FACTS AND ARGUMENTS
BY FRITZ MULLER.
WITH ADDITIONS BY THE AUTHOR.
TRANSLATED FROM THE GERMAN
BY W.S. DALLAS, F.L.S.,
ASSISTANT SECRETARY TO THE GEOLOGICAL SOCIETY OF LONDON.
LONDON: JOHN MURRAY, ALBEMARLE STREET. 1869.
MR. DARWIN'S WORKS.
A NATURALIST'S VOYAGE ROUND THE WORLD; BEING A JOURNAL OF RESEARCHES INTO THE NATURAL HISTORY AND GEOLOGY OF COUNTRIES VISITED. Post 8vo. 9 shillings.
THE ORIGIN OF SPECIES, BY MEANS OF NATURAL SELECTION; OR, THE PRESERVATION OF FAVOURED RACES IN THE STRUGGLE FOR LIFE. WOODCUTS. Post 8vo. 15 shillings.
THE VARIOUS CONTRIVANCES BY WHICH BRITISH AND FOREIGN ORCHIDS ARE FERTILISED BY INSECTS, AND ON THE GOOD EFFECTS OF INTERCROSSING. Woodcuts, Post 8vo. 9 shillings.
THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION. Illustrations. 2 volumes, 8vo. 28 shillings.
My principal reason for undertaking the translation of Dr. Fritz Muller's admirable work on the Crustacea, entitled 'Fur Darwin,' was that it was still, although published as long ago as 1864, and highly esteemed by the author's scientific countrymen, absolutely unknown to a great number of English naturalists, including some who have occupied themselves more or less specially with the subjects of which it treats. It possesses a value quite independent of its reference to Darwinism, due to the number of highly interesting and important facts in the natural history and particularly the developmental history of the Crustacea, which its distinguished author, himself an unwearied and original investigator of these matters, has brought together in it. To a considerable section of English naturalists the tone adopted by the author in speaking of one of the greatest of their number will be a source of much gratification.
In granting his permission for the translation of his little book, Dr. Fritz Muller kindly offered to send some emendations and additions to certain parts of it. His notes included many corrections of printers' errors, some of which would have proved unintelligible without his aid, some small additions and notes which have been inserted in their proper places, and two longer pieces, one forming a footnote near the close of Chapter 11, the other at the end of Chapter 12, describing the probable mode of evolution of the Rhizocephala from the Cirripedia.
Of the execution of the translation I will say but little. My chief object in this, as in other cases, has been to furnish, as nearly as possible, a literal version of the original, regarding mere elegance of expression as of secondary importance in a scientific work. As much of Dr. Muller's German does not submit itself to such treatment very readily, I must beg his and the reader's indulgence for any imperfections arising from this cause.
LONDON, 15TH FEBRUARY, 1869.
It is not the purpose of the following pages to discuss once more the arguments deduced for and against Darwin's theory of the origin of species, or to weigh them one against the other. Their object is simply to indicate a few facts favourable to this theory, collected upon the same South American ground, on which, as Darwin tells us, the idea first occurred to him of devoting his attention to "the origin of species,—that mystery of mysteries."
It is only by the accumulation of new and valuable material that the controversy will gradually be brought into a state fit for final decision, and this appears to be for the present of more importance than a repeated analysis of what is already before us. Moreover, it is but fair to leave it to Darwin himself at first to beat off the attacks of his opponents from the splendid structure which he has raised with such a master-hand.
DESTERRO, 7TH SEPTEMBER, 1863.
CHAPTER 1. INTRODUCTORY.
CHAPTER 2. THE SPECIES OF MELITA.
CHAPTER 3. MORPHOLOGY OF CRUSTACEA.
CHAPTER 4. SEXUAL PECULIARITIES AND DIMORPHISM.
CHAPTER 5. RESPIRATION IN LAND CRABS.
CHAPTER 6. STRUCTURE OF THE HEART IN EDRIOPHTHALMA.
CHAPTER 7. DEVELOPMENTAL HISTORY OF PODOPHTHALMA.
CHAPTER 8. DEVELOPMENTAL HISTORY OF EDRIOPHTHALMA.
CHAPTER 9. DEVELOPMENTAL HISTORY OF ENTOMOSTRACA, CIRRIPEDES, AND RHIZOCEPHALA.
CHAPTER 10. ON THE PRINCIPLES OF CLASSIFICATION.
CHAPTER 11. ON THE PROGRESS OF EVOLUTION.
CHAPTER 12. PROGRESS OF EVOLUTION IN CRUSTACEA.
HISTORY OF CRUSTACEA.
CHAPTER 1. INTRODUCTORY.
When I had read Charles Darwin's book 'On the Origin of Species,' it seemed to me that there was one mode, and that perhaps the most certain, of testing the correctness of the views developed in it, namely, to attempt apply them as specially as possible to some particular group of animals. such an attempt to establish a genealogical tree, whether for the families of a class, the genera of a large family, or for the species of an extensive genus, and to produce pictures as complete and intelligible as possible of the common ancestors of the various smaller and larger circles, might furnish a result in three different ways.
1. In the first place, Darwin's suppositions when thus applied might lead to irreconcilable and contradictory conclusions, from which the erroneousness of the suppositions might be inferred. If Darwin's opinions are false, it was to be expected that contradictions would accompany their detailed application at every step, and that these, by their cumulative force, would entirely destroy the suppositions from which they proceeded, even though the deductions derived from each particular case might possess little of the unconditional nature of mathematical proof.
2. Secondly, the attempt might be successful to a greater or less extent. If it was possible upon the foundation and with the aid of the Darwinian theory, to show in what sequence the various smaller and larger circles had separated from the common fundamental form and from each other, in what sequence they had acquired the peculiarities which now characterise them, and what transformations they had undergone in the lapse of ages,—if the establishment of such a genealogical tree, of a primitive history of the group under consideration, free from internal contradictions, was possible,—then this conception, the more completely it took up all the species within itself, and the more deeply it enabled us to descend into the details of their structure, must in the same proportion bear in itself the warrant of its truth, and the more convincingly prove that the foundation upon which it is built is no loose sand, and that it is more than merely "an intellectual dream."
3. In the third place, however, it was possible, and this could not but appear, prima facie, the most probable case, that the attempt might be frustrated by the difficulties standing in its way, without settling the question, either way, in a perfectly satisfactory manner. But if it were only possible in this way to arrive for oneself at a moderately certain independent judgment upon a matter affecting the highest questions so deeply, even this alone could not but be esteemed a great gain.
Having determined to make the attempt, I had in the first place to decide upon some particular class. The choice was necessarily limited to those the chief forms of which were easily to be obtained alive in some abundance. The Crabs and Macrurous Crustacea, the Stomapoda, the Diastylidae, the Amphipoda and Isopoda, the Ostracoda and Daphnidae, the Copepoda and Parasita, the Cirripedes and Rhizocephala of our coast, representing the class of Crustacea with the deficiency only of the Phyllopoda and Xiphosura, furnished a long and varied, and at the same time intimately connected series, such as was at my command in no other class. But even independently of this circumstance the selection of the Crustacea could hardly have been doubtful. Nowhere else, as has already been indicated by various writers, is the temptation stronger to give to the expressions "relationship, production from a common fundamental form," and the like, more than a mere figurative signification, than in the case of the lower Crustacea. Among the parasitic Crustacea, especially, everybody has long been accustomed to speak, in a manner scarcely admitting of a figurative meaning, of their arrest of development by parasitism, as if the transformation of species were a matter of course. It would certainly never appear to any one to be a pastime worthy of the Deity, to amuse himself with the contrivance of these marvellous cripplings, and so they were supposed to have fallen by their own fault, like Adam, from their previous state of perfection.
That a great part of the larger and smaller groups into which this class is divided, might be regarded as satisfactorily established, was a further advantage not to be undervalued; whilst in two other classes with which I was familiar, namely, the Annelida and Acalephae, all the attempted arrangements could only be considered preliminary revisions. These undisplaceable groups, like the sharply marked forms of the hard, many-jointed dermal framework, were not only important as safe starting points and supports, but were also of the highest value as inflexible barriers in a problem in which, from its very nature, fancy must freely unfold her wings.
When I thus began to study our Crustacea more closely from this new stand-point of the Darwinian theory,—when I attempted to bring their arrangements into the form of a geological tree, and to form some idea of the probable structure of their ancestors,—I speedily saw (as indeed I expected) that it would require years of preliminary work before the essential problem could be seriously handled. The extant systematic works generally laid more weight upon the characters separating the genera, families and orders, than upon those which unite the members of each group, and consequently often furnished but little employable material. But above all things a thorough knowledge of development was indispensable, and every one knows how imperfect is our present knowledge of this subject. The existing deficiencies were the more difficult to supply, because, as Van Beneden remarks with regard to the Decapoda, from the often incredible difference in the development of the most nearly allied forms, these must be separately studied—usually family by family, and frequently genus by genus—nay, sometimes, as in the case of Peneus, even species by species; and because these investigations, in themselves troublesome and tedious, often depend for their success upon a lucky chance.
But although the satisfactory completion of the "Genealogical tree of the Crustacea" appeared to be an undertaking for which the strength and life of an individual would hardly suffice, even under more favourable circumstances than could be presented by a distant island, far removed from the great market of scientific life, far from libraries and museums—nevertheless its practicability became daily less doubtful in my eyes, and fresh observations daily made me more favourably inclined towards the Darwinian theory.
