THE TRANSACTIONS OF THE LINNEAN SOCIETY OF LONDON. SECOND es ae . b~ VOLUME ES PART THE SIXTH. MISSOURI BOTANICAL GARDEN. LONDON: PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET: SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE ; AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER- ROW. Mboóo.uxxix. CONTENTS. PART VI.—January 1879. XX. On Mycoidea parasitica, a new Genus of Parasitic Alge, and the Part which it plays in the Formation of certain Lichens. By Davip DOUGLAS CUNNINGHAM, MS F.LS., Surgeon HA Indian Army . . . . . . +... page 301 XXI. On the Self-fertilization of Plants. By the Rev. GEoRcE HENsLOw, M.A., PLS IUE 1 uon WU Dcus hs EO Vou d XXII. List of Fungi from Brisbane, Queensland ; with Descriptions of New Species. By the Rev. M. J. BERKELEY, M.A., F.L.S., and C. E. BROOME, PELO FRR XXIII. On the Occurrence of Conidial Fructification in the Mucorini, illustrated by Choanephora. By D. D. Cunninenam, M.B., F.L.S., Surgeon H.M.’s Indian AME | 1 ig dou son oro A e os e AE Qe 7, ab, (8 8o i iud Sas Gur Bet bt r E ug, Omg. (vo. b. Ab [ 301 ] XX. On Mycoidea parasitica, a new Genus of Parasitic Alge, and the Part vu: plays in the Formation of certain Lichens. By Davip Dorota CUNNINGHAM M.B., FILS, Surgeon H.M. Indian Army. (Plates XLII. & XLIII.) Read June 21st, 1877. I. Introductory ; Descriptive Characters ; and .Life- History of Mycoidea. NOW that the recent researches of Cohn and others have directed attention to the . occurrence of Algal parasites, it may be of interest to record the occurrence in India of a form which not only appears to be the highest yet observed to play a truly parasitic part, but which possesses even greater points of interest in the nature of its characters as compared with Dia of its nearest allies, and in its relation to epiphyllous Lichens. At intervals during the past eight years I have met with specimens of the plant in various stages of development; but it was only during the course of the last six months that I have been enabled to work out the subject so as to obtain materials for a con- sistent narrative of its life-history. The first specimens were obtained on the leaves of a mango-tree in 1869. On these leaves an eruption of bright orange patches or pustules was observed. "These patches were at first sight regarded as of a fungal nature, but on close examination presented peculiarities of structure preventing them from being referred to any determined genus of Fungi. Subsequently other specimens were obtained on Rhododendron-leaves in the Nilgiris, on the leaves of species of ferns, Crotons, and various other plants in Caleutta, and on Tea-leaves in the Rumaun hills. 'The discovery of their presence on the latter host plant attracted renewed attention to their nature; and when they were again encountered in abundance in Calcutta as a destructive blight on plants of Camellia japonica, an opportunity was afforded of care- fully studying their real nature and modes of development. As to the specific identity of the Algæ occurring on all these various hosts, I am as yet not in a position to form a definite opinion, as, unfortunately, beyond drawings of one form of fructification, I have not preserved the materials necessary for comparison in many instances. With regard to the specific identity of the plant, in many cases there can be no doubt; and I am strongly inclined to believe that this holds good for all specimens yet obtained, however various the nature of the host. If this really prove to be the case, it is, no doubt, remarkable, and in a sense lowers the parasitism of the Alga. One important feature common to most, if not to all, of the leaves affected was the existence of a firm coriaceous texture and a thickened epidermal covering. Leaving the discussion of any further general questions until a detailed description of the species observed on the Camellia has been given, I shall now proceed to describe the structure and life-history of the parasite, and the nature of the lesions which it produces in . SECOND SERIES.—BOTANY, VOL. I. ; 2U S 302 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA, its host. On examining one of the affected plants, the injuries due to the presence of the parasite are visible at once (Pl. XLII. fig. 1). The leaves in varying number, according to the length of time which the plant has suffered, and dependent on the time of year and the nature of the weather, are seen to be irregularly eaten away at the margins, penetrated by circular patches of decay or of absolute loss of substance throughout their surfaces, or spotted with circular pustules of various sizes and various shades of colour, ranging from pale green to bright orange. On examining the pustules more closely, they are found to be slightly elevated, and of a more or less distinctly marked radiating structure ; and on looking at the margins of the holes and other deficiencies in the laminz of the leaves, they are generally found to be bounded by a distinct rim of like colour and strueture. Af certain seasons of the year numerous very minute, elevated, green spots and an abundanee of barely perceptible orange specks will also be encountered scattered over the surface of the leaves. The pustules and spots are almost invariably confined to the upper surfaces of the leaves; but in some rare instances they may be found on the inferior surfaces also. The latter phenomenon, however, is generally due to the extension of a pustule spreading round the margin from above. The size of the patches naturally varies greatly with the age of the plant causing them. On an average, patches which have not yet begun to die off in the centre do not exceed 5:0 to 6:25 millims. in diameter. The extent to which the leaves are affected varies greatly, but in advanced cases is sufficient very seriously to disfigure the plant. The very young leaves are seldom affected, owing, no doubt, in great part to the slow growth of the parasite. This is a matter of considerable economic importance in connexion with any form of parasitic growth affecting the Tea-plant. . On removing one of the pustules (which is easily done by the aid of a sharp knife), a brown discoloured patch of leaf-tissue is exposed, corresponding closely with the site of the pustule, and of various degrees of depth of colouring from a reddish brown to blackish. These discoloured patches in many cases penetrate the entire thickness of the leaf, and render the distribution of the Alga recognizable on the lower surface. In some instances the pustules are uniform in colour throughout, and end in a smoothly rounded margin; butin others the centre of the patch is pale, surrounded by a ring of deeper colour, or the margins, in place of being even, are more or less distinctly divided into a series of irregular radiating lobes. Even to the naked eye the appearance of the pustules varies much in different instances—in some being smooth and comparatively uniform, in others. covered with a crop of erect orange-coloured filaments, or variegated with minute orange points and specks. The structure of the patches always points to a peripheral growth from a central point; but their appearance varies greatly with the ` season of the year and the nature of the weather. | Commencing with the rains (the season when the vegetative growth of the plant is at a maximum), we find the following to be the anatomical features presented by fully developed specimens. They are then covered with a dense crop of erect filaments of various tints of orange and green. These filaments on close examination may be seen to proceed from beneath the epidermis of the host plant, and to arise from a flattened cushion of mediating filaments situated there, pp are net ane divided into a A NEW GENUS OF PARASITIC ALGA. 803 series of cells by transverse septa, and terminate superiorly either in blunt points or in more or less developed spherical capitella bearing a few large spore-like cells on short curved processes (Pl. XLII. fig. 2). The appearance of a mass of such filaments, with their transparent glassy walls and brilliantly coloured contents, is strikingly beautiful. Besides such filaments, numerous older ones are seen—some prostrate and colourless, others still erect and with coloured contents remaining in their cells and capitella, but with only empty spore-cases adherent to the latter. Some with more or less developed heads are also encountered, which are distinguished by the presence of greenish rounded masses adhering to them laterally, whilst in many the cell-contents, in place of being of an orange colour, are vividly green. On detaching such a pustule from the leaf, and examining it under higher powers, it is found to consist of a flattened subepidermal disk of radiating filaments, from which numerous ascending twigs arise, which, breaking through the epidermis, are developed into the fructifying filaments just described. Careful vertical sections of the leaves, such as may be obtained by means of paraffin as an embedding medium, show very clearly the arrangement of the various portions of the pustule and its relations to its host (Pl. XLII. fig. 8). Before proceeding further, however, it may be well briefly to describe the structure of the mature Camellia-leaf. On examining a good vertical section we find the entire thick- ness of the leaf to measure on an average about 0°3125 millim. Proceeding from above downwards, there is first a very dense layer of epidermal cells. These are colourless, flattened, and on treatment with iodine frequently show a very distinctly stratified structure in their walls. They have sinuous margins laterally and beneath, by means of which they dovetail into one another and into the irregularities of the subjacent stratum. This layer measures about 0:0100 millim. in its thicker and 0:0075 millim. in its thinner portions. The cells of the next stratum are also very thick-walled, and appear . on section as a series of cubes. They form a continuous stratum, about 0:02 millim. in thickness, and are distinguished by the small amount of their contents, and by hardly ever showing the presence of any chlorophyll. As a rule, in fact, they appear almost empty, containing only a peripheral layer of protoplasm, or in rare instances a small oil- globule. Beneath this comes a stratum composed of from one to three rows of prismatic elongated cells, measuring about 0°050 millim. in long diameter, and full of chlorophyll granules. The continuity of this stratum is interrupted by the intercalation of a series of very remarkable cells at uncertain intervals. These are the remains of a system of ‘cells which at one stage in the development of the leaf form the greater part of its substance, and are gradually thrust aside and separated by the development of the common cells of the leaf-tissue. They are of great size, in some cases attaining a length ` of 0°25 millim., and a diameter of 0:035 millim. at their wider basal extremity. They are rendered very conspicuous by the great thickness of their walls, as well as by their colourless finely granular contents. They are usually only found proceeding from the upper towards the lower surface of the leaf, being inserted immediately beneath the subepidermal layer; but occasionally they occupy a reverse position, and are then - inserted on the inferior subepidermal layer. They are broad at the extremity of 2v2 304 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA, - insertion, and frequently expand there into one or two blunt diverging processes, giving them a somewhat funnel-shaped aspect on section. They taper off from the base, and sometimes divide into two or three diverging branches, ending freely among the loose tissue of the interior of the leaf in blunted tips. Their thick walls appear to be channelled by a number of minute canals, leading from the cavity of the cell to its exterior surface ; and the cell-contents consist of a colourless fluid full of minute granules. Beneath the stratum of prismatic cells is the loose tissue of the centre of the leaf, con- taining air-spaces and an abundance of large spheraphides of oxalate of lime; and this is succeeded by one or two layers of tabular or cubical cells, and finally by a thin layer of flattened epidermal cells. There are no stomata on the upper surface of the leaf; but they occur abundantly below, ranging from 1000 to 1600 per square inch. On examining a vertical section of a leaf, including young pustules of the parasite, appearances similar to those shown in the drawing are observed (Pl. XLII. fig. 8). The filaments of the disk are seen to lie between the epidermis and the subepidermal layer of cells. According to the degree of development of the patch, the epidermis over it is more or less elevated and separated from the tissue below. In a very young patch, such as that to the left hand of the figure, the displacement is comparatively slight; but where an abundant growth of ascending filaments has taken origin from the disk, it becomes very considerable, and may amount to as much as 0:125 millim. or even more. Simultaneous with and proportionate to the displacement of the epidermis, changes begin to occur in the subjacent cells of the leaf. A certain amount of sclerosis of their walls appears to take place ; but the most conspicuous and considerable alteration occurs in the nature of their contents, which become gradually thickened and discoloured, passing through various shades of yellow and brown, and ultimately assuming a dry granular consistence and bright burnt-sienna colour. These changes at first only affect the layer of cells immediately beneath the Alga; but as time goes on they gradually advance deeper and deeper into the substance of the leaf, until they extend throughout its entire thickness, the affected area corresponding closely with the algal disk. As the disease advances the filaments in the centre of the disk become emptied of their contents and die; they then dry up, together with the portion of leaf-substance beneath them ; and the dry and withered slough, readily breaking off, leaves a hiatus in the leaf, which, according to its situation, appears either in the form of a hole punched through the surface, or as an irregular space eaten out of the margin. Whilst this destruction of the centre of the patch is taking place, the parasite continues to extend peripherally ; so that in many instances a great portion of the leaf is ultimately destroyed. The filaments of the algal disk appear, as a rule, to be confined between the epidermis and the sub- epidermal layer; occasionally, however, branches are given off which force their way downwards between the cells of the latter and reach the prismatic layer beneath (Pl. XLII. fig. 8). The disk is originally composed of a single layer of dichotomous filaments, which force their way outwards, between the epidermis and the subepidermal cells. They are thick-walled, and at this time (so long, at all events, as they are in active growth) are filled with a bright green protoplasm. Whilst the disk continues: to increase at the A NEW GENUS OF PARASITIC ALGA. 305 margins by means of the dichotomous growth of the filaments, ascending branches are given off by many of the cells of the older central portion; and these, forcing their way upwards, tend still further to elevate the epidermis. From the dense resistant nature of the latter they are generally more or less bent laterally, and in certain instances may even come to follow a course almost parallel to that of those of the disk from which they arise. As they ascend towards the epidermis their contents, in place of retaining their bright-green colour, become first yellowish and then rich golden orange. Some of them become enlarged above, and, again ascending, finally penetrate the epidermis and appear on the surface of the leaf. On examining a patch at this stage it is found to be sprinkled over with erect blunt-pointed filaments, which generally project in small groups through openings in the epidermis (Pl. XLII. fig. 3). This grouping of the aerial fila- ments is in many cases due to several of them arising at the termination of one of the subepidermal ones, but sometimes appears to be owing to filaments from several different origins making their exit through a common rupture in the epidermis. ‘They continue to increase by apical growth and the gradual separation of cells by transverse partition, and ultimately attain a height of LO millim., with a breadth varying from 0:05 millim. at the base to 0:025 millim. towards the apex. The terminal cell now, in place of re- maining pointed, begins to swell out, and to assume a clavate, and ultimately a more or less spherical figure, forming a rounded capitellum about 0*1 millim. in diameter. A great accumulation of orange protoplasm now occurs within it; and a number of smaller clavate processes arise from it, into which much of the contents passes. These processes in course of time become developed into oval spore-like bodies, supported on narrow curved stems; and the process is completed by the accumulation of the greater part of the protoplasm in the former and the formation of septa between them and the partially emptied stems, which remain adherent to the capitellum (Plate XLII. fig. 4). The growth of a filament may cease permanently here; but in many cases, after the formation of the capitellum, the cell immediately beneath it begins anew to grow, shoot- ing upwards at one side of the capitellum and forcing it aside. In the course of deve- lopment this displacement advances so far that the new portion of the filament comes to lie more or less in the axis of the older one, and appears as a direct continuation of it ; whilst the old capitellum, bearing the remains of the spore-cases, and containing more or less orange or green protoplasm, appears as a lateral swelling, attached to the filament in its course (Pl. XLII. fig. 2). This further development, however, is by no means uni- versal; and in no instance has it been seen to proceed beyond the formation of a second head. It occurs most frequently during the height of the rains, when the leaves are ‘almost constantly wet and all the vegetative processes of the parasite at a maximum. Although the growth of the filaments appears to be apical only, there are in many instances evidences of what may perhaps be a tendency to intercalary growth; for, on examining the cells under high powers, processes may frequently be seen projecting from the walls into the cell-cavity, and in some cases proceeding so far as to give rise to marked constriction of the protoplasmic contents (Pl. XLII. fig. 7). When under what may be regarded as normal conditions, the contents of the filaments and spores are of a brilliant orange colour and granular consistence, and at once strike a deep blue or black 306 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA, with solutions of iodine. When, however, the specimens are submerged, or when, during a continuance of wet weather, they are constantly saturated with moisture, the orange-colour is gradually replaced by a bright green, and the contents eventually come to resemble exactly those of the subepidermal disk-cells. The mature spores, if they may be so termed, vary somewhat in size; one of medium size, which was measured, was 0°0379 x 0°027 millim. Their contents appear at first as a uniform granular mass; but as time goes on a process of segmentation occurs in this, ultimately dividing it into from twelve to twenty-four, or even more, oval bodies; and whilst this process is being completed a rounded orifice forms in the thick wall of the spore (Pl. XLII. figs. 4, 5). This orifice is generally situated laterally, but occasionally occurs towards one or the other end of the spore. "When such ripe spores are wetted by the addition of water, by dew, or by rain, active swarming rapidly commences among the included oval bodies ; and after this motion has continued for a short time, they separate from one another, and emerge in rapid succession through the orifice in their mother cell as active zoospores. These are at first pear-shaped, measuring about 0:00825 x 0:0052 millim., and are provided with a couple of long slender flagella, arising from the ante- rior pointed extremity (Pl. XLII. fig. 6). This extremity is almost colourless; but the rest of the body is of a reddish-orange colour by transmitted, and greenish by reflected light. They swim about actively for a short time, and then gradually become spherical and cease to move. The minute anatomy and subsequent history of these zoogonidia is identical with that of those developed in the sexual fructification, and will be further referred to in describing them. In some cases all the zoospores do not escape, one or two remaining behind in the mother cell and there undergoing various changes (Pl. XLII. fig. 5). Among the commonest of these is one in the course of which they gradually become circular green cells. The process of the formation of zoospores is at any time liable to be arrested by the addition of excessive moisture, which, as in the case of the filaments, causes the contents of the mother cells to become green, even after the process of division has advanced so far as to have caused the formation of distinct masses. Such green cellules do not usually appear to be capable of movement, but become gradually invested by distinct cell-walls, and remain within their mother cell until freed by acci- dental violence or by a process of gradual decay. "They are met with in great abundance during the height of the rains; but the age events i in their history have not been observed. After the escape of the zoospores from their mother ecliar $ the latter remain as sidity colourless appendages adherent to the capitellum. The filament, if it does not undergo further development, remains for some time erect, and then, falling over, lies prostrate on the surface of the pustule. The number of zoospores produced in this way is very - great during the continuance of moist weather, and amply suffices for the propagation of the parasite at such times. Towards the close of the rainy season, however, the formation of filaments gradually diminishes, and, as the cold and dry season advances, ultimately ceases altogether. The patches of the parasite are now entirely subepidermal, and hardly extend at all peripherally. They are of a bright orange colour; and many of ` 3 them dry up entirely and die off. Were there no special Mm c securing the pre- E A NEW GENUS OF PARASITIC ALGA. | 307 servation of the parasite under these circumstances, the amount of it surviving from one season to another would be comparatively small. . Such an arrangement is, however, provided in the sexual fructification. During the height of the rains, and whilst vegetative growth is actively progressing, only the asexual fructification is produced; but subsequently the sexual form begins to appear, and, gradually increasing in abundance, eventually more or less completely replaces it. The sexual organs, in place of taking origin from the aerial filaments, are developed on those of the subepidermal disk. These filaments, in place of containing an abundance of brilliant green protoplasm, as at first, begin to assume a yellowish tint, and ultimately come to contain only masses of globules and granules of as vivid an orange colour as that of the contents of the aerial branches (Plate XLII. figs. 9, 10). Whilst this change of colour is taking place, the contents also become greatly condensed and contracted, so as in many cases to be reduced to one or two isolated masses, or to a mere band along the centre of the cell. Their coarsely granular consistence also disappears; and a number of separate globules of large size and oily aspect are ultimately formed. Whilst these changes are advancing, the organs destined for sexual fructification begin to appear. Certain of the filaments, in place of, as before, continuing to grow by a process of dichotomous division, resulting in the formation of two nearly similar branches, give origin to only one filament at the site of division, whilst the other member of the dichotomy, in place of elongating, swells up into an obovate dilatation (Pl. XLII. fig. 11). This is sometimes sessile, but is generally situated on a short process of the mother cell. A septum now forms at the base of. the dilatation ; and the latter rapidly increases in size, and becomes filled with a great accumulation of orange protoplasm. The new cell now appears as a large thick-walled sac inserted between the neighbouring filaments of the disk, which are displaced laterally by its growth, and, curving along its margins, come again into contact at its distal extremity. The thickness of the cell-wall is very considerable, amounting in many cases to as much as 0:004 millim., and frequently shows distinct evidences of stratification. "The cells, when mature, vary considerably in size ; but average specimens may measure about 0-0625 x 0:0415 millim. Due to the dense nature of the disk, to its subepidermal site, and to the fact that, when detached from the leaf, only retrograde changes, tending to a recurrence to pure vegetative growth, occur in the developing fructification, I have been unable continuously to follow out the further steps in the development of these cells, or oogonia as they now are. In so far, however, as very numerous examinations of separate specimens are capable of throwing light on the matter, the following appears to be the order of events. The contents ot the oogonia, which were at first in close relation to the walls of the cell, become gradu- ally removed from them as the cell enlarges, and form an oospheric mass, separated from its case by a distinct interval, save towards the basal extremity, immediately over the septum dividing the oogonium from its mother cell. Whilst these changes have been occurring in the oogonia, numerous slender-branched filaments have arisen from the neighbouring cells of the disk. Some of these become dilated at the extremity ; and the large terminal cell becomes applied and closely adherent to an oogonium (Pl. XLII. fig. 12). These filaments appear, as a rule, to arise from the under surface of the disk ; 308 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA, and those which are developed into pollinodia are usually attached to the oogonia towards their bases. The contents of the terminal adherent cell appear next to be emptied into the oogonium, and to blend with the oosphere. Owing to the reasons previously men- tioned, this process has never been actually observed to occur; but the contents of the ` pollinodial cells in many eases almost entirely disappear, and many examples of oogonia have been met with containing a mass of protoplasm independent of, or only partially blended with, the oosphere. The oosphere now begins to show a distinct cell-wall, and is soon converted into a large spherical or oval oospore, which lies free in the cavity of the oogonium (Pl. XLIII. fig. 2). The cell-wall is at first thick and soft, but gradually becomes very thin and delicate. Another series of changes has been occurring in the oogonium itself. Its position, which, like that of the filaments of the disk, was originally horizontal, has been gradually exchanged, in the process of growth, for a more or less vertical one, the base of the organ being depressed beneath the plane of the disk, whilst its apex ascends and approaches the epidermis of the leaf. Its shape also exhibits alterations, frequently becoming more or less acuminate at the apex, and losing its original smooth obovate outline. Changes also begin to oceur in the walls. At one point or other towards the apex a thinning of the wall begins to occur ; and this process, confined to a limited area, gradually advances until the entire thickness is perforated, and a circular opening of considerable size and sharply-defined outline (by means of which the interior of the oogonium communicates freely with the subepidermal space) is formed (Pl. XLIII. fig. 2, & Pl. XLII. fig. 14). With this the development of the oogonium itself ceases. In many instances they remain in this condition; but in others the mature organs are more or less completely invested, save around the ostiolum, by a loose coating of fine filaments, similar in ap- pearance and origin to those bearing the pollinodial cells. In any case the original filaments of the disk now die off, and the oogonia, with their contained oospores, are left persistent among the dried-up débris of the disk beneath the thick epidermis of the leaf. How long such oospores may retain their vitality in this condition has not been definitely determined; but that they do retain it for a considerable time is certain, as they have frequently been obtained in a lively condition from the under surface of dried-up flakes of epidermis on the site of old patches of the parasite—where hardly any other evidences of its previous presence persisted, and where the leaf-tissues among which they were encountered were thoroughly dried up and destroyed (Pl. XLIII. fig. 1). i The mature oospores are spherical, and on an average measure about 0:035 to 0-031 : millim. in diameter. Sooner or later, when exposed to favourable circumstances, the last _ events in the course of their development occur. The dried-up epidermis eventually cracks, and in doing so leaves channels of communication open between the external surface of the leaf and the interior of the oogonia, as the open mouths of the latter communicate, as before mentioned, with the subepidermal space, and in many cases, after the rupture of the epidermis has occurred, even project free on the surface (Pl. XLII. fig. 15). On the addition of moisture the mass of the oosporic contents now breaks upintoa multitude of oval bodies like those developed in the aerial spores; and, the amount of fluid being — A NEW GENUS OF PARASITIC ALG. — 309 sufficient, these soon commence to swarm actively, rupture the thin wall of the oospore, and, escaping in succession through the ostiolum of the oogonium, swim off freely into the subepidermal space and over the surface of the leaf. In size, colour, and general ` appearance, these zoospores are precisely similar to those developed in the asexual fruc- tification. Like them, they are provided with two elongated flagella, one of which, so long as the body retains its original oval form and active motion, is usually trailed behind, whilst the other is employed in progression. Occasionally they become adherent to one another, or to other solid bodies, by their trailing filaments. Sometimes a pair of them may be seen to become adherent to one another by the extremities of these filaments, and, continuing their rapid rotating movement, to twist them so tightly up as to bring their bodies into close contact with one another. Under such circumstances, an actual fusion or conjugation appears in some cases to occur, resulting in the formation of one large biflagellate zoospore. Such a phenomenon, however, is very rare, and seemingly accidental. The zoospores, when first emitted, are of an oval or pear-shaped form, and average about 0:00825 x 0°0052 millim. in size. As they become less active and begin to change their form, they frequently increase in size considerably. Eventually they cease to move and become spherical, the flagella continuing to vibrate for some time, and, after becoming motionless, persisting for a considerable period as two delicate hair-like fila- ments, attached to a portion of the cell, which, by its comparative freedom from colour- ing matters, indicates the site of the original hyaline extremity. When examined under a high power at this time, the cells sometimes appear to be invested by a delicate halo of gelatinous matter (Pl. XLII. fig. 16, a). They are provided with a very delicate mem- branous wall, and contain a fluid with numerous reddish granules, and a mass of green- ish colour investing from ten to twenty, or even more, oval particles of considerable size. When confined beneath a cover-glass, the greater number of the cells next undergo the following changes :—A swarming movement begins among the reddish granules of the cell-fluid ; this attains a great intensity, the cell-wall bursts, an escape of granules occurs, and the green mass or globule is partially protruded from the cell. The body now pre- sents an appearance similar to that shown in the figure (Pl. XLII. fig. 16, 6). After remaining for some time in this condition, the oval particles within the green mass begin in their turn to swarm, and, ultimately escaping from it one by one, swim off and are dispersed in the fluid, leaving the green mass adherent to the fragments of the original cell. At the time of exit the oval particles measure about 0:001 x 0:0005 millim. Under normal circumstances and favourable conditions, however, many of the zoo- spores proceed to give origin to new plants of the parasite. The various stages in deve- lopment may be traced by means of specimens which may be obtained in abundance from the surface of the leaves, near patches of the parasite containing mature oospores, and which have been exposed to sufficient moisture. On examining such portions of the leaves under a low power, or even, in some cases, with the naked eye, they may be seen to be sprinkled with minute yellowish or orange specks. These are readily detached from the surface, and consist of the spores and young plants derived from them. The spores, on ceasing to move, become spherical and increase slightly in size. The. con- SECOND SERIES.—BOTANY, VOL. I. 2x 310 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA, tents melt into a homogeneous mass of an orange or sometimes of a green colour, and a certain amount of thickening of the cell-wall occurs. The cell-wall and coloured con- tents now become separated by a distinct interval. The coloured mass next begins to assume a lobed or more or less distinctly cruciate outline, due to the development of marginal projections; and these continuing to grow, and becoming divided. dichoto- mously, a flattened disk or rosette is gradually formed (Pl. XLII. fig. 17). Whilst this division of the coloured contents is advancing, processes are developed from the cell- wall, which, shooting inwards, and giving off branches as the division of the contents progresses, ultimately coalesce with one another, and divide the disk into a series of cells arranged in a radiant fashion. The formation of new cells now continues by the repe- tition of the process described above; and as the original cell-wall of the spore does not rupture, for some time at all events, but appears to stretch with the development of its _ contents, a series of coherent, flattened, cellular disks are gradually formed. Although the septa marking out the constituent cells are at first common to their adjacent cavities, they ultimately divide into two layers, so as to transform the originally continuous disk first into a series of segments, and ultimately more or less completely into radiating filaments, which, although remaining closely in contact, may be separated from one another, and in some cases even broken up into their constituent cells. These disks vary in size from mere points to 0-4 millim. in diameter, or even more in some cases, and adhere closely to the surface of the leaves (Pl. XLIII. figs. 3, 6). These disks differ from those of the mature plant both in their position, which is superficial to, not beneath, the epidermis of the host, and in their much denser structure (Pl. XLIII. fig. 4). In order conveniently to distinguish the two sets of disks from one another, those formed directly from the germinating zoospores may be termed germinal or primary disks. We have next to follow the steps by means of which the primary disks come to be replaced by the subepidermal disks of the mature plant. Many of them never are so replaced, but, after persisting for some time, dry up and disappear from the surface of the leaves, or are utilized by parasitie fungal elements in a fashion which will be subsequently described. Were it not for these limits to the spread of the disease, the destruction of leaf-tissue would be incalculably greater than it is; for the number of primary disks originally formed is often excessive, not unfrequently being generally dif- fused over the entire surface of the leaf. It is a matter of some difficulty to follow the progress of development in those specimens which pass beyond this epiphytic condition ; but a series of horizontal and vertical sections shows the process to be of the following nature. Some of the cells of the primary disk, in place of merely growing outwards by the formation and separation of peripheral lobes, give origin to buds from their under surface, which gradually penetrate the thickened epidermis, and ultimately reach the subepidermal space, or, more correctly, the line of separation between the epidermis and the subepidermal layer of cells (Pl. XLIII. fig. 7. Having done so, they take on an active growth, and, forcing their way along beneath the epidermis, and dividing dichotomously as they advance, soon form a mass of radiating filaments. Owing to their mode of origin, these filaments never occur in such regular and coherent disks as the primary disks; but, as they continue to spread and to become crowded upon one A NEW GENUS OF PARASITIC ALGA. nag 311 another, they form dense masses of radiating structure; such as are found in the mature plant. Under favourable circumstances, large numbers of such secondary disks are formed, and appear as elevated green patches and spots on the surface of the affected leaves. The primary disks cease to grow, and, after remaining for some time recog- nizable as small brownish patches on the surface of the epidermis of the new plant, gradually dry up and disappear. The relation of the primary and secondary disks to one another, and to the epidermis of the leaf, may sometimes be very clearly determined in transverse sections (Pl. XLIII. fig. 5). When the weather is dry the young plants, which are originally bright green, soon assume.an orange colour; and they then remain dormant and protected by the epidermis until favourable conditions of temperature and moisture rouse them to go on to increased growth and the ultimate development of the various forms of fructification. : | The various processes described in the previous pages seem to constitute the most important features in the life-history of this plant; and it now remains to consider some more general questions regarding it. As to its truly parasitic nature there can be little doubt. The situation of the mature plant, within the tissues of its host, and protected by a thick and highly cuticularized epidermis, would, even at first sight, lead to the conclusion that it was dependent on these tissues for its nourishment ; and this conclu- sion is confirmed by the destructive effects which it produces. Whilst its essential parasitism is thus rendered clear, it remains an open question, how far the Alga makes use of the organized materials of the tissues for its nutrition, and to what extent it pro- duces its prejudicial effects by merely appropriating inorganie elements of nutrition normally destined for the tissues of its host. That it acts, mainly at all events, in the latter way is rendered probable by the very various nature of the host plants on which specimens are found to occur; and much of the destructive effect which it produces may be ascribed to the large amount of water which it draws off whilst in active growth. In regard to the precise nature and alliances of this Alga, there are several points calling for consideration. So far as the vegetative growth is concerned, it agrees very closely with the genus Coleochete, the primary disks resembling the disks of C. scutata, whilst the secondary ones approach in characters those of some of the more loosely branched species. There is, however, an entire absence of the bristles characteristic of Coleochete. When we come to consider the reproductive organs, the resemblance ceases, and very striking differences make their appearance. The asexual zoospores, in place of being produced indifferently in any cells of the disk, are here deve- loped only on highly specialized filaments; and the sexual fructification, in place of being a well-developed carpospore, the result of fertilization by motile antherozoids, is rather an oospore than a carpospore, and is the result of fertilization by means of anthe- ridial filaments. There is never a formation of true carpospores, although in many cases a tendency to such formation is indicated by the investment of the fertilized oogonium by a mass of cellular filaments. So in regard to the ultimate development of the sexual process, in place of the formation of a cellular mass, the individual cells of which give origin to zoospores, there is here an immediate resolution of the contents of the oospore 2x 2 312 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA, into zoospores, and the subsequent development of each of these into a compound cellular body—the so-called primary disk. | In certain respects it appears to be more closely allied to the genus Phycopeltis than to any other. The development of the zoospore into the primary disk is of the same nature as that by which the disks of Phycopeltis are formed ; there is the same absence of setz, the same colouring of the cell-contents, and the habitat (on the leaves of living plants) is similar. It is, however, distinctly marked off from that genus by the deve- lopment of the secondary subepidermal disk, and by the peculiar arrangement of the cells which produce the asexual fructification. In spite of the peculiarities which it presents, the close relation of this Alga to Coleo- chete and the allied genera is unmistakable; and all the departures from the ordinary type which it presents are explicable as the result of modifications adapting it to its parasitic habit and peculiar locality. The production of zoospores by the common cells of the disk would be useless, where such zoospores would be imprisoned beneath the epidermis of the host, and unable to find a fit site for their further development. So in regard to the sexual fructification; a process of fertilization by motile antherozoids is replaced by one effected by means of pollinodia, where the sexual organs, in place of lying free in a fluid, are buried in the tissues of a host plant. MYCOIDEA, nov. gen. Thallus entophyticus, disciformis, e filis articulatis formatus et filis erectis, aeriis, sporiferis przeditus. Cytioplasma aureum aut viride. Propagatio zoogonidiis et oosporis fit. Zoogonidia in cellulis matricalibus, que extremis filorum aeriorum superimposite sunt, formata. MYCOIDEA PARASITICA, Cunningham. Generis que differentia ezedem speciei. Zoogonidiis numerosis, diametro 0°00825 x 0:0052 m. m. Oosporis globosis. Habitat, Indize Orientalis, in fruticum et arborum foliis. Il. Its relations when attacked by Parasitic Lichens and Fungi. The first part of this paper has been occupied with the description of the characters and life-history of this Alga per se; it now remains to give some account of ‘its relations as a gonidia-former when in its turn subjected to the attack of parasites in the guise of fungal filaments. As far as my observations have yet gone, it may enter into the formation of various species of lichens, according to the stage of development in which it is attacked; but at present attention will be confined to one species, the development of which has been comparatively fully worked out, and which occurs on the leaves of Camellia japonica, of Nephelium, and of various other plants, all of which are — to the attacks of the Alga. It has been already mentioned that only a relatively small number of the primary disks give origin to subepidermal filaments, and that many of them dry up and disappear A NEW GENUS OF PARASITIC ALG&. 313 after having attained a more or less considerable size. A still larger number, however; are utilized by fungal filaments, and in association with these go to the development of patches of a heteromerous lichen. The leaves on which this occurs begin, shortly after the onset of the dry weather of the winter months, to show conspicuous dry superficial patches of a grey colour, and which, on close inspection, are seen to be composed of colonies of minute whitish circular disks (Pl. XLIII. fig. 8). In some cases almost the entire surface of the leaf is covered; but in general the groups are more or less isolated, circular, and mingled with distinct patches of the Alga in various stages of growth. On moistening such lichenoid patches the grey colour disappears in great part; and the separate disks now appear as circular green spots shining through a semitransparent veil, which invests them and unites them to one another. The patches come in course of time to be besprinkled with minute elevated black specks, and ultimately show a greater or less number of flattened circular apothecia, with black raised margins and brownish contents (Pl. XLIII. fig. 9). On removing such a patch (a process which is readily accomplished, as it adheres but slightly to the surface of the leaf), it is found to be composed of a basis of delicate colour- less membrane, passing continuously over the grey disks and the intervening spaces, and in many instances readily separable from the disks save towards their centres. On examining such a detached flake of membrane and its attached disks, the former is found to consist of a dense web of empty interwoven hyphze, whilst the latter are readily recognizable as primary disks of the Alga, which have undergone greater or less modi- fications (Pl. XLIII. fig. 10). The cells of the disk are in great part empty and colourless ; but beneath the colourless framework or skeleton is a great mass of circular cells. These are quite distinet from one another ; and many of them become detached from their places in the process of preparing the specimen, and may be found floating free in the sur- rounding fluid (Pl. XLIII. fig. 18). The larger number of them, however, remain in situ, and appear to be embedded in a granular matrix. Entangled in the flake, in some cases, other primary disks may be encountered, which are as yet completely unaltered, or exhibit various stages of transition to those just described. The black specks in the patch are now resolved into spermogonia full of minute spermatia, and the apothecia present all the characters of those of gymnocarpous lichens (Pl. XLIII. figs. 9 & 10). The following account of the details of structure and the history of development of these lichenous patches is founded on very numerous careful examinations of specimens . in every stage of formation. Before any of these lichenous patches had appeared, and whilst the study of the development of the primary disks of the Alga occupied attention, it was frequently observed that fungal filaments had become attached to the latter (Pl. XLIII. fig. 12). These filaments were colourless, ramified over the surface of the leaf, and, where they came into contact with the disks, were frequently bent at right angles to their previous course, becoming closely applied to the margins of the disks, adhering firmly to them, and sending numerous branches over their surfaces. On following out the subject more closely in connexion with thelichenous patches, the subsequent history of these adherent filaments was ascertained to be as follows :—After becoming adherent to the disks the hyphz proceed to ramify abundantly, and eventually form the densely 314 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA, interwoven film of empty filaments (corresponding with the cortical layer of other lichens) which covers the disks and the intervals between them (Pl. XLIII. fig. 19). Whilst this film is forming, changes occur in the disk beneath. The cells multiply greatly by the formation of tangential septa dividing the rays into rows of short spaces; and their contents pass from orange into bright green. Each of the cells now buds out ` below, and forms a rounded prominence on the inferior surface of the disk; and these buds, gradually absorbing all the contents of their parent cells, are ultimately detached as independent bright green circular cells, and form accumulations beneath the now empty framework of the disks (Pl. XLIII. fig. 16). The process begins from the centre of the disks, and gradually extends peripherally until the disks are left as empty cellular shields covering a mass of detached circular cells. The hyphæ of the cortical layer, after ramifying over the upper surface of the disks, finally force their way downwards between the constituent cells of the latter towards the central point from which the rays diverge. At the point of entrance a dense cellular mass of fungal cells is developed, eausing an appearance, very frequently to be observed in the disks of the mature lichen, of a central colourless space in the groups of gonidial cells (Pl. XLIII.fig.14). Having gained an entrance, the hyphs then ramify among the projecting buds on the under surface of the disk, and, becoming adherent to them, retain them in position when freed ` from their parent cells. The green gonidial cells continue to multiply by the division of their contents, and frequently form several strata, beneath which a colourless layer of hyphee is more or less distinctly developed. Owing to the ease with which the lichenous patches are detached from their position on the surface of the leaf, it is a matter of some difficulty to obtain good demonstrations of their structure; but the accompanying illustrations show the appearances present in successful vertical sections, as well as in cases in which masses of gonidia and hyphæ have been teased out from beneath their investing disks (Pl. XLIII. figs. 11, 17). | I At various points of the patch the filaments, sometimes those of the film, and more rarely those apparently of the intergonidial hyphæ, give origin to an inferior layer of loose tissue. This rapidly multiplies and forms dense whorls of tangled filaments, from which the spermogonia and apothecia are gradually developed; and it is only around these structures that any loose tissue corresponding to a medullary layer can be encountered (Pl. XLIII. fig. 20). | The spermogonia, as previously mentioned, are developed anteriorly to the apothecia. They ultimately appear as slightly elevated circular spots of a greenish-brown hue, measuring 0°175 millim. in diameter, and discharge their contents on the surface of the patch through minute rounded ostiola. The spermatia are colourless, slightly curved, and measure 0:004 by 0:001 millim. (Pl. XLIII. fig. 13). The apothecia measure about 0-7 millim. in diameter by 0:2 millim. in height, and appear to the naked eye as minute blackish cups on the surface of the patch. They are invested by a distinct rim, derived from the cortical layer, and show the usual features presented by the apothecia of gymnocarpous lichens—being anatomically separable into excipulum, subhymenial layer, - hymenium, and nucleus, consisting of the remains of the whorls from which they are developed (Pl. XLIII. fig. 15). The asci contain eight curved uniseptate spores, and are A NEW GENUS OF PARASITIC ALGE. | 815 surrounded by an abundance of slender somewhat capitate paraphyses (Pl. XLIII. fig. 21). I am uncertain whether any of the spores yet obtained were mature; but those showing most indications of being so were colourless or very pale yellow, somewhat con- stricted at the septum, and measuring 0:0227 x 0:0075. Numerous experiments were made as to the reaction of the asci with solution of iodine. In no instance was there any distinct indication of blue staining as encountered in most lichens observed. The walls of the asci, on the contrary, appeared to be unaffected į but the presence of a large quantity of the material styled epiplasma by De Bary was demonstrated around the spores, which, under such circumstances, appeared as pale yellow bodies embedded in a deep-red-brown basis. i The history of the development of this species of epiphyllous lichen appears in itself to afford a complete demonstration of the composite nature of such structures. There can be no doubt here as to the source of the gonidial cell, and that that source is an Alga capable of independent existence and of producing perfect forms of fructification. On the other hand, it is only in association with these algal elements that the fungal fila- ments form a tissue capable of giving origin to spermogonia and apothecia. Various other points might be touched upon ; but in the mean time I would merely suggest that the anomalous subepidermal site and so-called cephaluroid conditions of the various species of Strigula are probably explicable as owing to their gonidial elements belonging to Algæ identical with or similar to the species here described. The sub- epidermal site is at once explicable by the supposition of the entrance of hyphæ along with the filaments of the Alga going to the. formation of the secondary disks; whilst the cephaluroid condition may, so far as can be judged from figures at all events, be referred to cases in which the algal element has partially retained or regained the ascendancy, and has given origin to a crop of the asexual fructification normally belonging to it. In conclusion, it may be well to call attention to the fact that the existence of structures closely resembling the primary disks of the Alga forming the subject of the present paper has been already indicated by various authors. The organisms described by Mettenius as occurring on the leaves of ferns, and referred by Millardet to the genus Phycopeltis *, may with as much propriety be ascribed to the present genus. Even the relation of the disks to fungal elements would seem to have been noticed. For example, the Rev. Mr. Berkeley, in a paper on “ The Thread Blight of Tea,” mentions the occurrence on tea-leaves of ** minute shield-like bodies, consisting of cells radiating from a central aperture containing spores," and gives a figure which might well pass for that of one of the primary disks f. Mr. Archer also, in exhibiting some of Dr. Bornet's preparations, commented on one showing a minute lichen with its hyphe investing a form of Phyllactidiwm or Coleochete, and pointed out the singular habitat of the Alga, ** on the leaves of living trees in the tropics” 1. * o De la Germination des Zygospores dans les genres Closterium et Staurastrum et sur un genre nouveau d'algues Chlorosporées,” par M. A. Millardet, Mémoires de la Société des Sciences Naturelles de Strasbourg, 1870. T Quarterly Journal of Microscopical Science, 1875, p. 132. t Ibid. p. 334. 