m li THE EOTIFEEA. VOLUME I. PRINTED BY BPOTTISWOODE AND CO., NEW-STREET SQUARE LONDON H THE ROTIFEBA; OR WHEEL-ANIMALCULES, BOTH BEITISH AND FOBEIGN. BY C. T. HUDSON, LL.D. CANTAB., F.R.S. ASSISTED BY P. H. GOSSE, F.E.S. IN TWO VOLUMES, WITH SUPPLEMENT. VOLUME I. WITH ILLUSTRATIONS. LONDON : LONGMANS, GKEEN, AND CO. AND NEW YOEK : 15 EAST 16th STEEET. 1889. All rights reserved. Caecam mihi in cunctis fidem haberi hand postulo ; id tantum optans, ut continua indagatione ac studio mea aliquando confirmentur, aut me a vero aberrasse demonstretur. Perscrutatoris vel exactissimi, et quamvis Bummam adhibeat, attentionem fugere aliquando qusedam possunt ; et casus nonnunquam fortuito nobis offert, quse intensissima ssepe cura frustra qusesivimus. — J.' EASTER. C'est dans les livres de la Nature, qu'on doit lire, quand on veut travailler sur 1'Histoire Naturelle ; mais on ne peut pas y lire, quand on veut. II faut des lieux, des saisons et des circonstances favorables pour faire des observations ne"cessaires. Quelques fois a la verite* on peut aider a faire naitre des circonstances heureuses, mais plus souvent il faut qxie le hazard nous serve. — EKAUMUR. PEEFACE. NEARLY fifty years have passed, since Ehrenberg's successful invasion of the unseen world was made known by the publication of his noble work, Die In- fusionsthierchen ; and twenty-five since Dr. Arlidge's edition of Pritchard's excel- lent History of Infusoria brought together, in a convenient form, the results of the long and patient observation of many investigators. Meanwhile memoirs on various species of Eotifera, as well as the number of species themselves, kept slowly increasing ; and, what is of greater moment, the prolonged study of these interesting atoms, by several acute observers, has made clear some of the perplexing difficulties in their structure, and swept away many mistakes both of observation and inference. Under these circumstances the authors of this work believed that such a book as the present was much wanted ; and they hoped that their prolonged study of the Eotifera (continued, in the case of each author, almost daily, for upwards of thirty years), as well as their invariable habit of drawing from life all that they had observed, would enable them, by means of their long accumu- lated stores of drawings and notes, to meet this want. They venture to think, from the manner in which, so far, the book has been received, that this belief and hope have both been justified ; and in confirmation of the favourable opinions of it that have already been expressed, would point out (they trust with pardonable pride) that this work contains more than 120 species which were unrecognized when Dr. Arlidge wrote ; that nearly the whole of these have been added to science by the authors themselves ; that about eighty of these new species, chiefly among the Plo'ima, have been found by one of the authors during the last fifteen months ; and that the other by the discovery, among other remarkable forms, of Pedalion minim, has put beyond question the fact that the ROTJFERA, in one point at least, are closely linked to the ARTHROPODA. The thirty coloured plates have been divided nearly equally between the two colleagues ; the small uncoloured plates A, B, C are by Dr. Hudson. The figures in plate D are taken from various sources ; but in every case, both here and in the coloured plates, the original authority has been indicated ; the initials G or H being attached to those figures which have been drawn from the life by Mr. Gosse or Dr. Hudson. Mr. Gosse's independent portions of the text are always inclosed in square brackets [ ], and marked at the end with his initials, P.H.G. : the portions not so marked are by Dr. Hudson, including the first four chapters. vi PREFACE. Throughout the whole of the work Dr. Hudson has had the invaluable assistance of Mr. Gosse's MS. notes, and of his close and constant revision of the proofs. The hearty thanks of the authors are due to Mr. Frank Crisp, one of the secretaries of the Royal Microscopical Society, and editor of its Journal, for the great service that he has rendered them by forwarding early notices of all the pamphlets and papers published on the subject : anyone who is, or has been, engaged in a task similar to this will know what time and labour have thus been spared, by his thoughtful kindness. They are, too, greatly indebted to numerous kind correspondents for living specimens, often obtained with no little expenditure of time and trouble, and for the records of their observations on them ; as well as for valuable preparations of rare species, which could not be sent alive : to these correspondents the authors tender their grateful thanks. In each case where the specimen was new or rare the name of the place in which it was found, and (when permitted) the name also of the finder, has been added to the description. Thf,y wish, however, to make special mention of how much they owe to Miss Saunders of Cheltenham, Miss Davies of Woolston, Dr. F. Collins, and Mr. John Hood of Dundee, for their constant kindness in sending a profuse number of specimens of many species, some of which were of unusual interest. It is so natural to recommend one's own favourite pursuit that the recom- mendation often carries but little weight ; and yet there is much to be said in favour of the study of the Rotifera, that cannot be gainsaid. They are to be found almost everywhere ; they cost nothing ; they require neither expensive lenses nor an elaborate apparatus ; they tempt us to explore the country, and to take pleasant walks ; they are beautiful themselves ; and they suggest all kinds of difficult questions on life and being. Moreover there is happily still a great store of scientific ignorance concerning them, thus leaving an ample field for fresh discoveries. Nor is this all. The study of these animated specks (in which teeth, stomach, muscles, and even a brain lie hidden in the compass of an invisible mote) irre- sistibly leads the mind to the contemplation of Him, whose almighty hand is as visible in an atom of this animated dust, as it is in the myriad sparkles of the starlit heavens. C. T. II. CONTENTS OF T H E F I 11 IS T VOLUME CHAPTEE I. PAGE INTRODUCTION 3 CHAPTEE II. HISTORY OF THE LITERATURE CONCERNING THE ROTIFERA . .15 CHAPTEE III. ON THE CLASSIFICATION OF THE ROTIFERA CHAPTEE IV. ON THE HAUNTS AND HABITS OF THE ROTIFERA CHAPTEE V. RHIZOTA, (FLOKCTLAKIAU.*:) ........ 43 CHAPTEE VI. RHIZOTA, (MELICEKTAD/Ti) ........ 67 CHAPTEE VII. BDELLOIDA . . . . . . . . 9o CHAPTEE VIII. PLOi'MA, (IL-LOKICATA) , . . . . , . .117 THE EOTIFEEA CHAPTBE I. INTRODUCTION. Contemplatio Naturoj pra-gustus est voluptatis cn-lestis, constans aninii gaudium, perfectique ejus solatii inithuu, summusque fclicitatis luuuana; apex. Cum Anima hujus particcps fuerit, ex gi-avi quasi soporo excitata, in luce ambulat, sui ipsius obliviscens, iu citlesti, ut ita dicam, terra, inque terrestri coelo. — J. BASTEII. Inest in explicatione Naturae, insatiabilis qmtdam e cognoscendis rebus Voluptas, in qua uua, confectis rebus necessariis, vacui iiegutiis, honeste ac liberaliter possuiuus vivere. — CICKUO, DC Finibus, lib. iv. cap. 5. CHAPTEE I. INTRODUCTION. ON the Somersetshire side of the Avon, and not far from Clifton, is a little comhe, at the bottom of which lies an old fish-pond. Its slopes are covered with plantations of beech and fir, so as to shelter the pond on three sides, and yet leave it open to the soft south-western breezes, and to the afternoon sun. At the head of the combe wells up a clear spring, which sends a thread of water, trickling through a bed of osiers, into the upper end of the pond. A stout stone wall has been drawn across the combe from side to side, so as to dam up the stream ; and there is a gap in one corner, through which the overflow finds its way, in a miniature cascade, down into the lower plantation. The pond's smooth surface is prettily diapered with the green leaves of many a water-plant, and with the sharp images of three famous beeches growing close to its edge : but to a naturalist's eye the old wall is the more charming object. Time has crumbled away the mortar near the water's edge, and made a thousand nooks and crannies ; which, densely clothed with algre, are the haunts of myriads of living creatures. If we approach the pond by the gamekeeper's path from the cottage above, we shall pass through the plantation, and come unseen right on to the corner of the wall ; so that one quiet step will enable us to see at a glance its whole surface, without disturbing any living thing that may be there. Far off at the upper end a water hen is leading her little brood among the willows ; on the fallen trunk of an old beech, lying half-way across the pond, a vole is sitting erect, rubbing his right ear ; and the splash of a beech husk just at our feet tells of a squirrel, who is dining somewhere in the leafy crown above us. But see ! the water rat has spied us out, and is making straight for his hole in the bank, while the ripple above him is the only thing that tells of his silent flight. The water hen has long ago got under cover, and the squirrel drops no more husks. It is a true ' Silent Pool,' and without a sign of life. But if, retaining sense and sight, we could shrink into living atoms and plunge under the water, of what a world of wonders should we then form part ! We should find this fairy kingdom peopled with the strangest creatures : — creatures that swim with their hair, that have ruby eyes blazing deep in their necks, with telescopic limbs that now are withdrawn wholly within their bodies and now stretched out to many times their own length. Here are some riding at anchor, moored by delicate threads spun out from their toes ; and there are others flashing by in glass armour, bristling with sharp spikes or ornamented with bosses and flowing curves ; while, fastened to a green stem, is an animal convolvulus that by some invisible power draws a never-ceasing stream of victims into its gaping cup, and tears them to death with hooked jaws deep down within its body. Close by it, on the same stem, is something that looks like a filmy heart's-ease. A curious wheelwork runs round its four outspread petals ; and a chain of minute things, living and dead, is winding in and out of their curves into a gulf at the back of the B 2 4 THE EOTIFEBA. flower. What happens to them there we cannot see ; for round the stem is raised a tube of golden-brown balls, all regularly piled on each other. Some creature dashes by, and like a flash the flower vanishes within its tube. We sink still lower, and now see on the bottom slow-gliding lumps of jelly that thrust a shapeless arm out where they will, and, grasping their prey with these chance limbs, wrap themselves round their food to get a meal ; for they creep without feet, seize without hands, eat without mouths, and digest without stomachs. Time and space, however, would fail me to tell of all the marvels of the world beneath the waters. They would sound like the wild fancies of a child's fairy tale, and yet they are all literally true ; and, moreover, nearly all of them are true of that roti- ferous world which it is my purpose to describe. But it will be naturally asked by those of my readers to whom the subject is new, " What is a Eotiferon ? " and no doubt one would say that a book about Eotifera ought to begin at the beginning, and define precisely what a Eotiferon is. Precise definition is, however, in such a case, quite out of the question ; for, though it is easy enough to define the typical form of a natural group of animals, or even to include in the definition forms that must be placed not far off from the central one, yet in the ambitious attempt to frame a definition that shall include many families, we find (as we get farther away from the typical form) that one by one all the positive statements are disappearing from our definition ; and at last we have nothing left but the mere shell of a proposition, with everything worth the stating struck out of it. The Eotifera, then, are small aquatic animals varying from £ to ^|)Tr of an inch in length, and deriving their name from a wheel-like appearance produced by fine circlets of hairs seated on the front of their heads. A few species are marine ; but the great majority known to us belong to fresh water, and are to be found in ditches, ponds, reser- voirs, lakes, and slowly running streams, sometimes attached to the leaves and stems of water plants, sometimes creeping on the alga), sometimes swimming freely through the water. Although the greater number of the genera resemble each other in the chief features of their internal organization, so as to form a very natural group of animals, yet there are several aberrant forms which would render it a difficult matter to include them all in one precise definition. This indeed could be done only by introducing so many qualifications and exceptions to every statement, that the portrait would be rendered too vague for any reader but one already familiar with the whole subject. Of the greater number, however, it is enough to say : (1) That they swim by means of hairs on the front of their heads. (2) That they possess a simple stomach and intestine ; and peculiar jaws. (8) That they have muscles which are sometimes striated, and which often pass freely through the cavity of the body. (4) That they have a well-developed vascular system. (5) That their nervous system consists of one ganglion, with nerve threads radiating to their organs of sense. (0) That they are dioecious ; have ova of two kinds; and do not pass through any distinct metamorphosis. Though the above six statements are precise enough, and in the main true, yet it will be as well for those, who are not versed in the subject, to pass them over for the present, and first to master the structure of some one typical Eotiferon ; as, when this has been done, the general conception of a Eotiferon will be easily grasped, and the variations from the type readily followed and understood. For this purpose I have selected Brachionus rubcns, whose figure is given much magnified in PL A, fig. 1. The genus Brachionus is to be met with almost every- where. It is hardly possible in summer to take a dip of water from a garden-pond, or to gather the algfe from its walls, without bringing up some specimen of the genus. Brachionus rubcns is a fairly common species. It is comparatively a large handsome INTRODUCTION. 5 animal, very suitable for the purpose of description, and one which bears the tem- porary captivity of a compressorium remarkably well. The Female. Fig. 1, PI. A, represents the dorsal aspect of the female of this Brachionus, and fig. 2 the upper part of the ventral aspect. The drawings are from life ; but the out- lines of the various organs have been made unnaturally sharp and distinct, for the sake of clearness. The dorsal and ventral surfaces may be distinguished from each other in the great majority of the Eotifera by the following considerations : First, as to the dorsal surface : (1) It is arched (fig. 5). (2) The stomach (fig. 1, s] passes down it ; between it and the ovary (fig. 1, oy}. (8) The cloaca (fig. 1, d) is on it ; in the median line. (4) There is almost invariably one antenna (fig. 1, a) (or a coalesced pair) on it ; placed anteriorly on the median line. (5) The eye or eyes (fig. 1, e) are towards the dorsal surface. (6) In swimming over objects the Eotiferon keeps the dorsal side upwards. Secondly, as to the ventral surface (fig. 2) : (1) It is comparatively flat. (2) The entrance to the mouth lies on it (figs. 2, 5, bf). (3) The ovary is placed close to it (fig. 2, 5, oy}. In the case of those Eotifera whose dorsal and ventral surfaces have much the same contour, the above considerations present points of difference enough to decide between the two.1 B. rubens is inclosed in a case or lorica (figs. 3, 4, 5) which is both hard and trans- parent. The internal structure can be readily seen through it ; and, by suffering the animal to dry on a glass slip, and then dropping on it a solution of caustic potash, the softer portions of the body may be dissolved away, and the lorica left unharmed. It will then be seen to be closed above and below, with an opening at each end, like the shell of a tortoise. From the front opening the head is protruded, and from the hind the pseudopodium or foot (fig. I,/). The lorica has a glassy shining surface, and is armed with six short sharp spines in front, of which the central pair is the longest. Four of them are distinctly on the dorsal surface ; but the outmost pair belongs as it were to both surfaces, being on the edge where they meet. The front edge of the ventral surface (the mental edge as it is sometimes termed) 2 is hollowed out symmetrically into graceful curves (fig. 4). The lorica widens from the front backwards, till, at about two-thirds of its length, it reaches its maximum breadth ; and is then rounded off by two ogee curves that are separated, by a square notch on the dorsal surface, but by a nearly circular one on the ventral. In consequence, it is often said that the lorica has two blunt spines behind on the dorsal surface ; but this is somewhat misleading, as these so-called spines are merely the sides of the excavation. Strong ridges from each of the four central front spines run down about one-third of the dorsal surface (fig. 3), and still longer ridges mark the ventral surface with sweeping curves (fig. 4). The median portion of the lorica is by far the deepest, and in it the internal organs mainly lie. The dorsal surface of the lorica slopes upwards from the head to its line of greatest w'dth (fig. 5), and then abruptly falls to meet the under surface ; the whole lorica thinning off there into closely approaching plates, through the excavation in which the foot can pass. Each side, too, of the median portion of the lorica thins off in a similar manner ; so that the dorsal and ventral surfaces meet everywhere (except at the head) in a sharp edge. 1 Cf. Dr. Moxon, Trans. Linn. Soc. vol. xxiv. 1864, p. 455. 2 From mcntum, the chin. 6 THE EOTIFEEA. The head is shaped somewhat like a truncated cone, with the larger end forward ; posteriorly it is studded with several small rounded lobes ; while from its anterior surface rise three fleshy protuberances, crowned with stout vibrating hairs called styles. Each side of the front of the head, or corona,1 is rounded into a nearly circular lobe, and along the rim of each lobe runs an unbroken row of smaller vibrating hairs, called cilia, which are continued so as to meet each other on the dorsal surface. It is by means of this apparatus that Brachionus both swims and procures its food. As the head is seldom withdrawn into the lorica for more than a moment or two, and as the cilia begin to play the instant it is protruded, Brachionus would have been condemned to almost perpetual motion if it had not been for the foot. This organ is provided at its extremity with two pincer-like processes, or toes (fig. 1, t) ; which, however, do not pinch, but which can adhere even to glass by means of a viscid secretion that flows through their tips. The foot-glands (fig. 1, fg], which secrete this substance, are two club-shaped organs running down the whole length of the foot. They are to be met with in nearly all the Eotifera. The cilia, which are set closely round the edge of the corona, lash the water with such fury that it is impossible usually to follow the action of any individual cilium ; but, by selecting an animal whose corona is close to the covering glass of the live box, some spot can often be found where the action of the cilia is checked by their striking against the glass ; and, under these circumstances, it is easy to understand their action. Each cilium lashes sharply downwards (like a whip) on the corona, and then rises gradually into its place again, to repeat the action continuously, so long as the corona is expanded. As, however, the cilia do not do this simultaneously, but in turn, one after the other, in very rapid succession, those that can be seen together at any given moment are in every phase, from complete extension to complete depression; thus giving rise to various wave- like illusory appearances, according to the illumination, and also to the plane on which the objective is brought to focus. One of the most common of these appearances is that of a toothed wheel, which is so well imitated by the Philodinadcs, that early observers thought such wheels existed, and drew them like the escapement-wheel of a watch.2 If a little carmine be mixed with the water, two beautiful coloured spirals will be produced by the action of the ciliary wreaths, one on each side of the head leading down to the buccal funnel. The orifice of the buccal funnel, or, as it will be termed, the buccal orifice, lies in a niche on the ventral surface ; it is fringed by the ciliary wreath, which here dips down on either side of the corona, and passes round the V-shaped opening of the buccal funnel (fig. 2, bf)— that is, of the passage leading from the niche to the mastax. The atoms brought by the ciliary currents pass down the buccal funnel, which itself is lined with cilia ; and, if uninterrupted, enter the mastax (figs. 1, 2, mx], a muscular bulb containing the troplii (fig. 1, ti) or teeth. But it is not every atom whirled down the buccal funnel that is suffered to reach the mastax ; for there are two lip-like processes (fig. 2, Ip) rising from the mastax, which can be seen every now and then thrust up and down the buccal funnel ; and which by closing prevent the passage of morsels that are not to the Eotiferoii's taste. The sudden check, produced by the lips on the inflowing current, always sweeps out of the buccal funnel whatever the animal desires to reject ; and a constant stream of rejected particles may be seen issuing from the buccal funnel midway between the spirals caused by the corona. The Mastax. The mastax (figs. 1,2, mx ; fig. 6) is the muscular covering of the jaws or trophi. It has thick walls, and is slightly three-lobed, each lobe investing one of the three prin- cipal parts of which the trophi consist. There is an opening in front towards the ven- tral surface at the bottom of the buccal funnel, whose walls here merge into those of 1 A name suggested by Mr. Cubitt in lieu of trochal disk. 2 Baker on the Microscope, vol. i. Plate VIII. fig. 6, 1785. INTRODUCTION. 