In determining to state the arguments which I derived from the consideration of our Crustacea in favour of Darwin's views, and which (together with more general considerations and observations in other departments), essentially aided in making the correctness of those views seem more and more palpable to me, I am chiefly influenced by an expression of Darwin's: "Whoever," says he ('Origin of Species' page 482), "is led to believe that species are mutable, will do a good service by conscientiously expressing his conviction." To the desire expressed in these words I respond, for my own part, with the more pleasure, as this furnishes me with an opportunity of publicly giving expression in words to the thanks which I feel most deeply to be due from me to Darwin for the instructions and suggestions for which I am so deeply indebted to his book. Accordingly I throw this sand-grain with confidence into the scale against "the load of prejudice by which this subject is overwhelmed," without troubling myself as to whether the priests of orthodox science will reckon me amongst dreamers and children in knowledge of the laws of nature.
CHAPTER 2. THE SPECIES OF MELITA.
A false supposition, when the consequences proceeding from it are followed further and further, will sooner or later lead to absurdities and palpable contradictions. During the period of tormenting doubt—and this was by no means a short one—when the pointer of the scales oscillated before me in perfect uncertainty between the pro and the con, and when any fact leading to a quick decision would have been most welcome to me, I took no small pains to detect some such contradictions among the inferences as to the class of Crustacea furnished by the Darwinian theory. But I found none, either then, or subsequently. Those which I thought I had found were dispelled on closer consideration, or actually became converted into supports for Darwin's theory.
Nor, so far as I am aware, have any of the NECESSARY consequences of Darwin's hypotheses been proved by any one else, to stand in clear and irreconcilable contradiction. And yet, as the most profound students of the animal kingdom are amongst Darwin's opponents, it would seem that it ought to have been an easy matter for them to crush him long since beneath a mass of absurd and contradictory inferences, if any such were to be drawn from his theory. To this want of demonstrated contradictions I think we may ascribe just the same importance in Darwin's favour, that his opponents have attributed to the absence of demonstrated intermediate forms between the species of the various strata of the earth. Independently of the reasons which Darwin gives for the preservation of such intermediate forms being only exceptional, this last mentioned circumstance will not be regarded as of very great significance by any one who has traced the development of an animal upon larvae fished from the sea, and had to seek in vain for months, and even years, for those transitional forms, which he nevertheless knew to be swarming around him in thousands.
A few examples may show how contradictions might come forth as necessary results of the Darwinian hypotheses.
It seems to be a necessity for all crabs which remain for a long time out of the water (but why is of no consequence to us here), that air shall penetrate from behind into the branchial cavity. Now these crabs, which have become more or less estranged from the water, belong to the most different families—the Raninidae (Ranina), Eriphinae (Eriphia gonagra), Grapsoidae (Aratus, Sesarma, etc.), Ocypodidae (Gelasimus, Ocypoda), etc., and the separation of these families must doubtless be referred to a much earlier period than the habit of leaving the water displayed by some of their members. The arrangements connected with aerial respiration, therefore, could not be inherited from a common ancestor, and could scarcely be accordant in their construction. If there were any such accordance not referable to accidental resemblance among them, it would have to be laid in the scale as evidence against the correctness of Darwin's views. I shall show hereafter how in this case the result, far from presenting such contradictions, was rather in the most complete harmony with what might be predicted from Darwin's theory.
(FIGURE 1. Melita exilii n. sp., male, enlarged five times. The large branchial lamellae are seen projecting between the legs.)
A second example.—We are already acquainted with four species of Melita (M. valida, setipes, anisochir, and Fresnelii), and I can add a fifth (Figure 1), in which the second pair of feet bears upon one side a small hand of the usual structure, and on the other an enormous clasp-forceps. This want of symmetry is something so unusual among the Amphipoda, and the structure of the clasp-forceps differs so much from what is seen elsewhere in this order, and agrees so closely in the five species, that one must unhesitatingly regard them as having sprung from common ancestors belonging to them alone among known species. But one of these species, M. Fresnelii, discovered by Savigny, in Egypt, is said to want the secondary flagellum of the anterior antennae, which occurs in the others. From the trustworthiness of all Savigny's works there can scarcely be a doubt as to the correctness of this statement. Now, if the presence or absence of the secondary flagellum possessed the significance of a distinctive generic character, which is usually ascribed to it, or if there were other important differences between Melita Fresnelii and the other species above-mentioned, which would make it seem natural to separate M. Fresnelii as a distinct genus, and to leave the others united with the rest of the species of Melita—that is to say, in the sense of the Darwinian theory, if we assume that all the other Melitae possessed common ancestors, which were not at the same time the ancestors of M. Fresnelii—this would stand in contradiction to the conclusion, derived from the structure of the clasp-forceps, that M. Fresnelii and the four other species above-mentioned possessed common ancestors, which were not also the ancestors of the remaining species of Melita. It would follow:—
1. From the structure of the clasp-forceps: that M. exilii, etc. and M. Fresnelii would branch off together from a stem which branches off from M. palmata.
2. From the presence or absence of the secondary flagellum: that M. palmata, etc. and M. exilii, etc. would branch off together from a stem which branches off from M. Fresnelii.
As, in the first case, among the Crabs, a typical agreement of arrangements produced independently of each other would have been a very suspicious circumstance for Darwin's theory, so also, in the second, would any difference more profound than that of very nearly allied species. Now it seems to me that the secondary flagellum can by no means furnish a reason for doubting the close relationship of M. Fresnelii to M. exilii, etc., which is indicated by the peculiar structure of the unpaired clasp-forceps. In the first place we must consider the possibility that the secondary flagellum, which is not always easy to detect, may only have been overlooked by Savigny, as indeed Spence Bate supposes to have been the case. If it is really deficient it must be remarked that I have found it in species of the genera Leucothoe, Cyrtophium and Amphilochus, in which genera it was missed by Savigny, Dana and Spence Bate—that a species proved by the form of the Epimera (Coxae Sp. B.) of the caudal feet (uropoda Westw.), etc., to be a true Amphithoe* possesses it (* I accept this and all the other genera of Amphipoda here mentioned, with the limits given to them by Spence Bate ('Catalogue of Amphipodous Crustacea').)—that in many species of Cerapus it is reduced to a scarcely perceptible rudiment—nay, that it is sometimes present in youth and disappears (although perhaps not without leaving some trace) at maturity, as was found by Spence Bate to be the case in Acanthonotus Owenii and Atylus carinatus, and I can affirm with regard to an Atylus of these seas, remarkable for its plumose branchiae—and that from all this, at the present day when the increasing number of known Amphipoda and the splitting of them into numerous genera thereby induced, compels us to descend to very minute distinctive characters, we must nevertheless hesitate before employing the secondary flagellum as a generic character. The case of Melita Fresnelii therefore cannot excite any doubts as to Darwin's theory.
CHAPTER 3. MORPHOLOGY OF CRUSTACEA—NAUPLIUS-LARVAE.
If the absence of contradictions among the inferences deduced from them for a narrow and consequently easily surveyed department must prepossess us in favour of Darwin's views, it must be welcomed as a positive triumph of his theory if far-reaching conclusions founded upon it should SUBSEQUENTLY be confirmed by facts, the existence of which science, in its previous state, by no means allowed us to suspect. From many results of this kind upon which I could report, I select as examples, two, which were of particular importance to me, and relate to discoveries the great significance of which in the morphology and classification of the Crustacea will not be denied even by the opponents of Darwin.
Considerations upon the developmental history of the Crustacea had led me to the conclusion that, if the higher and lower Crustacea were at all derivable from common progenitors, the former also must once have passed through Nauplius-like conditions. Soon afterwards I discovered Naupliiform larvae of Shrimps ('Archiv fur Naturgeschichte' 1860 1 page 8), and I must admit that this discovery gave me the first decided turn in Darwin's favour.
(FIGURE 2. Tanais dubius (?) Kr. female, magnified 25 times, showing the orifice of entrance (x) into the cavity overarched by the carapace, in which an appendage of the second pair of maxillae (f) plays. On four feet (i, k, l, m) are the rudiments of the lamellae which subsequently form the brood-cavity.)
The similar number of segments* occurring in the Crabs and Macrura, Amphipoda and Isopoda, in which the last seven segments are always different from the preceding ones in the appendages with which they are furnished, could only be regarded as an inheritance from the same ancestors.
(* Like Claus I do not regard the eyes of the Crustacea as limbs, and therefore admit no ocular segment; on the other hand I count in the median piece of the tail, to which the character of a segment is often denied. In opposition to its interpretation as a segment of the body, only the want of limbs can be cited; in its favour we have the relation of the intestine, which usually opens in this piece, and sometimes even traverses its whole length, as in Microdeutopus and some other Amphipoda. In Microdeutopus, as Spence Bate has already pointed out, one is even led to regard small processes of this tubular caudal piece as rudimentary members. Bell also ('British Stalk-eyed Crustacea' page 20), states that he observed limbs of the last segment in Palaemon serratus in the form of small moveable points.