316 DR. D. D. CUNNINGHAM ON MYCOIDEA PARASITICA. DESCRIPTION OF THE PLATES. PLATE XLII. Fig. 1. Leaf of Camellia japonica affected by Mycoidea parasitica. 2. Aerial filaments bearing the asexual fructification : x 92. Fig. 3. Group of young aerial filaments emerging from beneath the epidermis: x 180. 4 5 . Upper extremity of an aerial filament, with stipitate spores arising from it: x 480. . A spore from which the zoospores have escaped, showing the rounded opening of exit and four zoospores yet remaining in the cell-cavity : x 960. Fig. 6. A free zoospore in its original condition : x 840. Fig. 7. Aportionofone of theaerial filaments, showing projectionson theinner surfaceof thecell-wall: x 960. ` Fig. 8. A vertical section through a portion of a Camellia-leaf affected by two patches of Mycoidea parasitica: x 180, 3 Fig. 9. Filaments from the subepidermal disk in their original condition: x 180. Fig. 10. Similar filaments from a plant with sexual fructification: x 180. Fig. 11. A young oogonium: x 180. Fig. 12. An oogonium with attached pollinodium : x 980. Fig. 13. Mature oogonium with escaping zoospores: x 960. Fig. 14. Portion of an oogonium showing development of ostiolum : x 980. 1 Fig. 15. Portion from the surface of a leaf, showing the ruptured epidermis and protruding ostiolum of ` — the oogonium: x 1960. ; Fig. 16. (a) A zoospore in the circular condition: x 1960. (b) A zoospore after rupture of the cell-wall. (c) Oval particles contained in the interior of the zoospores. Fig. 17. Zoospores which have become circular and are beginning to germinate: x 1960, Pirate XLIII. Fig. 1. Group of mature oospores with filaments ramifying around the oogonia: x 180. Fig. 2. Mature oogonium with its ostiolum and oospore: x 960. Fig. 3. Young primary disk, resulting from the germination of a zoospore: x 700. Fig. 4. Vertical section showing the relation of the primary disk to the epidermis: x 180. Fig. 5. Portionof epidermis, with aprimary disk andthemassof subepidermal filaments arisingfromit: x 180. Fig. 6. Primary disk as seen from the surface : x 37. Fig. 7, Vertical section, showing germinating cells of the primary disk, and penetration of the epidermis by the buds arising from them : x 960. Fig. 8. Lichenous patch on the surface of a Nephelium-leaf : of natural size. Fig. 9. Portion of a patch with spermogonia and apothecia, viewed laterally : x 23. Fig. 10. Detached portion of a patch, showing the film of E Siem? the spermogonia, apothecia, and circular groups of gonidia: x 23. j Fig. 11. Section through a portion of a gonidial patch, showing the filamnets of the cortical layer, the . remains of the algal disk, the green gonidial cells, and the intergonidial hyphæ : x 960. Fig. 12. Portion of a primary disk, seen from below, with fungal filaments attached to it: x 960. Fig. 13. Spermatia: x 960. Fig. 14. Portion of dense cellular tissue from the point where the fungal filaments penetrate the disk: x 960. Fig. 15. Apothecium in vertical section: x 44. Fig. 16. Portion of the skeleton of a primary disk detached from a group of gonidia, with one or two goni- dial buds still connected with the cells: x 960. Fig. 17. Detached mass of hyph: and gonidia from one of the groups: x 960. Fig. 18. Free gonidia; two of them showing division of their contents : x 960. Fig. 19. Hyphz of the cortical layer: x 960. 3 Fig. 20. Medullary fibres: x 960, Fig. 21. Asci and paraphyses : x 960. * Trans. Linn.Soc Ser 2 Bor Vou ] Tas 42 | \ X 4960 | Zig. WN N zal f à > eee A ME. : Í Í | " » — n. , = Xe 96Ccomccccao 6C]! oco e JoCoCICoC OCG En S ^ pius nA Ped ees del MYCOIDEA PARASITICA. | Lud EUNT ENEE A . DDCunningham del ; : MYCOIDEA PARASITICA Taans Lane Soc SER 2.Bor Vou 1 Tap 43 Fig 4 1 ! = anana A le WOOO ICI OC Keleleleleleleleld x 4/80 Fig 6. | F3] Fig 18 em bx p E \ / Sch ) | [ 37 ] XXI. On the Self-fertiligdfion of Plants. By the Rev. GEoRaE HeEnstow, M.A., F.L.S., POS (Plate XLIV.) Read November 1st, 1877. INTRODUCTION.—No one can venture to treat of the fertilization of plants without being deeply indebted to the laborious investigations of Mr. Darwin, and, I would add, no one ought to do so without expressing his profound gratitude to that great author for the aid derived from his many works. Such, at least, are my own feelings, that while I differ in some respects from his conclusions with reference for the subject of this paper, I cannot proceed without first acknowledging a debt of gratitude for the vast stores of facts which are to be found in his writings. In order to establish what I believe to be the real value and effects of self-fertilization, I shall be obliged to refer frequently to Mr. Darwin’s latest work on the ‘Cross and Self-fertilisation of Plants,’ because he has therein several times stated his belief in what he calls ‘the evil effects and *'injuriousness" of self-fertilization—terms which, I have reason to think, not only fail to express accurately, but which are liable to misrepresent the real value of the pro- cess. It is true, there may have occurred in his experiments some individual instances of cultivated plants to which such terms might seem applicable; but it is not so much with isolated and exceptional cases that I purpose dealing as recognizing a broad and general principle in the vegetable kingdom, taken in its entirety—one, in fact, not necessarily limited to the flowering plants, but applicable to ni Aap qure as well, though I am not at present concerned with the latter*. Before addressing myself to the subject proper, I think the reader will not consider it out of place if I state, as briefly as possible, what are the results Mr. Darwin has secured to science by his latest observations—as, in order to show what I conceive to be the true value of self-fertilization, it will be necessary to point out the "n value of other kinds of crossing, so as to make the results comparative. There are, at least, five kinds of union, apart from hybridization proper, or the union of distinct species or genera :—(1) self-fertilization, or the fertilization of a pistil by the pollen from the same flower; (2) crossing different flowers on the same plant; (3) cross- ing flowers on different plants of the same stock ; (4) crossing flowers of different plants, * I had purposed writing this paper before I was aware that Mr. Darwin's book was coming out, and contributed an article on self-fertilization to * Nature,’ vol. xiv. p. 543, in which number Mr. Darwin’s book was, I think, first mentioned as being shortly about to appear. On subsequently restudying the question, with the aid of Mr. Darwin’s work, I found it necessary to thoroughly recast my paper, as he had recorded a vast number of facts quite new to me, and of which I have now taken advantage. I would also add that I did not study Miiller’s ‘Befruchtung der Blumen durch Insekten’ until I had written out all my own observations, so that wherever I have quoted or alledag to that acute observer’s writings, such sentences are later additions to this paper, SECOND SERIES.—BOTANY, VOL. I. 9v — 318 REV. GEORGE HENSLOW ON THE but of different stocks, all the preceding being of exactly the same form of species; (5) crossing different varieties of the same species. Dean Herbert, in his work on the Amaryllidee (p. 371), published in the year 1836, makes the following remark :— * I am inclined to think that I have derived advantage from impregnating the flowers from which I wished. to obtain seed with pollen from another individual of the same variety, or, at least, from another flower rather than its own, and especially from an individual grown in a different soil or aspect.” As far as I am aware, such a surmise has been generally accepted by horticulturists and botanists to the present time, but more as a general assumption than as one based on strictly scientific experiments; and it is the great merit of Mr. Darwin that, just as he established, on scientific grounds, the value of intererossing by insect agency, so well illustrated by Sprengel in 1790, so, in his present work, he has done the same for Dean Herbert’s inferences. He has thus placed on a scientific basis the exact value of | such crossings. Of the results thus anticipated by Dean Herbert, Mr. Darwin has proved incontestably that the greatest benefits are derived by crossing individuals which have grown in dif- ferent localities, so as to acquire different constitutional peculiarities, though they may not be perceptible in their external appearance, or else, if differences be visible, they consist of some slight characters, which render one a subvariety of the other. When such plants are crossed, the offspring raised exhibit, as a rule, greater weight, more vigour in branch- ing, a darker foliage, more blossoms, an earlier period in flowering, &c., than those of their parents. Such beneficial results, however, are not absolutely invariable. Mr. Darwin LS [ 1 cases where but slight or even no benefit accrued to the plants raised from the seed 1 of the crossed plants; but that, on the other hand, the self-fertilized plant beat its - crossed opponent in height &c., with which it was put in competition. Such, for ex- ample, was the case with a tall, highly self-fertile individual of Ipomea purpurea, to which Mr. Darwin gave the name of * Hero." Eschscholtzia californica, Petunia vio- lacea, and Mimulus luteus are other instances. pe: The next degree of advantage of crossing was with individuals of the same stock, i.e. grown for many generations in the same soil and under similar conditions; and which therefore were probably all descended from the same parentage. With these the advantages of crossing were apparent at first, but after a few generations the com- parative results between such intercrossing and self-fertilization gradually approximated — uniformity. The third kind of union is between flowers on the same plant. And here Mr. Darwin | has corrected a hitherto widespread supposition, that it was beneficial for flowers to be thus crossed. It would seem, however, that such may do little or no good at all. Un- fortunately Mr. Darwin has not experimented to any great extent upon such a union ; and it would be unwise to generalize from so few instances. Collecting the various . items from his work under this head, they may be tabulated as follows, where 100 is | T placed as the standard for height or weight of the intercrossed :— ‘SELF-FERTILIZATION OF PLANTS. 319 Intercrossed. Self-fertilized. Ipomea purpurea—ratio of heights . . . . . 100 106 ia e So WEIS. ^ «o5. d^ bora 400 124 Mimulus luteus - ,; leighis oce ad mes bas OO 101 ` SE ‘j Horoa tsoias et od: andy 100 103 Digitalis purpurea — ,, BEEN | —. . —. IU0 94. » »* Wi og weight: 1.105.751 100 78 A slight benefit was gained by the intercrossed "m of Eschscholtzia californica, Corydalis cava, and Oncidium (sp.). No benefit followed from intercrossing flowers on Origanum vulgare, Pelargonium (sp.), Reseda odorata, and Abutilon Darwinit. From these cases it appears that no benefit resulted in four cases, and with all the rest, excepting the weight of Ipomea, the advantage is so slight as to be comparatively insignificant; for the instances are too few,to be trustworthy for deducing broad or general results. It is, however, worth while observing, that the flowers of these species of Eschscholizia, Corydalis, Oncidium, and Digitalis are more or less usually self-sterile, the first three physiologically, the last morphologically so. That individual buds on a plant may be more or less, but probably rarely very greatly differentiated, is a fact well admitted, and treated by Mr. Darwin elsewhere under the title of bud-variation: and when such flowers are crossed, some benefit is likely to accrue to the resulting seeds ; for when flowers are more or less self-sterile, it shows that their sexual organs have become more highly differentiated than usual. Hence one would, à priori, expect that more benefit would result from their crossing than with flowers which had been less differen- tiated, as is doubtless the case with most flowers on one and the same plant; so that we have a reasonable explanation from crossing different flowers on the same plant of Esch- scholtzia californica, Corydalis cava, Oncidium (sp.) *. The fourth kind of union is self-fertilization; and although Mr. Darwin seems on several occasions to be strongly inclined to recognize some benefits as resulting from this process, as in the passages to be quoted presently, yet he repeatedly insists upon, not merely what I should prefer to call the negative results, but the “ evil effects” of self- . fertilization. It is here, therefore, where I join issue with him ; and it will be the main object of this paper to give positive evidence to show that there are decided advantages, rather than disadvantages, accruing to such plants as can, or rather do, habitually fer- tilize themselves. * Such an explanation of the nature of self-sterile plants is given by Mr. Darwin in his work on * Animals and Plants under Domestication,’ vol. ii. p. 140; but he appears to have now changed his views, for in ‘Cross and Self- fertilisation of Plants’ (p. 345) he says:—** When such plants [which are capable of complete self-fertilization] are taken to another country, and become in consequence self-sterile [as did Eschscholtzia in Brazil], their sexual elements and organs are so acted on as to be rendered too uniform [?]for such interaction [why not become too highly differentiated ?], like those of a self-fertilised plant long cultivated under the same conditions" [but such con- ditions do not render self-fertilizing plants self-sterile]. ; I cannot but think this change of view unfortunate ; for the earlier interpretation, to my mind, certainly carries more weight with it; for had such flowers been too little differentiated, it would imply they were under a more pri- mitive condition; and one would reasonably ie some correlative “ lowness" of structure elsewhere; but eh is far from being the case. : 2¥2 320 REV. GEORGE HENSLOW ON THE I shall now quote three passages from his ‘ Cross and Self-Fertilisation of Plants,’ in which Mr. Darwin inclines to the belief that there is some good in self-fertilization :— “ We should always keep in mind the obvious fact that the production of seed is the chief end of the | act of fertilisation ; and that this end can be gained by hermaphrodite plants with incomparably greater certainty by self-fertilisation, than by the union of the sexual elements belonging to two distinct flowers or plants ” (p. 3). In speaking of the superiority of self-fertilized seedlings of Ipomea over those raised | from flowers fertilized br pollen taken from other flowers on the same plant, he says:— “ This is a remarkable fact, which seems to indicate that self-fertilization is in some manner more i advantageous than crossing, unless the cross bring with it, as is generally the case, some decided and i preponderant advantage ” (p. 61). Lastly, Mr. Darwin observes :— 1 “The most important conclusion at which I have arrived is that the mere act of crossing by itself 1 | does no good. The good depends on the individuals which are crossed differing slightly in their bh constitution ” (p. 27). H Before proceeding with my own observations, it will be advisable to give the opinions ` of at least two eminent botanists, who regard self-fertilization as an important principle l in nature. I allude to Dr. H. Müller and Mr. T. Meehan. 1 The former, alluding to Mr. Darwin's well-known aphorism that ‘ Nature abhors per- — petual self-fertilization," says that it was exaggerated by his successors, as Hildebrand - in Germany, and Delpino in Italy; while, on the other hand, he refers to Axell, who 1 propounded the doctrine that— | E " The development of the fertilizing arrangements in phanerogams has been always in advance, and 1 still continues in advance, in one and the same direction, towards a perfection which affords more and 1 more facilities for self-fertilization.” He then says that— “He is convinced that neither Hermann, Darwin, nor Axell's opinion is a thoroughly adequate one, | but that, under certain conditions, the facility for self-fertilization is most advantageous to a plant, 1 while, under other conditions, the inevitableness of cross-fertilization by the visits of insects is the more — advantageous." Similarly Mr. Thomas Meehan, of Germantown, Philadelphia, in a paper entitled, “ Are Insects any material Aid to Plants in Fertilization ? " *, says that he does not regard this supposed necessity for cross-fertilization, or this supposed injury to plants, as at all established, nor is the injuriousness to animals from in-and-in breeding by any means proved, but that there are abundant evidences to the contrary. He further adds his belief that undoubted self-fertilized plants have existed as long, and are every way as healthy, as those that cannot now fertilize themselves. His admirable paper, which was reproduced in the ‘Gardeners’ Chronicle’ for Sept. 11, 1875, is, in fact, an “apology” for self-fertilization. * Proceedings of the American Association for the Advancement of Science, vol. xxiv, p. 243 (1875), Also ‘ Nature,’ Sept 25, 1873, : SELF-FERTILIZATION OF PLANTS. 321 Lastly, Mr. Darwin’s expressions, “evil effects" and “injuriousness”’ of self-fertiliza- tion, seem to imply something of the nature of disease; but they show no consonance with Mr. Berkeley’s definition of disease, which is as follows :— * Tn all cases where the necessary functions of the species are impeded, and vital action is impaired, whether tending to actual decay or to the obstruction of the main ends of cultivation [aud I would add that the sole recognizable end of all plant-life in a state of nature is the propagation of the species by seed], we must assume that diseased action is going on” *. I hope this essay will clearly show that nothing of the kind can be predicated of the innumerable and habitually self-fertilized plants in nature. I will now state the nature of the observations which have led me to believe particular plants to be normally self-fertilizing (and by that I mean, though they may in some cases be cross-fertilized by insects, especially perhaps by small moths at night, yet there is every reason to believe such to be exceptional, and that they can and do fertilize themselves habitually) :— (a) Inconspicuousness of the flowers even when fully expanded. (b) Calyx and corolla, often only partially expanded, or not at all. (c) White or pale colour of the corolla; while specially coloured streaks, specks, or “ guides," Ee, are more or less reduced or absent. (d) Partial or total arrest of the corolla. (e) Little or no honey, though the disk or glands may be present. The glands found in allied species, however, may be suppressed in the self-fertilizing. (f) Little or no scent. (g) The mature stamens of the expanded flower retaining the incurved position they had in bud. ; (K) Stamens often reduced in number and the pollen in quantity. (i Pollen-tubes visibly penetrating the stigma, either from grains still within the anther-cells, or evidently derived from those of the same flower. (j) The stigma situated appropriately for direct pollinization from the anthers of the same flower. | (k) The order and rates of development of the different whorls, the pistil maturing with comparative rapidity. — | (1 The early maturation of the stigma so as to be ready before or simultaneously with the dehiscence of the anthers. (m) Great (absolute) abundance of fruit or seeds, or both. (n) Great rapidity of maturation of seed. [fertilization. (o) Special contrivances and adaptations of the stamens and pistil to secure self- (p) Cleistogamy. ` $ (q) Netting to exclude insects. (r) Growing them indoors and free from insect visits t. * « Vegetable Pathology,” Gard. Chron. (1854), p. 724. + Of these two experimental methods (g and r) I have little experience, and am dependent upon Mr. Darwin's and H. Müllers observations. Having no facilities in London, I have been almost entirely compelled to forego these important methods. Still I venture to think that the cumulative evidence gathered under the preceding heads affords sufficient proof for conviction, which is quite equivalent to an absolute demonstration by such methods. 322 ,REV. GEORGE HENSLOW ON THE (s) Self-fertilizing plants are very often annuals, while their allies or intercrossing species of the same genus are perennials. In this respect Stellaria media may be compared with S. Holostea, Geranium pusillum. Ze, with G. pratense, Polygonum aviculare with P. Bistorta and many others. There are, of course, many exceptions, as Malva rotundi- Jolia ; but even this species is said to become annual in New Zealand. 1 (t) ** Weeds” are probably all self-fertilizing or else anemophilous. A weed is simply . - an unattractive plant, and possessing no feature worthy of cultivation. The following are the chief facts which may be regarded as occurring correlatively with self-fertilization, some being actual causes which directly or indirectly bring it about :— l. The majority of flowering plants can fertilize themselves. 2. Few plants are known to be physiologically self-sterile with their own pollen. 3. Many plants bear flowers which are morphologically or structurally self-sterile; the — pollen of any such flower, however, is effectual if artificially placed on the stigma of - the same flower. 4. Both physiologically and more or less morphologically self-sterile plants may become highly self-fertile under certain conditions, self-fertilization being then correlated with or even caused by one or more of the following peculiarities :— i. The withering of the corolla after or without any expansion. ii. The excision of the corolla with a portion of the stamens (?). iii. The partial or even total arrest in development of the corolla, normally present. iv. Absence of colour in the corolla. v. The corolla closing after expansion. vi. The flower-bud never opening. vii. In the absence of the proper visiting insects. ` [ climate. viii. The reduction of temperature, as in autumn, or by transportation to a cooler ix. By grafting on a new stock. Highly self-fertile varieties may arise under cultivation or in nature. Inconspicuous flowers are highly self-fertile. . Cleistogamous flowers are always self-fertilized. . Special adaptations occur for securing self-fertilization. . Conservation of energy is seen in the reduction of the number of stamens and the quantity of pollen in self-fertilizing flowers. St The relative fertility may equal or surpass that of crossed plants. d l. The fertility does not decrease in successive generations of a plant perpetually Been | 12. The fertility may increase in successive generations, when a more or less self- sterile plant has acquired self-fertilizing powers. 13. Free from competition, self-fertilized plants may be equal to the intercrossed com- pared with them either (1) as seedlings, or (2) when planted in the open ground. 14. Self-fertilized plants sometimes gain no benefit from a cross with the same or even a different stock. | 15. Self-fertilized EN are ə perfectly rcs SG © CO vi E or SELF-FERTILIZATION OF PLANTS. 323 16. They may be much more productive than flowers dependent upon insects. 17. The most widely dispersed of British plants are almost, if not quite, always self- fertilizing. 18. Naturalized abroad they often gain great vigour. 19. They are **the fittest to survive in the struggle for life." I will now proceed to treat of each of these facts in succession. l. The majority of flowers are self-fertile. It is to Mr. Darwin's and H. Müller's investigations that I am partly indebted for this fact. Until their works were published I had, perhaps in common with others, a general belief that conspicuous flowers were much more self-sterile from dichogamy or other cause than is really the case. The former gives two lists of forty-nine species in each, one of self-sterile and the other of self-fertile plants; but he adds (J. c. p. 370):— * T do not, however, believe that if all known plants were tried in the same manner, half would be found to be sterile within the specified limits; for many flowers were selected for experiment which presented some remarkable structure; and such flowers often require insect aid." The limit was the production, when covered, of less than about half the number of seeds produced by unprotected plants. So that while 49, in Mr. Darwin's experiments, were generally highly self-fertile, the other 49 were not absolutely self-sterile when protected. Moreover, a large number of inconspicuously flowering plants from a great variety of Orders may be mentioned which are habitually self-fertile. Hence the above statement may be considered as established notwithstanding Asclepiadacez and Orchid- acee being large Orders and exceptional in requiring almost universally insect aid for the cross-fertilization of their species. 2. Comparatively few flowers are known to be physiologically quite self-sterile. When the pollen of a self-sterile flower is placed upon the stigma of the same flower it is impotent. Mr. Darwin has dealt with this subject in his work on ‘ Animals and Plants under Domestication,’ vol. ii. p. 132, seqq., to which the reader must be referred for details. It appearsthat such plants are chiefly Orchids and of the genus Onicidium ; ‘but examples are to be met with from widely different Orders: thus Corydalis cava, Lobelia fulgens, Verbascum nigrum, species of Passiflora, Ee, are in this condition; and I repeat that I cannot but think Mr. Darwin’s interpretation given in the above work is correct, and not that which he has lately advanced in his ‘ Cross and Self-fertilisation of Plants.’ In the former work he says, ** The sexual elements of the same flowers have become differentiated in relation to each other almost like those of two distinct species ;” but in the latter book he says, such sterility is ** due to the sexual elements not having been sufficiently differentiated " (p. 456). All the flowers here mentioned have their perianths in a highly differentiated state, as witnessed by their great irregularities, and it is at least reasonable to infer that the pollen is correlatively changed as well. This idea is certainly corroborated by the fact that the pollen of some species of Oncidium actually affects the stigma as a poison. But what is particularly noticeable is that, in the first place, allied species of such 324 REV. GEORGE HENSLOW ON THE self-sterile plants are perfectly self-fertile; thus while Verbascum nigrum is quite sterile, V. Thapsus and V. Lychnitis are highly self-fertile. Again, though Corydalis cava and C. solida are self-sterile, C. Halleri is only slightly self-sterile, and C. intermedia notat — - allso. Lastly, while several species of Passiflora are self-sterile, P. gracilis, an annual, is 1 “ nearly as fertile with its own pollen as with that from a distinct plant.” Secondly, phy- : siologically self-sterile plants may themselves become highly self-fertile under certain con- | ditions. Thus Mr. Darwin quotes the experience of Kélreuter, who found Verbascum - pheniceum to be sometimes self-sterile, at other times self-fertile. Some plants of — Lobelia fulgens, according to Gärtner, are self-sterile, others not so; but none of ` Mr. Darwin's specimens were in this condition (J. c. p. 179). Eschscholtzia californica is 1 self-sterile in Brazil, but when introduced into England it became self-fertile. This species, ` however, is also much more self-sterile in Germany; and a plant of English parentage | sent to Brazil, and exposed for two seasons to that climate, * proved quite self-sterile, ' like a Brazilian plant" (l c. p. 333). 1 These two classes of facts show that physiological self-sterility is only a conditional | phenomenon; and probably any usually self-sterile plant will regain self-fertility if — grown under changed circumstances, as, for example, was the case with the Brazilian | Eschscholtzias described by Mr. Darwin, who says, * their self-fertility had evidently 1 increased greatly by being reared for two generations in England." 1 Hence as many of the so-called self-sterile plants become self-fertile in other localities, ] | and then, as with Eschscholtzia, show very great vigour, we cannot infer that self-sterility — injures the constitution in any way, but is, as Mr. Darwin says, an “ incident ;” for a degrees of self-sterility and want of vigour show no necessary correlation whatever, and 1 the former involves no detriment to the plant beyond the loss of fertility. We may 1 therefore with tolerable safety, infer that those others which have not yet been shown to — be self-fertilizing might become so under proper conditions for evincing such reversion. | 3. Many plants are morphologically self-sterile. Besides physiologically self-sterile plants, there are many of which the pollen of any 1 flower, although it be quite effectual on the stigma of the same flower, is prevented by cer- - tain mechanical obstructions from having access to it, Thus if species of Lupinus have their — petals artificially moved, seed is set, but not usually otherwise. Now it appears probable | that if such plants be habitually neglected by insects, they will either perish entirely — or else become self-fertilizing, or else, as another but more rare alternative, become ` anemophilous, as it has probably been with Poferium and Plantago*. The above ` appears to have been the case with such conspicuous flowers as Pisum sativum and ` Lathyrus. odoratus, which are mostly perpetuated in this country by self-fertilization — alone. Phaseolus vulgaris is also highly self-fertile; but P. multiflorus, though closely - allied to it, is still morphologically more or less self-sterile. ` 3 Similarly, certain inconspicuous and irregular flowers, which may therefore be re- d * Mr. W. B. Clarke informs me that he has found certain small-flowered species of Combretum to be protero- 1 gynous, like Plantago, with the style protruding through a circular orifice of the unexpanded flower before the stamens appear. = bey i: pod inposita * : SELF-FERTILIZATION OF PLANTS. 325 garded as certainly descended from insect-fertilized plants, have regained and retained the power of self-fertilization. Thus while Trifolium pratense seems generally to require the aid of humble-bees, some very small-flowered species, as 7’. minus, are self-fertile. Again, while conspicuous species of Dicentra and Corydalis are adapted for insect agency, Fumaria officinalis is highly self-fertile. A Both physiologically and morphologically self-sterile plants may become highly self- fertile under certain conditions. I have enumerated nine proximate causes which appear to be more or less capable of accounting for the self-fertilization of certain flowers, or they are at least phenomena cor- related with it. Some of these require additional observations to establish them, as a sufficient number of cases has not yet been observed; but at all events they are all based upon direct observations of usually many more than one instance. 4. i. The first cause stated is the withering of the corolla. Mr. Darwin says of Viola tricolor that a large covered plant set only 18 capsules, several of which contained from only one to three seeds; but an uncovered plant set 105 . fine capsules. " The few flowers which produce capsules when insects are excluded are perhaps fertilised by the curl- ing inwards of the petals as they wither, for by this means pollen-grains adhering to the papille might be inserted into the cavity of the stigma” (J. c. p. 124). I can corroborate this fact, for I have found a flower of this plant with the calcarate petal ‘glued’ to the stigma with plenty of pollen-grains penetrating the orifice; the stamens were detached, and remained suspended above the ovary, just as in cleistogamous violets (Tab. XLIV. fig. 6 d). In September, 1876, I found a plant of Tradescantia erecta at Kew producing an abundance of capsules, but in every instance from unopened flower-buds. The corollas were completely shrivelled, and had never expanded, and were matted down upon the anthers together with the long style, so that the stigma was impregnated by the pollen beneath the withered corolla. Several of the anthers had withered immaturely. I gathered several stalks, which, when placed in water, lived for several days, and opened their capsules successively. The embryos were perfectly well formed (Tab. XLIV. fig. 36). The seeds germinated in 1877, and grew into vigorous plants. Mr. Darwin records that Zpomea purpurea is highly self-fertile, while Convolvulus arvensis can fertilize itself. `C. sepium closes at night, but not, like C. arvensis, in wet weather. It has no smell, and is comparatively little visited by insects. Now all these plants wither rapidly, the corolla becoming twisted upon the essential organs; and I strongly suspect such a process is highly favourable to the securing of self-fertilization, although the stamens do not appear to be proterandrous. The stigmas are somewhat elevated, at least at first, beyond the anthers. H. Müller also alludes to Hypericum perforatum as being specially assisted in the process of self-fertilization by the withering of the corolla. He observes that the anthers and stigmas mature together, and that although there is no honey, yet the conspicuousness of the flowers will attract pollen-seeking insects which might cross SECOND SERIES.—BOTANY, VOL. I. 22 326 REV. GEORGE HENSLOW ON THE them ; still the abundance of pollen renders the flowers very liable to be self-fertilized. The innermost stamens develop first, while the outer ones are delayed. This is contrary to the order in Buttercups. Perhaps it is correlated to the elongated styles, which project outwards, and whose stigmas will thus be fertilized more probably by the last or outermost dehiscing anthers, just as the carpels are in Buttereups, though in this latter case by the innermost stamens. In addition, therefore, to the probability of self- fertilization occurring in the expanded flower, the withering of the corolla which binds stigmas and anthers together will inevitably ensure it. Many other flowers retain their corollas or perianths for some time in a withered or * marcescent ' condition, such as Gentianee, Campanula, Ze, and I would suggest the probability of such being in all cases at least an aid to secure self-fertilization. 4. ii. The excision of the corolla, and stamens in part(?). It occurred to me on observing, I may say, the almost universal tendency to reduce the size of the corolla or to suppress it altogether when flowers are habitually or solely self-fertilized, and that it was also the case with the female flowers of polygamous species as well as of * gynodiecious" plants, as Mr. Darwin calls them (* Forms of Flowers,’ pp. 287-309), that there might be some sort of compensating process between the pistil and the corolla—that when the latter is arrested the former grows rapidly, so that if it were early cut away a normally proterandrous flower might become homogamous ; but experi- ments with strongly proterandrous flowers gave mostly negative results when the corolla alone was removed from the bud. Hence I am led to regard the reduction of the corolla as a secondary result issuing from some tendency to check the energy of the stamens, which may convert a proterandrous flower into a self-fertilizing one, or a normally hermaphrodite into a female. The reduced corolla, and sometimes a reduced calyx *, must therefore be regarded as correlative results issuing from certain hidden causes affecting the essential organs.. I have tried several experiments of removing the corolla both with and without some or all of the stamens in Pelargonium; the style and stigma then appeared to gain vigour, and to mature from two to five days earlier than usual. 'lhis, however, requires corroboration. Mr. Darwin records the fact that certain proterandrous or self-sterile flowers, which had their corollas removed, set seed, such. as Delphinium Consolida and Viola tricolor (* Cross and Self-fertilisation,’ p. 420, note) ; and this led me to regard ‘such cases as E strengthening my deduction; but I am now inclined to withdraw it, and substitute the _ belief that the reduction of the corolla is an outward index, as it were, of some con- — stitutional peculiarities which are affecting more especially the stamens. I have else- A where observed of diplostemonous flowers that if a whorl of stamens be suppressed itis 4 usually that opposite the petals, and in the Paronychiew the corolla often goes with it, showing some hidden bond of connexion between these two whorls. Hence I would -agree with Mr. Darwin, who thus speaks of the smallness of the corolla of the female ; forms of usually hermaphrodite plants :—‘ It seems therefore probable that the de. ` * Rhamnus catharticus (t Forms of Flowers,’ p. 308) and Amphicarpac monoica have cleistogamous flowers without even “ the semblance of a e e (I. a) SELF-FERTILIZATION OF PLANTS. 327 creased size of the female corollas in the foregoing cases is due to a tendency to abortion spreading from the stamens to the petals ” (* Form of Flowers,’ p. 308). On the other hand, the Corymbiferze of the Composite furnish exceptions ; for the ray- florets are female, and have larger (ligulate) corollas than have the disk-florets ; while Centaurea and Viburnum Opulus carry us further; for the corollas are relatively even still larger, the pistil being arrested as well as the stamens. 4. iii. The partial or total arrest of development of the corolla. The tendency to arrest of the corolla in many flowers, which, however, set seed very freely, has often been observed. Thus M. Boisduval says* that ** Viola palustris is often apetalous in the mountains, notably at Lautaret. In the environs of Paris this species has petalous flowers, which fructify very well [probably by insect agency], but never apetalous. It is the same with V. biflora, an essentially Alpine plant." He adds that E Zuppi a species of Northern Italy, never bears * with us" conspicuous flowers, which does not hinder it from fructifying. He also cites Prismatocarpus and some Campanulas, whose flowers have no corolla, which fact does not hinder them from fructifying. Mr. Berkeley also alludes to the want of a corolla being no hindrance to the setting of seed (* Gard. Chron.’ 1855, p. 36). “ Varieties occur, as, for instance, in the Sweet William, where not a trace of petal exists, and yet every ovary is impregnated ; but the contrary condition is not infrequent. In several species of Violet the early petaliferous flowers are often barren, while those which appear later in the season are pro- ductive [i. e. the cleistogamous apetalous flower-buds]. In Ajuga Iva fertile flowers occur indifferently with or without a corolla. In Ononis minutissima, in contradistinction to the Violets, the earlier flowers are apetalous, and equally fertile with the petaliferous flowers which are produced later in the season; and the same equal fertility belongs to both kinds of flowers in Helianthemum, in which genus, Lespedeza, &c.they are generally simultaneous. In some of these cases the stamens seem very imperfect.... A very limited number of pollen-grains may sometimes be found." The general conclusion I would draw from all such cases where the corolla is more or less reduced or entirely absent is, that the flowers are readapting themselves to self- fertilization ; the conspicuous corolla being no longer required for insect agency, the energy usually expended in developing it is now diverted into the pistil, while a not unusual degeneracy or “ contabescence” of some of the stamens accompanies the process, involving a reduction of the quantity of pollen, which I regard, not as any sign of “ injuriousness," but as a form of less expenditure of wasted energy. 4: iv. A partial or entire loss of colour from the corolla is often correlated with self- fertilization. Generalizing from several observations, I came to the conclusion that what- ever cause may be at work to abstract or withhold energy from, it may be, the staminal whorl, together with the corolla and adjacent parts, tends to promote self-fertilization ; so that asthe absence of colour may be regarded as indicating a poverty of nutriment in the corolla, whenever some colour is normally present in the species, and of which white or pale-coloured forms are varieties, such would seem to be, if not a proximate cause, at least a concomitant of self-fertility. Several instances are recorded by Mr. | * Bull. Soc. Bot. de Fr. iii. p. 569. i 222 7998 REV. GEORGE HENSLOW ON THE Darwin of such a correlation. Thus pale-flowering Pelargonia, such as,“ Christine," — according to Mr. J. Denny, are “ great seeders.” That observer remarks (as quoted by ` Mr. Darwin, l. c. p. 142, from the * Florist and Pomologist, Jan. 1872, p. 11) :— * There are some varieties, especially those with petals of a pink colour, or which possess a weakly ] constitution, where the pistil [stigmas] expands as soon as or even before the pollen-bag bursts, and in — which also the pistil is frequently short, so when it expands it is smothered, as it were, by the bursting 1 anthers ; these varieties are great seeders, each pip being fertilized by its own pollen." (Tab. XLIV. fig.12.) - That albinism is due to want of energy is corroborated by some experiments of 1 Mr. B. T. Lowne, who found that garden Balsams became white if ammonia be with- — held from the soil, but if they were manured with it the colour returned. | 1 Another instance of a usually strongly proterandrous flower becoming self-fertile by — losing its dichogamy is Dianthus caryophyllus. Mr. Darwin cultivated a dark crimson — variety, which became highly self-fertile in the third generation :—‘ The proportional - number of seeds per capsule produced by the plants of crossed origin to those produced ^ by the plants of self-fertilized origin (both lots being spontaneously self-fertilized) was as ] 100 to 125.” On p. 309 we learn that ** the self-fertilized plants of the third generation 1 all bore flowers of exactly the same pale rose-colour ;" and Mr. Darwin thinks that “it | is not improbable that some of the parent plants which were first self-fertilised may have — borne flowers thus coloured.” If, as is also probable, others were of the crimson kind, 3 then such must have acquired the paler tint. E Of Antirrhinum majus, the relative self-fertility of the red and white varieties is ` as 98:20. Of Lobelia ramosa, while the coloured variety was quite sterile, Mr. Darwin 4 found the white or **snow-flake " form partly self-fertile, and produced when protected 3 about one third as many seeds as when uncovered. Mimulus luteus produced under cul- 3 tivation a highly self-fertile white or nearly white variety. Its fertility, as compared | with that of intercrossed plants, which are yellow, was as 147: 100 (l.c. p. 348). Ker- - bascum Lychnitis is highly self-fertile, and has a very pale-coloured corolla, as well as — white hairs on the filaments ; but V. nigrum is bright yellow with purple filaments, and 1 is self-sterile. Lastly, the Petunia "violacea cultivated by Mr. Darwin was of a dingy | purple, but the self-fertilized plants assumed a flesh-colour. E While writing I have before me a blossom from a variety of the crimson Chinese Pring 1 rose. The colour is approximating a brick-red, The greenish spot at the base of each petal ` is not well defined as in the purer breeds, but extends upwards with an indented outline - spreading over the lower third part of the petal. There is also a minute white speck at ` the base of each incision of the margin. It belongs to the short-stamened form, as they ` are situated halfway down the tube; but the pistil is entirely below them. It seeds very 1 freely. The foliage and petals are somewhat smaller than in the normal forms. P A white variety of Polygala vulgaris. This I found to be self-fertilizing, as will be 1 described hereafter. (Tab. XLIV, figs. 7 a, b.) | Pinguicula. The larger species, with dark purple blossoms, are adapted for inter- 1 crossing; but P. lusitanica, which is pale lilac or white, is self-fertilizing. pans XLIV figs. 34 a, b, c.) Lysimachia vulgaris. Dr. H. Müller Fecords the fact that, the self-fertilizing form of 1 SELF-FE RTILIZATION OF PLANTS. (829 this plant has a paler yellow colour than the intercrossing form, and, moreover, the red tint at the base of the latter is also wanting. Fumaria capreolata, var. pallidiflora, is, according to Müller's belief, restricted to self*fertilization *. A fact connected with white varieties is that they generally transmit their colour much more truly than any other variety (Anim. and Pl. under Domest. ii. p. 20), and rarely vary into any other colour. Mr. Darwin says he has found the white varieties of Delphinium Consolida and of the Stock are the truest; so is the white var. of the Sweet Pea according to Dr. Masters. Now the annual species of Delphinium are easily self- fertilized, though by Mr. Darwin’s experiments are not quite so fertile as when crossed, and the Sweet Pea is highly self-fertile. And when we remember that the most widely dispersed British plants are white, and also self-fertile, we seem to arrive at the clue to the fact stated at the beginning of this paragraph, namely, that white colour is cor- related with a more or less facility to self-fertilize. Hence the stability of the colour; for Mr. Darwin has shown how any strain can probably be retained by perpetual self- fertilization, all crossing being carefully prohibited. 4. v. Self.fertilization may be secured by the flowers closing after expansion. It has been suggested that some flowers close at such periods of the day or night when they willnot be visited by their proper insects, so as to preserve the nectar or honey. I would, however, suggest another and much more important object as being gained, and that is self-fertilization. Buttercups which are self-fertile, as Ranunculus acris, which Mr. Darwin records as producing plenty of seeds under a net, cannot fail to press the stamens down upon the pistil as they close, and thus secure the impregnation of the . earpels. Many flowers, e. g. Mesembryanthema, close as soon as the direct sunlight ceases to shine upon them, but long before insects would fail to see and visit them. Our own Anagallis arvensis, though very conspicuous, may be also aided by the rapid closing of the corolla; it appears, however, to be self-fertilizing irrespective of that fact. The order of development of the whorls is calyx, stamens, pistil, corolla. The pistil overtakes the stamens, and the stigma matures at the dehiscence of the anthers which cluster round it, especially by the closing of the corolla. Pollen-tubes were easy of detection. Erythrea Centaurium and Chlora perfoliata both appear to facilitate or bring about self-fertilization by the corolla folding round the anthers and stigmas which are thus brought into immediate contact. Other plants close in the evening. Light would seem to be the chief cause at work; for flowers which have closed at dusk will often reopen in artificial light, as do Crocuses. Mr. T. Meehan has arrived at exactly the same con- clusion as myself, our observations having been quite independent. (‘The ` Sleep of Plants’ as an Agent in Self-fertilization," Proc. of Acad. of Nat. Sc. of Philadelphia, 1876, p. 84.) * Mr. J. Traherne Moggridge drew attention to the fact that the colour of the corolla heightened after fertiliza- tion (* Nature, April 2, 1874). This raised a discussion as to the use of colours not being solely for insect agency. May not the interpretation be that as the ovule develops into seed, the heightening of the corolla was an accidental correlation with the increase of nourishment now determined to the flower? 390 REV. GEORGE HENSLOW ON THE This condition of subsequently closing after flowering may be regarded as a particular. 1 ! instance of a more general principle, of which the next (no. 4. vi.) as well as the case no. 7 i are special examples. | Another remark may be, perhaps, appropriate here, namely, concerning the use of i stamens being epipetalous or epiphyllous, as the case may be. When a corolla expands 1 under sunlight, and is thus prepared to receive the visits of insects, it necessarily will — cause the stamens to spread away from the pistil; but on closing again, as if it had — never expanded at all, the corolla brings the stamens forward, and the anthers now come in contact with the stigma. ` 4. vi. In some cases flower-buds never open. That flowers can and do fertilize them- — selves by not opening at all, is a fact of frequent occurrence. It often occurs in cold — and inclement weather, and late in the season, as autumn, and frequently in the winter — if it be mild. I examined a large number of unopened and half-opened buds in the Kew ] : Gardens in the autumn of 1876, such being merely the last attempts of the plants that — bore them to blossom. The pollen was abundant, and escaping from the anther-cells, — but not from the buds; and the pollen-tubes were in nearly all cases penetrating the — stigmas freely. Thus of a plant of Gnothera biennis, a half-opened flower, about half an — inch long, had its anthers clustering round the base of the stigmas, which, in a normally — developed flower with a fully expanded corolla, are elevated considerably above them. 3 : The anthers had dehisced without the corolla expanding. The stigmas were viscid ` throughout their whole length, and the pollen-tubes were freely penetrating. Mr. Meehan observes (Gard. Chron. 1875, p. 327) :— "A large number of plants have their pistils covered by their own pollen before the flowers open. Of these species among Wistaria, Lathyrus, Colutea, Cercis, Genista, Phaseolus, Pisum, Linaria, Ballota, ` Leonurus, and some others. This is particularly the case early in the season ; later the pollen-sacs burst — more generally about the same time with the opening of the corolla." I have italicized one sentence, as it corroborates some of Mr. Darwin's observations on — Papaver vagum, Reseda odorata, and also Mr. Bennett's and my own, that a colder 3 temperature appears to favour or induce self-fertilization in such plants as are more 3 inclined to be proterandrous with a relatively warmer temperature. 1 Mr. Meehan goes on to remark :— : “ It may be objected that the covering of the stigma with pollen is not fertilisation, as it requires à 1 peculiar condition of the pistil to receive it. But pollen has a long vitality. Carriére has found its ` 4 fertilising power unimpaired after three months old on one species, and other cases have been recorded." .