7 the mastax ; and there is a posterior opening towards the dorsal surface, just above the oesophagus. The contained hard parts, or trophi, consist of two hammer-like bodies, the mallei (fig. 6, ms ; fig. 9), and of a third anvil-piec:: called the incus (fig. 6, is). Each malleus has for its head or uncus (figs. 6, 7, 9, us) a piece which, when spread out by pressure, is like a comb with five unequal teeth (fig. 8, us), but which, under ordinary circumstances, is much curved, so as to bring the teeth close together. The handle of each of these hammers is a single stout piece called the manubrium (figs. 6, 7, 8, 9, mm). The anvil or incus roughly resembles a triangular prism, of which one end (the dorsal one) has been tapered to a point (fig. 10). It is divided into two equal pieces, the rami (figs. 6, 7, rs), by a cut from the dorsal to the ventral surface, so as to leave them just attached to each other at the bottom of the broader end. The two rami rest there on what looks like a stem, i\\e fulcrum (figs. 6, 7, fm), but which viewed laterally is seen to be a slender plate (fig. 10, fm). Muscles, springing from the walls of the mastax, are attached to various parts of the mallei and rami, and act so as to cause the unci to approach and recede from each other. But each uncus is fastened to the corresponding ramus ; and, in consequence, as the mallei recede the rami are opened, and when the mallei approach each other the rami are closed.1 From these arrangements it results that all the food which falls from the buccal funnel into the mastax, is first torn by the sharp points of the unci and then crushed between the opposing surfaces of the rami. On passing the rami, the food enters at once into the oesophagus. The Oesophagus. The oesophagus (fig. 2, 02) is a short tube, with thick walls, connecting the mastax with the stomach. The thickness of the walls may be readily seen by feeding Brachionus with carmine, and watching the stream of coloured particles trickle through the oeso- phagus into the stomach. Their course, in a narrow central line, shows clearly the small dimensions of the passage. There is always an appearance of water flowing through the oesophagus, even when the animal has withdrawn its head into the lorica, and when of course the action of the ciliary wreath has ceased. This appearance seems to be due to minute cilia lining the oesophagus, which by their joint rhythmical action give rise to illusory waves much larger than any real waves could be. That this explanation is not a mere guess will be shown farther on in the case of the Floscules. The Glands. Seated near the top of the mastax, and on either side of the buccal funnel, are two clear vesicular organs ; which are possibly salivary glands (fig. 2, sg), and are unusually large in Brachionus. They consist of two or three lobes on either side, and are best seen from the ventral side, as the buccal funnel enters the mastax, a little below the summit, on that side. There is a pair of glands called the gastric glands (figs. 1, 2, gg) placed at the anterior end of the stomach, and 011 either side of it. They exhibit cells with central nuclei imbedded in a granular substance. Each has a long, wide, nucleated duct connecting it with the upper end of the stomach. They probably perf&rm the func- tion of a rudimentary liver, and possibly cause, by their secretion, the peculiar yellow- brown colour which is so frequently met with in the stomach. The foot-glands have already been described in p. 6. The Stomach and Intestine. The stomach (figs. 1 , 2, s) is a simple sack, with thick cellular walls, which are frequently 1 For a full description of the complex action of the trophi see Mr. Gosse's paper " On the Struc- ture, Functions, and Homologies of the Manducatory Organs in the class llotifera," Phil. Trans. 1856, p. 419. 8 THE EOTIFEEA. studded with what appear to be bright yellow oil-globules. It is divided by an in- vagination at its lower third into an upper portion, the true stomach, and a lower which may be considered to be a short intestine (fig. 1, i). This lower portion frequently lies transversely to the longer axis of the stomach. Both are thickly ciliated on the inner surface, but the cilia of the intestine are larger, and more readily seen. When a portion of digested food has been transferred from the stomach to the in- testine, it is kept slowly revolving by the cilia, till it is suddenly expelled through the cloaca (fig. 1, cl). The intestine is connected with the cloaca by a short and very dilatable tube or rectum (fig. 1, r), and ends (as has been already said) on the dorsal surface, in the median line, just at the commencement of the foot. The rectum also is ciliated, so that the whole of the alimentary tract from the top of the buccal funnel down to the cloaca, with the exception perhaps of the passage through the mastax, is lined with cilia. The Vascular System. At the right of the intestine (viewed dorsally), and just under the line of the lorica's greatest width, lies the contractile vesicle (fig. 1, cv). This is a delicate bladder which alternately dilates and contracts, and with some regularity. The contraction is produced by fine muscular threads, which ramify in its walls, and cause it to empty its contents through a duct into the cloaca. Its distension is most probably due to the fluid poured into it by two looped and twisted tubes (fig. 1, Ic), which may be seen passing to it, one on each side of the body down from the head. This is, however, a much disputed question, which will be discussed fully in another place, along with the probable function of the whole apparatus. The tubes appear to be surrounded with a granular floccose material, which here and there dilates into irregular masses. Attached to the tubes on each side, at tolerably regular intervals, are five little tag-like bodies (fig. 1, vt), in which a flickering motion may be constantly seen, sometimes presenting the appearance of a waving cilium. There is much difference of opinion about the true structure of these tags — the vibratile tags, as they are termed — but it is probable that their office is to direct the perivisceral fluid into the tubes, and along them into the contractile vesicle, whence it is driven at intervals through the cloaca. The Muscles. The dorsal muscles are shown in fig. 3, and the ventral in fig. 4. From the posterior dorsal surface of the head, on each side of the cephalic ganglion, and close to it, a stout muscle (fig. 3, 1, i) slopes backward towards the dorsal surface, and is attached by a broad base to the lining membrane of the lorica. Outside this pair is a second (fig. 3, 2, 2), similarly attached, and running rather obliquely underneath the first pair, but not quite so stout. A similar pair (fig. 4, 4, 4) is attached to the posterior ventral surface of the head, and to the lining of the ventral surface of the lorica. The united action of these three pairs of muscles withdraws the head into the lorica. When it is so withdrawn, a pair of diverging muscular threads (not given in the figure) can be seen fixed to the lorica, -just below its central notch, with their other ends fastened to the head. These evidently oppose the action of the three other pairs (figs. 3, i, 1, 2, 2 ; 4, 4, 4) and help to draw out the head again. They are assisted in this by a further pair of muscles (also omitted from figs. 3 and 4), each of which is fastened at one end to the base of one of the outermost anterior spines, and at the other to a side lobe of the head. But the principal part in driving out both the head and the foot is borne by transverse muscles, which are attached to the lorica at the side, and are closely applied through- out their length to the soft organs of the body. Their sudden contraction compresses the perivisceral fluid, and so forces out the retracted head or foot. Nothing could be INTRODUCTION. 9 more effective than this hydrostatic pressure ; and under it the retracted foot darts out of the lorica with amazing swiftness. When the head is protruded, and the cilia are all in full play, Brachionus may often be seen to move its head, without withdrawing it, first to one side and then to the other, depressing the side lobes alternately. This action is effected by two pairs of short muscles (figs. 3, 4, 3, 3). one on each side, attached to the lorica at about one-third of its length from the front, and at their anterior ends inserted into the side lobes. Just as the muscles attached to the dorsal surface control the action of the head, so do the majority of those of the ventral surface give its various motions to the foot. They are six in all. First, two central muscles, closely parallel (fig. 4, 5, 5), and each divided into two branches at its upper end, run from nearly the middle of the lorica down to the bottom of the foot, where they appear to be attached each to one of the toes. Next, two slighter ones (fig. 4, 6, e), which pass up from the base of the toes, one on each side of the foot, and then diverge right and left from the central pair to points on the lorica about half-way between the median line and its edge. Besides these, there is, attached to the upper end of the foot and on each side of it, a muscle (fig. 4, 7, 7) which diverges still farther from the median line, and is fastened not far from the margin. All these six can act together, so as to draw the foot suddenly within the lorica ; while, by contracting some and relaxing others, the animal can whisk the foot about, or, if the toes be adherent to any substance, can shake its whole body vigorously from side to side — a favourite action. The Nervous System and Organs of Sense. The nervous system is represented by a bean-shaped cephalic ganglion (figs. 11, 1, gn) seated within the head on its dorsal side. Its substance is marked with what appear to be the hexagonal boundaries of cells. It is two-lobed posteriorly, and on the niche between the lobes lies the crimson eye (fig. 11, e). The pigment is distinctly curved round both sides of the niche so as to lie on each lobe, and to give rise to the notion that the eye may really represent a pair fused into one. I have not detected in this Rotiferoii the clear spherical lens which is so plainly visible in some of the others.1 Attached to the upper portion of the cephalic ganglion on the dorsal side is a conical and very flexible tube (fig. 1, a), whose broad base rests on the ganglion, and whose free end passes through the sinus in the lorica between the two longest spines. This is the dorsal antenna. A bulb armed with motionless setae completely closes the orifice of the tube ; and is so attached to its rim, that when this bulb is withdrawn by the contraction of a muscular thread fastened to its base, the tube is drawn down also by being infolded like the finger of a glove (fig. 15). There are two other setigerous bodies, close to the dorsal surface of the lorica, and with the setae protruding through the surface. They are near the margin on each side, a little above the line of greatest breadth (fig. I, a'}. They are rocket-shaped struc- tures, the cylindrical heads carrying on their blunt, rounded, outer ends, radiating setas ; and giving off, from their inner and pointed ends, cords which can be traced but a little way below the surface. But the list of the tactile organs of BracJiionus is not yet exhausted. From the two spaces on the dorsal side of the corona lying between the three large setigerous promi- nences (fig. 1, sp] rise two papillae, each bearing a long and very flexible style (fig. 1, ts). A similar style (fig. 2, t's') is placed on the ventral side, just within the rim of each ciliary circlet ; and another pair on the top of the central prominence. The whole six are very mobile ; and, from the way in which they seem to be used to explore in all direc- tions, there is little doubt that they are organs of touch. 1 As in Pcdalion, Conochilus, &c. 10 THE ROTIFERA. The Reproductive System. The reproductive system of the female is only too conspicuous ; as the presence of a large ovary (figs. 1, 2, oy), and of one or two opaque ova in different stages of growth, frequently obscures the sight of the other organs. The ovary is studded with large and rather irregularly shaped germs (figs. 1,2,0); and the ova (fig. 1, om), so long as they are within the body, are dark, granular, and homo- geneous. The ovary opens doubtless by an oviduct into the cloaca, but this I have not been able to see. This is the arrangement that exists in other cases wherever I have seen the oviduct. The mature ovum is expelled very quickly ; and the egg often remains attached to the animal by a thread till the young escapes by rupturing the shell (if it may be so termed) in which it is inclosed. If, however, a Brachiomis, with two or three eggs attached, be held in the compressorium, it will frequently free itself by pushing with its foot against the eggs, and so breaking the threads. The Egg. Of the eggs and their development it is unnecessary here to say more than that B. rubens has two kinds of female eggs, nearly of the same size ; of which the one has a smooth, transparent, membranous covering or shell, while the other has a thick opaque shell, ornamented with hemispherical knobs. The former is the ordinary " summer " egg, and the latter the so-called " lasting," " winter," or " ephippial " l egg. In shape, too, the ephippial egg differs from the ordinary female one. It is much bigger at one end than the other, and at the smaller end there is a projecting neck and cover (fig. 16). The male eggs are smooth and semi-transparent like the ordinary female egg ; but are somewhat rounder in shape, and barely half the size. They occur, too, in larger clusters ; for while it is iisual to see only two or three smooth female eggs, or one ephippial egg, attached to the mother, no fewer than eight or ten male eggs may be seen carried about together. The Male. The young female Brachionus when hatched resembles its mother ; but the young male is a widely different animal (figs. 13, 14). It is about one-third of the length of the adult female's lorica, and it has neither lorica, mastax, jaws, nor stomach. The head bears a simple circle of long cilia, and there is a red eye on a cephalic gan- glion placed just as it is in the female. The vascular system, with its tortuous canals, vibratile tags, and contractile vesicle, is also present ; and the foot is furnished with the usual pair of glands ; but of a nutritive system there is not the slightest vestige. Nearly half the body is occupied by a great sperm-sack (figs. 13, 14, ss), in which under favourable circumstances the spermatozoa themselves may be seen in motion. The sperm-sack ends in a short protrusile tube, ihepenis (figs. 13, 14, _p), ciliated at the end, and placed just above the foot. In the larger and more transparent males of other genera — notably in that of Asplanchna Ebbesbornii— there are special muscles for drawing back the protruded penis ; and, though I have failed to see these in the male of B. rubens, it probably possesses a similar structure. This strangely unfurnished creature leads a brief life of restless energy, now darting from place to place, so swiftly that the eye can scarcely follow it, and now whirling round as if anchored by its curved foot and penis. It often circles round the female, attaching itself now here, now there, and forcing its companion to waltz round and round with it, from the top of the phial to the bottom. With animals so active and so small it is difficult to be certain of having seen actual 1 So named by Prof. Huxley from the resemblance to the lasting-eggs of the Daphniro. These latter were called " ephippial" by Mailer from their shape, which is that of an ephippium, or saddle. INTRODUCTION. 11 coitus, but Mr. Gosso had tins good fortune in the case of a closely allied species, B. pala. He says : " I collected about a dozen females, half-grown and adult, and placed with them two lively males that had been hatched during the night. I directed my attention principally to one of these, as I could not watch them both. It soon came near one of the females, when it seemed to become animated by a sort of frenzy ; describing with excessive rapidity a circle, of which its head formed the circumference and its foot the centre. The extremities were incurved in the direction of its circular movement. After a while it left off, and began to play about the body of a female, moving over and round the lorica, while she whisked about the foot, as if to lay hold of him ; at length she drew in her foot, and that of the male appeared to adhere to it ; and I distinctly saw the thick penis presented to the cloaca, and for a moment inserted about half its own length ; then it was instantly drawn out, and the male began his frenzied gyrations again." l It is obvious, even from the brief account here given, that several highly interesting questions arise concerning the reproductive system of the Eotifera. For instance, in what respects, besides outward form and covering, do the ephippial eggs differ from the smooth female eggs ? and what leads to their production ? What part, if any, does the male play in these differences ? and how is it that one female produces none but male, while another has none but female eggs ? Again, why are the males of so strange a structure ? and why do they appear only for a short time during the year ? and is their appearance due to external causes, or are they the inevitable completion of a cycle of reproductive changes ? Unfortunately it is much easier to ask these questions than to answer them. The observations that have been made on these points are but scanty ; and, to some extent, contradictory ; and the difficulties in the way of persistent investigation are by no means slight. 1 " On the Dioecious Character of the Eotifera." Phil. Trans. 1857, f 19. CHAPTER II. THE HISTORY OF THE LITERATURE CONCERNING THE ROTIFERA. MiU turn egerunt, qui ante nos fueriint, sed non peregcmnt. Multum adhuc restat operis, multumque restabit ; nee ulli, nato post mille secula, prsecludetur occasio aliquid adhuc adjiciendi. — SENECA, Epist. Ixiv. Cffiterum millius in verba jurans, aliortun inventa consarcinare haud institui ; quic ipse qusesivi, reperi, debitaque attentione et patientia repetitis vicibus, diver- soque tempore annoruiu serio observavi, propono. — O. F. MULLER, Venn. Terrcst. et Fluv. pracf. 15 CHAPTEK II. THE HISTORY OF THE LITERATURE CONCERNING THE ROTIFER A. IT is nearly 200 years since Mr. JOHN HARRIS, F.E.S., Eector of Winclielsea, pub- lished in the " Philosophical Transactions" } the following " Microscopical Observations" on a drop of some rain water which " had stood in a gallipot in his window" for about two months : "I saw here an animal like a large maggot, which could contract itself up into a spherical figure, and then stretch itself out again ; the end of its tail appeared with a forceps like that of an earwig ; I could plainly see it open and shut its mouth, from whence air-bubbles would be frequently discharged. Of them I could number about four or five, and they seemed to be busy with their mouths as if in feeding." This description is but vague ; and yet it is very probable that the animal which the rector found in his gallipot was a Rotifer : and if so, this is the earliest notice that we have of the class. A few years later, in 1703, LEUWENHOEK published, in the same "Transactions,"2 an account of some animalcules, living in sheaths, that he had found at Delft, on green weed brought by the flooding of the Maes from Delft Haven. These little creatures were almost certainly Limnias ceratophylli. Leuwenhoek gives figures of one, and notices its bearing " two wheels thick set with teeth as the wheel of a watch." In a later paper 3 in the " Philosophical Transactions" he gives a much better account of Limnias, with greatly improved figures. He states that when the two wheels are viewed frontally they are seen to be continuous and to form but one ; and he adds a figure (PI. B, 4) of the corona so seen, and notices that the ciliary waves run right round the whole wreath in the same direction. Leuwenhoek next describes an animal " that has a receptacle or little house com- posed of round bubbles," and furnished with " surprising wheelwork " of four parts, three of which only were shown in his figure, " the fourth being almost hid from sight." One glance at the figure of the animal (PL B, 1), and at its corona (PL B, 2), shows us that he has had Melicerta ringcns under his microscope. His next paper 4 on the same Eotiferon is pleasant reading ; for it vividly recalls the shock of delighted surprise with which every naturalist first enters into the enchanted world beneath the waters. " I took notice," he says, " of the surprising figure of an animalculum, fixed in a little scabbard or sheath, fastened to some of the small green weeds found in ditches of water. And, as often as I viewed these animalcula and showed them to others,' we could not satisfy ourselves with looking on such surprising objects ; and the more because we could not conceive how so strange a motion, as they all had, could be performed ; as also what should be the use of such a motion." Leuwenhoek also relates how Meli- certa makes its tube pellet by pellet ; and this is his account of it. "I viewed one of these animalcula a good while together ; and observed, several times, one after another, that when the animalculum thrusts its body out of the sheath or case, and that the 1 Phil. Trans, vol. xix. 1696. 2 Phil. Trans. No. 283, 1703. 3 Phil. Trans. No. 295, 1705. 4 Phil Trans. No. 337, vol. xxviii. 1713. 1C THE KOTIFEEA. wheel-like or indented particles moved in a circle, at the same time out of a clear and transparent place a little round particle appeared, which, without nicely viewing, could hardly be perceived ; which particle growing larger, moved with great swiftness as it were about its own axis, and continued without any alteration in its place, till the animalculum had drawn part of its body back into its sheath ; in doing which it placed the said round particle on the edge of its sheath, which thus became augmented with a round globule ; and whereas the animalculum had placed the said globule on the east part of its sheath, another time it fixed it on the south or north side ; by which means the sheath was regularly increased on all sides." Of course this is but a rough sketch of the machinery and actions of the famous tube- maker ; but it is a vigorous one, and true to the life as far as it goes. With equal truth and vigour does Leuwenhoek describe, in the same paper, the transformations of Philodina roseola ; — that creature, whose powers of lying dormant for rotiferous ages, and of then coming to life again, have made it as famous as the l< Seven Sleepers." To anyone with a sense of humour it must be delightful to read the following para- graphs of Leuwenhoek's paper ; and then to reflect how his discoveries have been re- peatedly re-discovered; and how again and again they have been challenged, con- firmed, forgotten, and once more discovered. In fact, the Philodine has been the cause of a dispute which has all the marvellous properties of the Eotiferon itself. For it periodically goes to sleep and revives again, just as P. roseola does ; but with a difference. The Eotiferon, when it awakes after its long sleep, takes up its life at the point where it left it off ; and ultimately " gives up its murmuring breath " after an existence of three or four years ; but the dispute concerning it invariably begins again both de novo and de ovo ; and having already lasted, with periodical intermissions, for nearly 200 years, evidently bids fair to last for 200 years more ; as it has only a short time ago awakened once more, as fresh and as vigorous as ever, and found its way into our daily papers. " I discovered," says Leuwenhoek, " several animalcula that protruded two wheels out of the fore-part of their body as they swim, or go on the sides of the glass. . . . This sort I found, in great numbers, in the gutter water which had stagnated some days in the small pits or cavities of the lead. ... In October 1702, I caused the dirt of the gutters, when quite dry, to be gathered together, and taking a small quantity of it, I put it into a paper on my desk ; since which time I have often taken a little of it, and poured on it boiled water after it had stood still till it was cold, that I might obviate any objection that should be made, as if there were living creatures in that water. These animalcula, when the water runs off or dries away, contract their bodies into a globular or oval figure. . . . In the month of September I put a great many of the last-mentioned animalcula into a wide glass tube, which presently placed themselves on the sides of the glass ; whereupon, pouring off the water, I then observed that several of the animalcula, to the number of eighteen or nineteen, lay by each other in a space of coarse sand; all of which, when there remained 110 more water, closed themselves up in a globular figure. Some of these animalcula were so strongly dried up that one could see the wrinkles in them, and they were of a reddish colour ; a few others were so transparent, as if they had been little glass balls, that, if you held them up between your eye and the light, you might move your fingers behind them, and see the motion through their bodies. After these animalcula had lain thus dried up a day or two, in an oval or globular form, I poured some water into the glass tube ; whereupon they presently sank to the bottom, and after the space of about half an hour began to open and extend their bodies, and, getting clear of the glass, to swim about the water. ... In the month of October, before the dirt of the leaden gutter was quite dried up, I took a handful of it, and laid it on a glazed earthen dish in order to preserve it. ... Upwards of twenty-one months after, I took some of this dry stuff and infused it, both in cold water that had been boiled, and in rain water newly fallen ; whereupon the animals began to show themselves in great numbers." The only points of this much vexed question that Leuwenhoek passes over are : HISTORY OF LITERATURE CONCERNING THE ROTIFERA. 