The attempt has often been made to divide the body of the higher Crustacea into small sections composed of equal numbers of segments, these sections consisting of 3, 5 or 7 segments. None of these attempts has ever met with general acceptance; my own investigations lead me to a conception which nearly approaches Van Beneden's. I assume four sections of 5 segments each—the primitive body, the fore-body, the hind-body, and the middle-body. The primitive body includes the segments which the naupliiform larva brings with it out of the egg; it is afterwards divided, by the younger sections which become developed in its middle, into the head and tail. To this primitive body belong the two pairs of antennae, the mandibles and the caudal feet ("posterior pair of pleopoda," Sp. B.). Even in the mature animal the fact that these terminal sections belong to one another is sometimes betrayed by the resemblance of their appendages, especially that of the outer branch of the caudal feet, with the outer branch (the so-called scale) of the second pair of antennae. Like the antennae, the caudal feet may also become the bearers of high sensorial apparatus, as is shown by the ear of Mysis.
The sequence of the sections of the body in order of time seems originally to have been, that first the fore-body, then the hind-body, and finally the middle-body was formed. The fore-body appears, in the adult animal, to be entirely or partially amalgamated with the head; its appendages (siagonopoda Westw.) are all or in part serviceable for the reception of food, and generally sharply distinguished from those of the following group. The segments of the middle-body seem always to put forth limbs immediately after their own appearance, whilst the segments of the hind-body often remain destitute of feet through long portions of the larval life or even throughout life (as in many female Diastylidae), a reason, among many others, for not, as is usual, regarding the middle-body of the Crustacea as equivalent to the constantly footless abdomen of Insects. The appendages of the middle-body (pereiopoda) seem never, even in their youngest form, to possess two equal branches, a peculiarity which usually characterises the appendages of the hind-body. This is a circumstance which renders very doubtful the equivalence of the middle-body of the Malacostraca with the section of the body which in the Copepoda bears the swimming feet and in the Cirripedia the cirri.
The comprehension of the feet of the hind-body and tail in a single group (as "fausses pattes abdominales," or as "pleopoda") seems not to be justifiable. When there is a metamorphosis, they are probably always produced at different periods, and they are almost always quite different in structure and function. Even in the Amphipoda, in which the caudal feet usually resemble in appearance the last two pairs of abdominal feet, they are in general distinguished by some sort of peculiarity, and whilst the abdominal feet are reproduced in wearisome uniformity throughout the entire order, the caudal feet are, as is well-known, amongst the most variable parts of the Amphipoda.)
And if at the present day the majority of the Crabs and Macrura, and indeed the Stalk-eyed Crustacea in general, pass through Zoea-like developmental states, and the same mode of transformation was to be ascribed to their ancestors, the same thing must also apply, if not to the immediate ancestors of the Amphipoda and Isopoda, at least to the common progenitors of these and the Stalk-eyed Crustacea. Any such assumption as this was, however, very hazardous, so long as not a single fact properly relating to the Edriophthalma could be adduced in its support, as the structure of this very coherent group seemed to be almost irreconcilable with many peculiarities of the Zoea. Thus, in my eyes, this point long constituted one of the chief difficulties in the application of the Darwinian views to the Crustacea, and I could scarcely venture to hope that I might yet find traces of this passage through the Zoea-form among the Amphipoda or Isopoda, and thus obtain a positive proof of the correctness of this conclusion. At this point Van Beneden's statement that a cheliferous Isopod (Tanais Dulongii), belonging, according to Milne-Edwards, to the same family as the common Asellus aquaticus, possesses a carapace like the Decapoda, directed my attention to these animals, and a careful examination proved that these Isopods have preserved, more truly than any other adult Crustacea, many of the most essential peculiarities of the Zoeae, especially their mode of respiration. Whilst in all other Oniscoida the abdominal feet serve for respiration, these in our cheliferous Isopod (Figure 2) are solely motory organs, into which no blood-corpuscle ever enters, and the chief seat of respiration is, as in the Zoeae, in the lateral parts of the carapace, which are abundantly traversed by currents of blood, and beneath which a constant stream of water passes, maintained, as in Zoeae and the adult Decapoda, by an appendage of the second pair of maxillae, which is wanting in all other Edriophthalma.
For both these discoveries, it may be remarked in passing, science is indebted less to a happy chance than immediately to Darwin's theory.
Species of Peneus live in the European seas, as well as here, and their Nauplius-brood has no doubt repeatedly passed unnoticed through the hands of the numerous naturalists who have investigated those seas, as well as through my own,* for it has nothing which could attract particular attention amongst the multifarious and often wonderful Nauplius-forms. (* Mecznikow has recently found Naupliiform shrimp-larvae in the sea near Naples.) When I, fancying from the similarity of its movements that it was a young Peneus-Zoea, had for the first time captured such a larva, and on bringing it under the microscope found a Nauplius differing toto coelo from this Zoea, I might have thrown it aside as being completely foreign to the developmental series which I was tracing, if the idea of early Naupliiform stages of the higher Crustacea, which indeed I did not believe to be still extant, had not at the moment vividly occupied my attention.
And if I had not long been seeking among the Edriophthalma for traces of the supposititious Zoea-state, and seized with avidity upon everything that promised to made this refractory Order serviceable to me, Van Beneden's short statement could hardly have affected me so much in the manner of an electric shock, and impelled me to a renewed study of the Tanaides, especially as I had once before plagued myself with them in the Baltic, without getting any further than my predecessors, and I have not much taste for going twice over the same ground.
CHAPTER 4. SEXUAL PECULIARITIES AND DIMORPHISM.
Our Tanais, which in nearly all the particulars of its structure is an extremely remarkable animal, furnished me with a second fact worthy of notice in connection with the theory of the origin of species by natural selection.
When hand-like or cheliform structures occur in the Crustacea, these are usually more strongly developed in the males than in the females, often becoming enlarged in the former to quite a disproportionate size, as we have already seen to be the case in Melita. A better known example of such gigantic chelae is presented by the males of the Calling Crabs (Gelasimus), which are said in running to carry these claws "elevated, as if beckoning with them"—a statement which, however, is not true of all the species, as a small and particularly large-clawed one, which I have seen running about by thousands in the cassava-fields at the mouth of the Cambriu, always holds them closely pressed against its body.
A second peculiarity of the male Crustacea consists not unfrequently in a more abundant development on the flagellum of the anterior antennae of delicate filaments which Spence Bate calls "auditory cilia," and which I have considered to be olfactory organs, as did Leydig before me, although I was not aware of it. Thus they form long dense tufts in the males of many Diastylidae, as Van Beneden also states with regard to Bodotria, whilst the females only possess them more sparingly. In the Copepoda, Claus called attention to the difference of the sexes in this respect. It seems to me, as I may remark in passing, that this stronger development in the males is greatly in favour of the opinion maintained by Leydig and myself, as in other cases male animals are not unfrequently guided by the scent in their pursuit of the ardent females.
Now, in our Tanais, the young males up to the last change of skin preceding sexual maturity resemble the females, but then they undergo an important metamorphosis. Amongst other things they lose the moveable appendages of the mouth even to those which serve for the maintenance of the respiratory current; their intestine is always found empty, and they appear only to live for love. But what is most remarkable is, that they now appear under two different forms. Some (Figure 3) acquire powerful, long-fingered, and very mobile chelae, and, instead of the single olfactory filament of the female, have from 12 to 17 of these organs, which stand two or three together on each joint of the flagellum. The others (Figure 5) retain the short thick form of the chelae of the females; but, on the other hand, their antennae (Figure 6) are equipped with a far greater number of olfactory filaments, which stand in groups of from five to seven together.
(FIGURE 3. Head of the ordinary form of the male of Tanais dubius (?) Kr. magnified 90 times. The terminal setae of the second pair of antennae project between the cheliferous feet.
FIGURE 4. Buccal region of the same from below; lambda, labrum.
FIGURE 5. Head of the rarer form of the male, magnified 25 times.
FIGURE 6. Flagellum of the same, with olfactory filaments, magnified 90 times.)
In the first place, and before inquiring into its significance, I will say a word upon this fact itself. It was natural to consider whether two different species with very similar females and very different males might not perhaps live together, or whether the males, instead of occurring in two sharply defined forms, might not be only variable within very wide limits. I can admit neither of these suppositions. Our Tanais lives among densely interwoven Confervae, which form a coat of about an inch in thickness upon stones in the neighbourhood of the shore. If a handful of this green felt is put into a large glass with clear sea-water, the walls of the glass are soon seen covered with hundreds, nay with thousands, of these little, plump, whitish Isopods. In this way I have examined thousands of them with the simple lens, and I have also examined many hundreds with the microscope, without finding any differences among the females, or any intermediate forms between the two kinds of males.
To the old school this occurrence of two kinds of males will appear to be merely a matter of curiosity. To those who regard the "plan of creation" as the "free conception of an Almighty intellect, matured in the thoughts of the latter before it is manifested in palpable, external forms," it will appear to be a mere caprice of the Creator, as it is inexplicable either from the point of view of practical adaptation, or from the "typical plan of structure." From the side of Darwin's theory, on the contrary, this fact acquires meaning and significance, and it appears in return to be fitted to throw light upon a question in which Bronn saw "the first and most material objection against the new theory," namely, how it is possible that from the accumulation in various directions of the smallest variations running out of one another, varieties and species are produced, which stand out from the primary form clearly and sharply like the petiolated leaf of a Dicotyledon, and are not amalgamated with the primary form and with each other like the irregular curled lobes of a foliaceous Lichen.