| H. Müller has observed the same fact; for example, he speaks of Chelidonium majus — as having the stamens and stigmas mature together, though it be often crossed, but that | in wet weather self-fertilization is secured in half-opened buds. Besides the, so to say, accidental occurrence of normally conspicuous flowers not open- ing in autumn &c., there are many inconspicuous flowers which ripen seed all the year round, excepting in very extreme weather and when covered with snow, and do so by retaining their buds in an unopened condition. Mr. A. W. Bennett has recorded some instances which he observed in the winter of 1860 (' Nature, Jan. 1860); and I can corroborate his remarks by instancing several other cases, e. g. Stellaria media. This A HRS ^ EC NN iz m SS Si MEET xix á Se es a Tee CR UN MERE ee AUS a. R SELF-FERTILIZATION OF PLANTS. 331 plant seeds abundantly as long as the weather is at all ** open." The rapidity in doing this, as with many other self-fertilizing plants, is very astonishing. Any flower assumed to be no. 1 will be shedding its ripe seed, while no. 3, in order of development, is a self- fertilizing bud. Spergula arvensis also seeds freely while in bud; but as soon as a warm sun-shining day comes in January the flowers will expand. Cerastium glomeratum, however, I found produced nothing but closed self-fertilizing buds, even in an intensely and exceptionally hot day in June 1876. "The plants were growing, too, in the middle of an exposed hay-field. Polygonum Convolvulus seems never to open its flower-buds. Hence many flowers, though opening in warm sunshine, often fail to do so, but still set seed freely on other occasions; and in all such circumstances self-fertilization is the object. gained. 4. vii. Flowers may become habitually self-fertilizing in the absence of insects specially adapted for intercrossing them. Although there is reason to suspect that when an annual or biennial, with dichogamous flowers, fails to receive the visits of insects, as when it is transferred to another country, it will become extinct, yet there are some cases where it appears, at least when under cultivation, that they may become independent of the visits of insects, and so propagate themselves by self-fertilization. The Papilionacez are specially constructed for facilitating the intercrossing of different flowers by the agency of insects. Mr. Darwin quotes the description of Mr. Farrer (1. c. p. 160) given in ‘ Nature,’ Oct. 10, 1872. But as all irregular flowers may be assumed to have been developed in adaptation to insects, I take this for granted. In England and N. Germany, however, the Garden Pea, Piswm sativum, is rarely, if ever, visited by insects; and Mr. Darwin observes, “It does not follow that the species in its native country would be thus circumstanced.” Lathyrus odoratus is invariably self-fertilizing in this country; and even when visited by tiumble-bees these insects do not appear able to depress the keel-petals sufficiently so as to expose the anthers and stigmas; but at Florence “it is the fixed opinion of gardeners there that the varieties [of the Sweet Pea] do intereross, and that they cannot be preserved pure unless they are sown separately." Phaseolus is another genus illustrating the same fact. P. multiflorus produced with . Mr. Darwin from 4 to 4 the number of pods when covered which were formed on uncovered plants. It is only morphologically sterile, for it can produce “remarkably fine pods " when the flowers are simply and mechanically moved. P. vulgaris, on the other hand, is * quite fertile " when insects are excluded. Mr. Darwin observes that *this difference in self-fertility between P. vulgaris and multiflorus is remarkable, as these two species are so closely related that Linnæus thought that they formed one.” Lastly, Lupinus luteus and L. pilosus seeded freely with Mr. Darwin when insects were excluded ; but the flowers of some species of Lupine will not do so in New Zealand unless artificially disturbed (see Z. c., note, p. 150). These, and such like cases, seem to indicate that certain plants, with conspicuous flowers, and specially adapted for intercrossing by insect agency, may, by transportation to foreign countries, and of cooler climate, where their particular insects are absent, 332 REV. GEORGE HENSLOW ON THE regain, or at least acquire, the power of self-fertilization. The probable immediate causes ` will be dwelt upon more fully hereafter. f Hence we see, on the one hand, that flowers adapted for intercrossing by insects may 1 become self-fertilizing in their absence, and yet retain their conspicuousness. On the other, as I surmise, they may in the course of many generations give rise to self-fer- | tilizing forms (species) by the retention of an arrested corolla and other structural alterations; as will be referred to in the case of Zpilobiwm parviflorum as compared with | E. hirsutum, and species of Spergularia Ke, l There would seem to be two other alternatives which may happen to such plants on the © failure of insects to visit and cross them: the one is to become anemophilous or wind- ! fertilizing. Such appears to be the case with Poterium, Littorella, Plantago, the Amen- f tifere (mostly), Grasses and Sedges. The other alternative is to perish entirely. And — this last has, I suspect, largely taken place under migration, as will be shown more fully — hereafter: ! 1 A propos of the connexion between the conspicuousness of flowers and the presence of | insects, Sir J. D. Hooker informs me that, speaking generally of the Arctic flora, it is remarkable for its many conspicuous flowers when compared with the Antarctic, and that the former is associated with insects, whereas the latter has but comparatively few. ` The Rocky Mountains would seem to illustrate an intermediate condition; for thus speaks Mr, Meehan (‘ Gardeners’ Chronicle,’ Sept. 11, 1875, p. 327):— | “ The flowers of the Rocky-Mountain region are beautifully coloured; but Fremont pathetically de- scribes the solitary Bee that rested on his shoulder at the top of Pike's Peak. On my first visit the. comparative absence of insects proved very annoying to the entomologists who accompanied me. It was a frequent subject of conversation, whether Fremont’s Bee was not apocryphal; and though, on. a visit some years later, some Humble-bees on Polygonum Bistorta, on Gray’s Peak, enabled me to do justice to the veteran explorer, the incident shows how rare such insects are. Indeed, the paucity of animal life of all kinds in the Rocky Mountains is well known; but there is no more scarcity of seed. in the coloured flowering plants than in similar ones elsewhere." Mr. Meehan also justly calls attention to the fact of spring flowers being abundant, but blossoming at a time when few or no insects are about; yet they all, without any remarkable exception, seed well. 4. vill, Plants may become self-fertilizing on the reduction of temperature. This has already been alluded to in speaking of flower-buds not opening, and so becoming self- fertilizing in the autumn and winter. But besides such cases of indigenous plants, Mr. Darwin records the fact of Eschscholtzia californica being physiologically self-sterile in Brazil, but which, when transferred to and grown from seed in England, became self- fertile. Moreover, when transported to Germany, it again became self-sterile, as also did English-grown plants when sent to Brazil Similarly, Abutilon Darwinii, * which is self-sterile in its native home of Brazil, became moderately self-fertile in a single generation in an English hot-house.". In allusion to the Poppy, Papaver, he found some plants are self-sterile during the early part of the year, but later in the season become self-fertile. ** P. vagum produced plenty of capsules in my garden when insects were excluded, but only late in the season" (p.108). Mr. Meehan, as quoted SELF-FERTILIZATION OF PLANTS. 333 above, noted a corresponding fact of plants being self-sterile in the spring or early in the summer. Müller remarks the following curious fact about Cardamine pratensis (* Befruchtung, Ee p. 134) :— “In fine weather the anthers of the tallest stamens twist outwards on dehiscing, and are thus favourably situated for the removal of the pollen by insects. In cold and rainy weather, however, one often finds flowers in which the twist is much slighter, or does not exist at all; consequently the pollen of the taller stamens then falls on the stigma.” He, moreover, adds that, in most flowers, one finds both the shorter stamens below the. stigma; but in some they are rather longer, and in others of the same height as the stigma. In these latter flowers, therefore, even the shorter stamens secure self- fertilization. Scrophularia nodosa, Müller observes, sets seeds abundantly in cold weather and in the absence of wasps. Mr. Darwin, in an article on ** The Sterility of Plants from changed Conditions of Life and from other Causes " (Anim. and Pl. under Domest. ii. p. 163), says that ** Linnzus long ago observed that Alpine plants, although naturally loaded with seed, produced either few or none when cultivated in gardens." There are, however, exceptions. One cause may be suggested besides the absence of necessary insects, namely, that the warmer temperature of the garden may have brought about sterility, as was the ease with Eschscholtzia. 4.ix. Grafting. This appears to be a means by which self-fertilization may be reco- vered; for Mr. Darwin records the fact that Passiflora alata, from being naturally self- sterile, became self-fertile when grafted upon another species (Anim. & Pl. under Domest. ii. p. 164). 4*. On the origin of conspicuous flowers, and. the causes which induce reversion to self- fertilization. Having enumerated the different ways by which self-fertility is regained, it will be advisable to explain here what I believe to be the immediate causes ofits recovery. In speaking of the “immediate cause ” of self-sterility, Mr. Darwin attributes it in most cases to * the conditions" to which the plants have been subjected. These should, I believe, be rather called the “ proximate causes,” the immediate being, as it seems to me, the undue preponderating influence of the exterior whorls of the flower—that is to say, the stamens and corolla especially, and, in addition to them, the glandular secreting organs. These, by being greatly stimulated by the repeated visits of insects, tend to become hypertrophied.’ Hence the corolla enlarges, becomes more brightly coloured, the nectariferous organs increase the quantity of secretion, and the stamens develope more pollen. Such being the case, nourishment is withheld from the pistil, which is delayed in its development; consequently such a flower is very generally proterandrous. . Mr. Darwin rejects the notion that Natural Selection has favoured self-sterility in order to prevent self-fertilization, and finally comes to the conclusion that * we must look at it as an incidental result." Now this somewhat tallies with his modified views respecting Nature that she ** abhors perpetual self-fertilisation." He remarks (J. c. p. 8), * If the word perpetual had been omitted, the aphorism would have been false. As it stands, I believe that it is true, though perhaps rather too strongly expressed." Mr. Meehan goes further, and I am prepared to go with him; for that observer remarks :— SECOND SERIES.—BOTANY, VOL. I. SA H 334 REV. GEORGE HENSLOW ON THE * We thought Nature had a horror of in-and-in breeding. Our selected breeds of cattle are the result of this sort of selection, and they have proved just as healthy and productive as the veriest scrub. But it was thought they would at least revert to their original form when the hand of man was taken away. Prof. Brewer, however, showed that this was also a mistake **. Quite recently Mr. George Darwin has shown, in a remarkable paper, made up of an extensive study of old families among the English — nobility, where marriages among relations have been a source of social necessity for ages, that the popular ` idea is erroneous. These intermarriages have resulted as productively and as healthily, mentally and — morally, as the average marriages of the rest of the world” t. I am not in a position to be able to refute or endorse these latter remarks, upon which | I am not competent to form an opinion; though perhaps it is worthy of observation | that plants and vertebrate animals are not on a parallel stage of development; for the ` highest plant, by its composite character of powers of budding, each bud being prac- tically a new individual, would seem to be more on a level with a low invertebrate i animal; and therefore what may be true for the highest plants may be equally true for 1 the lowest animals, though the highest of the vertebrates may be far from exhibiting ` similar or parallel phenomena with plants. : Respecting the general prevalence of self-fertilization in the vegetable kingdom, how- ever, I agree entirely with Mr. Meehan; and with reference to the immediate cause of ` self-fertility or sterility, I believe they are due to the existence of and want respectively | of what I would call homogamic equilibrium between the andreecium and gynecium, | at least as far as morphologically self-sterile plants are concerned. The peculiarities of ` pollen which is impotent upon the pistil of the same flower (that is, when the flower is - physiologically self-sterile) are too subtle for any analysis at present known, and can only | be described by the indefinite phrase of undue or excessive differentiation. What led me to arrive at the above conclusion was a series of observations on the | emergence and growth of the whorls of flowers. I found the following to be the usual 1 order of development with flowers having conspicuous corollas. First, and invariably 80, appeared the sepals, which always grew to a size very considerably in advance of and much larger than the remaining organs. Secondly, the stamens opposite the sepals; and if there be two whorls, then the stamens opposite the petals. Thirdly, the pistil, and, — last of all, the corolla. It is, however, a matter of extreme difficulty to detect the exact | moment of the emergence or origin of the corolla; and in the majority of cases it pro- - bably emerges immediately after the calyx; but when this is the case, it seems to be immediately arrested, or at least the stamens grow so rapidly, the anthers enlarging, ` especially while every thing else is rudimentary, that the corolla is much delayed in com- parison with the stamens. It is not until the latter are nearly completed, and the pistil much advanced, that the corolla regains its power, and then rapidly enlarges. : On the other hand, with inconspicuous flowers, such as small-flowered Crucifere, the pistil may emerge simultaneously with or even before the stamens; and then it grows very rapidly, and matures its stigma either just before or synchronously with the dis- persion of the pollen. * Prof. Wrightson, of the Roy. Agr. Coll. Cirencester, informed me that in-breeding i is constant! pesto with short-horns, but with no deteriorating effects. d T ‘Gardeners’ Chronicle,’ Sept. 11, 1875. SELF-FERTILIZATION OF PLANTS. 335 These observations led me to see a special correlation between the pistil and the other floral whorls. Hence, if a flower be proterandrous, whatever may tend to lessen the energy of growth of these outer whorls tends at the same time to direct it into the pistil, which now advances, and is able to develope simultaneously with the Manon; and so self-fertilization is the result. Now, on referring to the headings 4, i. and 4, iv., it will be seen that these two causes of self-fertilization would seem due to the reduction or arrest of the corolla. Such, how- ever, may not be the sole cause, but only the index of an impaired condition or, at least, a lowering of the general vitality of the plant. Thus the withering of the corolla and part of the andreecium of Tradescantia implied a want of energy in those parts, in eonsequence of the lowness of the autumn temperature. Similarly, the partial arrest or complete loss of colour in the corolla is another indieation of lessened vegetative vigour, just as albinism in the animal kingdom indicates the same fact. There is reason to believe such flowers are more inclined to be self-fertile than others, as they are also more permanent. It must be remembered that while feebleness of constitution may induce self-fertility, the converse is not true. Self-fertility does not induce feebleness. If this be the true rationale, we see that self-fertilization and self-sterility are resolved, so far at least, into a simple question of compensation, which, in turn, is a question of nutrition; and it has nothing to do with any supposed injuriousness whatever. In fact -it is an absolute loss (and therefore an “injury ") to a plant to be self-sterile, for its facility for propagating is largely, if not entirely, checked *. This is notably the case with the Scarlet Runner (Gard. Chr. vol. x. p. 561). Though I agree with Mr. Darwin, “ that the inefficiency of a plant's own pollen [phy- siologically] is in most cases an incidental result [due to differentiation], and has not been specially acquired for the sake of preventing self-fertilization," I cannot agree with him in drawing a different conclusion for morphologically self-sterile plants; for he adds :— * On the other hand, there can hardly be a doubt that dichogamy ... . that the hete- rostyled condition of certain plants, and that many mechanical structures, have all been acquired, so as both to check self-fertilisation and to favour cross-fertilisation.” And yet he proves that when plants lose their dichogamy they regain self-fertility, which Mr. Darwin then recognizes as an advantage to the plant; while elsewhere he says, “ it is difficult to avoid the suspicion that self-fertilisation is in some respects advantageous.” My impression is that flowers were primordially hermaphrodite, inconspicuous, and self- fertile. If I might venture further into the regions of speculation, I would suggest, as a feasible hypothesis, that insects having been attracted to the juicy tissues of such flowers, by perpetually withdrawing fluids have thereby kept up a flow of the secretion, which has increased, and so developed into glandular and regularly secreting organs. The mere puncture and lesion caused by an insect would not of itself be hereditary, just as galls do not form spontaneously ; still even with them there may be, for all we know, a predisposition to form them, and they may be now perhaps much larger than they were * Since this paper was read, one by Mr. Meehan has been printed in the Journal of the Linn. Soc., in which he shows how Wistaria, though it never sets seeds, when “trained,” yet, when grown as a “ tree," does at expend ita energy in forming long branches, and consequently fruits abundantly. 8a 2 336 REV. GEORGE HENSLOW ON THE when insects of any particular species first punctured oaks. It is well known that in q | the human subject there may be a predisposition for tumourous or cancerous growths ` which is hereditary ; and there would seem to be a very close resemblance between 1 | tumours and galls, both being hypertrophied conditions of certain normal tissues. For ~ example, Sir B. C. Brodie thus describes a fatty tumour :—“ There is no distinct boun- - | dary to it, and you cannot say where the natural adipose structure ends and the morbid 1 growth begins." Such is very much like the growth of galls, which is due to cell-division 1 | setting in at certain points of the epidermis and subjacent tissues. But although lesions 1 | and mutilations will not, as a rule, prove to have any hereditary effects, yet the constant 1 | drain upon a secreting organ may. A mutilation being once made, the place heals, and | there is an end of all vital action; but if a constant drain be kept up upon an organ 1 | thus irritated, and made to secrete, there will be a corresponding flow of nutriment to 1 the place, which must also affect more or less the adjacent parts. j Mr. Darwin, in speaking of the Cow, observes :—‘‘ We may attribute the excellence 4 of our cows, and of certain goats, partly to the continued selection of the best milking 4 animals, and partly £o the inherited effects of the increased action, through man’s art, of — the secreting glands” (Anim. & Pl. under Domest. ii. p. 300). 1 This fact, recorded in the last sentence, which I have italicized, I take as the basis of 3 my hypothesis :—That insects again and again visiting the same flowers will thus cause a 1 constantly repeated flow of nutrition, which affects (I assume) not only the glands them- 4 selves, which thus become hypertrophied, but the adjacent parts, such as the corolla and ` stamens ; and there would be an à priori probability of vegetable structures being thus ` more likely to be affected than animal, because there is no well-defined or bounded ` channel to conduct the sap solely to the one point of irritation. Such, then, I would d advance, hypothetically, as a vera causa for the origin of all conspicuous flowers. E Natural selection might come into play and determine the final result. E We might, perhaps, go a step further, and speculate as to the origin of irregularity 13 in flowers, by regarding them, so to say, as having a kind of plasticity which will enable ` them to respond to repeated irritation, in some sort of way analogous to that of tendrils of ` the Virginian Creeper, which also respond to mechanical irritation, then develope secreting organs and become hypertrophied. So that as long as insects visit a flower they are continually keeping up a sort of irritation at that region, the whole weight of the insect being often thrown upon the corolla. If it be terminal, the insect alights on any petal or petals, and nothing induces the flower to become irregular* ; but if the flower be axil- lary, it alights on the anterior side, and so (I assume theoretically until it can be demon- strated or disproved) brings a stimulus which is responded to by the inherent forces of the plant, and which causes the flower to become bilateral, by determining a super abundant flow of nutriment to that part. What it is that determines the peculiarities of structure in each flower respectively, causing irregular flowers to be so very different, ` it is quite impossible to say. This constant irritation and the continual drain iu the | secretive organs must stimulate them to develope more and more, just as à man’s arm | ... * Of course I do not mean Ge any single flower is thus affected, but that the ros is "€ acquired after many generations. SELF-FERTILIZATION OF PLANTS. . 337 increases by work, or as the mamme may be made to secrete for prolonged periods ; so that, in my view, it is not that insects have gone to the flowers, because they were first conspicuous, but have actually themselves determined their conspicuousness. The final result has been that nourishment has been delayed from the pistil, and the flowers have become dichogamous and proterandrous. | That there should be no à priori objection to the idea of an external mechanical structure giving rise to organie changes in the tissue of a plant is obvious from such well- known cases as tendrils and carnivorous plants, as also from Mr. H. Spencer's experi- ments with Cacti, which showed an increase of vascular tissue when subjected to unusual mechanical strains. Perhaps these and other * responsive" activities may hereafter prove to be due to a widely extended principle of reflex action, to which Mr. Darwin attributes the curvature of the tentacles of Drosera. On the other hand, in the absence of insects, there is no such increase of energy in the corolla, &c., and the balance is restored with the result of self-fertilization. — A restoration to a homogamic equilibrium, then, I take to be the true explanation of pale varieties of Pelargonium being great seeders, and of Dianthus and other dicho- gamous flowers becoming highly self-fertile. Mr. Darwin records of Dianthus caryg- phyllus that it became highly self-fertile in three generations, and that the value of its fertility, and that of intercrossed plants, was as 125: 100. It was similarly with varieties of Primula veris and P, Sinensis (to be alluded to again). And I have myself found a wild Primrose with the style (of a “ short-styled form”) elongated so that the stigma was surrounded by the anthers at the orifice of the tube, and which was “setting” seed abundantly. Ido not, therefore, see how the conclusion can be avoided that self-fertilization is per se a decided advantage, and that intercrossing, as far as the production of seed is only regarded, a compensatory process for the loss of self-fertility. A similar explanation will, I think, apply to proterogynous flowers. If we regard homogamie equilibrium as the normal condition of hermaphroditism, then the pro terandrous state, as shown, results from the energy preponderating in the whorls external to the pistil; but if these be more or less suppressed, as seen in what is called contabescence of the anthers (Anim. and Pl. under Domest. ii. p. 165), which will be spoken of again below, there is a corresponding gain to the pistil, so that it may emerge con- jointly with the stamens, as with Nasturtium officinale, Lepidium campestre, or it may precede them in emergence, but mature together, as in Cerastium glomeratum, Arenaria trinervis and serpyllifolia. If, however, the pistil not only starts first in the race but retains the lead and ultimately matures its stigma first, then proterogyny is the result. Now as far as it has been observed as yet, proterandry is recognized as being by far the commonest condition amongst conspicuously flowering plants; white proterogyny is the exception with large-flowering dicotyledonous genera. Thus Helleborus viridis is, according to Hildebrand, proterogynous,. and here the corolla is replaced by small nectariferous tubes. Geraniwm pratense and other large-flowered species are pro- terandrous. G. pyrenaicum matures its pistil conjointly with the inner whorl of stamens; 338 REV. GEORGE HENSLOW ON THE but in G. molle it matures it before that whorl, and is therefore proterogynous; and 1 these stages correspond exactly with the decreasing size of the corollas. | 1 Scrophularia nodosa is proterogynous and has a very small flower when compared 1 with the conspicuous forms of Digitalis and Linaria &c., which are very strongly pro- j terandrous. Armeria Statice and Plantago, whose corollas rapidly dry up and become ` scarious, would seem to allow of nourishment being soon directed into the pistil, which q in these genera is decidedly proterogynous. The same remark applies to the Juncacee. 1 Asarum and Aristolochia, again, as also some plants of Corylus Avellana, are protero- d gynous, this last clearly showing that it is not solely due to such a compensatory 4 process as I have supposed, though it may be still a question of.nutrition. Arum, Pota- q mogeton, Triglochin, and Paris are ‘all proterogynous and inconspicuously flowering 1 plants. On the other hand, Colchicum is said to be proterogynous, though bearing a. 1 large flower, but Butomus is proterandrous. Lastly, Prunus spinosa and Padus, as well 7 as Orategus oxyacantha, are proterogynous. Perhaps the lateness of the season in which 1 Colchicum flowers, and the earliness in spring when the genus Prunus is in bloom, may | account for the comparative delay in the stamens, thus bringing about a relatively | earlier development of the pistil, resulting either in self-fertilization or proterogyny. 1 "Finally, I would observe that the causes which may bring about self-fertility appear ` to be very complicated, and that very likely no one cause stands alone. Therefore, while 1 I have enumerated what I could designate prowimate causes, yet they may be perhaps: 4 better regarded as correlative phenomena than as being so many distinct causes. The | primary or fundamental cause (both of them, e. g. loss of colour and of self-fertility) - lies apparently too deep for discovery in the constitution of the entire plant. We must, however, be on our guard against interchanging causes and effects. It is one thing to say that continued self-fertilization may bring on a deterioration in the constitution (which, I think, is not proved to be the case), another thing to believe that a seemingly less vigorous condition of vegetative growth may induce a return to self-fertility i in dicho- gamous and other flowers, such being in some cases, I believe, the real interpretation. The intimate relationship between vegetative and reproductive vigour is too well known to require any elucidation; and all I would add to that compensatory process is, that in proportion as the former is lessened so does the return to self-fertilization follow, that being the special form of reproductive energy which, I believe, Nature vmm strives toregain. (See also Anim. and Pl. under Domest. ii. p. 163.) . Moreover, it must be borne in mind that great vegetative * vigour' is not NAR a good sign of any lasting benefit to the plant, just as giants among men are often. not so well favoured by nature as men of mean height. This idea of the existence of a compensatory process between the outer whorls e the pistil is further countenanced by an analogous one between the ovary and the seed. An interesting and important paragraph by Mr. Berkeley illustrates this fact so well that I quote it in full from the * Gard. Chron.’ (1855, p. 36) :— PTT * Though all the parts of a plant may be perfect, and impregnation may take place, and an embryo ! be formed, the plant may be WE from constant abortion. This is especially the case where the — of SELF-FERTILIZATION OF PLANTS. 339 the ovary are highly developed, insomuch that in some of the finer varieties of Pears &c. perfect seed is of comparatively rare occurrence; while, on the contrary, in groups attacked by mildew the seeds are in general multiplied. In such cases, as the properties which make the fruit valuable are altogether inde- pendent of the seed, it is sufficient for the purpose of the cultivator if impregnation have taken place to such an extent as to ensure the swelling of the sarcocarp. It is to this abortion that many cases of sterility aredue. The process of impregnation is so far successful as to stimulate the contents of the embryo-sac and the fleshy walls of the fruit ; but after a time the embryo ceases to grow and the sarco- carp withers... . . The sterility is by no means due to deficient impregnation, but apparently to a greater degree of heat than its growth requires, which stimulates other parts at the expense of the embryo.” Mr. Berkeley also remarks, that * The influence of the pollen is not the only influence which will cause the succulent portion of the organs of fructification toswell. The process of caprification * is an instance in point, as also the touching the orifice of a fig with oil, or piercing the young walls with an oiled straw." 5. Highly self-fertile varieties may arise under cultivation. For this important fact I am indebted to Mr. Darwin; and I propose giving under this heading a brief summary of certain facts collected from his work on Cross and Self- fertilisation of Plants. Although in the majority of cases it appears undoubtedly true that the offspring of cultivated plants erossed by pollen from a different plant of the same, and still more of a different stock, become more fertile than such plants when self-fertilized, yet Mr. Darwin raised during his experiment some highly self-fertile forms, which “ yielded more seed and produced offspring growing taller than their self-fertilized parents, or than the intererossed plants of the corresponding generation." ‘The following examples will illus- trate this fact. 3 In cultivating Zpomea purpurea, with the purpose of contrasting the heights of the intercrossed and self-fertilized plants, it was not until the sixth generation was raised that a single plant of the latter beat its competitor; that is to say, about forty pairs of such plants had been cultivated before one of the self-fertilized surpassed its rival, the heights of these two being respectively 87 and 86:5 inches, or as 100:99:4. Mr. Darwin was so much surprised at this case, that he saved the self-fertilized seeds of this plant, which he called the ‘ Hero,’ and experimented on its descendants, with the following results, summarized :— (a) Its descendants inherited a power of growth equal to the ordinary intercrossed ; for the first generation, i.e. the children of Hero, self-fertilized, were to the intercrossed as 100 : 95. (b) A cross between the grandchildren of Hero did no good. (c) The descendants became more fertile than was usually the case; for the self- fertilized grandchildren of Hero had a higher average of seeds per capsule than was observed in any other case with self-fertilized plants. (d) No benefit was derived from intercrossing with plants of the same stock. * Gasparrini, however, denies that caprification has any effect other than impregnation (Journ. Hort. Soc. Lond. iii. p. 185). 340 REV. GEORGE HENSLOW ON THE (e) No benefit was derived from crossing with plants of a distinct stock. 1 Besides this Hero, which appeared in the sixth [aj rbi and which, as stated, beat 1 its opponent by about half per cent., two other “ heroes” appeared in the eighth gene- 1 ration, the ratio of whose heights to those of their competitors were as 111:3 : 100, and 3 as 140:5 : 100; but in other respects than height they do not appear to have been such fine plants as their opponents. Their offspring were not cultivated. 4 The next remarkably self-fertile form to be noticed was a white or pale variety of. Mimulus luteus blotched with purple. This first appeared in the third generation, and. 1 Mr. Darwin observes on it :—* From the tallness of this variety, the self-fertilised plants | 1 exceeded the crossed plants in height in all the generations from the fifth to the seventh | inclusive." Moreover, this tall pale variety increased in the later self-fertilized gene- ` 4 rations, owing to its great self-fertility, to the complete exclusion of the original kinds. 1 The average ratio of the heights of this variety to that of the intercrossed was about. 126 :100; and the ratio of fertility in the sixth generation was as 147: 100, i. e. the ] production of capsules; while the number of seeds of the former “ appeared decidedly - more numerous than those from the crossed plants." As with the descendants of Hero 1 intercrossing with plants of the same stock did not benefit this variety. But when this highly self-fertile variety was crossed by a new stock, unlike Hero, it was vastly ben fited, as Mr. Darwin shows by the following results :— Weight of seed produced by the same number of ** Chelsea-crossed " and intererossed | plants, as 100 : 4. Weight of seed produced by the same number of Chelsea-erossed | 1 and self-fertilized plants, as 100: 3. Weight of seeds produced by the same number of intercrossed and self-fertilized plants, as 100 : 73. Nicotiana tabacum is a third example; for the flowers on the parent plants, whid were self-fertilized by Mr. Darwin, “ yielded half again as many seeds as did those whiel were crossed ; and the seedlings raised from these self-fertilised seeds exceeded in heigh those raised from the crossed seeds to an extraordinary degree." Reseda odorata and R. lutea. Some plants of these two species are incompara : more self-fertile than other individuals. Pisum sativum. | ** The cultivated varieties of the common Pea are highly selí-fertile although they have been self-fertilised for many generations; and they exceeded i height seedlings from a cross between two plants belonging to the same co in th ratio of 115 : 100." Four pairs only, however, were compared. Let _ Primula veris. Of this species Mr. Darwin observes, “ The self-fertility increase dus several generations of illegitimate fertilization, which is a process closely analogou to self-fertilization, but only as long as the plants were cultivated under the same favou able conditions." He adds, ** I have also elsewhere * shown that with Primula veris and sinensis, equal-styled varieties E appear which possess the sexual organs of the two forms combined in the same flower. Consequentl they fertilise themselves in a legitimate manner and are highly self-fertile ; but the remarkable factis | that they arerather more fertile than ordinary plants of the same species legitimately fertilised by] len. from a distinct individual." uc * Best d Dun. Soc., Bot. x. 1867, pp. 417-419. - " SELF-FERTILIZATION OF PLANTS. 941 I can also personally corroborate part of the above facts, and have already described above a form of P. Sinensis which was highly self-fertile. I have, however, never met with an equal-styled P. veris; but I found a plant of P. vulgaris this year (1877) highly self-fertile, and which had an elongated style, so that the stigma was surrounded by the anthers at the orifice of the tube; that is to say, the position of the stamens indicated that it was of the short-styled form, but had become equal-styled and homogamous. It is, perhaps, worth adding, that just as there arise equal-styled self-fertilizing indi- viduals of normally heteromorphie species, so in Primula we have, besides many cases of the latter form, others normally equal-styled. Thus P. mollis is non-dimorphie and highly self-fertile, ** nearly every flower producing a capsule filled with good seed” *. Such being examples of the appearance of highly self-fertile varieties recorded by Mr. Darwin, a few inferences on the same subject may be drawn from a study of the tables given in his work. Table A contains 99 ratios of the heights of the intercrossed plants (always represented by 100) and those of their self-fertilized opponents. There are 54 species of 30 natural Orders represented. Deductiiig 3 cases of equality, there are 17 in which the self- fertilized exceeded the intercrossed, or over 17 per cent. Again, reviewing the separate tables of each plant, there are 8 out of 57 in which some one or more of the self- fertilized plants beat its intercrossed opponent, or nearly 65 per cent. The number of pairs cultivated varied from two to over thirty, and it is worth while examining the five highest numbers. Lobelia fulgens, 2nd generation, 34 pairs of plants were grown in 9 pots. In pots i, ii, and vi (4. c. p. 181), containing in all 12 pairs of plants, all the self-fertilized beat the intercrossed in height in the mean ratio of 116: 100. In pots-iii, iv, vii, viii, and ix, containing 18 pairs of plants, all the intercrossed beat the self-fertilized in the ratio of 100: 74 (nearly). Digitalis purpurea, plants raised from a cross between different flowers on the same plant were grown with plants raised by self-fertilization. Of 28 pairs, 10 self-fertilized plants beat their opponents, or nearly 36 per cent. beris umbellata, of 30 pairs, 5 self-fertilized plants beat their opponents. Reseda odorata, seedlings from a highly self-fertile plant were grown in 5 pots. Of 19 pairs, only 2 self-fertilized plants exceeded the intercrossed; but of 8 other pairs (of the same lot of seeds) grown in open ground, 5 self-fertilized plants beat their opponents. R. odorata, seedlings from a semi-self-sterile plant were grown in 5 pots.. Of 20 pairs, 7 self-fertilized plants beat their opponents; in one pot all the tallest were self-fertilized plants. Nemophila insignis, Cyclamen persicum, and „Limnanthes Douglasii, 12 pairs of each of the first two and 16 of the third were grown, and not a single plant of the self- fertilized beat its opponent. Of the second generation, however, of WV. insignis, out of 7 pairs, 6 self-fertilized plants were taller than their rivals. Certain inferences appear to be deducible from these facts. Recalling to mind how of 77 pairs of plants of Ipomea purpurea, only 3 self-fertilized plants were taller than their rivals, and that when Mr. Darwin cultivated one of these three especially (and which beat its rival only by about 5 per eent he raised a highly self-fertile form, which proved * This statement is on the authority of Mr. J. Scott, Journal of Linn. Soc., Bot. vol. viii. 1864, p. 119, . SECOND SERIES.— BOTANY, VOL. I. | 9B. 342 . REV. GEORGE HENSLOW ON THE. * to be superior to the intererossed plants in évery way, and, moreover, was not benefited l | by a cross with a new stock, one is led to infer that others of these numerous indi — viduals of the self-fertilized plants which beat their rivals might, on selection and cul — tivation, have proved equally as vigorous as “ Hero." It is not only when large 1 numbers of pairs of plants are cultivated that some one or more of the self-fertilized 1 individuals were superior to their rivals, but even when very few pairs were grown. 1 Thus, of 2 pairs of Passiflora gracilis, 1 self-fertilized plant was the taller. Of 4 pairs | of Borago officinalis, 2 such were taller. Of 5 pairs of Nolana prostrata, 4 such were ^ taller. Of 7 pairs of Pelargonium, 8 self-fertilized were also the taller ; and many other q cases might be mentioned. Yet another important inference may be drawn from such a case as Lobelia felgen 1 Of this plant there were 12 pairs in 3 pots, all the self-fertilized of which were superior; — and of 18 pairs in 5 pots, all the intercrossed were superior. Supposing Mr. Darwin had ~ only cultivated the first 3 pots, he would have inferred that intercrossing was injurious; 7 and had he cultivated only the last, he would not have known that self-fertilized plants 1 | could ever beat their rivals. Now, of about thirty-five cases he has not cultivated more 1 than 12 pairs of plants, and in only about ¢welve cases has he cultivated more than ` 12 pairs of plants according to the tables. Hence, while recognizing the value of inter- ` crossing generally in imparting height to the offspring, such a case as Lobelia fulgens 1 throws a certain amount of doubt over these thirty-five in respect to thé proportional 1 number of successful self-fertilized plants that would have arisen had "A numbers | of pairs been grown. q . Lastly, from the first table of Nemophila insignis, containing 12 pairs, we learn that 1 crossing does great good, as not 4 single individual of the self-fertilized beat its rival, the ratio being as 100 : 60; but when certain of these pairs of plants were placed under a net, and all allowed to fertilize themselves spontaneously, we find that in 6 out of 7 pairs, the self-fertilized offspring of two generations beat their crossed plants self- fertilized on the second generation. Mr. Darwin does not think the averages deduced from their heights trustworthy ; but as both kinds appear to have been equally cir- cumstanced, the heights alone may probably be so. DE Re 6. Inconspicuous flowers are almost invariably self-fertilizing, or else anemophilous. . . In chap. x. of his work on Cross and Self-fertilisation, Mr. Darwin has written section on inconspicuous flowers, and makes the following statement on p. 382 :— “There can hardly be a doubt that dichogamy .... —the heterostyled condition of certain plants, and that many mechanical structures—haye all been acquired so as both to check self-fertilisation. sitig favour eross-fertilisation.” | It does not seem quite clear whether Mr. Darwin would mean by “so as to” ' to signify a purpose or a consequence. 'To my mind it would be the latter; and the fol- lowing sentence strengthens this view :— "It might perhaps have been expected that plants having their flowers thus peculiarly constructed : would profit in Se ge acinis cdm wem WI bue FUSCO UU ER em Ray OR QUIE d E SELF-FERTILIZATION OF PLANTS. 343 And he compares Tropeolum minus and Salvia coccinea with Limnanthes Douglasii and others, as examples. Such * irregularities ;" as these flowers possess I regard as simply necessary consequences of the visits of insects which have brought them about; but the amount of * profit" gained by intercrossing would depend upon the degree of benefit derived from the infusion of new constitutional elements by the aid of foreign pollen. © The larger and more conspicuous a flower, the more frequent are probably the visits of insects, while inconspicuous flowers are rarely and, in many cases, never visited by them. This led H. Müller to say that they must be self-fertilizing or they would ‘become extinct; and Mr. Darwin supports his conclusion that small and inconspicuous flowers are completely self-fertilized. My own observations fully corroborate the state- ment. Mr. Darwin adds, ‘The converse of the rule that plants bearing small and in- conspicuous flowers are self-fertile, namely, that plants with large and conspicuous flowers are self-sterile, is far from true." This is, of course, equivalent to my former statement that very few flowers are self-sterile; and this speaks volumes in favour of the enormous advantage, as far as propagation by seed goes, of self-fertilization ; and let us not forget that propagation is not only, to use Mr. Darwin's expression, of “ paramount importance," but is the sole end of plant life. There are several reasons why inconspicuous flowers are not likely to be intercrossed by insects. 1, their unattractiveness; 2, the absence of honey-secreting organs; 3, the want of scent; 4, they frequently do not expand, or at most remain but half-open, espe- cially in cold or inclement weather, while perfectly cleistogamous flowers are, of course, never open; 5, their structure sometimes would seem absolutely to prevent the ingress of insects (such appears to be the case with Polygonum Convolvulus and P. Hydro- piper, the flowers of which seem to be always closed, and with many others). ~ Such being the case, whence is the origin of inconspicuous flowers? On inspecting a — list of British plants only, it will be seen how nearly, if not absolutely, every large order has some genera with inconspicuous species; or else there are some genera with incon- spicuous flowers in an order otherwise characterized by conspicuous flowers. The idea that they are further differentiated forms at once recommends itself. This is especially the case with such plants as Fumaria, Trifolium, Salvia, and many others; for certain species are highly self-fertile and quite inconspicuous, but yet have retained the form of the corolla so perfectly adapted for insect-agency in their more conspicuous allies. Hence I would venture to generalize, and say that all of our existing inconspicuous flowers are more differentiated than the latter * and are not primitive forms. I would herein add my belief that the Incomplete of Dicotyledons do not represent, nor are they survivals of primitive types, but further advanced states of “degradation.” Thus of the “ Cyclo- spermez" t, the orders usually included in the Incomplete represent very degraded forms of Caryophyllez. In other words, I believe self-fertilization was the primitive condition, * I will use the word “ degraded," though it be liable to misconception ; for the popular sense of the term implies some kind of “ degeneration." I shall regard it, however, as equivalent to the expression in the text, involving, moreover, the idea of reversion, so far as the reacquiring self-fertilization is concerned. ** Degradation” will there- . fore solely refer to the more or less Ee condition of the Um &e., and not at all imply degeneracy or impaired constitutional phenomena. T* Éléments de Botanique; par P. Duchartre, 2™ éd., p. 1135. aw "o 8522 344 REV. GEORGE HENSLOW ON THE and that the necessity for intercrossing, as far as propagation by seed is alone ĉon- i cerned, has arisen by insect agency, and that whenever or wherever the proper insects fail, one of the following results may happen :—(1) The plant will die out, especially if it be - an annual. (2) If a perennial, it may propagate itself vegetatively, and so maintain its | existence. (3) It may become readapted to other insects. (4) It may revert to self- : fertilization, (a) with a retention of a conspicuous corolla, as Pisum sativum, or (b) with a dwarfing of it, as in the case of Fumaria and Trifolium, or (e) lose it ge, or (a 1 become absolutely cleistogamous” — I strongly suspect No. 1 or No. 4 (b) to be usually the case with seedlings fe: ` to distant countries, as will be more particularly adverted to hereafter. Prof. Dyer, in his review of Mr. Darwin's work, ‘Cross and Self-fertilisation, in * Nature, thinks that cleistogamous flowers represent the primitive condition. I take an opposite view, | and regard them, as well as inconspicuous flowers, to be in all cases (excepting gymnda ] sperms) degraded conditions, and for the following reasons :— 3 (a) Had such features as are borne by cleistogamous flowers been characteristic of. primordial conditions, they would have most probably been correlated with embryonic | or primitive states of other organs; or, at least, such plants would be of a relatively low | grade. They are, however, found to exist in widely different orders, which are amongst the most highly differentiated, and in no other respect showing any thing whatever of a primitive character about them. (b) Every degree of degradation can be found between the normal Bowtie and the cleistogamous on the same plant. Thus in strong cultivated plants of Viola odorata, comparatively large cleistogamous buds occur with five petals, the posterior spurred, and the stamens with their nectariferous appendages, but completely self-fertilizing, while on wild specimens these flowers may be entirely apetalous. (Tab. XLIV. figs. 4a, b, and 5 a, b.) (c) Many plants have flowers in an intermediate condition, either opening when th weather is warm and favourable, but remaining closed if it be inclement, as do many of : the Alsinee, Illecebracee, Polygonum, Ze, P. Convolvulus and Hydropiper even appear to be already perfectly cleistogamous, though P. aviculare still opens its minute honeyless ` blossoms. Similarly, Cerastium glomeratum sometimes never opens its buds, at other times, and indeed scarcely ever, is more than half-open. Fumaria officinalis and. some small-flowered clovers, which are self-fertilizing, might almost be ealled permanently cleistogamous ; for, as in their more conspicuous allies, which are intercrossed, the sexual organs are completely concealed, but in their case do not require insect agency. As such flowers, though extremely inconspicuous, have yet irregular corollas, they are clearly derived from some ancestral form, from which their conspicuous allies have also been descended ; and it becomes therefore a moral conviction that they are ee con- ditions from more highly conspicuous forms. 3 I do not think the Amentifere form any exception to this rule; for in many there is a regular calyx, with the stamens opposite the sepals, which seem to indicate that they are degraded conditions with the corolla gone. Thus, if we take Ulmus as our starting point, we have a abr five-lobed itr and fixe: stamens suet, the s a * SELF-FERTILIZATION OF PLANTS. 345 Next, selecting Alnus, the sexes are divided; but of the four sepals each has a stamen opposite to it. In Betula, the male flowers are heaped together in an irregular sort of manner; but still a stamen is in front of each sepal. In Populus, the limb of the calyx is now arrested, and we get a still further degradation, which culminates in Saliz*. The Cupulifere furnish another case of degradation, but from an offshoot from the Amentiferous genealogical tree. This group had originally advanced to a more highly differentiated state, as is proved by the inferior ovary; but here, again, the calyx of Quercus and Fagus, with stamens opposite to the sepals, indicates the same loss of a corolla, . If it be asked what is the evidence of the corolla having been totally arrested instead of its never having been formed, on the assumption that these were primitive types rather than degraded ones, the reply is that existing analogies seem to prove it. Thus we see the loss of it in members of the Alsinee, such as Sagina, Spergularia, &c., being both petalous and apetalous. In the latter condition the stamens opposite the petals disappear as well as the petals. They are the last to develop and are the first to go. Sueh (and many others might be quoted, where some species are apetalous, others petalous) would seem to support the idea that both a whorl of stamens and the corolla have been long lost in the case of the Amentiferee ; and in the Monochlamydez generally, wherever two whorls of stamens occur, as in Daphne, it seems to indicate the loss of the corolla only, while the different heights of the two staminal whorls in this plant may point to an ancestral heterostyled condition which perhaps no longer exists. I cannot therefore accept Mr. Darwin’s conclusion, that some plants “ have actually had their flowers reduced, and purposely rendered inconspicuous;" for I take it to be simply and purely, or at least mainly, a result consequent upon the absence of insects— just as, conversely, the conspicuousness of corollas, the development of secretive organs, dichogamy, and unisexuality are direct consequences of the disturbance in the floral equilibrium brought about by the actual visits of the insects themselves +. I think we should be very cautious not to confound means with ends. If I under- stand Mr. Darwin aright, he seems sometimes to look at the machinery for intercrossing as a purpose for benefiting the plant; elsewhere he shows and states clearly that inter- erossing per se does no good unless it bring constitutional differences; so that we must keep clear these two facts :—(1) The act of fertilization per se, as having its sole end the propagation by seed; (2) The constitutional benefits acquired through the agency of crossing. . With regard to the first, self-fertilization, as Mr. Darwin acknowledges, “is incom- parably the surest method ;” and for that purpose intercrossing is hazardous, if we do not use the word injurious. With regard to the second fact, it is clear that self-fertilization cannot introduce new * Authors regard the little prominence at the base of the stamens and of the pistil in Salix either as the calyx or ‘else as the axis. It appears to me to be simply a cellular gland for secreting honey, which they do abundantly. 