204 DR. N. T. MASTERS ON THE DETELOPMEXT OF

On tlic Development of the dndrecium in CochZiostenzn, Lem. By M. T. ~IASTERS,KD., P.R.S., F.L.S., &c. [Rend March 7, 1S’iS.I

[PL LTE m.] INthe ‘ Gardeners’ Chronicle’ for 1S63, pp. 264, 323, I had occa- sion to describe that most remarkable and beautiful Coch- liostenm Jucobiniiuitt, Lcm., and to give the history of its intro- duction into this country. At that time I was only enabled to exailline adult , and, with seine imprudencc it must be con- fcssed, I hazarded a theory as to the peculiarities of the stamiiinl apparatus. I have since studied the course of devclopment of the in question ; and, as a result, I find that while my iiescrip- tioii of the adult flowrJJXS in the main accurate, my theoretical es- planation of its peculiarities rras in niany respects erroneous. It is vith the TieJT of correcting these errors and induciug others to incestigate the peculiady reinarkable structure of this splendid plant, that I norr beg to lay before the SocietF the follon-ing par- ticulars as to the development of the flon er, and especially of the andrcecium. The floirer first appears in the form of a globular tubercle (Pl. IT. fig. I), on one side of wliich shortly appears a thick semiluuar pro- minence, which is the origin of the posterior (fig. 2). The two remaining are formedin succession, one after thc other(fig. 3). I have Iierer absolutely secn the flower with two sepals only ; but, from the small relatire size of one of the thee sepals (that placed nest the bract) and from analogy Kith other florrers, I haye no doubt that the three sepals are formed in succession. If the three sepals be turned back rrith the dissecting-n’eedle, as shorrn in fig. 4, the central gron-ing point inay be seen to haTe lost its glo- bulatj foriii, to haye assuiiied a triangular outline, and to be some- what depressed at the top. From the three corners of the central growing point protrude simultaneously the thee (fig. a), which only precede the ap- pearance of the starnew by a very short interval-so Simt, indeed, that one inay esainine a great many flovers before fincling one in which the petals alone arc formed nithout indications of the . At this period the superiority in relatire size of the posterior and first-formed sepal is Tery mnrl~ccl. AS to the andrcfciuin, it is perfectly cleni. that this consists of six eleinents ; oud all analogy would suggest that thesix portions THE AXDZECICX IX COCHLIOSTEXA. 20.5 should be in txo i*ows, one within the other, the innermost deve- loped bubsequently to the outer. I cannot assert positiTely that this is not so ; all that I can say is, that after examining several floners I liare never been able to see that the andrcecium follows this caurse. It has always appeared to iiie that the three posterior stamens are clm eloped at about the same period aid in advance Of the three anterior ones (figs. 6,S, 9) ; at myrate, shortly after the , have nianifestcd themdws, the anihceciuni niay be seen to consist of three tubercles at the back of the flower, all three of about equal size, and of thee anterior tubercles much sqaller than the posterior ones, and of uuequal size, the central one being much smaller than the other trio (figs. 8, 9). The position of the staminal tubercles is as follow: the odd posterior is opposite the first-formed sepal j the two postero-lateral stamina1 tubercles are opposed to two corresponding petals. Of the three tubercles in the anterior part of the flower, the tiyo lateral are opposite the sepals, the central one is opposite the anterior petal. The arrmgement, then, of the stamens is quite consistent with their disposition in ti\ o ron s formed one after the olher, though, as before stated, I hare not been able to satisfy myself that, in point of time, there is any such regularity of appearance. Fol- loxing the coiirx of the andrecium from this point, it will. be seen that the superior size of the thee posterior staminal tubercles is maintained, that they become notched at the summit (figs. 10, ll),and speedilF present traces of anthers. The three anterior tubercles, on the other hand, progress very slowly, and never form authers at their summits (fig. 12). It is now evident that the two antero-lateral tubercles are tho lateral " staminodes '' of the adult flower. The central tubercle is the least developed of the thee ; in fact it often ceases to grow, so that in the adult floner it fre- quentlphappens that it is oyerlookcd, or that it is really absent. Up to thi, time the six stamina1 tubercles hue been dibtinct ; but now, by the further growth of the entire andrcecium at the base, a shal- lon- tube is formed, the margin of n-hich bears the stamens and staininodes as just described. On ing to the relatively increased gronth on the posterior side of the flomi- as compared with that on the anterior half, the staminal tube is much deeper on the foorincr (fig. 13) than on the latter side (fig. 12); and it retaius this inequality to n wry marked degree in the adult flower, vhere? indeed, the three posterior stamecs niay be seen alxays inseparate for nearly half their length, Thile the two antero-lateral stami- Q2 206 DR. 31. T. MISTERS