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Chapter II: Front Matter (2)

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The scutes which protect the _ventral_ side of the body, from the throat backwards, are four-sided and similar in their ornamentation to the dorsal scutes; but they exhibit neither ridge nor angulation, their outer and inner surfaces being parallel, and either nearly flat or evenly curved. Each forms, in fact, a segment of a large cylinder, inasmuch as the whole ventral shield is convex transversely, being nearly flat in the middle and much bent up at the sides. The dorsal shield, taken as a whole, is, on the contrary, nearly flat. The lateral edges of the ventral scutes interlock suturally; and their anterior and posterior edges are overlapped and overlap, just like the dorsal scutes. The outer edges of the outermost ventral scutes thin off and are not united with any bony element; and the ventral, like the dorsal scutes, are usually arranged symmetrically on either side of the median sutural line. There may be as many as twenty-two scutes united by their lateral sutures into a single strong, curved, transverse, bony, bar-like segment of the ventral armour.

Throughout the neck and body, and as far as the commencement of the tail, the ends of the dorsal and ventral bony bars, whose sum may be regarded as a dorsal and a ventral shield respectively, are separated by an interval of integument, in which only small scattered scutes are visible. The physiological import of this arrangement becomes obvious when we consider in what manner the animal breathes; and indeed the integumentary interval answers very precisely to the leather which connects the two boards of a bellows. Again, though the limbs are themselves covered with articulated scutes, they are afforded free play upon the body by this flexible interspace. Immediately behind the hind legs, the ventral and dorsal shields unite; and the tail is from that point surrounded by a succession of bony hoops, each of which corresponds with a vertebra, the segments of the exoskeleton answering to those of the endoskeleton.

The most remarkable feature about the ventral scutes, however, and that in which they differ most widely from the dorsal ones, consists in the fact that each scute is composed of two distinct pieces, an anterior and a posterior, which unite together by a transverse serrated suture. The anterior piece or 'semi-scute' may attain to three-quarters the length of the posterior, and it has exactly the same width. The anterior semi-scute bears the articular facet and the transverse pitted groove, whose posterior wall is just in front of its hinder edge, or in other words, of the suture, when the two semi-scutes are united.

Such are the general characters and mode of arrangement of the dorsal and ventral armour of _Jacare_. But there remain many noteworthy peculiarities in the disposition and number of the components of each band of the armour.

Thus, in the _dorsal shield_ there are two rows of nuchal scutes, each containing eight separate keeled bony plates; and of cervical scutes there are five rows, the two anterior of which contain four angulated and carinated scutes each, while the three posterior contain only two scutes each. All these scutes, except the anterior row, have articular facets; and all those of each row are united suturally. Of dorsal scutes there are thirty transverse rows up to the median keel of the tail, which commences with the thirty-first row. The number of scutes in each row is as follows:--

Rows. Scutes.
1, 2, 3, 4 6
5, 6, 7, 8, 9, 10, 11 10
12, 13 8
14, 15 6
16, 17, 18 4
19 6
20 8
23, 24 6
25, 26 5
27, 28 4
29, 30 4

31, 32, 33, 34 5
The rest of the tail is wanting.

Throughout the dorso-lumbar and sacral regions (_i. e._ up to the nineteenth row), the median scutes are hardly keeled at all, while the outer ones are the more strongly carinate the more external they lie.

In the caudal region, the second scute from the middle line, in the twenty-third row, has a strong keel and angulation, which grows stronger in the corresponding scutes up to the thirtieth inclusive, until the superior and lateral faces of these scutes, in the twenty-ninth and thirtieth rows, are inclined to one another at a right angle and very strongly keeled. I have said that, as a rule, the median line is occupied by a suture between two median scutes; but in the caudal region[8], in the twenty-fifth row (which corresponds with the sixth caudal vertebra) the two median scutes are replaced by one flat scute, so that there is no suture in the middle line. In the twenty-sixth row there is a similar arrangement, but the flat scute is smaller; and in the twenty-seventh no trace of it is left, so that the strongly keeled lateral scutes meet in the middle line, which is again occupied by a suture. This continues up to the thirty-first row, when the median scute reappears as a thin vertical plate, broader below and in front, where it articulates with the median lateral scutes, than above and behind, where it exhibits a free edge only covered by the horny epidermis. It is thus that the serrated dorsal crest of the tail is formed. The scutes of the crest exhibit only very small round and distant pits.

The _ventral shield_ begins in the neck just behind the level of the anterior margins of the orbits: the fifteen anterior rows may be termed subcervical, as they lie in front of the thorax. In the first six rows the scutes are very small, and increase in number up to twelve in a row. In the next six rows there are ten scutes in a row, and in the last three, twelve. All these rows are symmetrically divided by the median line. In the three hinder rows the inner scutes are longer than the outer ones; and this is most markedly the case in the fifteenth row, whose innermost scute is half as long again as the corresponding one of the preceding row, and more than three times as long as the outermost of its own row.

[8] The second and third cervical rows in _Caiman palpebrosus_ and
_trigonatus_ also contain a median scute, and consequently an odd
number of scutes. In _Caiman trigonatus_, the third to the ninth
supra-caudal rows have each a median single scute.

The sixteenth row differs from its predecessors and successors, and may be termed the axillary row. It is bent upon itself with an angle open forwards, and is divided into two halves (each of which contains seven scutes) by the union of the middle scutes of the fifteenth subcervical with those of the first row of what may be termed the subdorsal scutes, or those which lie under the thorax and abdomen. Of subdorsal and subcaudal scutes there are, up to the broken-off end of the tail, thirty-seven rows, with the following numbers of scutes:--

Rows. Scutes.

