft nature of their bodies. We might add to the Protista from the still living lowest groups of organisms—the Fungi; and in so doing should make a very large addition to its domain. Provisionally we shall leave them among plants, though many naturalists have separated them altogether from the vegetable kingdom.

The pedigree of the kingdom Protista is still enveloped in the greatest obscurity. The peculiar combination of animal and vegetable properties, the indifferent and uncertain character of their relations of forms and vital phenomena, together with a number of several very peculiar features which separate most of the subordinate classes sharply from the others, at present baffle every attempt distinctly to make out their blood relationships with one another, or with the lowest animals on the one hand, and with the lowest plants on the other hand. It is not improbable that the classes specified, and many other unknown classes of Protista, represent quite independent organic tribes, or phyla, each of which has independently developed from one, perhaps from various, Monera which have arisen by spontaneous generation. If we do not agree to this polyphyletic hypothesis of descent, and prefer the monophyletic hypothesis of the blood relationship of all organisms, we shall have to look upon the different classes of Protista as the lower small offshoots of the root, springing from the same simple Monera root, out of which arose the two mighty and many-branched pedigrees of the animal kingdom on the one hand, and of the vegetable kingdom on the other. (Compare pp. [74], [75].) Before I enter into this difficult question more accurately, it will be appropriate to premise something further as to the contents of the classes of Protista given on the next page, and their general natural history.

Fig. 8.—Protamœba primitiva, a fresh-water Moneron, much enlarged. A. The entire Moneron with its form-changing processes. B. It begins to divide itself into two halves. C. The division of the two halves is completed, and each now represents an independent individual.

It will perhaps seem strange that I should here again begin with the remarkable Monera as the first class of the Protista kingdom, as I of course look upon them as the most ancient primary forms of all organisms without exception. Still, what are we otherwise to do with the still living Monera? We know nothing of their palæontological origin, we know nothing of any of their relations to lower animals or plants, and we know nothing of their possible capability of developing into higher organisms. The simple and homogeneous little lump of slime or mucus which constitutes their entire body (Fig. [8]) is the most ancient and original form of animal as well as of vegetable plastids. Hence it would evidently be just as arbitrary and unreasonable to assign them to the animal as it would be to assign them to the vegetable kingdom. In any case we shall for the present be acting more cautiously and critically if we comprise the still living Monera—whose number and distribution is probably very great—as a special and independent class, contrasting them with the other classes of the kingdom Protista, as well as with the animal kingdom. Morphologically considered, the Monera—on account of the perfect homogeneity of the albuminous substance of their bodies, on account of their utter want of heterogeneous particles—are more closely connected with anorgana than with organisms, and evidently form the transition between the inorganic and organic world of bodies, as is necessitated by the hypothesis of spontaneous generation. I have described and given illustrations of the forms and vital phenomena of the still living Monera (Protamœba, Protogenes, Protomyxa, etc.) in my Monograph of the Monera,[(15)] and have briefly mentioned the most important facts in the eighth chapter (vol. i. pp. [183]-187). Therefore, only by way of a specimen, I here repeat the drawing of the fresh-water Protamœba (Fig. 8). The history of the life of an orange-red Protomyxa adrantiaca, which I observed at Lanzerote, one of the Canary Islands, is given in Plate [I]. (see its explanation in the Appendix). Besides this, I here add a drawing of the form of Bathybius, that remarkable Moneron discovered by Huxley, which lives in the greatest depths of the sea in the shape of naked lumps of protoplasm and reticular mucus (vol. i. p. [344]).

The Amœbæ of the present day, and the organisms most closely connected with them, Arcellidæ and Gregarinæ, which we here unite as a second class of Protista under the name of Amœboidea (Protoplasta), present no fewer genealogical difficulties than the Monera. These primary creatures are at present usually placed in the animal kingdom without its in reality being understood why. For simple naked cells—that is, shell-less plastids with a kernel—occur as well among real plants as real animals. The generative cells, for example, in many Algæ (spores and eggs) exist for a longer or shorter time in water in the form of naked cells with a kernel, which cannot be distinguished at all from the naked eggs of many animals (for example, those of the Siphonophorous Medusæ). (Compare the figure of a naked egg of a bladder-wrack in Chapter xvii. p. [90].) In reality every naked simple cell, whether it proceeds from an animal or vegetable body, cannot be distinguished from an independent Amœba. For an Amœba is nothing but a simple primary cell, a naked little lump of cell-matter, or plasma, containing a kernel. The contractility of this plasma, which the free Amœba shows in stretching out and drawing in its changing processes, is a general vital property of the organic plasma of all animal as well as of all vegetable plastids. When a freely moving Amœba, which perpetually changes its form, passes into a state of rest, it draws itself together into the form of a globule, and surrounds itself with a secreted membrane. It can then be as little distinguished from an animal egg as from a simple globular vegetable cell (Fig. 10 A).

Fig. 10.—Amœba sphærococcus, greatly magnified. A fresh-water Amœba without a contractile vacuole. A. The enclosed Amœba in the state of a globular lump of plasma (c) enclosing a kernel and a kernel-speck (a). The simple cell is surrounded by a cyst, or cell membrane (d). B. The free Amœba, which has burst and left the cyst, or cell-membrane. C. It begins to divide by its kernel parting into two kernels, and the cell-substance between the two contracting. D. The division is completed, and the cell-substance has entirely separated into two bodies. (Da and Db)

Naked cells, with kernels, like those represented in Fig. 10 B, which are continuously changing, stretching out and drawing in formless, finger-like processes, and which are on this account called amœboid, are found frequently and widely dispersed in fresh water and in the sea; nay, are even found creeping on land. They take their food in the same way as was previously described in the case of the Protamœba (vol. i. p. [186]). Their propagation by division can sometimes be observed. (Fig. 10 C, D.) I have described the processes in an earlier chapter (vol. i. p. [187]). Many of these formless Amœbæ have lately been recognized as the early stages of development of other Protista (especially the Myxomycetæ), or as the freed cells of lower animals and plants. The colourless blood-cells of animals, for example, those of human blood, cannot be distinguished from Amœbæ. They, like the latter, can receive solid corpuscles into their interior, as I was the first to show by feeding them with finely divided colouring matters (Gen. Morph. i. 271). However, other Amœbæ (like the one given in Fig. 10) seem to be independent “good species,” since they propagate themselves unchanged throughout many generations. Besides the real, or naked, Amœbæ (Gymnamœbæ), we also find widely diffused in fresh water case-bearing Amœbæ (Lepamœbæ), whose naked plasma body is partially protected by a more or less solid shell (Arcella), sometimes even by a case (Difflugia) composed of small stones. Lastly, we frequently find in the body of many lower animals parasitic Amœbæ (Gregarinæ), which, adapting themselves to a parasitic life, have surrounded their plasma-body with a delicate closed membrane.

The simple naked Amœbæ are, next to the Monera, the most important of all organisms to the whole science of biology, and especially to general genealogy. For it is evident that the Amœbæ originally arose out of simple Monera (Protamœbæ), by the important process of segregation taking place in their homogeneous viscid body—the differentiation of an inner kernel from the surrounding plasma. By this means the great progress from a simple cytod (without kernel) into a real cell (with kernel) was accomplished (compare Fig. 8 A and Fig. 10 B). As some of these cells at an early stage encased themselves by secreting a hardened membrane, they formed the first vegetable cells, while others, remaining naked, developed into the first aggregates of animal cells. The presence or absence of an encircling hard membrane forms the most important, although by no means the entire, difference of form between animal and vegetable cells. As vegetable cells even at an early stage enclose themselves within their hard, thick, and solid cellular shell, like that of the Amœbæ in a state of rest (Fig. 10 A), they remain more independent and less accessible to the influences of the outer world than are the soft animal cells, which are in most cases naked, or merely covered by a thin pliable membrane. But in consequence of this the vegetable cells cannot combine, as do the animal cells, for the construction of higher and composite fibrous tracts, for example, the nervous and muscular tissues. It is probable that, in the case of the most ancient single-celled organisms, there must have developed at an early stage the very important difference in the animal and vegetable mode of receiving food. The most ancient single-celled animals, being naked cells, could admit solid particles into the interior of their soft bodies, as do the Amœbæ (Fig. 10 B) and the colourless blood-cells; whereas the most ancient single-celled plants encased by their membranes were no longer able to do this, and could admit through it only fluid nutrition (by means of diffusion).

Fig. 11.—A single Whip-swimmer (Euglena striata), greatly magnified. Above a thread-like lashing whip is visible; in the centre the round cellular kernel, with its kernel speck.

The Whip-swimmers (Flagellata), which we consider as a third class of the kingdom Protista, are of no less doubtful nature than the Amœbæ. They often show as close and important relations to the vegetable as to the animal kingdom. Some Flagellata at an early stage, when freely moving about, cannot be distinguished from real plants, especially from the spores of many Algæ; whereas others are directly allied to real animals, namely, to the fringed Infusoria (Ciliata). The Flagellata are simple cells which live in fresh or salt water, either singly or united in colonies. The characteristic part of their body is a very movable simple or compound whip-like appendage (whip, or flagellum) by means of which they actively swim about in the water. This class is divided into two orders. Among the fringed whip-swimmers (Cilioflagellata) there exists, in addition to the long whip, a short fringe of vibrating hairs, which is wanting in the unfringed whip-swimmers (Nudoflagellata). To the former belong the flint-shelled yellow Peridinia, which are largely active in causing the phosphorescence of the sea; to the latter belong the green Euglenæ, immense masses of which frequently make our ponds in spring quite green.

Fig. 12.—The Norwegian Flimmer-ball (Magosphæra planula) swimming by means of its vibratile fringes, as seen from the surface.