Let us suppose that the males of our Tanais, hitherto identical in structure, begin to vary, in all directions as Bronn thinks, for aught I care. If the species was adapted to its conditions of existence, if the BEST in this respect had been attained and secured by natural selection, fresh variations affecting the species as a species would be retrogressions, and thus could have no prospect of prevailing. They must rather have disappeared again as they arose, and the lists would remain open to the males under variation, only in respect of their sexual relations. In these they might acquire advantages over their rivals by their being enabled either to seek or to seize the females better. The best smellers would overcome all that were inferior to them in this respect, unless the latter had other advantages, such as more powerful chelae, to oppose to them. The best claspers would overcome all less strongly armed champions, unless these opposed to them some other advantage, such as sharper senses. It will be easily understood how in this manner all the intermediate steps less favoured in the development of the olfactory filaments or of the chelae would disappear from the lists, and two sharply defined forms, the best smellers and the best claspers, would remain as the sole adversaries. At the present day the contest seems to have been decided in favour of the latter, as they occur in greatly preponderating numbers, perhaps a hundred of them to one smeller.
To return to Bronn's objection. When he says that "for the support of the Darwinian theory, and in order to explain why many species do not coalesce by means of intermediate forms, he would gladly discover some external or internal principle which should compel the variations of each species to advance in ONE direction, instead of merely permitting them in all directions," we may, in this as in many other cases, find such a principle in the fact that actually only a few directions stand open in which the variations are at the same time improvements, and in which therefore they can accumulate and become fixed; whilst in all others, being either indifferent or injurious, they will go as lightly as they come.
(FIGURE 7. Orchestia Darwinii, n. sp. male.)
The occurrence of two kinds of males in the same species may perhaps not be a very rare phenomenon in animals in which the males differ widely from the females in structure. But only in those which can be procured in sufficient abundance, will it be possible to arrive at a conviction that we have not before us either two different species, or animals of different ages. From my own observation, although not very extensive, I can give a second example. It relates to a shore-hopper (Orchestia). The animal (Figure 7) lives in marshy places in the vicinity of the sea, under decaying leaves, in the loose earth which the Marsh Crabs (Gelasimus, Sesarma, Cyclograpsus, etc.) throw up around the entrance to their borrows, and even under dry cow-dung and horse-dung. If this species removes to a greater distance from the shore than the majority of its congeners (although some of them advance very far into the land and even upon mountains of a thousand feet in height, such as O. tahitensis, telluris, and sylvicola), its male differs still more from all known species by the powerful chelae of the second pair of feet. Orchestia gryphus, from the sandy coast of Monchgut, alone presents a somewhat similar structure, but in a far less degree; elsewhere the form of the hand usual in the Amphipoda occurs. Now there is a considerable difference between the males of this species, especially in the structure of these chelae—a different so great that we can scarcely find a parallel to it elsewhere between two species of the genus—and yet, as in Tanais, we do not meet with a long series of structures running into one another, but only two forms united by no intermediate terms (Figures 8 and 9). The males would be unhesitatingly regarded as belonging to two well-marked species if they did not live on the same spot, with undistinguishable females. That the two forms of the chelae of the males occur in this species is so far worthy of notice, because the formation of the chelae, which differs widely from the ordinary structure in the other species, indicates that it has quite recently undergone considerable changes, and therefore such a phenomenon was to be expected in it rather than in other species.
(FIGURES 8 AND 9. The two forms of the chelae of the male of Orchestia Darwinii, magnified 45 times.)
I cannot refrain from taking this opportunity of remarking that (so far as appears from Spence Bate's catalogue), for two different kinds of males (Orchestia telluris and sylvicola) which live together in the forests of New Zealand, only one form of female is known, and hazarding the supposition that we have here a similar case. It does not seem to me to be probable that two nearly allied species of these social Amphipoda should occur mixed together under the same conditions of life.
(FIGURE 10. Coxal lamella of the penultimate pair of feet of the male (a), and coxal lamella, with the three following joints of the same pair of feet of the female (b) of Melita Messalina, magnified 45 diam.
FIGURE 11. Coxal lamella of the same pair of feet of the female of M. insatiabilis.)
As the males of several species of Melita are distinguished by the powerful unpaired clasp-forceps, the females of some other species of the same genus are equally distinguished from all other Amphipoda by the circumstance that in them a peculiar apparatus is developed which facilitates their being held by the male. The coxal lamellae of the penultimate pair of feet are produced into hook-like processes, of which the male lays hold with the hands of the first pair of feet. The two species in which I am acquainted with this structure are amongst the most salacious animals of their order, even females which are laden with eggs in all stages of development, not unfrequently have their males upon their backs. The two species are nearly allied to Melita palmata Leach (Gammarus Dugesii, Edw.), which is widely distributed on the European coasts, and has been frequently investigated; unfortunately, however, I can find no information as to whether the females of this or any other European species possess a similar contrivance. In M. exilii all the coxal lamellae are of the ordinary formation. Nevertheless, be this as it will, whether they exist in two or in twenty species, the occurrence of these peculiar hook-like processes is certainly very limited.
Now our two species live sheltered beneath slightly tilted stones in the neighbourhood of the shore: one of them, Melita Messalina, so high that it is but rarely covered by the water; the other, Melita insatiabilis, a little lower; both species live together in numerous swarms. We cannot therefore suppose that the loving couples are threatened with disturbance more frequently than those of other species, nor would it be more difficult for the male, than for those of other species, in case of his losing his female, to find a new one. Nor is it any more easy to see how the contrivance on the body of the female for insuring the act of copulation could be injurious to other species. But so long as it is not demonstrated that our species are particularly in want of this contrivance, or that the latter would rather be injurious than beneficial to other species, its presence only in these few Amphipoda will have to be regarded not as the work of far-seeing wisdom, but as that of a favourable chance made use of by Natural Selection. Under the latter supposition its isolated occurrence is intelligible, whilst we cannot perceive why the Creator blessed just these few species with an apparatus which he found to be quite compatible with the "general plan of structure" of the Amphipoda, and yet denied it to others which live under the same external conditions, and equal them even in their extraordinary salacity. Associated with, or in the immediate vicinity of the two species of Melita, live two species of Allorchestes, the pairs of which are met with almost more numerously than the single animals, and yet their females show no trace of the above-mentioned processes of the coxal lamellae.
These cases, I think, must be brought to bear against the conception supported with so much genius and knowledge by Agassiz, that species are embodied thoughts of the Creator; and, with these, all similar instances in which arrangements which would be equally beneficial to all the species of a group are wanting in the majority and only conferred upon a few special favourites, which do not seem to want them any more than the rest.
CHAPTER 5. RESPIRATION IN LAND CRABS.
Among the numerous facts in the natural history of the Crustacea upon which a new and clear light is thrown by Darwin's theory, besides the two forms of the males in our Tanais and in Orchestia Darwinii, there is one which appears to me of particular importance, namely, the character of the branchial cavity in the air-breathing Crabs, of which, unfortunately, I have been unable to investigate some of the most remarkable (Gecarcinus, Ranina). As this character, namely, the existence of an entrance behind the branchiae, has hitherto been noticed, even as a fact, only in Ranina, I will go into it in some detail. I have already mentioned that, as indeed is required by Darwin's theory, this entrant orifice is produced in different manners in the different families.
In the Frog-crab (Ranina) of the Indian Ocean, which, according to Rumphius, loves to climb up on the roofs of the houses, the ordinary anterior entrant orifice is entirely wanting according to Milne-Edwards, and the entrance of a canal opening into the hindmost parts of the branchial cavity is situated beneath the commencement of the abdomen.
The case is most simple in some of the Grapsoidae, as in Aratus Pisonii, a charming, lively Crab which ascends the mangrove bushes (Rhizophora) and gnaws their leaves. By means of its short but remarkably acute claws, which prick like pins when it runs over the hand, this Crab climbs with the greatest agility upon the thinnest twigs. Once, when I had one of these animals sitting upon my hand, I noticed that it elevated the hinder part of its carapace, and that by this means a wide fissure was opened upon each side above the last pair of feet, through which I could look far into the branchial cavity. I have since been unable to procure this remarkable animal again, but on the other hand, I have frequently repeated the same observation upon another animal of the same family (apparently a true Grapsus), which lives abundantly upon the rocks of our coast. Whilst the hinder part of the carapace rises and the above-mentioned fissure is formed, the anterior part seems to sink, and to narrow or entirely close the anterior entrant orifice. Under water the elevation of the carapace never takes place. The animal therefore opens its branchial cavity in front or behind, according as it has to breathe water or air. How the elevation of the carapace is effected I do not know, but I believe that a membranous sac, which extends from the body cavity far into the branchial cavity beneath the hinder part of the carapace, is inflated by the impulsion of the fluids of the body, and the carapace is thereby raised.