1 12
2 10
3, 4, 5, 12
6, 7, 8, 9, 14
10 16
11 14
12-17 14
18-20 12
21 14
22 18
23 22
24 22
25 20
26-28 18
29-31 16
32-34 14
35 12
36, 37 10

It will be noticed that there are three more rows of ventral than of dorsal scutes. On endeavouring to ascertain how this came about, I observed that the first subdorsal was a good deal behind the first dorsal row, though the eighth to the twelfth dorsal corresponded exactly with the eighth to the twelfth ventral rows. In the anterior part of the body, therefore, there is a clear general correspondence between the segments of the dorsal and those of the ventral armour.

In the caudal region, again, I found that the twenty-fourth ventral row, which is the first of the caudal rows not excavated by the vent, corresponded exactly with the twenty-first dorsal row. It was clear, therefore, that three ventral rows wore interpolated somewhere between the twelfth and twenty-first dorsal rows; and on close examination I found this interpolation to arise from the doubling of the fourteenth, fifteenth, and sixteenth ventral rows.

I have examined _Jacare fissipes_ and _nigra_, _Caiman trigonatus_, and _C. gibbiceps_, in the British Museum; and I find, in all, dorsal and ventral armour having the same essential arrangement as that just described. A specimen of _Caiman palpebrosus_ about two feet long, the opportunity of examining which I owe to Dr. Grant, exhibits the dorsal and ventral shields (whose scutes are in the main similarly arranged) very beautifully; and a young _Jacare_ of about 18 inches in length, for which I am indebted to the kindness of the same gentleman, proves that the scutes are developed even in specimens of this age. I have no hesitation therefore in expressing my belief that this singularly complete dermal armour will be found to be characteristic of all the species of the genera _Caiman_ and _Jacare_. On the other hand, I have examined _Alligator Mississipiensis_, _Crocodilus vulgaris_, _C. biporcatus_, _C. Americanus_, _C. rhombifer_, and _C. bombifrons_, _Mecistops cataphractus_, and _Gavialis Gangeticus_, of various ages and sizes, without having been able to discover a trace of ventral scutes. This is the more remarkable, as the well-marked ventral and dorsal shields of many of the ancient _Teleosauria_ would lead one to expect a corresponding exoskeleton (if anywhere) in their nearest allies, the modern _Gavialidæ_. However, _Goniopholis_, with its strong armour, is more like an ordinary Crocodile; and I have recently discovered that a true Crocodile in some respects curiously similar to _C. bombifrons_ (_C. Hastingsiæ_) was covered with scutes exceedingly like those of the modern _Caiman_ and _Jacare_.

In minute structure the bony scutes of _Jacare_ closely resemble those of such a fish as a Sturgeon: a middle layer, containing so many canals as to appear almost cancellated in longitudinal or transverse section, is covered externally by a thin, and internally by a thick, layer composed of bony lamellæ, nearly parallel to the plane of the scute. Round the canals of the middle layer, the bony lamellæ are disposed concentrically, to a greater or less extent. The lacunæ are of very various shapes; and there are perhaps as many short as elongated forms. The canals of the middle layer communicate by large branches with the inner, by smaller and fewer branches with the outer surface of the scute.

In the young _Jacare_ mentioned above, I found the dermis to be distinguishable into two layers. The more superficial of these is thin, made up of irregular or formless connective tissue, and contains many ramified pigment-masses. Its smooth outer surface underlies the rete mucosum. Internally, it passes into the second or deep layer, which consists of successive layers of distinctly fibrous connective tissue, disposed in definite parallel bundles, and having a very regular arrangement. Throughout a space corresponding with the area of each scale, in fact, the bundles of each layer cross those of the succeeding layer at right angles; and the successive tiers of bundles are tied together by short cords disposed perpendicularly to the planes of the tiers. A corresponding arrangement of the bundles of connective tissue has long been known to obtain in the dermis of Fishes and _Batrachia_. At each end of this small "mat" of connective tissue, the bundles, if I may so say, fray out; and at the anterior end, the layers, loosened in texture, bend upwards, spreading out at the same time to become continuous with the fibres of the "mat" in front. In consequence of the matting under the quadrate surface of each scale, the dermis has a peculiar facetted aspect, quite apart from any osseous deposit. Where bony scutes are formed, they appear as very thin perforated plates in the most superficial portion of the deep layer of the dermis; so that there is a single thin layer of dense connective tissue above them, while below them are all the rest of the denser and deeper lamellæ of the dermis. Through the apertures in this primitive osseous plate (the rudiment of the middle layer of the future scute), bundles of connective tissue extend, connecting the deep with the superjacent lamellæ.

If a thin section is made and decalcified with weak acid under the microscope, the calcareous matter, as it is dissolved away, leaves an obscurely fibrous matrix of a different aspect from the surrounding connective tissue, and the endoplasts, or nuclei, of this matrix are seen each to have occupied the centre of a lacuna.

Again, the rudimentary scute lies in the dermis as in a sort of pocket, the superficial and deep walls of which separate from it with great ease; and in good thin sections made through the dermis and scute, there seems to be no direct connexion between the substance of the scute above and below, and the connective tissue with which it is in contact. Nor could I satisfy myself that the margins of the scute were continuous with the surrounding bundles of connective tissue. However, the specimen had been a very long time in spirit; and I am unwilling to lay too much stress upon these observations, which tend to negative the supposition that the scute proceeds from the direct calcification of the connective tissue of the dermis.

On the other hand, I must remark that horizontal sections of the scutes have presented oblique parallel fissures, sometimes crossing one another, which might readily be supposed to correspond with the lines of separation of ossified bundles of connective tissue.

