apical, often independently jointed; (2) with the outer ends of the rami articulate two lateral pieces (mallei), and again composed of a distal longitudinal piece (manubrium) and an apical transverse piece (the uncus), the whole recalling, as the name implies, a single-clawed hammer. For the varieties and modifications of the trophi we simply refer to Hudson's figure above. The relative size of the crop to the trophi varies greatly; it is small where the trophi are well developed and complex, as well as in Bdelloidea; but in Flosculariaceae it is large, and so it is in Asplanchnaceae. Eversible trophi of the forcipate or virgate type, which can be used for nibbling, are common in Ploima, notably Rattulidae, and are used for attachment to the host in the parasitic Seisonaceae, &c. In Asplanchnaceae also, From H. S. Jennings in American Naturalist, vol. xxxv., by permission of Ginn & Co.
FIG. 3. - a, Stephanoceros eichornii in gelatinous tube; b, Acyclus inquietus in gelatinous tube, with eggs; c, Floscularia coronetta in gelatinous tube, with eggs; d, A psilus bucinedax, showing lateral (distal) antennae funnel of mouth hanging into enormous crop, stomach at apical end with gastric glands, anus on postero-ventral surface, large coiled kidneys at proximal end, uniting into median duct; e, Melicerta ringens in tube; f, same, proximal end enlarged, showing a pellet in the cup proximal to the paired lateral antennae; g, Melicerta janus, tube formed of faecal pellets.
where the whole crop is strengthened by a framework of bars, the incudate mastax lies in a little postero-ventral pouch which can be everted through the crop and mouth. The stomach is generally large; its wall consists of a layer of very large ciliated cells, which often contain fat globules and yellowish-green or brown particles, and outside these a connective tissue membrane; muscular fibrillae have also been described. Very constantly a pair of simple sacklike glands open into the stomach, and probably represent the hepato-pancreatic glands of other Invertebrates.
Following upon the stomach there is a longer or shorter intestine, which ends in the cloaca. The intestine is lined by ciliated cells. In forms living in a tube the intestine turns round and runs forward, the cloaca being placed so as to debouch over the margin of the tube. The cloaca is often very large; the nephridia and oviducts may open into it, and the eggs lodge there on their way outwards; they are thrown out, as are the faecal masses, by an eversion of the cloaca. Asplanchna, Notommata seiboldii, and certain species of Ascomorpha are devoid of intestine or anus, excrementitious matters being ejected through the mouth. The body cavity (archicoele) contains a fluid in which very minute corpuscles have been detected. There is no trace of a true vascular system. The nephridia (fig. 1, B, n) present a very interesting stage of development. They consist of a pair of tubules with an intracellular lumen running up the sides of the body, at times merely sinuous, at others considerably convoluted. From these are given off at irregular intervals short lateral branches, each of which terminates in a flame-cell (f) precisely similar in structure to the flame-cells found in Planarians, Trematodes and Cestodes; here as there the question whether they are open to the body cavity or not must probably be answered in the negative. At the base these tubes open either into a permanent bladder (fig. 1, bl) which communicates with the cloaca, or directly into the cloaca. They have the same functions as the contractile vacuole of freshwater Protozoa. Nervous System. - There is a large ganglion lying in close contact with the pharynx, proximal to the crop and on its antero-dorsal side; in Bdelloidaceae at least it is united by short connectives with a smaller postero-ventral ganglion to form a nerve collar. From this simple nerve fibres are given off to the body-wall, especially 4uk t A ,x °C FIG. 5. - Pedalion mira. A, lateral surface view of an adult female: a, median ventral appendage; b, median dorsal appendage; c, distal ventro-lateral appendage; d, dorso-lateral appendage; f, dorsal antenna; g," chin "; x', cephalotroch. B, lateral view, showing viscera: oc, eye-spots; n, nephridia; e, ciliated toes; other letters as above. C, ventral view: x', trochus; x, cingulum; other letters as above. D, ventral view, showing the musculature (cf. text). E, dorsal view of a male: a, lateral appendages; b, dorsal appendage. F, lateral view of a male. G, enlarged view of the antenna f. H, enlarged view of the median ventral appendage. (All after Hudson.) to the ciliated cells of the corona, to the foot, and also to the muscles and sense organs.