A very remarkable new form of Protista, which I have named Flimmer-ball (Magosphæra), I discovered only three years ago (in September, 1869), on the Norwegian coast (Fig. 12), and have more accurately described in my Biological Studies[(15)] (p. 137, Plate V.). Off the island of Gis-oe, near Bergen, I found swimming about, on the surface of the sea, extremely neat little balls composed of a number (between thirty and forty) of fringed pear-shaped cells, the pointed ends of which were united in the centre like radii. After a time the ball dissolved. The individual cells swarmed about independently in the water like fringed Infusoria, or Ciliata. These afterwards sank to the bottom, drew their fringes into their bodies, and gradually changed into the form of creeping Amœbæ (like Fig 10 B). These last afterwards encased themselves (as in Fig. 10 A), and then divided by repeated halvings into a large number of cells (exactly as in the case of the cleavage of the egg, Fig. 6, vol. i. p. [299]). The cells became covered with vibratile hairs, broke through the case enclosing them, and now again swam about in the shape of a fringed ball (Fig. 12). This wonderful organism, which sometimes appears like a simple Amœba, sometimes as a single fringed cell, sometimes as a many-celled fringed ball, can evidently be classed with none of the other Protista, and must be considered as the representative of a new independent group. As this group stands midway between several Protista, and links them together, it may bear the name of Mediator, or Catallacta.

Fig. 13.—Labyrinthula macrocystis (much enlarged). Below is a large group of accumulated cells, one of which, on the left, is separating itself; above are two single cells which are gliding along the threads of the retiform labyrinth which form their “tramways.”

The Protista of the fifth class, the Tram-weavers, or Labyrinthuleæ, are of a no less puzzling nature; they were lately discovered by Cienkowski on piles in sea water (Fig. 13). They are spindle-shaped cells, mostly of a yellow-ochre colour, which are sometimes united into a dense mass, sometimes move about in a very peculiar way. They form, in a manner not yet explained, a retiform frame of entangled threads (compared to a labyrinth), and on the dense filamentous “tramways” of this frame they glide about. From the shape of the cells of the Labyrinthuleæ we might consider them as the simplest plants, from their motion as the simplest animals, but in reality they are neither animals nor plants.

Fig. 14.—Navicula hippocampus (greatly magnified). In the middle of the cell the cell-kernel (nucleus) is visible, together with its kernel speck (nucleolus).

The Flint-cells (Diatomeæ), a sixth class of Protista, are perhaps the most closely related to the Labyrinthuleæ. These primary creatures—which at present are generally considered as plants, although some celebrated naturalists still look upon them as animals—inhabit the sea and fresh waters in immense masses, and offer an endless variety of the most elegant forms. They are mostly small microscopic cells, which either live singly (Fig. 14), or united in great numbers, and occur either attached to objects, or glide and creep about in a peculiar manner. Their soft cell-substance, which is of a characteristic brownish yellow colour, is always enclosed by a solid and hard flinty shell, possessing the neatest and most varied forms. This flinty covering is open to the exterior only by one or two slits, through which the enclosed soft plasma-body communicates with the outer world. The flinty cases are found petrified in masses, and many rocks—for example, the Tripoli slate polish, the Swedish mountain meal, etc.,—are in a great measure composed of them.

Fig. 15.—A stalked fruit-body (spore-bladder, filled with spores) of one of the Myxomycetes (Physarum albipes) not much enlarged.

A seventh class of Protista is formed by the remarkable Slime-moulds (Myxomycetes). They were formerly universally considered as plants, as real Fungi, until ten years ago the botanist De Bary, by discovering their ontogeny, proved them to be quite distinct from Fungi, and rather to be akin to the lower animals. The mature body is a roundish bladder, often several inches in size, filled with fine spore-dust and soft flakes (Fig. 15), as in the case of the well-known puff-balls (Gastromycetes). However, the characteristic cellular threads, or hyphæ, of a real fungus do not arise from the germinal corpuscles, or spores, of the Myxomycetes, but merely naked masses of plasma, or cells, which at first swim about in the form of Flagellata (Fig. 11), afterwards creep about like the Amœbæ (Fig. 10 B), and finally combine with others of the same kind to form large masses of “slime,” or “plasmodia.” Out of these, again, there arises, by-and-by, the bladder-shaped fruit-body. Many of my readers probably know one of these plasmodia, the Æthalium septicum, which in summer forms a beautiful yellow mass of soft mucus, often several feet in breadth, known by the name of “tan flowers,” and penetrates tan-heaps and tan-beds. At an early stage these slimy, freely-creeping Myxomycetes, which live for the most part in damp forests, upon decaying vegetable substances, bark of trees, etc., are with equal justice or injustice declared by zoologists to be animals, while in the mature, bladder-shaped condition of fructification they are by botanists defined as plants.

The nature of the Ray-streamers (Rhizopoda), the eighth class of the kingdom Protista, is equally obscure. These remarkable organisms have peopled the sea from the most ancient times of the organic history of the earth, in an immense variety of forms, sometimes creeping at the bottom of the sea, sometimes swimming on the surface. Only very few live in fresh water (Gromia, Actinosphærium). Most of them possess solid calcareous or flinty shells of an extremely beautiful construction, which can be perfectly preserved in a fossil state. They have frequently accumulated in such huge numbers as to form mountain masses, although the single individuals are very small, and often scarcely visible, or completely invisible to the naked eye. A very few attain the diameter of a few lines, or even as much as a couple of inches. The name which the class bears is given because thousands of exceedingly fine threads of protoplasm radiate from the entire surface of their naked slimy body; these rays are quasi-feet, or pseudopodia, which branch off like roots (whence the term Rhizopoda, signifying root-footed), unite like nets, and are observed continually to change form, as in the case of the simpler plasmic feet of the Amœboidea, or Protoplasts. These ever-changing little pseudo-feet serve both for locomotion and for taking food.

The class of the Rhizopoda is divided into three different legions, viz. the chamber-shells, or Acyttaria, the sun-animalcules, or Heliozoa, and the basket-shells, or Radiolaria. The Chamber-shells (Acyttaria) constitute the first and lowest of these three legions; for the whole of their soft body consists merely of simple mucous or slimy cell-matter, or protoplasm, which has not differentiated into cells. However, in spite of this most primitive nature of body, most of the Acyttaria secrete a solid shell composed of calcareous earth, which presents a great variety of exquisite forms. In the more ancient and more simple Acyttaria this shell is a simple chamber, bell-shaped, tubular, or like the shell of a snail, from the mouth of which a bundle of plasmic threads issues. In contrast to these single-chambered forms (Monothalamia), the many-chambered forms (Polythalamia)—to which the great majority of the Acyttaria belong—possess a house, which is composed in an artistic manner of numerous chambers. These chambers sometimes lie in a row one behind the other, sometimes in concentric circles or spirals, in the form of a ring round a central point, and then frequently one above another in many tiers, like the boxes of an amphitheatre. This formation, for example, is found in the nummulites, whose calcareous shells, of the size of a lentil, have accumulated to the number of millions, and form whole mountains on the shores of the Mediterranean. The stones of which some of the Egyptian pyramids are built consist of such nummulitic limestone. In most cases the chambers of the shells of the Polythalamia are wound round one another in a spiral line. The chambers are connected with one another by passages and doors, like rooms of a large palace, and are generally open towards the outside by numerous little windows, out of which the plasmic body can stream or strain forth its little pseudo-feet, or rays of slime, which are always changing form. But in spite of the exceedingly complicated and elegant structure of this calcareous labyrinth, in spite of the endless variety in the structure and the decoration of its numerous chambers, and in spite of the regularity and elegance of their execution, the whole of this artistic palace is found to be the secreted product of a perfectly formless, slimy mass, devoid of any component parts! Verily, if the whole of the recent anatomy of animal and vegetable textures did not support our theory of plastids, if all its important results did not unanimously corroborate the fact that the whole miracle of vital phenomena and vital forms is traceable to the active agency of the formless albuminous combinations of protoplasm, the Polythalamia alone would secure the triumph of that theory. For we may here at any moment, by means of the microscope, point out the wonderful fact, first established by Dujardin and Max Schulze, that the formless mucus of the soft plasma-body, this true “matter of life,” is able to secrete the neatest, most regular, and most complicated structures. This secretive skill is simply a result of inherited adaptation, and by it we learn to understand how this same “primæval slime”—this same protoplasm—can produce in the bodies of animals and plants the most different and most complicated cellular forms.

It is, moreover, a matter of special interest that the most ancient organism, the remains of which are found in a petrified condition, belongs to the Polythalamia. This organism is the “Canadian Life’s-dawn” (Eozoon canadense), which has already been mentioned, and which was found a few years ago in the Ottawa formation (in the deepest strata of the Laurentian system), on the Ottawa river in Canada. If we expected to find organic remains at all in these most ancient deposits of the primordial period, we should certainly look for such of the most simple Protista as are covered with a solid shell, and in the organization of which the difference between animal and plant is as yet not indicated.

We know of but few species of the Sun-animalcules (Heliozoa), the second class of the Rhizopoda. One species is very frequently found in our fresh waters. It was observed even in the last century by a clergyman in Dantzig, Eichhorn by name, and it has been called after him, Actinosphærium Eichhornii. To the naked eye it appears as a gelatinous grey globule of mucus, about the size of a pin’s head. Looking at it through the microscope, we see hundreds or thousands of fine mucous threads radiating from the central plasma body, and perceive that the inner layer of its cell-substance is different from the outer layer, which forms a bladder-like membrane. In consequence of its structure, this, the little sun-animalcule, although wanting a shell, really rises above the structureless Acyttaria, and forms the transition from these to the Radiolaria. The genus Cystophrys is of a nature akin to it.