1 could detect no spiral vessels in them at all. + If we say that such structures appeared spontaneously, in anticipation, as it were, of the visits of insects, and so determined their coming, we at once fall into the old and objectionable teleological methods of interpretation. 946 REV. GEORGE HENSLOW ON THE constitutional elements, though, as with British weeds in New Zealand, they may acquire — by their introduction into new climatal conditions fresh constitutional vigour, superior to the native vegetative population; for the White Clover is actually ousting the New- ` Zealand Flax, Phormium tenax. Similarly, if the process had been reversed, analogy l would lead us to suspect that the New-Zealand vegetation might prove to have the mastery in competition, perhaps, with the very same genera and species now growing in ` this country. Anacharis Alsinastrum supports this idea. — — E Plants which have acquired the property of intercrossing gain immense advantage in 1 securing the introduction of new constitutional vigour by the infusion, so to say, of fresh | blood into their system. And this vigour may show itself in larger foliage, more | branches, and therefore more flowers, and in more fruit and seed; but there still 1 remains a remarkable fact, which Mr. Darwin has shown, that the number of seeds per | capsule often remains the same. In some orders and genera this, of course, must be so, | as with the Labiate, Boraginee, and Linum ; but it was also true with many others of 3 which the seeds were nüierous. On the other hand, even with conspicuous plants, several ` in Mr. Darwin's experiments became more fertile with their own pollen, though adapted ` for intercrossing, as did Mimulus luteus, Ipomea purpurea, and Nicotiana; while Ophrys 1 apifera produces very large capsules, with an enormous améunt of seed. This last - species is clearly a self-adapted form of what represents in other Orchids invariably inter- | crossing peculiarities *. Lastly, all inconspicuous self-fertilizing forms, as elsewhere ` stated, are great seeders. Mr. Darwin says (7. c. p. 386) :— “ It seems to me highly improbable that plants bearing small and inconspicuous flowers have been or 1 should continue to be subjected to self-fertilisation for a long series of generations. I think so, not . from the evil which manifestly follows from self-fertilisation. . . . . . . . But, if plants bearing | small and inconspicuous flowers were not occasionally intercrossed, and did not profit by the process, - all their flowers would probably have been rendered cleistogene, as they would thus have largely benefited — by having to produce only a small quantity of safely-protected pollen ” [my italics]. : But plants do not pass from one extreme to the other all at once. We do not know - all the conditions requisite for producing completely cleistogamous flowers; but as, in ` the case of Violets and Oxalis, they are concealed under dense foliage, and with Lamium E amplexicaule it is the early and late blossoms, it would appear to be certain external : conditions of a lessened amount of light and heat which are requisite. Small flowering ` plants, though having flowers more or less approximating the cleistogamous state, are S only partially so, because they are still in an intermediate condition from their habit of growth, the conditions not being requisite to -reduce them to complete cleistogamy. ` Thus Stellaria media and Spergula arvensis wil open in sunshine, but in shade or in ` winter are often as completely cleistogamous as Ozalis, —. E In the heading to this section I have introduced the qualification ** almost invariably ;” | * Great stress is laid by writers on the increased fertility gained by intercrossing. But it must be borne in mind | that this increase is relative, not absolute. It is quite reasonable to suppose that plants with conspicuous flowers, and | demanding inscct aid, were originally far more self-fertile, and that the increase of fertility gained by intercrossing - now only elevates it to the degree it ought to have, and probably had before insect visitation disturbed the sexual i : equilibrium. Cn the other hand, the absolute fertility of self-fertilizing weeds is undeniably : SELF-FERTILIZATION OF PLANTS. 347 for there are some flowers which one might, on à priori grounds, assume to be self- fertilizing, but are, nevertheless, proterandrous ; such, for example, is Lycopus europeus, with its stamens reduced to two. Hydrocotyle vulgaris, as far as concerned the spe- cimens I examined, was decidedly proterandrous ; nevertheless the flowers were com- pletely concealed beneath dense foliage of heath, grasses, &c., and not at all likely to be seen by insects; yet every fruit was “set”’*. Again, absence of scent is not an invariable rule ; for Salvia clandestina, though adapted for self-fertilization, is still stron gly perfumed, These cases compel caution in anticipating results which may prove not to occur. 7. Cleistogamous flowers + are always self-fertile. Mr. Darwin experimented upon two plants, Vandellia nummularifolia and Ononis minutissima. The following is a brief summary of his results. With regard to the first, the ratio of the height (100) of intercrossed conspicuous flowers and of self-fertilized conspicuous, and of the same with intercrossed and self-fertilized cleistogamous flowers, are as follows:—100:99 and 100: 94 respectively, a second comparison of the ratios of the heights of the former pair being as 100:97. "These results are practically ratios of unity. The crossed plants were, however, inferior in fertility to the self-fertilized cleistogamous flowers. Six of the finest plants of each kind yielded respectively 598 and 752 capsules, all being from cleistogamous flowers, as the crossed plants did not produce conspicuous flowers that year. The number of seeds per capsule was as 100 : 106. As with Vandellia so with Ononis, the cleistogamous flowers were highly self-fertile. The crossed and self-fertilized conspicuous flowers yielded seed in the ratio of 100 : 65 ; but the ratio between the crossed and cleistogamous was 100: 111, and the seeds them- selves looked finer even than those from the erossed perfect flowers. Further details on. certain cleistogamous flowers will be found in my descriptions of British self-fertilizing plants (postea, pp. 351, 359, 374), 8. Special adaptations Sor securing self-fertilization, mainly of British plants. RANUNCULACER.—Ranunculus hydrocharis, Spenner. Mr. Hiern (Journ. of Bot. vol. ix. p. 45) has made an excellent analysis of the many forms of this species of Ranun- culus, and deduced geometrical representations of the twelve species given in Babington's ‘Manual of British Botany. But it appears to me that another desirable investigation would be to test the raison d'étre of these different forms ; and I would suggest the fol- lowing interpretation as being more or less probable. Mr. Hiern finds that the twelve species can be grouped under three heads, and that if each species be placed ina given plane with reference to two axes of coordinates, the abscissa being the same number of units of length as the normal number of stamens, and the ordinate being the number of veins on each petal, then the species heterophyllus, confusus, Baudotii, trichophyllus, and Drouetii lie in a straight line whose equation is z—45--11—0; the species peltatus, flori- * Perhaps minute flies may assist, as I observed such diligently sucking the honey from Hydrocotyle americana in Kew Gardens. Indeed, Ido not deny that any flower, however minute, may not be visited, as Chickweed is sometimes, by bees (Meehan); but my object is toshow that it is quite immaterial, even if such be capable of self- fertilization. ` * For lists see Darwin's * Forms of Flowers,’ p. 312, and Bot. Zeit. xxv. p. 67. See also some excellent remarks by Prof. Oliver in the * Natural History Review,’ 1862, p. 238 seqq. ^ $48 -— | REV. GEORGE HENSLOW ON THE bundus, and tripartitus in a line whose equation is z—4y 4-6—0 ; and, lastly, the species puse circinatus, cenosus, and hederaceus lie in a line whose equation is y—y=4*. - The question, however, arises, What is the significance of these facts ? First, we may observe that the number of veins varies with the size of the petals and the amount of yellow colour, or “ guide,” at their base, as well as with the number of stamens ; therefore, generally with the conspicuousness of the flower. And as glands are well developed in the petals of the larger flowers, we may safely assume that such are adapted to attract insects, and so secure cross-fertilization. On the other hand, as the veins of the petals, as well as their yellow bases and the number of stamens, decrease, so the flowers become proportionately less conspicuous, and are more probably self-ferti- lizing, as they certainly are according to my observations in R. hederaceus. Hence I would infer that R. heterophyllus (25, 9)f, R. peltatus (30, 9), R. fluitans- (18, 14), stand at the head of the three groups, and are cross-fertilized, all having large flowers, On the other hand, the R. Drouetii (9, 5), R. tripartitus (6, 8), R. hederaceus — (7, 8) stand respectively as the last of each group, and which have very small flowers and are probably self-fertilizing. "The intermediate forms of each of these three groups will represent gradations possibly in each individual ease, being more or less liable to be intererossed, but yet have acquired a corresponding less or greater facility in self- fertilization. j Two species of this order are mentioned by Mr. Darwin as producing plenty of seed under a net, viz. Adonis e@stivalis and Ranunculus acris. R. bulbosus, R. acris, and R. repens, according to Müller, agree with R. flammula, which he describes in detail, with figures (Befrucht. &c. p. 114). The facts recorded exactly correspond with my own. observations in summer, that the anthers dehisce centripetally ; the outer stamens spread out towards the petals, the carpels being at that time quite immature, but are fully ` | developed before the last series of stamens have shed their pollen. B. muricatus has the : stamens reduced to about ten, and the carpels fully matured, while the first two or three 1 anthers only were dehiscing. I found no honey in the glands of the petals. The flowers are small and doubtless self-fertilizing. Certain differences obtain in the position of the line of dehiscence, so that while R. bulbosus dehisces laterally, R. >». repens and R. flammula are more inclined to be extrorse. “As a general rule, it would seem that extrorse anthers are specially concerned in intercrossing, but may by no means prevent self-fertilization; _ for the petals receiving the pollen from such anthers close at night, and so throw it on ` to the carpels. R. sceleratus and R. hederaceus, which have a reduced number of stamens — —a common feature in self-fertilizing { flowers—are certainly homogamous. The petals 4 are frequently, but not always, without honey; while the filaments bend down upon the ` 1 carpels, so that the anthers are close pressed upon the stigmas. This is the usual position - of the stamens in self-fertilizing polyandrous or regular flowers f. It is particularly well — ` * R. Lobbii, a new species added by Hiern from N. America, and intermediate between these last two forms and ` ` the aquatilis group, has the same equation; for he records the number of veins to be 3, and the number of stamens from 5-to 9. If, therefore, 7, the equation is e—y=4. -t These numbers are that of the stamens (25), and that of the veins in the petals (9). t I noticed in a branch of Crateegus owyacantha that every flower with incurved stamens had set its fruit, whereas every flower with the stamens ae’ had failed to do so. This observation needs corrobora tion for any general, ` ` zation, SELF-FERTILIZATION OF PLANTS: 849 seen in Potentilla Fragariastrum, and often in Stellaria media. After dehiscing, the anthers spread away. The order of development of the whorls of R. sceleratus is—calyx, pistil, stamens, or corolla. In the preceding species the majority of the stamens mature, as stated, before the pistil. R. tuberosus, from Kerguelen’s Island, has only a few stamens, is small-flowered, and doubtless self-fertilizing also. . The larger-flowered Ranunculi are probably all proterandrous as far as the outer series of stamens are concerned ; but as they mature centripetally, the inner may mature simultaneously with the carpels, if the outer have then shed their pollen. Myosurus minimus has also a reduced number of stamens and is self-fertilizing. Müller has described this in detail in * Nature,’ xi. p. 129, from which the following is abridged and here inserted, as this plant affords an illustration of an unusual method of self- fertilization :— Myosurus is as remarkable for the great variability in the size of its flowers, and in the number of its parts, as for the enormous growth of the cone of carpels, which affords no other benefit to the plant than the self-fertilization of the greater part of the numerous carpels by the small number of anthers [about 5], in case it be not aided by insects. The scentless and very inconspicuous flowers are scantily visited by small Diptera, not 90 p. c. of the flowers being thus favoured. "The axis of the flower, extending gradually, during the blooming-time, into a long cone, brings a great part of the stigmas into contact with the lateral pollen-grains of the anthers. "Those ovaries which now are in contact with the anthers are soon afterwards elevated above them, while others, previously below the anthers, now reach them, Thus a numerous succession of stigmas, by grazing the anthers during the growth of the long cone, are self-fertilized with about five or more pollen-grains apiece; besides, also, the lowest stigmas of the flowers are fertilized by many pollen-grains falling down from the anthers. Consequently only those carpels are never fertilized which are already situated above the anthers before the opening of the flower, Müller contrasts this growth of the gynzcium with the elevation of the stamens in Myosotis versicolor by the gradual growth of the corolla, which will be found described below (p. 375). | NyurmzACEX.—Some species are quite self-sterile, but others quite self-fertile. In Nymphea alba the innermost stamens are closely adpressed upon the stigma; but I do not know whether this species is self-fertilizing or not. According to Delpino, it is fertilized by bees *. ; .. Paraveracex.—Species of Poppy, Glaucium luteum and others, are by no means self- sterile; but Corydalis is remarkable for having some species (e. g. C. cava and C. solida) self-sterile, others (e. g. C. intermedia) self-fertile, but evidently adapted for insect- - fertilization by the remarkable structure of the corolla. From the latter one infers that the small-flowering Fumitory is a degradation or reversion to self-fertilization from an intererossing condition. The stamens and pistil of Fumaria are completely inclosed Within the petals. 'The stigmas are two in number, ‘and project laterally like blunt horns, and are enveloped by the three anthers of each group respectively, which thus form, as it were, a three-sided box. The pollen-tubes may be seen entering the stigma | in great profusion, the anthers being still in situ. (Tab. XLIV. figs. 3 a, b, c.) The . A curious observation was made by one of my pupils, that the stamens can be made to spread out if sir be — foiosd tap the peduncle, by Mowing at the cnt end, "on decre _ SECOND SERIES.—BOTANY, VOL. I. : 299 850 REV. GEORGE HENSLOW ON THE development of the whorls corresponds with this self-fertility. At first the concave’ 1 bract protects the flower-bud. The two sepals, which are a£ first.clearly anterior and q posterior in position, cover the rest of the flower. The two concave lateral petals next. grow, then the two smaller, anterior and posterior. Next in order comes the pistil, and: lastly the stamens are matured. ; ` F. capreolata, var. pallidiflora. Müller believes this species to be restricted to self- fertilize tion, and remarks :—* It has lost, probably from permanent disuse, the elasticity of the cap formed by the inner petals, which in other Fumitories secures cross-fertiliza- tion in case of the repeated visits of insects” (* Nature,’ xi. p. 461). CRUCIFERÆ.—The relatively larger and more conspicuously flowering species are obviously adapted for insect agency. The small orifices above or in front of the shorter ` stamens are correlated with the proboscis of an insect. The longer stamens, however, have their anthers clustering round the stigma, and probably, if the flower be not crossed, can fertilize the pistil in the majority of instances, as described by Miller in the case of Cardamine pratensis (Befrucht. &c. p. 134). The notched stigma is adapted to ` catch the pollen from the proboscis as it glides over it. The order of development of the whorls of larger-flowered Crucifere is—calyx, stamens, pistil, corolla. The stamens enlarge : and mature while the pistil remains more or less rudimentary. On the other hand, with ` small-flowered and often white species, such as Capsella Bursa-Pastoris, Nasturtium offi- cinale, Sisymbrium Alliaria, Lepidium campestre, e, the stamens and pistil appear to. emerge and begin to develope together; but the latter soon grows more rapidly, and ultimately matures its globular stigma contemporaneously with the anthers of the tallest ` — stamens, and is thus self-fertilized (see Tab. XLIV. figs.1 and 2). I have found speci- ` 3 mens of Shepherd's Purse in winter with the pollen shed in half-opened buds, and many | pollen-tubes penetrating the green stigma, while the grains were still in the anther-cells. In Lepidiwm campestre Y have found the stigma quite glutinous in half-opened buds, while the taller stamens were rather shorter than the stigma; they ultimately reach the latter, pollinate it, and then the pistil rapidly elongates. Such is generally the case with all the small-flowered self-fertilizing members of the — . Crucifere; while the order of development is usually, as stated :—first the calyx, then ` 4 all the stamens, together with the pistil, grow simultaneously ; and, lastly, the corolla; ` 4 or else the pistil at once takes the lead before the stamens. Possibly these latter slight ` ` differences may not prove constant, but vary with accidental circumstances. The great E difference, however, between the rapid growth of the pistil of self-fertilizing species ie its long delay in the intercrossing forms is very apparent. | The globular stigma appears to be correlated with, for it is at least characteristic of, — the self-fertilized species of this Order, just as the clavate stigma of Epilobium is found id the self-fertilizing species, but a branched stigma in those which are intercrossed, as de- —— scribed at p. 364. That the four latter stamens are specially concerned in self-fertilization ` ` is borne out by the fact of their being reduced to two only in Senebiera didyma. In 3 this species the stigma is globular and highly self-fertilizing ; there are no glands. Pringlea antiscorbutica of Kerguelen and Marion E is supposed by some to be - wind-fertilized; but as the stigma is not feathery, but globular, I feel no hesitation in ` SELF-FERTILIZATION OF PLANTS. . B51 saying that it is most probably self-fertilizing.. This is quite in keeping with the profusion of siliquas it bears. | An exception to the above rule appears to be afforded by Cakile maritima. I found this growing at Felixstowe, and it was certainly proterandrous; for the stigma (which, however, is capitate) was mature, while the anthers had quite shrivelled. It matured only about 20 p. c. of seeds. The order of development was—calyx, all the stamens together, pistil, corolla. Hence it seemed to represent an intermediate condition. VionACEA.— Herman Müller has well described the self-fertilizing form of Viola tri- color; and I would regard it as an instance of a “ species" having been derived from an intercrossing form by adaptation to self-fertilization. It may not yet differ sufficiently from the typical form to be worthy of the title of ** species; " yet it would seem to be at least in the first stage towards the formation of one. Viola tricolor is constructed for fertilization by insects, and, as Müller has shown, very self-sterile if they do not visit it. It, however, may accidentally become self-fertile by the corolla withering, and so pressing the collected pollen-grains into the stigmatic chamber, as represented in Tab. XLIV. fig. 6 d. I have found, moreover, specimens in which the so-called “lip” was much elongated, and the stigmatic tissue which lines the * throat" extending outwards to the tip, which was highly glutinous. The consequence was that this ** tongue," as I should prefer to call it, had licked up the pollen, the tubes from which were abundantly penetrating the throat, and the ovary much enlarged with fertilized ovules. (Tab. XLIV. fig. 6c.) i I have met with other small self-fertilizing individuals of Viola tricolor in which there was a curious development of conducting or stigmatic tissue, not recorded by Müller. The globular ** head ” of the pistil of Viola is usually quite hollow, but containing some fluid, the pollen-grains passing into the ' head; but in the cases I am alluding to a pillar-like structure issued from the top of the style, and, curving forwards, protruded slightly from the orifice of the * head," and the extremity was there somewhat enlarged. This was highly glutinous, and a large quantity of pollen-grains were adherent to it, with quite a considerable bundle of tubes passing down the “ pillar,” they having fallen directly upon it from the anther-cells below. (Tab. XLIV. figs. 6a and b.) The existence of cleistogamous and highly self-fertile flowers in the genus [Viola is well known. A brief description may be adv isable. The five sepals are normal, but very minute. The petals are rudimentary purplish-green scales or absent. The stamens are from 2 (V. canina)* to 5 (V. odorata) in number. The filaments are slender, but the orange “flaps” large and rounded, while the anther-cells are small, oval at the basal part. The anthers are all alike, with no appendages, and are closely adpressed upon the summit of the ovary. The style is short, curved, and bears a truncated stigmatic end, which lies concealed amongst the anther-cells. The pollen-tubes penetrate, while the grains are still in situ, within the cells. As the ovary swells, the filaments are detached, and the anthers, becoming raised, form a stellate mass on the summit. (See Tab. XLIV. figs. b, c, d.) In * double” Violets I find the cleistogamous flowers become double also t. $ Three stamens are sometimes present as petaloid staminodia. + This description was written some time ago; and Mr. Darwin, in his latest work, ‘ Forms of Flowers’ mm nee Lee corroborates all I have seen, but adds further details, which should be n a 352 REV. GEORGE HENSLOW ON THE Mr. Darwin has devoted a chapter, in his ‘Forms of Flowers, to a description of these — peculiar and cleistogamous flowers, to which the reader is referred for full details. I | would only remark that I am fully convinced that they are, in the first place, arrested flower-buds; but the arrest is accompanied with, on the one had, the suppression of special structures adapted for insect agency ` and, on the other, it is supplemented by special facilities for seeuring self-fertilization. In no case do I believe they represent primitive forms of flowers, as Mr. Dyer suggested (‘ Nature,’ vol. xv. p. 331), who says :— “The view by which flowers are regarded as originally hermaphrodite instead of, as Mr. Darwin sug- gests, monccious, further supplies a very simple explanation of the otherwise almost inexplicable.nature - of cleistogene flowers. These being inconspicuots, and self-fertilizing, are probably survivals of the ` original type." : That flowers were originally hermaphrodite, I fully believe, but certainly not neces- sarily cleistogamous, nor that existing cleistogamous flowers are survivals. They appear — to me to be undoubtedly degradations, for at least two, if not more, reasons. One is the ` presence of the corolla; the other that peculiarities of structure formerly concerned with ` insect fertilization are not always completely lost, as, for example, the lip of the corolla | in Lamium amplexicaule and its four stamens. Moreover, transitional conditions may often be found between ordinary flowers and the cleistogamous. Thus in strongly grown garden-plants of Viola odorata the cleistogamous buds of the summer often have — spurs, and the stamens retain the nectariferous appendages, where they can be of no ` possible use whatever. Transitional conditions also occur with other plants that bear - cleistogamous flowers, fully proving that they are really transformed states of the normal 3 blossoms. (See Tab. X LIV. figs. 4 a, b, c, d, and 13 a, b.) | PoryeALEX.— The remarkable structure of Polygala is clearly correlated to i fertilization, yet the British species appear to be adapted to self-fertilization as well. -Hildebrand has described it in * Bot. Zeit., Sept. 6, 1867, from whose figures I have made ` the drawings in Tab. XLIV. figs. 7 d & e. Fig. e represents a case where an insect has deposited the pollen on the anterior side of the stigma; fig. d one in which the spoon- — shaped extremity has caught the pollen poured into it from the anthers, which have grasped the edges of the spoon and, as it were, emptied themselves into it. The stigma has recurved upwards, to bring the viscous surface in contact with the pollen. Ina white variety of which I found a few specimens, the corolla was completely wrapped round the anthers and stigma, as seen in fig. 7a; and the anthers gripped the two sides of the pistil just on a level with the very.viscid stigma, into which many pollen-tubes were penetrating, so that the pollen adhered to it directly, without falling into the. spoon i at all (fig. 7). Hence, if my observations were correct (but I had too few to render them quite satisfactory), there was a slight difference between the method of self-fertilization in my ease to that described by Hildebrand. It may be observed here, however, that slight variations such as this are not unique. Mr. Darwin records variations in the Jength of the stamen in Canna (1. c. p. 230) ; and I have reasons for suspecting individual plants of species vary the processes under different circumstances; and a contrivance which may occur on one plant may not necessarily be found in another individual of ` SELF-FERTILIZATION OF PLANTS. : 353 the same species. Other cases seem to bear out this somewhat important fact, such as the case of the Viola tricolor with the protruding placentiferous process described above; as also that of the peculiar form of the same plant with the prolonged * tongue," enabling it to “lick” up the pollen from the spurred petal. Again, the arching of the styles outwards, and the filaments inwards, in the self- fertilizing condition of Alisma plantago, as also the subsequently inflexed condition of the stamens of Agrimonia, seem to point to what may be called individual adaptabilities for securing self-fertilization; and such structures as these may not precisely or ever occur again, while other peculiarities may be found adopted to secure the same end. — I would invite the attention of observers to this point, which appears important, but requires further corroboration. CARYOPHYLLACEJX.—The larger-flowered species of this Order are probably always pro- terandrous, and adapted for insect-fertilization ; yet, as Mr. Darwin has shown in the case of Dianthus Caryophyllus, which is very strongly proterandrous, there appears to be no difficulty in their becoming self-fertilizing by losing their dichogamy and so becoming homogamous; that is, the anthers and stigmas mature together. Mr. Darwin found, in his first experiment with Carnations, that there was little difference in the number of seeds produced by cross-fertilization and self-fertilization (artificially produced) in this plant. But in the second generation the crossed was more fertile than the self-fertilized in the ratio of 100: 65, both being grown much crowded. But when plants of the third generation were placed under a net, and both the crossed and self-fertilized were allowed to set seed spontaneously, the self-fertilized plants now produced more seeds than the self-fertilized ** crossed " plants, in the ratio of 125:100; and he adds :—“ This anomalous result is probably due to some of the self-fertilised plants having varied so as to mature their pollen and stigmas more nearly at the same time than is proper to the species,” . It must be borne in mind that proterandry in flowers admits of degrees; that is to say, there is no absolute length of time between the maturation of the pollen and that of the stigma. When the period is relatively long, as in Dianthus, some species or varieties of Pelargonium, Geranium pratense, and Malva sylvestris, every grain of pollen will have been shed, and sometimes the anthers fallen off as well, before the stigmas are nearly ready. On the other hand, Stellaria Holostea and Geranium pyrenaicum develope their two whorls of stamens in succession; and the stigmas maturing about the same time as the second whorl, these flowers are only partly purum upon insects. This condition of things accounts for many flowers retaining, or else regaining, their self- fertilizing properties, though still being adapted for intercrossing. ` The point, then, I wish to bring forward as partieularly well shown by members of this Order, is that complete self-fertility may be regained with great ease, namely, by shortening the time between the periods of maturity of the anthers and stigmas, The causes to bring this about are ënn various, though somewhat difficult to isolate and specify exactly, With Dianthus Mr. Daititin records that it was of a pale’ pink or rose-colour, which mek indicate the approximate cause, assisted by continual and artificial homogamy ; so ~ 354 REV. GEORGE HENSLOW ON THE that a self-fertilizing form of Dianthus was artificially produced analogous to the garden and sweet peas, which appear to have become so in this country on their own account. : ^ Several genera in this Order have both conspicuous and probably all proterandrous flowers, as well as inconspicuous and self-fertilizing species. Thus, for example, while the large-flowered Cerastium arvense and Stellaria Holostea are both proterandrous, the small-flowered C. glomeratum and Stellaria media Ze, are, self-fertilizing. With regard to C. glomeratum, it is perhaps worth mentioning that the flowers often remain very imperfectly open ; and on one occasion, in June 1876, I found a number of plants in an exposed meadow with all the flower-buds actually eleistogamous, although exposed to full sunlight and an intensely high temperature. Stellari ia media and Spergula arvensis, although they usually expand their blossoms in sunlight if it be warm, yet in January I have found them self-fertilizing abundantly but with the buds completely closed. In all these cases the filaments are bent down over the ovary, with their anthers in close proximity to the stigmas. (Tab. XLIV. figs. 8 a, b, & 9.) Stellaria media may show analogous features. Thus in a larch-wood I found specimens permanently cleistogamous and apetalous and with three stamens only; on the same plant were flowers with rudimentary petals and with four stamens. The petals being the last organs to be developed are often the first to go, the order of development being as follows :—First stage, calyx, pistil, stamens, corolla; secondly, the pistil is equal to the stamens in height; thirdly, the corolla, which has long delayed, starts again. The same order occurs in the self-fertilizing Sagina apetala and Polycarpon tetraphyllum. So also in Spergularia marina (subsp. neglecta), the order of development is—calyx, pistil, stamens opposite the sepals, stamens opposite the petals, corolla. The ten stamens close round the pistil, and the anthers dehisce while the bud is opening. ` ` I have already remarked that the reduction of the size of the corolla, with often a loss of colour, if not an immediate cause, is at least closely correlated with self-fertilization, and that as some of the stamens are often entirely arrested, so too may the corolla be arrested as well. This will account for some species of this Order becoming apetalous, as the genus Sagina, as also Herniaria and Scleranthus of the Paronychiez. Thence we may pass to the incomplete members of the group called Cyclospermere, and so on, as I believe, to all other members of the Monochlamydez: and Achlamydee of Dico- _tyledonous Angiosperms, that the arrest of the corolline whorl, indicated by the stamens being opposite to the sepals, has issued from a correlation with self-fertilization. Further remarks, however, will be added when I treat specially of these divisions. ; Of the Illecebrace:e, the few that I have examined are self-fertilizing, often in unopened buds, as. Paronychia Bonariensis, Corrigiola littoralis, Scleranthus annuus, and Herniaria | glabra; and I would venture to express it as my opinion that this Order is "simply a degraded form of Caryophyllaceze by becoming self-fertilizing. If I may offer as a suggestion of the process how it came about, I should imagine that seeds of certain . insect-visiting conspicuously flowering species became located where they were neglected by insects; and having never entirely lost the power of self-fertilization, as is the case with Stellaria Holostea &c., which is partially proterandous only, reacquired it, but with the ECH reduction of the size of the corolla, its colours and also of the eee SELF-FERTILIZATION OF PLANTS. 355 and so degraded into self-fertilizing forms which we now recognize as distinct species or even genera. Compare, e. g., the form Spergularia marina (proper), with large corolla and ten stamens, with the subsp. neglecta, having small petals, three or four stamens, and which is highly self-fertile. In support of this view, I may remark that while it often happens that the number of stamens and the quantity of pollen is much reduced in self-fertilizing flowers, or, on the contrary, the number of ovules may be reduced, compensated, however, by the pro- fusion of blossoms, yet the full number of stamens and a large quantity of pollen is not unfrequently retained, which therefore indicates the retention of a former condition, when a larger amount of pollen was required for insect agency. "Thus there is only one seed in Fumaria officinalis, but the quantity of pollen which becomes fixed to both stigmas by their pollen-tubes is very great. Similarly with Chenopodium, the five anthers are retained, but there is only one seed to each flower. Conversely, cleisto- gamous flowers of Viola canina and buds of Stellaria media produce many seeds ; but the anthers may be reduced to two. | The Caryophyllacez and its allied Orders are therefore very instructive as far as fer- tilization is concerned. Dianthus, which is very strongly proterandrous, may be placed at one end of the scale and Sagina apetala, or some incomplete member, at the other ; and between these extremes there are degrees of differentiation from the utter impos- sibility of securing self-fertilization and complete self-fertility. Yet since such an extreme form as Dianthus can become highly self-fertile in two or three generations, as Mr. Darwin has shown, we may be assured that all others could readily become self- fertilizing under fitting conditions. Plants of this Order also well illustrate the extreme rapidity of maturation of seed resulting from self-fertilization. If, for example, we call the terminal flower of a cyme of Cerastium triviale as No. 1, then if this be shedding ripe seed, the two capsules (No. 2) will be nearly ripe, while the four (No. 3) will be self-fertilizing buds; and lastly, the eight buds (No. 4) will be in a very rudimentary stage, with the corolla quite unde- veloped. The same rapidity may be noticed in the case of Poa annua, which is mostly, I suspect, self-fertilizing, and by the extraordinary rapidity of its seeding and growing so rapidly clothes bare places in our London parks, to the general surface of which, indeed, it contributes in no slight degree. This extreme rapidity of seeding com- pensates, where needed, for any comparative paucity in the number of seeds produced in a single capsule, though, as a rule, there is not even any particular deficiency in that respect at all. Other ‘points of importance are well illustrated by this Order, namely, the actually beneficial effects of size and duration of existence in these self-fertilizing plants. As a rule, they are small and annuals. Thus while Cerastium arvense and Stellaria - Holostea are perennial, the self-fertilizing Cerastia and Stellaria media are annuals. Similarly in other Orders, Geranium sanguineum, pratense, &c. are large-flowering and perennials, but the small-flowered species are annuals and more or less decidedly self- fertilizing. It is by no means invariably the case; e. g., Malva PONAM and mec Tn of ME are EENG and pique. 356 REV. GEORGE HENSLOW ON THE In estimating the importance of relative size, whether of the entire plant, its: foliage, &c., or of the flowers only, we must be mindful not to be misled by our own impressions of what we may deem a superiority. We must start with some end in view, it is true; and I agree with Mr. Darwin in regarding the propagation by seed as the one and only end of plant-life that we may legitimately recognize. This is clearly the case with an annual; its whole life is directed to this end, and as soon as the end is secured, 7. e, | the seed matured, the plant perishes. Now, as Mr. Darwin observes, self-fertilization secures this end with ** incomparably greater certainty " than by any other means. Hence self-fertilization is so far a positive * good." Now that observer has proved that much “ finer ” plants are reared from inter- crossing; hence we may rest assured that what is artificially produced would occur also in nature; but as a result of intercrossing, dichogamy and other sexual differentiations have been established, and the security of setting seed is not enhanced but lessened by such. adaptations to insect-agency. Indeed the more highly a flower becomes dif- ferentiated the less is the chance of securing seed on the failure of certain insects to yisit it. We must, too, carefully remember that smallness of size is no detriment to a plant in the struggle for life, but the reverse. This is proved not only by the immense extent of low plant-life, such as unicellular Algee, but by the wide dispersion of ** weed-like " herbs, as will be more fully diseussed hereafter, and by the fact that small herbs of one country may drive out their more formidable opponents in another, as, e. g., thelittle Duteh Clover is vigorously competing with its gigantic foe Phormium tenax in New Zealand. The cycle of its existence is often run through in a few weeks, so that crop after crop, descended from one and the same individual, ean be produced in a single year. Hence the justly framed expression of ** troublesome weeds." Therefore I maintain that, for the “ good " of the plant itself, there is no special advantage in its being a perennial, or in being a large or, so-called, “ finer " plant, nor in having conspicuous flowers requiring insects for their fertilization. But, on the other hand, small self-fertilizing species can establish them- selves almost anywhere. They produce, probably, far more offspring, first, on account of their being capable. of self-fertilization, and therefore not dependent upon insects, secondly, by rapidly running through their life-cycle, producing crops in succession in the same season. So that “ weeds," such as Stellaria media, prove themselves to be best fitted to survive in the struggle for life; and any idea of “ injuriousness”’ arising from their self-fertility is purely a subjective impression, which is not correlated with any Mice fact. | MaLvACEX.—Malva sylvestris, is very strongly proterandrous. Every grain of pollet: appears to have escaped and the anther-cells more or less shrivelled before the stigmas have protruded and are mature for xs the pollen, in this case necessarily, from other flowers. M. rotundifolia, which is a stich smaller plant altogether, and with pale pink incon- spicuous flowers, is self-fertilizing, as has been described by H. Müller (Befrucht. Ze, p.171) Other small-flowered species, as M. crispa &c., are also self-fertilizing in a similar manner, and, cupere furnish an exception to therale first mentioned, that. self- SELF-FERTILIZATION OF PLANTS. 357 fertilizing plants are usually small, in that this species is a tall finely foliaged plant. The development of the flower is as follows :—The bracts of the epicalyx are at first free and much larger than the calyx, which is densely hairy. The stamens arise as a circular rim at first, then follows the pistil, while the corolla delays its development; the pistil then begins to grow rapidly, while the corolla at first remains small, but sub- sequently covers the elongated styles, which thus are compelled to grow downwards amongst the anther-cells. The curling amongst the latter is therefore simply due to the styles elongating under the constriction of the corolla, which keeps them down, and they subsequently retain that position when the corolla has expanded. This case, therefore, again illustrates the general principle, noticed under Cruciferz, that in self-fertilizing flowers the pistil grows rapidly in the bud, whereas in conspicuous inter- crossing flowers it is delayed, so that the flower becomes proterandrous in consequence. Althea Armeniaca is a comparatively small-flowering species (1 inch diam.), and furnishes an intermediate condition, in that the sti gmas begin to protrude from amongst the anthers before they dehisce, whereas in M. sylvestris they are quite invisible. They do not, however, curl back, and consequently are not so well adapted for self-fertilization as in M. rotundifolia. A genus analogous in the above features to Mallow is Anoda; for A, hastata is de- cidedly proterandrous, the anthers dehiscing centrifugally, while the branches of the style, terminating with globular green stigmas, are concealed within the tube. After the anthers have all dehisced the branches become erect and the stigmas red; the flower is pink, and 14 inch in diameter. 4. Wrightii is much smaller, salmon-coloured, and developes its pale yellow stigmas simultaneously with the anthers, which cover them with pollen. The branches are notrecurved, as the stigmas only rise to the level of the anthers. . Linacem.—Like Caryophyllacez, this Order furnishes us with extreme instances. Linum grandiflorum, as Mr. Darwin has shown, is not merely morphologically but physiologically self-sterile (‘Forms of Flowers, p. 264)*. On the other hand, Linum catharticum, though secreting honey, and frequently visited by insects, is capable of self- fertilization, not only in the absence of insects, but an insect visiting the flower is very likely to press the anthers against the stigma (Tab. XLIV. fig. 10). This plant can also fertilize itself by the corolla closing at night; and Radiola millegrana is also, as might be almost inferred from the profusion of its fruit, quite self-fertilizing. Moreover the flower-buds remain nearly closed, retaining the stamens pressed down upon the ovary. The anthers contain but a small quantity of pollen. GERANIACEJ.—Like the preceding, this Order furnishes us with transitional species between strong dichogamy and entire self-fertilization. Sir John Lubbock has so well described this, and it quite confirms my own observations, that I will transcribe a short passage from his * British Wild Flowers in Relation to Insects, pp. 42—44. * The genus Geranium affords us an instructive example. There are a number of species, which differ much in the size of the flowers. Thus those of G. pratense are nearly twice as large as those of * Linum perenne is also dimorphic (* Forms of Flowers,’ p. 92), but, like Dianthus, is quite capable of acquiring the powers of self-fertilization (Meehan). oe : | : SECOND SERIES. -—BOTANY, Y Jh ii — pn x. 3D 358 REV. GEORGE HENSLOW ON THE G, pyrenaicum; which, again, are much larger than those of G. malle, while those of G. pusillum are still'smaller. These differences of size appear to be connected with other remarkable differences between the species. In G. pratense five of the stamens raise themselves and stand upright, and surround the still immature pistil. ` When they have shed their pollen they sink back and shrivel up, when the other five raise themselves. At a later stage these in their turn fall back and shrivel up; but the stigma does not become mature until all the stamens have shed their pollen. Under these circumstances G. pratense has lost the power of self-fertilisation, and is absolutely dependent on the visits of insects. G. pyrenaicum is also proterandrous; but while in G. pratense the pistil is not mature until the stamens have shed all their pollen and fallen back, in G. pyrenaicum the second series of stamens are still upright when ` the stigmatic lobes unfurl; the flower is consequently less absolutely dependent on insects, and we see ` that the corolla is much smaller. In the third species, G. molle, the pistil matures before the second series of stamens, and the corolla is still smaller; while in G. pusillum the pistil matures before any of the stamens.” Thus, then, these four species may be arranged in atable (slightly modified) as below :—- G. pratense. G. pyrenaicum, ; G. molle. G. pusillum. Flower large. Flower small. ; Flower smaller. S Flower smallest, First exclusively male, | First exclusively male, | First exclusively male, | First exclusively female, then exclusively female. then homogamous*. then homogamous. very soon homogamous. Incapable of self-fertiliza- | Generally fertilized by Often self-fertilized. Probably entirely self- | tion. | inseets. fertilized. I have quoted this passage, for it both entirely corroborates and so well expresses what I have myself observed, both in regard to Geranium as of many other genera ; and | it illustrates what I believe to be the origin of a large number of * species’ of plants, namely, the change from a state requiring intercrossing to self-fertilizing conditions. 1t would be rash to assert that the above four species represent actual linear descendants, but they doubtless typify stages of descent from forms like @. pratense to G.. pusillum. The description of G. pusillum as being * at first exclusively female,” the reader will see, is exactly paralleled by several cases already mentioned, as of the Cruciferze, and will be again illustrated by following ones, and e ae P the origin of Proterogynous flowers generally. G. Robertianum. In this very common species I have found specimens growing in à wood, and not likely to be visited by insects, with the pollen-grains caught by the — stigmas in the unopened buds, and self-fertilization apparently being secured. In some flowers the anthers of the lower stamens were spread out and the stigmas likewise; the ` latter also were curling backwards oe the anthers, : much in the same way as ‘occurs : in small-flowered Mallows. Erodium cicutarium. Though sometimes proterandrons, this can completely ‘fertilize | itself. The loss of five stamens in this eed is probably correlated with this rever- sion to complete self-fertility. Pelargonium. Ihave given figures opa XLIV. figs. 11 a, b) to show the strongly. ZS Sir J. Lubbock used the wad “ hermaphrodite,” for which I have substituted “ homogamons.” The former “merely means that stamens and min are in the same flower; the latter that, in E to this, ee V ean be dioi, | ; SELF-FERTILIZATION OF PLANTS. 359 proterandrous condition of some species, as well as the self-fertilizing condition of many varieties of the so-called “ Scarlet Geranium " (fig. 12). | Oxalis Acetosella, Mr. Darwin alludes to Michalet’s description of the cleistogamous flowers of this species (given in Bull. Soc. Bot. de Fr., tom. vii. 1860, p. 465), and adds some observations of his own (‘ Forms of Flowers,’ p.321). He quotes an observation of Michalet’s, that the five shorter stamens are sometimes quite aborted, a condition I have not met with myself, but one quite in keeping with the common process of reduction of stamens in self-fertilizing flowers. He also adds this interesting observa- tion, * In one case the tubes, which ended in excessively fine points, were seen by me stretching upwards from the lower anthers towards the stigmas, which they had not as yet reached. My plants grew in pots, and long after the perfect flowers had withered they produced not only cleistogamic but a few minute open flowers, which were in an intermediate condition between the two kinds." This last remark is clearly in . accordance with the true origin of these flowers, that they are in all cases degradations from the conspicuous forms normally characteristic of the genera which produce them. The figs. 134, b, c, d, Tab. XLIV., clearly show that in Qzalis Acetosella the cleistogamous state is simply a normal flower-bud, which has become adapted to self-fertilization; and the intermediate conditions alluded to by Mr. Darwin I should suspect were analogous to the permanent forms of flowers of O. corniculata, which I at first inferred, from its wide distribution, must be habitually self-fertilizing *, and subsequently I discovered the pollen-tubes' penetrating the stigma of each unopened bud (fig. 14). M. E. Michalet's description (/. c. p. 467) is much to the point, as the subjoined free translation testifies. The cleistogamous flowers are about the size of the head of a pin, often subterranean, on short curved peduncles, with included petals. They arise from the same points of the rhizome, but not simul- ` taneously, and continue through summer and even to autumn. The sepals are closely applied during fertilization, and conceal hermetically the essential organs. They increase to double their size; but the capsule soon takes on a relatively enormous size, considering the minuteness of the ovary. The petals, 5, are shorter than the sepals ; they rarely are wanting. The andrecium is composed of 10 stamens; 5 are large, inserted upon a narrow disk which surrounds the base of the ovary. The anthers of the smaller stamens appear infertile, or even entirely abortive. The 5 fertile stamens are inclined towards the stigmas, and are bound to them by fine threads; they certainly play some part in the fecundation ; but the nature of their functions is obscure. [Consult Tab. XLIV. fig. 13d.] As in Violets, I have vainly looked to find the emission of the pollen from the anthers upon the stigmas. The pollen appears to be slightly deliquescent ; the cells which contain it have seemed to me to remain closed and intact after fecundation has already taken place, and is manifested by the growth of the capsules. In a very young ovary the 5 carpellary leaves are nearly free, as in Sedum ; later the carpels unite by their lateral walls and form a 5-celled ovary. The stigmas (which in the spring flowers are on long styles) are sessile on the ovary. The capsule is shorter and more rounded at the summit. The seeds do not appear to * While examining O. Acetosella I found the petioles of the leaves articulated near to, but not absolutely at the base. ‘The small thickened portion which remains has its cellular tissue highly charged with very large starch- grains. Dr. Masters tells me that this also occurs at the bases of other deciduous leaf-stalks, apparently as nutri- ment for the formation of the cells which give rise to the transverse division which causes the leaf to fall; but in the: case of the Wood-Sorrel the leaf Aas fallen, so that they must be regarded as little reservoirs of nutriment for some ` * 360 |. REV. GEORGE HENSLOW ON THE differ from those of the ordinary flowers; but as they ripen under the moss or vegetable detritus which covers the rhizome, they cannot be projected around the mother plant, but remain in their place, My own observations do not agree entirely with the last sentence; for I have often found the fruit-capsules elevated and the seeds with quite as elastic a coat as those derived from the fully developed flowers. Impatiens fulva and I. Noli-me-tangere have also eleistogamous flowers. For full de- seriptions the reader is referred to Mr. Darwin's work, ‘Forms of Flowers,’ p. 327, where ` reference to other describers is given. —Zmpatiens parviflora is undoubtedly self-fertilizing. The stigma lies amongst the anthers, which burst even before the flower is opened, and I have detected pollen-tubes penetrating in that condition. 