The sense organs are eyes, antennae, sensory styles and a statocyst in a few species. The eyes are refractive globules set in a cup of red pigment traversed by a nerve fibre, and lie on the proximal side of the body, directly on the postero-dorsal surface of the brain, or at a little distance from it, on the neck, often within the circle on the corona, and usually well within the transparent body. There may be one, a pair, or rarely more, the outer ones being more or less rudimentary. The antennae are short tubular extensions of the body wall, sometimes retractile with a depressed tip from which protrudes a tuft of fine stiff bristles. They are possibly organs of external taste (smell) as well as of touch. Typically there are two pairs - a proximal, more or less approximated on the postero-dorsal surface, and a distal pair, more widely separate. But the proximal pair are often fused into a single median antenna (supplied, however, by two nerves), and in one case at least the distal pair may be similarly fused. Additional paired antennae may occur within the coronal surface, which is the seat of the sensory styles, of less complex structure, which occur in many genera. The statocyst (retro-cerebral organ of P. Marius de Beauchamp) is a sac filled with highly refractive granules soluble in dilute acids, and opening by a slender duct (or a pair) to the surface: its function is doubtless that of an organ of equilibrium, and it resembles in its opening to the surface the primitive internal ear of even Vertebrates, for the duct to the surface persists through life in the sharks.
Locomotor Organs
Most free rotifers swim by the corona, aided by the ciliated auricles when present. In Bdelloidaceae this may alternate with a leech-like gait; the corona being withdrawn, the cupped end of the proboscis serves as a sucker for attachment alternately with the adherent foot, so that the animal loops its way along. In two families motile articulated rods occur; in Triarthridae they probably simply expand the dimensions of the body in adaptation to life at the surface; or as a protection against being swallowed by their smaller foes. In Polyurthra and Pteroessa they are numerous, pinnated (feathered), and are doubtless used for active swimming by jerks; they can be moved up or down by special muscles attached to their bases, which project into the body.
FIG. 6. - Male Rotifers. 1, Euchlanis deflexa; 2, lateral; 2a, dorsal views of Colurus bicuspidatus; 3, Notops brachionus; 4, Diglena permollis; 5, Gastropus minor; 6, Anuraea serrata; 7, Ascomorpha parasita; 8, Notholca heptodon. (Drawn from specimens by F. R. Dixon Nuttall.) In Pedalion (fig. 5), a remarkable form discovered by Dr C. J. Hudson in 1871 and found in numbers several times since, these appendages have acquired a new and quite special development. They are six in number, median, ventral and dorsal, and two unequal lateral pairs. The largest is placed ventrally at some distance distal to the mouth. Its free extremity is a plumose fan-like expansion (fig. 5, Aa and H). It is, in common with others, a hollow process into which run two pairs of broad, coarsely transversely striated muscles. Each pair has a single insertion on the inner wall - the one pair near the free extremity of the limb, the other near its attachment; the bands run up, one of each pair on each side, and run right round the body forming an incomplete muscular girdle, the ends approximating in the median line. Above this point springs the large median dorsal limb, which terminates in groups of long setae. It presents a single pair of muscles attached along its inner wall which run up and form a muscular girdle round the body in its posterior third. On either side is attached a dorsolateral and ventro-lateral appendage, each with a fan-like plumose termination consisting of compound hairs or setae, found elsewhere only among arthropods (q.v.); each of these is moved by muscles running upwards towards the neck and arising immediately under the trochal disk, the inferior ventro-lateral pair also presenting muscles which form a girdle in the hind region of the body. It bears a group of long setose hairs the bases of which are connected with the nerve fibre. There are also two pairs of distal antennae. Pedalion presents a pair of ciliated toes in the posterior region of the body (fig. 5, B, C, and D, e), which it can apparently use as a means of attachment; Dr Hudson states that he has seen it anchored by these and swimming round and round in a circle.