* * * * *

NOTE.--During a recent visit to Paris, my friend Mr. Busk was kind enough to examine the specimens of recent _Crocodilia_ in the Museum of the Jardin des Plantes, with reference to certain points to which I requested his attention. Mr. Busk informs me that there is no doubt about the transverse direction of the premaxillo-maxillary suture in _Crocodilus rhombifer_; and his statements lead me to entertain no question that _C. bombifrons_ is a synonym of _C. palustris_.

In the typical specimens of _C. marginatus_ and _C. suchus_ of Geoffroy St.-Hilaire, the premaxillo-maxillary suture extends back to the level of the seventh tooth.

Mr. Busk has furthermore pointed out to me the existence of another American species of Crocodile--_C. Morelettii_, which has been described by M. Auguste Duméril in his "Description des Reptiles nouveaux ou imparfaitement connus," &c., 'Archives du Muséum,' t. vi. 1852.

This species inhabits lake Flores, in Yucatan; and it is said by M. Duméril to approach _C. Americanus_, from which it differs in the proportions of the skull and in the characters of the dermal armour.

June 21st, 1859.

* * * * *

On the Habits of the "Aye-Aye" (_Cheiromys madagascariensis_, L., Cuv.). By the Hon. H. SANDWITH, M.D., C.B., Colonial Secretary of the Mauritius. Communicated by Prof. OWEN, F.R.S., V.P.L.S.

[Read April 7th, 1859.]

"Mauritius, Jan. 27, 1859.

"MY DEAR MR. OWEN,--After very great difficulty and much delay, I have at length obtained a fine healthy male adult _Aye-Aye_; and he is now enjoying himself in a large cage which I have had constructed for him.

He is a most interesting little animal; and from close observation I have learnt his habits very correctly. On receiving him from Madagascar, I was told that he ate bananas; so of course I fed him on them, but tried him with other fruit. I found he liked dates,--which was a grand discovery, supposing he be sent alive to England. Still I thought that those strong rodent teeth, as large as those of a young Beaver, must have been intended for some other purpose than that of trying to eat his way out of a cage--the only use he seemed to make of them, besides masticating soft fruits. Moreover, he had other peculiarities,--_e.g._, singularly large, naked ears directed forward, as if for offensive rather than defensive purposes; then, again, the second finger of the hands is unlike anything but a monster supernumerary member, it being slender and long, half the thickness of the other fingers, and resembling a piece of bent wire. Excepting the head and this finger, he closely resembles a Lemur.

Now as he attacked, every night, the woodwork of his cage, which I was gradually lining with tin, I bethought myself of tying some sticks over the woodwork, so that he might gnaw these instead. I had previously put in some large branches for him to climb upon; but the others were straight sticks to cover over the woodwork of his cage, which _alone_ he attacked. It so happened that the thick sticks I now put into his cage were bored in all directions by a large and destructive grub called here the _Moutouk_. Just at sunset the Aye-Aye crept from under his blanket, yawned, stretched, and betook himself to his tree, where his movements are lively and graceful, though by no means so quick as those of a squirrel. Presently he came to one of the worm-eaten branches, which he began to examine most attentively; and bending forward his ears, and applying his nose close to the bark, he rapidly tapped the surface with the curious second digit, as a woodpecker taps a tree, though with much less noise, from time to time inserting the end of the slender finger into the worm-holes, as a surgeon would a probe. At length he came to a part of the branch which evidently gave out an interesting sound, for he began to tear it with his strong teeth. He rapidly stripped off the bark, cut into the wood, and exposed the nest of a grub, which he daintily picked out of its bed with the slender tapping finger, and conveyed the luscious morsel to his mouth.

I watched these proceedings with intense interest, and was much struck with the marvellous adaptation of the creature to its habits, shown by his acute hearing, which enables him aptly to distinguish the different tones emitted from the wood by his gentle tapping; his evidently acute sense of smell, aiding him in his search; his secure footsteps on the slender branches, to which he firmly clung by his quadrumanous members; his strong rodent teeth, enabling him to tear through the wood; and lastly by the curious slender finger, unlike that of any other animal, and which he used alternately as a pleximeter, a probe, and a scoop.

But I was yet to learn another peculiarity. I gave him water to drink in a saucer, on which he stretched out a hand, dipped a finger into it, and drew it obliquely through his open mouth; and this he repeated so rapidly, that the water seemed to flow into his mouth. After a while he lapped like a cat; but his first mode of drinking appeared to me to be his way of reaching water in the deep clefts of trees.

I am told that the _Aye-Aye_ is an object of veneration at Madagascar, and that if any native touches one, he is sure to die within the year; hence the difficulty of obtaining a specimen. I overcame this scruple by a reward of £10.

I quite despair of obtaining the bones of the _Dinornis_ or _Dodo_, though I have made every effort. I shall always be proud to be of service.

Believe me, yours very faithfully,

H. SANDWITH."

* * * * *

On the Moulting of the common Lobster (_Homarus vulgaris_) and Shore Crab (_Carcinus mænas_). By S. JAMES A. SALTER, M.B., F.L.S., F.G.S.

[Read April 7th, 1859.]

I am induced to bring this subject before the Linnean Society, on account of the singularly perfect specimen of the thrown-off slough of a Lobster which I have now an opportunity of exhibiting, and because the process by which it was shed was witnessed and carefully watched by two competent observers--by my friend Mr. Robert Cooke, of Scarborough, a Fellow of this Society, and by the intelligent wife of the Curator of the Scarborough Museum, in an aquarium in which institution the occurrence took place.

The methods by which certain of the Decapod Crustaceans cast their old shells in the process of renewal and growth have already been made the subject of observation and record.

Réaumur, as early as 1712, and again in 1718, saw and described the sloughing of the common freshwater Crayfish (_Astacus fluviatilis_).