17 (i) how the Rotiferon, when drawn up into a ball, resists the persistent baking of a summer's sun on the housetop, or the long drought of twenty-one months on the naturalist's desk, without parting with its own internal moisture ; and (ii) why only eighteen or nineteen of his Rotifera (those near the coarse sand) succeeded in rolling themselves up and sur- viving, while the rest perished.1 The investigations thus worthily begun by Leuwenhoek were carried on with much spirit by many other observers ; and, during the hundred and thirty-five years that elapsed before 'the publication of Ehrenberg's famous work, " Die Infusionsthierchen," no fewer than sixty of Ehrenberg's species, contained in thirty of his genera, were entered on the list of known Rotifera. I have arranged the more striking forms of these in the following table, which classifies them under the heads of some of the families into which I have divided the Rotifera ; 2 and I have added the names and dates of their discoverers ; it will be thus seen how wide a ground had been covered by the early naturalists, since more than half the families have representatives in the table. f Stephanoceros Eichhornn . . Eichhorn, 1701 FLOSCULAKIAD^; ..-,_,.,. „. ,, -,r,nn 1 Floscularia ornata . . . Eichhorn, 1767 A List of some of the Rotifera discovered before 1838. Stephanoceros Eichhornii . . Eichhorn, ] Floscularia ornata . . . Eichhorn, 3 I Melicerta ringens .... Leuwenhoek, 1703 Limnias ceratophylli . . . Leuwenhoek, 1703 MELICEBTAD.E . . •< T . , . . , . ^ , .,„-/• Lacinularia sociahs . . . Brady, 1755 I Megalotrocha alboflavicans . . Eosel, 1755 , Eotifer macrurus . . . Baker, 1753 PHILODINAD.E • . 1™ -i n- T u i ir,n0 ( Philodma roseola .... Leuwenhoek, 1702 HYDATINAD;E . . Hydatina senta .... Miiller, 1773 SYNCH^TAD^: . . Synchseta Baltica .... Baster, 1759 NOTOMMATAD;E . . Notommata tigris .... Miiller, 1786 TKIABTHBAD.K . . Triarthra longiseta . . . Eichhorn, 1775 / Brachionus pala .... Joblot, 1718 BKACHIONID^E . . J Brachionus urceolaris . . . Hill, 1751 lAnurasa striata .... Miiller, 1776 PTERODINAD;E . . Pterodina patina .... Eichhorn, 1775 EtTCHLANiD^E . . Euchlanis dilatata . . . Eichhorn, 1775 EATTULID/'E . . . Mastigocerca carinata . . . Miiller, 1786 , Dinocharis pocillum . . . Miiller, 1776 DINOCHABID;E . . \ , , „ . ........ -__,, I Stephanops lamellaris . . . Muiler, 178b It will be seen that the names of Eichhorn and Miiller occur much more frequently than that of any other observer in this list, and their works on the Rotifera deserve, I think, a special notice. JOHANN CONRAD EICHHORN was the pastor of St. Catharine's church at Danzig, and his book on the " Natural History of the Smallest Aquatic Animals " 3 was published in 1781. Though small, it is a most interesting work. He gives figures and descriptions of about a dozen Rotifera that can be identified, including Stephanoceros, Floscularia, Melicerta, Triarthra, Dinocharis, Actinurus, Euchlanis, and Pterodina ; and of most of his species he was the discoverer. His enthusiasm is delightful. " I have devoted myself," he says, " to this invisible world, which yields itself to our ken only under the magnifying glass ; and I have, for eleven years, spent my leisure moments on it, so far as my professional duties would permit, in order to know God in His smallest and invisible works ; and I have found Him very great therein. Not the great works only, those vast heavenly bodies — 1 The question will be found fully discussed farther on in the general account of the family Philodinadro. 2 See chap. III. On the Classification of the Eotifera. 3 Beitrdge zur Naturgeschichte der kleinsten Wasserthiere. C 18 THE EOTIFEEA. not those huge animals, on earth as well as in sea, who can scarcely drag the weight of their frames— not these alone declare the glory of the Almighty. No ! the smallest also show, just as distinctly, the perfections of their Creator. Yea ! one may say, these even more than those ! A great church clock is certainly a wonderful machine, but a pocket watch — a watch in a ring — is yet more so, and conduces to the greater fame and glory of its maker." His description of his chief discovery, that of his Crown Polyp (Stephanoceros Eichhornii) is very amusing. " I found," he says, " this extraordinary and marvellously formed animal first in 1761, on July 20, on a water plant, which had been standing some weeks in water. I saw that there was something on the plant which was quite unknown to me. I moved the glass, in order to see if it was something alive, and if it would draw itself together, which happened, to my delight ; therefore I examined it through a lens, but it appeared to me, through this, just like an orange flower which was not yet closed, but which now drew itself together, and now outspread itself. All this stirred up in me a great desire to see this new animal under the glass, but that required skill to get it out, as the glass vessel in which it was, was nearly an ell high, and this animal was right at the bottom. I tried first with the quill of a feather to bring it to the top, but it was continually lost to the eye by shutting itself up. At last I succeeded with a little wire hook in drawing out the plant on which it obviously was, and as soon as I could reach it with the scissors I snipped off a tiny stem, and that brought me out the whole animal unharmed. I placed it at once under the magnifying glass,' and saw this matchless creature as it is shown in the engraving." l What a pleasant picture this is of the grave pastor fishing away with a quill pen to fetch up Steplianoccros from the bottom of a glass beaker a yard and a quarter high ! About the same time as Eichhorn, flourished the great Danish naturalist, OTHO FREDERIC MULLER. He was an excellent botanist and zoologist, and published works on many subjects. He wrote on the Flora and Fauna of Denmark, on Fungi, on the Hydraclmae, and on Fresh-water and Marine Worms ; but his chief delight was in the Infusoria, and his posthumous work, " Animalcula Infusoria Fluviatilia. et Marina, &c." 178G, was the first that brought this new kingdom to the knowledge of the naturalist. The " Animalcula Infusoria " contains the descriptions and figures of about fifty Eotifera, among which are Lacinularia, Hydatlna, Scaridium, Triarthra, Bracliionus, Anurcea, Pterodlna, Euchlanis, Dinocharis, Stephanops, and Mastigocerca. More than half of Miiller's species were new when published ; and his figures, taken from life, are beautifully drawn on copper by himself. Of course there is a great lack of detail in the drawings of the internal structure of the animals, but they are an immense advance on those of Eichhorn, the outlines being usually both spirited and faithful. Miiller's text, too, is as good as his figures. It is the work not only of a naturalist, but of a thoughtful and learned man ; and both the " Animalcula Infusoria " and his pre- vious work, " Vermium Terrestrium et Fluviatilium," abound with admirable and striking passages. In the latter, he thus begins his dissertation on the Infusoria : " The world of the invisible, a world shut to our ancestors, was first entered about a hundred years ago. It breeds monsters of unheard-of form and manner of life, it abounds in miracles as much as do the remote Indies ; but is explored with lesser peril, for it lies everywhere at our very feet, and is not sought out for gold. " Each was explored with great slaughter of its inhabitants ; the one often resisted by wasting the lives of its aggressors, the other had no defence but patience. " This we owe to the needle, which joined two hemispheres together ; that to the lens, which images alike the solar spots and the infusoria, the widest apart of all things. " In this interval what indeed is great, what little ? Man: for he thinks and suffers." L. JOBLOT styles himself, " Professeur Eoyal en Mathematiques, de 1' Academic Eoyale de Peinture et Sculpture, demeurant sur le Quayde 1'Horloge du Palais, au grosEaisin." 1 See PI. B, fig. 14. HISTORY OF LITERATURE CONCERNING THE ROTIFERA. 19 His book, published in 1718, consists of two parts. In the first he describes various forms of microscope, and the best way of using them ; in the second he details, from his diary, the results of a series of experiments made with infusions of various plants and substances. The list of his infusions is long and curious. He tried pepper, roses, rhubarb and senna, maize, violets, mushrooms, hay, raspberry stalks, celery, knapweed, fennel, straw, marigolds, melons, tea, oak bark, &c. &c., and even found an animalcule that, — for saving charges, A peeled sliced onion eats, and tipples verjuice. He scoffs at the notion that living animals could be produced by the putrefaction of any- thing, and is confident that the infusion of each substance produces its own peculiar animals. He supposes that eggs are laid on these substances "by a countless number of very little animals that fly or swim in the air close to the ground," and who " let fall their eggs and little ones as they course backwards and forwards in the air," particularly in the spots where they were stopped by the vapours escaping from their favourite plants. He was the first to discover the genus Brachionus (B. pala and B. amphiceros), and he found in his infusions various species of the genus Rotifer (probably B. vulgaris and B. citrinus] as well as, possibly, a Lepadella and a Monostyla. His figures are grotesque enough ; and he damaged his reputation as a sober naturalist by sketching a six-legged creature with " tout le dessus de son corps couvert d'un beau masque bien forme, de figure humaine, parfaitement bien fait." A fierce moustachioed face it is, and, as Joblot adds, " couronne d'une coeffure singuliere." His names for the animals are as odd as his figures : he has " top-knots," " bagpipes," "dandies," " tor- toises," " kidneys," and even " crowned and bearded pomegranates " — the last strange title being given to his new discoveries, the Brachioni. In spite of all these absurdities his written descriptions are often vivid and accurate, and he is a shrewd observer. For instance, he notices how cleverly the Rotifera swim without jostling each other ; and he concludes that " though we cannot see them, they must have eyes, and those very good ones." He describes the restless movement of Brachionus swaying from side to side as it thrusts about its long foot, and observes that " there are some females who carry only one egg, some that carry two, and some as many as six, which, however, is not common ; and when there are so many eggs they are smaller in size than they are when they are fewer." It is thus clear, both from his description and his figure, that this first discoverer of Brachionus pala had already lighted on a female carrying a cluster of male eggs. Again, he discerns the difference, in size and colour, between Eotifer vulgaris and B. citrinus • and describes their leech-like movements, their telescopic joints, and their constantly moving jaws. I need hardly add that he considers the latter to be the heart. His comments on his discoveries are as characteristic as his figures and theories. We have seen the Danish naturalist exulting in the human intellect which, armed with one simple weapon, attacks alike the distant planets and the invisible infusoria ; and find- ing even greater reason for his pride in the very weakness and ills that flesh is heir to. The Prussian pastor, too, is as enthusiastic after eleven years' study as he was when he first began ; and, as if he would add another verse to the Benedicite, cries, to all the creatures of the invisible world, " Bless ye the Lord ; praise Him and magnify Him for ever." And Professor Joblot is also enthusiastic, but his strain is pitched in a different key. He says of one of his infusions that " it gives rise to a most delightful spectacle, so curious to see and watch that I do not think that the diversion of the play, of the opera with all its magnificence, of rope-dancing, tumbling, or of the animal-combats which we see in this superb city, ought to be preferred before them." It would be unfair to M. Joblot not to add that he is capable of better things, as the following extract shows : " There is nothing despicable in nature ; and all the works of God are worthy of our C 2 20 THE EOTIFEKA. respect and admiration, especially if we take heed to the simplicity of the means by which God has made and preserved them. The smallest gnats are as perfect as the hugest animals, the proportions of their limbs are equally just, and it seems as if God had even wished to give them more ornaments than He has to the greater creatures, in order to make up to them for the srnallness of their bodies. They have crowns, tufts, and other adornments on their heads, which surpass all that female luxury has invented ; and we may say that those who have looked only with unaided eyes have seen nothing so beautiful, so fitting, nor even so magnificent, in the palaces of the greatest princes, as that which the microscope shows on the head and body of a simple fly." About forty years after Joblot, HENRY BAKER, F.K.S., published a somewhat similar work. The first volume, " The Microscope made Easy," treats of the instrument itself ; while the second volume, " Employment for the Microscope," describes the various things that may be seen with it. In the second volume he gives an elaborate account, with figures, of what I believe to be Philodina roseola ; as well as descriptions and drawings of Rotifer macrums, Brachionus pala, B. urceolaris, B. Bakeri, and probably also of Euchlanis triquetra ; and of these six species the second and last had not been described before. His drawings are vastly superior to those of Joblot, especially his figures of the Brachioni. He notices and introduces into his figure the long vibrating styles which crown the head of B. pala, as well as its winter eggs. He failed, indeed, to understand the lorica of Euchlanis ; but that is no wonder, for he has had many to bear him company. It is unnecessary for me to say more of a book that is still within everyone's reach ; but there is one admirable passage in his preface that I must give myself the pleasure of quoting. " That man is certainly the happiest who is able to find out the greatest number of reasonable and useful amusements, easily attainable and within his power ; and, if so, he that is delighted with the works of nature, and makes them his study, must un- doubtedly be happy ; since every animal, flower, fruit, or insect, nay, almost every particle of matter, affords him an entertainment. Such a man never can feel his time hang heavy on his hands, or be weary of himself, for want of knowing how to employ his thoughts ; each garden or field is to him a cabinet of curiosities, every one of which he longs to examine fully ; and he considers the whole universe as a magazine of wonders, which infinite ages are scarce sufficient to contemplate and admire enough." In Plate B, I have given copies of some of the old figures drawn by these authors, and if the reader will compare them with EH REN BERG' s drawings of the same animals, he will see at a glance why the Prussian naturalist's work ' swallowed up as it were the very memory of all his predecessors. Instead of feeble, inaccurate drawings, in which the internal structure was represented by mere blots and patches, Ehrenberg gave excellently drawn figures full of accurate details ; and at the same time described the animals them- selves with wonderful exactness, considering the very great number that he studied unaided. Nor was this all : he had such a grasp of the whole subject, such a minute personal knowledge of the living animals themselves, that he invented a system of classification which has held its own for nearly fifty years. In addition to its other merits, Ehrenberg' s splendid work added more than a hundred new species to those already known, containing among them such remarkable forms as Conochilus volvox, Notommata clavulata, N. copeus, N. centrum, Diglena grandis, Polyarthraplatyptera, Noteus quadricornis, Microcodon clavus , (Ecistescrystallinus, &c. Three years after the publication of " Die Iiifusionsthierchen," DUJARDIN published his " Iiifusoires " as one of the volumes of the Histoire Naturelle des Zoophytes in the " Suites a Buffon." The last part of this volume, being one-sixth of the whole, is devoted to the " Systolides " or Eotifera. His book is mainly critical, and, so far as I can find, con- tains little on the Eotifera that was new, except his observations on Albertia and Lindia. 1 Die Lifzisionsthicrclien ; Leipzig, 1838. HISTOKY OF LITERATURE CONCEENING THE ROTIFEEA. 21 His criticisms are shrewd and often just ; he points out that Ehrenberg's respiratory tube is probably an antenna ; suggests that the convoluted tubes, flickering tags, and contractile vesicle of the vascular system have a respiratory function ; calls atten- tion to the varying forms of the mastax and trophi as good characters for classification ; and conjectures that the perivisceral fluid is the true analogue of the blood. On the other hand, he could not see Floscularia' 's tube ; could not make out the striated muscles in any Eotiferon, even in Pterodina patina, of which he gives a figure ; could see indeed no difference between the muscles and the nerves ; doubted the exist- ence, as specialised structure, of either the one or the other ; and from want of personal acquaintance with them, affirmed the identity of many of Ehrenberg's species, which are undoubtedly distinct. But although he has small claim to be considered either an original or an accurate observer of the Eotifera, he made one happy hit in his attempted classification, which will be detailed elsewhere. Since Dujardin's time the more noteworthy essays that have been published on various portions of our present subject are by Mr. P. H. Gosse, F.E.S. ; Dr. F. Leydig ; Professor T. H. Huxley; Herr C. Vogt ; Dr. F. Cohn; Dr. W. Moxon, F.L.S. ; Dr. W. Salensky ; Dr. S. Bartsch ; and Herr Karl Eckstein. Mr. GOSSE, in his paper, " On the Anatomy of Notommata aurita," l described with minuteness the organization of this common species, so that the essay became, as it were, a key to the structure of the majority of free-swimming Eotifers. His next treatise, " On the Structure, Functions, and Homologies of the Manducatory Organs in the Class Eotifera," 2 is illustrated with a great many drawings of the mastax and trophi of various species ; and discusses the changes that they undergo, in passing from the typical to the most aberrant forms. It is in this treatise that Mr. Gosse contends that the dental organs of the Eotifera are true mandibulae and maxillae ; and that the mastax is a mouth ; and assigns to the Class a position among the Articulata. In a subsequent paper, " On the Dicecious Character of the Eotifera," 3 Mr. Gosse extended this character from a single genus, Asplanchna, to five others ; and trebled the number of the known dioecious species. Some years later, Mr. Gosse began, in " Contributions to the History of the Eotifera," 4 a general account of the whole class, arranged according to a classification of his own, and continued it so far as the Flosculariadce, Melicertadce, and Notommatina, illus- trating each family with descriptions and figures of certain selected species. This work, however, owing to the cessation of the periodical, was never completed. Dr. F. LEYDIG, in " Ueber den Bau und die systematische Stellung der Eaderthiere," 5 after a full description, accompanied with figures, of many species, three of which are new, proceeds to deal with the structure of the Eotifera as a class, and to arrange them in a system of his own. He further discusses their true position in the animal kingdom, and assigns them a place among the Crustacea. Professor HUXLEY, in his paper, " On Lacinularia socialis ; a Contribution to the Anatomy and Physiology of the Eotifera,"6 takes this Eotiferon as his text, and, while minutely describing its structure, discusses various questions concerning that of the whole class. He enters into the general relations of the Eotifera to other animals, and arrives at the conclusion that they are permanent forms of Echinoderm larvae. Herr C. VOGT, in his treatise, " Eiiiige Worte iiber die systematische Stellung der Eaderthierchen," 7 combats Leydig's reasonings and conclusion oil the position of the Eotifera, and affirms that they must be classed among the Vermes. In Dr. F. COHN's essay, "Ueber die Fortpflanzung der Eaderthiere,"8 the males and females of three species are minutely described, especially with reference to their reproductive organs ; and the general question of the reproductive system of the whole class is also discussed. Dr. W. MOXON's 'Notes on some Points in the Anatomy of the Eotatoria"9 call 1 Trans. Micr. Soc. 1852. 2 Phil. Trans. 1856. 3 Phil. Trans. 1857. 4 Popular Sci. Rev. 1862 and 1863. 