The Basket-shells (Radiolaria) form the third and last class of the Rhizopoda. Their lower forms are closely allied to the Heliozoa and Acyttaria, whereas their higher forms rise far above them. They are essentially distinguished from both by the fact that the central part of their body is composed of many cells, and surrounded by a solid membrane. This closed “central capsule,” generally of a globular shape, is covered by a mucous layer of plasma, out of which there radiate on all sides thousands of exceedingly fine threads, the branching and confluent so-called pseudopodia. Between these are scattered numerous yellow cells of unknown function, containing grains of starch. Most Radiolaria are characterized by a highly developed skeleton, which consists of flint, and displays a wonderful richness of the neatest and most curious forms. Sometimes this flinty skeleton forms a simple trellice-work ball (Fig. 16 s), sometimes a marvellous system of several concentric trelliced balls, encased in one another, and connected by radial staves. In most cases delicate spikes, which are frequently branched like a tree, radiate from the surface of the balls. In other cases the whole skeleton consists of only one flinty star, and is then generally composed of twenty staves, distributed according to definite mathematical laws, and united in a common central point. The skeletons of other Radiolaria again form symmetrical many-chambered structures, as in the case of the Polythalamia. Perhaps no other group of organisms develop in the formation of their skeletons such an amount of various fundamental forms, such geometrical regularity, and such elegant architecture. Most of the forms as yet discovered, I have given in the atlas accompanying my Monograph of the Radiolaria.[(23)] Here I shall only give as an example the picture of one of the simplest forms, the Cyrtidosphæra echinoides of Nice. The skeleton in this case consists only of a simple trelliced ball (s), with short radial spikes (a), which loosely surround the central capsule (c). Out of the mucous covering, enclosing the latter, radiate a great number of delicate little pseudopodia (p), which are partly drawn back underneath the shell, and fused into a lumpy mass of mucus. Between these are scattered a number of yellow cells (l).

Fig. 16.—Cyrtidosphæra echinoides, 400 times enlarged. c. Globular central capsule. s. Basket-work of the perforated flinty shell. a. Radial spikes, which radiate from the latter. p. The pseudo-feet radiating from the mucous covering surrounding the central capsule. l. Yellow globular cells, scattered between the latter, containing grains of starch.

Most Acyttaria live only at the bottom of the sea, on stones and seaweeds, or creep about in sand and mud by means of their pseudopodia, but most Radiolaria swim on the surface of the sea by means of long pseudopodia extending in all directions. They live together there in immense numbers, but are mostly so small that they have been almost completely overlooked, and have only become accurately known during the last fourteen years. Certain Radiolaria living in communities (Polycyttaria) form gelatinous lumps of some lines in diameter. On the other hand, most of those living isolated (Monocyttaria) are invisible to the naked eye; but still their petrified shells are found accumulated in such masses that in many places they form entire mountains; for example, the Nicobar Islands in the Indian Archipelago, and the Island of Barbadoes in the Antilles.

As most readers are probably but little acquainted with the eight classes of the Protista just mentioned, I shall now add some further general observations on their natural history. The great majority of all Protista live in the sea, some swimming freely on the surface, some creeping at the bottom, and others attached to stones, shells, plants, etc. Many species of Protista also live in fresh water, but only a very small number on dry land (for example, Myxomycetes and some Protoplasta). Most of them can be seen only through the microscope, except when millions of individuals are found accumulated. Only a few of them attain a diameter of some lines, or as much as an inch. What they lack in size of body they make up for by producing astonishing numbers of individuals, and they very considerably influence in this way the economy of nature. The imperishable remains of dead Protista, for instance, the flinty shells of the Diatomeæ and Radiolaria and the calcareous shells of the Acyttaria, often form large rock masses.

In regard to their vital phenomena, especially those of nutrition and propagation, some Protista are more allied to plants, others more to animals. Both in their mode of taking food and in the chemical changes of their living substance, they sometimes more resemble the lower animals, at others the lower plants. Free locomotion is possessed by many Protista, while others are without it; but this does not constitute a characteristic distinction, as we know of undoubted animals which entirely lack free locomotion, and of genuine plants which possess it. All Protista have a soul—that is to say, are “animate”—as well as all animals and all plants. The soul’s activity in the Protista manifests itself in their irritability, that is, in the movements and other changes which take place in consequence of mechanical, electrical, and chemical irritation of their contractile protoplasm. Consciousness and the capability of will and thought are probably wanting in all Protista. However, the same qualities are in the same degree also wanting in many of the lower animals, whereas many of the higher animals in these respects are scarcely inferior to the lower races of human beings. In the Protista, as in all other organisms, the activities of the soul are traceable to molecular motions in the protoplasm.

The most important physiological characteristic of the kingdom Protista lies in the exclusively non-sexual propagation of all the organisms belonging to it. The higher animals and plants multiply almost exclusively in a sexual manner. The lower animals and plants multiply also, in many cases, in a non-sexual manner, by division, the formation of buds, the formation of germs, etc. But sexual propagation almost always exists by the side of it, and often regularly alternates with it in succeeding generations (Metagenesis, vol. i. p. [206]). All Protista, on the other hand, propagate themselves exclusively in a non-sexual manner, and in fact, the distinction of the two sexes among them has not been effected—there are neither male nor female Protista.

The Protista in regard to their vital phenomena stand midway between animals and plants, that is to say, between their lowest forms; and the same must be said in regard to the chemical composition of their bodies. One of the most important distinctions between the chemical composition of animal and vegetable bodies consists in the characteristic formation of the skeleton. The skeleton, or the solid scaffolding of the body in most genuine plants, consists of a substance called cellulose, devoid of nitrogen, but secreted by the nitrogenous cell-substance, or protoplasm. In most genuine animals, on the other hand, the skeleton generally consists either of nitrogenous combinations (chitin, etc.) or of calcareous earth. In this respect some Protista are more like plants, others more like animals. In many of them the skeleton is principally or entirely formed of calcareous earth, which is met with both in animal and vegetable bodies. But the active vital substance in all cases is the mucous protoplasm.

In regard to the form of the Protista, it is to be remarked that the individuality of their body almost always remains at an extremely low stage of development. Very many Protista remain for life simple plastids or individuals of the first order. Others, indeed, form colonies or republics of plastids by the union of several individuals. But even these higher individuals of the second order, formed by the combination of simple plastids, for the most part remain at a very low stage of development. The members of such communities among the Protista remain very similar one to another, and never, or only in a slight degree, commence a division of labour, and are consequently as little able to render their community fit for higher functions as are, for example, the savages of Australia. The community of the plastids remains in most cases very loose, and each single plastid retains in a great measure its own individual independence.

A second structural characteristic, which next to their low stage of individuality especially distinguishes the Protista, is the low stage of development of their stereometrical fundamental forms. As I have shown in my theory of fundamental forms (in the fourth book of the General Morphology), a definite geometrical fundamental form can be pointed out in most organisms, both in the general form of the body and in the form of the individual parts. This ideal fundamental form, or type, which is determined by the number, position, combination, and differentiation of the component parts, stands in just the same relation to the real organic form as the ideal geometrical fundamental form of crystals does to their imperfect real form. In most bodies and parts of the bodies of animals and plants this fundamental form is a pyramid. It is a regular pyramid in the so-called “regular radiate” forms, and an irregular pyramid in the more highly differentiated, so-called “bilaterally symmetrical” forms. (Compare the plates in the first volume of my General Morphology, pp. 556-558.) Among the Protista this pyramidal type, which prevails in the animal and vegetable kingdom, is on the whole rare, and instead of it we have either quite irregular (amorphous) or more simple, regular geometrical types; especially frequent are the sphere, the cylinder, the ellipsoid, the spheroid, the double cone, the cone, the regular polygon (tetrahedron, hexahedron, octahedron, dodecahedron, icosahedron), etc. All the fundamental forms of the pro-morphological system, which are of a low rank in that system, prevail in the Protista. However, in many Protista there occur also the higher, regular, and bilateral types, fundamental forms which predominate in the animal and vegetable kingdoms. In this respect some of the Protista are frequently more closely allied to animals (as the Acyttaria), others more so to plants (as the Radiolaria).

With regard to the palæontological development of the kingdom Protista, we may form various, but necessarily very unsafe, genealogical hypotheses. Perhaps the individual classes of the kingdom are independent tribes, or phyla, which have developed independently of one another and independently of the animal and the vegetable kingdoms. Even if we adopt the monophyletic hypothesis of descent, and maintain a common origin from a single form of Moneron for all organisms, without exception, which ever have lived and still live upon the earth, even in this case the connection of the neutral Protista on the one hand with the vegetable kingdom, and on the other hand with the animal kingdom, must be considered as very vague. We must regard them (compare p. [74]) as lower offshoots which have developed directly out of the root of the great double-branched organic pedigree, or perhaps out of the lowest tribe of Protista, which may be supposed to have shot up midway between the two diverging high and vigorous trunks of the animal and vegetable kingdoms. The individual classes of the Protista, whether they are more closely connected at their roots in groups, or only form a loose bunch of root offsets, must in this case be regarded as having nothing to do either with the diverging groups of organisms belonging to the animal kingdom on the right, or to the vegetable kingdom on the left. They must be supposed to have retained the original simple character of the common primæval living thing more than have genuine animals and genuine plants.