It was witnessed in the common edible Crab (_Cancer Pagurus_) by Mr. Couch, in 1833.

Subsequently the moulting-process was observed by Mr. Gosse, in the Spinous Spider-crab (_Maia Squinado_).

Beyond these three recorded examples, I believe that the actual operation of moulting in Decapods has never been seen, though the sloughs of our common Crustacea, and the animals themselves but recently emerged from their old shells, are familiar to all marine zoologists.

There is no recorded account of the moulting of the Lobster, that I have been able to discover.

The Lobster from which the slough was obtained, and whose operations are the subject of this communication, was an inhabitant of a large marine aquarium in the Museum at Scarborough. The period was July 1857. The aquarium contained the ordinary assemblage of sea-shore animals, and a considerable collection of vegetation, which consisted of _Ulva_, _Fucus_, and other common sea-weeds.

For two days previous to its throwing off the shell, the Lobster was observed in a very peculiar attitude, and to be very busily engaged. Its abdomen was permanently and stiffly erected and straight; while the animal, in this rigid attitude, was hard at work detaching and carrying all the soft sea-weed it could collect to one end of the aquarium, where it thus accumulated a large mass of vegetation, which was afterwards destined to become a screen and protection for its soft body. At the same time, and by the same means, a clearing was made at the other end of the tank, in which it had space for the evolutions which were subsequently necessary for the extrication of its body.

The Lobster remained in the peculiar rigid attitude I have described, during the entire two days previous to the moult. On the third day, a crack was observed along the membrane which unites the dorsal surface of the first abdominal ring with the carapace; and when these parts became separated by about half an inch, the bright-blue membrane of the new shell being plainly visible beneath, the operation of extricating the abdomen commenced. By a strong vibratory action of the whole abdomen, principally in a lateral direction, one segment was, at first, protruded through the split; and this was followed by an interval of complete repose, during which the animal remained quite motionless. Then, by another vibratory action, the second segment was extricated; then followed an interval of repose, when the third was withdrawn; and so on till, at last, the entire abdomen, after having been bent double upon itself, was turned completely out backwards, and then, elongated and compressed, remained above and parallel to the empty shell that it had occupied, and which was still attached to the under surface of the cephalothorax. Hitherto the only orifice of escape consisted in the transverse splitting of the first abdominal segment from the carapace, on the dorsal surface. None of the abdominal segments separated from each other.

Thus far the extrication had commenced at the front of the abdomen, and had progressed from before backwards. It was now observed that the carapace had split from behind forwards, the fissure commencing posteriorly at the transverse split between _the_ carapace and the first abdominal segment, and reaching forwards to the apex of the rostrum, which, however, it did not absolutely divide. The two halves of the carapace then separating posteriorly, the interval between them, together with the original transverse slit, constituted a trifid opening, through which the rest of the animal escaped.

The escape of the cephalo-thoracic portion was effected from behind forwards. First the posterior ambulatory legs were loosened and withdrawn; then followed the next pair; and this process was continued from behind forwards, pair by pair--the withdrawal of each pair of legs being followed by an interval of repose. The limbs were withdrawn very readily from the old shell, slipping out of it as a leg would from a loose boot. No apparent effort accompanied these operations so far.

The extrication of the claws, however, was attended with much and violent exertion. This consisted of two powerful and sudden tugs, the soft abdomen of the Lobster pressing by its under surface upon the upper surface of the empty shell. By this means the soft chelæ were drawn through the narrow joints of the old shell, exhibiting strong, unmistakeable marks of the violence and pressure to which they had been subjected. The escape of the chelæ from their unyielding incasement was not aided by any splitting of the old shell, the large soft hands being drawn by compression through the narrow joints, as a wire is drawn through the contracting holes of a draw-plate.

The efforts for the withdrawal of the chelæ were the last, and succeeded in completely freeing the Lobster from its old case. Not only the claws, but the parts of the mouth, the antennæ, and the eyes, were all unsheathed; and with the last tug the regenerate Lobster plunged backwards, and entirely escaped, above and behind the now empty shell--its former tenement.

The operation, from first to last, occupied about twenty minutes, and was performed entirely in view, in that part of the aquarium which the Lobster had cleared of sea-weed.

Immediately after emerging from the old shell, the Lobster, was much deformed: there was a general elongation of the whole animal; but this was most remarkably the case with the claws, which were quite drawn out of shape. During the few subsequent hours, both the body and the claws became shorter and much enlarged. This increase of size did not result from any unfolding of membrane of the shell previously plicated, as no folds were observable immediately after the emergence of the animal, but from a simple distension, apparently from the imbibition, either by swallowing or by endosmosis, of considerable quantities of water. The membrane of the new shell was perfectly soft, and of a bright blue colour. At first the Lobster was shy and quite inactive, retiring to and remaining concealed among the accumulated sea-weed; but in a few hours it emerged from its retreat, and moved freely about the aquarium. The membrane of the new shell remained soft for some days, but on the seventh it appeared to have become perfectly calcified.

These are the details of the exuviation of the Lobster whose cast-off shell is before the Society. By a happy accident, the same observers had an opportunity of witnessing the sloughing of another Lobster, in the month of November following. The process was identically the same in every particular; but it was observed that the subsequent calcification of the shell did not take place till after the lapse of about fourteen days,--a circumstance probably dependent on a lower temperature and a less active nutrition. These are, I believe, the only two instances in which the exuviation of the Lobster has been actually witnessed; but there exist specimens of sloughs which are entirely in keeping with this description. In the fish-house of the Zoological Society of London there are two specimens which were cast in the tanks there; and in each there is the same transverse splitting of the carapace from the abdomen, and the longitudinal splitting of the carapace itself, without any other opening for the escape of the animal.

