Part 2

A classification according to structure goes then a long way towards mapping out the main lines of the family-tree of organisms. We are further assisted in the task by the fossil remains of extinct organisms which sometimes give to us the actual ancestors of forms now living. But the most remarkable aid to the correct building-up of the pedigree of _animals_ at least (and the remarks which follow are confined to that division of the organic world), is afforded by the changes—the phases of development—which every animal exhibits in passing from the small shapeless egg to the adult condition. The aid which we here obtain depends on the following facts. Just as we suppose any one animal—say a dog—to have developed by slow change through an immense series of ancestors which become simpler and simpler as we recede into the past until we reach a small shapeless lump of living matter devoid of structure, so do we find actually as a matter of fact, which any one can see for himself, that every individual animal begins its individual life as a structureless particle which is thrown off from its parent, and is known as the _egg-cell_ (Fig. 1). Gradually passing through a series of more and more elaborated conditions of structure, that egg grows into the adult dog. The changes which have taken countless ages in the one case, are accomplished in a few weeks in the other.

And now we have to note the important fact which makes this process of development so intensely interesting in relation to the pedigree of the animal kingdom. There is very strong reason to believe that it is a general law of transmission or inheritance, that structural characteristics appear in the growth of a young organism in the order in which those characteristics have been acquired by its ancestors. At first the egg of a dog represents (imperfectly, it is true,) in form and structure the earliest ancestors of the dog; a few days later it has the form and structure of somewhat later ancestors; later still the embryo dog resembles less remote ancestors; until at last it reaches the degree of elaboration proper to its immediate forefathers.

Accordingly the phases of development or growth of the young are a brief recapitulation of the phases of form through which the ancestors of the young creature have passed. In some animals this recapitulation is more, in others it is less complete. Sometimes the changes are hurried through and disguised, but we find here and there in these histories of growth from the egg most valuable assistance in the attempt to reconstruct the genealogical tree. The history of the development of the common frog is a good illustration of the _kind_ of evidence in question.

The frog’s egg first gives rise to a little aquatic creature with external gills and a tail—the tadpole—which gradually loses its gills and its tail and acquires in their place lungs and four legs (Fig. 2), so as now to be fitted for life on dry land. From what we otherwise know of the structure of the frog and the animals to which it is allied, we are justified in concluding that the tadpole is a recapitulative phase of development, and represents to us more or less closely an ancestor of the frog which was provided with gills and tail in the adult state, and possessed neither legs nor lungs.

A less familiar case is that of a certain kind of shrimp, which is illustrated in the wood-cuts (Fig. 3 and right lower corner of Fig. 4). The little creature which issues from the egg of this shrimp is known as the “Nauplius form.” Many animals very different in appearance from this shrimp make their first appearance in the world as Nauplii; and it appears probable that the Nauplius-phase is the recapitulative re-presentation of an ancestor common to all this set of animals, an ancestor which was not _exactly_ like the Nauplius, but not very different from it.

The Nauplius of our shrimp gradually elongates. At first it has but three pair of limbs, but it soon acquires additional pairs, and a jointed body, and thus by gradually adding to its complexity of structure as seen in Figs. 5 and 6, it approximates more and more to the adult form from the egg of which it originated.

And now we are approaching the main point to which I wish to draw the reader’s attention. In attempting to reconstruct the pedigree of the animal kingdom and so to exhibit correctly the genetic relationships of all existing forms of animals, naturalists have hitherto assumed that the process of natural selection and survival of the fittest has invariably acted so as either to improve and elaborate the structure of _all_ the organisms subject to it, or else has left them unchanged, exactly fitted to their conditions, maintained as it were in a state of _balance_. It has been held that there have been some six or seven great lines of descent—main branches of the pedigree—such as that of the Vertebrates, that of the Molluscs, that of the Insects, that of the Starfish, and so on; and that along each of these lines there has been always and continuously a progress—a change in the direction of greater elaboration.