But if we adopt the polyphyletic hypothesis of descent, we have to imagine a number of organic tribes, or phyla, which all shoot up by spontaneous generation out of the same ground, by the side of and independent of one another. (Compare p. [75].) In that case numbers of different Monera must have arisen by spontaneous generation whose differences would depend only upon slight, to us imperceptible, differences in their chemical composition, and consequently upon differences in their capability of development. A small number of Monera would then have given origin to the animal kingdom, and, again, a small number would have produced the vegetable kingdom. Between these two groups, however, there would have developed, independently of them, a large number of independent tribes, which have remained at a lower stage of organization, and which have neither developed into genuine plants nor into genuine animals.

A safe means of deciding between the monophyletic and polyphyletic hypotheses is as yet quite impossible, considering the imperfect state of our phylogenetic knowledge. The different groups of Protista, and those lowest forms of the animal kingdom and of the vegetable kingdom which are scarcely distinguishable from the Protista, show such a close connection with one another and such a confused mixture of characteristics, that at present any systematic division and arrangement of the groups of forms seem more or less artificial and forced. Hence the attempt here offered must be regarded as entirely provisional. But the more deeply we penetrate into the genealogical secrets of this obscure domain of inquiry, the more probable appears the idea that the vegetable kingdom and the animal kingdom are each of independent origin, and that midway between these two great pedigrees a number of other independent small groups of organisms have arisen by repeated acts of spontaneous generation, which on account of their indifferent neutral character, and in consequence of their mixture of animal and vegetable properties, may lay claim to the designation of independent Protista.

N.B.—The lines marked with a ♱ indicate extinct tribes of Protista, which have arisen independently by repeated acts of Spontaneous Generation.

Thus, if we assume one entirely independent trunk for the vegetable kingdom, and a second for the animal kingdom, we may set up a number of independent stems of Protista, each of which has developed, quite independently of other stems and trunks, from a special archigonic form of Monera. In order to make this relation more clear, we may imagine the whole world of organisms as an immense meadow which is partially withered, and upon which two many-branched and mighty trees are standing, likewise partially withered. The two great trees represent the animal and vegetable kingdoms, their fresh and still green branches the living animals and plants; the dead branches with withered leaves represent the extinct groups. The withered grass of the meadow corresponds to the numerous extinct tribes, and the few stalks, still green, to the still living phyla of the kingdom Protista. But the common soil of the meadow, from which all have sprung up, is primæval by protoplasm.

CHAPTER XVII.

PEDIGREE AND HISTORY OF THE VEGETABLE KINGDOM.

The Natural System of the Vegetable Kingdom.—Division of the Vegetable Kingdom into Six Branches and Eighteen Classes.—The Flowerless Plants (Cryptogamia).—Sub-kingdom of the Thallus Plants.—The Tangles, or Algæ (Primary Algæ, Green Algæ, Brown Algæ, Red Algæ.)—The Thread-plants, or Inophytes (Lichens and Fungi.)—Sub-kingdom of the Prothallus Plants.—The Mosses, or Muscinæ (Water-mosses, Liverworts, Leaf-mosses, Bog-mosses).—The Ferns, or Filicinæ (Leaf-ferns, Bamboo-ferns, Water-ferns, Scale-ferns).—Sub-kingdom of Flowering Plants (Phanerogamia).—The Gymnosperms, or Plants with Naked Seeds (Palm-ferns = Cycadeæ; Pines = Coniferæ.)—The Angiosperms, or Plants with Enclosed Seeds.—Monocotylæ.—Dicotylæ.—Cup-blossoms (Apetalæ).—Star-blossoms (Diapetalæ).—Bell-blossoms (Gamopetalæ).

Every attempt that we make to gain a knowledge of the pedigree of any small or large group of organisms related by blood must, in the first instance, start with the evidence afforded by the existing “natural system” of this group. For although the natural system of animals and plants will never become finally settled, but will always represent a merely approximate knowledge of true blood relationship, still it will always possess great importance as a hypothetical pedigree. It is true, by a “natural system” most zoologists and botanists only endeavour to express in a concise way the subjective conceptions which each has formed of the objective “form-relationships” of organisms. These form-relationships, however, as the reader has seen, are in reality the necessary result of true blood relationship. Consequently, every morphologist in promoting our knowledge of the natural system, at the same time promotes our knowledge of the pedigree, whether he wishes it or not. The more the natural system deserves its name, and the more firmly it is established upon the concordance of results obtained from the study of comparative anatomy, ontogeny, and palæontology, the more surely may we consider it as the approximate expression of the true pedigree of the organic world.

In entering upon the task contemplated in this chapter, the genealogy of the vegetable kingdom, we shall have, according to this principle, first to glance at the natural system of the vegetable kingdom as it is at present (with more or less important modifications) adopted by most botanists. According to the system generally in vogue, the whole series of vegetable forms is divided into two main groups. These main divisions, or sub-kingdoms, are the same as were distinguished more than a century ago by Charles Linnæus, the founder of systematic natural history, and which he called Cryptogamia, or secretly-blossoming plants, and Phanerogamia, or openly-flowering plants. The latter, Linnæus, in his artificial system of plants, divided, according to the different number, formation, and combination of the anthers, and also according to the distribution of the sexual organs, into twenty-three different classes, and then added the Cryptogamia to these as the twenty-fourth and last class.

The Cryptogamia, the secretly-blossoming or flowerless plants, which were formerly but little observed, have in consequence of the careful investigations of recent times been proved to present such a great variety of forms, and such a marked difference in their coarser and finer structure, that we must distinguish no less than fourteen different classes of them; whereas the number of classes of flowering plants, or Phanerogamia, may be limited to four. However, these eighteen classes of the vegetable kingdom can again be naturally grouped in such a manner that we are able to distinguish in all six main divisions or branches of the vegetable kingdom. Two of these six branches belong to the flowering, and four to the flowerless plants. The table on page 82 shows how the eighteen classes are distributed among the six branches, and how these again fall under the sub-kingdoms of the vegetable kingdom.

The one sub-kingdom of the Cryptogamia may now be naturally divided into two divisions, or sub-kingdoms, differing very essentially in their internal structure and in their external form, namely, the Thallus plants and the Prothallus plants. The group of Thallus plants comprises the two large branches of Tangles, or Algæ, which live in water, and the Thread-plants, or Inophytes (Lichens and Fungi), which grow on land, upon stones, bark of trees, upon decaying bodies, etc. The group of Prothallus plants, on the other hand, comprises the two branches of Mosses and Ferns, containing a great variety of forms.

All Thallus plants, or Thallophytes, can be directly recognized from the fact that the two morphological fundamental organs of all other plants, stem and leaves, cannot be distinguished in their structure. The complete body of all Algæ and of all Thread-plants is a mass composed of simple cells, which is called a lobe, or thallus. This thallus is as yet not differentiated into axial-organs (stem and root) and leaf-organs. On this account, as well as through many other peculiarities, the Thallophytes contrast strongly with all remaining plants—those comprised under the two sub-kingdoms of Prothallus plants and Flowering plants—and for this reason the two latter sub-kingdoms are frequently classed together under the name of Stemmed plants, or Cormophytes. The following table will explain the relation of these three sub-kingdoms to one another according to the two different views:—

The stemmed plants, or Cormophytes, in the organization of which the difference of axial-organs (stem and root) and leaf-organs is already developed, form at present, and have, indeed, for a very long period formed, the principal portion of the vegetable world. However, this was not always the case. In fact, stemmed plants, not only of the flowering group, but even of the prothallus group, did not exist at all during that immeasurably long space of time which forms the beginning of the first great division of the organic history of the earth, under the name of the archilithic, or primordial period. The reader will recollect that during this period the Laurentian, Cambrian, and Silurian systems of strata were deposited, the thickness of which, taken as a whole, amounts to about 70,000 feet. Now, as the thickness of all the more recent superincumbent strata, from the Devonian to the deposits of the present time, taken together, amounts to only about 60,000 feet, we were enabled from this fact alone to draw the conclusion—which is probable also for other reasons—that the archilithic, or primordial, period was of longer duration than the whole succeeding period down to the present time. During the whole of this immeasurable space of time, which probably comprises many millions of centuries, vegetable life on our earth seems to have been represented exclusively by the sub-kingdom of Thallus plants, and, moreover, only by the class of marine Thallus plants, that is to say, the Algæ. At least all the petrified remains which are positively known to be of the primordial period belong exclusively to this class. As all the animal remains of this immense period also belong exclusively to animals that lived in water, we come to the conclusion that at that time organisms adapted to a life on land did not exist at all.

For these reasons the first and most imperfect of the great provinces or branches of the vegetable kingdom, the division of the Algæ, or Tangles, must be of special interest to us. But, in addition, there is the interest which this group offers when viewed by itself. In spite of the exceedingly simple composition of their constituent cells, which are but little differentiated, the Algæ show an extraordinary variety of different forms. To them belong the simplest and most imperfect of all forms, as well as very highly developed and peculiar forms. The different groups of Algæ are distinguished as much by size of body as by the perfection and variety of their outer form. At the lowest stage we find such species as the minute Protococcus, several hundred thousands of which occupy a space no larger than a pin’s head. At the highest stage we marvel at the gigantic Macrocysts, which attain a length of from 300 to 400 feet, the longest of all forms in the vegetable kingdom. It is possible that a large portion of the coal has been formed out of Algæ. If not for these reasons, yet the Algæ must excite our special attention from the fact that they form the beginning of vegetable life, and contain the original forms of all other groups of plants, supposing that our monophyletic hypothesis of a common origin for all groups of plants is correct. (Compare p. [83.])