3 Lreuminos&.—That this order has its flowers generally adapted for insect-agency * all E will'admit. Mr. Farrer's elaborate description of Pisum, quoted by Mr. Darwin, ‘Cross — and Self-fertilisation, p. 160, will furnish the reader with details. Nevertheless there — are a great many very inconspicuously flowering species, as well as at least twelve "E a cleistogamous forms (‘ Forms of Flowers,’ p. 313). | 4 Of the different species of Trifolium, Mr. Darwin thus writes :— 7. incarnatum. “The a | flowers whieh were visited by bees produced between five and six times as many seeds E as those whieh were protected." E T. pratense. ‘ One hundred flower-heads on plants protected by a net did not produce a single seed, whilst one hundred heads on plants growing outside, which were visited by bees, yielded 68 grains weight of seeds; and as 80 seeds weighed 2 grains, the 100 heads must have yielded 2720 seeds.” (Cross and Self-Fert. p. 361.) . T. repens. The crossed and self-fertilized plants of Mr. Darwin yielded seeds in one © year in the ratio of 10:1, and in the second year twenty heads, unprotected, yielded 2290 — seeds, while twenty protected beads had “ only a single aborted seed.” | With regard to Clover Mr. Meehan observes (Gard. Chron. Sept. 11, 1875) :—* I am satisfied that in all eases I examined flowers just before expanding, and before any insect had interfered with them, the pistil had received its own pollen .... I covered a patch ` of clover with a sieve, having $ inch meshes. No Bees could get to gem: I think I may — say every flower perfected seed. Unfortunately, I found on one examination a small | Sand-wasp had ventured through, and was collecting pollen from a flower, Ido not — think any but this one entered; still it diminished seriously the value of the ex- — periment.” Mr. Meehan gives some further statements upon the Red and White Clovers which, if trustworthy, are opposed to Mr. Darwin; for while the latter found, on protected - Trifolium pratense, no seed was set, the former observed that Humble-bees, which alone ` apparently fertilize the red clover, would not visit a field when the white clover, 7. re- pens, was in blossom; and yet the red clover-fields “bore seed as fully as most insect- frequented fields would do." Assuming both of these accurate observers to be correct, the inference is unavoidable that the red clover, although highly differentiated, 80 as to. 3 _* I have described the peeuliar construction of Medicago sativa (Journ. Linn. Soc., Bot. ix. pp. 327 & 355) onl of 2 Genista tinctoria (Journ. Linn. Soc., Bot. x. p. 468) ; reproduced by Müller in his * Befrucht. Ze? pp. 225 and 235. The methods of fertilization have been grouped by H. Müller into four series, fee. Lubbock's. ‘British wild : Flowers in Relation to — p 90. . ; vitet SELF-FERTILIZATION OF PLANTS. 361 become morphologically self-sterile here, had, where Mr. Meehan observed it, slightly changed, and so recovered its self-fertilizing powers, as was the case with -Eschscholízia. T. arvense and T. procumbens, and, I would add, 7. minus, appear to be highly self-fertile, according to Mr. Darwin, though he suspeets the latter may be visited by small nocturnal moths. With regard to 7. minus, the order of development, viz., calyx, pistil, stamens, corolla, agrees with self-fertilizing plants, Medicago sativa. This plant, when protected, yielded seeds, as compared with unpro- tected, in the ratio of 101:.77. Hence it is highly self-fertile, though specially modified, in having * irritable" stamens, for cross-fertilization (note, p. 360). M. denticulata. This species appears to be quite self-fertile. The position of the anthers clustering round the stigma is seen in Tab. XLIV. fig. 15. Phaseolus vulgaris. While this species is quite self-fertile, P. multiflorus, when ** pro- tected from insects, produced on two occasions about one third and one eighth of the full number of seeds... .. The flowers are not visited by insects in Nicaragua; and, according to Mr. Belt, the species is there quite sterile." Cross and Self-Fert. p. 200 (Zhe Naturalist in Nicaragua, p. 10. See also a paper by myself in Gard. Chron. Nov. 3, 1878.) Lathyrus grandiflorus is in'this country more or less sterile. It never sets pods unless the flowers are visited by humble-bees, or artificially fertilized. L. odoratus and LL, Nissolia are fully self-fertile. ^ Pisum sativum is also fully self-fertile. Vicia faba. Unprotected plants were between three and four times more fertile than the protected plant. Vicia sativa and V. hirsuta are both perfectly self-fertile. The above cases, mainly taken from Mr. Darwin’s lists of sterile and self-fertile plants, seem to bring out the following facts :—(1) Conspicuous papilionaceous flowers may be absolutely self-sterile, or only partially so, or not at all Whether any species are physiologically self-sterile I am not sure; but many are certainly more or less morpho- logically, as Lupinus, sp., Phaseolus multiflorus, &c. (see * Cross and Self-fertilisation,' pp. 150, note, and 152). (2) One species of a genus may be highly self-fertile, while another species may be barren (Phaseolus and Trifolium). Even with regard to the self- sterile P. multiflorus, Mr. Darwin found that “the advantage gained by a cross is very small" or, practically, none, as the heights were as 100 : 96, and there was little or no difference in their fertility. The Sweet Pea, though crossed in the south of Europe, is entirely self-fertile in this country, though L, grandiflorus is self-sterile ; on the contrary, L. odoratus and, we may add, P. sativum, appear rather to agree with Phaseolus vulgaris in being plants which have become] highly self-fertile by being transformed from warmer climates to this country. If so, they exactly parallel the case of Eschscholtzia, which was actually proved to be so. From these intermediate stages we pass to species which are as fertile when unvisited as when fertilized by bees and other insects. Lastly, there are extremely small-flowered species, but with perfect blossoms, which are probably never visited, as Vicia hirsuta ; and also several with cleistogamous flowers, which are, as is always the case, highly self-fertile. - Mr. Darwin’s descriptions of several cleistogamous flowers (‘Forms of Flowers,’ P. 310 seqq.) are particularly interesting, as they so manifestly prove these closed 362 REV. GEORGE HENSLOW ON THE blossoms to be degraded, but differentiated, forms of conspicuous allied ones. Thus of Lathyrus Nissolia (p. 826) he remarks that this species “apparently offers a case of the first stage in the production of cleistogamous flowers, for on plants growing in a state of nature, many of the flowers never expand, and yet produce fine pods. Some of the buds are so large that they seem on the point of expansion ; Gen are much smaller, but none so small as the true cleistogamic flowers of other species.’ Rosace&.—‘“ Our three species of Prunus,” writes Sir John Lubbock (7. c. p. 90), «differ somewhat in the relations of the anthers to the stigma. In P. Cerasus (the Cherry) both: mature at the same time, while in P. spinosa (the Black Thorn) and P. Padus (the Bird Cherry) the stigma reaches maturity before the anthers; though, as it retains the capa- bility of fertilisation after the anthers have opened, the flowers are doubtless often self- ` fertilised, which, from the position of the anthers, probably happens more frequently 1 Im ` the Bird Cherry than in the Black Thorn.” This case of Prunus is an interesting one, for it is opposed to.the rule that protero- ` gynous flowers are inconspicuous and usually unattractive to insects. The cause of the ` species of this genus maturing the pistil early is probably in consequence of their flower- . ing early in spring; the temperature not being high, there is no special tendency to stimulate the staminal and coronal whorls, or possibly the glands. It has been elsewhere shown that flowers, usually perhaps intercrossed, will. become self-fertilizing in cold weather; so that what takes place abnormally with proterandrous flowers becomes normal in the case of Prunus. The same remarks apply to the Apple and Hawthorn, which are proterogynous. ! Spireas, if not visited by Bees for pollen —dis they contain no honey—are, without | little doubt, self-fertilizing. Geum. Though both British species are mellifluous, yet the smaller, G. urbanum, a probably mostly self-fertile; for the position of the stamens, arching over the carpels ` and dehiscing upon them, is quite in keeping with the small-flowered Ranuneuli, as also ` with Potentilla Fragariastrum. As with the Ranunculi, the stamens mature centri- ` petally, thus affording a considerable time to elapse, during which the carpels may suc cessively mature their stigmas; so that while in the more conspicuously flowering ` species, as Geum rivale aud the Blackberry, intercrossing is chiefly effected, it does not preclude the possibility of self-fertilization, which probably snpittodpa it Kess im the small-flowering more or less inconspicuous flowers. T Agrimonia Eupatoria.. In this flower the stamens are spreading on first expansion, but it is probably self-fertilizing afterwards, as the anthers become strongly recurved over ` the pistil. (See Tab. XLIV. figs. 16 a; 5.) : : . Potentilla reptans. This will, I think, illustrate the "deii dn above. The corolla is usually from three quarters to one inch in diameter, consequently it is very conspicuous and probably usually intercrossed. I found, however, specimens in flower _ on the 29th Sept. 1876, with the blossoms not half an inch in diameter; while in some ` ` buds unexpanded the pistil was covered with a dense mass of pollen, of which the tubes . 4 were to be detected penetrating the stigmas. If, then, my interpretation be correct, the ` ` inference I would draw is, sad Shin, was a process of EE induced SH the SELF-FERTILIZATION OF PLANTS. 363 lateness of the season, just as the Sloe is similarly, but normally, self-fertilizing because it blossoms so early in the summer; conversely, that conspicuously flowered plants with much honey are correlated with a high temperature and brilliant sunlight. It may not, perhaps, be too wide a generalization to conclude that all flowers with very numerous stamens, as in Ranunculacee, Rosaceæ, Myrtaceae, &c., are thus adapted for both intercrossing and self-fertilization, the former process being effected by the outer and first dehiscing anthers, and before the stigma is mature, the latter by the inner and later ones, which dehisce at the same time that the stigma is ready to receive their pollen. Lyraraceæ.—The genus Lythrum gives us another instance of a transition from a high state of differentiation in its adaptation to insect-fertilization in its trimorphic species to the degraded state seen in L. hyssopifolia, which is undoubtedly self-fertilizing. The intermediate condition is seen in L. thymifolia, which is only dimorphic. The stamens are reduced to six, which are homologous with the longest stamens, namely, those opposite the sepals. As observed in the Caryophyllacee, it is the rule that the whorl of stamens opposite the petals should disappear first. With L. hyssopifolia a still further point is reached. It is no longer heterostyled. As in the preceding, the six shorter stamens are often suppressed. Mr. Darwin says (‘ Forms of Flowers,’ p. 166) :— * The stigma is included within the calyx, and stands in the midst of the anthers, and would generally be fertilised by them ; but as the stigma and anthers are upturned, and as, according to Vaucher, there is a passage left in the upper side of the flower to the nectary, there can hardly be a doubt that the flowers are visited by insects." Whether this be the case or no, it seems to prove that this form is a degraded one, which has retained the relative positions of the stigmas and anthers necessary in L. Salicaria, but has acquired self-fertilizing powers as well. Its habit, too, agrees with homogamy, in being an annual and solitary and not like the other mentioned species, which are social. 5 Nesea verticillata. This, Mr. Darwin states, is trimorphic, while, according to Fritz Müller, a species of this genus in St: Catharina, in Southern Brazil, is homostyled. Cuphea purpurea “ was highly fertile with its own pollen when artificially aided, but sterile when insects were excluded” (* Forms of Flowers, p.168). It is therefore a case of morphological, and not physiological, sterility. Peplis Portula. I have not the slightest doubt but that this plant i is invariably self- fertilizing ; for all the features one is familiar with in Lythrum are entirely gone. The petals are reduced to a minute size, or else are wanting. The six stamens opposite the sepals are alone present. They incurve over the pistil, as is so thoroughly characteristic of self-fertilizing plants, and the capsule sets an abundance of seed, as is usually the case with such. Lastly, it is a small inconspicuous weed-like annual. ONAGRACE &. —JAEpilobium is an instructive genus from the point of view under con- sideration. E. angustifolium was first observed by Sprengel in 17 90* to be proteran- * This is quoted: on the authority of Sir John Lubbock (7. c. p. 27), and is certainly the case. Hence Mr. M. C. Cooke’s description in his ‘Manual of Structural Botany,’ as quoted by Prof. Dyer in ‘Journal of Botany,’ vol. ix. p. 22, must be wrong ; for it is described as being specially adapted to self-fertilization, ! *. 364 REV. GEORGE HENSLOW ON THE drous. Æ. hirsutum is, I think, so also; at all events the order of developmentis —calyx, stamens opposite the petals, pistil, corolla, which is the usual one for intercrossing *. In Æ. hirsutum the whole inner surface of the lobes are stigmatiferous or papillose and recurved ; but in Æ. parviflorum the lobes remain erect, and are papillose only at the edges. Moreover, they are not elevated above the anthers, and thus this species is self: fertilizing. In fact I feel disposed to regard it as a degraded form of E hirsutum, having adapted itself to homogamy. The club-shaped stigma, characteristic of other small- flowered species, as E roseum and E. tetragonum, is a yet further advanced stage, in which the lobes are completely welded together; they are, I need hardly say, self- fertilizing, especially, it would seem, by the longer stamens. The order of development is—calyx, pistil, st. opp. sep., st. opp. pet., petals. 4 Circea lutetiana. This species is both intercrossed as well as self-fertilizing. There — | is an annular disk surrounding the base of the style; but in no instance did I find any fluid within it in a number of plants growing, partly concealed, in a shrubbery. The filaments have a bend about halfway up, and the anthers. at first approximate the | stigma; they shed their pollen just as the corolla commences to expand. Pollinization | then takes place. Subsequently the filaments diverge, and not unfrequently the style is carried to one side by cohesion of the pollen-grains to the stigma (a fact, I see, Müller has also observed), into which I have seen the tubes penetrating while the grains were still lodged in the anther-cells (see Tab. XLIV. figs. 17 a, b). Gaura parviflora is even further degraded than the 2-merous Circea; for it has no corolla, and is cleistogamous, in that it is self-fertilizing in bud, as I found in See 4 growing at Kew. The order of development is—calyx, pistil, stamens. | GInothera biennis. In this species the style rapidly outgrows the stamens in Lett and is adapted for intercrossing. Late in September 1876, I found flowers not half an inch long with the anthers all clustered round the basal parts of the stigmas. "They dehisced before the corolla expanded. The stigmas were quite viscid throughout their whole length, and pollen-tubes were freely penetrating. In normal flowers the stigmas are elevated one fourth to one half inch above the anthers. Now this accidental condition, attributable, I presume, to the lateness of the season, was, however, exactly like the normal condition of another American species, (E. parviflora, in which the petals are not half an inch in length, the stamens are erect, and it appears to be self- fertilizing. May not this case, again, throw light upon the origin of some species, viz., that by adapting themselves to colder climates, the corolla becomes dwarfed, and the sexual organs mature more nearly together, and so a self-fertilizing form or =e is produced a Œ. bistorta. Tn this species the stigma is globular, the pistil does not Em? the stamens, but both mature together, and the plant is self-fertilizing. The globular stigma is here analogous to the club-shaped stigmas of Epilobium and the globular ones of homo- gamous Cruciferze. Ludwigia palustris, with the habit of Pelpis, i is, without doubt, also self-fertilizing. * I find Miiller has observed 26 species of insects visiting E hirsutum, and believes: self-fertilization to be ës sible; but he notices how E. = fertilizes itself (* Nature,’ vol. ix. ia E = EE EE EE EE e SEET ica stich abe EE p USUS ER SELF-FERTILIZATION OF PLANTS, e 365 Like that plant, the corolla of four minute petals is sometimes wanting. The stamens are reduced to four (opposite the sepals). The stigma is capitate, and it seeds very freely. So many genera have some one or more species with conspicuous flowers, and one or more other species which have them inconspicuous and self-fertilizing, that it is, I think, at least suggestive of the possibility of the latter having differentiated from the former by becoming self-fertilizing. A few out of many that might be mentioned will remind the reader of this fact. Cardamine hirsuta is small-flowered and highly self- fertile; but C. pratensis is larger-flowered, brightly coloured, and often barren. Stellaria Holostea is very conspicuous and proterandrous, but Stellaria media is profusely self- fertile. Malva sylvestris is highly conspicuous and proterandrous; but M. rotundifolia is inconspicuous and self-fertile. Lastly, large-flowered Geraniums are proterandrous, but smaller-flowered species are mostly or entirely self-fertile. Such cases might be multiplied almost indefinitely ; and I shall have occasion to allude to them again hereafter. The general conclusion I would draw is that the self-fertilizing species are derived from certain intercrossing forms (many of which may be now alto- gether extinct) by adaptation to self-fertilization, the probable cause of this being the neglect of insects to visit them. SAXIFRAGACEJE.— The species of Saxifraga are mostly more or less conspicuous, honey- bearing, and proterandrous; but Chrysosplenium has insignificant flowers, and the anthers and stigma mature together. Müller observes of C. alternifolium that, although itis thus adapted for self-fertilization, yet the general golden aspect of the clusters of flowers and leaves render it more or less attractive to insects. Drosera rotundifolia has cleistogamous flowers, while with D. anglica * the still folded petals on some plants in my greenhouse opened just sufficiently to leave a minute aperture; the anthers dehisced properly, but the pollen-grains adhered in a mass to them, and thence emitted their tubes, which penetrated the stigmas. These flowers, therefore, were in an intermediate condition, and could not be called either perfect or cleistogamie " (* Forms of Flowers,’ p. 329). UMBELLIFERÆ.—Of this extensive Order, the majority, with conspicuous umbels of flowers, are much visited by insects, are highly mellifluous, and for the most part pro- terandrous, "There are, however, some very insignificant forms, which one may suspect would prove self-fertilizing, but I have not had an opportunity of examining them, such as Bupleurum &c.; but Scandix Pecten-Veneris, with its minute flowers and very small umbels, is certainly self-fertilizing. The umbel consists of about five or six flowers only. The stamens remain incurved when dehiscing. There are two or three male flowers in the centre, the anthers of which burst later, so that it would appear that if the herma- phrodite flowers do not fertilize themselves, they can be pollinated by the subsequent male flowers. The retention of the incurved position of the stamens shows clearly that this method of self-fertilization, and which is particularly common, as in Potentilla Fragariastrum, is usually nothing more than the position of the stamens when in the bud retained after the corolla has expanded, though in Agrimonia and Alisma im are subsequently incurved. . SECOND SERIES.— BOTANY, TOL. I. 3E 366 REV. GEORGE HENSLOW ON THE Apium petroselinum. This is also self-fertilizing; for Mr. Darwin records that | covered plants apparently were as productive as uncovered. PASSIFLORACEH.—Passiflora gracilis. This species, unlike other Passion-flowers, is an annual, a feature characteristic of self-fertilizers, and * produces spontaneously numerous fruits when insects are excluded, and behaves in this respect very differently from most of the other species in the genus, which are extremely sterile unless fertilised with pollen from a distinct plant” (‘Cross and Self-Fertilization, p. 171), or even species. It is worth noting that this species differs from other members in the young internodes having the power of revolving. “It exceeds all the other climbing plants | which I have examined in the rapidity of its movements, and all tendril-bearers in the sensitiveness of the tendrils ” (* Climbing Plants, p.153). Such would seem hardly compatible with Mr. Darwin's idea of self-fertilization being injurious. Mr. Darwin also records the fact that “flowers on a completely self-impotent plant of Passiflora alata fertilised with pollen from its own self-impotent seedlings were quite fertile” (* Cross and Self-Fertilisation,’ p. 330). ! CAPRIFOLIACEX.—A. curious difference obtains between Sambucus nigra and S. Ebulus. In both the anthers are extrorse, while in the latter species the filaments are extraordi- narily thick and corrugated (Tab. XLIV. figs. 18 a, b). They stand erect in the middle of - the flower, and completely conceal the stigma and obstruct all entrance of the pollen. In order to be fertilized the corolla and stamens fall off, leaving the now viscid stigma exposed. As the flowers are in a dense corymb, they must get dusted by wind, or else later corollas fall upon them. With S. nigra the stamens spread away from the centre, the filaments are slender, and the stigma is fully exposed. They are sweet-scented, hence the blossoms “ are visited by several insects, but often fertilize themselves, as the stamens and pistil ripen simultaneously ” (* Wild Flowers in Relation to Insects,’ p. 109) but since the stamens are extrorse there is a greater probability of the pollen being transferred from one flower to another. : Symphoricarpus, or Snowberry. This is much visited by wasps and bees when the shrub is in an exposed situation, but it seems well adapted for self-fertilization. The pistil is mature just before the expansion of the corolla; the papillae are long and glutinous. The anthers are large, and arch over the stigma. I could detect no honey in the flowers, though there is a sort of inverted cone-like disk at the base of the style. Pollen-tubes were abundant in the stigma; but whether the grains had been brought by — bees or had fallen from the anthers, which shed their pollen as soon as the flower opens, I cannot speak positively. When the flowers happen to be ern there would ve less likelihood of self-fertilization taking place. 1 RunrACEX.—Some species of Galium, e. g. G. verum and o Mollugo, are proteran- drous, but G. Aparine and, I have no doubt, G. tricorne also are self-fertilizing. Mr. Darwin observed that the former species when protected Fraise quite as many seeds. as when unprotected. | Asperula Cynanchica. This flower has the stamens situated at a higher level iban ihe stigma ; puel it would seem to fulfil the oi once bcn to be ee SELF-FERTILIZATION OF PLANTS. 367 rule, but now found to have many exceptions, in that the pollen can fall from the anthers upon the stigma and so secure self-fertilization. © VALERIANACES.—Of this Order Valerianella supplies us with minute flowering species, of which V. dentata only have I had an opportunity of examining. In this species the anthers are incurved and dehisce very early, just before expansion of the corolla ; the stigma enlarges and curls backwards. I detected the pollen-tubes. The order of development is—corolla, stamens, pistil. The corolla is early developed, because it acts as a protecting organ, the calyx being almost obsolete. Composir£.—The great majority of this Order are undoubtedly favourable for cross-fertilization by insects, at least as far as the “ heads" are concerned. The con- trivance by which the pollen is swept out of the anther-tube by the growing style is also well known. There appear, however, to be many species with inconspicuous flowers, as well as, especially perhaps, of the conspicuously flowering Cichoracee, which are highly self-fertile. Though the structure is mainly the same as in the florets of the heads which are crossed, the stigmatic branches curl back to a much greater extent, re- sembling miniature rams' horns, and so insert their extremities amongst the pollen- grains of even their own florets as well as those of their neighbours. Such appears to be the ease with the Dandelion, Chicory, Hieracia, Ae Mr. Meehan notices that although the stigmatic branches of the Dandelion are closely adpressed at first, yet as they expand the pollen-grains which are attached “ to the line of the cleft” fall in between the branches, and thus secure self-fertilization. This, therefore, is an addi- tional security for being self-fertilized. Mr. Meehan also observes how the Chicory has all its florets fertilized before 8 o'clock, and by 9 A.M. have faded away [Tragopogon porrifolium does much the same]; but after the elevation of the pistil “there is not a cloven pistil that has not some pollen on the interior stigmatic surfaces.” He further adds, * I have observed the same in Dandelion and the Ox-eyed Daisy (Chrysanthemum Leucanthemum), as well as, I am sure, thousands that flower and perfect seed which no insect visits." I find my observations on the curling-back of the branches of the Style are corroborated by H. Müller, who alludes to more than one case and figures Hieracium umbellatum (* Befruchtung’ Ze p. 406). (Tab. XLIV. figs. 19 a, b.) Another and additional method obtains in Senecio vulgaris; for sometimes the stig- matic branches, though slightly separated, do ‘not protrude beyond the anther-tube or the corolla at all; the anthers, too, are often very slightly coherent, the ‘use’ of their being syngenesious is apparently no longer wanted, and so the original freedom is retained. Lastly, both conditions may exist in the same head, some florets having the styles protruding and recurved, others are not visible at all. (Tab. XLIV. fig. 20.) - Erigeron canadense. In this species, which has established itself about Kew, the inconspicuous outer florets have elongated styles and long branching stigmas, which all curl over the disk-florets and almost entirely conceal them. The stigmas of the latter do not spread out, but remain loosely in contact; these having thrust out the pollen, some of which adheres to the stigmas themselves, the ray (female) flowers get fertilized. Now although the ray-florets are fertilized by the disk, and the disk-florets are self- EE, m there ean; I rs be no doubt that the effect is the same in both = ! | co 3 E2 ES a: 368 REV. GEORGE HENSLOW ON THE the offspring of either impregnation being identical. Hence it would seem justifiable to consider all the florets of such members of the Compositz as do not receive the visits of insects as equivalent to being self-fertilized. This is quite in keeping with Mr. Darwin’s observations on the fertilization of flowers by pollen taken from another flower, but on the same plant, in that, in the majority of instances, such impregnation does no good. Erigeron canadense also illustrates the wide dispersion of self-fertilizing plants described below ; for although its native country is North America, yet it is “ now estab- lished in the greatest abundance as a roadside weed in almost all temperate and hot countries" (Bentham). A point perhaps of some significance with reference to the style of the Cichoracee is, that it is not only “ uniform in the tribe, but is also precisely be one most general in the female florets of the Order ” (Bentham). Lastly, the suggestion may be made that one reason why the Composite is such a predominant order resides in the fact of their being capable of both intercrossing and self-fertilization. : | CAMPANULACEJE.—The genus Campanula, with the remarkable collecting-hairs on the style, is highly differentiated for intercrossing ; but Specularia speculum produced almost as many capsules when covered as uncovered (Darwin) The stigmas of this genus are recurved, excessively like those of the Dandelion; and as the anthers reach to a great height, indeed almost to the summit of the pistil, the stigmas are evidently easily able to secure the pollen from them. M. Brongniart says that some species of Prismatocarpus (included under Specularia by Benth. and Hook.), as well as some Cam- panule, have no corollas, which “ does not hinder them from fructifying" (quoted by Michalet à propos of cleistogamous flowers, in Bull. Soc. Fr. vii. 467). I need hardly say that the absence of petals, on the contrary, is a sure indication of their being probably self-fertile. Bpecularia perfoliata has cleistogamous flowers; for a description of them I must refer the reader to Mr. Darwin’s ‘Forms of Flowers,’ p. 330. Prof. Oliver says that he has seen flowers on Campanula colorata in an intermediate condition between cleistogamous and perfect, witnessing thereby to the belief that the former are degraded states of the latter, but yet with special adaptations to self-fertilization. GENTIANACEJE.— Müller has made an elaborate study of the many Alpine species of Gentiana, and finds that while the majority are intercrossed, yet a few are adapted for self-fertilization. Thus he remarks of G. nivalis :— j “ The flowers are much smaller than in G. verna &c. and more distant one from another. One or some of the anthers commonly come into contact with the margin of the stigma and effect self-fertiliza- tion, in case cross-fertilization by insects is wanting " (‘ Nature,’ March 29, 1877). G. Andrewsi, according to Mr. Meehan, “ never opens at all” in America. Erythrea is one of the many plants whose corollas rapidly close, and by so doing wrap up the stamens and pistil, thus forcing the anthers into contact with the stigma, most probably therefore effecting self-fertilization (Tab. XLIV. figs. 21 a, b, c). - CoNYOLYULACEE.— Ipomea purpurea, or Convolvulus major. This proved to be highly self-fertile with Mr. Darwin. The self-fertility of this, as of other plants of this Order, is machi facilitated z the -er withering of the S Zeen BE the ant ers ‘SELF-FERTILIZATION OF PLANTS. 369 and stigmas together. Of Mr. Darwin’s experiments, Ipomæa was cultivated for the longest period, or ten years, This plant ought therefore to give the best results for showing the advantages of crossing. Omitting the tenth *, it is found that the ratio of the heights of the intercrossed to the self-fertilized does not steadily increase in suc- cessive generations, as one would à priori suppose it “ ought,” from the continued infu- sion of new vigour by a fresh cross every year; but it fluctuates, so that when repre- sented geometrically we do not obtain a regularly ascending curve but a series of * maxima." Hence, to ascertain the true result, it is advisable to take the mean of two or three years’ growth together. Dividing, therefore, the nine years into groups of three each, the mean ratios thus obtained of the heights of the intercrossed (given as 100) to those of the self-fertilized are as follows : —For 1st three generations, as 100 : 74:3; for the 2nd three, as 100: 77:6; and for the last three, as 100 : 81:6. That is, the ratio was approximating unity or equality. Again, of the years in which Mr. Darwin has recorded the ratio of fertility, ascertained from the number of seeds per capsule; they are as follows :—1st year, 100 :93 ; 3rd and 4th years, 100 : 94; 5th, 100 : 106:9, and the 8th year as 100: 1147, which shows a steady increase in favour of self-fertilization. Hence, whether we look to “heights” or “fertility” as standards, there does not appear to be any evidence of intercrossing giving a yearly increasing, much less a permanent advantage. Moreover, as with Mimulus so with Ipomea, a strongly self- fertilizing variety sprang up, which, like Piswm and Canna, proved very impatient of cross-impregnation. (See remarks on Ophrys apifera, p. 978.) Mimulus luteus, which is self-fertilizing, gives somewhat analogous results; for, taking the ratios of | heights as before, the mean for the first two years is as 100 : 63:5, of the third generation (the mean of all the heights given, p. 66) 100 : 76, and of the fourth and fifth as 100 : 103; after which the “ white variety," which was so highly self-fertile, became so abundant as to render further experiments unnecessary, and perhaps impossible. In almost all the other experiments the number of generations are too few to be analyzed in this way; but, as a rule, they all prove that a first cross imparts some sort of stimulus ; but there is nothing to show that the effects would be permanent. Moreover, in all cases, some plant or plants of the self-fertilized beat its opponent. Henee I do not see how we can avoid the conclusion that the stimulus to growth afforded by crossing is not at all lasting, but gradually disappears, till the self-fertilized prove to be either equal or superior to them in the long run; so that I am led to accept Mr. Meehan's interpretation as follows :— “ Mr. Darwin’s artificial experiments seem to show, not that self-fertilization produces any injury to the race, but that cross-fertilization brings about a more excitable condition of growth and reproduction” f. The effect may be, perhaps, compared to a tonic or many other drugs, which, however * I omit the tenth, as the difference between the heights of the intercrossed and self-fertilized plants was so great as to be probably “ accidental,” so that Mr. Darwin thinks it does not give the true ratio. + This is deduced from a statement on p. 39, that a self-fertilized plant produced “ the large average number of 5:1 seeds per capsule." But as Mr. Darwin does not give that of the intercrossed, I have taken the number 4:46 from a previous year (p. 36), which gives the ratio 100 : 114. t From the Pennsylvania Monthly, Philadelphia, June 1877, 370 REV. GEORGE HENSLOW ON THE potent at first, if continued for a long while completely lose their effect by the system becoming attuned to them. Sotanacea.—Solanum dulcamara is said to be not much visited by insects, yet it sets its berries very freely and is more or less proterandrous. S._nigrwm, however, is decidedly self-fertilizing. The flowers are tolerably inconspicuous and pendulous; the anthers are close round the style, but are not * sub-syngenesious," as in S. dulcamara. They dehisce by slits all the way down, and so shower their pollen over the stigma, which sets its berries in great profusion. (Tab. XLIV. fig. 22.) The anthers do not | appear to have the spirals in the cells of the walls, which indieates, therefore, that this method of dehiscence is a subsequent acquirement from the condition obtaining in S. dulcamara, the anthers of which dehisce by pores. A. nigrum, therefore, affords another proof that existing self-fertilizing plants are all de&radations from more con- spicuously flowering ancestors. S. nigrum is also white, the commonest colour of self- fertilizing plants. > Nicotiana tabacum proved to be fully self-fertile with Mr. Darwin. A plant growing | in Kew Gardens was clearly proterandrous; but W. rusticum, which has smaller dingy green flowers, was as obviously self-fertilizing. The pollen-tubes were abundant. SCROPHULARIACEX.— The many genera with conspicuous flowers, and all being more ` or less irregular, are obviously adapted for insect-fertilization ; but there are several inconspieuous ones which are habitually seif-fertilizing. "Thus of the genus Veronica, V. spicata, blue garden-variety, is proterandrous, and so much so, that in some instances it is not until the corolla and stamens have actually fallen off that the style elongates * (Tab. XLIV. figs. 26 a, b, c). "This and V. Chamedrys, with its brilliant blue corolla, are intercrossed. The order of development of This species is—calyx, stamens, pistil, corolla ; and though the anthers and stigmas mature apparently quite or nearly simultaneously, yet the stamens spread away, while the style projects forwards and is not, therefore, specially adapted for self-fertilization. On the other hand, V. hederifolia, V. serpylli- folia, and V. agrestis, which is, perhaps, the self-fertilizing form of V. Buxbaumii, as well as V. Anagallis, are self-fertilizing (Tab. X LIV. fig. 27). In these the order of development is the same, viz. calyx, stamens, pistil, corolla. In V. serpyllifolia the ` pistil rapidly elongates, even beyond the stamens, just before expansion; the stamens - * Such, at least, applied to some specimens I found growing in a much-shaded garden, and have recorded in my notes that the corolla had the tube densely clothed with upturned hairs. The style, only just or not at all protruded» is green and the stigma immature, while the anthers, on elongated filaments, are turned vertically downwards. As soon as they have shed their pollen the corolla is ejected. The style then elongates and curves slightly downwards, becomes bright blue, and the stigma enlarges. The relative length of the style is now the same as that of the stamens, or one fourth of an inch. The development of the stigma subsequent to the ejection finds its parallel in Sambucus Ebulus, antea, p. 366. Much the same occurred in a white variety, only the style was of the same length ` as the stamens when the anthers dehisced, but the stigma still immature after the ejection of the corolla; the style a elongated to about half of its former length, then shrivelled. No seeds were detected in any capsules. The ` order of development is—calyx, stamens, corolla, pistil. Since this note was penned, I find that Müller has also dis- - covered this plant to be very peculiar; for he has met with plants which present exactly the reverse conditions, in ` that they are are while = like those seen by me, were Leg Zeie ad) He Bat both. kinds i c. p. 287).. ae SELF-FERTILIZATION OF PLANTS. 371 then in their turn elongate, and so both stamens and pistil are of the same height and mature together. Moreover, instead of spreading out, the filaments are erect and parallel with the style, so that the anthers and stigmas are in contact. The corollas of the self- fertilizing species have all a tendency to remain only partially expanded; and as the anthers burst when in this condition, self-fertilization is secured. The extraordinary difference between the rapid development of the style in these species and its retardation in V. spicata, more especially the blue variety, is very curious. The fertilization is secured by bees and large Diptera visiting the flowers which have got their corollas; but in so doing they sweep their bodies over the long styles m project from the lower “ corollaless" flowers. ! Serophularia nodosa, with its small flowers, is proterogynous, and is “much frequented and fertilised by wasps” (Sir J. Lubbock, 7. c. p. 137). This fact is quite in keeping with the dwarfed size of the corolla, and moreover harmonizes with S. canina, dried specimens of which from Castleton, America, bore very small blossoms, of which the long style was much recurved, and had the stigma lying between the anthers. It had therefore all the appearance of being self-fertilized. The preceding was written before I found that Mr. T. Meehan had described it; for he thus speaks :— “ The pistil protruded while the anthers were’ still rolled back in the throat of the corolla. One by one the stamens were straightened out, the anther coming into close proximity with the stigma, when it burst, and by the contraction of the sacs the pollen was ejected, falling on to the stigma” (Proc. of Acad. of Nat. Sc. of Philadelphia, p. 13, 1876). The early growth of the pistil here mentioned quite corresponds with my general observations of self-fertilizing plants, and which have been several times alluded to. Linaria vulgaris. Mr. Darwin found this plant to be extremely sterile when covered up. I found some plants, in September 1876, with small spurred as well as spurless corollas. The pollen-tubes were penetrating the stigma from both the anthers above it as well as from below (Tab. XLIV. fig. 28); and Mr. C. B. Clarke informs me he has found similar aborted corollas, but of self-fertilizing flowers like the above, early in the season. Müller appears to infer that self-fertilization may take place at any time, from the relative positions of the essential organs, though Darwin's experiments do not, as stated, corroborate this. It may be, therefore, only in checked buds where self-fertiliza- tion occurs; for Müller alludes to this being the case with Scrophularia nodosa, which in warmer weather is fertilized by wasps; and he remarks that the capsules are just as replete with seeds from self-fertilization in cold and wet weather as by the aid of wasps when it is fine. The inference to be drawn is that self-fertilization was being attempted in correlation with the dwarfing and more or less arrested condition of the corolla, the two phenomena being so generally associated in plants which are habitually homo- gamous, L. minor. In this species the position of the stigma is also just between the anthers, and is. s abundantly self-fertilizing. Antirrhinum majus. Mr. Darwin records the fact that uncovered plants of the red e Kee of ES Sege bore more than twice the weight of seed as compared, with covered: | 372 REV. GEORGE HENSLOW ON THE plants; but of the white variety with a pink mouth, “ fifty pods, of which only a very — few were empty, in a covered-up plant contained twenty grains weight of seed; so that ` this variety seems to be much more self-fertile than the previous one." "This case is | already shown in harmony with the fact that loss of colour seems to be often correlated — with self-fertilization. Again, as will be noticed further on, the majority of the British | species of plants, which are generally diffused over the world, and are for the most part ` self-fertilizing, are also white. Mr. Darwin observes that the “ peloric variety” of - this plant is morphologically, but not physiologically, self-sterile, * as humble-bees ` cannot crawl into the narrow tubular flowers.” Ido not know whether peloric varieties : of other genera, as Linaria and Calceolaria, are thus self-sterile ; if so, it would seem to 4 be a case where plants varied so as to bring about an injurious result; for in Antirrhinum — it appears to have excluded bees, and at the same time has not become self-fertilizing. Verbascum is another very interesting genus. V. pheniceum and V. nigrum Mr, Darwin regards as ‘‘ quite sterile," that is, without insect aid; on the other hand, | y. Thapsus is “ perfectly self-fertile” if insects are excluded. V. Lychnitis is rather ` e self-fertile * than the preceding " (p. 89), though Mr. Darwin elsewhere describes it ` * quite" and “ highly self-fertile" (p. 369). ** Kolreuter” (as quoted by Darwin, : itia &e.' p. 830), “long ago described plants of V. pheniceum which during two years were sterile with their own pollen, but were easily fertilised by that of four other species; these plants, however, afterwards became more or less self-fertile in a strangely fluc- — tuating manner." This case, then, elearly corroborates the general inference that, just as 3 the different species of a genus may vary from complete sterility to complete self-fertility — in their normalstates, so individual species can do the same under temporary changed | environments. This genus, therefore, resembles Corydalis in the former peculiarity, and Eschscholtzia californica and Papaver vagum in the latter. Calceolaria (greenhouse variety) is, according to Mr. Darwin, highly self-fertile. The position of the stigma on the erect style, which stands exactly between the two or three — ` anthers, or a little below them, on their erect filaments, is obviously favourable to self- ` fertilization, though the flowers are much crossed by insects, and it is therefore difficult ` — to keep varieties true (Tab. XLIV. fig. 20d). Some species, as C, eAelidonoides, O. pin- — nata, and C. glutinosa (Tab. XLIV. figs. 29 a, b, ei, with viscid foliage, have a remarkable — modification in the structure of their stamens. They are, in faet, similar to those of —— . Salvia, in the Labiatze ; so that these species are probably not readily self-fertilized. They ` constitute the section Aposecos (Gen. Pl. p. 930). The fertile anther-cells are concealed beneath the hood on the posterior side, the connectives, which are elongated, and with, in some cases, barren anther-cells, project outwards. The filaments are, as in Salvia, very short. On depressing the exposed anther-cells, the fertile ones swing round p as in that genus. ~- Mimulus luteus. Mr. Darwin has down that this species is | highly self-fertile when insects are excluded, though the bright yellow corolla, didynamous stamens, and irritable ` ` flap-like stigmas are evidently adaptations for intercrossing. This species is also notice- - : able for the tall white-flowered variety raised by Mr. Darwin, which proved so vigorous | and self-fertile. * Inthe fifth me the crossed pes were in sisi e to fui SELF-FERTILIZATION OF PLANTS. 373 fertilised as 100 to 126; in the sixth [in number of capsules]* as 100 to 147 ; and in the seventh [in height] as 100 to 137. This excess of height may be attributed not only to this variety naturally growing taller than the other plants, but to its possessing a peculiar constitution, so that it did not suffer from continued self-fertilisation " (p. 80). Con- ~ Geen it appeared to thrive by it, and unmistakably shows that there is no “ injurious- ness" per se in self-fertilization at all. Euphrasia officinalis, Dr. Miller thus describes the small self-fertilizing form of this species :—‘‘ Whilst in the flowers of the larger form the anthers remain soldered together, and do not scatter their pollen unless the hairs are shaken, in the flowers of the smaller form the anthers separate from each other, and scatter nearly all their pollen long before the corolla has fully opened.” Moreover, in this form, the deflexed hairs on the anthers are mostly wanting. Mr. Darwin’s experiment appears to have been with the intererossing form, as he alludes to the “ bristles which project from the anthers; " yet, when covered up, this plant “ produced plenty of seed." If this be a correct surmise, doth forms are highly self-fertile. Ascherson regards them as varieties. Rhinanthus Crista-Galli has also two forms, « and p, L., Rh. major and Rh. minor, Ehrh. i á Melampyrum: M. americanum is thus described by Mr. Meehan :—* The curved apex of the pistil is clasped by the stamens, and held in contact with the pollen just as in a cleistogamous violet," and is, he assumes, self-fertilizing. In these and other allied forms the self-fertilization is generally secured by the style curving down over the front of the stamens, and so reaching the anthers (Tab. XLIV. fig. 30 b); or else, as in Prunella, being too short to arch over them, stands below, and so allows the pollen to fall down upon them (Tab. XLIV. figs. 24 a, b, c, illustrate this). Vandellia nummularifolia has cleistogamous flowers, and proved more fertile when self-fertilized than when erossed by Mr. Darwin in the ratio of 752:598 or 100 : 79:5. Limosella aquatica. "The minute blossoms, globose stigma between the anthers, and the ‘abundance of seeds all point to the conviction that it is self-fertilizing ; but I have had no opportunity of examining it in the living state. .. OnosANCHACEJE.— Some plants of Orobanche Hedere I found growing on the St. Vin- eent’s Rocks at Clifton, having in the early stage of the flower a well-developed ond glutinous stigma projecting forwards before the anthers dehisced, as shown in Tab. XLIV. fig. 31a. A later stage is seen in fig. 31 b, in which the anthers were recurved about the stigma, and self-fertilization apparently secured. (This requires corroboration.) Epiphegus virginiana. This is described in the * Genera Plantarum ' as having the supe- rior flowers hermaphrodite, very often, but not always, sterile; the inferior cleistanthic female, fertile, with abortive stamens, but the ovary perfect. "This, if correct, would seem to record a case of parthenogenesis ; but in a specimen sent me by Mr. Meehan, with the lowermost cleistogamous buds subterranean, I found that each had two stamens, the anthers of which adhered to the stigma and were carried up by the enlarging eni (as * The ratio 100: 147 does not refer to heights, as Mr. Darwin has inadvertently included it in the text, but to capsules. For further inferences from this plant see above, p. 369 (l. c. p. 69). SECOND ——— NR) SEE Ae ee 974 REV. GEORGE HENSLOW ON THE in Violets), and lay concealed under the little knob at the top of the calyptriform corolla (Tab. XLIV. fig. 32a). The seeds in each capsule were innumerable. Lasiata.—Like the Scrophulariacee, the plants of this Order are mainly constructed for intercrossing, yet some small-flowered species are undoubtedly self-fertilizing, while a few are cleistogamous. Prunella vulgaris, according to Axell (quoted by Miiller), is self-fertile in the absence of insects. I had suspected this to be the case before I was aware of Axell's opinion, because of its very wide dispersion (see below, § 17, p.393). I have since repeatedly ex- amined it, and quite concur with the view, though Müller himself does not appear to have observed it. He gives a figure (Befrucht. Ze p. 318), in which the stigma is represented as having its lower branch between the anther-cells. I found a considerable amount of variation * in this respect; for while many flowers were as he has figured it, several had the stigmas considerably below them, and often with the posterior lobe curled upwards between the anther-cells, and becoming thus pollinated, as in Tab. XLIV. figs. 240, c. Perhaps the most interesting genus is Salvia. The stamens, as have been described by different observers, are levers of the “ first kind," oscillating in a vertical plane, exactly like Calceolaria glutinosa and other species of the section Aposecos. The details of size . &e. vary in different species; but the action is much the sanie in most of them f. In the following self-fertilizing species, however, the adaptations are completely changed. S. clandestina. In this species the two anther-cells, instead of having their lines of dehiscence looking downwards over the lip, face each other, so that the broad surfaces are in a vertical plane. The stigmas are greatly elongated, and curl backwards between the anther-cells, and thus get pollinated. Both anthers and stigmas are almost, if not quite, concealed within the hood. The flowers are altogether very inconspicuous as compared with other species (Tab. XLIV. figs. 23 a, b, c). Lastly, Sir J. D. Hooker informs me it is a particularly common species on the continent, which is quite in keeping with its being self-fertilizing. . Müller refers to three others species, A. Grahami, S. lanceolata, and S. hirsuta, as described by Hildebrand, as being also homogamous and self-fertilizing ; and the description given above seems much the same as for these latter. One species, S. cleistogama, has cleistogamous flowers, which is therefore a more advanced stage in degradation. ; Lamium. Though this genus has highly differentiated flowers adapted for insect nines yet "A amplexicaule has cleistogamous blossoms which appear in the early spring and again in autumn, in addition to the normal flowers. Here, I think, we have evidence of- the influence of temperature which, when reduced, strongly affects the development and expansion of corollas, but without influencing the sexual organs, except as m" to » estate? in length of filaments and styles is of common occurrence in plants, and bring about different methods of fertilization accordingly, some involving the necessity of intercrossing, others sélf-fertilization. _ t For a full description of Salvia see a paper by Dr. Ogle in the * Popular Science Review,’ July 1869. Many years ago I “discovered” this remarkable adaptation; and in the vain imagination that it was “ new,” called Mr. Darwin's attention to it. He kindly referred me to the description and figure by Sprengel, published i in 1790, of a Humble-bee oe inte a es flower im Salvia aad being smitten on the ! . SELF-FERTILIZATION OF PLANTS. 