Each of these great branches of the family-tree is held to be independent—they all branch off nearly simultaneously from the main trunk like the leading branches of an oak. The animal forms constituting the series in each of these branches are supposed to gradually increase in elaboration of structure as we pass upwards from the main trunk of origin and climb further and further towards the youngest, most recent twigs. New organs have, it is supposed, been gradually developed in each series, giving their possessors greater powers, enabling them to cope more successfully with others in that struggle for existence in virtue of which these new organs have been little by little called into being. At the same time _here and there_ along the line of march, certain forms have been supposed to have “fallen out,” to have ceased to improve, and being happily fitted to the conditions of life in which they were long ago existing, have continued down to the present day to exist in the same low, imperfect condition. It is in this way that the lowest forms of animal life at present existing are usually explained, such as the microscopic animalcules, Amœbæ and Infusoria. It is in this way that the lower or more simply-made families of higher groups have been generally regarded. The simpler living Mollusca or shellfish have been supposed necessarily to represent the original forms of the great race of Mollusca. The simpler Vertebrates have been supposed necessarily to represent the original Vertebrates. The simpler Worms have been supposed necessarily to be the stereotyped representatives of very ancient Worms.

That this is, to a certain extent, a true explanation of the existence at the present day of _low_ forms of animals is proved by the fact that we find in very ancient strata fossil remains of animals which differ, ever so little, from particular animals existing at the present day; for instance, the Brachiopods (lamp-shells), Lingula and Terebratula, the King-crabs, and the Pearly Nautilus are found living at the present day, and are also found with no appreciable difference in very ancient strata of the earth’s crust; strata deposited so long ago that most of the forms of life at present inhabiting the earth’s surface had not then been brought into existence, whilst other most strange and varied forms occupied their place, and have now for long ages been extinct.

Whilst we are thus justified by the direct testimony of fossil remains in accounting for _some_ living forms on the hypothesis that their peculiar conditions of life have been such as to maintain them for an immense period of time _in statu quo_ unchanged, _we have no reason for applying this hypothesis, and this only_, to the explanation of all the more imperfectly organised forms of animal or plant-life.

It is clearly enough possible for a set of forces such as we sum up under the head “natural selection” to so act on the structure of an organism as to produce one of three results, namely these; to keep it _in statu quo_; to increase the complexity of its structure; or lastly, to diminish the complexity of its structure. We have as possibilities either BALANCE, or ELABORATION, or DEGENERATION.

Owing, as it seems, to the predisposing influence of the systems of classification in ascending series proceeding steadily upwards from the “lower” or simplest forms to the “higher” or more complex forms,—systems which were prevalent before the doctrine of transformism had taken firm root in the minds of naturalists, there has been up to the present day an endeavour to explain every existing form of life on the hypothesis that it has been maintained for long ages in a state of Balance; or else on the hypothesis that it has been Elaborated, and is an advance, an improvement, upon its ancestors. Only one naturalist—Dr. Dohrn, of Naples—has put forward the hypothesis of Degeneration as capable of wide application to the explanation of existing forms of life;[4] and his arguments in favour of a general application of this hypothesis have not, I think, met with the consideration which they merit.

The statement that the hypothesis of Degeneration has not been recognised by naturalists generally as an explanation of animal forms, requires to be corrected by the exception of certain kinds of animals, namely, those that are parasitic or quasi-parasitic. With regard to parasites, naturalists have long recognized what is called _retrogressive metamorphosis_; and parasitic animals are as a rule admitted to be instances of Degeneration. It is the more remarkable whilst the possibility of a degeneration—a loss of organisation making the descendant far _simpler_ or _lower_ in structure than its ancestor—has been admitted for a few exceptional animals, that the same hypothesis should not have been applied to the explanation of other simple forms of animals. The hypothesis of Degeneration will, I believe, be found to render most valuable service in pointing out the true relationships of animals which are a puzzle and a mystery when we use only and exclusively the hypothesis of Balance, or the hypothesis of Elaboration. It will, as a true scientific hypothesis, help us to discover causes.

We may now examine a few examples of undeniably degenerate animals, and first, I may call to mind the very remarkable series of lizard-like animals which exist in the south of Europe and in other countries, which exhibit in closely related genera a gradual loss of the limbs—a local or limited Degeneration. We have the common Lizard (_Lacerta_), with five toes on each of its well-grown fore and hind limbs; then we have side by side with this a lizard-like creature, _Seps_, in which both pairs of limbs have become ridiculously small, and are evidently ceasing to be useful in the way in which those of _Lacerta_ are useful; and lastly, we have _Bipes_, in which the anterior pair of limbs has altogether vanished, and only a pair of stumps, representing the hinder limbs, remain.