OX THE DETELOP3TEJT OF nodes are quite detached. The anther, in the early stages of its existence, is an ordinary tn-o-lobed structure (fig. 12), eacli lobe beiiigagniu subdivided into two minor lobules bj-a slight depression. There is at first no trace of the spiral curvature which afterwards becoinzs so remarkable a feature of these . The connective in the odd posterior stamen remains horizontal, Khilst in the t1.i.o postero-lateral anthers it assumes gradually a rertical position. The direction of groTtli of the anther, as of the coil, is centrifugal, i. e. froni the centre outwards and up\~ards(fig. 14). Not till after the anthers begin to inaiiifest the spiral coil is there ally in- dication of those petal-like hoods \\ hich couceal the anthers in th adult flower, of which they constitute so remarkable n feature. The course of development slioivs them to be inere petaloid out- growths from the sides of each of the postero-lateral filnments, grad~iallyincreasing till they coillpletely euvelope the authers (fig. 13). During this yrocess the anther of the posterior centrd stamen, which heretofore had maintained its proper position, be- comes, as it irere, pressed to the front in advance of the tw-o lateral anthers, and thus in the adult flower may give rise to the false impression that it is really aid organically interior (figs. 16, 17), an impression I myself had denI first described the struc- ture of the flower. Not till the stanleiis have assumed the cha- racters and positions just mentioned is there any indication what- ever of that curiously lobed and ciliated process which I formerly regarded as a staminode belonging to an outer series of abortive stamens. It is, despite its pojitioii and prominence in the adult flower, a mere outgronth from the thalamus. at the back of the flower (fig. 18). TT'hat, if any: iuorphological significance, or what physiological office it may have, is at present uuknon-n. It may be connected in some "ray \\ ith the process of fertilization j but this is a mere guess. The de~elopmentof the pistil presents nothiilg notcn ortliF or dif- ferent from what occurs in many otlier Endogenous flowers. Its three conponeut carpels are formed simultaneously-one posterior, two autero-lateral. They make their appearance soon r.fter the for- mation of the staiiiinal tubercles, aiid before an!- iudicntion of anthers is prescut (fig.8). The carpels are closed, adthe stjle formed, before the occurrence of thoce remarkable changes in the stamens dread>-:illucd to. It is only after repeated exmiinations that I venture to lay before the Society the results of my observtions on the dwe- TIIC .4RDB(ECKJJI IS COCHLICSTE3L.L. 207 loplnent of this remarkable flowr. I liave the inore hesitatioii iii cl.oiug so, as the course of development iii the andrceciuin is so differcut from that described by Payer in Tradescniztia or Coii!- me?yizn*. For the sake of comparison I add a bricf abstract of the conclusions arrived at by the eminent French snvnnt. Payer dcscribes the sepals as deyeloped in succession, the latest being the anterior one or that superposed to the '' h-nctie mdre ;" the petals are described as simultaneous in their appearance-one placed internally, one esternally, aucl one, half iilternallg, half externally. So far niy observations are quite in accordance with those of Payer ; but in the case of the aitdroecium there are ma- terial discrepancies, as dlbe obvious from a coinparison of what has been previously stated with Payer's account of the develop- ment of the stamens iu the regular-flowered Trndescanfia. The andrcecium, according to this botanist, consists of six stamens in two rows : the stamens of each whorl appear all at the same time ; but the meinbers of the iiiiier series, those superposed to the petale, appear first, before those placed in front of the sepals, rl wholly esceptioiial occurrence. It is unnecessary to say more as to the development of the other parts of the flon.er, or as to the inflor- escence, as ins obserdions on these points coincide with those of PaFer. I may, Iio~erer,indicate that although Cochliosfenta in the adult state is: by reason of its sti-ucture, far removed from other genera of the same family, yet there are points in its conforination rrliich appear to connect it iyith other genera of the sniiie family. For instance, in ConzmeZynn itself, of the six stamens the three posterior only are fertile, i. e. one of tllc outer row and two of thc inner, exactly as iii Coc7~Ziosteiiza. 111 soine species of dneleiii~non the other hid, as also in Dichcrupern~zcnz,the three outer stameiis; placed in front of the sepals, are fertile, the three inuer oiies bring sterile. Dithyi-ocapmhas all six stamens fertile. The hairs ou the filaments of Tradescantias and Cotnmelynas may probably be looked on as having soine relation to the way iu which the flon-er gets fertilizcd ; but whether they liare the same ofice as the petal-like hoods of C'ochliosteiim is a matter of doubt. Again, the peculiarly dilated conncctire of many Tradescantias is clearly analogous to the cspanded filament of the plant under consideration. In conclusion, then, it m-odd appear from what has been above

