Facts and Arguments for Darwin by Fritz Muller (read along books .TXT) 📕

With regard to the Crabs, and also to the Isopoda and Amphipoda, we were led to the assumption that, about the period when these groups started from the common stem, a simplification of their process of development took place. This also seems to be intelligible from Darwin’s theory. When any circumstances favourable to a group of animals caused its wider diffusion and divergence into forms adapting themselves to new and various conditions of existence, this greater variability, which betrays itself in the production of new forms, will also favour the simplification of the development which is almost always advantageous, and moreover, exactly at this period, during adaptation to new circumstances, as has already been indicated with regard to fresh-water animals, this simplification will be doubly beneficial, and therefore, in connexion with this, a doubly strict selection will take place.

So much for the development of the higher Crustacea.

A closer examination of the developmental history of the lower Crustacea is unnecessary after what has been said in general upon the historical significance of the young states, and the application of this which has just been made to the Malacostraca. We may see, without further discussion, how the representation given by Claus of the development of the Copepoda may pass almost word for word as the primitive history of those animals; we may find in the Nauplius-skin of the larvae of Achtheres and in the egg-like larva of Cryptophialus, precisely similar traces of a transition towards direct development, as were presented by the Nauplius-envelope of the embryos of Mysis and the maggot-like larva of Ligia, etc.

It will be sufficient to indicate an essential difference in the process of development in the higher and lower Crustacea. In the latter all new body-segments and limbs which insert themselves between the two terminal regions of the Nauplius, are formed in uninterrupted sequence from before backwards; in the former there is further a new formation in the middle of the body (the middle-body), which pushes itself in between the fore-body and the abdomen in the same way, as these have done on their part between the head and tail of the Nauplius. Thus, that which appears probable even from the comparison of the limbs of the adult animal, finds fresh support in the developmental history, namely, that the lower Crustacea, like the Insects, are entirely destitute of the region of the body corresponding to the middle-body of the Malacostraca. It seems probable that the swimming feet of the Copepoda, as also of the pupae of Cirripedia and Rhizocephala, represent the abdominal feet of the Malacostraca, that is to say, are derived by inheritance from the same source with them.

It would be easy to weave together the separate threads furnished by the young forms of the various Crustacea, into a general picture of the primitive history of this class. Such a picture, drawn with a little skill, and finished in lively colours, would certainly be more attractive than the dry discussions which I have tacked on to the developmental history of these animals. But the mode of weaving in the loose threads would still in many cases be arbitrary, and to be effected with equal justice in various ways; and many gaps would still have to be filled up by means of more or less bold assumptions. Those who have not wandered much in this region of research would then readily believe that they were standing upon firm ground, where mere fancy had thrown an airy bridge; those acquainted with the subject, on the other hand, would soon find out these weak points in the structure, but would then be easily led to regard even what was founded upon well considered facts, as merely floating in the air. To obviate these misconceptions of its true contents from either side, it would be necessary to accompany such a picture throughout with lengthy, dry explanations. This has deterred me from further filling in the outline which I had already sketched.

I will only give, as an example, the probable history of the production of a single group of Crustacea, and indeed of the most abnormal of all, the RHIZOCEPHALA, which in the sexually mature state differ so enormously even from their nearest allies, the Cirripedia, and from their peculiar mode of nourishment stand quite alone in the entire animal kingdom.

I must preface this with a few words upon the homology of the roots of the Rhizocephala, i.e. the tubules which penetrate from its point of adhesion into the body of the host, ramify amongst the viscera of the latter, and terminate in caecal branchlets. In the pupae of the Rhizocephala (Figure 58) the foremost limbs (“prehensile antennae”) bear, on each of the two terminal joints, a tongue-like, thin-skinned appendage, in which we may generally observe a few small strongly refractive granules, like those seen in the roots of the adult animal. I have therefore supposed these appendages to be the rudiments of the future roots. A perfectly similar appendage, “a most delicate tube or ribbon,” was found by Darwin in free-swimming pupae of Lepas australis on the last joints of the “prehensile antennae.” From the perfect accordance in their entire structure shown by the pupae of the Rhizocephala and Cirripedia, there can be no doubt that the appendages of Sacculina and Lepas, which are so like each other and spring from the same spot, are homologous structures.