I have also observed the same elevation of the carapace in some species of the allied genera Sesarma and Cyclograpsus, which dig deep holes in marshy ground, and often run about upon the wet mud, or sit, as if keeping watch, before their burrows. One must, however, wait for a long time with these animals, when taken out of the water, before they open their branchial cavity to the air, for they possess a wonderful arrangement, by means of which they can continue to breathe water for some time when out of the water. The orifices for the egress of the water which has served for respiration, are situated in these, as in most Crabs, in the anterior angles of the buccal frame ("cadre buccal," M.-Edw.), whilst the entrant fissures of the branchial cavity extend from its hinder angles above the first pair of feet. Now that portion of the carapace which extends at the sides of the mouth between the two orifices ("regions pterygostomiennes"), appears in our animals to be divided into small square compartments. Milne-Edwards has already pointed this out as a particularly remarkable peculiarity. This appearance is caused partly by small wart-like elevations, and partly and especially by curious geniculated hairs, which to a certain extent constitute a fine net or hair-sieve extended immediately over the surface of the carapace. Thus when a wave of water escapes from the branchial cavity, it immediately becomes diffused in this network of hairs and then again conveyed back to the branchial cavity by vigorous movements of the appendage of the outer maxilliped which works in the entrant fissure. Whilst the water glides in this way over the carapace in the form of a thin film, it will again saturate itself with oxygen, and may then serve afresh for the purposes of respiration. In order to complete this arrangement the outer maxillipeds, as indeed has long been known, bear a projecting ridge furnished with a dense fringe of hairs, which commences in front near their median line and passes backwards and outwards to the hinder angle of the buccal frame. Thus the two ridges of the right and left sides form together a triangle with the apex turned forwards,—a breakwater by which the water flowing from the branchial cavity is kept away from the mouth and reconducted to the branchial cavity. In very moist air the store of water contained in the branchial cavity may hold out for hours, and it is only when this is used up that the animal elevates its carapace in order to allow the air to have access to its branchiae from behind.
In Eriphia gonagra the entrant orifices of the respiratory cavity serving for aerial respiration are situated, not, as in the Grapsoidae, above, but behind the last pair of feet at the sides of the abdomen.
(FIGURE 12. Posterior entrance to the branchial cavity of Ocypoda rhombea, Fab., natural size. The carapace and the fourth foot of the right side are removed.
FIGURE 13. Points of some of the hairs of the basal joints of the foot, magnified 45 diam.)
The swift-footed Sand-Crabs (Ocypoda) are exclusively terrestrial animals, and can scarcely live for a single day in water; in a much shorter period a state of complete relaxation occurs and all voluntary movements cease.* (* As this was not observed in the sea, but in glass vessels containing sea-water, it might be supposed that the animals become exhausted and die, not because they are under water but because they have consumed all the oxygen which it contained. I therefore put into the same water from which I had just taken an unconscious Ocypoda, with its legs hanging loosely down, a specimen of Lupea diacantha which had been reduced to the same state by being kept in the air, and this recovered in the water just as the Ocypoda did in the air.) In these a peculiar arrangement on the feet of the third and fourth pairs (Figure 12) has long been known, although its connexion with the branchial cavity has not been suspected. These two pairs of feet are more closely approximated than the rest; the opposed surfaces of their basal joints (therefore the hinder surface on the third, and the anterior surface on the fourth feet) are smooth and polished, and their margins bear a dense border of long, silky, and peculiarly formed hairs (Figure 13). Milne-Edwards who rightly compares these surfaces, as to their appearance, with articular surfaces, thinks that they serve to diminish the friction between the two feet. In considering this interpretation, the question could not but arise why such an arrangement for the diminution of friction should be necessary in these particular Crabs and between these two feet, leaving out of consideration the fact that the remarkable brushes of hair, which on the other hand must increase friction, also remain unexplained. But as I was bending the feet of a large Sand-Crab to and fro in various directions, in order to see in what movements of the animal friction occurred at the place indicated, and whether these might, perhaps, be movements of particular importance to it and such as would frequently recur, I noticed, when I had stretched the feet widely apart, in the hollow between them a round orifice of considerable size, through which air could easily be blown into the branchial cavity, and a fine rod might even be introduced into it. The orifice opens into the branchial cavity behind a conical lobe, which stands above the third foot in place of a branchia which is wanting in Ocypoda. It is bounded laterally by ridges, which rise above the articulation of the foot, and to which the lower margin of the carapace is applied. Exteriorly, also, it is overarched by these ridges with the exception of a narrow fissure. This fissure is overlaid by the carapace, which exactly at this part projects further downwards than elsewhere, and in this way a complete tube is formed. Whilst in Grapsus the water is allowed to reach the branchiae only from the front, I saw it in Ocypoda flow in also through the orifice just described.
In the position of posterior entrant orifice and the accompanying peculiarities of the third and fourth pairs of feet, two other non-aquatic species of the same family, which I have had the opportunity of examining, agree with Ocypoda. One of these, perhaps Gelasimus vocans, which lives in the mangrove swamps, and likes to furnish the mouth of its burrow with a thick, cylindrical chimney of several inches in height, has the brushes on the basal joints of the feet in question composed of ordinary hairs. The other, a smaller Gelasimus, not described in Milne-Edwards' 'Natural History of Crustacea,' which prefers drier places and is not afraid to run about on the burning sand under the vertical rays of the noonday sun in December, but can also endure being in water at least for several weeks, resembles Ocypoda in having these brushes composed of non-setiform, delicate hairs, indeed even more delicate and more regularly constructed than in Ocypoda.* (* This smaller Gelasimus is also remarkable because the chameleon-like change of colour exhibited by many Crabs occurs very strikingly in it. The carapace of a male which I have now before me shone with a dazzling white in its hinder parts five minutes since when I captured it, at present it shows a dull gray tint at the same place.) What may be the significance of these peculiar hairs,—whether they only keep foreign bodies from the branchial cavity,—whether they furnish moisture to the air flowing past them,—or whether, as their aspect, especially in the small Gelasimus, reminds one of the olfactory filaments of the Crabs, they may also perform similar functions,—are questions the due discussion of which would lead us too far from our subject. Nevertheless it may be remarked that in both species, especially in Ocypoda, the olfactory filaments in their ordinary situation are very much reduced, and when they are in the water their flagella never perform the peculiar beating movements which may be observed in other Crabs, and even in the larger Gelasimus; moreover, the organ of smell must probably be sought in these air-breathing Crabs, as in the air-breathing Vertebrata, at the entrance to the respiratory cavity.
So much for the facts with regard to the aerial respiration of the Crabs. It has already been indicated why Darwin's theory requires that when any peculiar arrangements exist for aerial respiration, these will be differently constructed in different families. That experience is in perfect accordance with this requirement is the more in favour of Darwin, because the schoolmen far from being able to foresee or explain such profound differences, must rather regard them as extremely surprising. If, in the nearly allied families of the Ocypodidae and Grapsoidae, the closest agreement prevails in all the essential conditions of their structure; if the same plan of structure is slavishly followed in everything else, in the organs of sense, in the articulation of the limbs, in every trabecula and tuft of hairs in the complicated framework of the stomach, and in all the arrangements subserving aquatic respiration, even to the hairs of the flagella employed in cleaning the branchiae,—why have we suddenly this exception, this complete difference, in connection with aerial respiration?
The schoolmen will scarcely have an answer for this question, except by placing themselves on the theologico-teleological stand-point which has justly fallen into disfavour amongst us, and from which the mode of production of an arrangement is supposed to be explained, if its "adaptation" to the animal can be demonstrated. From this point of view we might certainly say that a widely gaping fissure which had nothing prejudicial in it to Aratus Pisonii among the foliage of the mangrove bushes, was not suitable to the Ocypoda living in sand; that in the latter, in order to prevent the penetration of the sand, the orifice of the branchial cavity must be placed at its lowest part, directed downwards, and concealed between broad surfaces fringed with protective brushes of hair. It is far from the intention of these pages to enter upon a general refutation of this theory of adaptation. Indeed there is scarcely anything essential to be added to the many admirable remarks that have been made upon this subject since the time of Spinoza. But this may be remarked, that I regard it as one of the most important services of the Darwinian theory that it has deprived those considerations of usefulness which are still undeniable in the domain of life, of their mystical supremacy. In the case before us it is sufficient to refer to the Gelasimus of the mangrove swamps, which shares the same conditions of life with various Grapsoidae and yet does not agree with them, but with the arenicolous Ocypoda.
CHAPTER 6. STRUCTURE OF THE HEART IN THE EDRIOPHTHALMA.
Scarcely less striking than the example of the air-breathing Crabs, is the behaviour of the heart in the great section Edriophthalma, which may advantageously be divided, after the example of Dana and Spence Bate, only into two orders, the Amphipoda and the Isopoda.
In the Amphipoda, to which the above-mentioned naturalists correctly refer the Caprellidae and Cyamidae (Latreille's Laemodipoda), the heart has always the same position; it extends in the form of a long tube through the six segments following the head, and has three pairs of fissures, furnished with valves, for the entrance of the blood, situated in the second, third, and fourth of these segments. It was found to be of this structure by La Valette in Niphargus (Gammarus puteanus), and by Claus in Phronima; and I have found it to be the same in a considerable number of species belonging to the most different families.* (* The young animals in the egg, a little before their exclusion, are usually particularly convenient for the observation of the fissures in the heart; they are generally sufficiently transparent, the movements of the heart are less violent than at a later period, and they lie still even without the pressure of a glass cover. Considering the common opinion as to the distribution of the Amphipoda, namely, that they increase in multiplicity towards the poles, and diminish towards the equator, it may seem strange that I speak of a considerable number of species on a subtropical coast. I therefore remark that in a few months and without examining any depths inaccessible from the shore, I obtained 38 different species, of which 34 are new, which, with the previously known species (principally described by Dana) gives 60 Brazilian Amphipoda, whilst Kroyer in his 'Gronlands Amfipoder' was acquainted with only 28 species, including 2 Laemodipoda, from the Arctic Seas, although these had been investigated by a far greater number of Naturalists.)