* Payer, 1 Organogknie,' p. 663, t. 140. 205 DR. Af. T. NCISTERS 05 COCIILTOSTEMA. stated, that the andraxium of Cbch7iosfc,i~ndoes not differ in the number and position of its parts from that of other genera of tlie same Order. It does diEer in the order and manner in which the six elements are developed, in tlie congenital want of separation (union) of oue of the outer and tno oE the inner series of stamens, in the spirally tn isted anthers, in the petaloid outgrorrtli from the filaments, and in the derelopment at the back of the flower of that pecdiar tuft of hairs which in the adult floirer seems to foriii one of three staminodes, but which the course of development shows to be an acquired structure, long posterior to the true staniinodes in its development. It may be inferred froin this that the primitive or congenital number and arrangement of parts (in this case six in trro rows- one in front of the sepals, ouc in frout of the petals) is a fact of more n-eight than congenital irregularitp or congenital union (rrant of separation), and that the acquired 01’ adaptire characters (i. e. the spiral antherd, the petal-like hoods, and the posterior tuft of hairs) are, from a morphological point of Tievi-,altogether of secondary importance, greatly as they aEect the external ap- pearance of the floJrer in the adult state, and important as they are in all probability with reference to the physiological 1iistor~- of the flower. If these remarks aud the observations on which they are fonndecl be correct, they offer a styikiiig confirmation of the difference betvieeu, and the relative due of congenital and ac- quired characters, as ftxrther explained in my ~rorkon ‘ T-ege- table Teratology,’ and in a paper ou the “ Conformation of Passi- florace8 ” recently published in the Society’s ’ Transactions.’

ESPLASATIOS OF THE FIGCRES. Fig. 1. Earliest stage of flo~cr. 2. Shows the floral tubercle, with tho rodiiimit of tlic posterior sepal, s. 3. All thee sepals formed (s, s, s), the posterior one the largcst. 4. Posterior sepal turned back and the antero-lateral oues turned aside, to show the floral tiiberclc now triangular in outline. 5. Sepals turned back as before, to shon the tlmc petals (pzp~~)simd- taiieously dcrelopod. G. Section 1engt.hmise tlirongh n Ionng floner, sho\rii~gthe large posterior sepal. tno autcro-lateral sepals cut through, and. in the centre: indica- tions of the carpellary tubercles. 7. Side err of the same bud. 8. Bud. in nliich a portion only of the posterior eepal is shor;n : the petals are flattened; all six stameiis are non-iisible, three posterior oneslarge, VR. J. G. BIKER OX SCILLEX AKD CHLOBOGILEL. 200

thee Oihers smaller ; in tho centre are the1tlwee cxpellary tubercles as ret unclosed. 9. lndrmciuim of the same bud as 8, more highly magnified and seen from the back. 10. hclriecirim from a flower more fully adranced than 9, and sliorring the thrcc larger aud three smaller stamens surrouncling the pistil. 11. Back of the aiidrcc~iwfrom tho snme florrer as 10: p,13, the petals. 15. hichcecimn nncl ,zmium still further developed ; the antliers are now well dereloped. 13. IndrwSuui fiwn tho same Sorer 25 fig. 1.7, seen from the bncli. 14. Anthers beginning to sliorr the +ral tnist. 15. Andrcecium and pistil completed. 16. Posterior Tim of the same. 17. hdrcecium complete from another flower ; pwts forced open. 18. Andrieciuuti from tlie back shomng traces of the posterior staminode (?) at x. 19. Diagram of the flower. In the preceding figuros s stnnds for the sepals, p for the petals ; o indieates tho position of the axis, x tho situation of the postcrior appcnclage, and I the actual size of the orgau by the side of which it is plnecd.

Rerision of the Genera and Species of Scilles and Chlorogaleae. By 2. G. BIKER,Esq., F.L.S. [Xacl 1Iarch 7. 1872.1 THE present paper may be regarded as a continuation of the re- vision of the genera and species of Liliacere, the first part of which I had the lionour of submitting to the Society two years ago, and which mas priuted in the eleventh voliiine of the ' Proceedings,' pp. 349 to 436. In that part I dealt Kith the whole of the gamo- pliyllous capsular section of the order, exclusive of the tribe Aloinez. I aiii not prepared to suggest any better plan of classi- fying the capsular Liliacez than to arrange them, in the first; place. in trro series-one cliarncterizecl by n perianth Kith the di- risions more or less coiiiplete1~-uuited, and tlie other in which. the7 are free doxn to the base. If rre rtdopt this plan and take for the clingnostics of tribes the characters furnished by the nature of the rootstoclr and plan of arrangeiuent of the inflores- cence, regraiding a material difference in either of these tFo points as of sufficient iqortauce to constitute a distinct tribe, our tribes in the tn-o series Till run, to a large extent, parallel with. one anothey, there being pretty much the same range of vari at' ion in inflorescence and rootstock in the gamophyllous and polyphyl-