One or two general observations are suggested by the foregoing description. In the only examples of the exuviation of macrourous Decapod Crustaceans, there exists a singular diversity in the process itself. In _Astacus_, as described by Réaumur, the process commences with the escape of the cephalothorax; in _Homarus_, as I have now described it, it begins by the emergence of the abdomen. In _Astacus_ the carapace is detached and thrown off bodily and unbroken, being severed from its attachments with the lateral portions of the cephalothorax, as is the case in the Brachyura; whereas in _Homarus_ the lateral attachments of the carapace remain, whilst the plate itself is split up the centre. In _Astacus_, as is also the case in the Brachyura, the thrown-off slough is uniformly left resting on its dorsal surface; in _Homarus_ the reverse is uniformly the case. But the most striking dissimilarity is to be found in the circumstances _stated_ to attend the liberation of the chelæ. Prof. Bell, in the Introduction to his 'History of the British Stalk-eyed Crustacea,' remarks--"It is impossible to imagine that the crust of the legs, and especially of the great claws of the larger species, could be cast off, unless it were susceptible of being longitudinally split" (p. 35), and he then proceeds to give the account detailed by Réaumur of the longitudinal splitting of the shell in the neighbourhood of the joints of the claws in _Astacus_, so as to allow of the extrication of the hands. Nevertheless, however impossible it may appear for the chelæ to escape without this splitting, no such circumstance occurs in the exuviation of _Homarus vulgaris_; and when we consider that the hands of _Astacus_ are small in proportion to the wrist-joints, and that in _Homarus_ they are larger in proportion to those joints than in any other of the Macroura, this dissimilarity in the mode in which the claws escape is the more remarkable, and, I confess, to my own mind it suggests the suspicion that the distinguished and usually most accurate French naturalist to whom I have referred may possibly in this instance have been led to consider as a fact that which was to him a supposed necessity[9].

Since the foregoing account of the moulting of the Lobster was written, I have dredged a specimen of the common shore-crab (_Carcinus mænas_), in the act of casting its shell. This little crustacean had taken refuge, no doubt for the safe and secret performance of sloughing in a forest of _Zostera_, on one of the mud banks in Poole Harbour, and while scraping these weeds with a keer-drag it fortunately fell into my net. It shows how the Brachyura leave their old shells by the horizontal splitting away of the carapace from the other portions of the shell--the carapace itself remaining entire; and it also shows (and this was my principal object in exhibiting the specimen) the enormous amount of increase of size upon emerging from the shell, and the rapidity with which that increase takes place. The animal, as now seen, is in exactly the same state as when taken out of the water, and its bulk is probably some four times larger than the area of the shell in which it had been encased only a few minutes before. I retained the Crab in connexion with its old shell, and prevented its further escape by wrapping it in paper, so that it could not move its limbs. I thought such a specimen would be telling and illustrative, and that the old shell, being in contact with the new, would afford facilities for contrast. In this condition the Crab died, and, being out of water some time, it became dry, and the soft new shell collapsed and bulged in; but, upon placing the dead Crab in sea-water, the soft shell very speedily imbibed sufficient fluid to distend it to its previous dimensions. This of course was simply the effect of endosmosis. Mr. Couch, in describing the moulting of the common Edible Crab (_Cancer Pagurus_), speaks of its _drinking_ large quantities of water, and _thus_ becoming distended; but I rather think that the distension takes place by endosmosis, even during life. There are two circumstances which militate against Mr. Couch's opinion:--first, the rapidity with which the distension occurred in the Crab I have just exhibited, while still in the act of moulting; and secondly, that after death the same distension occurred when the Crab was immersed in sea-water; in which case it could only be by endosmosis. Indeed to me it seems very probable that this very endosmosis, when the water once comes in contact with the new, uncalcified shell, may, by distending it, be the main agent in the breaking open and dissevering of the elements of the old shell.

[9] The suspicion above expressed has been fully confirmed by
observations made by Mr. J. J. Bennett, the Secretary of the
Linnean Society. Mr. Bennett informs me that, in an aquarium in
his possession, an _Astacus fluviatilis_ has twice cast its shell,
and the process of moulting was on each occasion accomplished
without any splitting of the shell at the joints of the claws.

* * * * *

On the Shell-bearing Mollusca, particularly with regard to Structure and Form. By ROBERT GARNER, Esq., F.L.S.

[Abstract of a Paper read before the Society.]

The author commences the paper, of which the following is the substance, with some general observations on the morphology of animals. He thinks that the idea of an ascending and successive scale or chain of creation is, in the main, correct, when the great classes, and not species or genera, are made the links,--the disturbing or modifying influences being due to modes of life, food, habitat, &c., and causing a different (say the quinary) distribution. He is an advocate, too, for the doctrine of one fundamental plan of organization, and thinks that, in the zoophyte, there is a real union of both the animal and vegetable _nisus_.

The great divisions of this chain, the _radiate_, _articulate_, _molluscous_, and _vertebrate_, constitute an ascending series; the links of the chain, so to speak, being in each case, for such an extent, of a particular pattern; but, nevertheless, one of the highest mollusks may surpass in organization one of the lowest fishes, or an articulate creature a mollusk. The author considers such great divisions of animals, as well as minor ones--the gasteropodous mollusks, for instance--as realities, and not mere abstractions; and that they are independent of the circumstances of food, habitat, locomotion, &c., just referred to. So great, however, are these disturbing influences, that they often produce an extraordinary external resemblance or pseudo-analogy between animals of a very different nature, as between a _Chiton_ and an _Oniscus_, and they are connected intimately with, though not the cause of, what we call specific or generic distinctions. Aërial life, in contradistinction to aquatic, raises much the character of the locomotive organs; yet this is subordinate to type: hence the creeping Mollusk appears to have commonly a higher organization than the flying Insect.