The sole unimportant exception which I have hitherto met with is presented by the genus Brachyscelus,* (*According to Milne-Edwards' arrangement the females of this genus would belong to the "Hyperines ordinaires" and the previously unknown males to the "Hyperines anormales," the distinguishing character of which, namely the curiously zigzagged inferior antennae, is only a sexual peculiarity of the male animals. In systematising from single dead specimens, as to the sex, age, etc. of which nothing is known, similar errors are unavoidable. Thus, in order to give another example of very recent date, a celebrated Ichthyologist, Bleeker, has lately distinguished two groups of the Cyprinodontes as follows: some, the Cyprinodontini, have a "pinna analis non elongata," and the others, the Aplocheilini, a "pinna analis elongata": according to this the female of a little fish which is very abundant here would belong to the first, and the male to the second group. Such mistakes, as already stated, are unavoidable by the "dry-skin" philosopher, and therefore excusable; but they nevertheless prove in how random a fashion the present systematic zoology frequently goes on, without principles or sure foundations, and how much it is in want of the infallible touchstone for the value of the different characters, which Darwin's theory promises to furnish.) in which the heart possesses only two pairs of fissures, as it extends forward only into the second body-segment, and is destitute of the pair of fissures situated in this segment in other forms.* (* I find, in Milne-Edwards' 'Lecons sur la Physiol. et l'Anat. comp.' 3 page 197, the statement that, according to Frey and Leuckart, the heart of Caprella linearis possesses FIVE pairs of fissures. I have examined perfectly transparent young Caprellae (probably the young of Caprella attenuata, Dana, with which they occurred), but can only find the usual three pairs.)
Considering this uniformity presented by the heart in the entire order of the Amphipoda, it cannot but seem very remarkable, that in the very next order of the Isopoda, we find it to be one of the most changeable organs.
In the cheliferous Isopods (Tanais) the heart resembles that of the Amphipoda in its elongated tubular form, as well as in the number and position of the fissures, but with this difference, that the two fissures of each pair do not lie directly opposite each other.
(FIGURE 14. Heart of a young Cassidina.
FIGURE 15. Heart of a young Anilocra.
FIGURE 16. Abdomen of the male of Entoniscus Cancrorum. h. Heart. l. Liver.)
In all other Isopoda the heart is removed towards the abdomen. In the wonderfully deformed parasitic Isopods of the Porcellanae (Entoniscus porcellanae), the spherical heart of the female is confined to a short space of the elongated first abdominal segment, and seems to possess only a single pair of fissures. In the male of Entoniscus Cancrorum (n. sp.), the heart (Figure 16) is situated in the third abdominal segment. In the Cassidinae, the heart (Figure 14) is likewise short and furnished with two pairs of fissures, situated in the last segment of the thorax and the first segment of the abdomen. Lastly, in a young Anilocra, I find the heart (Figure 15) extending through the whole length of the abdomen and furnished with four (or five?) fissures, which are not placed in pairs but alternately to the right and left in successive segments. In other animals of this order, which I have as yet only cursorily examined, further differences will no doubt occur. But why, in two orders so nearly allied to each other, should we find in the one such a constancy, in the other such a variability, of the same highly important organ? From the schoolmen we need expect no explanation, they will either decline the discussion of the "wherefore" as foreign to their province, as lying beyond the boundaries of Natural History, or seek to put down the importunate question by means of a sounding paraphrase of the facts, abundantly sprinkled with Greek words. As I have unfortunately forgotten my Greek, the second way out of the difficulty is closed to me; but as I luckily reckon myself not amongst the incorporated masters, but, to use Baron von Liebig's expression, amongst the "promenaders on the outskirts of Natural History," this affected hesitation of the schoolmen cannot dissuade me from seeking an answer, which indeed presents itself most naturally from Darwin's point of view.
As not only the Tanaides (which reasons elsewhere stated (vide supra) justify us in regarding as particularly nearly related to the primitive Isopod) and the Amphipoda, but also the Decapod Crustacea, possess a heart with three pairs of fissures essentially in the same position; and as the same position of the heart recurs (vide infra) even in the embryos of the Mantis-Shrimps (Squilla), in which the heart of the adult animal, and even, as I have elsewhere shown, that of the larvae when still far from maturity, extends in the form of a long tube with numerous openings far into the abdomen, we must unhesitatingly regard the heart of the Amphipoda as the primitive form of that organ in the Edriophthalma. As, moreover, in these animals the blood flows from the respiratory organs to the heart without vessels, it is very easy to see how advantageous it must be to them to have these organs as much approximated as possible. We have reason to regard as the primitive mode of respiration, that occurring in Tanais (vide supra). Now, where, as in the majority of the Isopoda, branchiae were developed upon the abdomen, the position and structure of the heart underwent a change, as it approached them more nearly, but without the reproduction of a common plan for these earlier modes of structure, either because this transformation of the heart took place only after the division of the primary form into subordinate groups, or because, at least at the time of this division, the varying heart had not yet become fixed in any new form. Where, on the contrary, respiration remained with the anterior part of the body,—whether in the primitive fashion of Zoea, as in the Tanaides, or by the development of branchiae on the thorax, as in the Amphipoda,—the primitive form of the heart was inherited unchanged, because any variations which might make their appearance were rather injurious than advantageous, and disappeared again immediately.
I close this series of isolated examples with an observation which indeed only half belongs to the province of the Crustacea to which these pages ought to be confined, and which also has no further connexion with the preceding circumstances than that of being an "intelligible and intelligence-bringing fact" only from the point of view of Darwin's theory. To-day as I was opening a specimen of Lepas anatifera in order to compare the animal with the description in Darwin's 'Monograph on the Subclass Cirripedia,' I found in the shell of this Cirripede, a blood-red Annelide, with a short, flat body, about half an inch long and two lines in breadth, with twenty-five body-segments, and without projecting setigerous tubercles or jointed cirri. The small cephalic lobe bore four eyes and five tentacles; each body-segment had on each side at the margin a tuft of simple setae directed obliquely upwards, and at some distance from this, upon the ventral surface, a group of thicker setae with a strongly uncinate bidentate apex. There was above EACH of the lateral tufts of bristles a branchia, simple on a few of the foremost segments, and then strongly arborescent to the end of the body. The animal, a female filled with ova, evidently, from these characters, belongs to the family of the Amphinomidae; the only family the members of which, being excellent swimmers, live in the open sea.
That this animal had not strayed accidentally into the Lepas, but appertained to it as a regular and permanent guest, is evidenced by its considerable size in proportion to the narrow entrance of the test of the Lepas, by the complete absence of the iridescence which usually distinguishes the skin of free Annelides and especially of the Amphinomidae, by the formation and position of the inferior setae, etc. But that a worm belonging to this particular family Amphinomidae living in the high sea, occurs as a guest in the Lepas, which also floats in the sea attached to wood, etc., is at once intelligible from the stand-point of the Darwinian theory, whilst the relationship of this parasite to the free-living worms of the open sea remains perfectly unintelligible under the supposition that it was independently created for dwelling in the Lepas.
But however favourable the examples hitherto referred to may be for Darwin, the objection may be raised against them, and that with perfect justice, that they are only isolated facts, which, when the considerations founded upon them are carried far beyond what is immediately given, may only too easily lead us from the right path, with the deceptive glimmer of an ignis fatuus. The higher the structure to be raised, the wider must be the assuring base of well-sifted facts.
Let us turn then to a wider field, that of the developmental history of the Crustacea, upon which science has already brought together a varied abundance of remarkable facts, which, however, have remained a barren accumulation of unmanageable raw-material, and let us see how, under Darwin's hand, these scattered stones unite to form a well-jointed structure, in which everything, bearing and being borne, finds its significant place. Under Darwin's hand! for I shall have nothing to do except just to place the building stones in the position which his theory indicates for them. "When kings build, the carters have to work."
CHAPTER 7. DEVELOPMENTAL HISTORY OF PODOPHTHALMA.
Let us first glance over the extant facts.
Among the Stalk-eyed Crustacea (Podophthalma) we know only a very few species which quit the egg in the form of their parents, with the full number of well-jointed appendages to the body. This is the case according to Rathke* in the European fresh-water Crayfish, and according to Westwood in a West Indian Land Crab (Gecarcinus). (* Authorities are cited only for facts which I have had no opportunity of confirming.) Both exceptions therefore belong to the small number of Stalk-eyed Crustacea which live in fresh water or on the land, as indeed in many other cases fresh-water and terrestrial animals undergo no transformations, whilst their allies in the sea have a metamorphosis to undergo. I may refer to the Earthworms and Leeches among the Annelida, which chiefly belong to the land and to fresh water,—to the Planariae of the fresh waters and the Tetrastemma of the sparingly saline Baltic among the Turbellaria,—to the Pulmonate Gasteropoda, and to the Branchiferous Gasteropoda of the fresh waters, the young of which (according to Troschel's 'Handb. der Zoologie') have no ciliated buccal lobes, although such organs are possessed by the very similar Periwinkles (Littorina).