Reproduction Organs. - Rotifera are unisexual, with the sexes dimorphic. The ovary is, as in many Platyhelminthes, duplex; one part, the germary, being an organ for the production by cell multiplication of the germ-cells or eggs. proper, the other, the vitellarium, much more conspicuous and usually consisting of a definite number of large cells, producing yolk material for the growth of the egg. The whole ovary is unilateral and unpaired in most rotifers; symmetrical in Asplanchnaceae, Philodinaceae and Seisonaceae. In Asplanchnaceae the germary is median, continuous at the distal end with the middle of the transverse horseshoe-shaped vitellary. In Bdelloidaceae and Seisonaceae the whole organ is paired, the germary proximal, the vitellary next the cloaca. As a rule, the wall of the ovary is continued into a uterine tube opening into the cloaca; but in Philodinaceae this is absent, and the young are free in the body cavity and escape by perforating the cloacal walls. The male organs are usually a testis, a large seminal bladder and a protrusible penis. The males are unlike the females in most species; only in Eosphora digitata, Rhinops vitrea, Proales werneckii, and the Seisonaceae a complete digestive system is present. Frequently the foot is ciliated at the tip, as in the young of tubicolous forms.
The males of rotifers are of relatively rare occurrence, except in the genus A splanchna, where they were first recognized as such by Brightwell in 1841; though those of Hydatina had long since been seen and described as a distinct genus. Despite their rare occurrence, the males of over one hundred and twenty species have now been recognized, and we may well believe that all species will be found to present males. This statement may seem to need qualification; for the male of no Bdelloid has been seen, and there is but a doubtful record of" winter-eggs in this group. But possibly, as in Seisonaceae, the males resemble the females, and have escaped recognition. It may, however, well be that the capacity for wintering in the dry state has physiologically replaced the need for resistent fertilized eggs. Insemination takes place either by the introduction of the penis into the cloaca of the female, or by the puncture of the bodywall of the female by the penis, and the injection of the sperm into the body cavity, whence the single spermatozoa must make their way to the eggs. The females habitually produce eggs without impregnation, which again habitually develop into females, more rarely into males. These unfertilized eggs develop directly, often in the uterus. In other cases the eggs are liberated earlier and adhere to the foot, or are hatched within the tube (fig. 3, b, c). The impregnated eggs undergo a very partial development in the mother, and these pass into a state of rest, for which they are furnished with a dense shell. They always give rise to parthenogenetic females (see Reproduction). The thin-walled eggs are often termed "summer-eggs," the fertilized ones "winter" or "ephippial" eggs (by parity with the phyllopod Entomostraca, q.v.). But the appearance of males seems to be as much associated with those of summer drought as of winter cold. No adequate knowledge of the conditions under which males arise has been established. The phenomenon of seasonal dimorphism is of especial moment for the plankton dwellers. Not only is the appearance of males regular, but the forms of the females at different times of the year may be so distinct as to have led them to be classed as distinct species.
Relationships and Morphology
Passing over the earlier authors who regarded this group as allied to Infusoria, a view c ar first contested by Dujardin, T. H. Huxley viewed them as equivalent to and on a level with the larvae of Echinoderms, and of such other trocho phore larvae as resem- ???,, ally adopted. But eit u sh y ?e ? tt became more and more From Cambridge Natural History, vol. ii., "Worms,. apparent that the larvae &c.," by permission of Macmillan & Co. Ltd. of this category deFIG. 8. - Diagram of morphological rela veloped mouth, gut and tions of Rotifera. A, pilidium larva anus by the closure in of nemertine; B, Asplanchnapus the middle of such a slitschematized; C, a ploimal rotifer; like blastopore opening D, trochosphaera female (schematized into a sack-like stomach from Semper); E, veliger larva of mol as is seen in the larvae lusc; F, trochophore larva of annelid.