Most people living inland can form but a very imperfect idea of this exceedingly interesting branch of the vegetable kingdom, because they know only its proportionately small and simple representatives living in fresh water. The slimy green aquatic filaments and flakes of our pools and ditches and springs, the light green slimy coverings of all kinds of wood which have for any length of time been in contact with water, the yellowish green, frothy, and oozy growths of our village ponds, the green filaments resembling tufts of hair which occur everywhere in fresh water, stagnant and flowing, are for the most part composed of different species of Algæ. Only those who have visited the sea-shore, and wondered at the immense masses of cast-up seaweed, and who, from the rocky coast of the Mediterranean, have seen through the clear blue waters the beautifully-formed and highly-coloured vegetation of Algæ at the bottom, know how to estimate the importance of the class of Algæ. And yet, even these marine Algæ-forests of European shores, so rich in forms, give only a faint idea of the colossal forests of Sargasso in the Atlantic ocean, those immense banks of Algæ, covering a space of about 40,000 square miles—the same which made Columbus, on his voyage of discovery, believe that a continent was near. Similar but far more extensive forests of Algæ grew in the primæval ocean, probably in dense masses, and what countless generations of these archilithic Algæ have died out one after another is attested, among other facts, by the vast thickness of Silurian alum schists in Sweden, the peculiar composition of which proceeds from those masses of submarine Algæ. According to the recently expressed opinion of Frederick Mohr, a geologist of Bonn, even the greater part of our coal seams have arisen out of the accumulated dead bodies of the Algæ forests of the ocean.

Within the branch of the Algæ we distinguish four different classes, each of which is again divided into several orders and families. These again contain a large number of different genera and species. We designate these four classes as Primæval Algæ, or Archephyceæ, Green Algæ, or Chlorophyceæ, Brown Algæ, or Phæophyceæ, and Red Algæ, or Rhodophyceæ.

The first class of Algæ, the Primæval Algæ (Archephyceæ), might also be called primæval plants, because they contain the simplest and most imperfect of all plants, and, among them, those most ancient of all vegetable organisms out of which all other plants have originated. To them therefore belong those most ancient of all vegetable Monera which arose by spontaneous generation in the beginning of the Laurentian period. Further, we have to reckon among them all those vegetable forms of the simplest organization which first developed out of the Monera in the Laurentian period, and which possessed the form of a single plastid. At first the entire body of one of these small primary plants consisted only of a most simple cytod (a plastid without kernel), and afterwards attained the higher form of a simple cell, by the separation of a kernel in the plasma. (Compare above, vol. i. p. [345].) Even at the present day there exist various most simple forms of Algæ which have deviated but little from the original primary plants. Among them are the Algæ of the families Codiolaceæ, Protococcaceæ, Desmidiaceæ, Palmellaceæ, Hydrodictyeæ, and several others. The remarkable group of Phycochromaceæ (Chroococcaceæ and Oscillarineæ) might also be comprised among them, unless we prefer to consider them as an independent tribe of the kingdom Protista.

The monoplastic Protophyta—that is, those primary Algæ formed by a single plastid—are of the greatest interest, because the vegetable organism in this case completes its whole course of life as a perfectly simple “individual of the first order,” either as a cytod without kernel, or as a cell containing a kernel.

Among the primary plants consisting of a single cytod are the exceedingly remarkable Siphoneæ, which are of considerable size, and strangely “mimic” the forms of higher plants. Many of the Siphoneæ attain a size of several feet, and resemble an elegant moss (Bryopsis), or in some cases a perfect flowering plant with stalks, roots, and leaves (Caulerpa) (Fig. 17). Yet the whole of this large body, externally so variously differentiated, consists internally of an entirely simple sack, possessing the negative characters of a simple cytod.

Fig. 17.—Caulerpa denticulata, a monoplastic Siphonean of the natural size. The entire branching primary plant, which appears to consist of a creeping stalk with fibrous roots and indented leaves, is in reality only a single plastid, and moreover a cytod (without a kernel), not even attaining the grade of a cell with nucleus.

These curious Siphoneæ, Vaucheriæ, and Caulerpæ show us to how great a degree of elaboration a single cytod, although a most simple individual of the first order, can develop by continuous adaptation to the relations of the outer world. Even the single-celled primary plants—which are distinguished from the monocytods by possessing a kernel—develop into a great variety of exquisite forms by adaptation; this is the case especially with the beautiful Desmidiaceæ, of which a species of Euastrum is represented in Fig. 18 as a specimen.

Fig. 18.—Euastrum rota, a single-celled Desmid, much enlarged. The whole of the star-shaped body of this primæval plant has the formal value of a simple cell. In its centre lies the kernel, and within this the kernel corpuscle, or speck.

It is very probable that similar primæval plants, the soft body of which, however, was not capable of being preserved in a fossil state, at one time peopled the Laurentian primæval sea in great masses and varieties, and in a great abundance of forms, without, however, going beyond the stage of individuality of a simple plastid.

The group of Green Tangles (Chlorophyceæ), or Green Algæ (Cloroalgæ), are the second class, and the most closely allied to the primæval group. Like the majority of the Archephyceæ, all the Chlorophyceæ are coloured green, and by the same colouring matter—the substance called leaf-green, or chlorophyll—which colours the leaves of all the higher plants.

To this class belong, besides a great number of low marine Algæ, most of the Algæ of fresh water, the common water hair-weeds, or Confervæ, the green slime-balls, or Glœosphæræ, the bright green water-lettuce, or Ulva, which resembles a very thin and long lettuce leaf, and also numerous small microscopic algæ, dense masses of which form a light green shiny covering to all sorts of objects lying in water—wood, stones, etc.

These forms, however, rise above the simple primary Algæ in the composition and differentiation of their body. As the green Algæ, like the primæval Algæ, mostly possess a very soft body, they are but rarely capable of being petrified. However, it can scarcely be doubted that this class of Algæ—which was the first to develop out of the preceding one—most extensively and variously peopled the fresh and salt waters of the earth in early times.

In the third class, that of the Brown Tangles (Phæophyceæ), or Black Algæ (Fucoideæ), the branch of the Algæ attains its highest stage of development, at least in regard to size and body. The characteristic colour of the Fucoid is more or less dark brown, sometimes tending more to an olive green or yellowish green, sometimes more to a brownish red or black colour.

Among these are the largest of all Algæ, which are at the same time the longest of all plants, namely, the colossal giant Algæ, amongst which the Macrocystis pyrifera, on the coast of California, attains a length of 400 feet. Also, among our indigenous Algæ, the largest forms belong to this group. Especially I may mention here the stately sugar-tangle (Laminaria), whose slimy, olive green thallus-body, resembling gigantic leaves of from 10 to 15 feet in length, and from a half to one foot in breadth, are thrown up in great masses on the coasts of the North and Baltic seas.

To this class belongs also the bladder-wrack (Fucus vesiculosus) common in our seas, whose fork-shaped, deeply-cut leaves are kept floating on the water by numerous air bladders (as is the case, too, with many other brown Algæ). The freely floating Sargasso Alga (Sargasso bacciferum), which forms the meadows or forests of the Sargasso Sea, also belongs to this class.

Although each individual of these large alga-trees is composed of many millions of cells, yet at the beginning of its existence it consists, like all higher plants, of a single cell—a simple egg. This egg—for example, in the case of our common bladder-wrack—is a naked, uncovered cell, and as such is so like the naked egg-cells of lower marine animals—for example, those of the Medusæ—that they might easily be mistaken one for another (Fig. 19).

Fig. 19.—The egg of the common bladder-wrack (Fucus vesiculosus), a simple naked cell, much enlarged. In the centre of the naked globule of protoplasm the bright kernel is visible.

It was probably the Fucoideæ, or Brown Algæ, which during the primordial period, to a great extent constituted the characteristic alga-forests of that immense space of time. Their petrified remains, especially those of the Silurian period, which have been preserved, can, it is true, give us but a faint idea of them, because the material of these Algæ, like that of most others, is ill-suited for preservation in a fossil state. As has already been remarked, a large portion of coal is perhaps composed of them.

Less important is the fourth class of Algæ, that of the Rose-coloured Algæ (Rhodophyceæ), or Red Sea-weeds (Florideæ). This class, it is true, presents a great number of different forms; but most of them are of much smaller size than the Brown Algæ. Although they are inferior to the latter in perfection and differentiation, they far surpass them in some other respects. To them belong the most beautiful and elegant of all Algæ, which on account of the fine plumose division of their leaf-like bodies, and also on account of their pure and delicate red colour, are among the most charming of plants. The characteristic red colour sometimes appears as a deep purple, sometimes as a glowing scarlet, sometimes as a delicate rose tint, and may verge into violet and bluish purple, or on the other hand into brown and green tints of marvellous splendour. Whoever has visited one of our sea-coast watering places, must have admired the lovely forms of the Florideæ, which are frequently dried on white paper and offered for sale.

Most of the Red Algæ are so delicate, that they are quite incapable of being petrified; this is the case with the splendid Ptilotes, Plocamia, Delesseria, etc. However, there are individual forms, like the Chondria and Sphærococca, which possess a harder thallus, often almost as hard as cartilage, and of these fossil remains have been preserved—principally in the Silurian, Devonian, and Carboniferous strata, and later in the oolites. It is probable that this class also had an important share in the composition of the archilithic Algæ flora.

If we now again take into consideration the flora of the primordial period, which was exclusively formed by the group of Algæ, we can see that it is not improbable that its four subordinate classes had a share in the composition of those submarine forests of the primæval oceans, similar to that which the four types of vegetation—trees with trunks, flowering shrubs, grass, and tender leaf-ferns and mosses—at present take in the composition of our recent land forests.