All the marine forms of this section appear to be subject to a more or less considerable metamorphosis. This appears to be only inconsiderable in the common Lobster, the young of which, according to Van Beneden, are distinguished from the adult animal, by having their feet furnished, like those of Mysis, with a swimming branch projecting freely outwards. From a figure given by Couch the appendages of the abdomen and tail also appear to be wanting.
Far more profound is the difference of the youngest brood from the sexually mature animal in by far the greater majority of the Podophthalma, which quit the egg in the form of Zoea. This young form occurs, so far as our present observations go, in all the Crabs, with the sole exception of the single species investigated by Westwood. I say SPECIES, and not GENUS, for in the same genus, Gecarcinus, Vaughan Thompson found Zoea-brood,* which is also met with in other terrestrial Crabs (Ocypoda, Gelasimus, etc.). (* Bell ('Brit. Stalk-eyed Crust.' page 45) considers himself justified in "eliminating" Thompson's observation at once, because he could only have examined ovigerous females preserved in alcohol. But any one who had paid so much attention as Thompson to the development of these animals, must have been well able to decide with certainty upon eggs, if not too far from maturity or badly preserved, whether a Zoea would be produced from them. Moreover, the mode of life of the Land-Crabs is in favour of Thompson. "Once in the year," says Troschel's 'Handbuch der Zoologie,' "they migrate in great crowds to the sea in order to deposit their eggs, and afterwards return much exhausted towards their dwelling places, which are reached only by a few." For what purpose would be these destructive migrations in species whose young quit the egg and the mother as terrestrial animals?) All the Anomura seem likewise to commence their lives as Zoeae: witness the Porcellanae, the Tatuira (Hippa emerita) and the Hermit Crabs. Among the Macrura we are acquainted with the same earliest form principally in several Shrimps and Prawns, such as Crangon (Du Cane), Caridina (Joly), Hippolyte, Palaemon, Alpheus, etc. Lastly, it is not improbable, that the youngest brood of the Mantis-Shrimps (Squilla) is also in the same case.
The most important peculiarities which distinguish this Zoea-brood from the adult animal, are as follows:—
The middle-body with its appendages, those five pairs of feet to which these animals owe their name of Decapoda, is either entirely wanting, or scarcely indicated; the abdomen and tail are destitute of appendages, and the latter consists of a single piece. The mandibles, as in the Insecta, have no palpi. The maxillipedes, of which the third pair is often still wanting, are not yet brought into the service of the mouth, but appear in the form of biramose natatory feet. Branchiae are wanting, or where their first rudiments may be detected as small verruciform prominences, these are dense cell-masses, through which the blood does not yet flow, and which therefore have nothing to do with respiration. An interchange of the gases of the water and blood may occur all over the thin-skinned surface of the body; but the lateral parts of the carapace may unhesitatingly be indicated as the chief seat of respiration. They consist, exactly as described by Leydig in the Daphniae, of an outer and inner lamina, the space between which is traversed by numerous transverse partitions dilated at their ends; the spaces between these partitions are penetrated by a more abundant flow of blood than occurs anywhere else in the body of the Zoea. To this may be added that a constant current of fresh water passes beneath the carapace in a direction from behind forwards, maintained as in the adult animal, by a foliaceous or linguiform appendage of the second pair of maxillae (Figure 18). The addition of fine coloured particles to the water allows this current of water to be easily detected even in small Zoeae.
(FIGURE 17. Zoea of a Marsh Crab (Cyclograpsus ?), magnified 45 diam.
FIGURE 18. Maxilla of the second pair in the same species, magnified 180 diam.)
The Zoeae of the Crabs (Figure 17) are usually distinguished by long, spiniform processes of the carapace. One of these projects upwards from the middle of the back, a second downwards from the forehead, and frequently there is a shorter one on each side near the posterior inferior angles of the carapace. All these processes are, however, wanting in Maia according to Couch, and in Eurynome according to Kinahan; and in a third species of the same group of the Oxyrhynchi (belonging or nearly allied to the genus Achaeus) I also find only an inconsiderable dorsal spine, whilst the forehead and sides are unarmed. This is another example warning us to be cautious in deductions from analogy. Nothing seemed more probable than to refer back the beak-like formation of the forehead in the Oxyrhynchi to the frontal process of the Zoea, and now it appears that the young of the Oxyrhynchi are really quite destitute of any such process. The following are more important peculiarities of the Zoeae of the Crabs, although less striking than these processes of the carapace which, in combination with the large eyes, often give them so singular an appearance:—the anterior (inner) antennae are simple, not jointed, and furnished at the extremity with from two to three olfactory filaments; the posterior (outer) antennae frequently run out into a remarkably long spine-like process ("styliform process," Spence Bate), and bear, on the outside, an appendage, which is sometimes very minute ("squamiform process" of Spence Bate), corresponding with the antennal scale of the Prawns,* (* In a memoir on the metamorphoses of the Porcellanae I have erroneously described this appendage as the "flagellum.") and the first rudiment of the future flagellum is often already recognisable. Of natatory feet (afterwards maxillipeds) only two pairs are present; the third (not, as Spence Bate thinks, the first) is entirely wanting, or, like the five following pairs of feet, present only as a minute bud. The tail, of very variable form, always bears THREE pairs of setae at its hinder margin. The Zoeae of the Crabs usually maintain themselves in the water in such a manner that the dorsal spine stands upwards, the abdomen is bent forwards, the inner branch of the natatory feet is directed forwards, and the outer one outwards and upwards.
(FIGURES 19 TO 23. Tails of the Zoeae of various Crabs.
FIGURE 19. Pinnotheres.
FIGURE 20. Sesarma.
FIGURE 21. Xantho.
FIGURES 22 AND 23 of unknown origin.)
It is further to be remarked that the Zoeae of the Crabs, as also of the Porcellanae, of the Tatuira and of the Shrimps and Prawns, are enveloped, on escaping from the egg, by a membrane veiling the spinous processes of the carapace, the setae of the feet, and the antennae, and that they cast this in a few hours. In Achaeus I have observed that the tail of this earliest larval skin resembles that of the larvae of Shrimps and Prawns, and the same appears to be the case in Maia (see Bell, 'Brit. Stalk-eyed Crust.' page 44).
Widely as they seem to differ from them at the first glance, the Zoeae of the Porcellanae (Figure 24) approach those of the true Crabs very closely. The antennae, organs of the mouth, and natatory feet, exhibit the same structure. But the tail bears FIVE pairs of setae, and the dorsal spine is wanting, whilst, on the contrary, the frontal process and the lateral spines are of extraordinary length, and directed straight forward and backward.
(FIGURE 24. Zoea of Porcellana stellicola, F. Mull. Magnified 15 diam.
FIGURE 25. Zoea of the Tatuira (Hippa emerita), magnified 45 diam.
FIGURE 26. Zoea of a small Hermit Crab, magnified 45 diam.)
The Zoea of the Tatuira (Figure 25) also appears to differ but little from those of the true Crabs, which it likewise resembles in its mode of locomotion. The carapace possesses only a short, broad frontal process; the posterior margin of the tail is edged with numerous short setae.
The Zoea of the Hermit Crabs (Figure 26) possesses the simple inner antennae of the Zoea of the true Crabs; the outer antennae bear upon the outside on a short stalk a lamella of considerable size analogous to the scale of the antennae of the Prawns; on the inside, a short, spine-like process; and between the two the flagellum, still short, but already furnished with two apical setae. As in the Crabs, there are only two pairs of well-developed natatory feet (maxillipedes), but the third pair is also present in the form of a two-jointed stump of considerable size, although still destitute of setae. The tail bears five pairs of setae. The little animal usually holds itself extended straight in the water, with the head directed downwards.
This is also the position in which we usually see the Zoeae of the Shrimps and Prawns (Figure 27), which agree in their general appearance with those of the Hermit Crabs. Between the large compound eyes there is in them a small median eye. The inner antennae bear, at the end of a basal joint sometimes of considerable length, on the inside a plumose seta, which also occurs in the Hermit Crabs, and on the outside a short terminal joint with one or more olfactory filaments. The outer antennae exhibit a well-developed and sometimes distinctly articulated scale, and within this usually a spiniform process; the flagellum appears generally to be still wanting. The third pair of maxillipedes seems to be always present, at least in the form of considerable rudiments. The spatuliform caudal lamina bears from five to six pairs of setae on its hinder margin.
The development of the Zoea-brood to the sexually mature animal was traced by Spence Bate in Carcinus maenas. He proved that the metamorphosis is a perfectly gradual one, and that no sharply separated stages of development, like the caterpillar and pupa of the Lepidoptera, could be defined in it. Unfortunately we possess only this single complete series of observations, and its results cannot be regarded at once as universally applicable; thus the young Hermit Crabs retain the general aspect and mode of locomotion of Zoeae, whilst the rudiments of the thoracic and abdominal feet are growing, and then, when these come into action, appear at once in a perfectly new form, which differs from that of the adult animal chiefly by the complete symmetry of the body and by the presence of four pairs of well-developed natatory feet on the abdomen.* (* Glaucothoe Peronii, M.-Edw., may be a young and still symmetrical Pagurus of this kind.)