of Turbellaria and Ne- a, anus; ap, apical organ, correspond mertina. The extraing to foot of rotifers; at, median blastoporic opening of antenna, united by a nerve to br, brain the cloaca leads us to a (letter omitted in B); bl, bladder, re ver y different view, which ceiving ramified kidney in B, C, D; finds negative support f, foot, and f.g, its cement-gland;.
in the failure of previous g, ovary; k, kidney; m, mouth; n, supra morphologists to adapt oesophageal ganglion; nr, nerve ring in the details of developsection.
ment and of the struc ture of the disk to their identification of "trochus" and "cingulum" with the preoral and postoral wreaths of the trochophore larva. We homologize the rotifer with the Turbellarian larva (fig. 8, A), and with the preoral or upper part of the trochopore (fig. 8, E, F). Its adhesive foot is paralleled by a cup-shaped ciliated depression, possibly nervous, found in all the larvae cited, except some Echinoderms, and which in Asterids and Crinoids actually serves as an organ of attachment. This view obviates the deed for assuming the complicated flexures of the wreath which has to be done on other assumptions (see Rotifera, Encycl. Brit. ed. 9). Thus Trochosphaera (fig. 8, D) (which has a male of the same type as Melicerta, &c.) is an extremely modified type, and its resemblance to the trochophore larva of Lepadorhynchus or Polygordius is only superficial. We may note that it was long since shown that the apical organ (at first assumed to be the brain) of these larvae was innervated from an anterior thickening of the circular nerve ring, corresponding with the brain of Rotifers; the nerve cells immediately below the pit are the ordinary bipolar From H. S. Jennings in American Naturalist, vol. xxxv., by permission of Ginn & Co.
FIG. 7. - Loricate Rotifers. a, Notholca longispina, lorica only; b, Anuraea aculeata, like the former, a floating pelagic type (plankton proper); c, Synchaeta stylata; corona with accessory antennae and sensory styles; auricles for swimming - an actively swimming pelagic type (nekton); d, Pterodina patina, with bdelloid corona and retractile foot with terminal ciliated cup; e, Distyla gissensis partly extended; f, Rattulus tigris. a r rn t?
F nr x s.n ganglion cells below invertebrate sense-organs. Moreover, the body cavity of the rotifers is a primitive archicoele; the persistent or accrescent cleft between epiblast and hypoblast, traversed by mesenchymal muscular bands. Thus we regard Rotifers as an independent stem branching off at the outset of the rise from the Platode type to higher Invertebrata The Polyzoa (q v), which in many ways recall Rotifers, appear to be equally independent.
The following classification of rotifers is our modification of that of Hudson and Gosse, further altered through considerations put From H. S. Jennings in American Naturalist, vol. xxxv., by permission of Ginn & Co.
FIG. 9. - a, Microcodon clavus, showing corona, lateral antennae and jointed foot; b, Rhinops vitrea, corona from below, showing proboscidiform extension containing eyes; c, Philodina megalotrocha; d, head of Rotifer macroceros, postero-ventral view, showing lobes of corona, and antero-dorsal median antenna, telescopic with setae; e, Rotifer (Actinurus) neptunius, showing head with retracted corona, and protruded dorsal proboscis bearing median antenna, and telescopic foot with toes and spurs; f. Asplanchnopus myrmeleo, showing horseshoe-shaped germarium (left), blind saccate stomach (right), apical bladder, foot, &c.; g, Asplanchna ebbesbornii - the coiled tube at left is a kidney; h, i, incudate jaws of Asplanchna brightwellii and girodii chiefly formed of rami, with the rudimentary mallei parallel and external to them; j, Ascomorpha hyalina. forward by C. Wesenberg-Lund, which, however, we do not consider wholly convincing. He notably regards an oblique disk with uniform ciliation as primitive, a view which we cannot adopt. Classification: (A.) Disk usually with well-marked strong trochus, ciliated groove and more delicate cingulua interrupted by an antero-dorsal median gap, usually more or less bilobed.