5 Leipzig, 1854. 6 Trans. Micr. Soc. 1853. 7 Sieb. u. Koll. Zcits. 1855. 8 Sieb. u. Roll. Zcits. 1856. » Trans. Linn. Soc. 1864. 22 THE EOTIFEEA. attention to the right use of the terms "dorsal " and "ventral " as applied to the Eotifera ; to the existence, in many species, of three antennae, holding definite positions with respect to the dorsal and ventral surfaces ; to the true nature of the entrance to the crop of Floscularia ; and to the structure and function, of the vibratile tags. Dr. W. SALENSKY's paper, " Beitrage zur Entwicklungsgeschichte der Brachionus urceolaris," l traces the gradual changes in the ovum, from its first division into two unequal spheres, through its complete segmentation, to the formation of the germinal layers, and the evolution of the various organs of the completed embryo. Although the paper deals with only a single species, it is our principal contribution to the Em- bryology of the Eotifera. Dr. S. BARTSCH, in " Eotatoria Hungarise," 1877, and Herr KARL ECKSTEIN, in " Die Eotatorien der Umgegend von Giessen," 2 have published treatises containing descriptions and figures of local Eotifera (forty in Hungary and fifty at Giessen), in- cluding two or three new species ; as well as new classifications of the whole class KOTIFEBA. Of Dr. BARTSCH's work I can say but little, as it is written unfortunately in Hun- garian. His figures, though somewhat archaic, are well worth the studying ; and he gives drawings and descriptions (happily this time in Latin) of six new species, of which two had been recorded in England some years before. Herr ECKSTEIN also gives many interesting details of his local species, of which two are new ; and adds a general discussion of the structure, development, affinities, and classification of the whole class. His treatise also contains useful lists of synonyms, as well as a good bibliography of the subject. The last edition (1861) of PRITCHARD's " History of Infusoria," by Dr. Arlidge and others, is a work differing in character from any of the above. About one-sixth of it is devoted to the Eotifera, and contains descriptions of the whole of the then known species, illustrated by a great many figures. Both the descriptions and the figures have been mainly taken from Ehrenberg's work, which is closely followed throughout ; but they have been supplemented by others taken from the various treatises mentioned above. As a compilation, it is not only the best, but almost the only, English work on the subject. It contains, moreover, an admirable and exhaustive treatise on " The General History of the Eotifera " as a class, dealing minutely with their structure, reproduction, development, systematic position, and classification. This original and most valuable essay may be said to be indispensable to all students of the Eotifera, bringing together, as it does, into one point of view, the opinions of all the best observers, on the many vexed questions that these little creatures have given rise to, not only as to their organization and development, but as to their relations to the rest of the animal kingdom. A full list of works on the Eotifera, including numerous papers that have been published in various scientific periodicals, will be found at the end of this work. 1 Sieb. u. Kijll. Zcits. 1872. - Sieb. u. Koll. Zeits. 1883. CHAPTBE III. ON THE CLASSIFICATION OF THE ROTIFERA. Omnis enim systematica divisio clandicat lacunisque laborat ; optima est, quae paucioribus horret, documentum satis splendidum, mortales non e vero visionis piuicto Naturain contemplari. — 0. F. MULLER. Tous les ordres des etres naturels ne forment qu'une seule chaine, dans laquelle les differentes classes, comnie autant d'anneaux, tiennent si etroitement les ones aux autres, qu'il est impossible aux sens, et a Pimagination meme, de fixer precisement le point, ou quelqu'une commence ou finit.- — LEIBNITZ. Dum inter ea, quae determinatis characteribus discreta, et certis quasi limiti- bus inclusa sunt, semper intermediae quaedam species reperiuntur, quse, utriusque proxirne accedentis speciei, aliquid possideant, et ita copulationem quasi duarum diversarum specierum constituant ; coloram ad instar, qui ita commiscentur et quasi confluunt, ut nemo veros cujusque fines deterniinare possit. — J. BASTER. CHAPTER III. ON THE CLASSIFICATION OF THE ROTIFERA. FOUR attempts have been made to improve upon Ehrenberg's classification : viz. that of Dujardm in 1841, of Leydig in 1854, of Dr. S. Bartsch in 1877, and of Herr K. Eckstein in 1883. I do not intend to discuss here the various merits and faults of these five systems ; it is enough to say that they all seem to have the fault of needlessly bringing together animals that are different in structure, while separating others that closely resemble each other; I say "needlessly" because perfect classification appears to be an impossibility, except at that fleeting stage of our knowledge when none but the commoner genera are known to us. These usually differ from each other in a marked fashion, the very fact of their wide-spread co-existence being perhaps due to their differ- ing so as not to interfere with each other. When, however, continued search has brought to light the rarer forms, these usually prove to be links between the more common ones ; and then the troubles of the classifier begin. For these strange forms, which are the delight of the naturalist, are the classifier's despair. Do what he will, no system that he can devise will put into Nature those sharp divisions and well-marked gaps that are so dear to him, but of which she knows nothing. Nature has but one law, that of infinite variety ; and the utmost that the classifier can do is to group his animals as well as he can round certain typical forms, content to have the symmetry of his plans and the sharpness of his definitions marred by forms that perversely bear the characteristics of two or three of his types, in nearly equal proportions. He may take comfort, nevertheless ; for, even if he had been able to invent a thoroughly satisfactory classification, it is from the nature of the case written in sand- He can never say as he throws down his pen : Exegi monumentum sere perennius ; for it is almost certain that the fresh discoveries of the next ten years will require his work to be re-cast ; and no higher praise could be given to Ehrenberg's system than that, in spite of new discoveries and its own obvious faults, it has reigned alone for nearly five times the usual period. The Eotifera may first be divided into four natural orders, according to their modes of locomotion, and the structure of the foot. The first of these ideas appears in Dujardin's classification, and the second in Leydig's, and they are both excellent; for there are Eotifera that swim by means of their ciliary wreath, and skip by the help of their arthropodous limbs ; Rotifera that swim only with their wreath ; others that swim and creep like a leech ; and lastly, some that, when adult, are stationary. Moreover, in three of these four orders there is only one form of foot in each order, and that form is unlike those of the other two ; and although in one order there are more forms of the foot than one, still they are all unlike the forms of the other three. Nor is this all. The natural character of these four orders is further shown by there being other important points of structure, in which the animals comprised in each order at the same time resemble each other and differ from those of the other orders. 26 THE ROTIFERA. I have named these orders as follows : I. EHIZOTA (the rooted). Fixed when adult. II. BDELLOIDA (the leech -like). That swim with their ciliary wreath, and creep like a leech. III. PLOIMA (the sea- worthy). That only swim with their ciliary wreath. IV. SCIETOPODA (the skippers). That swim with their ciliary wreath, and skip with Arthropodous limbs. Now, the creatures contained in these orders, with a few exceptions, differ from each other, first in their habits, and secondly in the following points : I. In the structure of the foot. II. In the arrangement of the ciliary wreath. III. In the form of the trophi. I. In the structure of the foot. (1) The RMzotic foot is transversely wrinkled, and ends in a sort of sucking disk (fig. 16) or cup (fig. 17). It is not retractile within the body, it never has telescopic joints, nor is it ever furcate. Fig. 16.— Hliizotic foot. Fig. 1 7.— Rhizotic foot. (FlosculaHd campanulaia.) (AMicerla ringetu) Fig. 18.— Bdelloidic foot. {Rotifer citrinus) Fig. 19. — Scirtopodic foot. (I'edalion mirum) (2) The Bdelloidic foot (fig. 18), on the contrary, is telescopic, retractile, furcate, and is never transversely wrinkled, nor terminated by a sucking disk. (3) The Scirtopodic foot (fig. 19) is unique ; it is divided into two unconnected, smooth, jointless styles, each ending in a ciliated expansion. (4) The Plo'imic foot is various in shape, but is always unlike that of any other order ; for — (a) if transversely wrinkled, it is yet retractile within the body, and almost invari- ably furcate ; (b) if jointed and furcate, it is not also telescopic ; (c) occasionally it is absent altogether. II. In the arrangement of the ciliary wreath.1 (1) The Rhizotic ciliary wreath is of two forms : (a) The first encircles the body twice, by bending on itself; thus inclosing the mouth, and having a dorsal gap between the points of flexure, figs. 20, 21. Fig. 20.— RMzotic wreath («), front view. Fig. 21.— Rhizotic wreath («), side view. (Alelicerta ringens) (JJclicerta Hwjens) Fig. 22.— Rhizotic wreath (&), seen from above. (Flosculuria campanulata) 1 In figs. 20 to 25, cio is the ciliary wreath ; pw is the principal wreath ; sio is the secondary wreath bf the buccal funnel ; Ip the lips. ON THE CLASSIFICATION OF THE KOTIFEEA. 27 (b) The second form is a simple segment of a circle, placed on the ventral side above the mouth, fig. 22. (2) The Bdelloidic wreath is also of two forms : (c) The first, like the Ehizotic wreath (a), is a double wreath surrounding the body twice, and inclosing the mouth ; but, unlike the Ehizotic, it has two gaps instead of one, viz. a dorsal gap between the points of flexure, and a ven- tral gap in the upper wreath opposite to the mouth. (d) The second form of Bdelloidic wreath is a mere furring of the corona on its ventral surface, as shown in fig. 25. (8) The Ploimic wreath is very various in shape, but is never Ehizotic, while it is Bdelloidic only in one genus.1 Fig. 23.— Bdelloidic wreath (c), from above. (Rotifer citrimis) Pig. 24.— Bdelloidic wreath (c), side view. (Rotifer citnnus) Fig. 25.— Bdelloidic wreath (d). (Adineta vaga) (4) The Scirtopodic wreath is of Bdelloidic type. III. In the form of the trophi. If we disregard two genera 2 of the Plo'ima (not one-fifteenth of the whole number of Ploimic genera), we can then say that the first three orders differ also in the forms of their trophi. For — (1) The Ehizotic trophi are either malleo-ramate,3 fig. 26, or uncinate,3 fig. 27. (2) The Bdelloidic trophi are always ramate3 fig. 28. (3) The Plc'imic trophi are of various forms ; but are never Bdelloidic, and are Ehizotic only in two genera.2 (4) The Scirtopodic trophi are of a Ehizotic type, being malleo-ramate, fig. 26. Fig. 26. — Malleo-ramate. (Melicerta ringens) Fig. 27. — Uncinate. (Stephunoceros Eichhornii ) Fig. 28.— Ramate. (Rotifer citrinus) Now, in reviewing the points of agreement and of difference in the four orders, we may at once set aside the fourth order, the Scirtopoda, as unmistakably separated from the others. This order contains but one family, which has only one genus, and that genus itself consists of only one species.4 In fact, it has been formed to contain that remarkable creature Pedalion mirum, which I discovered at Clifton in 1871. This 1 Pterodina. 2 Triarthra and Pterodina. 3 For the explanations of these technical terms, see p. 29. 4 I pass over for the present Dr. Schmarda's Hexarthra polyptcra, and will discuss it under the Pedalionidce. 28 THE EOT1FEEA. Eotiferon has six hollow limbs continuous, in true Arthropodous fashion, with the body- cavity, and worked by opposing muscles passing down them, and is thus plainly linked to the Crustacea and Insecta. Pedalion, in fact, is a Nauplius larva, and is yet a Eotiferon. Order IV., then, is sufficiently separated from the rest by its Arthropodous limbs, and by the use made of them ; and of the other three orders, it has been shown above that, if we disregard some points of only two genera, we may say of orders I. II. III. that they differ inter se in their habits, and in the structure of their feet, trophi, and ciliary wreaths. This seems a satisfactory first step towards classification ; but it is only fair to the reader to warn him that it has been gained by omitting some parasitic Eotifera, as well as a few very troublesome forms, such as Trochosphcera, Apsilus, Microcodon, &c. I have dwelt on the differences in structure, as well as in habits, between the four orders, in order to show that these four groups are natural ; but I do not propose to use as ordinal characteristics any others than the mode of locomotion and the structure of the foot ; and for this reason, that each of the first three orders has more than one form of the trophi, or of the ciliary wreath, or of both. The Rhizota, for instance, have two forms of the trophi, and two of the ciliary wreath. The Bdelloida have two forms of wreath, and the Plo'ima have many different forms of both wreath and trophi. Before I proceed to divide these four orders into families, I must, however, digress a little to explain and name the various types of trophi, as the classification partly depends upon them. Mr. Gosse's treatise on " The Manducatory Organs in the Class Eotifera" essays to show that these organs present seven principal types of structure, distinguished from each other by the prominence of some particular part. To make this clear, it may be as well to re-state that, in the mastax of a Brachionus, there are two hammer-like bodies or mallei (fig. 29, ms), which work on a kind of split anvil or incus (fig. 29, is), and that each malleus consists of an upper part, the head or uncus (fig. 29, us], and of a lower part or handle, the manubrium (fig. 29, mm) ; while the incus consists also of two parts, the upper divided into two symmetrical halves, the rami (fig. 29, rs), which are supported on the lower piece or fulcrum (fig. 29, /m). Now, in Brachionus all the trophi are well developed, but the other typical manducatory organs may be arranged •fm " in a series in which the mallei are successively degraded, Fig. 29.-Maiieatc. while continually greater prominence is given to the incus ; at least in all but three types ; and in two of these the rami and unci are the prominent parts, while the third is distinguished by the close connection of the mallei and the rami. The typical trophi may, then, be named as follows : 1. Malleate (fig. 29). Mallei stout ; manubria and unci of nearly equal length ; unci 5- to 7-toothed ; fulcrum short ; as in Brachionus urceolaris. 2. Sub-malleate (fig. 30). Mallei slender ; manubria about twice as long as the unci ; unci 3- to 5-toothed ; as in Euclilanis deflexa. 3. Forcipate l (fig. 31). Mallei rod-like ; manubria and fulcrum long ; unci pointed or evanescent ; rami much developed and used as a forceps ; as in Diglena forcipata. 1 In Furciilaria, and in a few other genera, the rami as well as the mallei are rod-like ; and the ON THE CLASSIFICATION OF THE ROTIFERA. 29 4. Incudate (fig. 82). Mallei evanescent ; rami highly developed into a curved forceps ; fulcrum stout ; as in Asplanchna Ebbesbornii. 6. Uncinate (fig. 27). Unci 2-toothed ; manubria evanescent ; incus slender ; as in Stephanoceros Eichhornii. Fig. 30.— Sub-malleate. Pig. 31.— Forcipatc. Fig. 32.— Incudate. 6. Eamate (fig. 28). Rami sub-quadrantic, each crossed by two or three teeth ; manubria evanescent ; fulcrum rudimentary ; as in Philodina roseola. 7. Malleo-ramate (fig. 26). Mallei fastened by unci to rami ; manubria 3 loops soldered to the unci ; unci 3- toothed ; rami large, with many striae parallel to the teeth ; fulcrum slender ; as in Melicerta ringens. Now, the seven Rotifera, made use of above to yield examples of typical trophi, are very distinct from each other, and show that the form of the trophi is one good charac- teristic for separating the families. But a difference in the shape and disposition of the corona, and of its ciliary wreath, generally accompanies a difference in the manducatory organs ; and the three together will serve as good guides to a division of the four orders into families. In one of the sub-divisions of the Plo'ima, however, the corona, ciliary wreath, and trophi are often difficult of determination ; but just where these guides desert us, a new character, viz. the lorica, comes to our aid, and shows such well-marked differences in shape and structure, as to enable us to divide this sub-order (the Loricatd] into fairly natural groups. The Loricata are so called from the integument of the body ; which, from the distribution of chitine throughout the tissue, is hardened into a stiffened coat or shell (lorica, a coat of mail) inclosing, more or less completely, the internal organs. In the Il-loricata the integument is soft and flexible ; but there is, unfortunately, no very sharp division between the two sub-orders in this respect ; as every variety of integu- ment exists, from the hard, dense coat of Dinocharis, to the tough yet flexible covering of Rattulus, and the perfectly soft cuticle of Albertia. ' The following scheme, then, is an attempt to divide the four orders of Rotifera into families, by means of the various characters which I have just detailed : of course, there are some genera which do not readily fall into the arrangement ; but this is only what is certain to happen to every possible scheme of classification. Such difficulties must attend every attempt to marshal Nature's endless varieties into whole apparatus looks like one forceps within another, term virgate will be applied. To this variety of the forcipate trophi the 30 THE EOTIFEEA. , well-marked battalions. Nature knows no hard lines of separation; and the best of classifications can be only that which contains the fewest faults : as Miiller has forcibly said, ' Optima est, quae paucioribus horret.' Order I. EHIZOTA. Fixed when adult; foot transversely wrinkled, not retractile within the body, ending in a slicking disk or cup. Fam. 1. FlosculariadcB (PI. C, fig. 1). Corona produced longitudinally into setigerous lobes ; buccal orifice central ; ciliary wreath a single half-circle above the buccal orifice ; trophi uncinate.1 Fam. 2. Melicertadce (PI. C, fig. 2). Corona not produced into setigerous lobes ; buccal orifice lateral ; ciliary ivreath a ; marginal continuous curve, bent on itself at the dorsal surface, so as to encircle the corona twice, with the buccal orifice between its upper and lower curves, and having also a dorsal gap between its points of flexure ; trophi malleo-ramate.1 Order II. BDELLOIDA. That swim with their ciliary wreath, and creep like a leech ; foot wholly retractile within the body, telescopic, furcate. Fam. 3. Philodinadee (PI. C, fig. 3). Corona two transverse circular lobes ; ciliary ivreath a marginal continuous curve bent on itself at the dorsal surface, so as to encircle the corona twice, with the buccal orifice between its upper and lower curves, and having also two gaps, the one dorsal between its points of flexure, and the other a ventral gap in the upper curve opposite to the buccal orifice : trophi rarnate.1 Fam. 4. Adinetada (PI. C, fig. 4). Corona a flat ventrally placed surface ; ciliary wreath the furred ventral surface of the corona ; trophi ramate.1 Order III. PLO!MA. That swim ivith their ciliary ivreath, and (in some cases) creep with their toes. Sub-order. Il-loricata. Foot, when present, almost invariably furcate ; but not transversely wrinkled ; rarely more than feebly telescopic, and partially retractile. Fam. 5. Microcodidce. Corona obliquely transverse, flat, circular ; buccal orifice central ; ciliary wreath a marginal continuous curve encircling the corona, and two curves of larger cilia, one on each side of the buccal orifice ; trophi forcipate ' ; foot stylate. Fam. 6. Asplanchnadce (PL C. fig. 7). Corona two transverse, flattened, confluent cones, with their summits distinct ; ciliary wreath single, marginal; trophi incudate ' ; intestine, cloaca, and/ooi, absent. 1 For description of these technical terms, see pp. 28, 29. ON THE CLASSIFICATION OF THE ROTIFERA. 81 Fam. 7. Synchcetada (PI. C. fig. 6). Corona a transverse spheroidal segment, sometimes much flattened, with styligerous prominences ; ciliary wreath a single interrupted marginal curve, encircling the corona ; trophi forcipate l ; foot minute, furcate, or absent. Fam. 8. Triarthradce. Corona transverse ; ciliary wreath single, marginal, fringing the buccal orifice ; trophi malleo-ramate l ; foot absent. Fam. 9. Hydatinada (PI. C, fig. 5). Corona truncate, with styligerous or ciliated prominences ; ciliary wreath two parallel curves, the one marginal fringing the corona and buccal orifice, the other lying within the first, the styligerous prominences being between the two ; trophi malleate J \foot furcate. Fam. 10. NotommatadcB. Corona obliquely transverse ; ciliary wreath one of interrupted curves and clusters, usually with a marginal wreath surrounding the buccal orifice ; trophi forcipate l ; foot furcate. Sub-order. Loricata. Corona and ciliary wreath various in shape, but never Bhizotic, and Bdelloidic only in the Pterodinada : trophi of different types, but never Bdelloidic, and Khizotic only in the Pterodinadce. Division I. Foot jointed, stylate or furcate ; not transversely wrinkled, nor wholly retractile. Fam. 11. Rattulid®. Lorica entire, cylindrical, without angles ; trophi asymmetrical. Fam. 12. Dinocharida. Lorica entire, vase-shaped, sometimes facetted ; head distinct, with a chitinous covering ; trophi symmetrical. Fam. 13. SalpinadcB. Lorica compressed, cleft down the back, the two halves united by a membrane, so as to form a dorsal furrow. Fam. 14. Euchlanidce (PL C. fig. 10). Lorica depressed, of two dissimilar plates, one dorsal and one ventral, united by a membrane so as to form a lateral furrow. Fam. 15. LepadelladcB. Lorica depressed, broad, closed beneath ; head distinct, surmounted by a retractile, arched, chitinous plate. Fam. 1G. ColuridcB. Lorica compressed, open beneath ; head distinct, surmounted by a retractile, arched, chitinous plate. 1 For description of these technical terms, see pp. 28, 29. 