(FIGURE 27. Zoea of a Palaemon residing upon Rhizostoma cruciatum, Less., magnified 45 diam.)
The development of the Palinuridiae seems to be very peculiar. Claus found in the ova of the Spiny Lobster (Palinurus), embryos with a completely segmented body, but wanting the appendages of the tail, abdomen, and last two segments of the middle-body; they possess a single median and considerably compound eye; the anterior antennae are simple, the posterior furnished with a small secondary branch; the mandibles have no palpi; the maxillipedes of the third pair, like the two following pairs of feet, are divided into two branches of nearly equal length; whilst the last of the existing pairs of feet and the second pair of maxillipedes bear only an inconsiderable secondary branch. Coste, as is well known, asserts that he has bred young Phyllosomata from the ova of this lobster—a statement that requires further proof, especially as the more recent investigations of Claus upon Phyllosoma by no means appear to be in its favour.
The large compound eyes, which usually soon become moveable, and sometimes stand upon long stalks even in the earliest period, as well as the carapace, which covers the entire fore-body, indicate at once that the position of the larvae hitherto considered, notwithstanding all their differences, is under the Podophthalma. But not a single characteristic of this section is retained by the brood of some Prawns belonging to the genus Peneus or in its vicinity. These quit the egg with an unsegmented ovate body, a median frontal eye, and three pairs of natatory feet, of which the anterior are simple, and the other two biramose—in fact, in the larval form, so common among the lower Crustacea, to which O.F. Muller gave the name of Nauplius. No trace of a carapace! no trace of the paired eyes! no trace of masticating organs near the mouth which is overarched by a helmet-like hood!
(FIGURE 28. Nauplius of a Prawn, magnified 45 diam.
FIGURE 29. Young Zoea of the same Prawn, magnified 45 diam.
FIGURE 30. Older Zoea of the same Prawn, magnified 45 diam.
FIGURE 31. Mysis-form of the same Prawn, magnified 45 diam.)
In the case of one of these species the intermediate forms which lead from the Nauplius to the Prawn, have been discovered in a nearly continuous series.
The youngest Nauplius (Figure 28) is immediately followed by forms in which a fold of skin runs across the back behind the third pair of feet, and four pairs of stout processes (rudiments of new limbs) sprout forth on the ventral surface. Within the third pair of feet, powerful mandibles are developed.
In a subsequent moult the new limbs (maxillae, and anterior and intermediate maxillipedes) come into action, and in this way the Nauplius becomes a Zoea (Figure 29), agreeing perfectly with the Zoea of the Crabs in the number of the appendages of the body, although very different in form and mode of locomotion and even in many particulars of internal structure. The chief organs of motion are still the two anterior pairs of feet, which are slender and furnished with long setae; the third pair of feet loses its branches, and becomes converted into mandibles destitute of palpi. The labrum acquires a spine directed forward and of considerable size, which occurs in all the Zoeae of allied species. The biramose maxillipedes appear to assist but slightly in locomotion. The forked tail reminds us rather of the forms occurring in the lower Crustacea, especially the Copepoda, than of the spatuliform caudal plate which characterises the Zoeae of Alpheus, Palaemon, Hippolyte, and other Prawns, of the Hermit Crabs, the Tatuira and the Porcellanae. The heart possesses only one pair of fissures, and has no muscles traversing its interior like trabeculae, whilst in other Zoeae two pairs of fissures and an interior apparatus of trabeculae are always distinctly recognisable.
During this Zoeal period the paired eyes, the segments of the middle-body and abdomen, the posterior maxillipedes, the lateral caudal appendages and the stump-like rudiments of the feet of the middle-body are formed (Figure 30). The caudal appendages sprout forth like other limbs freely on the ventral surface, whilst in other Prawns, the Porcellanae, etc., they are produced in the interior of the spatuliform caudal plate.
As the feet of the middle-body come into action, simultaneously with other profound changes, the Zoea passes into the Mysis- or Schizopod-form (Figure 31). The antennae cease to serve for locomotion, their place is taken by the thoracic feet, furnished with long setae, and by the long abdomen which just before was laboriously dragged along as a useless burden, but now, with its powerful muscles, jerks the animal through the water in a series of lively jumps. The anterior antennae have lost their long setae, and by the side of the last (fourth) joint, endowed with olfactory filaments, there appears a second branch, which is at first of a single joint. The previously multi-articulate outer branch of the posterior antennae has become a simple lamella, the antennal scale of the Prawn; beside this appears the stump-like rudiment of the flagellum, probably as a new formation, the inner branch disappearing entirely. The five new pairs of feet are biramose, the inner branch short and simple, the outer one longer, annulated at the end, furnished with long setae, and kept, as in Mysis, in constant whirling motion. The heart acquires new fissures, and interior muscular trabeculae.
During the Mysis-period, the auditory organs in the basal joint of the anterior antennae are formed; the inner branches of the first three pairs of feet are developed into chelae and the two hinder pairs into ambulatory feet; palpi sprout from the mandibles, branchiae on the thorax, and natatory feet on the abdomen. The spine on the labrum becomes reduced in size. In this way the animal gradually approaches the Prawn-form, in which the median eye has become indistinct, the spine of the labrum, and the outer branches of the cheliferous and ambulatory feet have been lost, the mandibular palpi and the abdominal feet have acquired distinct joints and setae, and the branchiae come into action.
In another Prawn, the various larval states of which may be easily recognised as belonging to the same series by the presence of a dark-yellow, sharply-defined spot surrounding the median eye, the youngest Zoea (Figure 32), probably produced from the Nauplius, agrees in all essential particulars with the species just described; its further development is, however, very different, especially in that neither the feet of the middle, nor those of the hind-body are formed simultaneously, and that a stage of development comparable to Mysis in the number and structure of the limbs does not occur.
(FIGURE 32. Youngest (observed) Zoea of another Prawn. The minute buds of the third pair of maxillipedes are visible. The formation of the abdominal segments has commenced. Paired eyes still wanting. Magnified 45 diam.)
Traces of the outer maxillipedes make their appearance betimes. Then feet appear upon four segments of the middle-body, and these are biramose on the three anterior segments, and simple, the inner branch being deficient, on the fourth segment. On the inner branches the chelae are developed; the outer branches are lost before an inner branch has made its appearance on the fourth segment (Figure 32). The latter again becomes destitute of appendages, so that in this case at an early period four, and at a later only three, segments of the middle-body bear limbs. The fifth segment is still entirely wanting, whilst all the abdominal segments have also acquired limbs, and this one after the other, from before backwards. The adult animal, as shown by the three pairs of chelae, will certainly be very nearly allied to the preceding species.* (* The oldest observed larvae (see Figure 33) are characterised by the extraordinary length of the flagella of the outer antennae, and in this respect resemble the larva of Sergestes found by Claus near Messina (Zeitschr. fur Wiss. Zool. Bd. 13 Taf 27 Figure 14). This unusual length of the antennae leads to the supposition that they belong to our commonest Prawn, which is very frequently eaten, and is most nearly allied to Peneus setiferus of Florida. Claus's Acanthosoma (l.c. Figure 13) is like the younger Mysis-form of the larva figured by me in the 'Archiv fur Naturgeschichte,' 1836, Taf 2, Figure 18, and which I am inclined to refer to Sicyonia carinata.)
The youngest larva of the Schizopod genus Euphausia observed by Claus, stands very near the youngest Zoea of our Prawns; but whilst its anterior antennae are already biramose, and it therefore appears to be more advanced, it still wants the middle maxillipedes. In it also Claus found the heart furnished with only a single pair of fissures. Do not Nauplius-like states in this case also precede the Zoea?
(FIGURE 33. Older larva produced from the Zoea represented in Figure 32. The last segment and the last two pairs of feet of the middle-body are wanting. Magnified 20 diam.)
The developmental history of Mysis, the near relationship of which with the Shrimps and Prawns has recently again been generally recognised, has been described in detail by Van Beneden. So far as I have tested them I can only confirm his statements. The development of the embryo commences with the formation of the tail! This makes its appearance as a simple lobe, the dorsal surface of which is turned towards and closely applied to that of the embryo. (The young of other Stalk-eyed Crustacea are, as is well known, bent in the egg in such a manner that the ventral surfaces of the anterior and posterior halves of the body are turned towards each other,—in these, therefore, the dorsal, and in Mysis the ventral surface appears convex.) The tail soon acquires the furcate form with which we made acquaintance in the last Prawn-Zoea described. Then two pairs of thick ensiform appendages make their appearance at the opposite end of the body, and behind these a pair of tubercles which are easily overlooked. These are the antennae and mandibles. The egg-membrane now bursts, before any internal organ, or even any tissue, except the cells of the cutaneous layer, is formed. The young animal might be called a Nauplius; but essentially there is nothing but a rough copy of a Nauplius-skin, almost like a new egg-membrane, within which the Mysis is developed. The ten pairs of appendages of the fore- (maxillae, maxillipedes) and middle-body make their appearance simultaneously, as do the five pairs of abdominal feet at a later period. Soon after the young Mysis casts the Nauplius-envelope it quits the brood-pouch of the mother.* (* Van Beneden, who regards the eye-peduncles as limbs, cannot however avoid remarking upon Mysis: "Ce pedicule n'apparait aucunement comme les autres appendices, et parait avoir une autre valeur morphologique.")