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Ttaubenfqiii B. 61 Niederl)Ana TtaubenfQ III b. 61 nieDerl)ana ON A NEW CLASSIFICATION OF THE BDELLOID ROTIFERA. By DA VID BRYCE. (Read June 28th, 1910.) h has long been felt by those who are interested in the BDELLOID HOTIFERA that a revision of the classification of this group would considerably facilitate further investigation into a comparatively little-known corner of the animal kingdom. During the last eighteen years the number of known species has more than doubled, the great majority of the new forms being additions to the two genera Philoclina and Cctlliclina, which have conse­ quently become overcro:wded, unwieldy, and unsatisfactory. Besides this, a more intimate acquaintance with the diversities of structure and of habit of a gr'eatly extended array of species has proved that not only are the old generic definitions in­ f.dequate, but that they are also unreliable, and should no longer be accepted. The object of this paper is to place the classification of the BDELLOIDA on a more satisfactory basis, and it is hoped that the arrangement now put forward will provide a sound foundation, or, at the least, a new starting-point for future work, and that the lines on which it is framed will prove to be reliable and true to the natural relationships of the species with which it deals. Fl'Om the point of view of classification the BDELLOID ROTIFERA have already experienced a somewhat complicated career. Their history as a recognised group of allied species seems to have begun in 1830, when Ehrenberg published his first Classified Li"t of Micro-organisms (2), wherein he introduced the family Zygot1"ocha, comprising all Rotifera with a ciliary wreath of two similar parts. So far as regards the BDELLOIDA this earliest classification may be summarised thus: FA~IILY ZYGOTROCHA. Rotifera with corona of two similar parts (" ciliorum coronulis binis "). Loricate Section Bmchionaea. Illoricate Section Philodinaea. 62 D. BRYCE ON A NEW CLASSIFICATION m' THE BDELLOID ROTIFERA. D. BRYCE ON A NEW CLASSIFICATION OF THE BDELLOID ROTIFERA. 63 SECTION PHILODINAEA. use of the number of spurs and of toes on the foot. It has been Without eyes . Gen. Calliclinct. pointed out by lVIurray in a recent p}1per (63) that the dis­ tinction made was inaccurate as between Rotife?' and Actin~t?'~ts, With two eyes. since the foot is not thrice furcate in the species assigned to Eyes frontal: Rotifm·. But if inaccurate in that case, the phrase" cauda ter Foot thrice furcate (" cauda ter fur- furcata" is conect with regard to the genus Philodh~a, and cata ") Gen. Rotifer. clearly indicates the two spurs, the two dorsal toes, and the Foot ending in two spurs and three two terminal toes posse' ed by all but one of the species which toes (" caudae quinque apicibus") Gen. Actinun~s. were described by Ehrenberg as members of that genus. Eyes dorsal: In 1884 Hudson (17) recognised the distinctive character of Foot simply furcate (" cauda sim- the manner of creeping peculiar to the group, and proposed pliciter furcata ") Gen. jJ£onolabis. that the several genera should form a separate order, that of Foot thrice furcate (" cauda ter fur- the BDEJ,LOIDA, or I,eech-like Creepers, and this proposal was cata") Gen. Philodina. further established by its adoption in 'l'he Rotife?'a, published by him in 1886 in collaboration with Gosse (19). In this work In the following year, 1831, Ehrenberg published a more the new family Adinetadae and the new genus Aclineta were comprehensive arrangement (3), in which the Philodinaea were created for the reception of a species which differed markedly advanced to the rank of a family, and this position was again in the type of the corona from all others of the group included assigned to them in his great work of 1838 (4), based upon his by them at that time. The four recognisable genera of Ehren­ third and best-known system of classification. In these later bel'g were placed in the new family Philoclinadae, and were schemes the two genera Typhlinct and Hyclrias were added to distinguished as before by the presence and position of the eyes. the Family with the following characters: Earlier in the same year, 1886, the importance given by vVithout rostrum or spurs: Ehrenberg to the eyes in the generic distinction of the Philodinaea Trochal discs on pedicels Gen. Hyclrias. had bpen challenged by Milne (18), who proposed to arrange Trochal discs without pedicels Gen. l'yphlina. the various species into genera either new or redefined, and to It bas not yet been found possible to recognise any of the discard altogether all generic characters relating in any way species assigned to the genei'a jJ£onolabis, Hyd?'ias, and l'yphlinct, to the eyes. He claimed for his scheme that it did not dis­ and these genera have not been accepted by later writers, who sociate manifestly similar forms, at