We may suppose that the submarine tree forests of the primordial period were formed by the huge Brown Algæ, or Fucoideæ. The many-coloured flowers at the foot of these gigantic trees were represented by the gay Red Algæ, or Florideæ. The green grass between was formed by the hair-like bunches of Green Algæ, or Chloroalgæ. Finally, the tender foliage of ferns and mosses, which at present cover the ground of our forests, fill the crevices left by other plants, and even settle on the trunks of the trees, at that time probably had representatives in the moss and fern-like Siphoneæ, in the Caulerpa and Bryopsis, from among the class of the primary Algæ, Protophyta, or Archephyceæ.

With regard to the relationships of the different classes of Algæ to one another and to other plants, it is exceedingly probable that the Primary Algæ, or Archephyceæ, as already remarked, form the common root of the pedigree, not merely for the different classes of Algæ, but for the whole vegetable kingdom. On this account they may with justice be designated as primæval plants, or Protophyta.

Out of the naked vegetable Monera, in the beginning of the Laurentian period, enclosed cytods were probably the first to arise (vol. i. p. [345]), by the naked, structureless, albuminous substance of the Monera becoming condensed in the form of a pellicle on the surface, or by secreting a membrane. At a later period, out of these enclosed cytods genuine vegetable cells probably arose, as a kernel or nucleus separated itself in the interior from the surrounding cell-substance or plasma.

The three classes of Green Algæ, Brown Algæ, and Red Algæ, are perhaps three distinct classes, which have arisen independently of one another out of the common radical group of Primæval Algæ, and then developed themselves further (each according to its kind), and have variously branched off into orders and families. The Brown and Red Algæ possess no close blood relationship to the other classes of the vegetable kingdom. These latter have most probably arisen out of the Primæval Algæ, either directly or by the intermediate step of the Green Algæ.

It is probable that Mosses (out of which, at a later time, Ferns developed) proceeded from a group of Green Algæ, and that Fungi and Lichens proceeded from a group of Primæval Algæ. The Phanerogamia developed at a much later period out of Ferns.

As a second class of the Vegetable Kingdom we have above mentioned the Thread-plants (Inophyta). We understood by this term the two closely related classes of Lichens and Fungi. It is possible that these Thallus plants have not arisen out of the Primæval Algæ, but out of one or more Monera, which, independently of the latter, arose by spontaneous generation. It appears conceivable that many of the lowest Fungi, as for example, many ferment-causing fungi (forms of Micrococcus, etc.), owe their origin to a number of different archigonic Monera (that is, Monera originating by spontaneous generation).

In any case the Thread-plants cannot be considered as the progenitors of any of the higher vegetable classes. Lichens, as well as fungi, are distinct from the higher plants in the composition of their soft bodies, consisting as it does of a dense felt-work of very long, variously interwoven, and peculiar threads or chains of cells—the so-called hyphæ, on which account we distinguish them as a province under the name Thread-plants. From their peculiar nature they could not leave any important fossil remains, and consequently we can form only a very vague guess at their palæontological development.

The first class of Thread-plants, the Fungi, exhibit a very close relationship to the lowest Algæ; the Algo-fungi, or Phycomycetes (the Saprolegniæ and Peronosporæ) in reality only differ from the bladder-wracks and Siphoneæ (the Vaucheria and Caulerpa) mentioned previously by the want of leaf-green, or chlorophyll. But, on the other hand, all genuine Fungi have so many peculiarities, and deviate so much from other plants, especially in their mode of taking food, that they might be considered as an entirely distinct province of the vegetable kingdom.

Other plants live mostly upon inorganic food, upon simple combinations which they render more complicated. They produce protoplasm by the combination of water, carbonic acid, and ammonia. They take in carbonic acid and give out oxygen. But the Fungi, like animals, live upon organic food, consisting of complicated combinations of carbon, which they receive from other organisms and assimilate. They inhale oxygen and give out carbonic acid like animals. They also never form leaf-green, or chlorophyll, which is so characteristic of most other plants. In like manner they never produce starch. Hence many eminent botanists have repeatedly proposed to remove the Fungi completely out of the vegetable kingdom, and to regard them as a special and third kingdom, between that of animals and plants. By this means our kingdom of Protista would be considerably increased. The Fungi in this case would, in the first place, be allied to the so-called “slime moulds,” or Myxomycetes (which, however, never form any hyphæ). But as many Fungi propagate in a sexual manner, and as most botanists, according to the prevalent opinion, look upon Fungi as genuine plants, we shall here leave them in the vegetable kingdom, and connect them with lichens, to which they are at all events most nearly related.

The phyletic origin of Fungi will probably long remain obscure. The close relationship already hinted at between the Phycomycetes and Siphoneæ (especially between the Saprolegniæ and Vaucheriæ) suggests to us that they are derived from the latter. Fungi would then have to be considered as Algæ, which by adaptation to a parasitical life have become very peculiarly transformed. Many facts, however, support the supposition that the lowest fungi have originated independently from archigonic Monera.

The second class of Inophyta, the Lichens (Lichenes), are very remarkable in relation to phylogeny; for the surprising discoveries of late years have taught us that every Lichen is really composed of two distinct plants—of a low form of Alga (Nostochaceæ, Chroococcaceæ), and of a parasitic form of Fungus (Ascomycetes), which lives as a parasite upon the former, and upon the nutritive substances prepared by it. The green cells, containing chlorophyll (gonidia), which are found in every lichen, belong to the Alga. But the colourless threads (hyphæ) which, densely interwoven, form the principal mass of the body of Lichens, belong to the parasitic Fungus. But in all cases the two forms of plants—Fungus and Alga—which are always considered as members of two quite distinct provinces of the vegetable kingdom, are so firmly united, and so thoroughly interwoven, that nearly every one looks upon a Lichen as a single organism.

Most Lichens form small, more or less formless or irregularly indented, crust-like coverings to stones, bark of trees, etc. Their colour varies through all possible tints, from the purest white to yellow, red, green, brown, and the deepest black.

Many lichens are important in the economy of nature from the fact that they can settle in the driest and most barren localities, especially on naked rocks upon which no other plant can live. The hard black lava, which covers many square miles of ground in volcanic regions, and which for centuries frequently presents the most determined opposition to the life of every kind of vegetation, is always first occupied by Lichens. It is the white or grey Lichens (Stereocaulon) which, in the most desolate and barren fields of lava, always begin to prepare the naked rocky ground for cultivation, and conquer it for subsequent higher vegetation. Their decaying bodies form the first mould in which mosses, ferns, and flowering plants can afterwards take firm root. Hardy Lichens are also less affected by the severity of climate than any other plants. Hence the naked rocks, even in the highest mountains—for the most part covered by eternal snow, on which no plant could thrive—are encrusted by the dry bodies of Lichens.

Leaving now the Fungi, Lichens, and Algæ, which are comprised under the name of Thallus plants, we enter upon the second sub-kingdom of the vegetable kingdom, that of the Prothallus plants (Prothallophyta), which by some botanists are called phyllogonic Cryptogamia (in contradistinction to the Thallus plants, or thallogonic Cryptogamia). This sub-kingdom comprises the two provinces of Mosses and Ferns.

Here we meet with (except in a few of the lowest forms) the separation of the vegetable body into two different fundamental organs, axial-organs (stem and root) and leaves (or lateral organs). In this the Prothallus plants resemble the Flowering plants, and hence the two groups have recently often been classed together as stemmed plants, or Cormophytes.

But, on the other hand, Mosses and Ferns resemble the Thallus plants, in the absence of the development of flowers and seeds, and even Linnæus classed them with these, as Cryptogamia, in contradistinction to the plants forming seeds; that is, flowering plants (Anthophyta or Phanerogamia).

Under the name of “Prothallus plants” we combine the closely-related Mosses and Ferns, because both exhibit a peculiar and characteristic “alternation of generation” in the course of their individual development. For every species exhibits two different generations, of which the one is usually called the Prothallium, or Fore-growth, the other is spoken of as the Cormus, or actual Stem of the moss or fern.

The first and original generation, the Fore-growth, or Prothallus, also called Protonema, still remains in that lower stage of elaboration manifested throughout life by all Thallus plants; that is to say, stem and leaf-organs have as yet not differentiated, and the entire cell-mass of the Fore-growth corresponds to a simple thallus. The second and more perfect generation of mosses and ferns—the Stem, or Cormus—develops a much more highly elaborate body, which has differentiated into stalk and leaf (as in the case of flowering plants), except in the lowest mosses, where this generation also remains in the lower stage of the thallus.

With the exception of these latter forms the first generation of Mosses and Ferns (the thallus-shaped Fore-growth) always produces a second generation with stem and leaves; the latter in its turn produces the thallus of the first generation, and so on. Thus, in this case, as in the ordinary cases of alternation of generation in animals, the first generation is like the third, fifth, etc., the second like the fourth, sixth, etc. (Compare vol. i. p. [206]).

Of the two main classes of Prothallus plants, the Mosses in general are at a much lower stage of development than the Ferns, and their lowest forms (especially in an anatomical respect) form the transition from the Thallus plants through the Algæ to Ferns. The genealogical connection of Mosses and Ferns which is indicated by this fact can, however, be inferred only from the case of the most imperfect forms of the two classes; for the more perfect and higher groups of mosses and ferns do not stand in any close relation to one another, and develop in completely opposite directions. In any case Mosses have arisen directly out of Thallus plants, and probably out of Green Algæ.

Ferns, on the other hand, are probably derived from extinct unknown Mosses, which were very nearly related to the lowest liverworts of the present day. In the history of creation, Ferns are of greater importance than Mosses.