(i.) Trophi incudate: I. Asplanchnaceae; trochus circular; foot absent or minute; trophi incudate; stomach blind; males frequent, not very dissimilar to females. Asplanchna Gosse (fig. 9, g - i); Asplanchnopus Deguerne (fig. 9, f); Ascomorpha Perty (fig. 9, j). (ii.) Trophi malleoramal: 2. Melicertaceae; females tubicolous, usually attached, or forming spherical floating social aggregates; males free swimming. Melicerta Schranck (fig. 3, e, f); Oecistes Ehrenberg; Lacinularia Schweigger; Conochilus Ehrenberg, with gap postero-ventral and mouth anterodorsal (fig. 2, 5).
3. Trochosphaeraceae; female footless; subspherical, the corona bulging into a hemisphere which may equal the hemispherical body; anus apical; male as in Melicertaceae, Trochosphaere Semper (fig. 8, D).
4. Ploimoidaceae; subconical; corona bilobed; retractile foot absent or ciliated; motile appendages present in two families.
(a) Pterodinidea; foot a ciliated cup; cuticle forming flat lorica. Pterodina Ehr. (fig. 7, d). (b) Triarthridae; body with a pair of long cervical spines pointing distally and serving for leaping movements or to extend the body and make it too big for small enemies to swallow; Pedetes Gosse (no median spines); Triarthra Ehr., one postero-ventral spine; Tetramastix Zacharias, two unequal median spines.
(c) Pedalionidae, foot represented by two styles, sometimes ciliated; body provided with six hollow-jointed muscular fins for swimming and leaping. Pedalion Hudson (fig. 5).
(iii.) Trophi ramate: 5. Bdelloidaceae; foot with two toes and accessory spurs or a simple perforated disk; body telescopic at either end, with an antero-dorsal proboscis ending in a ciliate cup and bearing the proximal antenna; corona usually bilobed, very wheel-like. Males if present probably like the females. Germary and ovary paired; oviduct absent; young viviparous. Rotifer Schrank (fig. 9, d, e); Philodina Ehr. (fig. 9, c); Callidina Ehr. (eyeless); Adineta Hudson is eyeless with the corona uniformly ciliated, and proboscis adnate, hooked.
(iv.) Trophi uncinate: Flosculariaceae; disk a contractile cup, often lobed, the cingulum of long vibratile cilia, of very long motionless bristles or absent, rarely with an outer zone of fine cilia. Trochus a pair of ridges or horseshoe open in front. Oral funnel produced into a fine tube hanging freely into a pharyngeal cup, containing the uncinate trophi. Bodywall usually traversed by a network of canals serving by their contraction to expand the disk. Males and larvae with a ciliated pedal cup and a simple ciliated disk.
(a) Floscularidae; tubicolous, with a lobed disk, bearing stiff or vibratile setae. Floscularia Oken (fig. 3, b); Stephanoceros Ehr. (fig. 3, a).
(b) Acyclidae. Disk entire or tentaculate, not seti ferous; Acyclus Leidz (fig. 3, c). Foot repre sented by a button-like disk, carried far from the posterior surface; Apsilus Metchnikoff (fig. 3, d); Atrochus Wierzerski (fig. 3, c). (B) Ploimaeae; disk variable, often circular, sometimes with a lobed trochus bearing membranelles (vibratile styles); trophi complete, malleate, submalleate, virgate, or forcipate; anus subapical; foot usually short, and usually bearing two toes which may be much elongated.
Illoricata, cuticle soft; ciliated exsertile auricles above the disk sometimes present. Albertia Dujardin; Drilophagus Vejdovsky; Microcodon Ehr. (fig. 9, a); Rhinops Hudson (fig. 9, b); Synchaeta Ehr. (fig. 7, c); Hydatina Ehr. has no eye; Notommata Ehr. (restricted by Gosse); Copens Gosse; Notops Hudson (fig. 6, 3); Proales Gosse; Gastroschiza; Diglena Ehr. (fig. 6, 4).