32 THE ROTIFERA. Division II. Foot transversely wrinkled, wholly retractile, furcate or ending in a ciliated cup ; sometimes absent. Fam. 17. Pterodinad® (PI. C. fig. 9). Lorica greatly depressed, entire, of two nearly equal plates, soldered together at the edges ; corona and ciliary wreath those of the Philodinadce ; trophi malleo-ramate ] ; foot ending in a ciliated cup. Fam. 18. Brachionida (PI. C. fig. 8). Lorica depressed, entire, dorsally arched, generally armed with anterior and posterior spines ; corona transverse with styligerous prominences ; ciliary wreath single, marginal, fringing the buccal orifice ; trophi malleate ' ; foot furcate, or absent. Order IV. SCIRTOPODA. That sivim with their ciliary wreath, and skip with Arthropodous limbs ; foot replaced by two dorsal, stylate, unconnected appendages, ending in ciliated expansions. Fam. 19. Pedalionid® (PI. C. fig. 11). Corona truncate ; ciliary wreath a marginal continuous curve, bent on itself at the dorsal surface, so as to encircle the corofla twice, with the buccal orifice between its upper and lower curves ; having also two gaps, the one dorsal between its points of flexure, and the other a ventral gap in the upper curve opposite to the buccal orifice ; trophi malleo-ramate.1 The further subdivision of each family into genera will be given with the description of that family. 1 For description of these technical terms, see pp. 28, 29. CHAPTEE IV. ON THE HAUNTS AND HABITS OF THE ROTIFERA. Nonne vides, qusecunque mora fluidoque calore Corpora tabuerint, in parva animalia verti ? OVID, Metam. xv. 8G2. Equidem turn Naturae rerum gratias ago, cum illam non ab hac parte video, quae publica est, Bed cum secretiora ejtis intravi. Curiosus spectator excutit singula et quaorit. Quidni quacrat ? Scit ilia ad se pertinere. Quantum enim est, quod ante pcdcs jacet. SENECA, Nat. Quccst. praef. (adapted by 0. F. MULLER). 'T is born with all : the love of Nature's works Is an ingredient in the compound man, Infused at the creation of the kind. . . . It is a flame that dies not even there Where nothing feeds it : neither business, crowds, Nor habits of luxurious city life, Whatever else they smother of true worth In human bosoms, quench it or abate. COWPER, The Task. CHAPTEK IV. ON THE HAUNTS AND HABITS OF THE EOTIFEEA. THAT the first thing to be done is to catch your game, is a maxim as applicable to Rotifera as to hares ; and it is no less true of these that to hunt for them successfully requires some knowledge of their haunts and habits. To carry away from a pond's side a bottle of dirty water full of slimy weed, is by no means a good plan for catching these animals, even the commonest and coarsest. It is true that there are some fine forms which may be found in very dirty ponds, or even in dirtier puddles : for instance, there can hardly be too dark a farmyard puddle for Hydatina senta, which rejoices in the drainings of a manure-heap, even when the water is of so deep a colour that it is impossible to see the animals in it when you have got them. Triarthra, too, and the beautiful Notops l clavulata are to be met with in cattle ponds, where the water is like pea-soup ; and Brachioni of all kinds rejoice in such places, especially when green with Euglence, and alive with the motile seeds of algas. Indeed, there is one Brachionus, B. angularis, whose presence in a pond bids us put up our bottles and go elsewhere, as it likes water that will support hardly any Eotiferon but itself. Floscularia, Stephanoceros, Melicerta, Limnias, and (Ecistes are, of course, to be found only in such places as pond weeds will grow in healthily. Old ponds that have been left long undisturbed are their favourite haunts. Floscularia is a very wide-spread genus, at least so far as one or two species are concerned ; and these may be looked for with every prospect of success in any such pond. Most of the finer and rarer kinds have been found in the Scotch lakes by Mr. Hood, who during the last four years has doubled the number of recorded species by his discoveries in the lochs round Dundee. Stephanoceros, though by no means a rare Eotiferon, is more partially distributed ; it is found often enough in ponds near London and Birmingham ; but I have not heard that a single specimen has ever been met with in the neighbourhood of Clifton. It appears also to be rare in Scotland ; as Mr. Hood has found it only once or twice, in marsh-pools in Perthshire. Melicerta ringens is to be found almost everywhere. It has even been seen swarming in one of the aquaria in the parrot-house in the Clifton Zoological Gardens. The roots of duckweed, the fibres of alga3, the leaves of Myriophyllum, and of all sorts of water plants, bear this very common species, as they do also the tubes of Limnias and (Ecistes. Lacinularia and Megalotrocha have similar tastes, but are less frequently met with, especially the latter. This must be a comparatively rare genus, as it has been sent to me but three or four times in many years. Cephalo siphon is also rare. I once found a large colony of it, on a water weed at Nailsea in the big pond near the railway station, and it has been sent to me from Cheltenham and London ; 2 but I never met with it again. Conochilus is a lover of clear water. I have found it in Loch Lomond, and Dr. Imhoff has obtained it, in abundance, in the middle of Lake Zug. It is common enough 1 Notommata clavulata ; Ehr. 2 Miss Saunders, to whom I am indebted for the specimens, says : " It is curious I never before came across this tube-dweller in the hundreds of pools I have searched." 86 THE EOTIFEEA in clean ponds round about London, and Mr. Hood has met with it in the Perthshire lochs. The Notommatadiz and Philodinada have a very wide range. The algae of ponds always hold many species, and many haunt the sediment that lies on the mud of the bottom. Some of the latter are to be found in gutters of houses, in water-butts, on the blades of wet grass, anywhere indeed where dust can fall and moisture can follow. For the eggs of the Eotifera are blown away by the wind from dried-up puddles, and are scattered broadcast through the air ; and some of the creatures they give birth to can exist appa- rently under almost any kind of moist conditions. The Asplanchnadce fortunately are as indifferent to their quarters as they are large and handsome. I have found them in roadside ditches thickly covered with Lemna, in farmyard ponds, in the clear water of a miniature lake, and in a foul yellow-green duck puddle in which the fluid (it could not be called water) was so thick that it had to be diluted with five or six times its own bulk, before anything could be seen in it. An Asplanchna is the very beau-ideal of a Kotiferon for a beginner. It is very large and transparent ; it swims slowly ; and it generally occurs in great numbers. Moreover, its male is even more transparent than the female ; a mere living bubble, thinner and clearer than the finest blown glass. Unhappily, they are as capricious as they are charming; for a pool may be full of them to-day and deserted to-morrow ; and, so far as my own experience goes, they do not occur in the same spot year after year as many Kotifera do. The Pterodinada and Euchlanidcs are dwellers in clear ponds, and rather solitary in their habits. On a warm sunny day the latter may be captured by skimming off the Lemnce, and floating bits of leaves and stems, that are driven to the leeward corner of a pool. If the bottle be then allowed to stand a few minutes till .the water is clear, a Euclilanis will often be found slowly gliding up the glass with its long toes pressed against it. It is always worth while to capture it at once with a pipette, and put it into a small tube along with any others of the same kind, as a live Euchlanis, properly ex- hibited under dark-field illumination, is one of the choicest treats that the Eotifera afford. The Pterodinadce are almost always creeping about the algae on the sides of the pond. I never but once have seen any other than solitary specimens, but that exception was a notable one. I then found swarms of them in the small space in which the sluice gate of a pond worked. It was not more than a foot square by about four feet deep, and was mantled over by duckweed. Out of this unlikely spot they were dipped by hundreds, while not one could be found in the pond itself. Of course the duckweed came up with the Ptero- dina}, though as little as possible was taken, on account of the disagreeable way in which it clings to every pipette put into the bottle. On this occasion, however, I found it of great service ; for, on inspecting the catch at home with a hand-lens, I noticed that the Eotifera were attached in clusters to some of the roots of the duckweed. Watch- ing for a favourable opportunity, I whipped one of the Lemncz out of the water so suddenly, that the creatures had not time to let go their hold. Then cutting off the green head, I coiled the stem into a circle on a glass slip, and covered it with thin glass. It was impossible to conceive a more beautiful sight than this natural cage now afforded me. Thirty or forty shields of living glass were flashing across the field of view in every direction ; some were adhering to the stem, swaying backwards and forwards so as to present themselves in every point of view, while others were moored to the glass cover, thus giving an admirable opportunity of making out their structure. It was a memorable occasion, but I never had such a chance again. I have yet to speak of the SynclicBtada among the free-swimmers. Both the genera included in this family are to be found in open water ; and both alike shun dirty ponds ; though in different degrees, for Synchceta is absolutely intolerant of them ; while I have dipped up Polyarthra from. the hollows of a muddy bottom where once a pond had been. Both genera are tolerably common, and are often to be caught in considerable numbers. The habits of the two chief species of Synchceta, viz. S. mordax and S. ON THE HAUNTS AND HABITS OF THE BOTIFEEA. 87 tremula, are very different. The former is the swiftest and most restless of the Eotifera ; it is the very swallow of the waters, ever whirling round and round in endless spirals, and never still for a single instant from its birth till its death ; but the latter may be constantly seen drifting along in some gentle current, while twisting round at the end of a long thread spun from its toes, and fastened to some floating object. Of course, creatures with habits like these can be captured only by making random dips in the water, now at the surface, now deeper down — here in the sunshine, and there in the- shade ; for even Eotifera have their fancies, and are sometimes swarming in one particular spot, while all the rest of the pond is deserted by them. I have, however, noticed that they specially affect the neighbourhood of a forest of weeds growing up from the bottom ; waltzing up and down outside of them in myriads, like gnats under the trees in summer. There is yet another free swimmer that avoids the shore, and sails out into the open sea ; viz. Anurcea longispina. This curious creature has a lorica like a Greenlander's canoe, or a University eight, and it keeps off from the weeds and algse, as if fearing lest it should be entangled for life if it once got among them. It was discovered by Professor D. S. Kellicott in Niagara water at Buffalo, U.S., in 1879, and was found almost imme- diately afterwards in the Olton Eeservoir, near Birmingham and since then in Lake Zug in Switzerland. It appears to be a rare species, though its rarity may be due partly to the fact that it often requires a boat to catch it ; and an ordinary Eotifer hunter can hardly be expected to add this to his apparatus. The known habitats of the Pedalionidce are at present very few. I had the good fortune to be the first to light on Pedalion mirum. It was in July 1871 that I found it in a small roadside pool at the top of Nightingale Valley, close to Clifton. Soon afterwards I dipped it from a fine old pond at Abbot's Leigh, about two miles distant from Nightingale Valley. It reappeared in this pond in the following year, but since then it has not revisited* the neighbourhood. It has been met with several times near Chester and Birmingham, and on one occasion it was tolerably abundant in the warm water-lily tank in the Duke of Westminster's gardens at Eaton.1 Dr. L. K. Schmarda discovered in Egypt, in 1853, in some brackish pools near El-Kab, a six-limbed Eotiferon, Hexarthra polyptera, which evidently belongs to the same family, though it must be placed in a different genus. He says that there were great swarms of them distinctly ^visible to the naked eye, in a pool of very transparent, colourless water, of a strong brackish taste. Now, a Eotiferon that is equally at home in dirty puddles, clear ponds, warm-water tanks, and brackish pools, ought not to be a rare one : and yet Pedalion is rare. Possibly its apparent rarity is due to its being constantly mistaken for an Entomo- stracous larva. I was on the point of throwing the water away, when I first dipped Pedalion out of the pool in Nightingale Valley. Its skipping movement is so precisely that of the young of a Cyclops, that I thought I had caught nothing more valuable than these ever-present nuisances. Fortunately I noticed that, unlike them, my captives seemed to glide along after every skip, instead of stopping stock-still to gather breath for a fresh jump ; and so, thinking that they might possibly be some large sort of Polyarthra, took them home for further investigation. But it is very probable that Pedalion has been thrown away hundreds of times, and will be so again, as this happened to me after nearly twenty years' experience in catching Eotifera.2 » Eotifera may often be seen perched just under the plumed heads of one of the fresh - 1 M. J. Barrens described, in the Revue Scientifiquc, No. 13, 1877, p. 303, a marine Eotiferon under the name Pedalion, and gave an account of its embryology. His description, however, shows that the animal was of the genus Syncli/zta. 2 It is a pity that Pedalion is not more frequently met with, as there are some points in its structure that yet remain to be cleared up ; and as it is such a striking link between the Eotifera and the Arthropoda. Mr. T. Bolton, of Birmingham, has, however, succeeded in preserving specimens as micro- scopic objects, and they can generally be obtained from him. 38 THE KOTIFEBA. water zoophytes, wisely making use of the stronger currents produced by the ciliated tentacles of their hosts, in order to bring grist more easily to their own mills. I have had sometimes quite "a happy family " of them in the field of view at once ; a Brachionus, a Philodina or two, and a Melicerta, all attached to the neck of a Plumatella, and all eagerly whirling their wheels in order to divert to their own throats a portion of the currents that swept down to them from above. Nor was this all ; for the Melicerta in its turn had the top of its tube turned to the same use, and bore, as closely under its wheels as possible, the tiny case of one of its own offspring. Limnias ceratophylli and Melicerta ringens carry this semi-parasitical habit to a great extent. Clusters of two or three generations all attached to one tube are not at all uncommon in the former species ; and I once found in Nailsea pond a large Lim- nias bearing up no fewer than fourteen of its own descendants. Melicerta ringens, too, in America,1 is frequently met with in large adhering clusters, but in England it is usually a solitary species. However, this is not invariably the case ; for not long ago I had the pleasure of seeing as many as thirty-four live Melicertce attached to one another.2 They were of all ages and sizes, and were grouped round one large tube, so as to form a striking example of a natural co-operative society. Nor is this the only way in which the Eotifera show their capacity for fighting the battle of life. Every animal is limited by its own powers to a certain space, beyond which its excursions cannot possibly extend. Its food and its mate must be found within these limits ; and when these two imperious wants are satisfied, there is but little time or strength left for travelling. But it would be an obvious advantage to many creatures if they could be carried about from one spot to another without tiring their own muscles — ready to slip off, at any favourable opportunity, " to fresh woods and pastures new." Now this is precisely what some of the Brachioni and Philodince contrive to do ; for they may be seen riding in clusters on the backs and sides of the Entomostraca, or thickly fringing the legs and side plates of the water wood- louse.3 Whenever I have caught a water-flea 4 so encumbered, and have placed it in a live box to see the Eotifera it carried, they have soon deserted their captive steed, and have swum off as if to search for a more serviceable one. There are, too, some Eotifera whose structure has been adapted to give them a good grasp of their host, or even to enable them to pierce its skin, and so suck its juices for their own support. Balatro calvus, for example, has been found5 in the Seine (Canton de Geneve) creeping on the bodies of small water-worms Avhich it habitually infests, and having two greatly enlarged foot-processes, which probably enable it to take a firm hold. Another Eotiferon, Callidina parasitica, is always found attached to the thoracic or abdominal appendages of the fresh-water shrimp 6 and water wood-louse, and limits its journeyings to creeping about on the body of its host ; while the strange creature Drilopliaga Bucephalus holds on by its altered jaws to the hind segments of a fresh- water worm, Lumbriculus variegatus, and sucks the animal it clings to. This parasitic Eotiferon was discovered in North Bohemia in the great pond at Hirse- berg, in the banks of which the Lumbricidus is found in immense numbers. The worm buries the fore part of its body in the mud, and moves its naked hinder segments like a pendulum in the water. But no such gentle motion will unfasten the grip of the Drilopliaga, which is so firmly attached to the worm's skin that it can be dislodged only by using considerable force. 1 I am indebted to Mr. Galloway C. Morris, of Philadelphia, U.S., for this information. A cluster of tubes that he sent to me contains twenty-eight specimens of various sizes adhering together. - It was found near Clifton by Mr. E. C. Bousfield, and is drawn in PI. V. fig. 1, e. 3 Asellus vulgaris. 4 Daphnia pulex. 5 By M. Ed. Claparede. See Ann. Sci. Nat. Zool. 5 Ser. vol. viii. 1867. 6 Gammarus imlex. Mr. H. Giglioli, who discovered this species, says that he has never found it anywhere else ; and that out of 700 Gammari from different localities, not one was free from the para- site. Quart. J. Micr. Sci. N. Ser. vol. iii. 1863, p. 237 ON THE HAUNTS AND HABITS OF THE EOTIFEEA. 89 If the Eotifer hunter can rise to a pitch of enthusiasm, which I confess I have never been able to attain to, he may follow Dujardin's example, and, by making incisions l in the sides of earth-worms and slugs, obtain from the expressed fluids the entozoic Albertia vermiculus. The same creature has been seen inside Nais proboscidea,* and an allied species (Albertia crystallina) in the viscera of Nais litloralis,3 while the Synaptas of the Channel Islands have been found4 to carry within their body-cavity a minute Kotiferon only ^jj of an inch in length. But I have not yet exhausted the list of these strange dwelling-places. A pretty little Eotiferon, Notommata parasita, may be found swimming about within the beautiful spheres of Volvox globator, or lodged within the embryo globes when almost ready to escape from the parent sphere. " On examining several specimens of Volvox with a pocket lens we may frequently detect such as are thus tenanted, by perceiving a spot differing from the young clusters in form and colour. These spots are found to be the Notommata, snugly ensconced within the globe, in the spacious area of which it lives at ease, and swims to and fro like a gold fish in a glass vase. We see it for the most part, however, clinging to the inner surface of the circumference, engaged in devouring the green monads with which the gelatinous surface is studded, or else eating away the embryo clusters." 8 Volvox globator is common enough in most neighbourhoods, and may be met with it even in clear rain puddles in quarries and plantations ; while in two or three ponds near Clifton it is sometimes so abundant as to give a green hue to the water ; and yet I have never seen its guest here, nor do I know anyone in the neighbourhood who has ; so it can scarcely be a common species. The reproductive cells of Vaitcheria — a thread-like alga which grows on pond walls and in many moist places — are the homes of another Notommata, N. WernecJcii. This parasitic Eotiferon passes a small portion of its youth in the open water ; but it soon re- turns to a lifelong imprisonment in the green cells in which it was hatched, and where it undergoes very singular changes of form. Its presence in the Vauchcria may be detected by the unusual size and shape of the reproductive cells, and by their containing a black spot which is the animal's stomach.6 Some further means of obtaining Eotifera have yet to be mentioned. If a little of the mud or rotten leaves at the bottom of a dried-up pool, in which Eotifera have been observed, is brought home and allowed to lie in a vessel of water, the chances are in favour of there being in the mud some of their eggs, and of their ultimately hatching. I have often adopted this plan with success, especially when some rare species has been discovered in a little pool due only to the rain, and drying up after two or three days' fine weather. Unluckily the mud too frequently harbours an abundance of small worms also ; and these are disagreeable to see and troublesome to deal with, for they are liable to starve, die, and taint the water. Eotifera also may be produced at home by placing infusions of hay, leaves, &c. in some vessel out of doors. No very great variety is to be obtained by such methods ; but it is always as well to try it, and to have a good-sized pan in the garden, full of soft water, into which rubbish from pond-gatherings may occasionally be thrown. These, and the chance droppings from the air into the pan, will sometimes give the student, at his own door, species which he would otherwise have to travel far to find. Many of the Eotifera may be kept indoors in vessels in which there is a healthy growth of Myriophyllum, Anacharis, or other water- weed. Mr. Gosse has tried this plan with 1 Ann. Sci. Nat. Zool. 2 Ser. vol. x. 1838, p. 176. 2 By Mr. P. H. Gosse, in water from a pond at Walthamstow. 3 By M. Max Schulze. 4 By Professor Bay Lankester, Quart. J. Micr. Sci. N. Ser. vol. viii. 1868, p. 54. s Mr. P. H. Gosse, Trans. Micr. Soc. vol. iii. 1852, p. 143. 6 Prof. Balbiani, Ann. Sci. Nat. Zool. 6 Ser. vol. vii. 1878. I am not aware that Notommata Werneckii has been found in England. Probably it would be, were it deliberately searched for. Mr. F. W. Eoper, of Eastbourne, has found a similar species tightly rolled up in a ball inside the leaves of one of the liver-worts, Lejeunia minutissima. 