375 render them homogamous. That the cleistogamous blossoms of this species are degraded forms of the normal kind is obvious from the presence of the **lip," as well as by there being four and didynamous stamens. The style elongates very much, and, under the pressure of the closed summit, becomes bent, so that the stigma lies between the anther- cells, which thus pollinate it (Tab. XLIV. figs. 25a, b, c). BonAGINEJE.—Sir J. Lubbock (7. c. p. 131) observes of this Order:—** As Müller has well pointed out, there are the widest differences in the conditions of fertilisation. Pul- monaria officinalis is dimorphous and sterile, not only with its own pollen, but even in some cases with that of a different flower, unless it belong to the different form. Echium vulgare has lost the power of self-fertilisation, but, so far at least as we know, is fertile with the pollen of any other flower belonging to the species. Other species are generally fertilised by insects, but in their absence perform this office for themselves ; while, lastly, some species, such as Lithospermum arvense and Myosotis intermedia, habitually fertilise themselves." di With regard to the order of development of the floral whorls of some of the Boraginex, I find the pistil lags behind the others in a remarkable manner, but subsequently grows rapidly. This at least is the case with Borago officinalis *, the order of which is, at first, calyx, stamens, corolla, pistil. The corolla, however, soon overtakes the stamens, while the pistil remains rudimentary for a long while. It is the same with Anchusa officinalis . and Myosotis arvensis. With Myosotis versicolor I find there may be recognized three stages. In the first the pistil remains for a time very rudimentary, then it rapidly grows and overtops the stamens; but, thirdly, the corolla, by enlarging, lifts the anthers, so that the stigma now lies on a level with the middle of the anthers. Müller thus describes this flower (* Nature,’ x. p. 130) :— * The corolla, when opening, is not only still of a pale yellowish colour, like the buds of other species of Myosotis, but even when not yet fully developed ; the anthers and pistil are mature at the same time, and the stigma slightly overtops the corolla. Hence, when insects visit the flowers in this state, their probosces always touch the stigma sooner than the anthers, and consequently, when flying to another flower, always cross-fertilize it. But, by the gradual lengthening of the corolla-tube, the anthers affixed to its inner side are raised till they surround and fertilize the stigma, now enclosed in the corolla." I have drawn three stages of growth, which illustrate the relative heights of the stamens and pistils at successive periods (see Tab. XLIV. figs. 33 a, b, c). It is probable that several small-flowered gamopetalous species are self-fertilized during the fall of the corolla, as by that process the anthers are dragged over the stigma. Such, for et än probably occurs with Myosotis sp., Anchusa, &c. | Amsinkia angustifolia. A very small yellow-flowered plant from Kew. The order of x The ejection of the ‘tit of Borage is rather peculiar. As soon as the anthers have shed their pollen, the calyx-lobes contract, and the petal-lobes, from having been somewhat reflexed, become concave. These two actions combined throw off the corolla. The appendages to the stamens are outgrowths by “ enation," and are not developed till late. I am doubtful as to their purpose. They seem to press upon the anthers, and so form a hollow cone, with a circular hole at their apex. Bees stand at right angles upon the stamens when sucking the honey from the . Rectariferous appendages on the corolla; the pollen thus falls on one 2m on their abdomens; and as the stigma i is (meli over this ume spot, intererossing is effected: 376 REV. GEORGE HENSLOW ON THE development is exactly the same as in Geer versicolor, ait appears, like it, to be self-fertilizing. PRIMULACEH.—This order is most conspieuous for its heterostylism. Primula. Though probably the majority of species of this genus are dimorphic, there are some which are normally non-dimorphie *. Others, again, have been observed to fluctuate; thus Mr. Scott mentions P. pusilla and P. floribunda, both of which are, as far as his observatibli went, solely short-styled forms, yet he found an individual of each with stamens and pistil of the same length. With regard to the position of the anthers of non-dimorphie forms, they may be in either position.—P. scotica. Mr. Scott noticed that in native specimens of this species the anthers usually surround the mouth of the corolla-tube, while in cultivated specimens they are generally attached about one third of the length | of the tube below the orifice; the length of style varies accordingly.—P. sibirica. The native variety of this, with oblong entire leaves, was always dimorphic ; but the culti- vated specimens of the variety, with ovate crenate leaves, was always non-dimorphic, —P. verticillata. In both wild and cultivated specimens the structure is the same, viz. | the stamens are attached to the upper third of the tube. The length of the style is sub- ject to slight variations. With regard to the fertility of these plants, Mr. Scott found that when P. scotica was protected, it produced seeds per capsule in the ratio of 1:2: 24. , às compared with the seeds of artificially fertilized flowers. P. mollis greatly exceeded all other species which Mr. Scott examined in its * regular self-fertility," inasmuch as * nearly every flower produced a capsule filled with good seed." Other non-dimorphie species produce “ very generally a high percentage of abortive capsules, together with a- great variability in the number of seeds contained in those that do set seed." P. verti- cillata, contrary to the preceding, presents an imperfect functional dimorphism in con- junction with a non-dimorphie structure. In 1862 this plant proved to be self-sterile; but in 1863 “certain of the flowers, fertilized by their own pollen, yielded a considerable amount of seed ; " but the relative value of crossing, as compared with self-fertilization, appears in the ratio of 4:1. Primula veris is sometimes non-dimorphie, and, according to Mr. Scott, is then more fertile than by legitimate unions, thus :—Red cowslip, non- dimorphic form, yielded average number of seeds per capsule 34. Do., long-styled des do. 28. Do., short-styled form, do. 20 (7. c. p. 106). | Primula Sinensis. Mr. Darwin has given details respecting this slant (* Cross and Self-fértilization, p. 225), and records that although legitimate unions were more fertile than illegitimate, v * there was n difference in growth between the offspring fro legitimate and illegitimate union." Mr. Darwin accounts for this by the fact t | English plants are commonly raised from self-fertilized seeds; so that, as with common Pea, which is habitually self-fertilized, crossing does little good. “ Moreo many of the plants are now varying and changing their character, so as to become greater or less degree equal-styled, and in eonsequence highly self-fertile " (2. e. p. 225 ‘Such conditions, then, I ev to be reversions to an ancestral equal-styled state ; and — * Mr.J. Scott, in a paper on “The Functions and Structure of the Reproductive Organs in the io Donen? (Journ. Linn. Soc., Bot. viii. p. 7 8) gives. 36 Tas which are ger 7 msdn d ee oue pone SELF-FERTILIZATION OF PLANTS. 377 although the plant may perhaps lose its beauty (from our point of view), as the one I have described above, yet that is of no consequence to the plant—its end being not size or colour, but easily acquired, rapid, and abundant propagation ; and this it does acquire by becoming non-dimorphic. Primula vulgaris. I found in spring 1877 a plant of the Common Primrose which bore non-dimorphic flowers, the first, I believe, known in this species, It was the * short-styled " form, judging by the exserted position of the stamens; but the style was long, so that the stigma was situated in the middle of the stamen, and self-fertilization was taking place, Lysimachia. Miller has observed that L. vulgaris has extreme forms, on the one hand requiring insects, and on the other habitually self-fertile and inhabiting shady places, and that these forms are connected by intermediate links (J. Lubbock, 7. c. p. 126). Anagallis cerulea is smaller-flowered than A. arvensis and is homogamous. The ` flowers are probably seldom visited by insects, and it would appear that they generally fertilize themselves. Müller describes the structure of this flower, and adds much the same remarks as I have elsewhere made—that, as the corolla closes at 3 o'clock, the petals bring the stamens into contact with the pistil, so that self-fertilization is secured (l. c. p. 349). Glaux maritima. In this the style is much longer than the stamens, but curves back so as to bring the stigma into the midst of the anthers, and is consequently self-fertile and seeds profusely (Tab. XLIV. fig. 35). | Centunculus minimus and Samolus Valerandi are both regularly self-fertilized. PrneuicuLacr£.—tThe larger-flowered species of Pinguicula are doubtless adapted to intercrossing ; but P. lusitanica is self-fertilized by the stigma being recurved into the anther-cells (Tab. XLIV. figs. 34a, b, c). IxcouPLETA.— The prevailing absence of all bright colours in this division of Dicotyle- donous Angiosperms is clearly correlated with the absence of the visits of insects. Ex- ceptions of course occur, as with the Buckwheat. If we leave out of consideration the many diclinous forms which are wind-fertilized, we have a residuum which, there is but little doubt, are self-fertilized. CnENoPoDIACEX.— Beta maritima. Mr. Darwin says that the Beetroot is fully self- fertilized when netted. ~ Salicornia herbacea. The stamens protrude their anthers first from the nearly closed orifice of the perianth, and only one at a time; then the stigmas rise two or three to each pistil. As there are three flowers close together in each “article,” it is very difficult to detect whether the individual flowers are self-fertilized, or whether the three flowers do not fertilize one another. ‘The branched stigmas have the ‘‘anemophilous”’ character. Salsola Kali, as examined by me, was sometimes proterogynous, the stigma being then quite withered before the anthers dehisce. In other cases the style at first extends beyond the anthers; afterwards the latter overlap the stigma and cover it with pollen, - so that it is probably self-fertilizing. Hence this plant seems to be sometimes protero- * - 378 REV. GEORGE HENSLOW ON THE gynous and wind-fertilized, but at others self-fertilizing by the pistil being slightly delayed in its development. Sueda maritima. The order of development of this plant is—calyx, pistil, stamens; but the stamens outgrow the pistil, curve over it, and dehisce while the lobes of the calyx remain more or less unexpanded. It is thus self-fertilizing. PoLYGONACE®.— Polygonum is a genus affording considerable differences in the sexuality of the species. P. Fagopyrum Müller has shown to be dimorphic, and P. Bis- torta proterandrous; but I find P. Persicaria, P. aviculare, and P. Convolvulus and P. Hydropiper are all self-fertilizing. P. Persicaria secretes a little honey,but insect-visits are probably very rare; so that it may represent an intermediate condition between intercrossing and purely self-fertilizing forms. P. Convolvulus and P. Hydropiper appear to be always cleistogamous, at least I have never seen the perianth expanded. The stamens all arch over the pistil, and the anthers cover the stigmas with pollen. EvPHORBIACE.—Mr. Bennett found that Euphorbia Helioscopia and E. i. Peplus were self-fertilizing, i. e. if we may use the expression for the moncecious inflorescences of these plants. Other species, such as E. amygdaloides and some tropical forms, with brilliantly coloured bracts or glands, are strongly proterogynous. | MOoNOCOTYLEDONS.—l have not made many observations on plants of this class; but ihe same principles appear to hold good, namely, that conspicuous flowers are adapted for intererossing by insects, but inconspicuous ones are either self-fertilizing or anemo- - philous. : OncurpACE E.—Mr. Darwin has recorded that Ophrys apifera is highly self-fertile, and - is therefore an exception to the rule, of a conspicuous flower being not crossed by insects. Cephalanthera grandiflora is also described by that author as a case of “ perpetual self- fertilisation, but in an extremely imperfect degree" (Fert. of Orchids, p. 111). ps If we ask why the Bee Ophrys is so prolific, is one of the most abundant and vigorous : of Orchids, and is yet perpetually self-fertilized, I presume the answer would be that it is descended from an intererossing form which was crossed, and that the benefits of the ` cross have been inherited and become permanent, so that it no longer requires any more - extraneous aid. This, at least, is Mr. Darwin's argument to account for Andrew Knight's | varieties of Peas lasting for 60 years, though regularly self-fertilized (Cross &c. p. 305). My reply is that the entire argument is an à priori inference, based upon the. supposed necessity of crossing. It is an argument which, from the nature of the case, cannot be proved. On the other hand, none of Mr. Darwin’s experiments countenance the idea that the effects of crossing are permanent; whilst in many cases plants which either are ` habitually self-fertilized, or have acquired self-fertilization, showed great indifference to being erossed, and were, in fact, vastly superior to their intercrossed competitors. More over, plants like Polygonum Convolvulus, P. Hydropiper, and Hordeum murinum are, as | far as I have observed them, habitually cleistogamous, the flowers never expanding, and en ` cannot be crossed. Hence my argument that crossing is superfluous for all such plants ` is equally, if not more, pertinent. (See remarks under Ipomea and Mimulus, p. sag | AXISMACEA.—4lisma Plantago. This plant appears to me to resemble Agrimo in daos at first Late for eres ne the anthers spr A SELF-FERTILIZATION OF PLANTS. 879 stigmas, but afterwards, by becoming reflexed, as well as by the styles now spreading, to be self-fertile. The stamens and carpels mature well before the buds expand. The anthers are extrorse, but the filaments become strongly curved backwards, so that the anthers hang more or less over the stigmas, though dehiscing upwards. The styles are long, and bend backwards and outwards over the ovaries. The anthers burst just as the perianth expands, or even a little before, and the stigmas are mature to receive it. The flowers are not very conspicuous, and every carpel is “set” (Tab. XLIV. fig. 37). If it be self- fertilizing, then we may, I think, safely believe the other species of Alisma to be so as well; and I would include Scheuchzeria and Triglochin also, as being most probably golf fartiliziny’s but I have had no opportunity of examining them. I now find Dr. Müller has seen certain Diptera to be attracted by the small honey- glands between the stamens of Alisma Plantago; yet he adds, this species is very pro- bably self-fertilized as well. In his figures he draws the styles erect, though still possessing a bend near the base; while the anthers are drawn spreading far away from the carpels. In this condition shy are evidently specially adapted for intercrossing ; whereas in the later stage they appear to offer better facilities for self-fertilization. Under this interpretation we see the use of the * bend " in the style in the latter cases, but its “uselessness ” in the flowers figured by Müller. JUNCACEJE.—Members of this family are strongly proterogynous; but Juncus bufo- nius is remarkable by bearing in parts of Russia only cleistogamic flowers, which contain three, instead of six anthers found in the perfect flowers” (Darwin). It is the most widely dispersed species (see p. 393). GRAMINE®.—Grasses are mostly anemophilous ; but some are self-fertilizing, and others even cleistogamous*. Thus, *In the genus Hordeum it has been shown by Delpino that the majority of the flowers are ee some of the others expanding and apparently allowing of cross-fertilization.” (Tab. XLIV. fig. 38. See description, p. 398.) Leersia oryzoides. ‘The cleistogamic flowers are very small, and usually mature their seeds within the sheaths of the leaves” CG Forms of Flowers,’ p. 333). This appears to be the only species of this genus which bears cleistogamic flowers; and what is particularly interesting to me is that it ranges from Persia to North America; so “there can therefore be little doubt that this plant generally propagates itself throughout an immense area by cleistogamic seeds, and that it can hardly ever [never ?] be invigorated by eross-fertilisation ” (J. c. p. 335). Poa annua. This has already been alluded to as a good instance of a self-fertilizing plant ripening its seeds with astonishing rapidity. Our London parks are much in- debted to this little plant and its extraordinary powers of propagation. Hordeum murinum. The central floret of the spikelets appears always cleistogamous (Tab. XLIV. fig. 38). * Mr. A. S, Wilson, in a paper on the * Fertilization of Cereals,’ contributed to the Transactions of the Botanical Society of Edinburgh, thinks that the European cereals are self-fertilized, and that the act of fertilization in those eases in which the flower opens is probably performed in the opening, and is necessarily confined to the twenty or thirty minutes’ duration in which the flower remains open. Mr. Bennett, in his notice of this paper, controverts, en - = a ihn, mins xoa : A. 380 REV. GEORGE HENSLOW ON THE Summary of the chief adaptations of the various parts of plants to secure self-fertilization. | Bracts.—These may assist in those genera of the Compositze which close their heads — in the evening, as do Taraxicum &c. Calyx.—The same remark applies to the calyx; for the sepals afford no special aid, - but only indirectly by pressing on the corolla in bud, and so keeping the essential organs ` in contact with one another. Corolla.—This effects fertilization in some cases by withering, and so matting together the anthers and stigmas, either habitually, as in Hypericum, Convolvulus, &c., or perhaps - only occasionally, as in the Pansy, and accidentally, as in the Tradescantia mentioned | above; in others by closing at evening or in diffused sunlight, as is the case with Butter cups and Anagallis, Mesembryanthema, Erythrea, Ke, Stamens. —(a) By the clustering of their anthers around the stigmas, both, of course, ` maturing together. This is by far the commonest method, as, e. g., in Buttercups and with the long stamens of the Crucifere, Stellaria media, Spergula, cleistogamous Oxalis ` Acetosella, small-flowered Leguminose, Epilobium parviflorum and E. tetragonum, Ze (b) By their arching over the pistil. This is conspicuously the case in Potentilla Fra- gariastrum, as also in Galium Aparine, Alisma Plantago, as described above, in whic the anthers are even extrorse. (c) By having the anthers specially situated, wit reference to the stigma, as in Fumaria, cleistogamous Violets, Polygala vulgaris, Oro banche Hedere, Salvia clandestina, and garden species of Calceolaria, &c. 5 Pistil.—(a) By « curvature of style, so as to bring the stigma into contact with the anther-eells. This is effected by its growth under confinement in the unopened bud, e. g. cleistogamous Violets, Lamium amplezicaule, Rhinanthus and Euphrasia, Glaus mari- tima, and abnormally i in Tradescantia erecta. (b) By arrest of growth of the style, as in pale-flowering Pelargonium, non-dimorphie Primule with depressed stamens, Prunella vulgaris. (c) By elongating the style, as in non-dimorphie Primule with elevated sta mens. (d) By stigmas recurving: Malva rotundifolia and other small-flowering species Viola tricolor, var. arvensis, Taraxicum Dens- Leonis, Salvia clandestina, Pinigai lusitanica. (e) By the development of abnormal stigmatiferous tissue, as in the small flowering variety of Viola tricolor, var. arvensis. (f) By the flower being pendulous. Mr, Meehan alludes to this in Ranunculus abortivus, and Solanum nigrum may be added* 9. Conservation of energy is seen in the reduction of the number of Mni and of the quantity of pollen. decore ml "ree are generally subject to this phenomenon. The following speci ! e tis perhaps worth while observing that the old teleological idea of pendulous and erect flowers always having the anthers situated at a higher level than the stigmas, and thereby being specially designed for self-fertilization, i not only far from being the case, but when they are so, it may with tolerable certainty be suspected of being not adapted for self- but for cross-fertilization ; for had Nature intended the pollen to reach the stigma, the anthers would be placed in contact with it, and not at some distance above it, a position which involves the possibility of a loss of pollen by the wind without its "pr the stigma at all, Such Sowas as posso! A are owes, moreover _ icc cote : SELF-FERTILIZATION OF PLANTS. 881 will illustrate the former of the above degradations. Small-flowered Ranunculi have often less than twelve stamens; Myosurus has five; Senebiera didyma has only two; cleistogamous flowers of Viola canina two or three ; Stellaria media often three only, &c. This is in accordance with the fact that only a relatively small amount of pollen is really requisite for fertilization. An interesting experiment of Mr. Darwin's proves this. He placed a very small mass of pollen-grains on one side of the large stigma of Ipomea purpurea, and a great mass of pollen over the whole surface of the stigma of other flowers, and the result was that the flowers fertilized with little pollen yielded rather more capsules and seeds than did those fertilized with an excess (‘ Cross and Self- fertilisation,’ p. 25). That normally intercrossing flowers produce a great superfluity of pollen is well known. Thus Kólreuter found that sixty grains were necessary to fertilize all the ovules of a flower of Hibiscus, while he calculated that 4863 grains were pro- duced by a single flower, or eighty-one times too many (Darwin, l. e. p. 877). The latter says :— ** In order to compensate the loss of pollen in so many ways, the anthers produce a far larger amount than is necessary for the fertilisation of the same flower. ...... and it is still more plainly shown by the astonishingly small quantity produced by cleistogene flowers, which lose none of their pollen, in comparison with that produced by the open flowers borne by the same plants; and yet this small quantity suffices for the fertilisation of all their numerous seeds ” (/, c. p. 376). Mr. Darwin observed that when flowers were artificially self-fertilized for several successive generations, a degeneracy sometimes took place in the anthers and pollen; and he seems to attribute this to the “evil effects” of self-fertilization ; but from the above facts I am inclined to regard it as an illustration of a universal principle in Nature, namely, the preservation of energy wherever possible, and that such cases as appeared under his experiments were adaptations to this principle, as the flowers became habi- tuated to self-fertilization. It is perhaps worthy of note that while both the number of stamens and the quantity of pollen are thus often much reduced in some flowers the capsules of which produce many seeds, yet in others which set but one, as Fumaria, or at least but few seeds, the number of stamens may remain unreduced. Such seems to me to be an additional proof that such flowers are degradations from forms originally adapted to intercrossing when much more pollen was requisite. Hence the present forms are retentions of former ancestral conditions. The following cases will illustrate this :— Daphne Laureola has 8 stamens and 1 seed. Chenopodium has . . . . . 5 » l Medicago, Sis has ficia yx.» 10 D lc Vicia tetrasperma has . . .10 » 4 seeds. Scleranthus perennis has . . 10 i 1 seed. The phenomenon called contabescence by Gartner (Anim. and Pl. under Domest. ii. P. 165) would seem to have its rationale in this ze cup to self-fertilization. Mr. Darwin observes :— “ The anthers are affected at a very early period in the flower-bud, and remain in the same state (with ` one recorded exception) during the life of the plant. The affection cannot be cured by any See = n Temm and is propagated by layers, cuttings, Se, ren — In MG NE SS d a | SECOND SERIES — areir ae pd AMET ptt, : 382 REV. GEORGE HENSLOW ON THE the female organs are seldom affected, or merely become precocious in their development. The cause S of this affection is doubtful, and is different in different cases. ...... The contabescent plants of | Dianthus and Verbascum found wild by Wiegmann grew on a dry and sterile bank." Though there may be more than one cause, I should feel strongly inclined to think that in the majority of instances it is correlated with a lessened degree of dichogamy, ~ if not always with self-fertilization, which Mr. Darwin himself gives as an “ additional | cause." The last sentence above points, however, to the chief probable cause being - nutrition. 10. The relative fertility may equal or surpass that of crossed plants. The fertility of self-fertilized plants, observes Mr. Darwin (7. c. p. 326), * ranges from ` zero to a fertility equalling [or exceeding] that of the crossed flowers ; and of this fact no explanation can be offered." As examples of plants of which crossed and self-fertilized — flowers produced a nearly equal number of seeds are Ipomea purpurea, Gesnera pen- ` dulina, Salvia coccinea, Limnanthes Douglasii, Lobelia fulgens, and Nolana prostrata; — yet “ thé plants raised from the crossed seeds exceeded considerably in height those | raised from the self-fertilized seeds." He rightly considers that ** the average number of | seeds per capsule is a more valuable criterion of fertility than the number of capsules | produced.” By selecting all cases from his Table D, of ** Relative Fertility of Plants of Crossed and Self-fertilised Parentage,” which give the ratio of the number of seeds per ` capsule, I find it to be about 100 : 92 for the crossed to the self-fertilized, a result not far | from equality. The two examples where the self-fertilized surpassed the intercrossed ` were the cleistogamous flowers of Vandellia munmularifolia (100 : 106) and the third | ; generation of Dianthus Caryophyllus (100 : 125). It has already been shown how the ` ratio of the fertility of the self-fertilized plants of Ipomea purpurea and Mimulus luteus ` gradually equalled and then surpassed that of the intercrossed plants. | ll. The fertility or the health of either cultivated and artificially or wild and naturally self-fertilized plants does not necessarily decrease in successive generations. Mr. Darwin says, * There is no evidence at present that the fertility of plants goes — on diminishing in successive self-fertilised generations." He is here alluding to the ` results of his cultivated plants; but it is evidently equally true of all common self- - | fertilizing weeds. Chickweed, Groundsel, Shepherd's-purse, and Poa annua have probably — flourished for centuries just as they do now, and are as vigorous at the present day and : doubtless as fertile as ever they were. A rapid succession of ripening capsules may ` compensate for any want (if any) of a copious supply of seeds in individual capsules. Again, besides having the power to propagate adequately, the self-fertilized may be as healthy as the intercrossed after any number of generations. In support of this latter statement is the fact that the intererossed plants did not exceed the self-fertilized in — height in greater and greater degrees in successive generations; but the reverse was the case; for the mean ratio of the first three generations of the intercrossed to the | self-fertilized plants of Zpomea was 100:74°3; that of the second three as 3 100: IT 6 | that of the third three as 100 : 81-6. The ratios thus gradi ally a ate 5 80 that instead of the xatio noone a smaller fraction, etm u SELF-FERTILIZATION OF PLANTS. 383 plants deteriorated, it shows that ‘the difference between self-fertilization and inter- crossing was gradually diminishing. Such an effect is due, according to Mr. Darwin, to the fact that the intercrossed plants became by close in-breeding more and more ‘intimately related ; but we must not forget that self-fertilization is a still closer method of in-breeding. Therefore if that be a cause of deterioration to the intercrossed, it . should, à fortiori, be so to the self-fertilized. But it cannot be; for if it were, the ratio should either remain constant, or else vary progressively in favour of the intercrossed, as the self-fertilized should deteriorate faster than the intercrossed. On the other hand, if it do not so vary, and the intercrossed be really deteriorated, there is but one inter- pretation, viz. that the self-fertilized have not deteriorated in the same ratio as the former. Such want of any correspondence between ratios of heights and successive generations occurred also with others of Mr. Darwin’s experiments; for thus was it with Petunia violacea and Nicotiana tabacum. The difference between the heights of the intercrossed and of the self-fertilized plants of the first named, arranged in a decreasing order, would be as follows :—4th, 5th, 2nd, 4th, 1st, 3rd years. In the last case, the ratio for the intercrossed to the self-fertilized was 100 : 131, where the difference has again increased, but is now in favour of the self-fertilized. 12. The fertility of cultivated self-fertilized plants may increase in successive generations. Mr. Darwin gives some interesting illustrations of this fact (l: c. p. 112). Thus Eschscholtzia californica was self-sterile in Brazil, but in the first year's cultivation in England the ratio of the yield of seeds between the intercrossed and self-fertilized plants was 100: 15. Eight flowers on these self-fertilized plants (7. e. grandchildren of the plants which grew in Brazil) were again fertilized with pollen from the same plant, and produced five capsules with an average of 27:4 seeds; but in the previous year the average was only 12 seeds per capsule; hence the fertility of the self-fer- tilized plants was thus increased to more than double that of the first year. Dianthus Caryophyllus is naturally so strongly proterandrous, that Mr. Darwin was obliged often to use pollen from a different flower on the same plant for artificially self- fertilizing it. This species appeared to adapt itself to self-fertilization with great readiness by becoming homogamous. The proportion of seeds of parent plants inter- crossed and of self-fertilized plants was in the ratio of 100 : 92; and when the self- fertilized offspring was crossed by a new stock, the proportion of the latter to the again self-fertilized was in the ratio of 100 : 123. Similarly the third generation of the inter- crossed subsequently self-fertilized was in proportion to the fourth self-fertilized gene- ration as 100 : 125. The above results, taken from Mr. Darwin’s work, clearly show a steady increase of self-fertilizing power, which was acquired by the loss of dichogamy, the stamens and pistil maturing together. Looe Free from EV the self-fertilized planis may equal the intercrossed (1) as | seedlings, (2) when adult and planted in open ground. Bur. hist apps iom Ms Darwin's observations. E. = 210 Tuto m s 384 REV. GEORGE HENSLOW ON THE the general rule appears to be that as long as the pairs of seedlings planted in the same pot in his experiments were free from competition there was not much difference in their growth. As soon, however, as they began to compete, the crossed plants showed their superior vigour, and so checked the growth of their self-fertilized competitors. They would inevitably continue to do so by abstracting the nourishment which the others would otherwise have secured, and so would ultimately, as was so generally the case, beat the self-fertilized plants (7. c.[p. 285). Secondly, in some of Mr. Darwin’s experiments he grew the plants much more crowded in some pots than in others. In these latter the ratio of the heights of the intercrossed to the self-fertilized were almost invariably nearer unity than that deduced from the crowded plants. Lastly, when the plants were grown in open ground, and entirely free from competition, there was often, as Mr. Darwin states, but little difference in their growth *. Mr. Darwin planted pairs in the same pot in order to imitate nature ; but the struggling between individuals of the same species is not generally quite so common as that between. different genera; and though his experiments bring out well enough the superiority of the crossed to the self-fertilized under competition, they fail to show what would be their relative values when competing with other genera of other orders. It is a well- known fact that plants of different orders, and presumably of different constitutions, not requiring identically the same food, can live and flourish together or in succession when plants of the same species would fail to do so. This is seen in the flora of oceanic islands, in which the number of orders is large in comparison with the number of genera, and the latter large as compared with the number of species. It is also the basis of the principle. upon which the rotation of crops is founded; but for Mr. Darwin’s object perhaps the method adopted was the best. The question, however, arises, Why is it, if the intercrossed are so superior in vigour to the self-fertilized, that such a difference is noć so great when they are free from com- petition? If there were any evil or injurious effects, as Mr. Darwin supposes, to follow from self-fertilization, it is difficult to understand why the ratio does not keep constant under all circumstances; for an ample space and a good soil being common to both, one would, à priori, infer that the intercrossed would be benefited thereby just as much as, if not even in a higher degree than, the self-fertilized. This, however, does not appear to be the case. A few examples from Mr. Darwin’s experiments will illus- trate this. T . In several of his tables he has written ** plants crowded” to certain pots; and I find by calculating the ratios of these, as well as the ratios of all the rest not so crowded, that they are, with a solitary exception, nearer unity in the less crowded. Thus, omittin| j decimals :— ey Ipomea purpurea, Table-X., not crowded. sT sar sor «..s + 4.100: 90. b h crowded < recta i fap oc cx MN A EM 2 purpurea crossed on same plant, Table XII., not crowded. 100 : 111 » e e crowded. . 100: 97 Ditto crossed NEC Colchester enge? Table XIL, not crowded . 100: 83 5. ean. » . erowd s d ; “+ Animals and Plants undor Domest p. 128 SELF-FERTILIZATION OF PLANTS. | 385 Mimulus luteus, Table XIX., not crowded, . . . . . . . . 100: 109 ap e ctowded i ahaha of wooed HEUO £4148 Digitalie purpurea, Table XXIV., not crowded . . . . . . . 100: 94 » » crowded, nis os dioi asw i 1008: DO So also when pairs of plants grown in pots are contrasted with similar pairs. in open ground very similar results occur :— Reseda lutea in open ground `, . . . . . . . .. 100 e Së M e EE EE EE Obi Ss KR. odorata in open;ground.. bis | uiuos awe: och ENEE 0 s» mM SOBEL uo POGUE ROE qnt DOqu100vv ag Dianthus Caryophyllus in open g den: oles (RT ep (rr 260454 2120902 0:808 o mpow ““pawli vd. EE . 100: 58* Lastly, a like difference resulted in the ae of plants of Petunia violacea when crossed with a new stock; for the ratio of the weights of what Mr. Darwin called the ** Westerham-cerossed " to that of the self-fertilized was in pots as 100 : 22, but in open ground as 100 : 53, and to that of the normally intercrossed in pots as 100 : 101, while in open ground it was as 100 : 146 (Z. c. pp. 200, 201). Nicotiana tabacum illustrates the converse. This is a highly self-fertile plant. The ratio of the weight of seeds of the intercrossed to the self-fertilized was as 100 : 150, and that of their heights when young as 100: 189, but when adult as 100: 178. When, however, they were greatly crowded the ratio became one of equality, or 100 : 100. . Hence, whatever be the standard of measurement, the results appear to furnish the decided rule that self-fertilized plants regain their equivalency with or even an ascend- ancy over the intercrossed as soon as they are freed from competition ; consequently I do not see why Mr. Darwin should speak thus of Nicotiana :—“ It is a strange fact that the self-fertilised plants, which were subjected to very severe competition with the crossed, had on two occasions no advantage over them." I take the explanation to be this :—Tobacco is evidently a vigorous self-fertilizer, nevertheless crossing did some good ; but that good was not apparent, except under certain conditions, i. e. severe com- petition, when the introduction of some new constitutional elements had the opportunity of evincing their power, whereas the innate vigour of the self-fertilized Nicotiana com- pletely eclipsed it when free from competition. Finally, there are apparently but two alternatives to appeal to, in order to account for the fact that intercrossed plants are not so greatly superior to the self-fertilized when planted in open ground as when in competition in pots, viz. either the intercrossed plants become deteriorated on being planted in open ground, which would seem to be absurd, or else the self-fertilized must regain or acquire vigour in a relatively greater degree than do the intererossed, and thus would seem to evince what might be called a greater * elasticity " of growth than their intercrossed competitors. | 14. Some self-fertilized plants derived no benefit from a cross with another plant of the same stock, nor even from a cross with a plant of a distinct stock. . With regard to the first statement, it applies to plants long cultivated e coe eae | Dm o M DOM Darwin, see ‘ Cross and Self-fertilisation, p. 288.- eed 386 REV. GEORGE HENSLOW ON THE tion, and the two cases which are recorded by Mr. Darwin are Pisum sativum and Canna Warscewiczi. With regard to the Garden Pea, the papilionaceous corolla is obviously an adaptation to insect agency by which intercrossing is secured; but the Pea in this country is rarely, if ever, crossed, so that cultivated varieties keep true even when grown together. Some of Mr. Knight’s varieties kept true for more than sixty years ; but “ their glory is now departed,” says Mr. Darwin*. As to the effects of intercrossing, the latter remarks :— * Owing to the varieties having been self-fertilised for many generations, and to their having been subjected in each generation to nearly the same conditions, I did not expect that a cross between two such plants would benefit the offspring ; and so it proved on trial” (l. c. p. 162). The ratio of the heights were as 100 : 115, * so that the crossed plants, far from beating the self-fertilised, were completely beaten by them." : Canna Warscewiczi is also highly self-fertile, the pollen being shed before the flowers expand. Intercrossed plants showed no advantage over the self-fertilized ; the ratio of the heights of the intercrossed to self-fertilized for three generations taken together was as 100: 101, and Mr. Darwin observes (/. c. p. 233) :— * We may therefore conclude that the two lots possessed equal powers of growth; aii this I believe to be the result of long-continued self-fertilisation, together with exposure to similar conditions in each generation, so that all the individuals had acquired a closely similar constitution ? +. In speaking of Pisum (l. c. p. 264) he says, “as the plants have been long cultivatell | under nearly similar conditions, we can understand why a cross between two individuals of the same variety does not do the least good to the offspring either in height or fer- tility; and he compares Pisum with his own cultivated examples of Mimulus and * Hero.” But the words I have italicized are an assumption which will not hold good; for he does not show that the soil &c. was any degree more nearly exactly the same for Pisum than for any other plants of which the intercrossed beat the self-fertilized. But the matter or question cannot rest there. One still asks why it is so. A priori one would be led to suppose that a plant habitually self-fertilized would, on the contrary, immediately be benefited by even the slightest infusion of new blood by a cross of the same stock, and much more by a cross from a distinct stock or different varieties. In the last case it is true; for by crossing varieties of Peas the ratios became at once ` 100 : 60-70. But it is not clear why two individuals of the same variety do not benefit ` by a cross when they have been habitually self-fertilized. To say it is because they have ` * Why it is gone he does not explain ; and one would like to know whether it was from any gradually deteriorating - effect of self-fertilization, as he seems to imply (though the sixty years certainly afford strong presumption mE such being the case), or whether it was not that other marketable varieties had superseded them. 4 t Mr. Darwin says that the opinion of some persons that this flower is invariably self-fertilized is “ am extra- - ordinary conclusion, for it implies that a great amount of pollen is produced for no purpose." May not the real ` explanation be, that while the flower was formerly adapted for insect-fertilization, as indicated by its bright ` perianth, abundance of pollen, &c., in the absence of the proper (native) insects it has become self-fertilizing in ` this country, but has retained its customary habit of developing more pollen than is actually required for self- fertilization? Fumaria officinalis sets but one seed, yet the amount of pollen is relatively very great. Similarly ` Chenopodium has retained its five fertile anthers but yields only a single seed. Many other cases might be adduced, in addition to those given above, to show an apparently great tit NIA: Sion Mu dert the ono hand, and the mambas E atida ai we es = SELF-FERTILIZATION OF PLANTS. 387 been grown under “ nearly similar conditions" does not explain it; for Ipomea had been so grown; and in making his experiments, Mr. Darwin was most careful to make the soil, moisture, light, &c. as absolutely identical as possible for his pairs of plants, so that the benefit of crossing should be solely confined to the effect of the sexual process. On the other hand, Canna has been always grown in pots; and when no object has been in view beyond mere cultivation, it is a small chance that the soil should have been so identically the same in every case. The cause cannot then be looked for solely, if at all, in environment; it must be in the fact of habitual self-fertilization. It is perhaps worth observing that, if a single instance may be trusted, the mean ratio of all the intercrossed to the self-fertilized of Mr. Darwin's experiments is exactly the same ratio of the self- fertilized to the intercrossed of Pisum sativum, or 100 : 87. I do not see, therefore, how we can avoid the conclusion that, if a plant is habitually self-fertilizing, it can amply retain its numbers by propagation, and in no way deterio- rates in consequence of the process, though, on the other hand, it may derive immediate and great benefit from a cross with a new variety or stock. However, that a plant may derive no benefit from such an intercross was proved by Mr. Darwin in the case of ** Hero," the remarkable self-fertilizing individual of Ipomea, of which Mr. Darwin says (l. c. p. 51):— * No advantage, as far as we can judge, was derived from intercrossing two of the grandchildren of Hero, any more than when two of the children were crossed. It appears, therefore, that Hero and its descendants have varied from the common type, not only in acquiring great power of growth and increased fertility when subjected to self-fertilisation, but in not profiting from a cross with a distinct stock ; and this latter fact, if trustworthy, is a unique case, as far as I have observed in all my experiments." Mr. Darwin calls this a “ unique ease; " but in Table C, which treats of ratios derived from fresh stocks, offspring of Eschscholizia californica, being from Brazil, crossed by an English stock, the self-fertilized plant beat the crossed in height and weight, but only fell short of it in fertility. But as this plant was self-sterile in Brazil, yet acquired a power of self-fertility in England in the ratio of 100:15 in the first year, which was raised to 100:40 in the second generation (Table C), this seems to show that even in fertility the self-fertilized plants were rapidly gaining upon those crossed with a new stock, and would equal them, at the same rate of increase, in very few generations. Such exceptional cases as the above clearly prove that it can by no means be regarded as an absolute fact that intercrossing plants of the same stock, or crossing plants of different stocks, does necessarily benefit them. The process of crossing is, as Mr. Darwin clearly proves, solely a means to an end, that end being the introduction of new consti- tutional elements; and if a cross cannot do this, then the plant, so far from being bene- fited, is as much “ deteriorated " as a plant which is habitually intercrossed is “ impaired ” by self-fertilization. ** Deterioration” or “injuriousness” are only relative terms; for if a plant like Pisum sativum habitually fertilizes itself, and its average height be represented by 100, and if, by intercrossing different plants, that standard instantly becomes lowered to 87, we are as much justified in saying that the plant was “ deterio- “rated” by the cross as, on the other hand, that Zpomea is “ benefited " to the same v extent z2 aa ob Ces is true for the 4 one : must be logically true for the other. 388 REV. GEORGE HENSLOW ON THE So, conversely, if self-fertilization is “injurious " for Ipomea, intercrossing is " injurious” for Pisum. We may, moreover, go further, and observe that a cross may actually infect a plant and bring about deleterious results; for Berkeley, while observing that “the main object to which the whole system of the organs in any individual plant tends is the reproduction of the species by means of seed," adds :—** The embryo partakes, not only of the nature of the plant which bears the seed, but, if the impregnation have taken place from the pollen of some neighbouring plant of the same species, of that of the male parent also. The seed, therefore, even of a healthy, much less of an unhealthy, plant will not neces- sarily produce a healthy offspring " *. Whichever way, however, we use the term *'injuriousness," it seems to be mis- leading, for it implies some positive evil instead of a mere relative degree of vigour; so I would prefer to suggest the term “ negative ” for both cases where no increased powers are acquired by the plant. The facts recorded by Mr. Darwin, and quoted in this section, appear to me to have great significance, and might be expressed by such an aphorism as permanently self-fertilized plants refuse to be intercrossed. Mr. Darwin, as we have seen, attributed this to the fact of their having been so long cultivated by self-fertilization under nearly the same conditions; but this would not seem to be any real explanation; and I find that Mr. Meehan has arrived at the same conclusion; for he says in a letter to me :— f 33 “ There is another matter which has occurred to me, but which I have not yet had time to work out to a positive conclusion ; those species which seed the most readily are self-fertilizers ; and they seem to abominate crossing. Disemma aurantiaca, an Australian species of Passifloracee, seeds with extreme freedom. I took care, as I thought, to keep all pollen from it but that of P, cerulea: I felt sure of a hybrid; but the result was only Disemma. At that time I thought I must have made a mistake, but I have since had the same experience with Primula japonica and P. involucrata; and my friend Parkman with Lilium auratum . . . . Why should “these cases only occur in instances where the plant seeds abundantly by its own pollen ? If cross-fertilization is a benefit, we should suppose that those which seeded the most freely would be in the most need of it, and show a greater See to receive outside aid when they could get it.” i Such is my view also, as ST above. 15. Self-fertilized plants are perfectly healthy. Self-fertilized plants are, when in natural conditions, as healthy as intercrossed plan though exceptional cases may occur under cultivation. With regard to the first class, have but to remind the reader of the names of a few of the commonest and most troub je some “ weeds,” and he will doubtless have experienced their vigour and healthiness, suc as Groundsel, Sowthistle, Chickweed, Shepherd’s-purse, Knotweed, Buttereups, Solanum nigrum, species of Cerastium, Poa annua, Ee, Ee, all of which are highly and habitually self-fertile, and of which many propagate themselves with extraordinary rapidity, and are certainly far from warranting any belief in the injuriousness of self-fertilization. If we turn to Mr. Darwin’s pa several cases occurred where the self fertilized i * Yop: Ib? axir, $ 100 and aa $ 128, in‘ Gan. Cima? 1854 pp gon 3 SELF-FERTILIZATION OF PLANTS. 889 plants exhibited astonishing vigour. Thus he speaks of the white or pale-coloured variety of Mimulus luteus (l.c. p. 80) :— “ From the tallness of this variety, the self-fertilised plants exceeded the crossed plants in height in all the generations from the fifth to the seventh inclusive; and no doubt would have done so in the later generations, had they been grown in competition with one another.’ [My italics.] Again, of Hero, the sixth self-fertilized generation of Ipomea, he says :— “If the seeds produced by Hero had been as greatly in excess of those produced by the other plants as was the case with Mimulus, and if all the seeds had been mingled together, the offspring of Hero would have increased to the entire exclusion of the ordinary plants in the later self-fertilised generations, and from naturally growing taller would have exceeded the crossed plants in height in each succeeding generation." I quote these passages, for they show that, according to Mr. Darwin's estimation, these self-fertilized varieties would have been superior to others under competition, a fact which, as a rule, is opposed to the results of his experiments, as he, indeed, observes :— “ Thus we have a complete reversal of what occurred in the previous generation." The interpretation appears to be that they found certain ingredients in the soil which suited and strengthened their constitutions, and so they gained independently what was usually only to be acquired by means of a cross with a distinct stock; which seems to prove that if a self-fertilizing plant can secure new constitutional elements from a fresh soil, it then may show as much vigour as, or more than, one which may be intererossed with another plant growing under the same conditions. On the other hand, Mr. Darwin mentions some instances where the self-fertilized appeared to him to have actually suffered from self-fertilization. This, he thinks, was shown indirectly by the intercrossed plants withstanding certain adverse conditions, while the self-fertilized failed to do so; thus he says (l. c. p. 289), ** The crossed plants always withstood the injurious effects of being suddenly removed into the open air after having been kept in the greenhouse better than did the self-fertilised. On several occa- sions they also resisted much better cold and intemperate weather . . . . The offspring of plants of the eighth self-fertilised generation of Mimulus, crossed by a fresh stock, survived a frost which killed every single self-fertilised and intercrossed plant of the same old stock. Nearly the same result followed with some crossed and self-fertilised plants of Viola tricolor .. ... I have met with only one exception to the rule of crossed plants being hardier than the self-fertilised [that of EscAscholtzia]. . ... Independently of any external cause which could be detected, the self-fertilised plants were more liable to premature death than were the crossed.” A few further remarks will be found in l. c. pp. 290-1*. But allowing for these facts, which occurred under cultivation, self-fertili- zation, as carried on by Nature, does not support them. This, for example, is shown by such of our wild flowers as blossom in mild winters being (after eliminating any anemo- - philous cases) probably, without exception, self-fertilized. They can and do ripen their seeds in profusion in January as well as in July, and in that respect show a vast superiority over those plants which have to depend upon the visits of insects to set seeds at all. They iy EE ET slag Fertilization of Plants," in the ‘Gardeners’ Chronicle,’ —5 RA uno ee T SE i os 390 REV. GEORGE HENSLOW ON THE are for the most part humble and insignificant weeds, it is true, and are probably de- graded forms of conspicuous and formerly intererossed plants; but, as they exist now, there are no perceptibly evil effects or injuriousness (using these words in their ordinary sense) at all attributable to self-fertilization. On the contrary, if we regard, with Mr, Darwin, propagation as the one end and aim of plant life, then self-fertilization can only be regarded as an inestimable boon to the plant. That self-fertilization is in some way injurious appears to be the view of Sir J. D. Hooker; for he says, in alluding to different methods of fertilization :— ** Tn all these instances the double object of Nature may be traced; for self-impregnation (or * breed- ing in’), while securing identity of form in the offspring, and hence hereditary permanence, at the same — — time tends to weakness of constitution, and hence to degeneracy and extinction” *, He does not state on what grounds this opinion is based. 16. Self-fertilized plants may be absolutely much more productive than flowers dependent upon insects. The latter may fail in particular seasons, and may be entirely wanting if a plant migrate beyond the range of its habitual visitor. Under either of these alternatives the self-fertilizing plants would prove to be the best off. It is well known, for example, ` how clover-seed is dependent upon the visits of Humble-bees, and how a field may ` largely fail in inclement weather if these insects do not visit it. Moreover, humble-bees ` often do more harm than good, by perforating the tubes of flowers instead of entering ` the corolla. Plants habitually crossed in Hil native country may quite fail to secure any insects i a to accomplish the process elsewhere, as the Scarlet Runner in Nicaragua; yet if they K acquire self-fertilizing powers they will at once become independent of them. Such a appears to have been the case in this country under cultivation with Pisum sativum, ` Phaseolus multiflorus, and Canna Warscewiczi. If they do not do so, and no insects ` a visit them in their new abode, they must inevitably die out. On the other hand, there ` are many conspicuous and (but probably more) inconspicuous flowers which, being both independent of insects and highly self-fertile, are quite able to maintain their existence — ` and propagate freely wherever they may happen to be located. Now the above appears — to be actually the case, as far as negative evidence on the one hand, i. e. the absence of our intercrossing plants, and positive evidence on the other, i. e. the presence of our self- fertilizing plants in foreign countries, tends to prove it. A large number of European plants are very widely dispersed, not only ione am the northern hemisphere, but in the southern as well. And if we, for convenience, limit ` ` our observations to the distribution of British plants, what we find is, that they are ` certainly mostly, and probably all, self-fertilizing or wind-fertilizing plants. At all events, ` with rare exceptions, they are all inconspicuous plants; and experience leads me to con- : clude, I think justly, that all such are habitually self-fertilized or anemophilous. It is a _ remarkable fact that wherever there are two or more species to a genus, one or more of : which are conspicuous, and the other or others inconspicuous, it is a rule that the latter only have succeeded in establishing themselyes in foreign countries. | SCH Ce EE any otis Tie zd x. | cp Kan Ge SELF-FERTILIZATION OF PLANTS. 391 It is immaterial to consider how they got into far distant countries; for it is not the method of transportation, but the reason for their establishment that I am considering. Many weeds invariably accompany Europeans on migration; but many others are found located in places which do not warrant the idea that man has either intentionally or unintentionally imported them; and they must now be regarded as natives. Several are water-plants, which may be regarded as perhaps the easiest to transport, as aquatic birds will readily disperse them; but the question at issue is, although they may be transported with ease, why do they maintain their ground when in competition with the native indigenous plants? The question is equally pertinent for aquatic as well as terrestrial plants, as competition occurs in both cases. The reply I would give is, that it is the self- or wind-fertilizing plants which alone can maintain their ground, because they are independent of insects, and that having been once introduced into new climatic conditions they acquire new and often great constitutional vigour, and so beat their native rivals, who, having long since arrived at a standard of equilibrium, cannot acquire any new vigour at all proportionate to that infused into the new comers, which thus overcome them in every way. 17. The world-wide distribution of self-fertilizing British plants. In the following enumeration I will limit myself, for brevity sake, to such species only as are recorded as growing in at least four distinct localities, i. e. as far as my researches into “ floras” have enabled me to discover. Many other species are in one, two, or even three countries; but the point I particularly wish to bring out is, that whenever British plants are found very widely dispersed they are probably almost always self-fertilizing or else anemophilous plants. How they have become located where they now are is a question with which I am not now concerned; but I only wish to show that such plants, having somehow reached far distant localities, have succeeded in establishing themselves ; and the method of doing so is, I believe, mainly because they were self-fertilizing and consequently independent of insects *. | RANUNCULACEX. Ranunculus hydrocharis: forms of this species are recorded from S. Austr., Tas., S. Af., S. Am., N.E. Af, N.E. Asia, Cal. R. sceleratus T, Trop. As., S. Af., Hong., Jap., N.E. Asia. Crucirerm. Brassica oleracea, Jap., Mad., N.Z.; B. rapa, Trop. As., Jap., N. Z., And. | Cardamine hirsuta, Chili, S. Austr., A. & C., N. Z., F. & F., Tas, 8. Af, Trop. Asia, N.E. Af., Hong., Mad., N.E. Asia, Kam. [For the sake of comparison, I will add C. pratensis (doubtfully according to Hooker), Tas.; C. impatiens, Temp. Asia and Jap.) * In the list which follows I have used the following abbreviations:—S, Austr. (South Australia); Tas. (Tasmania); S. Af. (South Africa); S. Am. (South America); N.E. Af. (i. e. North-east Africa); Trop. As. (Tropical Asia, south of Himalayas); A. & C. (Auckland and Campbell's Islands) ; N. Z. (New Zealand); F. & F. (Falkland and Fuegia); Hong. (Hongkong); Mad. (Madeira); And. (Andaman Isles); Jap. (Japan); Kerg. (Kerguelen Isle); Soe. I. (So- ciety Isles); Norf. I. (Norfolk Isle); Cal. (California); Kam. (Kamtschatka). _ t Many of the forms of our British species found in foreign countries are not identically the same as ours, but varieties of them. This is no more than might be expected (antea, p. 389); still it does not affect the question of p COO NE ja o 392 REV. GEORGE HENSLOW ON THE Nasturtium officinale, N. Z., N.E. Af., Jap., Mad., Jam., S. Am. ; N. palustre, S. Austr. - S; Am., N.E. Af., Jap., N. Z., N.E. Asia. Capsella Bursa-Pastoris, Chili, N. Z., Trop. Asia, N.E. Af., S. Austr., Hong., Jap. Kam. Sisymbrium officinale, Chili, S. Austr., N. Z., Mad. Lepidium ruderale, S. Austr., Tas., N. Z., Trop. Asia, S. Af. CARYOPHYLLACER. Cerastium vulgatum or viscosum, F. & F., N. Z., S. Af., S. Austr. Trop. Asia, Mad., Jap., N.E. Af., Marion I., Tristan d'Acunha. Stellaria media, A. & C., Kerg., N. Z., Trop. Asia, Mad., S. Af., S. Am., S. Austr Hong. Jap. S. uliginosa, Hong., Jap., Mad: Morocco. TA. Holostea, only W. Asia.] Arenaria serpyllifolia, Trop. Asia, N.E. Af., Jap., N. Z. Spergularia rubra, Chili, S. Austr., Tas., N. Z., N.E. Af. PortuLacace®. Montia fontana, A. & C., F. & F., Tas, N. Z., Kerg., S. Am, Marion I., Jap., N.E. Asia, N. Am., Andes. MarvacEms. Malva rotundifolia, N. Z., N.E. AT, Soc. I., Sand. I., Jap., Cal. GrnANIACEX. Of the small-flowered annual species several are found in the southern ` hemisphere ; but the larger-flowered proterandrous perennials are mostly W. and N. Asia. - Erodium cicutarium, Chili, S. Austr., N. Z., Cal., N.E. Af. Oxalis corniculata, S. Austr., N.E. Af., Soc. I., And. I., Hong., N. Z., Cal. LEGUMINOSA. Medicago lupulina, N. Z., N.E. Af., Hong., Jap. Medicago denticulata, Chili, N.E. Af., Jap., N. Z., Cal. Lotus corniculatus, S. Austr., Tasm., N.E. Af., N. Z. Vicia sativa, N.E. Af., Jap., Mad., N. Z. ROosACEJE. Potentilla anserina, F. & F., Tas., N.Z., S. Am., Jap., N.E. Asia, Kam, Cal. Geum urbanum, Tasm., 8. Am., N. Z., N.E. Asia, Fuegia. LyruRACEX. Lythrum Salicaria, S. Austr., Tasm., S. Af., S. Am., Jap., N.E. Asia, Central Asia, N. Am. (L. hyssopifolia, S. Aust., Tasm., N.Z] ` ONAGRACEÆ. Epilobium tetragonum, S. Aust., P. & F., Tasm., N. Z., S. Am., Jap. N.E. Asia, Cal. HaroRAGrACEX. Callitriche verna, S. Austr., A. & C., F. & E., Tasm., N. Z., S. Ama N.E. Af., Marion I. UMBELLIFERE. >- T. s sri Di serie Bang feuctiligation eet «samen EE Chlamydospores. | The Speer. and d clam dospori FER bodies See t to, be more nal FRUCTIFICATION OF CHOANEPHORA. 419 allied to one another than to the conidia. They are new and independent cells formed within the tubular system of the parent plant; but the conidia are merely isolated portions of that system, being, in fact, merely the tips of the terminal filaments of the aerial portion of the plant. The distinction, moreover, is not merely an anatomical one; the difference in the nature of the conditions favouring the development of the various forms of fructification indicates a physiological distinction also, The conidial fructifica- tion is the form characteristic of active nutrition and vegetative growth. Given a very rich nutritive medium and fully developed normal conidia, a luxuriant development of mycelium occurs, followed, sooner or later, by an abundance of conidial fructification. With diminishing nutrition there is progressively poorer mycelial development and less- developed conidial fructification. When this degeneration has reached its utmost limit, when the conidial fructification is reduced to its poorest and simplest type, the sporangia begin to make their appearance; and when conditions of nutrition are too greatly lowered even to allow of this, we find the chlamydosporic fructification providing for the preservation and diffusion of the plant. The fact that the occurrence of various forms of fructification may be determined by conditions of nutrition should be constantly borne in mind in the study of organisms with polymorphic reproductive apparatus. The observation of the fact isnot a new one; but the present instance appears to be one of the clearest demonstrations of it which has yet been afforded. It is not, however, merely in respect of this, although no doubt closely associated with it, that the phenomena exhibited by Choanephora deserve atten- tion. They appear, in addition, to be capable of affording a possible explanation of certain conflicting conclusions which have been arrived at as the result of recent observations on the Mucorini conducted by highly competent authorities. I allude especially to those of MM. Brefeld, Van Tieghem, and Le Monnier. The former author has been led by the result of his observations to divide the Mucorini, or Zygomycetes as he prefers to style them, into two subfamilies, one of which he distinguishes as characterized by sporangial, the other by conidial asexual fructification. MM. Van Tieghem and Le Monnier, on the other hand, deny that true conidial fructification ever occurs in the Mucorini, and affirm that in those cases where it has been stated to occur, the supposed conidia were either monosporous sporangia or aerial chlamydospores. The phenomena presented by Choanephora show both these conclusions to be incor- rect; for, on the one hand, they demonstrate that true conidia really do occur in the Mucorini, and, on the other, that true conidia may occur in the same species with sporangia. Whilst thus proving the incorrectness of the conclusions of the distinguished observers just named, the present observations afford a means of reconciling the appa- , rently conflicting results upon which these conclusions, are founded. They do so by showing that although the observations of MM. Brefeld, Van Tieghem, and Le Monnier may have been correct, their conclusions are not so, merely because of the assumption that all the phenomena presented by the species under observation had presented them- Selves to each observer. M. Brefeld, because he never encountered any but a conidial form of asexual fructification in Chetocladium and Piptocephalis, assumes that sporangia * "never are produced in these genera ; whilst MM. Van Tieghem and Le Monnier, believing ` 420 DR. D. D. CUNNINGHAM ON THE CONIDIAL that the forms of asexual fructification observed by them were either sporangial or chlamydosporic, at once conclude that conidial forms never can occur. In reference to Brefeld’s observations, it is of importance to note that he knew Chetocladium and Piptocephalis, his conidial genera, only as parasitic organisms, that — is, only under conditions in which their nutrition was at its highest,—conditions which ` the phenomena of Choanephora justify us in regarding as those securing the production ` of a luxuriant mycelium, occasionally of the sexual fructification, and constantly of the conidial form of asexual fructification. MM. Van Tieghem and Le Monnier, however, studied one of these genera, Cheto- — cladium, under different circumstances. They observed it not only as a parasite on other Mucorini, but as growing independently and deriving its nourishment directly — from the surrounding media. In reference to the conclusions of these observers regarding the nature of the fructification in this genus and in Piptocephalis, three possi- — bilities suggest themselves in endeavouring to reconcile their statements with those of — Brefeld. It may be (1) that in both genera they only met with true sporangial fructi- fication; or that (2) they encountered both sporangial and conidial forms; or possibly even (3) that in certain cases they met with conidial forms alone. It may appear to be presumptuous to suggest the latter two possibilities; but when carefully studied in connexion with the present series of observations, their interpretations of many of the phenomena actually observed appear to have been in some degree influenced by a fore- gone conclusion of the impossibility of the occurrence of conidia. In reference to the possibility of the form of fructification observed by MM. Van Tieghem and Le Monnier | having been of a different nature from that obtained by Brefeld in his cultivations, it may be noted that the former authors give an account of the phenomena attending germination in Chetocladium differing materially from that furnished by Brefeld. They affirm that germination occurs with hardly any antecedent enlargement of the spores ; Brefeld describes the conidia as enlarging considerably prior to germinating. The grounds on which there is reason to suspect that the interpretation of phenomena given by MM. Van Tieghem and Le Monnier has occasionally been somewhat influ- enced by a foregone conclusion, and by a desire to avoid the complication incident on the recognition of two subfamilies of Mucorini, as proposed by Brefeld, must be considered somewhat in detail in order to be appreciated. In reference to Chetocladium, these writers allow that the bodies which they regard as sporangia invariably contain only a single spore. Here the question at once arises, How are monosporous sporangia to be distinguished from conidia? MM. Van Tieghem and Le Monnier apparently distinguish them by the fact that in germinating, or when exposed to pressure, the contents of the cell, if of a sporangic nature, escape, partially or completely, as a homogeneous spore-like body. But such a phenomenon may surely occur in conidia‘ or spores in which there has been a distinct formation of a firm exospore including an endosporic sac. If one look for examples of such an occurrence among kibori; although cases of complete escape of the endospore are perhaps unknown, there is no great difficulty in obtaining examples of partial escape. Tuniak to MM. Yon Tieghem and vs Monnier’ s own account of the germination of the spora | es of FRUCTIFICATION OF CHOANEPHORA. 421 Phycomyces, we find it stated :—** Si la spore jeune n'avait pas acquis de double contour, il ny a pas d'exospore percée par le tube, et le contour externe de la spore est seule- ment plus noir que celui du filament qui en procède ; mais si la membrane s'était déjà séparée du protoplasme par un contour interne, la spore, en se dilatant, brise un exospore, qui se décolle souvent sur tout le pourtour et continue à l'envelopper en partie." If we compare this description and the illustration accompanying it with those referring to the phenomena occurring in the case of the so-called sporangia of Chetocladium, it is evident that the difference in the two cases is one of degree and not of kind. In regard to the phenomena presented by Piptocephalis, it is evident that further observation is yet wanting. To those not already convinced of the necessary absence of eonidia, neither the descriptions nor illustrations furnished by M.M. Van Tieghem and Le Monnier afford satisfactory evidence of the sporangial nature of the fructification. That the sporoid cells are not formed successively by continuous budding, like the conidia of Penicillium and Aspergillus, but by the segmentation of previously continuous rod-like bodies, all observers are agreed; but that this affords definite proof of the sporangial nature of the fructification is certainly not a fair conclusion. A very similar process occurs in the filaments of Oidium lactis and in the ascal spores of certain Spheria, where the filaments or cells breaking up into a series of conidia or secondary spores, as the case may be, can in no sense be regarded as sporangia. There is another difficulty, moreover, in the way of accepting the interpretation proposed by MM. Van Tieghem and Le Monnier. This lies in the fact that they affirm that the spores are held together by a substance similar to that occupying the interspaces between the spores in the spo- rangia of other Mucorini. It is somewhat difficult to realize in what the similarity lies, seeing that in other Mucorini the special function of the material appears to be to ensure the separation and diffusion of the spores due to its property of swelling up in contact with water, whereas here it is supposed to hold them together although immersed in a globule of water. The same tendency to insist on the impossibility of the occurrence of conidial fructi- fication appears more or less in the account given by the same authors of the phenomena observed by them in JMortierella and Syncephalis. The difficulties in the way of accepting their conclusions regarding the nature of the fructification in the latter genus are precisely those alluded to in connexion with Piptocephalis. With regard to both genera it may, moreover, be remarked, that even if the observers be correct in denying the conidial nature of the fructification, it is not clear on what grounds they distinguish it as sporangial rather than chlamydosporous. As to Mortierella, the doubtful inter- pretation does not refer to sporangia, but to certain bodies which they regard as aerial chlamydospores. No doubt their observations regarding the occasional formation of such bodies beneath the termination of the parent filament would, if confirmed, be practically conclusive. Confirmation is, however, certainly required, as the whole character and history of the development is otherwise strongly suggestive of the conidial nature of the bodies in question. It is in this respect very significant that the aerial . . ehlamydospores are described as occurring under conditions of high nutrition, not on the ` |. same mycelium with the other recognized form of chlamydospores, and even, to a certain. — . SECOND SERIES.— BOTANY, VOL. I. 3M 422 CONIDIAL FRUCTIFICATION OF CHOANEPHORA. extent, in an inverse relation to the sporangia. The description, too, of the different degrees ` of development of the filaments and dilatations bearing the aerial chlamydospores recalls very forcibly the phenomena occurring in the case of the unequivocally conidial frueti- ` fication of Choanephora. ; The above observations have not been made with any desire to detract from the value ` — of the brilliant series of observations recorded by MM. Van Tieghem and Le Monnier, but simply with a view to pointing out that the subject cannot yet be regarded as satis- factorily decided. It is not, of course, to be assumed that because one genus of Mucorini has been shown to possess both sporangial and conidial forms of fructification, all other genera must necessarily do so also. Still as one has been shown to do so, and as the production of the various forms of fructification has been shown to be closely connected with variations in the conditions of the nutrition of the plant, there are some grounds ` ` for suspecting that the phenomenon may not be an exceptional one, and there is cer- tainly reason for renewed investigation in order to determine the point. DESCRIPTION OF PLATE XLVII. Fig. 1. Mature conidial head of Choanephora. x 76. Fig. 2. Head after the fall of the conidia. x 180. Fig. 3. Immature capitellum with young conidia. x 800. Fig. 4. Capitellum after fall of conidia. x 800. Fig. 5. Diagram of the various types of conidial heads. Fig. 6. Mature conidium. x 960. Fig. 7. Conidium and young germinal tube. x 960. Fig. 8. Conidium and portion of young mycelium. x 960. 9 . Sporangial fructification. x 180. . Ruptured sporangium and mature spores. x 840. . Commencement of sexual process ; formation of club-shaped dilatations on the mycelium. x 360. Fig. 12. Conjugation of sexual cells; isolated sexual cell and empty dilatations. x 420. Fig. 13. Sexual cells in contact. x 840. Fig. 14. Mature sexual cell, showing arcuate form. x 480. Fig. 15. a, Conjugated cells ; 6, commencing growth of the zygospore; c, continued development of the zygospore; d, mature zygospore. x 180. Fig. 16. Ruptured zygospore, with escape of oil and granular protoplasm. x 180. Fig. 17. Chlamydospore. x 960. Fig. 18. Chlamydospores and mycelial filaments. x 180. Fig. 19. Origin of conidiiferous filaments. x 180. Fig. 20. Mycelial filaments. x 180. J ru &&i kel Lm TO Fig.15.x190, - D IFICATION OF CHOANEPHORA. eg? oh? XXIV. On a new Species of Helvella. By Wu PuiruiPs, P.L.S. (Plate XLVIII.) Read June 5th, 1879. IN a collection of Fungi gathered in 1876 on the Sierra Nevada mountains, in California, and sent for determination by Dr. Harkness of San Francisco, was a species of Helvella presenting characters essentially different from those of any species hitherto described. The specimens have evidently been dried with great care, and arrived here in a most satisfactory condition, which is not always the case with large and fleshy species. It is to be hoped that Dr. Harkness will continue to prosecute his researches in that highly interesting country, as there can be little doubt that it possesses many treasures to reward the labour of diligent investigation. HELVELLA CALIFORNICA, Phillips, n. sp. (Pl. XLVIII. figs. 1-6.) Pileus campanulate or saddle-shaped, deflexed, sublobate, free, veined beneath, purplish brown ; stipes longitudinally costato-lacunose, rosy pink; asci cylindrical, nar- rowed towards the base; sporidia 8, elliptic, binucleate ("017 x 009 millim.); paraphyses linear, clavate, and brown at the apices. | On the earth in dense forest, near rocks, Sierra Nevada mountains, California (Dr. Harkness, no. 1005). The pileus is from two to six inches in diameter, the stem from two to six inches high and about three quarters of an inch to one and a half inch in diameter. Its nearest ally is Helvella crispa, Fr., from which it differs in the colour of the hymenium and the stem, and in being a larger species. DESCRIPTION OF PLATE XLVIII. Figs. Lë 2. Helvella californica. Two medium-sized specimens, of natural size. Fig. 3. A median perpendicular section of a smaller-sized specimen. Fig. 4. A transverse section of the stem, viewed partly sideways, and of natural size. Fig. 5. The asci and paraphyses, magnified about 400 times. Fig. 6. The sporidia, magnified to the same extent as the asci. SECOND SERIES.—BOTANY, VOL. I. 3N X TRAxs.Limm.Soc.Srn.2.Bor Vor 1 Pr 48. WPhillips del. H C A PLEA IP n Titch imp Freq. g. HELVELLA CALIFORNICA, ZAzZzps. wap. me This page number was omitted in the original pagination. No page or information was missed during scanning. L [ 425 ] XXV. A Review of the Ferns of Northern India. By CHARLES BARON CLARKE, M.A., F.L.S. (Plates XLIX.-LXXXIV.) Read June 19th, 1879. INDIA proper (that is, exclusive of the Malayan or Trans-Gangetic Peninsula) was divided by Kurz into three main regions, viz. :—(1) the Himalaya, extending from Kashmir to Bhotan and Chittagong ; (2) the Peninsula with Ceylon, extending as far north as the tableland extends; and (3) the great plain between, the home of the Hindoos, Hin- doosthan. The area included in the present paper comprises the first and third of these divisions, i. e. all India proper except the Peninsula. Several considerations have induced me to confine myself to this area. First, I have collected Ferns myself from Kashmir to Bho- tan, Khasia, and Chittagong, for upwards of eleven years; I have only visited South India for a few weeks. Secondly, Col. Beddome has spent his Indian career in Southern India; and his aecount of the Ferns of Southern India is, in the judgment of Major Henderson, nearly unassailable; but Col. Beddome has, T believe, hardly collected at all himself in Northern India, and his invaluable plates of the Ferns of British India are, in the case of many species, founded on very scanty material As my own Herbarium contains more forms from Northern India than the whole Kew accumulations, I hope therefore to have been able to add somewhat to the previous A MAI of the North- Indian Ferns. I have drawn up the present paper at Kew, seated within a few yards of Mr. Baker and I believe I have taken advantage of his extreme courtesy to the extent of asking his opinion on every point as it turned up. While, therefore, he is not responsible for the view I may have adopted in each case, I may claim that I have had the full benefit of his experience. After the different tribes have been written out, Major Henderson has gone over the whole material at Kew with me; he has corrected several slips that I had made; and in the body of the paper I have in all cases mentioned particularly any point wherein he differs from what I have written. The present paper is therefore virtually a paper by Major Henderson and myself; but, as I have done all the writing, he declined to have it so entitled. The paper is in the form of a copious appendix to Hooker and Baker's ‘ Synopsis Filicum ;’ i. e. I have often given no diagnosis of a species; and the remarks on every Species are additions or corrections to the account in the ‘Synopsis Filicum. No person is likely to undertake the study of Indian Ferns without this book at his elbow ; and Ihave not wished to print more repetition than the large quantity always absolutely necessary in work of this kind. The first 48 pages of the ‘ Synopsis Filicum `" were done 3N 2 426 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. by Sir W. J. Hooker, the remainder by Mr. Baker; but for shortness I have quoted it as though a joint work. Botanists are pretty well agreed in England to give each Phænogam the specific name given it by him who first referred it to the right genus; but Pteridologists follow the ` contrary rule, giving each Fern the specific name given it by him who first referred it to the wrong genus or to any genus. Mr. Bentham, in the ‘Ferns of the Australian Flora,’ has submitted to this distinction; it seems to me untenable. It is argued, indeed, that the species of Ferns have been arranged in genera in so many ways that : less confusion on the whole arises by taking the oldest specific name; I think this isa very dangerous line of argument for any one to advance who follows the contrary rule ` with regard to Pheenogams. I doubt whether the genera of Indian Ferns have been ` more changed than those of the Indian Rubiaces. However, as the present paper is founded on Hk. & Baker's Syn. Fil., I have, to avoid confusion, followed the rule adopted ` ` in that work, and have only changed Hk. & Baker's names where I have supposed — myself compelled to do so in order to be consistent. I have quoted a large numberof | synonyms, which I believe a matter of great importance for the firm establishment of — species; I have quoted none that I have not looked up and satisfied myself to mean the — plant referred to: the only exceptions to this statement are references to Milde's d Equisetum, Luerssen's Ophioglossum, and a few others of the same kind, in which : T cases botanists will at once understand that it would take weeks to master such elaborate monographs, and that I have not done so. a I have endeavoured to fix more accurately the habitat of the North-Indian Ferns. To — read Hk. & Baker’s Syn. Fil., or even Beddome’s ‘Supplement to the Ferns of India, t it would appear that the vast moist tropical plain of Bengal was nearly destitute of ` ferns. This, I need hardly say, is not so— several ferns attributed by authors to the * Himalaya" being abundant in Bengal, and several of them indeed, so far as my knowledge extends, being quite unknown in the Himalaya. | The Plates appended to the paper are entirely supplementary to those of Col. Beddome, and are not intended to be complete in themselves. Col. Beddome has so nearly exhausted the Ferns of India, that the Plates are merely designed to bring out minor differences to assist in specific determination: e.g. when a Plate is given of a fern scarcely specifically different from the universal Pteris quadriaurita, I have not given an analysis of the fruit to show that the genus is Pteris. So of the critical Athyriums, | the Plates are designed merely to give an idea of the cutting, so difficult to define. ge words. | In the *Supplement to the Ferns of India,' Col. Beddome allows (species and vin with separate numbers) l Ap SOO. E ar eo O In Southern India . . . sous MEM £A In the Trans-Gangetic Peninsula sor te DEUS 1n In Northern DUE. senes usc ` LA The nu paper admits 363 species in Northern India, exelusive of 12 ric n MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 427 4 Equisetums, which I have added to make the paper include all the Acrogens having spores of one kind only. By far the largest genera in North India are Polypodium (67 species), Asplenium (56 species), Nephrodium (54 species). The new species described are 16, viz. :— Hymenophyllum Levingii. Nephrodium ingens, W. S. Atkinson (compound Lastrea). Cheilanthes albomarginata. Nephrodium Wightii, - iis Pteris subindivisa. Nephrodium de P stt Asplenium bellum, Allantodioid. Polypodium subtripinnatum (Phegopteris) . Asplenium torrentium, Polypodium chattagramicum (Dictyopteris) . Asplenium sikkimense, reng Polypodium subameenum (Goniophlebium). Asplenium succulentum, : Polypodium jaintense (Niphobolus). Nephrodium rhodolepis (compound Lastrea). Polypodium clathratum (Pleopeltis). A considerable number of species (especially of Wallich’s) that have been sunk or misunderstood are rehabilitated. More interesting than these or the new species are several cases where, by aid of the more abundant material at my command, I have altered the genus (or subgenus) to which the plant is referred: such are :— Hemitelia Brunoniana, formerly Alsophila Brunoniana, Wall. Diacalpe feniculacea, Si Aspidium feniculaceum, Hook. Davallia dareeformis, m Polypodium dareæforme, Hook. Lindsaya repens, St Davallia repens, Baker. A. (Euasplenium) longifolium, y A. (Diplasium) longifolium, Baker. A. (Pseudallantodia) procerum, ` ,, A. (Athyrium) umbrosum, var., Baker. N. (Lastrea) sikkimense, » Aspidium sikkimense, Baker. P. (Pleopeltis) erythrocarpum, ` P. (Goniophlebium) erythrocarpum, Baker. Several of these species, placed in wrong genera or subgenera, have been hitherto insoluble mysteries to many Indian botanists who had collected the plants, but looked for ` them in the book under other genera or subgenera. I have appended to the paper a complete reduction of the North-Indian Ferns in Wallich’s Herbarium. This will be of some value to this Society in showing the state of that Herbarium, and may also assist botanists to form a correct estimate of the weight to be attached to the quotations of the Wallichian numbers. In very many cases a mixture of ferns is pasted down under one number; and in numerous cases the same fern appears under different numbers. The rule as to quoting Wallich’s names is that the name applies to the plant on the large paper type-sheet (letter A). But this rule cannot be implicitly followed: the Wallichian no. 361 contained a large number of duplicate sheets; all these were one fern, Aspidiwm fuscipes, to which the specific name _ fuscipes has been applied; but I find that the type-sheet no. 361 is Nephrodiwm sage- nioides, Baker, belonging to a different subgenus. About 3 per cent. of the Wallichian Ferns are blank sheets, on which are pasted the Wallichian lithographed ticket. I learn from Mr. West that these Ferns arrived loose, and that when they came to be pasted down it was in many cases found impossible to discover the plant belonging to the ticket ; this seems to have been specially the case with the rarer and more critical species, of which probably only a small quantity was collected. The result of all these compli- cations is that I begin to doubt whether the great pains taken in quoting Wallich’s numbers are well bestowed ; for I find, in very many instances, that such quotations have merely puzzled or misled. Mr. Moore has already pointed out that the Aspleniùm Fin- 428 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA, laysonianum, Wall., of Hooker & Baker, is not Asplenium Finlaysonianum, Wall., nor even of the same subgenus, the fact being that the Wallichian duplicate sheets of Asple- nium Finlaysonianum, Wall, at Kew, are not the same as the Wallichian type-sheet; and if errors from this cause are not rare at Kew, botanists at a greater distance from head quarters cannot be expected to avoid them. Of the 379 species of North-Indian Ferns, 88 (so far as yet known) are confined to it. In the Himalaya westward from Nepaul 149 species occur, while in the Himalaya east of Nepaul 269 species occur; the Eastern region of Khasia and Chittagong has afforded 258, while 52 species occur in the plains. Out of 75 genera (as understood by Hooker and Baker) 50 are represented in North India. 1. GLEICHENTA, Smith. 1. G. evauca, Hook. Sp. Fil. i. 4, t. 3 B, not of Swartz; Mett. in Ann. Mus. Lugd. Bat. 1.48. G. longissima, Blume, Enum. Pl. Jav. Fil. 250; Hook. Sp. Fil. i. 4; Mett. Lei Hk. & Baker, Syn. Fil 12; Luerssen, Fil. Graeff. 248. G. gigantea, Wall. Cat. 157, as to the type-sheet ; Hk. & Bauer, Gen. Fil. t. 39; Hook. Sp. Fil. i. 5, t. 84 ; Bedd. Ferns Brit. Ind. t. 30. G. Bancroftit and excelsa, Hook. Sp. Fil. i. 5, t. 4 A, B. G. arachnoides, Mett. l. c. 47. OG japonica, Spreng. Syst. iv. 25. Poly- podium glaucum, Thunb. Fl. Jap. 338. Mertensia glauca, Swartz, Syn. Fil. 164, 390. M. glabra, Brack. U.S. Explor. Ferns, 1854. Sikkim and Bhotan; alt. 4500-7590 feet, in the dripping forest region, common. Khasia; alt. 8500-5000, common. Nepaul, Wallich.—Distrib. Extends eastwards through China, Japan, and Malaya, to Tropical Australia, Polynesia, and America. Not in the — Deccan. nor in Ceylon. EC Seandent over other jungle, sometimes covering a spur for several hundred feet. Fronds in pairs at the dichotomous branching of the main rhachis, often 6 feet long, pendent, curved. Texture of the pinnules hard, stout, margin recurved; veinlets ` ` 2-branched from near their base. Frond (sometimes on both surfaces) with the capsules often glistening, puberulous, from scattered minute subglobose glands. Rhachis and pinnules beneath glaucous or densely ferrugino-tomentose (in the Indian examples). 2. G. LINEARIS, C. B. Clarke. G. Hermanni, R. Br. Prodr. 161, and in Wall. Cat. 155; Blume, Enum. Pl. Jay. Fil. 248, not of Hk. & Grev. G. ferruginea, Blume, Enum. ` ` Fil. Jav. 249; Hook. Sp. Fil. i. 10. G. lanigera, Don, Prodr. Fl. Nepal. 17. Gd ` chotoma, Hook. Sp. Fil. i. 12; Bedd. Ferns South Ind. t. 74; Mett. in Ann. Mus. d Lugd. Bat. i. 50, t. 3; Carr. in Fl. Viti. 332; Hk. & Baker, Syn. Fil. 15 ; Benth. — Fl Austral. vii. 698; Luerssen, Fl. Graeff. 249. G. Klotzschii, Hk. Sp. Fil.i.19, ` t. 5, B. G. pteridifolia and crassifolia, Presl, Epimel. 23, t. 13, 14. Polypodium ` lineare, Burm. F1. Ind. 235, t. 67. fig. 2. D dichotomum, Thunb. Fl. Jap. 338, t. 37; Roxb. in Cale. Journ. Nat. Hist. iv. 493. Mertensia emarginata, Brack. U.S. Ex- ` plor. Ferns, 297, t. 42: M. dichotoma, Langsd. & Fisch. Voy. Russ. t. 29. MS Sikkim, Bhotan, and Khasia, alt. 0-5000 feet, common; extending Gene Nepal 3 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 429 to Kumaon, Strachey & Winterbotiom.—Distrib. Mts. of Malabaria and Ceylon, China, Japan, Malaya, to Tropical Australia, Polynesia, and America. Scandent over other jungle, sometimes for several hundred feet, often rooting. Fronds in divaricate pairs. Texture of the pinnules hard, stout, margin recurved; veinlets 3- (or more) branched from near the base. Pinnules beneath usually glaucous, with some ferruginous hair near the base of the pinna; sometimes this hair extends to the rhachis of the pinnules beneath. 2. CxATHEA, Smith. Involucre globose, uniform in texture, scarcely thinner at the apex, dehiscing by irre- gular lines when the capsules are ripe. At no stage of ripening, therefore, are the involucres to be found as hemispheres in which complete globes of unripe capsules are seated. (This definition is merely designed to separate the single North-Indian species from the closely allied Hemitelias.) l. C. serNULOSA, Wall. Cat. 178. Rhachis of pinnules beneath sparingly bullate-scaly, not pubescent; veinlets (in the segments of a pinnule from the middle of a barren pinna) 2-branched and frequently 3-branched; sori very large, the two rows occu- pying very nearly the whole breadth of the segment; involucre white, stout.—Hk. Sp. Fil. i. t. 12c. (Pl. XLIX: fig. 1.) Nepaul, Wallich; Jaintea Hills, alt. 4800-4500 feet, C. B. Clarke. Attains 30 feet. Prickly, to the rhachis of the pinnules. Pinnules rusty, hirsute- pubescent on the midrib above, otherwise nearly glabrous. Fertile and barren fronds somewhat dimorphous ; fertile segments much narrowed, especially in their fruit-bearing portion; the veinlets rarely 3-branched. The venation must be observed in the segments of the pinnules taken from the middle of a well-developed pinna from a barren frond. In many species of tree ferns with uniformly 2-branched veinlets, if an imperfectly-deve- loped pinnule be taken from a pinna near the top of the frond, the veinlets may be found 3-branched or subpinnate. Sir W. J. Hooker took his figures from Wall. Cat. 178, which are very faithful; but fig. 4 (taken probably from a segment with very unripe fruit) shows the sori too small, and, indeed, does not agree with fig. 3 (same place), which shows the sori nearly as broad as the segment. The accompanying description (Sp. Fil. i. 25) is unfortunately drawn up partly from Wight's no. 149, which is Alsophila late- brosa; and the diagnosis in Hk. & Baker, Syn. Fil. 23, is similarly compounded of two plants. Oyathea spinulosa, Bedd. Ferns South Ind. t. 57, represents a pinna of Hemi- telia Beddomei, C. B. Clarke, with analyses of segments copied from Hook. Sp. Fil. i. t. 12 c, which do not represent the Deccan species at all. Hemitelia Beddomei has the segments (of a pinnule from the middle of a well-developed pinna) elongate crenulate, veinlets frequently 3-branched, sori very small, involucre at first completely enclosing the sorus, but very thin and soon reduced to a hemispheric cup, and is not conspecific either with any Himalayan fern or any other than I can find in the Kew Herbarium. 430 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 9. HEMITELIA, R. Br. Involucre at first globose, thin at the vertex, usually disappearing from the vertex before the capsules are ripe; the capsules then appear as a globose mass, half-exsert from a hemispheric cup having a slightly toothed margin. As the capsules ripen the cup withers or becomes less than a hemisphere, but is usually to be found as a patelliform scale (not as a mere lateral bullate scale) completely surrounding the base of the sporo- phore after the capsules have fallen. 1. H. pEcretens, J. Scott, in Trans. Linn. Soc. xxx. 33, t. 14. Rhachis of pinnules beneath bullate-scaly, not pubescent; veinlets (in the segments of a pinnule from the middle of a barren pinna) 2-branched and occasionally 3-branched ; sori in two rows, rarely occupying the whole breadth of the segment ; involucre membranous, at first globose, but reduced before the capsules are ripe to a hemispheric cup with an obscurely toothed margin.—Hk. & Baker, Syn. Fil. 455. Alsophila decipiens, Bedd. Ferns Brit. Ind. t. 311, not good. Cyathea spinulosa, J. Scott, l.c. 32, t. 13 A; Kurz, For. Fl. Brit. Burma, ii. 572, not of Wall. Amphicosmia decipiens, Bedd. Ferns Brit. Ind. Suppl. 1. Sikkim; alt.1000—4000 feet, frequent. Bhotan; Kalimpoong, alt. 4000 feet, C..B. Clarke. Khasia; below Nungklow, Griffith, Hk. f- $ Thoms., and at the Bor Panee, Hk. Ji & Thoms. Attains 30-40 feet. Prickly, to the rhachis of the pinnules; pinnules glabrous or very nearly so; fertile segments narrowed in the fruit-bearing portion, or sometimes (in pinne from the same frond) not at all narrowed. This is specifically separated from ` T Cyathea spinulosa by the much rarer 3-branched veinlets and the young fruit, excellently figured by J. Scott, Z. c. t. 14. figs. 10, 11. I have no hesitation, with J. Scott's descriptions, figures, and type specimens before me, in uniting Cyathea spinulosa, J. Scott, to Hemitelia decipiens; the two are, so far as I can see, absolutely identical. Whether Cyathea spinulosa, Wall., really differs speci- fically may be questioned : it seems to have been collected only thrice; and it would be ` hazardous to conclude that the white globes of fruit, uniformly tough throughout their ` whole extent, are more than a casual variety of the fruit of the much commoner ` ` Hemitelia decipiens. 2. H. Brunontana, C. B. Clarke. Rhachis of pinnules beneath more or less crisped- — pubescent; veinlets (in the segments of a pinnule from the middle of a barren pinna) 2-branched, none 3-branched ; involucre membranous, at first (at least some- times) globose, reduced before the capsules are ripe to a hemispherie cup or sub- patelliform scale. —Alsophila Brunoniana, Wall. ; Hook. Sp. Fil. i. 52. A. latebrose, ` var. hemitelioides, J. Scott, in Trans. Linn. Soc. xxx. 34. Sikkim and Bhotan ; alt. 4000—7500 feet, very common. East Nepaul, Sir J. D. Hooker. Khasia ; alt. 3000-5000 feet, very common.— Distrib. Deccan. Usually 10-15 feet, attains 30-40 feet sometimes. POS eis muricated rhachis MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 431 of pinnze often free from prickles; 3-branched veinlets very rare. In the common Khasi plant the involucre appears as a half-cup with lacerate margin while the spores are dispersing; in the Sikkim form the involucre, though a complete thin globe in the earliest stage, is reduced ultimately to a patelliform scale, sometimes hidden by the ripe sorus, but seen as a disk surrounding the carpophore when the spores have been dis- persed. The whole of the Kew Cyathea spinulosa (except Wallich’s type specimen) is this plant; it is very easily distinguished therefrom by the 2-branched veinlets and the pubescent rhachis of the pinnules. There is no original specimen of Wall. Cat. 7073— the examples under that number in the Wallichian Herbarium having been “ taken from the specimen on the staircase," which it was supposed might have been 4. Bru- noniana, Wall.: these are Alsophila glauca, J. Smith. The trunk referred to by Sir W. J. Hooker remains in the Kew Museum; but the fronds on it described by him are gone; this trunk, however, is certainly not that of Alsophila glauca, as supposed in Hk. & Baker, Syn. Fil. 42. Griffith says (Private Journals, p. 170), “the Alsophila Brunoniana is apparently confined to the limestone hill at Cherra, while the tree fern Polypodium is found on sandstone." The limestone hill at Cherra, alt. 4200 feet, still supplies plentifully Hemitelia Brunoniana. By the tree fern Polypodium, Griffith doubtless meant the exinvolucrate Alsophila glauca, which extends from the Cherra Khud to Sylhet Station. Griffith also says (Private Journals, p. 7) that Wallich and himself found 4. Brunoniana during a trip from Cherra to Mamloo. The only tree ferns near this path are the Hemitelia Brunoniana across the brook on the limestone hill. Var.? Scottii. Segments of the fertile pinnules unusually large, deeply crenated, with many 3-branched veinlets. Sikkim.—A single fertile pinna, collected by J. Scott, and marked by him Alsophila latebrosa, var. B, of which it has exactly the fruit. The 3-branched veins are a con- comitant of the deeply crenate segments, which thus show an approach to a pinnatifid state. This is probably (as Scott evidently considered it) an unusually developed con- dition of the Sikkim form of Hemitelia Brunoniana; but I should not be surprised if it should prove a new species. 4. ALSOPHILA, R. Br. Involucre, if any, disappearing before the capsules are mature. In several species there are ovate or ovate-lanceolate bullate scales along the rhachis of the pinnules and segments. When the sorus is near the midrib of the segment, the bullate scale may be attached by its broad base almost under the sorus, but only laterally to the sorus on the inner side, and not continuous with the carpophore; so that when the capsules have dispersed it does not appear as a patelliform seale surrounding the base of the carpophore. The bullate scale is of lax areolar tissue, entirely different from the thin scale en} is the remnant of the involucre in Hemitelia, as J. Scott has remarked. ^ l. A. LATEBROSA, Hook. Sp. Fil. i. 37. Rhachis of pinnules beneath glabrous; veinlets (in the segments of a pinnule from the middle of a barren pinna) 2-branched; sori : . not nearly occupying the whole breadth of the me & Baker, Syn. Fil 45) — in (SECOND D SERTES.—BOTANY, Y VOL. LO -| 439 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. partly; Kurz, For. Fl. Brit. Burma, ii. 573. Polypodium latebrosum, Wall. Cat. 318, partly. Hemitelia latebrosa, Mett. in Ann. Mus. Lugd. Bat. i. 54? Malabaria; Pinang; throughout Malaya common. No example from Northern India. Var. ? Schmidiana, Kunze, in Linnea, xxiv. 294. Rhachis of pinnules beneath crispedly pubescent.—A. latebrosa, Bedd. Ferns South. Ind. t. 58; J. Scott, in Trans. Linn. Soc. xxx. 34, t. 18 B. A. khasiana, Moore; Kuhn, in Linnea, xxxvi. 154. Sikkim; alt. 3500-5000 feet, J. Scott. Assam, Mrs. Mack.—These are the only examples in the Kew Herbarium. I have several times collected the plant both in Sikkim, Bhotan, and Khasia; but it is far less common than Hemitelia Brunoniana. This species is a fine green, drying a rich brown, in texture so unlike the black-drying shining Malay A. late- brosa, that I strongly suspect it to be a different species. 2. A. GLAUCA, J. Smith, in Hook. Journ. Bot. iii. 419. Frond glaucous beneath; veinlets (in the segments of a pinnule from the middle of a barren pinna) 2-branched, often 3-branched; base of the carpophore, after the capsules have fallen, naked or surrounded by lax hairs.—M.ett. in Ann. Mus. Lugd. Bat. i. 54, with all syn. A. contaminans, Hook. Sp. Fil. i. 52, t. 18 B; Bedd. Ferns Brit. Ind. t. 85; J. Scott, in Trans. Linn. Soc. xxx. 35, t. 15; Hk. & Baker, Syn. Fil. 41; Kurz, For. FL — Brit. Burma, ii. 573. A. glaucescens, Wall. Cat. 7074. 4A. Wallichiana, Presl; Hook. Sp. Fil. i. 55. .4. Brunoniana, Bedd. Ferns Brit. Ind. t. 66, but not of Wall. d A. Sollyana, Griff. Notul. i. ii. 624, Ic. Pl. As. Rar. 180, fig. 3. Chnoophora glauca, ` Blume, Enum. Pl. Jav. Fil. 443. Polypodium contaminans, Wall. Cat. 320. Sikkim, Bhotan, Assam, Khasia, Cachar, Sylhet, Chittagong ; alt. 0—4000 feet, frequent. —Distrib. Burma, Malay Peninsula and Islands. | Attains 50 feet (J. Scott). More or less prickly; fruiting segments more or less - elongate; on the whole very uniform, and easily distinguishable from all other North- — S Indian Ferns. Neither of Beddome’s pictures is good; he shows an apparently well ` ` developed barren pinna without any 3-branched veinlets in the ultimate segments. - = J. Scott’s picture is excellent. 3. A. ORNATA, J. Scott, in Trans. Linn. Soc. xxx. 36, t. 16 A. Pinnules glabrous be- ` neath, or the rhachis slightly flocculose villous; fruit-segments linear-oblong; —— crenulate-serrate ; veinlets (in the segments of a pinnule from the middle of a barren pinna) 2-branched, a few 3-branched; base of the carpophore, after the | “ie have fallen, naked.—Bedd. Ferns Brit. Ind. t. 342; Hk. & Baker, EL] D Sikkim ; alt. 2500 feet, Government Cinchona-plantation, J. Scott. Collected by J. Scott twice apparently. Two fertile pinnæ at Kew, communicated by J. Scott. Otherwise unknown tome. Attains 20—40feet, with a somewhat slender trunk. Seems to me more nearly allied to A. latebrosa than to A. Oldhami, but well distinct ; ` the veinlets are much wider apart, sometimes 3-branched. Here, as in other species, there are no pinnate veinlets (as might be inferred from J. Scott’s words) ; at the tips of the frond Wie videtur being imperfectly developed, become confluent ; BI the m MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 433 of the imperfect segment be taken for a veinlet, and its veinlets for branches, then a . pinnate veinlet may be made out. 4. A. ANDERSONI, J. Scott, in Trans. Linn. Soc. xxx. 38, t. 17. Surface of the frond beneath with needle-like hairs; veinlets (in the segments of a pinnule from the middle of a barren pinna) 2-branched, none 3-branched ; carpophore hairy, without any involucre at its base after the capsules have fallen.—Bedd. Ferns Brit. Ind. t. 310; Hk. & Baker, Syn. Fil. 459. Sikkim; alt. 1000-2500 feet, J. Scott &c. Khasia, Dr. Jerdon. Attains 20-30 feet (J. Scott). Segments serrate ; sori small, the two rows wider apart at the base of the segment than at the apex. 5. A. OLDHAMI, Bedd. Ferns Brit. India, t. 343. Pinnules glabrous beneath, often with ovate adpressed scales on the rhachis; veinlets not branched, or a few 2-branched ; sori in two parallel rows, exinvolucrate.—A. Scottiana, Baker, in Gard. Chron. 1872, - 699, with fig.; Hk. & Baker, Syn. Fl. 460. Æ. comosa, J. Scott, in Trans. Linn. Soc. xxx. 36, t. 16 B, not of Wall. Sikkim ; alt. 5000-6000 feet above the Cinchona-plantation, and above Khursiong. Khasia, Dr. Oldham, Sir J. D. Hooker; alt. 4000 feet, C. B. Clarke.