The cartilages of _Sepia_ have a true resemblance to those of a Skate, and the Cirrhipede truly connects the Mollusk with the Crustacean. The author regards _Dentalium_ as a gasteropod, differing in this respect from Lacaze-Duthiers, whose beautiful paper, however, renders it supererogatory to say anything more on this animal, except that the author believes that the presence of the spiniferous tongue, of a proboscis, and the nature of the food, are favourable to his view: he also takes the feathery tufts to be the branchiæ.

The anatomy of _Aspergillum_ is similar to that of _Pholas_; its mantle, however, is all but closed in front, and ends in an obliquely-set muscular disk, applied to the internal surface of the rose of the so-called _arrosoir_, the openings of this part of the shell giving exit to certain processes and fimbriæ of the fleshy disk,--a narrow slit being also left in both the muscular and shelly disks for the exsertion of the small, compressed and curved foot. The animal is enveloped within the shell by a rather horny, general membrane.

The author touches upon the anatomy of some other genera of Lamellibranchiata. _Solemya_ has its firm, horny, dark cuticle doubled inwards from the valves over the tubular mantle; behind, it has an anal opening, and a second fringed branchial slit lower down: the branchiæ and tentacles are single on each side, the former being remarkably feather-like. The foot is similar to that of the Solens, but crenate round its anterior disk. _Cyrenoidea_ has the mantle closed below, but with two openings behind, the upper one with a semicircular internal fringe, incomplete above; a callous rim and fringe surround the mantle, which has also a third opening for the long, compressed, bent, and blunt foot. This last has a remarkable crystalline body, directed from the stomach to the pedal pore, apparently, as in _Cardium_, subserving by its elasticity to the extension of the foot, and consequently to locomotion; at any rate, it is not a sexual distinction. The external branchiæ are short, and the upper or internal branchial cavity does not communicate with the lower one. The renal organ opens near the branchial nerve, and the ovary at the base of the abdominal mass. _Trigonia_ is remarkable for its beautifully fringed, open mantle, its pectinated pits for the secretion of the teeth, and the large scythe-shaped foot, trenchant before and peaked behind, and having a fringed disk. _Vulsella_ is allied to the Oyster, but more so to the Pectens, having a small cylindrical grooved foot and appended visceral mass, but no byssus; the rectum perforates the heart, and has a tentacle above its opening. _Perna_ has a similar foot, and a very bulky byssus, with a large muscle attached to their base; the lips resemble those of the Oyster. The anatomy of _Crania_ is little different from that of _Orbicula_, as described by Owen,--the beautiful arms folded in several coils, with a simple mouth at their base, the stomach and short intestinal canal surrounded by the liver and hearts, and terminating by a lateral bend; the ovaries ramifying in the mantle; the adductor muscles being four in number, with some bands to the mantle; and on the latter, glandular markings corresponding with the microscopic sculpture of the shell. With respect to _Anomia_, the author has again been anticipated by Lacaze-Duthiers, though he has already given, in another paper, most of its anatomy and morphology: he would simply call attention to its very long and curious crystalline stilette, unconnected with the minute foot.

With respect to that _quæstio vexata_, the sexes of the Lamellibranchiata, he observes that any number of individuals of _Cyclas_ may be examined, and young fry will be found in the branchial laminæ in all; that all Oysters have ova, and also all individuals of _Pecten maximus_, the subpedal mass being visibly composed of an ovary and a testis. He is obliged to believe that one species of British _Anodon_ is universally oviferous. But the common Edible Cockle appears to have the individuals of different sexes, and the same may be said with regard to _Mytilus edulis_ and _Patella_.

The spermatozoa in the Cockle are oblong and a little curved, and torulated, as it were, whilst they are pear-shaped in _Mytilus_; they are also extremely minute, and their appendages must be very fine, for with a power magnifying 500 diameters they are scarcely to be seen.

In the shell of a _Patella_, _Emarginula_, or _Haliotis_, we have the two conjoined valves of a lamellibranchiate mollusk; and through such forms as _Calyptræa_, _Hipponyx_, _Navicella_, and _Nerita_, we arrive at the ordinary form of the gasteropod with its operculum.

Then follows a disquisition on the progressive tendency to a spiral geometry in these animals, due to a varying plan of conformation, and not to the force of the heart, there being generally an atrophy of the left side of the body. In _Nautilus_ and _Argonauta_, the shell and mantle are reversed in position to what they are in the Gasteropods, whilst _Sepia_ and _Hyalæa_ agree rather with the latter. The symmetrical shell of the lower Gasteropods undergoes a lateral torsion in the higher, spiral forms, to become again symmetrical in the Cephalopoda. The branchiæ in _Patella_ retain a position analogous to that of the same organs in the Lamellibranchiata; in some Chitons they have a tendency to retract towards the anus, as in _Doris_; in _Fissurella_ they waste at the sides and become developed above the neck, as in the spiral Gasteropods; but in them, the right branchia, and right side of the mantle are principally developed. From this torsion arises the form and spire of the shell. In _Aplysia_, where the branchial fissure is far back and to the right side, the right respiratory nerve preserves a superior position, and passes backwards to form its ganglion at the front of the branchial opening; the left, on the contrary, passes under the oesophagus to form a second ganglion, not mentioned by Cuvier, behind the first. In the more spiral Gasteropod the torsion is greater; the right nerve, for instance, mounts upwards over the digestive canal to form its ganglion quite in the left flank, whilst the left goes below the digestive canal to attain the right flank. In _Sepia_ the branchiæ are again symmetrical and abdominal.