The branch of Mosses (Muscinæ, also called Musci, or Bryophyta) contains the lower and more imperfect plants of the group of Prothallophytes, which as yet do not possess vessels. Their bodies are mostly so tender and perishable that they are very ill-suited for being preserved in a recognizable state as fossils. Hence the fossil remains of all classes of Mosses are rare and insignificant. It is probable that Mosses developed in very early times out of the Thallus plants, or, to be more precise, out of the Green Algæ. It is probable that in the primordial period there existed aquatic forms of transition from the latter to Mosses, and in the primary period to those living on land. The Mosses of the present day—out of the gradually differentiating development of which comparative anatomy may draw some inferences as to their genealogy—are divided into two different classes, namely: (1) Liverworts; (2) Leafy Mosses.

The first and oldest class of Mosses, which is directly allied to the Green Algæ, or Confervæ, is formed by the Liverworts (Hepaticæ, or Thallobrya). The mosses belonging to them are, for the most part, small and insignificant in form, and are little known. Their lowest forms still possess, in both generations, a simple thallus like the Thallus plants; as for example, the Ricciæ and Marchantiaceæ. But the more highly developed liverworts, the Jungermanniaceæ and those akin to them, gradually commence to differentiate stem and leaf, and their most highly-developed forms are closely allied to leaf-mosses. By this transitional series the liverworts show their direct derivation from the Thallophytes, and more especially from the Green Algæ.

The Mosses, which are generally the only ones known to the uninitiated—and which, in fact, form the principal portion of the whole branch—belong to the second class, or Leafy Mosses (Musci frondosi, called Musci in a narrow sense, also Phyllobrya). Among them are most of those pretty little plants which, united in dense groups, form the bright glossy carpet of moss in our woods, or which, in company with liverworts and lichens, cover the bark of trees. As reservoirs, carefully storing up moisture, they are of the greatest importance in the economy of nature. Wherever man mercilessly cuts down and destroys forests, there, as a consequence, disappear the leafy mosses which covered the bark of the trees, or, protected by their shade, clothed the ground, and filled the spaces between the larger plants. Together with the leafy mosses disappear the useful reservoirs which stored up rain and dew for times of drought. Thus arises a disastrous dryness of the ground, which prevents the growth of any rich vegetation. In the greater part of Southern Europe—in Greece, Italy, Sicily, and Spain—mosses have been destroyed by the inconsiderate extirpation of forests, and the ground has thereby been robbed of its most useful stores of moisture; once flourishing and rich tracts of land have been changed into dry and barren wastes. Unfortunately in Germany, also, this rude barbarism is beginning to prevail more and more. It is probable that the small frondose mosses have played this exceedingly important part in nature for a very long time, possibly from the beginning of the primary period. But as their tender bodies are as little suited as those of all other mosses for being preserved in a fossil state, palæontology can give us no information about this.

We learn from the science of petrifactions much more than we do in the case of Mosses of the importance which the second branch of Prothallus plants—that is, Ferns—have had in the history of the vegetable world. Ferns, or more strictly speaking, the “plants of the fern tribe” (Filicineæ, or Pterideæ, also called Pteridophyta, or Vascular Cryptogams), formed during an extremely long period, namely, during the whole primary or palæolithic period, the principal portion of the vegetable world, so that we may without hesitation call it the era of Fern Forests. From the beginning of the Devonian period, in which organisms living on land appeared for the first time, namely, during the deposits of the Devonian, Carboniferous, and Permian strata, plants like Ferns predominated so much over all others, that we are justified in giving this name to that period. In the stratifications just mentioned, but above all, in the immense layers of coal of the Carboniferous or coal period, we find such numerous and occasionally well preserved remains of Ferns, that we can form a tolerable vivid picture of the very peculiar land flora of the palæolithic period. In the year 1855 the total number of the then known palæolithic species of plants amounted to about a thousand, and among these there were no less than 872 Ferns. Among the remaining 128 species were 77 Gymnosperms (pines and palm-ferns), 40 Thallus plants (mostly Algæ), and about 20 not accurately definable Cormophyta (stem-plants).

As already remarked, Ferns probably developed out of the lower liverworts in the beginning of the primary period. In their organization Ferns rise considerably above Mosses, and in their more highly developed forms even approach the flowering plants. In Mosses, as in Thallus plants, the entire body is composed of almost equi-formal cells, little if at all differentiated; but in the tissues of Ferns we find those peculiarly differentiated strings of cells which are called the vessels of plants, and which are universally met with in flowering plants. Hence Ferns are sometimes united as “vascular Cryptogams” with Phanerogams, and the group so formed is contrasted as that of the “vascular plants” with “cellular plants,”—that is, with “cellular cryptogams” (Mosses and Thallus plants). This very important process in the organization of plants—the formation of vessels—first occurred, therefore, in the Devonian period, consequently in the beginning of the second and smaller half of the organic history of the earth.

The branch of Ferns, or Filicinæ, is divided into five distinct classes: (1) Frondose Ferns, or Pteridæ; (2) Reed Ferns, or Calamariæ; (3) Aquatic Ferns, or Rhizocarpeæ; (4) Snakes Tongues, or Ophioglossæ; and (5) Scale Ferns, or Lepidophyta. By far the most important of these five classes, and also the richest in forms, were first the Frondose Ferns, and then the Scale-ferns, which formed the principal portion of the palæolithic forests. The Reed Ferns, on the other hand, had at that time already somewhat diminished in number; and of the Aquatic Ferns, we do not even know with certainty whether they then existed. It is difficult for us to form any idea of the very peculiar character of those gloomy palæolithic fern forests, in which the whole of the gay abundance of flowers of our present flora was entirely wanting, and which were not enlivened by any birds. Of the flowering plants there then existed only the two lowest classes, the pines and palm ferns, with naked seeds, whose simple and insignificant blossoms scarcely deserve the name of flowers.

The phylogeny of Ferns, and of the Gymnosperms which have developed out of them, has been made especially clear by the excellent investigations which Edward Strasburger published in 1872, on “The Coniferæ and Gnetaceæ,” as also “On Azolla.” This thoughtful naturalist and Charles Martins, of Montpellier, are among the few botanists who have thoroughly understood the fundamental value of the Theory of Descent, and the mechanical-causal connection between ontogeny and phylogeny. The majority of botanists do not even yet know the important difference between homology and analogy, between the morphological and physiological comparison of parts—which has long since been recognized in zoology—but Strasburger has employed this distinction and the principle of evolution in his “Comparative Anatomy of the Gymnosperms,” in order to sketch the outlines of the blood relationship of this important group of plants.

The class among Ferns which has developed most directly out of the Liverworts is the class of real Ferns, in the narrow sense of the word, the Frondose Ferns (Filices, or Phyllopterides, also called Pteridæ). In the present flora of the temperate zones this class forms only a subordinate part, for it is in most cases represented only by low forms without trunks. But in the torrid zones, especially in the moist, steaming forests of tropical regions, this class presents us with the lofty palm-like fern trees. These beautiful tree-ferns of the present day, which form the chief ornament of our hot-houses, can however give us but a faint idea of the stately and splendid frondose ferns of the primary period, whose mighty trunks, densely crowded together, then formed entire forests. These trunks, accumulated in super-incumbent masses, are found in the coal seams of the Carboniferous period, and between them, in an excellent state of preservation, are found the impressions of the elegant fan-shaped leaves, crowning the top of the trunk in an umbrella-like bush. The varied outlines and the feather-like forms of these fronds, the elegant shape of the branching veins or bunches of vessels in their tender foliage, can still be as distinctly recognized in the impressions of the palæolithic fronds as in the fronds of ferns of the present day. In many cases even the clusters of fruit, which are distributed on the lower surface of the fronds, are distinctly preserved. After the Carboniferous period, the predominance of frondose ferns diminished, and towards the end of the secondary period they played almost as subordinate a part as they do at the present time.

The Calamariæ, Ophioglossæ, and Rhizocarpeæ seem to have developed as three diverging branches out of the Frondose Ferns, or Pteridæ. The Calamariæ, or Calamophyta, have remained at the lowest level among these three classes. The Calamariæ comprise three different orders, of which only one now exists, namely, the Horse-tails (Equisetaceæ). The two other orders, the Giant Reeds (Calamiteæ), and the Star-leaf Reeds (Asterophylliteæ), are long since extinct. All Calamariæ are characterized by a hollow and jointed stalk, stem, or trunk, upon which the branches and leaves (in cases where they exist) are set so as to encircle the jointed stem in whorls. The hollow joints of the stalk are separated from one another by partition walls. In Horse-tails and Calamiteæ the surface is traversed by longitudinal ribs running parallel, as in the case of a fluted column, and the outer skin contains so much silicious earth in the living forms, that it is used for cleansing and polishing. In the Asterophylliteæ, the star-shaped whorls of leaves were more strongly developed than in the two other orders. There exist, at present, of the Calamariæ only the insignificant Horse-tails (Equisetum), which grow in marshes and on moors; but during the whole of the primary and secondary periods they were represented by great trees of the genus Equisetites. There existed, at the same time, the closely related order of the Giant Reeds (Calamites), whose strong trunks grew to a height of about fifty feet. The order of the Asterophyllites, on the other hand, contained smaller and prettier plants, of a very peculiar form, and belongs exclusively to the primary period.

Among all Ferns, the history of the third class, that of the Root, or Aquatic Ferns (Rhizocarpeæ, or Hydropteridæ), is least known to us. In their structure these ferns, which live in fresh water, are on the one hand allied to the frond ferns, and on the other to the scaly ferns, but they are more closely related to the latter. Among them are the but little known moss ferns (Salvinia), clover ferns (Marsilea), and pill ferns (Pilularia) of our fresh waters; further, the large Azolla which floats in tropical ponds. Most of the aquatic ferns are of a delicate nature, and hence ill-suited for being petrified. This is probably the reason of their fossil remains being so scarce, and of the oldest of those known to us having been found in the Jura system. It is probable, however, that the class is much older, and that it was already developed during the palæolithic period out of other ferns by adaptation to an aquatic life.