No naturalist doubts that Seps and Bipes represent two stages of Degeneration, or atrophy of the limbs; that they have, in fact, been derived from the five-toed four-legged form, and have _lost_ the locomotor organs once possessed by their ancestors. This very partial or local atrophy is not however that to which I refer when using the word Degeneration. Let us imagine this atrophy to extend to a variety of important organs, so that not only the legs, but the organs of sense, the nervous system, and even the mouth and digestive organs are obliterated—then we shall have pictured a thorough-going instance of Degeneration.

Degeneration may be defined as a gradual change of the structure in which the organism becomes adapted to _less_ varied and _less_ complex conditions of life; whilst Elaboration is a gradual change of structure in which the organism becomes adapted to more and more varied and complex conditions of existence. In Elaboration there is a new _expression_ of form, corresponding to new perfection of work in the animal machine. In Degeneration there is _suppression_ of form, corresponding to the cessation of work. Elaboration of some one organ _may_ be a necessary accompaniment of Degeneration in all the others; in fact, this is very generally the case; and it is only when the total result of the Elaboration of some organs, and the Degeneration of others, is such as to leave the whole animal in a _lower_ condition, that is, fitted to less complex action and reaction in regard to its surroundings, than was the ancestral form with which we are comparing it (either actually or in imagination) that we speak of that animal as an instance of Degeneration.

Any new set of conditions occurring to an animal which render its food and safety very easily attained, seem to lead as a rule to Degeneration; just as an active healthy man sometimes degenerates when he becomes suddenly possessed of a fortune; or as Rome degenerated when possessed of the riches of the ancient world. The habit of parasitism clearly acts upon animal organisation in this way. Let the parasitic life once be secured, and away go legs, jaws, eyes, and ears; the active, highly-gifted crab, insect, or annelid may become a mere sac, absorbing nourishment and laying eggs.

Reference was made above to the larval stage of a certain shrimp (Figs. 4, 5, 6). Let us now compare these with the young stages of a number of shrimp-like animals, viz., Sacculina, Lernæocera, Lepas, Cyclops, Limnetis, (all drawn in Fig. 4), some of which lead a parasitic life. All start equally with the recapitulative phase known as the Nauplius; but whilst the Nauplius of the free-living shrimp grows more and more elaborate, observe what happens to the parasites—they degenerate into comparatively simple bodies; and this is true of their internal structure as well as of their external appearance. The most utterly reduced of these parasites is the curious Sacculina (Fig. 7) which infests Hermit-crabs, and is a mere sac filled with eggs, and absorbing nourishment from the juices of its host by root-like processes.

Lernæocera again, which in the adult condition is found attached to the gills of fishes, has lost the well-developed legs of its Nauplius childhood and become an elongated worm-like creature (Fig. 8), fitted only to suck in nourishment and carry eggs.

Amongst these Nauplii—all belonging to the great group Crustacea, which includes crabs and shrimps—is one which gives rise to an animal decidedly degenerate, but not precisely parasitic in its habits. This Nauplius is the young of the ship’s Barnacle, a curious stalked body, inclosed in a shell of many pieces (Fig. 9). The egg of the Barnacle gives rise to an actively swimming Nauplius, the history of which is very astonishing. After swimming about for a time the Barnacle’s Nauplius fixes its head against a piece of wood, and takes to a perfectly fixed, immobile state of life (Fig. 10). The upper figures represent the Nauplius stage of animals closely resembling the Barnacle: the lower figures show the transformation of the Nauplius into the young Barnacle. Its organs of touch and of sight atrophy, its legs lose their locomotor function, and are simply used for bringing floating particles to the orifice of the stomach; so that an eminent naturalist has compared one of these animals to a man standing on his head and kicking his food into his mouth.

Were it not for the recapitulative phases in the development of the Barnacle, we may doubt whether naturalists would _ever_ have guessed that it was a degenerate Crustacean. It was in fact for a long time regarded as quite remote from them, and placed among the snails and oysters; its true nature was only admitted when the young form was discovered.

Other parasitic organisms, which exhibit extreme degeneration as compared with their free-living relatives, might be cited and figured in profusion, did our limits permit. Very noteworthy are the degenerate Spiders—the mites, leading to still more degenerate forms, the Linguatulæ.