least as regards some nineteen believe them to have been founded on imperfect observations species examined by him. His most valuable suggestion in this of animals which, if again seen, have been referred to other raper was that the genus Philodina should be distinguished by groups of the Rotifera. The four genera, Callidina, Philoclina, the possession of four toes, thus giving first place to the character Rotifm', and Actimwus, have fortunately proved to be recognis­ which Ehrenberg had indicated in 1830 in the phrase "cauda able, and the majority of the species, which have been discovered ter furcata." since 1838, have been assigned to one or other of them. In 1888 Milne (23) adduced fresh instance; in support of his As in the classification of 1830, so in his later schemes, previous contention, and proposed further that the genus Rotife?' Ehrenberg distinguished the four genera last named principally should be distinguished by the character" viviparous." upon characters afforded by the presence or abseoce of eyes, and, Another important advance was made in 1889, when Plate when present, by their position, either in the front of the head (27) pointed out that the Bdelloilla shared with the Seisonidae or in the neck. As a quite subsidiary character, to distinguish the peeuliarity of having two ovaries, whereas all other Rotifera Rotife?' from Actim~rus, and Pkilodinct from illonolabis, he made have one only. He proposed therefore to divide the class RO'l'Il'ERA 64 D. BRYCE ON A NEW CLASSIFICATION OF THE BDELLOID ROTIFERA. D. nRYCE ON A NEW CLASSIFICATION OF THE BDELLOID RO'l'JFERA. 65 into two sub-classes, the DIGONONTA (or two-ovaried), comprising In 1905 James Murray (55) announced the discovery of the the Bdelloida and the Seisonidae, and the MONOGONONTA (or one­ curious Bdelloid, Jlfic7'oclinc£ IJC£7'acloxct, for which he created the ovaried), including all other Rotifera. new family .Afic'l'odinadc~e. This and numerous other discoveries In a useful monograph on the Philoclinaec£, published in 1893, of Bdelloid forms hitherto unknown, and in all cases communi­ J anson (38) discussed at some length the views and suggestions cated to me before publication, led naturally to the discussion in of earlier writers and, in particular, those of Hudson and Gosse, our correspondence of the demerits of the current classification of and of Milne. On the one hand, he criticised the creation by the the group. The arrangement of the genera and species now ad­ former authors of the family of the Aclinetadae. On the other, vanced is in great measure the outcome of that discus,ion. To he admitted the contention of Milne that under the definition of some extent the lines on which it is mainly framed have been Ehrenberg many eyeless species would be classed as Callidinae, indicated by my correspondent in recent papers, notably in (56) although in respect of their structure they should clearly be "The Bdelloid Rotifera of the Forth Al'ea" (1905) and in (63) regarded as belonging to the genus Rotife?" Nevertheless he " Philodinc£ mc£crostylc£ and its Allies" (1908). hesitated to accept the genera proposed by Milne, and preferred In the fOl'mer of these he provision'tl1y redefined the genera fol' the time to abide by the Ehrenbergian family of Philod'inaea, Philodina, Calliclina, and Rotife7' as follows: which in his view covered all the various genera. He made the PJIILODINA.-Having four toes and a corona consisting mainly one correction of transferring to the genus Rotife7' the two species of a pair of wheel-like ciliated discs. which had been assigned to Actimwt~s, recent discoveries having shown the differences between these two genera to be less definite A. Eyes present; ovip~rous. than had previously appeared. B. Eyes absent; oviparous. In an important treatise publisMd in 1899 Wesenberg Lund O. Viviparous; eyes present or absent. (50) dealt in great detail with the wide question of the l'elation­ OALLlDTNA.-Having three toes or a perforate disc formed by ship to each other of all the various groups of the Rotifera, and, a union of the toes; oviparous; eyes present or absent. in conclusion, put forward a new classification based largely upon A. Food moulded into pellets. results afforded by his own investigations. At the outset he B. Toes bearing a number of cup-like suckers, or united followed Plate in dividing the class ROTIFERA into the sub-classes to form a broad disc. MONOGONONTA and DIGONONTA according to the number of ovaries O. Toes three; distinct, food not moulded into pellets. possessed by each species. So far as regards the MONOGONONTA, RO'l'TFER.-Viviparous; toes three. the subsequ!\nt grouping of the families and genera was carried out on principles essentially different from those of Hudson and In the latter paper he discusses exclusively the genus Philoclinc£, Gosse.
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