40 THE EOTIFEEA. success, and has lately had thousands of Stephanoceros, Melicerta, Pterodina, &c., thriving in tanks and jars in his study. Mr. J. Hood has been equally successful with Floscularia and Limnias ; and I know of other instances in which a literally constant stock of the tube-making Eotifera has been maintained in these home preserves. Indeed, if nothing more is desired than to watch the growth of a couple of generations or so, an ordinary zoophyte trough is aquarium enough. All that is necessary is, (1) to take great care that there are not many animals of any kind in it, (2) to keep it in a subdued light, (3) and at a moderate temperature, (4) and especially to provide the Kotifera with plenty of their natural food. For, in the great majority of cases, Kotifera die, when in captivity, of starvation : one moment's examination of their stomachs will make that point clear. Of course, what is their natural food must first be observed under the microscope, and then it must be provided to them every day by dropping a fresh supply of water containing it into the trough. It will, therefore, always be necessary to bring away from the pond, where they were found, a good supply of pond water free from all other kinds of animals. Sometimes, however, it is well to make an overfed Rotiferon starve a little, in order to see its internal structure. Hydatina senta, for instance, is frequently so gorged with dark green food, that little else can be seen but its distended stomach ; the organization of Pedalion mirum, too, is often a hopeless riddle, owing to its greedy habits ; but drop either of these creatures into a tube of clear soft water for an hour or so, and it may be fetched out again in delightful condition for microscopic investigation, and yet perfectly healthy. Summing up the various habitats that I have just recorded, we see that Eotifera may be found in rivers, lakes, reservoirs, ponds, ditches, puddles, gutters, and water- butts ; in the mud of dried ponds, in the dust of dried house-gutters, on wet moss and grass ; in the rolled-up leaves of liver-worts, in the cells of Volvox ylobator and of Vaucheria, in vegetable infusions ; on the backs of Entomostraca, and of fresh-water fleas, wood-lice, shrimps, and worms ; in the viscera of slugs, earth-worms, and Naiades, and in the body-cavities of Synaptce. Nor have I yet completed the list ; for several species have been found in the sea. Mr. Gosse says,1 " Synchata Baltica swims at large through the water, never resting ; it is self-luminous, and is one of the causes of the phosphorence of the sea. Brachionus Miilleri and Pterodina clypcata occur in brackish water at the mouths of rivers ; and other marine species may often be detected by searching with a pocket-lens the glass sides of a well-stocked aquarium." Since then these creatures have so wide a range of habitats, it is hardly possible for anyone, who will take the trouble, not to find some of them near his own home. To obtain some particular Eotiferon, at a particular given time, is often difficult enough, if not impossible ; but for one who is content to study these beautiful creatures as he finds them, there is always a never-ending supply of delightful amusement. 1 Marine Zoology, part i. 1855, p. 107. See also Tenby, 1856, p. 274. EXPLANATION OF THE LETTEKING OF THE PLATES. c I sp ch cc h Tip f t cw pw sw tr v ¥ IP t ep mx Corona .... Lobe of do. . Styligerous prominence . Chin .... Ciliated cup . Hook .... Horny processes Foot .... Toe .... Ciliary wreath Principal wreath Secondary wreath . Taster .... Vestibule Bticcal funnel . Lips .... Tube from lips to crop Crop .... Mastax Parts of head and body. Trophi ms malleus \ t^ . or Jaws is (K . (Esophagus S . Stomach . i . . Intestine r . . Eecturn . cl . . Cloaca sg . Salivary gland 99 • Gastric gland . fff • . Foot gland Ic . . Lateral canal . vt . . Vibratile tag . cv . . Contractile vesicle . oy . Ovary 9 • . Germ om . . Ovum ot . Oviduct . P • Penis ss . Sperm-sac Im . . Longitudinal muscle ttn . Transverse muscle . fjn . . Ganglion c . .Eye . a . Dorsal antenna a' . . Lateral antenna ts . . Tactile style . 11 . Nerve thread . manubrium \Nutritive system, uncus incus fm ramus fulcrum !• Secreting system, h Vascular system. Eeproductive system. [ Muscular system. -Nervous system. EXPLANATION OF PLATE A. %* In all cases througliaut the plates, both the plain and coloured, the figures tJiat have the initial H attached to them liave been drawn from life by Dr. Hiidson, and those with the initial G, also from life, by Mr. Gosse. BRACHIONUS RUBENS and the details of its structure. 1. Dorsal view . . . . . H 2. Ventral view ; upper half . . . H 8. Lorica and muscles ; dorsal view . H 4. „ „ ventral view . H 5. Side view ...... II G. Mastax and trophi ; dorsal view . . G 7. Trophi ; ventral view G 8. Malleus expanded by pressure . . G 9. Malleus, as in nature . . . G 10. Incus, side view G 11. Ganglion and eye . . . . H 12. Extremity of foot . . . . H 18. Male; penis withdrawn (after Dr. Cohn) 14. Male ; penis extruded (after Dr. Colin) 15. Dorsal antenna . . . . H 1C. Eplnppial egg (after Dr. Colin) In all the above figures the letters signify as follows up , . . styligcrous prominence cl f . foot StJ t . toe 1'J cw ciliary wreath fff ¥ . buccal funnel Ic IP . lips vt mx . mastax cv ti . trophi ay ms . malleus 0 us . unc us om mm manubrium V is . incus ss rs . ranius Int fm fulcrum gn u: . a'sophagus c s . stomach a i . intestine a' r . rectum ts cloaca salivary gland gastric gland foot gland lateral canal vibratilc tag contractile vesicle ovary germ ovum penis sperm sac longitudinal muscle ganglion eye dorsal antenna lateral antenna tactile stvle PL. A EXPLANATION OF PLATE B. 1. Molicerta ringens ....... 2. Trochal disc of Melicerta ringens, from above 3. Limnias ceratophylli ...... 4. Trochal disc of Limnias ceratophylli, from above . 5. Rotifer or Philodina ...... 6. Synchaeta Baltica ....... 7. Eotifer rnacrurus 8. Brachionus Bakeri 9. Philodina or Rotifer ...... 10. Euchlanis ........ 11. Triarthra 12. Actinurus Neptunius 13. Megalotrocha 14. Stephanoceros Eichhornii ..... 15. 16. Floscularia ornata 17. Brachionns pala IB. Philodina or Rotifer ...... 19. Lacinularia socialis ...... 20. Stephanops cirratus ...... 21. Brachionns Bakeri ...... 22. Pterodina patina ....... 23. Salpina 24. Euchlanis ........ 25. Dinocharis pocillnm ...... 26. Mastigoccrca carinata ...... 27. Notommata tigris ....... 28. 29. Hydatina senta 30. Notommata longiseta 31, 32. Lacinularia socialis ...... 33. Notommata aurita ...... 34. Anursra striata j-Leuwenhoek, 1703-1705 Easier, 1759 [ Baker, 1753 Eichhorn. 1775 Joblot, 1718 Brady, 1755 -MtUler, 1776-1786 PL.B PLATE C. Showing the Coronee, Tropbi, and Feet of some of the principal Families of the BOTIFEEA.1 Flosculariadae »» Melicertadse Order I. EHIZOTA. Fig. I. Floscularia campanulata. „ Stephanoceros Eichhornii. Fig. II. Melicerta ringens. Philodinadae Adinetadse Order II. BDELLOIDA. Fig. III. Rotifer citrinus. Fig. IV. Adineta vaga. Asplanchnadse Synchaetadse Hydatinadse Euchlanidae Pterodinadse Brachionidse Order III. PLO1MA. Sub-order IL-LORICATA. Fig. V. Asplanclina Ebbesbornii. „ Synchaeta mordax. „ Hydatina senta. Sub- order LORICATA. Division I. Fig. VI. Exichlanis dcflexa. Division II. Fig. VI. Pterodina patina. Brachionus urccolaris. Order IV. SCIRTOPODA. Pedalionidse Fig. VII. Pcdalion mirum. 1 See pages 30, 31, 32. PL.C ctu.-. Ciliary urrea£fv pur... principal wreath/ -. Secondary wreath PLATE I. *#* In all cases throughout the plates, where the sex is not mentioned, the animal is a female. 1. Floscularia carnpanulata side view H la. „ „ ..... obliquely ventral . . . H 16. „ .... male, same scale as 1 . . . H lc. „ „ .... male, enlarged H Id. „ „ .... female, trochal disc from above . . H 2. Floscularia ambigua side view H 2a. „ „ obliquely ventral . H 3. Floscularia algicola side view H 4. Floscularia longicaudata dorsal view H 5. Floscularia coronetta side view . . . . . H 5a. „ „ (after Cubitt) . . . trochal disc, from above 6. Floscularia Cyclops side view H 7. Floscularia cornuta side view H 8. Floscularia regalis ...... side view . . . . . H 8«. „ „ ...... dorsal view . . . . . H 9. Floscularia ornata obliquely dorsal . . . . H 9a. ,, ,, trophi side view G Qb. ,, ,, ...... obliquely from above . . G 9c. ,, „ young female just hatched . G 9(Z. the same a little older . . ....... G im^w^^^ --^ , •^H^ ! . ,; -^ ' • •-•>'• %r" \.>«- '- Q t~ P < < ^ O P£ 0° fc U. O cr> O UJ O o r*-> M S ^ < PLATE II. 1. la. 16. 2. 3. 4. G. Off,. Floscularia algicola Floscularia coronetta Floscularia campanulata (? ambigua) Diagram of a Floscule Floscularia Iloodii .... „ „ Floscularia trilobata .... a cluster. .... . H H dorsal view of head . H dorsal view of head . H diagram after Dr. Moxon . H side view .... . H ventral view .... . H dorsal view .... . H ventral view .... H y •/.' w LO o o OJ 0 3 < 5 |s e& G- " ^ S ^ < < r ^ 'u~ ^ ^ to ^ fj Q «2 ^ < - ^ E- O ^ *-: o &N gtr •'-- t^ o to o a: • < ra o w ^ < PLATE III. 1. Floscularia mira . . seen obliquely from above . . . . H 2. Floscularia mutabilis . side view ; swimming . . . . .II 2a. „ ,, . . side view ; at rest ...... H 26. „ „ young female, ventral view . . . . H 2c. ,, .... young male, side view . . . . . H 3. Floscularia calva . . side view H 3a. „ ,, young female, dorsal view (after Mr. J. Hood) . H yi. ,. ,, . . young male, side view (after Mr. J. Hood) ?ll '^Mff^^^jf^M^lK^^^M m$m mi$^ ' >iw!$K •'•-• ^sS Arw:i''f : >^:^\\'Kv--^>-¥^ d!|^;iii:rs^;;>^:^N?^ §• ' i \ N M A m f— < t- w u, o SO 10 QQ * <; < s ^'A-^JCSftC^ PLATE IV. 1. Stephanoccros Eichhornii, side view ........ H 8. Young female just hatched, adhering to the glass cover head downwards . G 9 & 10. the same a little later G 11, 12, & 13. 3, 8, and 11 hours after being hatched G 14 & 15. 18 and 30 hours after being hatched ....... G Details of Stc2)hanoceros Eichhornii. 2. Head, dorsal view ............ H 3. „ ventral view ........... H 4. „ side view ............ II 5. „ seen dorsally in the plane passing through the knobs of the ciliary wreath ............ H 6. Trophi H 7. Ovary (after Dr. Leydig) 16. A budding tentacle (after Mr. Rossiter) 17. The same a little more developed (after Mr. Rossiter) e z; < Efi PLATE V. 1. Melicerta ringens . . . ventral view H la,. ,, ,, ... side view II 16. „ ,, ... partially retracted, side view . . . H lc. „ ,, ... upper part of 16, enlarged . . . H Id. „ ,, ... young female, side view . . . . H le. „ „ ... cluster H If. „ „ ... trophi H 2. Melicerta conifera . . . ventral view H 2a. „ „ ... dorsal view . . . . . . H 26. „ „ ... side view H 2c. „ ,, ... details of head, ventral view . . . H 2d. „ ,, ... details of body, ventral view . . . H 2e. „ ,, ... longitudinal muscles to chin and neck . H 8. Melicerta tubicolaria . . ventral view ...... H 3a. „ „ ... side view H 36. „ „ ... young female, dorsal view . . . H 3c. „ „ ... young male, side view . . . . H 4. Diagram of a Melicerta H R. t- tO W <; o S C3 P O O I— « PQ to u, o s CO CO w o CHAPTEE V. FLOSCULARIAM. Difficultates, quibus laborat investigatio animalculorum microscopicorum, in- numerae ; eorandemque certa et distincta determinatio tantum temporis, tantum oculorum judiciique acumen, tantamque animi compositi et patientissirai praesentiam requirunt, ut vix aliud supra. Nihil facilitis quain aniinalcnla videre, eornmque motu et ludo delectari ; differentias vero in bestiolis simplicissimis, mobilissimis, mutabilibus, in area minimi campi conspectum omni memento effugientibus, per- cipere, perceptas variosque cujusvia motus verbis significantibus exprimere, hie labor, hoc opus. Hinc srcpe post lucubrationem plurium horarum, cum videre et rnirari lassus essem, defectu tameii verborum insolitos motus et imagines exprimentium, metu- que, ne quse ipse quidem oculo et mente percepi lectori obscura manerent, chai'tic nihil conimisi. — O. F. MULLER. CHAPTER V. Order I. EHIZOTA. Fixed when adult, usually inhabiting a gelatinous tube excreted from the skin; foot transversely wrinkled, not retractile within the body, ending in an adhesive disk or cup. Family I. FLOSCULAEIAD^E. Corona produced longitudinally into setigerous lobes ; buccal orifice central ; ciliary wreath a single half circle above the buccal orifice ; trophi uncinate. This family, like the one that follows it, contains some of the largest, handsomest, and most attractive of the Eotifera. It consists of only two genera, Floscularia, and Stephanoceros, which closely resemble each other in their habits and internal structure, but differ considerably in outward form. The latter genus, which is represented by only a single species, has its frontal lobes produced into long arms, having setae set round them in whorls ; while the former, which contains no fewer than sixteen species, has the lobes comparatively short and expanded, with the setae radiating from their summits, and frequently edging the whole rim of the corona. The two genera differ also in the kind of tubes that they secrete. The majority of the Floscules have some- what irregular tubes of slight consistency ; but Stephanoceros has a thicker tube, more regular in shape, and apparently of greater density. Both genera are to be found adhering to the common water-plants, frequently in the axils of the leaves ; though some species prefer more exposed positions on the stems, or on the leaves themselves. The Flosculariadce bear captivity fairly well, and may be easily bred, provided that they have a good- sized trough and a plentiful supply of food ; for they are greedy feeders. They live mainly on Monads, ciliated Protozoa, &c. ; but occasionally capture and swallow comparatively large animals, such as Stentors, or even free swimming Rotifera. In one respect, however, captivity often tells on them ; for the home-bred specimens, though healthy enough, and breeding freely, are frequently much inferior in size to those that are brought fresh from their native haunts. Genus FLOSCULAEIA, Oken. GENERIC CHARACTERS. — Frontal lobes short, expanded, or wholly wanting ; setae very long and radiating, or short and cilia-like; foot terminated by a non-retractile peduncle, ending in an adhesive disk. Neither pen nor pencil can do justice to the beauty of these animated flowers. It can only be properly appreciated when they are seen by dark-field illumination under the microscope. Then the eye is at once delighted with the filmy transparency of the petaloid head, with the flowing curves of the lobes, and with the pencils of delicate setas radiating from their summits in all directions, and often passing altogether out of the field of view. Should, moreover, the species under observation happen to be a social one, such as F. campanulata, four or five specimens may often be found with their 44 THE ROTIFERA. tubes in juxtaposition, and the whole group can then be well shown under a low power; the animals, in various positions and under different aspects, forming, with their delicate cases and interlacing setaa, a picture that can be hardly rivalled. F. campanulata, when fully expanded, has been compared to " a long tubular flower with a five-angled petal, the tube swollen, contracted below the lip, and seated on the end of a long stalk." ! This description applies very well in most respects to the other species, except that the number of petals is not always five ; for, owing to late discoveries, there is now a regular series of Floscules with seven, five, three, and two lobes ; and one species in which the corona is not divided into lobes at all. The seta? also, which crown the lobes, and are so highly characteristic of the better- known forms, vary quite as much in the newer species as do the coronas ; in some exceeding the Eotiferon's utmost length, and in others diminishing almost to the size of ordinary cilia. Indeed, if the strange genus Acyclus — which must be very closely allied to the Floscules — be also taken into account, as well as the equally curious genus Apfsilus, there is a tolerably complete series of forms showing a gradual change from a Floscule, with seven lobes, and long radiating setaa, to a Floscule-like Rotiferon in which the setaa have entirely vanished, the corona has degenerated into a very delicate protrusile cup, and even the foot itself has shrunk into a mere sucking disk. The Tube. —The Floscules inhabit a semi-transparent gelatinous tube, into which the animal when alarmed can contract itself with great swiftness. It is secreted by the creature itself, and moulded on its own body by its sudden contractions, and slow expansions. When free from diatoms and extraneous particles (which is seldom the case), it is difficult to be seen, especially by transmitted light : under the dark-field illumination, not only can its outline be seen, but the substance of which it is composed can be traced from the outer surface, far in towards the Rotiferon itself. The tube becomes thinner towards the top, and it is often difficult to trace it there ; but it will generally be found to close in neatly a little under the neck.2 A Floscule, emerging from its tube, after one of its contractions, presents the appear- ance of a pear-shaped body on a transversely wrinkled stalk, with a pencil of long parallel hairs rising from the puckered centre of the rounded upper end. It slowly stretches itself till the wrinkles of the foot have nearly disappeared ; and then, after a delay, sometimes provokingly long, the puckers round the setae relax, and the whole pencil is thrust forward, by the unfolding of the lobes of the corona ; which, as they rise, show that they had been drawn down into the body by inversion, as the tip of the finger of a glove may be drawn into it, by pulling it from within. After a little further hesitation, the lobes unfold, and expand into a wide cup, while " the setae seem to fall round it on all sides in a graceful shower." The now fully expanded Floscule consists of five well- marked portions; the corona, the vestibule, the crop, the trunk (including the viscera), and the foot. The corona is a delicate nearly hemispherical cup, whose free edge is cut into lobes varying much in si^e, shape, and number. There arc two main varieties of lobe ; in the one they are narrow, pointed, and ending in a spherical knob; in the other they are broad, bounded by low convex curves, and knobless. In almost all, the dorsal lobe is conspicuous by its greater size, or peculiar appendages. The corona is furled by the action of delicate muscular threads imbedded in its surface, and expanded by the upward rush of fluid between its outer and inner integuments, due to the contraction of the transverse muscles of the trunk. The setae are set either on the knobs that crown the summits of the icbes, or on a thick rim running round them ; and they sometimes form a continuous fringe on the 1 Gosse, Popular Set. Rev. vol. i. 18C2, p. 1GO. - Though the tube is of the flimsiest material, and lighter than water, it certainly protects the animal from its enemies. I have watched a sharp-jawed larva trying to bite through a Floscule's tube, and it was as completely foiled by its swaying about from side to side at every touch, as a boy at Hallow E'en is bullied by a floating apple, when trying to seize it with his teeth. FLOSCULARIAD/E. 45 rim of the corona. Owing to their great delicacy, and to their lying in different planes, it is impossible to see nearly all of them at once. They vary greatly in size, position and arrangement ; but their varieties, with those of the forms of the lobes, will be described in the account of each species. Volvocina, small Infusoria, and floating particles may constantly be seen to enter the bell-shaped corona, and to pass thence down towards the buccal orifice. The setae take no part in this process, beyond that of preventing the return of the captured prey, by interlacing in a close network over the top of the cup, or by individually lashing at a returning object, so as to throw it back again into the gulf. The interlacing of the seta? is accomplished by the heads of the lobes approaching each other, and, should the prey be large and vigorous, the lobes are pressed tightly together, so as to completely bar all chance of escape. In most of the species, the motion of the setae appears due to the fitful action of the cuticle, on which they are placed ; but in F. trilobata, F. Iloodii, and notably in F. mutabilis, a regular cilia-like motion occurs in the setae ; while in F. mira there is a still wider departure from the ordinary type ; since in this Floscule each seta has a constant, slow, independent, amoeboid motion. The Vestibule. — At the bottom of the corona is a second chamber (the vestibule), bounded above by a highly contractile collar, below by a diaphragm with a slit in its centre (the buccal orifice), and 011 the sides by thick walls. On the upper margin of the collar, and running half round it on the ventral side, is a horse-shoe-shaped ciliated rim, ending in two knobs, bearing long, slowly moving cilia ; and this rim is so set, that it slopes downwards from the dorsal side to the ventral. This true rotatory apparatus may be easily seen in the large Floscules F. Hoodii and F. trilobata.1 A current, due to the action of these cilia, sets down the coronal cup, in a plane at right angles to its base, and carries the food, past the collar, into the vestibule. When once an organism has reached the vestibule, there is no return for it to the upper world. The Floscule often suffers two or three small Infusoria &c. to wander about round the walls of the vestibule ; but at any attempt to pass the collar, that at once contracts on itself, and closes the passage. In the diaphragm, which is the base of the vestibule, there is a long slit, the buccal orifice, bounded by two chitinous lips (PI. I. fig. Id, Ip), from which there hangs into the next chamber, called the " crop," an elastic tube (PI. II. fig. 4, t), which may be seen always undulating above the mastax. When there are victims enough collected in the vestibule to make it worth while to swallow them, the collar contracts violently, the lips dart forward with a sort of snap, and the prey is forced down the tube into the crop. It is evident that this hanging tube is an admir- able contrivance for admitting fresh prey into the crop, while at the same time pre- venting the return of that previously captured. Naturalists plagiarise from the Floscules, when they drop their live specimens through a quill stuck into the cork of a bottle ; only the rigid quill is far inferior to the flexible and ever-moving tube. The crop (PL II. fig. 4, cp) is a rounded chamber just under the diaphragm at the base of the vestibule. It has very thick walls, which are strengthened externally by two granular spots one on either side of the Floscule's shoulders (PL I. figs. 4, 8a). Viewed as opaque objects they are white, like the similar oval knobs on MegalotrocJia alboflavicans. Under the action of small muscular fibres, the sides of the crop contract alternately, and throw the contained food from side to side ; by which means every part of it in turn is subjected to the action of the jaws (PL II. fig. 4, ti). These lie at the bottom of the 1 Dr. Dobie described the two ciliated knobs in F. campanula ta, Ann. Nat. Hist. 1849. Mr. Gosse inferred the existence and position of the true rotatory organ from the motion of particles in the coronal cup (Tenby, p. 307). Dr. Moxon says that " the alimentary canal above the gizzard is divided by a highly irritable cilium-clothed sphincter of irregular outline " (Trans. Linn. Soc. vol. xxiv. 186-1, p. 457). In 1867 I published a full description of the rotatory organ of F. campamilata in the Trans. Bristol Micr. Soc. In 1869 Mr. Cubitt did the same for Stephanoceros and F. coronet ta (Hon. Micr. J. vol. ii. 1869, p. 133). 