Now in three species of Lepas, in Dichelaspis Warwickii and in Scalpellum Peronii, Darwin saw, on tearing recently-affixed animals from their point or support, that a long narrow band issued from the same point of the antennae; its end was torn away, and in Dichelaspis, judging from its ragged appearance, it had attached itself firmly to the support. From this it follows that this appendage in Lepas australis can hardly be anything but a young cement-duct. If, therefore, the supposition that the appendages on the antennae of the pupae of Rhizocephala are young roots be correct, the roots of the Rhizocephala are homologous with the cement-ducts of the Cirripedia. And this, strange as it may appear at the first glance, seems to me scarcely doubtful. It is true that the act of adhesion of the Rhizocephala has never yet been observed, but it is more than probable that they attach themselves, just like the Cirripedia, by means of the antennae, and that therefore the points of attachment in the two groups indicate homologous parts of the body. From the point of attachment in the Rhizocephala the roots penetrate into the body of the host, whilst in the Cirripedia, the cement-ducts issue from the same point. The roots are blind tubes, ramified in different ways in different species. The cement-ducts in the basis of the Balanidae likewise constitute a generally remarkably complicated system of ramified tubes, with regard to the mode of termination of which nothing certain has yet been made out. Individual caecal branches are not unfrequently seen even in the vicinity of the carina; and, at least in some species, in which the cement-ducts divide into extremely numerous and fine branchlets, forming a network which gradually becomes denser towards the circumference of the basis, these seem nowhere to possess an orifice.

Now as to the question: How were Cirripedia converted by natural selection into Rhizocephala?

A considerable number of existing Cirripedia settle exclusively or chiefly upon living animals;—on Sponges, Corals, Mollusks, Cetaceans, Turtles, Sea-Snakes, Sharks, Crustaceans, Sea Urchins, and even on Acalephs. Dichelaspis Darwinii was found by Filippi in the branchial cavity of Palinurus vulgaris, and I have met with another species of the same genus in the branchial cavity of Lupea diacantha.

The same thing may have taken place in primitive times. The supposition that certain Cirripedes might once upon a time have selected the soft ventral surface of a Crab, Porcellana or Pagurus, for its dwelling-place, has certainly nothing improbable about it. If then the cement-ducts of such a Cirripede instead of merely spreading on the surface, pierced or pushed before them the soft ventral skin and penetrated into the interior of the host, this must have been beneficial to the animal, because it would be thereby more securely attached and protected from being thrown off during the moulting of its host. Variations in this direction were preserved as advantageous.

But as soon as the cement-ducts penetrated into the body-cavity of the host and were bathed by its fluids, an endosmotic interchange must necessarily have been set up between the materials dissolved in these fluids and in the contents of the cement-ducts, and this interchange could not be without influence upon the nourishment of the parasite. The new source of nourishment opened up in this manner was, as constantly flowing, more certain than that offered by the nourishment accidentally whirled into the mouth of the sedentary animal. The individuals favoured in the development of the cement-ducts now converted into nutriferous roots, had more than others the prospect of abundant food, of vigorous growth, and of producing a numerous progeny. With the further development, assisted by natural selection, of the roots embracing the intestine of the host and spreading amongst its hepatic tubes, the introduction of nourishment through the mouth and all the parts implicated in it, such as the whirling cirri, the buccal organs, and the intestine, gradually lost their importance, became aborted by disuse, and finally disappeared without leaving a trace of their existence. Protected by the abdomen of the Crab, or by the shell inhabited by the Pagurus, the parasite also no longer required the calcareous test, in which, no doubt, the first Cirripedes settling upon these Decapods rejoiced. This protective covering, having become superfluous, also disappeared, and there remained at last only a soft sack filled with eggs, without limbs, without mouth or alimentary canal, and nourished, like a plant, by means of roots, which it pushed into the body of its host. The Cirripede had become a Rhizocephalon.

If it be desired to form a notion of what our parasite may have looked like when half way in its progress from the one form to the other, we may consult the figures given by Darwin, (Lepadidae Plate 4 figures 1 to 7) of Anelasma squalicola. This Lepadide, which lives upon Sharks in the North Sea, seems, in fact, to be in the best way to lose its cirri and buccal organs in the same manner. The widely-cleft, shell-less test is supported upon a thick peduncle, which is immersed in the skin of the Shark. The surface of the peduncle is beset with much-ramified, hollow filaments, which “penetrate the Shark’s flesh like roots” (Darwin). Darwin looked in vain for cement-glands and cement. It seems to me hardly doubtful, that the ramified hollow filaments are themselves nothing but the cement-ducts converted into nutritive roots, and that it is just in consequence of the development of this new source of nourishment, that the cirri and buccal organs are in the highest degree aborted. All the parts of the mouth are extremely minute; the palpi and exterior maxillae have