The shell of the young _Sepia_ is composed of distant plates, only connected by minute transversely striated laminæ or flattened tubes, producing by their insertion a beautiful appearance of sinuous lines, very like those of a Baculite or Ammonite; and the spongy part of the shell, so constituted, is probably filled with air from the cavity of the body situated immediately in front, the intervening membrane having a peculiar structure. This cavity of the body exists in much lower mollusks; air being apparently secreted in it, to lighten the animal.

The author thinks that, in considering the anatomy and form of the body of the Gasteropoda, about ten species may be taken as types of corresponding families.

1. PATELLA _and its congeners_.--He claims to have been one of the first to show the termination of the oviducts and renal organs between the processes of the branchiæ in the Chitons. As they are commonly phytivorous, the intestine is often very long and disposed in large coils, in double apposition; the buccal apparatus is very remarkable. _Chitonellus_ differs but slightly from _Chiton_, the central elements of its tongue, however, being little developed, though having the same tessellated basement membrane. The tongue of _Emarginula_ differs much from that of _Patella_, having an immense number of serrated side-hooks and a dilated middle portion.

2. CALYPTRÆA, _&c._--The mollusks of this division have often supranuchal branchiæ, as have some of the last; the sexes also are frequently separate, rendering copulation necessary; and they are sometimes partially spiral, with a tendency to form an operculum. However, the little _Ancylus fluviatilis_ appears to be what is commonly called hermaphrodite, with a branchial lamina on the left side, together with the heart and openings of the genital organs; the stomach has a cæcum, and the penis a long filiform appendage; the female parts opening near the rectum and behind the male organs. It must respire by water rather than by air, for, in a rapid stream, the stones at the bottom are covered with _Ancyli_ (upon which also its round oothecæ, each containing four or five ova, are deposited), and it appears impossible for them to get to the surface to breathe. On the contrary, the lake-_Ancylus_, though the margin of its mantle is ciliated, may perhaps come to the surface, ascending the stalks of the Water Persicaria, on which it is mostly found, and on which its oothecæ are deposited. When the dark cuticle of this last minute creature is removed, its organs may be seen to be reversely disposed to those of the larger species, the heart being placed to the right, before the apex of the shell, and the rectum also on the same side.

3. DORIS, _&c._--The little _Doris aspera_ swims, back downwards, on the surface of a glass of sea-water, copulates, and deposits its semicircular oothecæ. The brain of the common Lemon Doris is of a fine orange colour, enveloped in a glandular matter, and is constituted by a complicated assemblage of ganglia: there are acoustic sacs and dark ocular spots upon it. There are six ganglia on the buccal mass, and about six or eight minute ones on the stomach. The anal sac appears to be a purple- or ink-bag; and the so-called matrix is composed of a peculiar substance, swelling enormously in water, of which it renders a large quantity viscid, and being also coagulable by alcohol and bichloride of mercury, but not by heat. Spermatozoa were found in the genital vesicle, as well as in the epididymis and its cæcum. The spines of the lingual plate are uniform, and in number about 10,000.

4. APLYSIA, _&c._--_Aplysia_ has been before alluded to. Cuvier, in his generally beautiful drawings, has scarcely done justice to (5) _Ianthina_, nor to its beautiful float and ootheca; it is peculiar for its fins, and the disk at the back of the foot. With respect to _Magilus_, it should be removed from the (6) Tubulibranchiata, its animal being a _Purpura_ in structure, with a bent horny operculum, and a very long linear appendage on the right side of the head, leading to the supposition that the animals are of different sexes, though there seem to be difficulties in the way of sexual congress. In the specimen examined, the spire of the shell was not solidified; the animal had a short proboscis, with rather bent subulate feelers, and eyes on the outside; it had also a rich purple secretion near the rectum on the right side.

7. TROCHUS, _&c._--Some of the species of _Trochus_ surpass even _Emarginula_ in the beauty of their lingual apparatus. The renal organ opens into the bottom of the branchial cavity, contrary to its disposition in _Helix_ and _Lymnæus_, where its exit is near the respiratory orifice. In _Planorbis_, that part of the respiratory cavity receiving the excretions seems separated by an imperfect valve from the right portion. With respect to the secretion of this organ, it consists, in both Gasteropoda and Lamellibranchiata, of numerous pellucid globular bodies, containing opaque earthy nuclei or granules, and presenting different appearances in _Anodon_ (for instance), _Cyclostoma_, _Buccinum_, and _Helix_. When these bodies are incinerated, lime is left, which in some cases appears to have been combined with oxalic acid. The little _Nerita litoralis_ presents the structure of the Turbonidæ very prettily and in small compass, particularly in the very long spiral tongue. _Delphinula_ has the fringed mantle and sides and very wonderfully armed tongue of the other Trochidæ. _Melania_ is of similar organization to our well-known _Paludina_, the stomach compound, the mantle and bilobed head fringed, and the latter marbled like that of _Paludina_. _Ampullaria_ appears to be truly amphibious.

8. BUCCINUM, _&c._--_Natica_ presents much the same structure as the common _Buccinum_, but has a muscular disk anterior to the mouth,--a disposition, with some variations however, found in other mollusks. The first and second stomachs are at a distance from each other, the tongue is little developed, and the branchiæ (often single in the Turbonidæ) two in number. _Purpura_ also differs but little from _Buccinum_. _Ovula_ is a less attainable mollusk: the foot is long and rather narrow, and subventral rather than subtrachelian, with a sinuosity on the right of the neck, where also is a short hooked penis in the male, receiving a vas deferens from near the rectum behind; there is a large and small branchia, and the reflected portion of the mantle is covered with tubercles and tentacles,--no doubt a fine garnish in the living animal; the mouth has a muzzle, and there are small eyes on the external sides of the curved, awl-shaped tentacles; the elements of the tongue are beautifully toothed and serrated.