46 THE EOTIFERA. crop, attached to the walls of the stomach ; and to reach the latter everything must pass hetweeii them. The Floscules are great feeders, and sometimes the crop becomes so distended with food, that the animal, unable to force it past the jaws quickly enough, seeks relief by expelling the contents of the crop right through the inverted tube. The lobes of the corona are folded back on the body, the diaphragm is pushed upwards and the tube is thrust inside out through the slit in the diaphragm ; while through it pours the unmanageable food. Dr. Moxon (loc. cit.) has seen this take place on two separate occasions, and I have seen it once : in each case the animal was F. campanulata.1 The appearances due to the tube have been variously described as caused by moving filaments, lamina?, vibratile cilia, and a waving membrane ; but these observations of Dr. Moxon, confirmed by my own, put the matter beyond doubt. The Trunk. — The outer wall of the trunk is a tough, elastic, and often shining cuticle, which has an inner and softer layer of varying thickness. This double covering interferes greatly with a clear view of the viscera, especially as it has intercommunicating cavities and channels containing fluid, which is driven upwards and downwards by the contraction of the muscles, and by the various motions of the body. Nor is this all ; for the fluid itself is often rendered semiopaque by granules floating in it. It is doubtless by means of this fluid that the lobes of the furled corona are pushed forward and expanded, the transverse muscles of the trunk forcing it into definite channels, which are thus rendered tight and stiff, like the ribs of an umbrella. Mr. Gosse 2 has described and figured these in the case of F. ornata ; and has noticed how the granules flow from the trunk over the neck into the various channels of the coronal cup. Mr. Hood, too, has watched a steady stream of granules passing down from the trunk into the foot, and returning again from a point about half way from its extremity. The granules may be frequently seen, in some degree, in specimens of almost every species, but occasionally they are in such abundance as to render the animal quite opaque ; and, by reflected light, of a dead white. The foot is very long and flexible, and is capable of great expansion and contraction, but cannot be drawn into the trunk. It consists of little else but muscles. The great longitudinal muscles pass down its whole length, and numerous fine muscular fibres encircle it everywhere, covering it with transverse rings of very variable thickness, from its junction with the trunk to its extremity. This latter contracts to an inextensible, and usually short cord or peduncle, which itself terminates in a sort of disk. In the foot are also the two club-shaped glands (so common in other genera) which probably secrete a viscous fluid for fastening the disk to some extraneous object. The Nutritive System. — The food is feebly pecked at by the jaws, while it is in the crop, but it evidently undergoes there some process of digestion. I once saw a Floscule bolt a small Salplna. When inside the crop it was still alive, and it charged from side to side, in the vain hope of escape. The sharp points of its lorica ought to have made its captor uncomfortable ; but the only result was, that its outline gradually grew dim, and that before long the whole animal faded into a shapeless mass. Mr. Gosse3 notices the absence of the mastax, and says of the trophi that " the jaws consist of a pair of curved unjointed but free mallei, with a membranous process beneath each. Each malleus (PL I. figs. 9a, 95) is an uncus of two slender arched divergent fingers, united by a subtle web ; the back of each curves downwards, where, expanding and becoming membranous, it is connected with some delicate but definite processes with rounded outlines, which I should have supposed to be muscular bulbs, but that they remain after treatment with potash." After passing between the jaws the food enters the stomach (PL II. fig. 4, s) appar- ently directly, as no oesophagus is visible. The alimentary canal is divided into stomach (s) intestine (i), and rectum (r), which latter is bent on itself, and ascends to the cloaca (cl) 1 Fig. 1, PI. I>, is a copy of Dr. Moxon's drawing, showing the tube turned inside out and pro- truded through the slit in the diaphragm : the lobes (I) are shown drawn down close to the body. 2 Popular /b'cj. Ecv. vol. i. 1802, p. 1GO, pi. ix. 3 Phil. Trails. 1856, p. -119. FLOSCULARIADjE. 47 on the dorsal surface. A partial separation (probably due to a sphincter muscle) is usually visible between the stomach and intestine, and the distinction is often made more obvious by the different colour of the contained food. The whole alimentary tract is richly ciliated ; and so is the tube (t) that hangs down into the crop. The cilia can be easily seen in the intestine, when it is partially empty ; and on the tube, when it is everted by the Floscule's disgorging its crop. Secreting System.— In the great majority of Eotifera there are two gastric glands, seated on the top of the stomach. I have, however, never been able to detect them in any Floscule, neither has anyone recorded their existence except Ehrenberg and Grenadier. The latter describes and figures them of unusual length in F. campanulata,1 but it is probable that in this matter he is mistaken. He also describes two club-shaped glands in the foot. Vascular System. — It is very difficult to trace this in most of the Floscules, owing to the optical difficulties due to their skins. But in 1804 Dr. Moxon (loc. cit.) published a complete account of it in F. campanulata. His figure of the contractile vesicle (PI. II. fig. 8, cv), the lateral canals (Ic), and of four of the vibratile tags (vt), is so clear as to render any verbal description unnecessary. A fifth vibratile tag was discovered by Grenadier in F. campanulata (loc. cit.), in the side of the coronal cup, near the spot where Dr. Moxon (loc. cit.) had anticipated that it would, some day, be found. Parts of this system have been seen in several other species, and doubtless it exists in all.2 The muscles consist of non-striated fibres. Below the bottom of each depression, between the lobes, a muscle runs downwards in the substance of the coronal cup and vestibule, and is lost on the surface of the body, to reappear again, as it nears and passes down the foot. The anterior portions of these muscles end in two or more branches which diverge to the thickened rim of the coronal cup, and often interlace, as seen in F. coronetta (PI. II. fig. 2), and in F. trilobata (PI. II. fig. G), where they may be seen reaching the summit of the dorsal lobe. There are some half-dozen transverse muscles imbedded in the integument of the trunk ; and the walls of the vestibule, with its upper ciliated rim, are all highly contractile. The Nervous System. — Dr. Moxon (loc. cit.) has described and figured the nervous ganglion in F. campanulata, and I have seen it in F.Hoodii. It is in nearly the same position as it is in Steplianoccros, namely, on the dorsal side of the vestibule ; and is, as usual, close to the organ of taste, and not far from the eyes and dorsal antenna ; to all of which doubtless it sends out nervous threads. Dr. Moxon has seen and figured such threads in F. campanulata (PI. II. fig. 8, n.) In the great majority of the Ilotifera the mastax is also not far from the nervous ganglion ; but in the Flosculariadcs the mastax almost vanishes, while the jaws and ganglion are far apart. Organs of Sense. — Two red eye-spots lie usually above the ganglion ; but, as in the adults they are deeply imbedded in the integument, they are not easily seen. In fact the ordinary way of attempting to see them, by transmitted light, will scarcely ever be successful ; but by treating the Botiferon as an opaque object, and concentrating a strong light on it, the eyes may often be seen glowing like rubies when all else is invisible.3 The eyes are conspicuous in the half-grown animals, and in the young within 1 Sieb. u. Koll. Zeits. Bd. xix. 18G9, p. 483. 2 Dr. Leydig saw the contractile vesicle in F. cornuta ; Ueb. d. Ban d. Rciderth. 1854. Dr. Bartsch has seen the contractile vesicle, lateral canals, and vibratile tags in his F. longilobata (F. coronetta) Rot. Hungaricc, 1877. 3 Mr. Gosse (Popular Sci. Rev. vol. i. 1862, p. ICG) observed one eye at a time in F. cornuta. Mr. Cubitt (Mon. Micr. J. vol. iii. 1870, p. 245) saw the eyes in F. coronetta ; and I have recorded (J. Roy. Micr. Soc. 2 Ser. vol. iii. 1883, p. 163) my having observed them in F. Hoodii and F. regalis. Herr K. Eckstein (Sieb. u. Koll. Zeits. Bd. xxxix. 1883, p. 347), unaware of the above observations, says "the Floscules have been hitherto regarded as eyeless," and records his having seen the eyes in F. cornuta. 48 THE KOTIFERA. the egg. There are three antennae in F. campanulata, F. coronetta, and jP. Iloodii; and no doubt the same three may with care be found in the other species. There is one on each side of the neck, and one on the median line near the middle of the dorsal lobe. The two lateral antenna are very short tube-like prominences each carrying a brush of divergent sette ; they are very apparent in F. coronetta (PL II. fig. 2) : the dorsal antenna is a mere setigerous pimple.1 The setae on the lobes act also as organs of touch, warning the creature of the approach of anything detrimental to its delicate cup ; and whipping back into it any animalcule that endeavours to escape from it. On the middle of the contractile collar, which is above the vestibule, and on the dorsal side of it, there is a round projection facing the concavity of the ciliary wreath. It can be easily seen in F. coronetta and F. Hoodii, and is probably an organ of taste, as it is constantly thrust forward to meet any particle which is passing into the vestibule. A very obvious and tongue-like organ holds a similar position, and acts in a similar way, in Stephanoceros. The Reproductive System. — The ovary, with its clear spherical germs, and frequently with an opaque egg in it, can be seen filling the greater part of the space between the stomach and the ventral surface. No other portion of the apparatus has been made out, owing no doubt to a habit that the Floscules have of contracting themselves sharply into their tube when about to lay an egg. When laid, the eggs are ranged above one another, between the foot and the tube. The ordinary number of female eggs is from two to five ; though as many as eight or ten have been seen at once. The male eggs, which are smaller, rounder, and more numerous, than the female, frequently amount to as many as nine or ten, and have occasionally been seen in a cluster of eighteen or twenty in the same tube. Both are inclosed in a delicate shell, which is left behind in the tube, when the young Floscule is hatched. Dr. Weisse and Mr. Hood agree in assigning six or seven days as the time from the extrusion of the egg to the birth of the young animal. The Young Female. — "The infant female Floscule is a white cylindrical maggot (PI. I. fig. 9c), blunt at the front end, with a central orifice, whence protrudes a short brush of cilia ; but the margins are capable of unfolding, when the cilia are seen to form a whorl around the truncate summit, swiftly rotating. The margin soon begins to bud forth the little knobs around which the cilia are gathered (PL I. fig. Qd) ; these quickly increase in length, and the angular flower-like corona gradually forms. Meanwhile the little creature, which was at first free, attaches itself by its hinder end, and assumes the condition as well as the form of the parent." : Mr. Hood has observed in F. calva, that the young animal fixes itself two or three hours after it has burst its shell, and soon begins to form its tube, which at first rises barely to half the height of the foot. By the time it is three days old (PL III. fig. 3a) the tube has attained fair proportions. The same observer noticed that the lobes of the young F. ambigua began to develop from a collar under the ciliary wreath, and were at first merely a dorsal and ventral lobe ; the latter with a small notch. In three or four days the notch deepened and widened so that there were three lobes ; but it was not till the fifth or sixth day that the rudiments of the small side lobes (the fourth and fifth) made their appearance. The young Floscule arrived at maturity at the twenty-fourth or twenty-sixth day, but continued to increase in size after it had deposited eggs : in fact, did not cease to grow till shortly before its death. The whole lifetime, in a trough, was from forty to forty-six days. Captivity, however, affected the growth of the animals, even when carefully attended to, and plentifully supplied with food. On one occasion, for instance, a large F. campanulata ,\s inch long, from one of the Scotch lochs, was placed in a tank ; and 1 Dr. Moxon (loc. cit.) first called attention to their existence in the Floscules. Herr Grenadier (loc. cit.) mentions his having discovered the median antenna in F. proboscidea (F. campanulata) ; but was unaware that Dr. Moxon had seen and described all three, five years before. '-' Mr. Gosse on F. campanulata. Popular Sci. Rev. vol. i. 1862, p. 166. FLOSCULARIA D.E. 49 the young reared from its eggs, though perfectly healthy and breeding freely, never exceeded -JT. inch in length : their eggs, too, were half the size of those of their parent. In F. trilobata, occasionally, the egg produces the living young in the body of the parent. Mr. Hood has seen the embryo alive in the egg, within the Floscule, and has witnessed its birth : yet Dr. Collins has seen the same Rotiferon deposit the usual eggs in its tube. The Male.— Until 1874 no male had been discovered among the Rhizota ; l and indeed some observers supposed this group to be monoecious ; but in that year I had the good fortune to find the male of Lacinularia socialis, and to study it thoroughly.2 Soon afterwards I found that of Floscularia campanulata (PL I. fig. Ic) and \ have since seen what I believe to be the male of F. mutabilis (PL III. fig. 2c). Mr. Hood has observed and figured the male of F. calva (PL III. fig. 36), and has seen that of F. ambigua actually hatched. The structure of the male Floscule has not yet been thoroughly investigated ; but, so far as it has been studied, it has been found to agree with that of other male Rotifera. The corona is an imperforate many-lobed cushion, surrounded by a simple circlet of long cilia. The nutritive system is wholly absent. Two red eyes are visible just under the surface of the corona ; and the longitudinal muscles, for withdrawing the head, are generally obvious. Nearly the whole of the body-cavity is filled with a large sperm-sac (PL I. fig. Ic, and PL IIL fig. 3b ; ss) from which the penis (p), a ciliated protrusile tube, proceeds to the dorsal surface, at the junction of the trunk and foot. The vascular system, ganglion, and antennae have not yet been seen ; but no doubt they are present, as in the males of other Rotifera. F. REGALIS, Hudson, (PL I. fig. 8.) Floscularia regalis , . Hudson, J. Roy. Micr, Soc, 2 Ser. vol. iii. 1883, p. 106, pi. iv. fig. 3. > SP. CH. Lobes seven, knobbed. The corona is a deep cup with a nearly circular rim, from which project four knobbed triangular processes on the ventral side, dividing that half of the rim into three equal spaces. The processes curve slightly outwards ; and, at the rim, their bases unite, so as to give that edge of the cup a semi-hexagonal appearance. In the middle of the dorsal side of the rim rises a large triangular knobbed lobe, bearing on each side a short recurved knobbed process. All seven knobs carry pencils of long radiating sets. A true ciliary wreath at the bottom of the trochal cup, and two red eyes, can be easily seen. This remarkable Rotiferon, the only seven-lobed species, was found by Mr. Thos. Bolton in September 1882, near Birmingham. Length, -^ to ^ inch.3 Habitat. Lakes and clear ponds. Birmingham (T.B.4); Perth (J.H.,4 and W. Dingwall) : not common. F. CORONETTA, Cubltt. (PL I. fig. 5 ; PL II. fig. 2.) Floscularia coronetta . . . Cubitt, Mon. Micr. J. vol. ii. 18G9, p. 133, pi. xxv. Stephanoceros Horatii . , . Cubitt, Mon. Micr. J, vol. vi. 1871, p. 106. Floscularia longilobata . . . Bartsch, Eot. Hungarian, 1877, p. 24, ii. Tab. fig. 11. 1 Mr. Gosse (loc. cit. p. 487) described some probably male eggs in M. ringcns. 2 Mon. Micr. J. vol. xiii. 1875, p. 45. 3 As the Rotifera vary from ^ to ~s inch, no attempt has been made to draw them to a fixed scale. The actual length of each species will be given at the end of its description. 4 Throughout the work the following initials will be used in the Habitat : — J. H. = Mr. John Hood ; T. B. = Mr. Thos. Bolton ; P. H, G. » Mr. Gosse ; C. T. H. = Dr. Hudson. F 50 THE ROTIFERA. SP. CH. Lobes five, linear, knobbed ; setae, non-extensile. The corona has five long narrow knobbed lobes, nearly all of equal length, sepa- rated by deep depressions, and forming a miniature coronet. The dorsal lobe is slightly the longest, and the lobes are so set on the front of the body that a plane touching the knobs would be oblique to its longitudinal axis ; the dorsal lobe being the furthest forward. All the knobs carry long radiating setae, and the setae are continued all along the edge of the trochal cup (PI. II. fig. 2). The true ciliary wreath and the eyes have been seen in the adult by Cubitt (loc. cit.} ; but the former with difficulty. The lateral antennae can be readily seen when the animal is favourably placed, as well as the delicate muscular threads by which the longitudinal muscles act on the corona (PL II. fig. 2). As many as seven male eggs have been seen in one tube. Length, ^ inch. Habitat. In ponds and marsh pools ; rare. Wandsworth Common (Cubitt) ; Forfar, Fife (J.H.). F. MIRA, Hudson. (PI. III. fig. 1.) Floscitlaria mira .... Hudson, J. Roy. Micr. Soc. 2 Ser. vol. v. 1885, p. G09. SP. CH. Lobes fw e, linear, knobbed ; setae extensile. The corona is very like that of F. ornata, which species the Rotiferon closely resembles in every respect but two. First, the tube is much more like that of a Stephanoceros than that of an ordinary Floscule. I have seen only one specimen, but Mr. Cocks (its dis- coverer) tells me that the tubes of the half-dozen specimens which he has seen were all of the same sort. Secondly, in its setae F. mira is not only unlike all other Floscules, but is unique among the Rotifera : for each seta is in constant independent motion, slowly extending or contracting like the pseudopodium of an Amoeba. When the retracted seta begins to extend again, it is often bent into a whip-like shape, a wave of motion overtaking, as it were, the resting anterior portion, and finally driving out the latter with a characteristic flourish of its tip. The setoa are of amazing length and abundance, exceeding the total length of the Rotiferon. This very rare and wonderful creature was discovered by Mr. W. G. Cocks in June 1884. Length, r^ inch. Habitat. Unknown ; found in an aquarium, in water that probably came from Epping Forest or Walton-on-Thames (W. G. Cocks). F. OKNATA, Ehrenberg. (PI. I. fig. 9.) Floscularia ornata. — Ehrenberg, Die Infus. 1838, p. 408, Taf. xlvi. fig. 2. Peltier, Ann. Sci. Nat. Zool. 2 Ser. t. x. 1838, p. 41, pi. iv. Dujardin, Hist. Nat. Zooph. 1841, p. G10, pi. xix. fig. 7. „ Gosse, Tenby, 1856, p. 307, pi. xx. „ Popular Sci. Rev. vol. i. 1862, p. 160, pi. ix. figs. 1-3. „ Pritchard, Infusoria, 1861, p. 675, pi. xxxii. figs. 384, 385, and xl. figs. 25, 26. Weisse, Sieb. u. K, Jan. 1872, p. 0, with fig. Floscularia trifolium . . . Hudson, J. Eoy. Micr. Soc. 2 Ser. vol. i. 1881, p. 4, pi. ii. SP. CH. Lobes three, large, broadly curved, separated by very deep and similarly curved depressions : dorsal lobe rather the largest; setse forming a continuous double fringe round the entire circumference of the corona ; the outer row arranged like those of F. campanulata ; the inner row short, slightly curved, aud arranged like cilia. This large and elegant Floscule was discovered by Dr. F. Collins in 1865, in a small pool near Sandhurst, Berks ; and he published a short account of it, with a figure (loc. cit.} in 1872. It was afterwards found by Mr. J. Hood, in Loch Lundie, near Dundee, in 1880 ; and I published a description of it (loc. cit.} naming it F. trifolium, as I had considerable doubt of its really being Dr. Collins' species. I have since seen the description and figures which Dr. Collins sent, with some live specimens, to Mr. Gosse in 1865 ; and I have now no doubt that F. trifolium arid F. trilobata are the same. 1 Kindly identified by Dr. Cooke in a letter to Mr. Gosse. FLOSCULARIADJE. 