The fourth class of ferns is formed by the Tongue Ferns (Ophioglossæ, or Glossopterides). These ferns, to which belongs the Botrychium, as well as the Ophioglossum (adder’s-tongue) of our native genera, were formerly considered as forming but a small subdivision of the frondose ferns. But they deserve to form a special class, because they represent important transitional forms from the Pterideæ and Lepidophytes towards higher plants, and must be regarded as among the direct progenitors of the flowering plants.

The fifth and last class is formed by the Scale Ferns (Lepidophytes, or Selagines). In the same way as the Ophioglossæ arose out of the frondose forms, the scale ferns arose out of the Ophioglossæ. They were more highly developed than all other ferns, and form the transition to flowering plants, which must have developed out of them. Next to the frondose ferns they took the largest part in the composition of the palæolithic fern forests. This class also contains, as does the class of reed ferns, three nearly related but still very different orders, of which only one now exists, the two others having become extinct towards the end of the Carboniferous period. The scaled ferns still existing belong to the order of the club-mosses (Lycopodiaceæ). They are mostly small, pretty moss-like plants, whose tender, many-branched stalk creeps in curves on the ground like a snake, and is densely encompassed and covered by small scaly leaves. The pretty creeping Lycopodium of our woods, which mountain tourists twine round their hats, is known to all, as also the still more delicate Selaginella, which under the name of creeping moss is used to adorn the soil of our hot-houses in the form of a thick carpet. The largest club-mosses of the present day are found in the Sunda Islands, where their stalks rise to the height of twenty-five feet, and attain half a foot in thickness. But in the primary and secondary periods even larger trees of this kind were widely distributed, the most ancient of which probably were the progenitors of the pines (Lycopodites). The most important dimensions were, however, attained by the class of scale trees (Lepidodendreæ), and by the seal trees (Sigillarieæ). These two orders, with a few species, appear in the Devonian period, but do not attain their immense and astonishing development until the Carboniferous period, and become extinct towards the end of it, or in the Permian period directly following upon it. The scale trees, or Lepidodendreæ, were probably more closely related to club-mosses than to Sigillarieæ. They grew into splendid, straight, unbranching trunks which divided at the top into numerous forked branches. They bore a large crown of scaly leaves, and like the trunk were marked in elegant spiral lines by the scars left at the base of the leaf stalks which had fallen off. We know of scale-marked trees from forty to sixty feet in length, and from twelve to fifteen feet in diameter at the root. Some trunks are said to be even more than a hundred feet in length. In the coal are found still larger accumulations of the no less highly developed but more slender trunks of the remarkable seal trees, Sigillarieæ, which in many places form the principal part of coal seams. Their roots were formerly described as quite a distinct vegetable form (under the name of Stigmaria). The Sigillarieæ are in many respects very like the scale-trees, but differ from them and from ferns in general in many ways. They were possibly closely related to the extinct Devonian Lycopterideæ, combining characteristic peculiarities of the club-mosses and the frondose ferns, which Strasburger considers as the hypothetical primary form of flowering plants.

In leaving the dense forests of the primary period, which were principally composed of frond ferns (Lepidodendreæ and Sigillarieæ), we pass onwards to the no less characteristic pine forests of the secondary period. Thus we leave the domain of the Cryptogamia, the plants forming neither flowers nor seeds, and enter the second main division of the vegetable kingdom, namely, the sub-kingdom of the Phanerogamia, flowering plants forming seeds. This division, so rich in forms, containing the principal portion of the present vegetable world, and especially the majority of plants living on land, is certainly of a much more recent date than the division of Cryptogamia. For it can have developed out of the latter only in the course of the palæolithic period. We can with full assurance maintain that, during the whole archilithic period, hence during the first and longer half of the organic history of the earth, no flowering plants as yet existed, and that they first developed during the primary period out of Cryptogamia of the fern kind. The anatomical and embryological relation of Phanerogamia to the latter is so close, that from it we can with certainty infer their genealogical connection, that is, their true blood relationship. Flowering plants cannot have directly arisen out of thallus plants, nor out of mosses; but only out of ferns, or Filicines. Most probably the scaled ferns, or Lepidophyta, and more especially amongst these the Lycopodiaceæ, forms closely related to the Selaginella of the present day, have been the direct progenitors of the Phanerogamia.

On account of its anatomical structure and its embryological development, the sub-kingdom of the Phanerogamia has for a long time been divided into two large branches, into the Gymnosperms, or plants with naked seeds, and the Angiosperms, or plants with enclosed seeds. The latter are in every respect more perfect and more highly organized than the former, and developed out of them only at a late date during the secondary period. The Gymnosperms, both anatomically and embryologically, form the transition group from Ferns to Angiosperms.

The lower, more imperfect, and the older of the two main classes of flowering plants, that of the Archispermeæ, or Gymnosperms (with naked seeds), attained its most varied development and widest distribution during the mesolithic or secondary epoch. It was no less characteristic of this period, than was the fern group of the preceding primary, and the Angiosperms of the succeeding tertiary, epoch. Hence we might call the secondary epoch that of Gymnosperms, or after its most important representatives, the era of Pine Forests. The Gymnosperms are divided into three classes: the Coniferæ, Cycadeæ, and Gnetaceæ. We find fossil remains of the pines, or Conifers, and of the Cycads, even in coal, and must infer from this that the transition from scaled ferns to Gymnosperms took place during the Coal, or possibly even in the Devonian period. However, the Gymnosperms play but a very subordinate part during the whole of the primary epoch, and do not predominate over Ferns until the beginning of the secondary epoch.

Of the two classes of Gymnosperms just mentioned, that of the Palm Ferns (Zamiæ, or Cycadeæ) stands at the lowest stage, and is directly allied to ferns, as the name implies, so that some botanists have actually included them in the fern group. In their external form they resemble palms, as well as tree ferns (or tree-like frond ferns), and are adorned by a crown of feathery leaves, which is placed either on a thick, short trunk, or on a slender, simple trunk like a pillar. At the present day this class, once so rich in forms, is but scantily represented by a few forms living in the torrid zones, namely, by the coniferous ferns (Zamia), the thick-trunked bread-tree (Encephalartos), and the slender-trunked Caffir bread-tree (Cycas). They may frequently be seen in hot-houses, and are generally mistaken for palms. A much greater variety of forms than occurs among the still existing palm ferns (Cycadeæ) is presented by the extinct and fossil Cycads, which occurred in great numbers more towards the middle of the secondary period, during the Jura, and which at that time principally determined the character of the forests.

The class of Pines, or coniferous trees (Coniferæ), has preserved down to our day a greater variety of forms than have the palm ferns. Even at the present time the trees belonging to it—cypresses, juniper trees, and trees of life (Thuja), the box and ginko trees (Salisburya), the araucaria and cedars, but above all the genus Pinus, which is so rich in forms, with its numerous and important species, spruces, pines, firs, larches, etc.—still play a very important part in the most different parts of the earth, and almost of themselves constitute extensive forests. Yet this development of pines seems but weak in comparison with the predominance which the class had attained over other plants during the early secondary period, that of the Trias. At that time mighty coniferous trees—with but proportionately few genera and species, but standing together in immense masses of individuals—formed the principal part of the mesolithic forests. This fact justifies us in calling the secondary period the “era of the pine forests,” although the remains of Cycadeæ predominate over those of coniferous trees in the Jura period.[2]

From the pine forests of the mesolithic, or secondary period, we pass on into the leafy forests of the cænolithic, or tertiary period, and we arrive thus at the consideration of the sixth and last class of the vegetable kingdom, that of the Metaspermæ, Angiospermæ, or plants with enclosed seeds. The first certain and undoubted fossils of plants with enclosed seeds are found in the strata of the chalk system, and indeed we here find, side by side, remains of the two classes into which the main class of Angiosperms is generally divided, namely, the one seed-lobed plants, or monocotylæ, and the two seed-lobed plants, or dicotylæ. However, the whole group probably originated at an earlier period during the Trias. For we know of a number of doubtful and not accurately definable fossil remains of plants from the Oolitic and Trias (sic) periods, which some botanists consider to be Monocotylæ, whilst others consider them as Gymnosperms. In regard to the two classes of plants with enclosed seeds, the Monocotylæ and Dicotylæ, it is exceedingly probable that the Dicotyledons developed out of the Gnetaceæ, but that the Monocotyledons developed later out of a branch of the dicotyledons.

The class of one seed-lobed plants (Monocotylæ, or Monocotyledons, also called Endogenæ) comprises those flowering plants whose seeds possess but one germ leaf or seed lobe (cotyledon). Each whorl of its flower contains in most cases three leaves, and it is very probable that the mother plants of all Monocotyledons possessed a regular triple blossom. The leaves are mostly simple, and traversed by simple, straight bunches of vessels or “nerves.” To this class belong the extensive families of the rushes, grasses, lilies, irids, and orchids, further a number of indigenous aquatic plants, the water-onions, sea grasses, etc., and finally the splendid and highly developed families of the Aroideæ and Pandaneæ, the bananas and palms. On the whole, the class of Monocotyledons—in spite of the great variety of forms which it developed, both in the tertiary and the present period—is much more simply organized than the class of the Dicotyledons, and its history of development also offers much less of interest. As their fossil remains are for the most part difficult to recognize, it still remains at present an open question in which of the three great secondary periods—the Trias, Jura, or chalk period—the Monocotyledons originated. At all events they existed in the chalk period as surely as did the Dicotyledons.

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?

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).