We have two of these represented in Figs. 11 and 12. The one (Fig. 11), as compared with a spider is seen still to possess the eight walking legs, small, it is true, whilst the palps and daggers of the spider have dwindled to a beak projecting from the front of the globular unjointed body. In the other the eight legs have become mere stumps, and the body is elongated like that of a worm.

The instances of degeneration which we have so far examined are due to parasitism, except in the example of the Barnacle, where we have an instance of degeneration due to sessile and immobile habit of life. We may now proceed to look at some sessile or immobile animals which are not usually regarded as degenerate, but which, I think, there is every reason to believe are the degenerate descendants of very much higher and more elaborate ancestors. These are certain marine animals, the Ascidians, or sea-squirts. These animals are found encrusting rocks, stones, and weeds on the sea bottom. Sometimes they are solitary (Fig. 13), but many of them produce buds, like plants, and so form compound masses or sheets of individuals all connected and continuous with one another, like the buds on a creeping plant (Fig. 14).

We will examine one of the simple forms—a tough mass like a leather bottle with two openings; water is continually passing in at the one and out at the other of these apertures. If we remove the leathery outer-case (Fig. 15), we find that there is a soft creature within which has the following parts:—Leading from the mouth a great throat, followed by an intestine. The throat is perforated by innumerable slits, through which the water passes into a chamber—the cloaca: in passing, the water aërates the blood which circulates in the framework of the slits. The intestine takes a sharp bend, which causes it to open also into the cloaca. Between the orifice of the mouth and of the cloaca there is a nerve-ganglion.

My object in the next place is to show that the structure and life-history of these Ascidians may be best explained on the hypothesis that they are instances of degeneration; that they are the modified descendants of animals of higher, that is more elaborate structure, and in fact are degenerate Vertebrata, standing in the same relation to fishes, frogs, and men, as do the barnacles to shrimps, crabs, and lobsters.

The young of some, but by no means of all these Ascidians, have a form totally different from that of their parents. The egg of Phallusia gives rise to a tadpole, a drawing of which placed side by side with the somewhat larger tadpole of the common frog is seen in the adjoining figure (Fig. 16). The young Ascidian has the same general shape as the young frog, but not only this; the resemblance extends into details, the internal organs agreeing closely in the two cases. Further still as shown by the beautiful researches of the Russian naturalist, Kowalewsky, the resemblance reaches absolute identity when we examine the way in which the various organs arise from the primitive egg-cell. Tail, body, spiracle, eye, and mouth, agree in the two tadpoles, the only important difference being in the position of the two mouths and in the fact that the Ascidian has one eye while the frog has two.

Now let us look at the internal organs (Fig. 17). There are four structures, which are _all four_ possessed at some time of their lives by all those animals which we call the Vertebrata, the great branch of the pedigree to which fishes, reptiles, birds, beasts, and men belong. And the combination of these marks or structural peculiarities is an overwhelming piece of evidence in favour of the supposition that the creatures which possess this combination are derived from one common ancestor. Just as one would conclude that a man whom one might meet, say on Salisbury Plain, _must_ belong to the New Zealand race, if it were found not only that he had the colour, and the hair, and the shape of head of a New Zealander, but also that he was tattooed like a New Zealander, carried the weapons of a New Zealander, and, over and above in addition to these proofs, that he talked the Maori language and none other; so here, in the case of the vertebrate race, there are certain qualities and possessions, the accumulation of which cannot be conceived of as occurring in any animal but one belonging to that race. These four great structural features are—first, the primitive backbone or notochord; second, the throat perforated by gill-slits; third, the tubular nerve-centre or spinal cord and brain placed along the back; and lastly, and perhaps most distinctive and clinching as an evidence of affinity, the myelonic or cerebral eye.

Now let us convince ourselves that these four features exist not only in the frog’s tadpole, as they do in all fishes, reptiles, birds, and beasts, but that they also exist in the Ascidian tadpole, and, it may be added, co-exist in no other animals at all.

The corresponding parts are named in Figs. 16 and 17, in such a way as to render their agreement tolerably clear, whilst in Fig. 18 a more detailed representation of the head of an Ascidian tadpole is given.