Loricata, cuticle hardened armour-like, often sculptured; Polyarthra Ehr.; Pedetes Gosse; Euchlanis Ehr. (fig. 6, I); Anuraea Ehr. (fig. 7, b) , Notholca Gosse (fig. 7, a) Distylis Eckstein (fig. 7, e); Rattulus Ehr. (fig. 7, f); Colurus Ehr. (fig. 6, 2); Taphrocam pa Gosse.
(C.) Seisonaceae. Body elongated with a narrow neck above the disk; foot ending in a terminal perforated disk. Trophi virgate exsertile; germary paired; genito-urinary cloaca opening above the neck in the male, subapically in the female. Gut blind (Paraseison), or opening into cloaca (Seison). Males resembling females, common. All known species are parasitic on the Crustacean Nebalia; Seison Claus; Paraseison Plate.
History and Bibliography
As rotifers are common in ponds, the first workers with the microscope observed them repeatedly, the first record being that of John Harris in 1696, who found a Bdelloid in a gallipot that had been standing in his window. Leeuwenhoek found and described some tubicolous species; and during the 18th century a fair number of species were observed, figured and described with names. During this time the illusion of a wheel or wheels produced by the ciliary action of the disk had puzzled all observers. C. E. Ehrenberg included the Rotifers in his Infusionsthiere, and described and figured with fair precision many of the genera and species. Dujardin gave a less detailed but more accurate account under the name of Zoophytes Systolides. The next full, work was a valuable compilation by W. C. Williamson (best known as a botanist) in Pritchard's Infusoria, in 1861. Much work was done with the gradual introduction of improved methods during the last forty years of the century. The discovery and recognition of the males was made, however, at the close of the fifties. P. H. Gosse collected and described many species, and elucidated the structure of the mastax in 1856. Zoologists of the standing of Huxley, Claus and Leydig added to our knowledge of the anatomy and to the theory of their relations. But the monumental monograph of C. T. Hudson and Gosse containing a new classification, an illustrated description of all the then known species and much information on habits and structure, provided students with an easy access to the domain and stimulated many to work hard at the group. Of these new-corners we may cite C. F. Rousselet, who has found many new species and many unknown males of known species, elucidated habits and faithfully kept record of the publications on the class in the Journal of the Royal Microscopical Society. He has moreover elaborated a method for preserving Rotif era for microscopic observation, so that the types of each observer are now as readily available for comparison as the plant-specimens of the botanist's herbarium. C. Zelinka has given us the most detailed anatomical accounts we possess for several Bdelloidaceae, and was the first to utilize modern methods of microscopic technique on a complete scale.
C. G. Ehrenberg, Die Infusionsthiere als vollkommenere Organis- (1838); F. Dujardin, Histoire naturelle des zoophytes (1841); T. H. Huxley, "Lacinularia socialis," Trans. Micr. Soc. i. (1853); P. H. Gosse, "Manducatory Organs in Class Rotifera," Phil. Trans. (1856); W. C. Williamson, "The Rotifera" in A. Pritchard's History of the Infusoria (1861); C. T. Hudson and P. H. Gosse, The Rotifera (1886), and supplement (1889); Marcus Hartog, "Rotifera," in Cambridge Natural History, vol. ii., reprinted 1901; H. S. Jennings, Synopses of North American Invertebrates, xvii., "The Rotifera," Amer. Nat. xxxv. (1901); C. F. Rousselet, numerous papers in Journ. Micr. Soc. and Journ. Quekett Club; C. Wesenberg-Lund, "Danmarks Rotifera," in Vid. Meddel. Nat. For. Kjiibenhavn (1899) C. Zelinka, "Studien fiber Rotiferen," in Zeit. Wass. Zool. xliv. (1886), xlvii. (1888), liii. (1891). (M. HA.)
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Bibliography Information
Chisholm, Hugh, General Editor. Entry for 'Rotifera'. 1911 Encyclopedia Britanica. https://www.studylight.org/encyclopedias/eng/bri/r/rotifera.html. 1910.
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