9. LYMNÆUS, _&c._--Of the air-breathing aquatic and (10) terrestrial gasteropods the most interesting particulars are their generative organs, which the author proposes to re-examine. The brain of _Helix aspersa_ is composed internally of pyriform or oval ganglionic vesicles, each giving origin to one or more nervous fibres. The acoustic sacs are similar to those of _Doris_. The nerves from the upper part of the ring are enveloped in a darkish neurilema, and comprehend no doubt olfactory, optic, and tactile twigs; there being the buccal ganglia for taste, and the acoustic sacs for hearing; the twigs, however, forming the buccal or pharyngeal ganglia have a broad double root on each side, near the origin of the above three nerves. The lower part of the brain is very analogous to that of _Sepia_, giving off nerves to the foot, and external and internal respiratory ones to the mantle, respiratory opening, branchiæ, &c. _Lymnæus_ has the cephalic ring formed by about twelve ganglia, exclusive of two large and two minute ones on the buccal mass. The upper portion of the ring has ganglionic swellings, but in other respects the nerves are as in _Helix_. Its lower portion consists of two pedal nerves, and has the acoustic spot and a minute ganglion upon it; behind, this lower portion consists of five ganglia connected with both the anterior and upper swellings by a cord, but separated from the former by the aorta, as usual, and giving nerves to the flanks, pulmonary orifices and sac, heart, stomach, and viscera. The lower ganglia are bright yellow.

With respect to the Pteropoda, the branchiæ in _Hyalæa_ exist as a delicate membrane under the swollen part of the shell, in structure much like the same part in the Ascidians, the inlet being through the anterior opening of the mantle. There are eyes at the fold of the mantle behind, and two small tentacles above the mouth; the heart and rectum being on the left side, and the generative opening at the base of the right ala. _Cleodora_ is a very beautiful creature, with the same disposition and structure of viscera; brain-spots but no eyes were visible; the mantle had beautiful muscular bands; the branchiæ as above; the buccal apparatus is imperfect in both. _Cleodora_ has similar membranous expansions with _Hyalæa_, and also a sort of triangular lip.

_Argonauta_ has a lachrymal pore before and beneath the eye. The beautiful and obvious respiratory mechanism in the Cephalopoda needs not to be described. There is a large sac behind the viscera of the Argonaut, which opens on each side; it is perhaps of some hydrostatic use. There are at least three pairs of salivary glands, of which four open on the floor of the mouth, and two or three at the commencement of the gullet. Several small shells of Pteropoda and fragments of Cephalopods were found in the stomach, on which was observed the large nervous ganglion found in all these, as well as in lower mollusks. The branchial nerves have each two ganglia, of which the last at the root of the branchiæ is rounder than the other; the branchial hearts have processes as in _Sepia_. In _Sepia_ two openings lead from the respiratory sac into the cavity containing the venæ cavæ and their secreting appendages often imbued with glittering crystalline particles, and from the above cavities a wider opening on each side leads into a second sac further back, situated in front of the shell. There are auditory sacs in the Argonaut. The oviducts have separate openings, but originate together. Both _Sepia_ and _Argonauta_ are infested with a subcutaneous filiform entozoon, hooked anteriorly and rolled up spirally in the former. _Loligo media_ and _Sepiola_ have but one oviduct, and the two large, glandular, laminated organs, opening at their summits, are wanting in _Argonauta_ and _Octopus_. In _Sepiola_ one would almost think that copulation takes place, for the author has taken what he supposes to be the capsules of Needham, with dilated oval ends, tubular and bent pedicles or processes, enclosed elastic filaments, and adhering zoosperms, from the oviducts of the female: he has made the same observation also in _Sepia_. The latter has very similar male organs to _Octopus_, as described by Cuvier. In the embryo _Sepia_, the yelk enters below the mouth and opens into the upper stomach, but the beak of the animal also appears to be inserted into it behind. The vitellus in reality therefore enters by the foot, as it does in _Bulimus_, and probably in all Bivalves.

* * * * *

On the Linnean Manuscript of the 'Museum Ulricæ.'

By SYLVANUS HANLEY, Esq., F.L.S.

[Read Dec. 3, 1858.]

Not the least important result of the investigations of the Committee appointed by the Linnean Society to examine the condition of the collections and manuscripts of Linnæus, was the rediscovery of a written copy of the 'Museum Ulricæ.' The volume was manifestly, from internal evidence, a legible transcript of the original manuscript of that work, with alterations and interpolations in the peculiar handwriting of the author. It was, indubitably, the unpublished catalogue so often mentioned in the tenth edition of the 'Systema,' and contains descriptions of certain species alluded to as defined, yet, strangely enough, omitted in the printed edition. It is worthy of notice for many reasons: it corrects the frequent misprints; explains the many fallacious allusions to preceding species, their sequence being very different; it exhibits those early synonyms, which, culled from comparison with the actually described specimens, had been eventually supplanted by supposed better representations; above all, it imparts to us those original headings, or diagnoses (condensed from the subsequent details), which had been suppressed, of old, in favour of those already published in the 'Systema.'

This wholesale substitution, adopted by Linnæus, as a ready method of avoiding a tedious revision of all the headings, when he absorbed in the more comprehensive groups of his 'Systema' the members of manuscript genera he had determined to reject, involved a serious amount of confusion; for, oftentimes, the species of the two works, although designated by the same appellations, were totally distinct; and the combination of the diagnosis of the one with the details of the other displayed an array of features not known to be associated in any object in nature.

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Journal of the Proceedings of the Linnean Society - Vol. 4Chapter II: Front Matter (2)

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