55 The peculiarity of a second fringe of set® lying within the principal fringe is shared with it by F. Iloodii alone. Unlike the ordinary sets, these smaller secondary seise possess a kind of joint action; for when any captive creature tries to escape from the coronal cup, and to pass the setae, those of the outer row either lash separately at it, or are drawn together over it by the converging lobes ; while a wave of motion, like a ciliary wave, runs once or twice round the inner row. The true ciliary wreath can be easily seen at the bottom of the coronal cup, owing to this Floscule's great size and transparency. I saw a small contractile vesicle, but I had no opportunity of tracing the rest of the vascular system. The two lateral antennas were also obvious. The first thing that strikes the observer, on watching the protrusion of the furled head, is the great size of the Floscule, and the curiously shrivelled appearance that the lobes of the coronal cup present, as they emerge from the opening head. They look exactly as if the animal were sickly or injured. In a few seconds, however, they gently swell out, the many folds and creases disappear, till at last the eye is gratified with the sight of a lovely transparent tulip, of three petals, their edges all fringed with delicate and motionless hairs. It is a creature of exquisite beauty ; from every point of view the flowing curves of the cup are charming, and its great transparency permits the whole of the outline of the rim to be seen at once. The dorsal lobe is rather larger than the other two, and is curved forward over the cup. Across each lobe run delicate muscular threads for furling it, which are specially visible on the back of the dorsal lobe (PI. II. fig. 6). Dr. Collins saw eggs, laid by his specimens, remaining attached to them within the tube ; but Mr. Hood observed that some specimens, which he reared in a trough, hatched the ova in the ovary, and then gave birth to the living young. They are voracious feeders on Infusoria and small animalcules, and are fond of stationing themselves in the axils of water plants. Length, ^g to f5 inch. Habitat. Lochs, marsh and boggy pools; Sandhurst (Dr. Collins and P.H.G.); Woolston, Hants (P.H.G.) ; Fife, Forfar, Perth (J.H.) : not common in England, abundant in Perthshire. F. HOODII, Hudson. (PI. II. fig. 5.) Floscularia Hoodii . . Hudson, J. Roy. Micr. Soc. 2 Ser. vol. iii. 1883, p. 161, pi. iii. figs. 1, 2. SP. CH. Lobes three; dorsal lobe much the largest, and carrying two, large, sleeve- like, flexible processes ; setae, short, cilia-like, in two parallel rows, fringing the entire circumference of the coronal cup. This is one of the largest of all the Eotifera ; adult specimens being quite ^ inch from the top of the dorsal lobe to the extremity of the peduncle. Its great size, and its curiously shaped three-lobed corona make it sufficiently remarkable ; but, in addition to these peculiarities, it has two extraordinary processes, perched one on each side of the back of the dorsal lobe. They appear to be hollow, and to communicate with two lenticular spaces lying between the two surfaces of the dorsal lobe. Fine muscular threads pass along and across them (PI. II. 5b], and the animal can contract and expand each independently of the other ; and throw them into all kinds of positions. The upper end of each seems to be separated from the lower portion by a constriction, from which a muscular thread runs down to the base. Each of these processes slowly and independently changes its shape and position, now sinking down on the dorsal lobe so as to be invisible, or again bending its free end at right-angles to the lower portion. I have never seen anything like them on any other Eotiferon : they may possibly be organs of touch, but I could detect no trace of setre on them. Mr. Hood tells me that both in young and adult specimens he has seen brown granular matter discharged from their free ends. The thickened rim of the three lobes carries its double fringe of seta? 66 THE KOTIFERA. set just as they are in F. trilobata, the larger row stretching outwards, and the smaller inwards ; and the same rapid flicker may be seen on occasions to run all round the edge of the coronal cup. The orifice of the coronal cup alters constantly, now opening in the characteristic way shown in PI. II. fig. 5a, and now reduced to a slit ; or even closed in puckers. Two dorsal ridges, as in F. ambigua, run like buttresses from the body to the back of the dorsal lobe, and in the lowest portion of the deep hollow between these lie the two pale pink eyes ; both in the neck, and one close to each buttress. The true ciliary wreath is distinctly visible throughout its whole length. It is a long horse- shoe-shaped and ciliated ridge, sloping sharply down from the bottom of the coronal cup into the vestibule. The contractile vesicle is unusually large and distinct ; close to it, and apparently situated in it, is a cluster of yellow globules, which look black by transmitted light. This strange and beautiful Floscule was discovered by Mr. J. Hood in December 1882, in a ditch on Tent's Muir, Fifeshire. Length, -^ inch. Habitat. Marsh pools ; Fife (J.H.) : rare. F. CALVA, Hudson. (PL III. fig. 8.) Floscularia calva .... Hudson, J. Roy. Micr. Soc. 2 Scr. vol. v. 1885, p. 610. SP. CH. Lobes two, short ; dorsal lobe the larger ; setae very short, radiating from the thickened summits of the lobes, incapable of cilia-like action; body unusually long and narrow, its outline confluent with that of the coronal cup, so that there is no neck ; eyes cervical. Mr. J. Hood discovered this species in 1884 on a Sphagnum leaf, in a mossy pool on Tent's Muir, only twelve inches deep, and on another occasion found it in Loch Lundie at a depth of ten feet. I have seen only two specimens of it, and those under disadvan- tageous circumstances ; as each had dropped from the plant on which it was found, and was lying in the sediment at the bottom of the tube. The creature appears to attach itself rather to its tube than to the stem of the plant which bears the tube, and so to be easily detached. I am indebted to Mr. Hood for drawings of the young male and female (PL III. figs. 3a, 36), each of which he saw hatched from eggs laid in the tube. The male is about ^\ 0 inch in length, and resembles that of F. campanulata. Length, *. to ^ inch. Habitat. Lochs and marsh pools, on Myriophyllum and Sphagnum; Forfar, Fife (J.H.) : rare. F. MUTABILIS, Bolton. (PL III. fig. 2.) Floscularia mulabilis . . Hudson, J. Roy. Micr. Soc. 2 Scr. vol. v. 1885, p. 609, pi. xii. figs. 1-3. SP. CH. Lobes two, well developed; dorsal lobe decidedly the larger ; setse rather short, set round the ivhole circumference of the disk, and capable of cilia-like motion; eyes near the summit of the dorsal lobe. F. mutabilis somewhat resembles F. calva, but is at once distinguished by its larger lobes, moveable setae, and by its unique habit of swimming. The animal has not as yet been found attached to any water plant. It looks, when resting in its case at the bottom of a live cell, just like an ordinary Floscule that had been knocked off its perch, as the seta? are straight and motionless. After a short rest it pulls down the two lobes to a level with the bottom of the depressions between them, and so alters the corona FLOSCULARIAD^E. 67 that it looks like that of an CEcistcs ; at the same instant the seta) l set up a vigorous cilia- like action ; and the animal, case and all, sails slowly, stern foremost, through the water. Two red eyes are very conspicuous in a most unusual position ; namely, near the top of the dorsal lobe. I have seen what I believe to to be the male (PL III. fig. 2c), but I failed to isolate it so as to make out its internal organs. Its length was about T|-ff inch. It appeared to have, in addition to the usual ciliary wreath, set* pointing backwards to the foot. Mr. T. Bolton discovered F. mutabilis in a pond of Button Park, near Birmingham, in May 1884. He described, named and figured it, soon afterwards, in one of the ily- leaves sent out with his specimen tubes. Length. About ^ inch. Habitat. A pond in Sutton Park, Birmingham (T.B.) : rare. F. EDENTATA, CollillS. (PL III. fig. 4.) Floscularia cdentata . . . Collins, Science Gossip, Jan. 1872, p. 9, with fig. ,, ,, ... Hudson, J. Roy. Micr. Soc. 2 Ser. vol. v. 1885, p. 611. Corona lobeless, transversely truncate ; setae very short, chiefly on the ventral and dorsal portions of the rim; body large and stout in proportion to the animal's total length, and nearly as long as tJic foot. Dr. Collins first discovered this ugly Floscule near Sandhurst in 1867. He says (loc. cit.) that it has no masticating organs, and that the food passes directly into a capacious stomach. As his specimen was a female (for it laid an egg while in captivity) this is very unlikely. My specimens were so gorged with food that no internal organs could be seen, except the stomach and a portion of the ovary. One of them was literally crammed full of specimens of Cocconema, which not only distended the real stomach and the crop, but even protruded above the rim of the coronal cup. How the animal contrived, with its feeble cilia, so to pack itself with these unmanageable diatoms, I cannot imagine. Length. My specimens, Js inch ; Dr. Collins', -^ inch. Habitat. Sandhurst, Berks (Dr. Collins) ; Woolston, Hants (P.H.G.) ; Blair Athol (W. Dingwall) : rare. Genus ACYCLUS, Leidy. GEN. CII. One dorsal, frontal lobe; setae absent, the coronal cup edged with a delicate festooned membrane; termination of foot truncate. * ACYCLUS INQUIETUS, Leidy.2 (PI. D. fig. 3.) Acyclus inquictus . . Leidy, Proc. Acad. Nat. Sci. Pa. 1882, p. 2-13, pi. ii. figs. 1-6. The structure of this species has been only imperfectly made out ; but, so far as it has been, the animal appears to be closely allied to the Floscularia ; and so also do the next two species, Apsilus lentifortnis, and Apsilus bipcra. The characters of Acyclus inquietus given by Prof. Leidy are as follows : — , " Body fusiform, tapering behind into a long narrow tail-like appendage, by which it is attached, not distinctly annulatcd, but becoming transversely ivrinkled in con- 1 It is possible that there may be (as Mr. Bolton says) a row of short cilia round the coronal cup, as well as the larger seta) ; but my impression is that there is not : I altered my opinion more than once, while watching the creature, but came at last to the conclusion that it swam by means of its setir, and not by a subsidiary row of cilia. - Throughout the work the species which are not known to be British will be marked with an asterisk. 58 THE EOTIFEEA. traction. A non-ciliated cup-like head prolonged into an incurved dig itiform appendage (as a substitute for the usual trochal disc), contractile and retractile." The Professor found eight specimens of this strange creature, each surrounded by a group of Megalotrocha alboflavicans , and all attached to the tubes of Plumatelladiffusa, in the Schuylkill river, U. S. It is considerably larger than M. alboflavicans, and can be readily distinguished with the naked eye, towering above the surrounding cluster of Megalotrocha " like a giant in a crowd." It is a very difficult animal to observe, as it bends abruptly in different directions ; suddenly contracting and slowly elongating, and scarcely ever for a moment remaining erect. It is translucent, whitish, with the thicker portion of the body of a yellow or brown hue, due to the colour of the alimentary canal. The corona is a cup prolonged on the dorsal side into an incurved lobe (PL D, fig. 3). It is capable of being expanded or contracted, protruded or retracted ; and when ex- panded, the dorsal lobe is also extended, but remains somewhat incurved. There are no cilia or setae on the edge of the cup or lobe, but both of them are bordered by a delicate festooned membrane. When contracted, the lobe is rolled up spirally (PI. D, fig. Qb). A narrow, transversely wrinkled neck lies between the cup and the body. No ciliary wreath has been noticed within the cup. There is generally no tube present ; but in two instances the animal has been seen in a " copious colourless gelatinous sheath." The cup converges into a pouch (the vestibule) occupying the neck, which is seen to expand and contract from time to time. Longitudinal muscles extend from the neck to the membrane surrounding the coronal cup, passing along its walls. Retractor muscles stretch from the body down the length of the foot. The secreting, vascular, and nervous systems have not been observed ; neither have any eyes or antennae in the adult female. The ovary is in the usual ventral position, and the ova are large, and unsegmented when extruded. Length, .,\ to ^V inch. Habitat. Schuylkill river, U. S. (Prof. Leidy) : rare. Genus APSILUS, Metschnikoff. GEN. CH. Coronal cup wholly membranous ; setae and foot absent. *Ar«JLUS LENTIFOKMIS, Nctschnikoff. (PI. D, fig. 4.) Dictyophora vorax (?) • • Leidy, Proc. Acad. Nat. Sci. Pa. 1857, p. 204, and 1882, p. 248, pi. ii. fig. 7. Apailus Icntiformis .... Metschnikof'f, Sicb. u. Koll. Zcits. Bd. xvi. 1866, p. 346, with figs. Cupelopagns bucincdax (?) . . Forbes, Amcr. Mon. Micr. J. 1882, pp. 102, 151, with fig. Apsilus vorax Foulke, Proc. Acad. Nat. Sci. Pa. 1884, p. 37 pi. i. figs. 2, 5. Apsilus Icntiformis .... Leidy, Proc. Acad. Nat. Sci. Pa. 1884, p. 50. Herr E.Metschnikoff found many specimens of this Rotiferon at Giessen in 1866, on the under side of the leaves of the yellow water-lily, to which they were attached by a chitinous ring on the ventral surface ; both in the young and adult female the foot was absent. The coronal cup is wholly membranous, and destitute of either cilia or setae. It is capable of having its edge all drawn close together into a point (PI. D, fig. 4a), and of being wholly withdrawn within the body, so that it acts as a net, closing over any prey that voluntarily enters it, and forcing it down into the chamber below it, which in the Floscules would be called the crop (fig. 4.a, cp}. At the bottom of the crop is a very peculiar set of trophi (fig. 4c). A broad stomach has a coecal appendage on each side, and a cloacal orifice on what appears to be the ventral side, but is really a portion of the dorsal, having been drawn round by the animal's curving its body when attached to the leaf. There are two pear-shaped glands attached by their narrow ends to the crop. There is a contractile vesicle opening into the cloaca, and from it, above, issues a FLOSCULARIAD^E. 59 duct which divides into two lateral canals. Each canal runs slantingly up to a coil at the side of the body below the cup, and thence sends a branch into the dorsal surface of the cup, anastomosing with its fellow above and below the nervous ganglion (PI. D, fig. 46), and bearing two vibratile tags on each side of it. The nervous ganglion is a four-side organ in the dorsal wall of the cup ; it sends out a nerve thread at each corner, the lower pair passing to two lateral antennae. No eyes are visible in the adult. The young embryo is developed in the egg in the body of the parent. When hatched, it is a free-swimming Eotiferon (PL D, fig. 4cZ), with a truncate, ciliated, anterior ex- tremity, and with the cloaca, at the ciliated posterior extremity, surrounded by a mem- branous ring. There are two red eyes, but the characteristic coronal cup is as yet undeveloped. The male, which has a ciliated foot, is so like those already described, that Herr Metsclmikoff s drawings supersede description (PI. D, fig. 4e). Prof. Leidy described in 1857 (loc. cit.) a new Eotiferon, " destitute of wheel- organs," which he named Dictyoplwra vorax. He obtained, however, some fresh specimens in 1884, and is now of opinion (loc. cit.} that the animal is identical with Apsilus lentiformis, and that the discrepancies between his account and Herr Metschni- koff's are due to the wrinkled condition of his first specimens, which had been forcibly removed from the glass sides of an aquarium. Mr. S. A. Forbes also described (loc. cit.} a Eotiferon found in a neglected aquarium, and "wholly destitute of cilia or other vibratile structure." He called it Cupelopagus bucinedax, and gave a very characteristic figure of its side view. I have little doubt that this also is Apsilus lentiformis. Length. Maximum about j,1,-,- inch. Habitat. On water plants, Giessen (Metsclmikoff) ; Fairmount Park, and Schuylkill river, U. S. (Leidy). * APSILUS BIPEIIA, Foiclkc. (PI. D, fig. 5.) Apsilus bipera .... Foulke, Proc. Acad. Nat. Sci. Pa. 1884, pp. 37, 50, pi. i. figs. 4, 7. Apsilus lentiformis . . . Leidy, Proc. Acad. Nat. Sci. Pa. 1884, p. 50. Miss S. G. Foulke, who discovered this Eotiferon, is of opinion that it differs from Apsilus lentiformis sufficiently to warrant its being regarded as a distinct species ; the points of difference being the shape of the cup, the absence of ganglion, the presence of a " second stomach," and the ciliation of the cup. If .1. bipera really has two stomachs, one above the jaws and the other below them, and each a closed sac with walls distinct from those of the body-cavity, then it would not only be a new species, but also a perfectly unique one among the Eotifcra. It is evident that further investigation is wanted on this and other points ; especially as Prof. Leidy is of opinion that Apsilus bipera, Dictyoplwra vorax, and Apsilus lentiformis are all the same animal. But whether Miss Foulke's species be a new one or not, to her is due the discovery of a true ciliary wreath within the coronal cup. It consists of two gradually narrowing ridges, fringed with long cilia, and running up the inside of the dorsal surface of the cup (fig. 5 a). Short diagonal lines of finer cilia can be indistinctly seen between the larger set. This ciliary apparatus is quite unique in position; and, if A. bipera and A. lenti- formis are the same, it is curious that Miss Foulke should have missed the nervous ganglion, and that Herr Metschnikoff should have missed the ciliary ridges. Length. Up to -T0 inch. Habitat. Water-plants in Fairmount Park, U.S. (Miss Foulke). 60 THE EOTIFEEA. Genus STEPHANOCEEOS, Ehrciibcrg. GEN. CH. Lobes long, slender, erect, convergent; setae set diagonally on the lobes in parallel bands ; foot terminated by an adhesive cup. S. EICHHOENII, Ehrenbcrg. (PL IV. fig. 1.) Stcplianoccros EicMwrnii . . Ehrenberg, Die In/us. 1838, p. 400, Taf. xlv. fig. 2. ,, ,, Dujardin, Hist. Nat. Zooph. 18-11, p. 612, pi. xix. fig. 8. „ ,, Gosse, Popular Sci. Rev. vol. i. 1802, p. 30, pi. iii. and iv. Stcplianoccros glacialis . . . Perty, Zur Kenntniss klcinst. Lcbcnsf. 1852, p. 47, Tab. i. fig. 1. Stcplianoccros Eichhornii . . Leydig, Ucb. d. Baud. Radcrtli. 1854, p. 5, Taf. i. figs. 1-4. ,, ,, Pritchard, Infusoria, 1801, p. 668, pi. xxxii. fig. 383, pi. xxxvii. figs. 1-4. „ „ Cubitt, Hon. Micr. J. vol. iii. 1870, p. 240, pi. Hi. ,, ,, . . . Ncwlin Peirce, Proc. Acad. Nat. Sci. Pa. 1875, p. 121. ,, „ ... Rosscter, J. Boy. Micr. Soc. 2 tier. vol. iv. 1884, p. 169, pi. v. figs. 1-3. Anyone who has seen Stephanoceros favourably placed, and properly lighted, can well understand the enthusiasm with which Eichhorn relates its discovery ! ; for it is a lovely creature, and as strange as it is beautiful. A small pear-shaped body, whose rich green and brown hues glow beneath a glistening surface, is lightly perched on a tapering stalk, and crowned with a diadem of the daintiest plumes : while the whole is set in a clouded crystal vase of quaint shape and delicate texture. The tube is denser than it is in the Flosculcs, is more symmetrical in shape, and is continuous in sub- stance from its outer surface almost to the creature's body. If an empty tube be examined, it will be found that it has a central hollow, which the body and foot will exactly fill. Mr. Gosse and Dr. Mantell have each seen a young Stephanoceros bore its way through its parent's tube by means of its cilia ; just as I have several times seen young Floscules do. The material, therefore, of which it is composed, must be of the flimsiest kind. The commencement of the formation of the tube has been described by Mr. Gosse (loc. cit.} as follows : " A specimen, which was hatched under my eye, swam for ten minutes, and then became permanently attached to the upper glass of the box, so that it was vertical in its position, with the foot next to the eye ; a favourable aspect for observing the development of the case. It presently began to dilate its body ; and, in about five minutes from its attachment, I perceived a distinct filmy ring around it, per- fectly circular, whose diameter was about twice that of the body (PL IV. fig. 8). The little animal now began to lean over to one side, and the ring soon had another segment additional, leaning in the same direction (fig. 9). The case, for such it was, looked like two broad hoops of glass, each swollen in the middle and set one on the other but not quite concentrically, at least to the eye of the observer. It was manifest that it was produced from an excretion from the body, owing its form and size to the animal's mov- ing round on the foot as on a pivot." Ehreiiberg's drawing of Stephanoceros has certainly been taken from a crushed or sickly specimen, and, indeed, in the majority of cases its portrait has been drawn too long after the creature had left its native haunts ; for when freshly caught and in vigorous health it arches its five plumes so that its crown almost forms a sphere.2 The 1 P. 18, supra. - Mr. Gosse has found that healthy specimens, removed from an aquarium and inspected at once, have their five arms more frequently produced into a cylindrical form, with their extremities incurved, than arched into a sphere. FLOSCULARIAD/E. 01 setae arc far longer than they appear at first sight, and are not stiff bristles as Ehrenberg has drawn them, but are gracefully curved, and taper off into lines of exquisite fineness. Those of one arm interlace with those of the arms on either side of it, so as to form a living cage of the finest network, through which it is hardly possible for anything to pass without striking some part of the sensitive meshes. The instant this happens band after band of the setas lashes at the runaway, a swift wave of motion runs along each band, and the captive is thrown back into the vortex produced by the wreath at the bottom of the coronal cup, the ciliary armature of which is precisely like that already described in Floscularia. I have also on more than one occasion detected a fitful ciliary wave running round the top of the coronal cup, just under the level of the lowest points of the depressions between its lobes'. This has not hitherto been noticed, but I am certain of the fact : the motion was of the briefest duration. There are considerable differences of opinion about the muscular system. Dr. Leydig (loc. cit.) says that there are four muscles which rise in the foot, and each of which divides into a pair, as it crosses the trunk, and then subdivides into smaller branches, as it passes over the coronal cup to the base of the lobes. Mr. Gosse makes them to be five pairs, and says that usually each pair runs up the trunk from the foot in a line with one of the arms ; and then, before reaching it, divides into diverging branches which, at remote points, are united to a muscular collar close to the base of the arms. He notices, how- ever, that he has seen cases where the muscles run down direct from the depressions between the lobes without uniting to form pairs. My own opinion, after prolonged observation of many specimens, is that there are really six pairs of muscles, and that they are arranged in the following fashion. Each pair runs up the foot looking like a single muscle ; and the reason why never more than four (pairs) are visible in the foot from any point of view, is that there is always a pair on each side of the animal (however viewed) which is there lost to sight. At the junction of the foot and trunk each pair begins to open a little ; and by the time they have reached the bottom of the coronal cup the constituents of each pair diverge obviously from each other, and terminate usually at the base of some one of the depressions between the lobes : but in such a fashion that the constituents of the same pair never end in the same depression (PI. IV. figs. 2, 3, 4, lm). There is, however, an exception to this in the case of the two pairs of dorsal muscles (PI. IV. fig. 2, lm). Here it will be seen that while the outer muscles in each pair end in a depression between the lobes, the inner muscles curve over towards each other and meet so as to form a fine arch, some distance below the base of the dorsal lobe. There are, too, fine hexagonal markings visible on this side of the coronal cup, which are probably the boundaries of large cells : oval nucleated cells are also easily seen in the wall of the coronal cup, when the animal is viewed from either side (fig. 4). The nutritive and reproductive systems are so similar to those of Floscularia, that they require no separate description. It is enough to call attention to Dr. Leydig's figure of the ovary treated with acetic acid (reproduced in fig. 7), and exhibiting the ova in various stages of growth, as well as its own delicate walls, and the oviduct (ot), which leads into the cloaca (cl). The Secreting System. — Neither salivary, gastric, nor foot glands have been observed in Stcplianoceros, but as the animal secretes a large and comparatively solid tube, it is clear that it must either have some organ for this purpose, or that i