—Distrib. Moulmein. Grows in groves. Unarmed, trunks 6-10 feet. Ultimate segments oblong or narrow- oblong, strongly crenulate-serrate nearly to their base. 6. A. GLABRA, Hook. Sp. Fil. i. 51. Pinnules glabrous beneath, often with lanceolate or linear lax scales on the rhachis; veinlets not branched, or a few 2-branched; sori exinvolucrate, in two rows approximating towards the apex of the segment.—Bedd. Ferns South. Ind. t. 60; Hk. & Baker, Syn. Fil. 43; Mett. in Ann. Mus. Lugd. Bat. i. 52; J. Scott, in Trans. Linn. Soc. xxx. 38, t. 18; Kurz, For. Fl. Brit. Burma, ii. 573. A. gigantea, Hook. Sp. Fil i. 53. A. Helferiana, Presl, Gefüssb. 33. A. Metteniana, Hance, in Seem. Journ. Bot. 1868, 175. Cyathea venulosa, Wall. Cat. 180. Gymnosphera glabra, Blume, Enum. Pl. Jav. Fil. 242. Polypodium giganteum, Wall. Cat. 321. P.? wmbrosum, Wall. Cat. 336. P. altissimum, Wall. MS. From Nepaul to Assam and Chittagong; alt. 0-1000 feet, abundant, ascending to 4000 feet alt.—Distrib. Mts. of Malabaria and Ceylon, Burma, the Malay Peninsula and Islands, South China. o i: Usually 10-20 feet, attaining 50 feet. The rhachis of the pinnule is usually glabrous beneath, sometimes with scattered very narrow scales or flocculose hairs of lax tissue, never (in the North-Indian examples) with needle-like hairs as shown in the enlarged fragment of a pinnule in Beddome’s picture. The Assam example at Kew with needle- like hairs I take to be Polypodium auriculatum, Wall. Receptacle, after the capsules have fallen, glabrous, or more often minutely pubescent; not unfrequently with many lax ~ hairs (paraphyses) in North-Indian exampl 2d 303. 434 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 5. DIACALPE, Blume. 1. D. AsPrDIOIDES, Blume, Enum. Pl. Jav. Fil. 241. Ultimate segments oblong, cuneate, nearly glabrous beneath, reddish or brown when dry.— Hk. & Bauer, Gen. Fil. t. 99; Hook. Sp. Fil.i 59; Bedd. Ferns South. Ind. t. 257; Hk. & Baker, Syn. Fil. 45. D. Hookeriana, Moore, in Gard. Chron. 1854, 135. Spheropteris Hookeriana, Wall. Cat. 775. Peranema aspidioides, Mett.; Kuhn, in Ann. Mus. Lugd. Bat. iv. 285. Aspidiwm foliolosum, Wall. Cat. 359, partly by admixture. Nepaul to Bhotan; alt. 6000-9000 feet, abundant. Khasia; alt. 4000—6000 feet, common.—Distrib. Ceylon, Moulmein, Tavoy, Malay Archipelago. Very large and compound at its upper limit (9000 feet). Rhachis of pinna with blackish or chestnut linear scales or lax hairs. Involucre often splitting regularly by a clean dehiscence into two lips, in which state it cannot be separated from Dicksonia. This species so exactly resembles Davallia nodosa, Hook., that it has been repeatedly mixed with it by very competent botanists, from Wallich downwards. The pinne are often subopposite, with large ovate thin scales at their base; the venation is similar; and if the inchoate sorus (as Mettenius maintains) is not exactly similarly placed on the vein, it is very nearly so. Davallia nodosa may generally be distinguished (apart from the involucral character) by the much greater glabrousness of the rhachis of the pinna. 2. D. FENICULACEA, C. B. Clarke. Ultimate segments linear-acute, with few scattered long lax hairs beneath, glossy green when dry.—Aspidium feniculaceum, Hook. Sp. Fil. iv. 36, t. 237; Hk. & Baker, Syn. Fil. 256. Lastrea faniculacea, Bedd. Ferns Brit. Ind. t. 36. Sikkim; alt. 7000—10,000 feet, east, west, and north, scattered, not common; but ` abundant at Buckeem, Upper Ratong, alt. 8000—9000 feet. Stipes tufted. Closely allied to D. aspidioides. The involucre is leathery, dark purple, spherical, attached by the one point under its centre, not stalked, splits irregularly from above into 2-5 triangular segments. 6. Oxocrza, Linn. di 1. O. omrzwTALIS, Hook. Sp. Fil. iv. 161; Hk. & Baker, Syn. Fil. 46. Séru£hiopteriS ` orientalis, Hook. 2nd Cent. Ferns, t. 4; Bedd. Ferns Brit. Ind. t. 171. x 2 Sikkim; Lachen, alt. 12,000 feet, Sir J. D. Hooker; alt. 9000 feet, H. J. Elwes. ` Khasia (probably near Shillong), Dr. Jerdon. Assam (probably near Shillong or pe D sibly the Gowhatty Teelas), Simons.—Distrib. Western China, Japan. 7. Woonsta, R. Br. 1. W. HYPERBOREA, R. Br. in Trans. Linn. Soc. xi. 173, t. 11; Hk. & Bauer, Gen. Fil. Xs t. 119; Hook. Sp. Fil. i. 64; Hook. Brit. Ferns, t. 7; Hk. & Baker, Syn. Fil 46. W. hyperborea, R. Br. A. form 8. subcordata, Mili Fil. we: 163. siden: Airimem M ue t. 2023. : ilr: ; MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 435 Kashmir; Sind valley, alt. 8000 feet, H. C. Levinge, once collected.—Distrib. Alpine : and Arctic Europe and North Asia. Fronds 5 by i-$ in.; lowest pinn 4 in. distant. Pinns ovate and cordate-ovate, crenate with rounded lobes; needle-like hairs plentiful near the sori, few on other por- tions of the lower surface of the pinna. H. C. Levinge's example is W. hyperborea type, in nowise verging towards Woodsia lanosa. 2. W. nANOSA, Hk. & Baker, Syn. Fil. 47; Bedd. Ferns Brit. Ind. t. 341. JV. mollissima, Hook. MS. Kumaon; Pindari alt. 12,000 feet, above Namik alt. 11,600 feet, Strachey § Winter- bottom. Sikkim; Lachen, alt. 14,000-16,000 feet ; Mt. Donkia, alt. 18,000 feet, Sir J. D. Hooker. | Like W. hyperborea, but the frond densely ferruginous, lanate beneath. The stipe is also villous or lanate in all the examples, though Beddome figures it glabrous; but the wool is deciduous, and seems likely to disappear altogether in age. d W. EzLoNGATA, Hook. Sp. Fil. i. 62, t. 21 c; Hk. & Baker, Syn. Fil. 47; Bedd. Ferns Brit. Ind. t. 14. Himalaya; from Dhurmsala to Sikkim, alt. 8000—12,000 feet, frequent. Frond 9 by 1-14 in., narrowly oblong; sori globose, much elevated; involucre thin but persistent, globose (though torn) in the over-ripe fronds. 8. PERANEMA, Don. - 1. P. cxarHEorpEs, Don, Prodr. Fl. Nep. 12; Bedd. Ferns South. Ind. t. 73. Sphero- pteris barbata, Wall. Cat. 183, Pl. As. Rar. t. 48; Hk. & Bauer, Gen. Fil. t. 22; Hook. Sp. Fil i. 58; Hk. & Baker, Syn. Fil. 49. Cyathea barbata, Wall. MS. Aspidium spectabile, Wall. Cat. 372, partly by admixture. Nepaul and Bhotan; alt. 6000-10,000 feet, plentiful. Khasia; alt. 4500-6000 feet, plentiful.—Distrib. Western Ghats, fide Bedd. l. c. 25. Main rhachis clothed with lanceolate-acuminate scales, and also laxly pubescent. Aspidium spectabile, Wall. Cat. 372, is Lastrea pulvinulifera, Bedd., mainly ; but some Peranema has got mixed. The two (when without fruit) are difficult to separate; but Aspidium spectabile has the rhachis (exclusive of the scales) nearly glabrous. 9. Drcksox1A, L'Hérit. 1. D. Baromerz, Hk. & Baker, Syn. Fil. 49. D. Baranetz, Link, Fil. Sp. 166. D. as- samica, Griff. Notul. i. ii. 607. D. Griffithiona, Griff. Ic. Pl. As. 186, fig. 2. Cibo- tium Barometz, J. Smith, in Hook. Lond. Journ. Bot. i 497. C. glaucescens, Kunze, Hook. Sp. Fil. i. 82. C. assamicum, Hook. Sp. Fil. i. 83, t. 295. C. glau- cum, Bedd. Ferns Brit. Ind. t. 83, not of Hook. & Arn. Polypodium Barometz, Linn. Sp. Pl. (ed. princeps) 1092. 436 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Mishmee, Griffith. Assam, Griffith, Mrs. Mack, Jenkins.—Distrib. Tavoy, Malay Archipelago, South China. Involucres large, persistent, almost or quite touching each other. The rhachises of the pinnnles are arachnoid or paleaceous in the Assam examples, but not hirsute as Griffith has depicted. | 2. D. scABRA, Wall. Cat. 2173; Hook. Sp. Fil. i. 80, t. 27 5; Hk. & Baker, Syn. Fil. 54. D. deltoidea, Hook. Sp. Fil. i. 80, t. 27 ^. .Dennstedtia deltoidea, Bedd. Ferns South. Ind. t. 258. | Himalaya; from Kumaon to Bhotan, alt. 4000-8000 feet, common. Khasia; alt. 4000- 6000 feet, common.—Distrib. Ceylon, Malay Peninsula. Rhizome creeping, villous. Main rhachis scabrous, often wavy. Frond ovate or del- toid; pinnz often distant; surface beneath with scattered glistening hairs. Very variable in size, and not, by that character, to be distinguished from D. appendiculata— — | As to the synonym Sitolobiwm strigosum, J. Smith (adduced here in Hk. & Baker, Syn. Fil. 54), J. Smith says (Genera of Ferns, 102) that he meant thereby Davallia strigosa, Swartz; and in Historia Fil. 261, he refers the same species to Microlepia: many examples of Dicksonia scabra have been marked Microlepia strigosa in the Herbarium ; but it does not appear clear that they were so marked by the hand of J. Smith. 3. D. ELWESII, Hk. & Baker, Syn. Fil. 54. Patania Elwesii, Bedd. Ferns, Suppl. t. 347. Sikkim; Lachen, alt. 8500 feet, H. J. Elwes; Choongtam, alt. 9000 feet, Sir J. D. -Hooker. = Glabrous, or the rhachis fibrillose. Frond large, lanceolate, elegantly cut. Ultimate segments oblong, entire, often somewhat clavate at the summit, which is entirely occupied by the sorus. Lower segments of the pinnule often divided; when barren, acute. 4. D. APPENDICULATA, Wall. Cat. 65; Hook. Sp. Fil. i. 79, t. 27 0; Bedd. Ferns Brit. Ind. t.82; Hk. & Baker, Syn. Fil. 54. | North-west Himalaya; Kumaon, Gori valley, alt. 5500 feet, Strachey & Winterbottom ; banks of Vishnugunga above Panchkisar, alt. 8000-9000 feet, Edgeworth. Nepaul, Wallich. Sikkim; Lachen valley, Blanford & Elwes; Sinchul, alt. 8500 feet, N. Gamble. ` Rhachis of frond hairy. Frond oblong; pinne close together, their rhachises parallel. ` ` Surface of frond beneath with scattered hairs. ! 10. HYMENOPHYLLUM, Linn. * Margin of frond entire, neither serrulate nor ciliate. ` B. 1. H. resp, Wall. Cat. 170, as to the type sheet. Rhachis beneath with scattered ` lax rufous hairs.—Hook. Sp. Fil. 109, t. 384 ; Bedd. Ferns South. Ind. t.9; Hk. & ` Baker, Syn. Fil. 58. HM. macroglossum, v. d. Bosch, Hymen. "E: ar H. densum, oS Wall. Cat. E eet he Lee DEE MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. | 437 . Himalaya; from Kumaon to Bhotan, alt. 4000-9000 feet, very common. Khasia; alt. 2000-5500 feet, common.—Distrib. Tenasserim, Mts. of South India and Ceylon. Frond lanceolate, oblong, ovate, or short-triangular.— H. ciliatum, Swartz, differs by the margin of the frond being ciliate. Wallich’s H. exsertum has some H. polyanthos mixed with it. 2. H. potyantuos, Swartz, Syn. Fil. 149. Glabrous or very nearly so, frond not crisped, stipe without a wing, or with an exceedingly narrow wing.—Hk. & Grey. Ic. Fil. t. 128; Hook. Sp. Fil. i. 106; Bedd. Ferns South. Ind. t. 280, not t. 267; Hk. & Baker, Syn. Fil. 60. H. polyanthos B. minor (= H. microsorum, v. d. Bosch ex Bedd. SuppL), Bedd. Ferns Brit. Ind. t. 306. H. abietinum, Hk. & Grev. Ic. Fil. t. 127. H. Blumeanum, Spreng.; Blume, Enum. Pl. Jav. Fil. 220; v. d. Bosch, Hymen. Jav. 46, t.36; Bedd. Ferns South. Ind. t. 266. H. badiwm, Wall. Cat. 172, not of Hk. & Grev. H. microsorwm, v. d. Bosch, Hymen. Suppl. 71; Hk. & Baker, Syn. Fil. 59. H. protrusum, Hook. Sp. Fil. 104, t. 37 B. H. pycnocarpum, v. d. Bosch, Hymen. Jav. 48, t.37. H. integrum, v. d. Bosch, Hymen. Jav. 49, t. 38. H. sphe- rocarpum, himalaianum, osmundioides, v. d. Bosch, Hymen. Suppl. 83, 72, 80. Himalaya; from Kumaon to Bhotan, alt. 1000—12,000 feet, abundant. Khasia; alt. 2000-6000 feet, very common.—Distrib. Mts. of Malabaria and Ceylon, Burma. In nearly the whole world in tropical and subtropical moist regions. Frond (in the Himalayan examples) varies from triangular to linear; involucres variable in size, often smaller than in v. d. Bosch's typical H. microsorum.—Only the species which v. d. Bosch has founded on North-Indian specimens are included in the above quotation of synonyms. None of these can be ranked even as à variety according to the scale adopted in the present paper. Not only may they all be collected in a short walk from Darjeeling, but a large number (I might say an indefinite number) of addi- tional species (such as they are) will be collected during the same walk. 3. H. JAvANICUM, Spreng. Syst. iv. 132. Glabrous or very nearly so; frond more e less crisped, especially the wing of the main rhachis, which is carried down the stipe.— Blume, Enum. P]. Jav. Fil. 222; Hook. Sp. Fil. i. 106; Hk. & Baker, Syn. Fil. 60; v. d. Bosch, Hymen. Jay. 50, t. 40; Benth. Fl. Austral. vii. 705. H. serpens, Wall. at.173. H. fimbriatum, J. Smith ; Hook. Sp. Fil. 102. t. 360; v. d. Bosch, Hy- men. Jav. 55, t. 44. H. flexuosum, A. Cunningham ; Hook. Sp. Fil. i. 105; Hook. Ic. Pl. t. 962. H crispatum, Wall. Cat. 169; Hk. & Grev. Ic. Fil. t. 77 ; Hook. Sp. Fil. i. 105; Bedd. Ferns South. Ind. t. 207. H. micranthum, v. d. Bosch, Hy- men. Jav. 52, t. 41. H. erosum, Blume, Enum. PI. Jav. Fil. 221; Hook. Sp. Fil. i. 108; v. d. Bosch, Hymen. Jav. 54, t. 43. H. dedaleum, Blume, Enum. Pl. Jav. Fil. 222; Hook. Sp. Fil. i. 108. H Reinwardtii, v. d. Bosch, Hymen. Jav. 52, t. 42. Himalaya ; from Nepaul to Bhotan, alt. 5000-8000 feet, common. Khasia ; alt. 3500- . 5500 feet, common. Distrib. Mts. of Malabaria and Ceylon; Burma, Malay Peninsula, . Australia, New Zealand, Mauritius, and Bourbon. E 438 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Var. badium, (sp.) Hk. & Grev. Ic. Fil. t. 76. Frond hardly crisped.—Hook. Sp. Fil. i. 105; Bedd. Ferns Brit. Ind. t. 282; Hk. & Baker, Syn. Fil. i. 60, not of Wall. The figure of Hk. & Grev. is taken from an example with large ultimate segments, and the stipe winged to the base, not the same as Wall. Cat. 172. ** Margin of frond serrulate. 4. H. Srmonstanum, Hook. 2nd Cent. Ferns t. 13. Stipe with few ferruginous hairs, frond pinnatifid to the winged rhachis; the primary segments 1—4-lobate; ultimate segments broadly oblong, serrulate, often 2 millims. broad.—Hk. & Baker, Syn. Fil. 68; Bedd. Ferns Brit. Ind. t. 281. Khasia, Dr. Simons. Sikkim; alt. 4000-10,000 feet, common; extending from the plains to Kinchinjunga, and from East Nepaul to West Bhotan (and doubtless much more widely). ? Stipe winged at the top or not. Frond 6 in. linear oblong, or round-ovate, 1l in. in — diam., and then full of fruit. Surface of the frond beneath usually glabrous; but ferru- — - ginous long hairs occur sometimes. Involucre subquadrate, often 2 millims. long. This fine species is at once recognized in Sikkim by its large size, broad segments, and large involucres. 5. H. DENTICULATUM, Swartz, Syn. Fil. 148 & 375. Frond slightly crisped, pinnatifid to the winged rhachis; primary segments pinnatifid ; secondary segments oblong, some- ` times again divided, serrulate on the margin, the serrulation often continued to the ` e wing of the main rhachis; involucre ovate, with serrulate valves.—H ook. Sp. Fili 101; Bedd. Ferns Brit. Ind. t. 278; v. d. Bosch, Hymen. Jav. 39, t. 29; Hk. & ` Baker, Syn. Fil. 71. Trichomanes denticulatum, Blume, Enum. DL Jay. Fil. 226. . Khasia ; alt. 4000-5000 feet, frequent ; plentiful in and round Cherra Station. Bhotan, — among the Duphlas, Col. Hutchinson.—Distrib. Burma, Malay Peninsula and Islands. ` Stipe usually with some fine scattered ferruginous hairs (even in the Malacca ex- — — amples), rarely naked. Wing of the main rhachis usually produced to the base of the — — frond, and more or less upon the stipe. Frond varying from ovate to narrowly oblong, | sometimes glabrous, sometimes with scattered hairs, especially on the main rhachis. _ Margin of the ultimate segments more or less acutely serrulate, generally through their ` ` whole outline. Involucres usually (in the Khasia examples) glabrous on their backs, — - sometimes scabrous, rarely with one or two hairs; the margin of the valves strongly ser- rulate, or often pectinate. Var. bride: sp., Y. d. Bosch ; Bedd. Ferns Brit. Ind. t. 276. Main rhachis often with ` much ferruginous hair beneath; frond flaccid, hardly at all crisped; ultimate seg- ` ments slightly serrulate in their upper portion only, the main rhachis quite entire; ` valves of the involucre slightly serrulate at the summit. H. khasianum, Hk. » Baker, Syn. Fil. 464. (Pl. KLIX. fig. 2.) | Khasia ; Cherra Station, growing with the type. - MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 439 This var. may be related to H. denticulatum as H. badium is to H. javanicum, repre- senting the fully developed uncrisped form of the plant. Every variation in denticula- tion and hairiness, and in serrulation of the involucre-valves, from H. flaccidum up to H. denticulatum may be seen in my copious examples. The fronds vary in shape from ovate to linear-oblong.—H. Smithii, Hook. Sp. Fil. i. 97, t. 35 B, slightly differs from H. denticulatum, var. flaccidum, by the ultimate segments being more narrowly oblong, the involucre-valves more elongate, less serrate, the carpophore frequently long exsert. It is a fern of the Malay Peninsula and Islands; and the Khasia example referred to it is typical H. denticulatum, var. flaccidum.—H. ciliatum, Swartz, is stated (Hk. & Baker, Syn. Fil. 63) to have been found in the Hast Himalaya, and by Beddome (Ferns Brit. Ind. t. 305) to come from Sikkim. There is no example of it at Kew from Northern India; nor have I ever seen an Indian specimen. Col. Beddome obtained his North- Indian Ferns second-hand ; and many of the collectors mixed in their private collections ferns from all parts of the world. I am not sure that Col. Beddome’s picture represents the true plant, which exhibits needle-like, stellate, sessile and stalked hairs, not there indicated (see Hk. & Grev. Ic. Fil. t. 35). 6. H. Levinen, C. B. Clarke. Frond small, narrowly oblong, not crisped, pinnatifid to the winged rhachis; primary segments 1l-4-lobate; ultimate segments oblong, remotely serrate, their midrib with many hairs and lanceolar scales of the same texture as the frond. (Pl. XLIX. fig. 3.) Sikkim; Yoksun and Neebay, alt. 7000 feet, C. B. Clarke. Very delicate in texture. Stipe lin. with moniliform hairs. Frond 1-2 in. long, more or less covered with moniliform hairs. The lanceolar scales on the midrib beneath are attached by their whole base; they are sometimes rare, sometimes very numerous, so as to form a thick coat beneath the frond. Involucres usually 1-2 at the end of the segment, small, glabrous, subquadrate, valves separating nearly to the base, entire or slightly toothed at the apex; capsules of Hymenophyllum 2-4 to each involucre, carpo- phore included.—This is not much like any other species of the genus. 11. TRICHOMANES, Smith. 1. T. wuscorpzs, Swartz, Syn. Fil. 141. Frond undivided, with a submarginal nerve.— Hk. & Grey. Ic. Fil. t. 179; Bedd. Ferns Brit. Ind. t. 304. In the tropics of nearly the whole world and in Ceylon; not yet known from Northern India. Var. sublimbatum, sp. C. Muell. in Bot. Zeit. 1854, 737. No trace of a submarginal nerve.—Microgoniwm sublimbatum, v. d. Bosch, Hymen. Jav. 6, t. 2. Khasia, alt. 5000 feet; sandy rocks near Surureem, Griffith; once collected, Nov. 2, 1835, which day Wallich and Griffith halted in the Surureem (i.e. Sohra Reen) Bun- galow and devoted to collecting. ; SS, Griffith’s examples are very perfect; they agree altogether with the Java 7. sublim- . SECOND SERIES.—BOTANY, VOL. x. ? ? id 440 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. batum and not with the Ceylon plant (Microgonium bimarginatum, v. d. Bosch), which is a point in geographic distribution whether 7. sublimbatum be estimated a species, or a var. as in Hk. & Baker, Syn. Fil. 75. Col. Beddome's figure, though given to illustrate the North-Indian plant, was not, it is presumed, drawn from a North-Indian example; it represents the Ceylon plant. 2. T. Kunzir, Bedd. Ferns Brit. Ind. t. 286. Frond under 1 in. long, deeply pinnatifid ; primary segments 1-3-lobate, ultimate segments linear.— 7' nanum, Hk. & Baker, Syn. Fil. 77. Crepidomanes nanum, v. d. Bosch, Hymen. Suppl. 122. Assam, Griffith.—Distrib. Andamans. A note in the Kew Herbarium on this species runs :—** Very near T. gracile, v. d. Bosch, Hymen. Suppl. 23." There is no authentic example of T. gracile for comparison ; but from the description it appears very near. [Trichomanes parvulum, Poir. (Hk. & Baker, Syn. Fil. 75; Bedd. Ferns Brit. Ind. t. 179), is stated by Beddome, in Ferns Suppl. 3, to be a native of North India. There ` is no example at Kew thence. ] [Trichomanes digitatum, Swartz (Hk. & Baker, Syn. Fil. 76; T. corticola, Bedd. Ferns South. Ind. t. 264), is also stated by Beddome, in Ferns Suppl. 3, to be a native of North India. The scraps of Griffith were collected, in all probability, in the Malay Penisula; and it is not known on what authority Col. Beddome locates the species in North India.] 3. T. BIPUNCTATUM, Poir. in Encyc. Méth. Bot. viii. 69. Frond tripinnatifid; lips of the involucre wider than the tube, ovate-acute or subacute.—Luerssen, Fil. Graeff. 241. T. Filicula, Bory, in Duperrey, Voy. Bot. i. 283; Hook. Sp. Fil. i. 124; Carr. in Fl. Viti. 344; Bedd. Ferns Brit. Ind. t. 283; Hk. & Baker, Syn. Fil. S1. T. insigne, Bedd. Ferns Brit. Ind. t. 284. 7T. plicatum, Bedd. Ferns Brit. Ind. t. 285. Hymeno- phyllum Filicula, Willd. Sp. Pl. v. 528. H. densum, Wall. Cat. 171, as to the top specimen on the type-sheet. H. alatum, Schkuhr, Fil. t. 135 5. Didymoglossum Fili- cula, v. d. Bosch, Hymen. Jav. 35, t. 26. D. racemulosum, late-alatum, plicatum, euphlebium, v. d. Bosch, Hymen. Suppl. 58, 54, 55, 58. Himalaya; from Kumaon to Bhotan, alt. 1000-6000 feet, common. Khasia ; alt. 2000- — 5000 feet, common.—Distrib. Malay Peninsula, and the tropics of the whole world.—The _ Malabar examples from Bombay to Ceylon are somewhat intermediate between the . Himalayan T. bipunctatum and the Chittagong T. pyzidiferum, having the lips of the ` involucre as broad or broader than long, and rounded. The Himalayan type has the — — lips of the involucre ovate subacute, more so even than as shown in Beddome's plate. T There is no good line between T. bipunctatum and T. pyzidiferum. A T. PYXIDIFERUM, Linn. Sp. Pl. 1561. Frond tripinnatifid ; lips of the involucre wider than the tube, obtuse, broader than long, or sometimes almost truncate.—Hk. & Grey. Ic. Fil. t. 206; Hook. Sp. Fil. i. 124; Bedd. Ferns South. Ind. t. 7; Hk. & ` Baker, Syn. Fil. 81; Benth. Fl Austral vii. 703. P. Schmidianum, Zenk.; Taschner, — MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 441 Trichom. 34, t. 1. fig. 1. T. proliferum, Thwaites, Enum. 397. no. 3329, not of Blume : see G. W. Cat. Ferns Ceylon, 1. East Bengal; from Cachar to Chittagong, alt. 0—1500 feet, common.—Distrib. Deccan and Malay Peninsulas, and tropical and warm temperate regions throughout the world. There is no Himalayan example of this at Kew, except a scrap communicated by Levinge, named 7. pyxidiferum (by Beddome, correctly), and said to be from Darjeeling ; but this has travelled through many hands, and the original collector does not appear. Var. limbatum, Bedd. Ferns Brit. Ind. t. 348. Fronds up to 6-8 in. long; pinnæ larger and less eut than in the type. Khasia; alt. 6000 feet, Hk. f. & Th.—Beddome says (doubtless correctly) that this is T. limbatum, Wall.; but there is no example of Wallich's so named at Kew. 5. T. RADICANS, Swartz, Fl. Ind. Or. 1736. Frond.3-4-pinnatifid ; main rhachis naked, or winged sometimes to the base of the stipe; ultimate segments oblong, l-nerved ; lips of the involucre scarcely wider than the tube, often altogether trun- cate.—Hook. Sp. Fil. i. 125, with syn. ; Hook. Brit. Ferns, t. 42; Bedd. Ferns Brit. Ind. t. 181; Hk. & Baker, Syn. Fil. 81, not Hk. & Grev. Ic. Fil. t. 218. 7. um- brosum, Wall. Cat. 165. T. Kunzeanum, Hook. Sp. Fil. i. 127, t. 39 D. Himalaya; from Nepaul to Bhotan, alt. 2000-7000 feet, common. Khasia; alt. 2000- 5500 feet.—Distrib. Mergui (not from the Deccan), and scattered throughout warm and warm-temperate regions of both hemispheres. Often climbing trees to the height of 10 feet ; 1-2-pinnate, with finely divided fronds, and then very distinct from all other Indian ferns, but varies so as to be with difficulty separated from 7T. pyaidiferum on the one hand, T auriculatum on the other. Var. anceps, sp., Wall. Cat. 166. Frond smaller; stipe often winged to the base; primary segments pinnatifid, or somewhat 2-pinnatifid ; lips of the involucre slightly broader than the tube. Sikkim and Khasia, frequent.—This is separated from 7. pyxidiferum, var. limbatum, by the lips of the involucre being much shorter, and the stipe often winged to the base. Maximowiez has sent exactly the same plant from Japan marked T. radicans, Sw.? Capt. Henderson is inclined to admit it to specific rank. 6. T. AURICULATUM, Blume, Enum. Pl. Jav. Fil. 225. Frond 1-2-pinnatifid, scarcely 3-pinnatifid; main rhachis more or less winged; ultimate segments ovate entire, with flabellate nerves, or narrowly oblong, 1-nerved ; lips of the involucre scarcely wider than the tube, often altogether truncate.—Hook. Sp. Fil. i.183; Hk. & Baker, Syn. Fil.82. 7. Belangeri, Bory, in Bélanger, Voy. Bot. t. 8. fig. 1. T. dissectum, J. Smith; Hook. Sp. Fil. i. 140; Bedd. Ferns Brit. Ind. t. 182. Cephalomanes auriculatum, v. d. Bosch, Hymen. Jav. 34, t. 25. Sikkim and Bhotan; alt. 2000-7000 feet, frequent. Khasia ; alt. 3000-5500 feet, . Common. Cachar, Æ. Z. Keemam—Distrib Malaya to Japan, Guiana, ` 442 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Often scandent on trees or under damp rocks. In the Khasi common form the pinnse are often undivided with the margin nearly entire. This state is represented by Bélanger’s plate; but the fern varies, becoming more dissected till it approaches T. radicans. None of the Indian examples are however so much cut as the Javan; and the North-Indian material may be satisfactorily sorted out as between T. radicans and T. auriculatum. There is no real difference in the venation between the two; in the divided pinna of T. auriculatum the alternate segments are oblong 1-nerved. 7. T. JAvANICUM, Blume, Enum. Pl. Jav. Fil. 224. Frond simply pinnate; pinne oblong serrate or linear fimbriate.— Hk. & Grev. Ic. Fil. t. 240; Hook. Sp. Fil i 130; Hook. Garden Ferns, t. 37; Bedd. Ferns Brit. Ind. t. 180; Hk. & Baker, Syn. Fil. 83; Luerssen, Fil Graeff. 242; Benth. Fl. Austral. vii. 702. T. rigidum, Wall. Cat. 161, not Swartz. T. setigerum, Wall. Cat. 158. Cephalomanes javanicum, Zollingeri, rhomboideum, v. d. Bosch, Hymen. Jav. tt. 22, 23, 24. Cachar, R. L. Keenan. Chittagong; alt. 0-1000 feet, plentiful.— Distrib. Malay Peninsula and Islands to Australia and Polynesia, Madagascar. 12. DAVALLIA, Smith. Sect. I. Humata. Rhizome long-creeping ; stipes solitary, distant, articulated upon the F rhizome; involucre thick, coriaceous, semicircular, attached by the base only. : 1. D. PEDATA, Smith, Tentam. Gen. Fil. 15; Blume, Enum. Pl. Jay. Fil. 280; Wall. Cat. 250; Hook. Sp. Fil. i. 154, t. 45 a; Hook. Garden Ferns, t. 7; Hk. & Baker, Syn. Fil. 89; Benth, Fl. Austral. vii. 716. D. subimbricata, Blume, Enum. Pl. Jav. Fil. 231. Adiantum repens, Linn. f. Suppl. 446. Humata pedata, J. Smith; Bedd. Ferns South. Ind. t. 12. Sikkim, Dr. Jerdon.. Bhotan, Griffith. Khasia and Jaintea; alt. 4000 feet, generally scattered, not common.—Distrib. Malayan Peninsula and Islands, Ceylon, extending to North Australia, South China, and Japan, and to the Mascarene Islands. Easily separable from all other North-Indian Ferns, but closely allied to several ` Malayan. Not distinct from D. alpina, Blume, in the judgment of Col. Beddome. Sect. II. Leucostegia. Rhizome long-creeping ; stipes solitary, distant, articulated upon ` ` the rhizome (except in D. nodosa); involucre thin, membranous, semicircular OP ` ` ovate, attached by the base only. | 2 2. D. MEMBRANULOSA, Wall. Cat. 255. Scales of the rhizome subulate from a narro w : x . lanceolate base; main rhachis beneath laxly hairy.—Hook. Sp. Fil. i. 159, t. 5834; —— Hk. & Baker, Syn. Fil. 91. Leucostegia membranulosa, Bedd. Ferns Brit. Ind. t. 98. d SE Wallich. Kumaon; Moharguri Pass, alt. 6500 feet, Strachey §& Winterbottom. ` _ There are only TON three pieces of this fern in the Kew Herbarium. It is very near 5 D. excu but differs in kthe points stated. The whole frond i is more or lese, d ry MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 443 beneath. Wallich’s example is glabrous above, even on the main rhachis. Sori and involucres small, and as in D. multidentata. 9. D. MULTIDENTATA, Hk. & Baker, Syn. Fil. 91. Scales of the rhizome ovate acute ; main rhachis beneath glabrous or glandular subflocculose. Acrophorus Thomsoni, Moore, Ind. Fil. ii. 4. Microlepia pteropus, Bedd. Ferns Brit. Ind. t. 313. Aspi- dium multidentatum, Wall. Cat. 346. Himalaya; from Nepaul to Bhotan, alt. 5000-8000 feet, frequent. Khasia; alt. 4000— 5000 feet, frequent. Rhizome usually stouter and frond larger than in D. membranulosa. Ovate scales often scattered about the rhachis, especially about the base of the pinnae, as in D. nodosa and some other species. Rhachis of frond above pubescent. Frond often glandulose beneath, sometimes even to the involucres. 4. D. assamica, Hk. & Baker, Syn. Fil. 467. Scales of the rhizome ovate acute or acuminate; frond glabrous, glistening, subcoriaceous. D. micans, Mett. in Griff. no. 2790. Acrophorus assamicus, Bedd. Ferns Brit. Ind. t. 94. Bhotan; Mishmee, and by the Dihong, Griffith. : There are four sheets of this, all from Griffith’s collections, in the Kew Herbarium. In one example the scales of the rhizome are narrower, lanceolate acuminate. Involucre rather large, broader than long, much resembling that of D. immersa. 5. D. rwwERsA, Wall. Cat. 256. Frond glabrous; sori large, impressed, clearly visible from the upperside of the frond; involucres large, broader than long.—Hook. Sp. Fil. i. 156; Hook. Fil. Exot. t. 79; Hk. & Baker, Syn. Fil. 91. Leucostegia immersa, Hk. & Bauer, Gen. Fil. t. 52 a. .Aerophorus immersus, Bedd.Ferns South Ind. t. 11. Himalaya; from Mussoorie to Bhotan, alt. 3000-6000 feet; plentiful in Sikkim. Khasia ; alt. 4000-5000 feet, common. Behar; Parasnath summit, alt. 4200 feet, Sir J. D. Hooker.—Distrib. Mts. of Malabaria (rare). Malay Peninsula and Java (seems not common). | Rhizome much underground, and there without scales; the tips above ground with chestnut lanceolate-acute scales. 6. D. DAREÆFORMIS, Levinge, MS. Scales near the ends of the rhizome spreading acuminate caudate from an ovate or lanceolate base; frond finely eut; ultimate segments narrow, not very acute; involucre fugacious; sorus finally large, often broader than its segment.—D. Clarkii, Hk. & Baker, Syn. Fil. 91. derophorus Hookeri, Moore, Ind. Fil. ii. 2; Bedd. Ferns Brit. Ind. t. 95. Polypodium daree- forme, Hook. Sp. Fil. iv. 256 ; Hook. 2nd Cent. Ferns, t. 24; Bedd. Ferns Brit. Ind. t. 174; Hk. & Baker, Syn. Fil. 339. ` Gymnogrammitis, Griff. Ic. Pl. As. t. 129. ip l. ! Himalaya ; from Nepaul to Bhotan, alt. 5000-11,500 feet, frequent. Khasia; alt. 4000F- . 5500 feet, frequent.—Distrib. Moulmein. 444 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. This belongs to the section Pseudocystopteris. The involucres are sometimes across a vein, very often at the bifurcation of a vein, very rarely subterminal on a vein. 7. D. PULCHRA, Don, Prodr. Fl. Nep. 11. Scales of the rhizome adpressed, obtuse, often peltately attached ; ultimate segments of the leaves small, lanceolate, not distant, often not very acute; involucre prominent; sorus usually about as broad as its Segment D. cherophylia, Wall. Cat. 259; Hook. Sp. Fil. i. 157, t. 514; Hk. & Baker, Syn. Fil. 92. Cystopteris squamata, Dene. in Jaequem. Voy. Bot. 178. Acrophorus pulcher, Bedd. Ferns South. Ind. t. 10. Himalaya ; from Nepaul to Bhotan, alt. 2000-9000 feet, abundant. Khasia; alt. 3000- 6000 feet, common.—Distrib. Moulmein ; Mts. of Malabaria and Ceylon. Belongs to sect. Pseudocystopteris, the involucre being often at the bifurcation of a vein. . The venation, however, only very slightly tends to that of Cystopteris. The sori may be considered terminal on the veinlet, as they are slightly above the bifurcation. In the typical form the scales of the rhizome are closely sessile, attached by their middle, their whitened edges being imbricate when fresh, slightly recurved when dry. Var. pseudo-cystopteris, sp. Kunze, in Bot. Zeit. 1850, 68. Scales of the rhizome more spreading; ultimate segments often very acute.—Acrophorus pseudo-cystopteris, SE Bedd. Ferns Brit. Ind. t. 92. Cystopteris dimidiata, Dene. in Jacquem. Voy. Bot. — — t. 178. (Pl. XLIX. fig. 4.) go Himalaya; from Dalhousie to Nepaul, alt. 4000-8000 feet.—This seems only a north- west var. of D. pulchra. Some of the examples of D. pulchra collected by Sir J. D. Hooker in North Sikkim at 11,000 feet alt. seem to run into the var. D. pseudo- eystopteris. (Hk. & Baker, Syn. Fil. 92, give East Himalaya as a habitat for D. affinis, Hook. ; but I have never seen that species from North India. The Himalayan examples so named in the Kew Herbarium are D. pulchra, Don, type. Mr. Baker further adds that — i D. affinis is probably a var. of D. pulchra ; but Beddome (in Ferns South. Ind. t. 252) rightly remarks that the rhizomeis very different. The two species appear to me clearly. distinct; but they will not be so if Himalayan examples are called D. affinis.] : 8. D. noposa, Hook. Sp. Fil. i. 157. Frond large, broad, compound, usually 4-pinnatifid ; lower pinnæ often opposite or subopposite; ovate scales scattered on the rhachis, — especially at the base of the pinnsz and pinnules; ultimate segments oblong or — obovate-oblong, not acute.—Hook. Journ. Bot. 1857, t. 10; Hk. & Baker, Syn. Fil. i: E ; 92. D. stipellata, Wall. Cat. 260.. D. ligulata, Wall. Cat. 254. Aspidiwm nodosum, a T Blume, Enum. Pl. Jav. Fil. 171. A. foliolosum, Wall. Cat. 359. Acrophorus nodosus, Presl, Tent. Pterid. t. 3. fig. 2; Bedd: Ferns Brit. Ind. t. 93. bag E nodosa, Mett. in Ann. Mus. Lugd. Bat. i. 24]. E. Himalaya ; from Nepaul to Bhotan, alt. 3000—7000 feet; abundant in Sikkim. Ehasia 3 de alt. 3000-6000 feet, common.—Distrib. Malacca, Java. Rhizome underground, — creeping, the summit with lax ovate not acute scales MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 445 stipes clustered, not articulate. This fern does not resemble Alsophila except in the size of the fronds, which are not rarely 6 feet in diam. It so closely resembles Diacalpe aspidioides that, as to some fragments in over-ripe fruit, I am not sure whether they are Diacalpe or Davallia nodosa. An example with rather larger segments and sori than usual has been separated by Mr. Baker, and marked D. spheropteroides provisionally, but is perhaps only meant to be a variety. Mettenius places this fern in Cystopteris, alleging that the sori in their evolution are not truly terminal on the vein. But if on that ground this fern is to be placed in Cystopteris, it will be better to unite Cystopteris and Davallia. The present fern is unlike all other Leucostegias, and by habit, as well as by the character of the rhizome, belongs rather to sect. Microlepia. Sect. III. Hudavallia. Rhizome long-creeping; stipes solitary, distant, articulated upon the rhizome; involucre subcoriaceous, cylindric, attached by the sides as well as by the base. 9. D. DivaricaTa, Blume, Enum. Pl. Jav. Fil. 237. Scales of the rhizome long-lan- ceolate, caudate, chestnut-coloured ; involucres as long as broad.—Hook. Sp. Fil. i. 167; Hk. & Baker, Syn. Fil. 96, not of Schldl. D. polyantha, Hook. Sp. Fil. i. 168, t. 59 a; Bedd. Ferns Brit. Ind. t. 107. Sikkim, WV. S. Atkinson. Mishmee and Khasia, Griffith. Seems very rare in North India.—Distrib. Malaya, South China. 10. D. GRIFFITHIANA, Hook. Sp. Fil. i. 168, t. 49 B. Scales of the rhizome long-lan- . ceolate, caudate, white or yellowish; involucres broader than long.—Bedd. Ferns Brit. Ind. t. 106; Hk. & Baker, Syn. Fil. 96. Bhotan and Mishmee, Griffith. Khasia and Jaintea; alt. 3000-5000 feet, Griffith, Hk. f. & T., C. B. Clarke.—Distrib. South China. (Baker adds Malay Peninsula; but I do not find any examples thence at Kew.) The involucres in this species are very shortly attached on the sides, and the species . might be placed in the section Leucostegia (as in a field-note of Sir J. D. Hooker). 11. D. suLLATA, Wall. Cat. 258. Scales of the rhizome hair-pointed, from a lanceolate base, chestnut-coloured; involucres longer than broad.— Hook. Sp. Fil. i. 169, t. 50 B; Bedd. Ferns South. Ind. t. 17; Hk. & Baker, Syn. Fil. 97. D. dimidiata, Dene. in Jacquem. Voy. Bot. t. 178. Himalaya ; from Nepaul to Bhotan, alt. 2000-6000 feet, common. Khasia; alt. 2000- 5000 feet, common.—Distrib. Bombay Ghats, Wynaad, Ceylon, Malay Peninsula and Islands, South China, Japan. 2 In all these Davallias the scales of the rhizome must be taken for examination from near the growing or stipe-bearing extremities. In D. bullata the scales on the older . portions of the rhizome are sometimes ovate-obtuse, exceedingly like those of D. pul- ` chro, Don. ` ` : : K ZE 44.6 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Sect. IV. Microlepia. Rhizome very shortly creeping; stipes tufted, not articulated from the rhizome; involucre triangular or subsemicircular, not quite marginal. . 12. D. HookERIANA, Wall. Cat. 2684. Pinne narrowly oblong, acuminate, entire or crenate-serrate, not pinnatifid; ultimate veinlets parallel.—Hook. Sp. Fil. i. 172, t. 47 B; Bedd. Ferns Brit. Ind. t. 101; Hk. & Baker, Syn. Fil. 97. Sylhet, Wallich. Upper Assam, Griffith. Khasia, F. Henderson. Mikir Hills, Simons.—Distrib. South China. "m Frond thinly hispid on both surfaces; pinnze often auriculate at base on the upper ` — margin, the rhachis beneath laxly villous, not closely strigose, subhirsute. Baker and Beddome give Kumaon as a locality for this species. There is no example thence (there are only four Indian examples altogether at Kew); it seems not probable that it was ever collected there. Wallich is not safe in the matter of localities, as he in several cases mixed Kumaon and Tenasserim collections together, sorted them by hand and eye, and distributed sheets compounded of Kumaon and Tenasserim species. 13. D. PLATYPHYLLA, Don, Prodr. Fl. Nep. 10. Large, glabrous, bipinnate, tripinnatifid ; secondary pinnæ coriaceous, shining on both surfaces, lanceolate-linear, often cau- date.—Hk. & Baker, Syn. Fil. 99.. D. lonchitidea, Wall. Cat. 240; Hook. Sp. Fil. i. . 173, t. 46 B; Hook. Fil Exot. t. 19. JMicrolepia platyphylla, J. Smith; Bedd. ` ` Ferns South. Ind. t. 13. | Himalaya; from Nepaul to Bhotan, alt. 83000-5500 feet, plentiful in Sikkim. Khai alt. 3000-4000 feet.—Distrib. Mts. of Malabaria and Ceylon. Rhizome horizontal; the stipes approximate near its apex. Young frond with lax E long hairs scattered often on both surfaces; "roug frond and fully developed barren ` — frond altogether glabrous. e There is a very fine new species, D. Kurzii, C. B. Clarke, obtained by Kurz in Burma, ` — uniformly pubescent beneath, the ultimate segments triangular, subobtuse, with bluntish — sera tures, otherwise like D. platyphylla. 14. D. vnoPHYLLA, Hook. Fil. Exot. t. 19, note. Large, bipinnate, tripinnatifid, secon- — ` dary pinnze coriaceous, shining above and beneath, but pubescent on the rhachises ` beneath, lanceolate-linear, very finely cordate.—Hk. & Baker, Syn. Fil. 99, not of — Wall. Cat. 2683. Microlepia caudigera, Moore in Herb. Not Mi crolepia pe p phylla, Bedd. Ferns Brit. Ind. t. 103. | Bhotan, Griffith, nos. 1449, 2795; 4 sheets in the Kew Herbarium. (PLL) Closely allied to D. platyphylla. Ultimate segments lanceolate, secund, quite glabrous S beneath. 15. D. MARGINALIS, Hk. & Baker, Syn. Fil. 98. Pinne narrowly oblong-linear, sub- entire, crenulate or pinnatifid halfway to the midrib, but not more deeply; rha- chises beneath strigose, subhirsute; veinlets and involucres more or less villous.— D. scabra, Don, Prodr. Fl. Nep. 9. D. villosa, Wall. Cat. 244; Hook. Sp. Fil. i. MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. AAT 172, t. 484. Microlepia scabra, Bedd. Ferns Brit. Ind. t. 102. M: urophylla, Bedd. Ferns Brit. Ind. t. 103, not of Wall. Polypodium marginale, Thunb. Fl. Jap. 337. HO Nepaul and Kumaon, Wallich. Khasia, Griffith. Mikir Hills, Simons. In the Kew Herbarium only 8 sheets from India.—Distrib. Formosa, Japan. Pinne above shining, somewhat coriaceous; the midrib pubescent or hirsute; beneath the veinlets (in the dried example) are much raised, as in D. polypodioides, Don.—In the form of this species which has the pinne subentire the sori are wide apart and at some distance from the edge, the involucres hirsute; so that it is easily separated from D. Hookeriana, where the sori are in a quasi-continuous row very near the margin, the involucres glabrous. The present species is only separated here from D. polypodioides, Don, in that its pinné are pinnatifid not more than halfway down, while in the least-divided forms of D. polypodioides, Don, the pinne are divided very nearly, if not quite, to the midrib; i. e. the frond is called bipinnate. Var. calvescens, (sp.) Hook. Sp. Fil. i. 172, t. 485. .Pinnz glabrous beneath, except the strigose hirsutulous midrib.— Davallia urophylla, Wall. Cat. 2683, not of Hook. Kumaon, JZallich.—Wall. Cat. 2983, cited by Hook. l: c, is not a fern. It is perhaps through some mixture of the numbers here that D. urophylla has been supposed to grow in Kumaon. 16. D. POLYPODIOIDES, Don, Prodr. Fl. Nep. 10. Frond bipinnate, sometimes tripinnate ; rhachis of the pinnz beneath strigose hirsutulous; veinlets of the ultimate seg- ments raised beneath (in the dried examples).—Hook. Sp. Fil. i. 181, var. y and -part 3. D. Khasiyana, Hook. Sp. Fil. i. 178, tt. 47 A, 574. D. hirta, Kaulf. ; Hook. Sp. Fil. i. 181; Hk. & Baker, Syn. Fil. 100. D. strigosa, Hk. & Baker, Syn. Fil. 80. D. Roxburghii, Wall. Cat. 2218. D. rhomboidea, Wall. Cat. 257. D. pilosula, Wall. Cat. 263. D. prorima, Blume, Enum. Pl: Jav. Fil. 238. Dicksonia polypo- divides, Swartz, Syn. Fil 356. Trichomanes strigosum, Thunb. Fl. Jap. 339. Microlepia strigosa, Moore; Bedd. Ferns South. Ind. t. 255. M. hirta, Bedd. Ferns Brit. Ind. t. 256. ! H^ Himalaya, from Kumaon eastwards ; Khasia, Chittagong, abundant; most common at 3000-5000 feet alt.—Distrib. Malabaria and Ceylon, Malay Peninsula and Islands, China, Japan, Polynesia, and Tropical America. j: No Indian fern, in the opinion of W. 8. Atkinson and Major F. Henderson, is more difficult to limit or to subdivide than this. There are no good breaks in the series from D. scabra, Don, to D. flaccida, R. Br. I here follow exactly Mr. Baker, but unite his D. strigosa and hirta into one species. They differ solely in the degree to which they are divided. This will depend largely (as W. S. Atkinson has shown) on tlie age of the rhizome; the same rhizome which in its early years produces 2-pinnate fronds will in full strength produce 3-pinnate. I have D. flaccida, R. Br. type once-pinnate scarcely twice-pinnate, the fronds being in scanty fruit as usual with such from weak rhizomes. - | I separate D. polypodioides, Don, from D. flaccida, R. Br., by a combination of two J SECOND SERIES.—BOTANY, VOL. I. | 3Q 448 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. characters. D. polypodioides, Don, besides the raised veinlets giving the segments a stout plicate appearance, always has the rhachis of the pinne beneath strigose hirsu- tulous; i. e. the hairs are close together, straight, somewhat stiff, slanting forwards. Beddome’s figures, above quoted, do not show these; but I presume he has got the true plant, and that the lax sparse villousness of the rhachis in his figures is the work of his native artist. As to the varieties, which graduate completely into each other, and which are exceedingly alike in texture, venation, and the nature of the indumentum, the fol- lowing have been noted :— Var. 1. strigosa. Frond 2-pinnate, the secondary pinne not pinnatifid.— 2. strigosa, var. a, Hk. & Baker, Syn. Fil. 93. Var. 2. rhomboidea. Frond 3-pinnate; ultimate segments in their outline subentire, lightly crenate. D. rhomboidea, Wall. Cat. 257. Var. 3. pilosula. Frond 3-pinnate; ultimate segments deeply notched, often more hirsute and some thin, straight, patent hairs added. D. pilosula, Wall. Cat. 263. Var. 4. hirta. More hairy than var. 3.—Hook. & Baker, Syn. Fil. 100. Besides the foregoing are the numerous forms of fronds from young rhizomes.— Benth. Fl. Austral. vii. 717, judging from his diagnosis, from his synonymy, and from the Aus- tralian material at Kew, has included under Davallia spelunce the whole of the present D. polypodioides, Don, as well as D. flaccida, R. Br.; and I suspect that is the course which ultimately pteridologists will take. 17. D. FLACCIDA, R. Br. Prodr. 157. Frond usually 3-pinnate; rhachis of the pinne beneath with flaccid hairs, or sometimes very villose, but not strigose; ultimate segments thin, flat; the veinlets not raised beneath.—Don, Prodr. Fl. Nep. 10; Blume, Enum. Pl, Jav. Fil. 237, exl. syn. D. pilosa, Roxb. in Cale. Journ. Nat. Hist. iv. 515, t.32. D. puberula, Wall. Cat. 262. D. pyramidata, Wall. Cat. 261. D. spe- lunce, Hk. & Baker, Syn. Fil. 100; Luerssen, Fl. Graeff. 218; Benth. Fl. Austral. vii. 717. D. jamaicensis and trichosticha, Hook. Sp. Fil. i. 188. Aspidiwm pilosulum, Wall. Cat. 337, partly. .4. puberwm, Wall. Cat. 338. — Microlepia spelunce, Moore; Carr. in Fl. Viti. 340; Bedd. Ferns Brit. Ind. t. 353. Polypodium speluncæ, Linn. Sp. Pl. 1555? From Kumaon to Upper Assam and Chittagong, alt. 0-4000 feet, common. —Distrib. Deccan Peninsula and Ceylon; Malay Peninsula and Islands; and in the Tropics nearly throughout the world. Cyathevid, Griff. Ic. Pl. As. 131, i. fig. 1’, 2, 3’, 4’, Notul. i. ii. 625, perhaps belongs here. As to 131, i. fig.1, it appears from the Notul. Z. c. not to be the same; it may be Hypolepis, and, if so, not North-Indian, There are two Deccan and one Ceylon specimen of this at Kew; and Wall says it is common in Ceylon, though Beddome doubts whether it is a South-Indian fern. It must be very doubtful therefore whether it is the fern Linnseus described from Ceylon as — Polypodium spelunce. Fronds flaccid, usually large, sometimes 10 feet long. This fern ` MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 449 grows at low elevations, and is one of the few ferns that spread out a hundred miles from the hills over the plains of East Bengal, as mentioned by Roxburgh. It is thin in texture, with weak glandular pubescence, and often more or less villous also. Several varieties of this were distinguished by Wallich; but they run completely into one another. Var. 1. pubera. Rhachis of the pinne slightly glandular-pubescent beneath, or nearly glabrous.—Aspidiwm puberum, Wall. Cat. 338.—The least-common variety. Var. 2. pilosula. Rhachis of the pinne beneath with lax scattered hairs not straight. —Aspidium pilosulum, Wall. Cat. 337. Var. 3. pyramidata. Rhachis of the pinne densely villous beneath.— Wall. Cat. 261. Alsophila Grevilleana, Wall. Cat. 7075.—This is the critical form; the ultimate segments are often very villous beneath; and this state is, I find, usually ticketed D. hirta; but the texture is thin, the veinlets not raised beneath, and the pu- bescence is not strigose. | -I would not pretend to lay down the law concerning these difficult forms, con- cerning which Col. Beddome does not agree with Mr. Baker; but having collected this fern in numerous forms on many occasions, and having a large series, I am clear that D. pyramidata, Wall., runs into D. flaccida, R. Br., rather than into any form of D. strigosa or hirta, Baker. I communicated a very flaccid example of D. flaccida from young rhizomes sparsely in fruit to Col. Beddome, which he returned, saying he had no name for it. Another still weaker example (less divided) of mine I found in the D. strigosa Kew bundle. Both these were from young rhizomes among a bed of D. flac- cida typica. Sect. V. Stenoloma. Rhizome creeping ; stipes tufted, not articulated upon the rhizome ; involucres terminal on the segments, often uniting. 18. D. cuivensts, Swartz, Syn. Fil. 183; Roxb. in Cale. Journ. Nat. Hist. iv. 517; Langsd. & Fisch. Voy. Russ. t. 27; Hook. Sp. Fil. i. 187; Carr. in Fil. Viti. 338. D. tenuifolia, Swartz; Blume, Enum. Pl. Jav. Fil. 239; Wall. Cat. 245; Presl, Tentam: Pterid. t. 4. fig. 27; Hook. Sp. Fil. i. 186; Bedd. Ferns South. Ind. t. 16; Hk. & Baker, Syn. Fil. 102. Trichomanes chinensis, Linn. d: P1. 1562. Lindsaya chinensis, Mett.; Luerssen, Fil. Graeff. 224. Himalaya; from Kumaon to Bhotan; alt. 1000-4000 feet, plentiful. Khasia; alt. 1000-3000 feet, common.—Distrib. South India and Ceylon, Malaya, China, Japan, Polynesia, East-African Islands. There is no good line here between Davallia and Lindsaya. Mett, Fil. Hort. Lips. 108 says that in Davallia the veinlets are carried free to the edge, while in Lindsaya they form a lodged marginal vein near the edge. In D. tenuifolia it is common to find confluentsori. Mettenius (7. c.) left the present species in Microlepia ; but it exists in the Kew Herbarium marked “ em tenuifolia, Mett. 1; and I do not — why it is not a Lindsaya. SCH Q 450 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 18. CYSTOPTERIS, Bernh. 1. C. sETosA, Bedd. Ferns Brit. Ind. t. 312. Lasírea setosa, Bedd. Ferns Brit. Ind, t. 262. Davallia setosa, Hk. & Baker, Syn. Fil. 468. Sikkim ; alt. 5000-8000 feet, very common.—Distrib. Moulmein. Tufted, the stipes rising from nearly one point. Frond 1-3 feet; both surfaces more or less scattered with white lax hairs, whieh consist of a single row of cells varying in size. Veinlets carried straight (often undivided) beneath the sorus to the margin of the leaf. The genus must be Cystopteris if that genus is retained. Mr. Baker has probably. placed C. setosa in Davallia because of its large size. If the two genera are united, the place of C. setosa will not be in Leucostegia (where Mr. Baker has put it), but in Micro- lepia near D. spelunce. The involucre in C. setosa is small, subquadrate, white, of lax tissue, attached by the base, and at a very early stage scarcely attached elsewhere—is in all respects that of Cystopteris. The involucre is usually directly across an undivided vein, not as in D. pulchra, var. pseudo-cystopteris. 2. C. FRAGILIS, Bernh. in Schrader, Neu. Journ. ii. 27, t. 2. fig. 9; Hk. E Bauer, Gen. Fil t. 525; Hook. Sp. Fil. i. 197; Hook. Brit. Ferns, t. 23; Bedd. Ferns Brit. Ind.t. 91; Hk. & Baker, Syn. Fil. 103; Benth. Fl. Austral. vii. 752. C. fragilis, A form 12 Huteri, Milde, Fil. Europ. 149. C. retusa, Dene. in Jacquem. Voy. Bot. t. 177. North-west Himalaya ; from Kashmir and Baltistan to Kumaon, alt. 10,000—15,000 feet, not rare. Sikkim, 7. Thomson.—Distrib. Central Asia, and in almost the whole globe in cold regions. Fronds 4-6 in., sometimes a foot, always weak, glabrous. Requires to be separated with care from the small high-level Athyriwms; examples in over-ripe fruit can harig be safely distinguished. 14. LINDSAYA, Dryand. E? CULTRATA, Swartz, Syn. Fil. 119. Frond simply pinnate ; ; scales of the rhizome linear; pinne unequal-sided, lower edge nearly straight near the main nerve; veins free, or uniting only at the base of the sori.—Hk. & Grev. Ic. Fil. t. 144; Hook. Sp. Fil. i. 203; Blume, Enum. Pl. Jay. Fil. 216; Hook. Fil. Exot. t. 67; Bedd. Ferns South. Ind. t. 23; Hk. & Baker, Syn. Fil. 105; Benth. Fl. Austral. vii. 719. LZ. lucida, Blume, Enum. Pl. Jav. 216 ; Hook. Sp. Fil. i. 206 ; Wall. Cat. 145. L. gracilis, Blume, Enum. Pl. Jav. Fil. 216; Hook. Sp. Fil. i. 207. L. odorata, Roxb. in Calc. Journ. Nat. Hist. iv. 511. L. Lobbiana, Hook. Sp. Fil. i. 205, t. 620; . Bedd. Ferns Brit. Ind. t. 28. ZL. attenuata, Wall. Cat. 151. ZL. pallens, Wall. ` Cat. 148. : From Nepaul to Mishmee and Chittagong, alt. 0-4000 feet, very common.—Distrib. Mts. of the South Deccan and Ceylon, Malay Peninsula and Islands, North Australia, Formosa, Japan, East-African Islands. y . Rhizome wiry, tangled, subterranean; stipes approximate. Taking the two extreme ` MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 451 forms, var. 1 may be 2-4 in. high, growing in river-sand, the pinne small, very coriaceous and hard, all slanting towards the vertex of the frond; var. 2 in rich soil may be 18 in. high, with large herbaceous pinnz spreading horizontally. All inter- mediate forms may be found.—Lindsaya attenuata, Wall Cat. 2192, is quite remote from L. attenuata, Wall. Cat. 151, and comes from Herb. Finlayson, probably collected in Siam or Cochin-China. 2. L. REPENS, Bedd. Ferns South. Ind. tt. 209, 214. Frond simply pinnate; scales of the rhizome lanceolate ` pinnee unequal-sided, lower edge nearly straight near the main nerve; veins free, or uniting only at the base of the sori—JZ. pectinata, Blume, Enum. Pl. Jav. Fil. 217; Hook. Sp. Fil i. 207; Hk. & Baker, Syn. Fil. 106. L. oblongifolia, Hook. Sp. Fil. i. 206, t. 61. Lindsaya, sp., Griff. Ic. Pl. As. 115 v, Notul.i.ii. 614. Davallia Boryana, Presl; Hook. Sp. Fil.i.175; Hk. & Grev. Ic. Fil. t. 143. D. hemiptera, Hook. Sp. Fil. i. 176. ZL. repens, Desy.; Hk. & Baker, Syn. Fil i. 93. Dicksonia repens, Swartz, Syn. Fil. 188. Odontoloma repens, J. Smith; Hk. & Bauer, Gen. Fil. t. 1145. O. Boryanum, Fée, Gen. Fil. t. 26 a. fig. 2. Mishmee and Khasia, Griffith. Sikkim Terai; Dulkajhar, alt. 1000 feet, N: Gamble.— Distrib. Ceylon, Malay Peninsula and Islands, Polynesia, Mauritius. Rhizome stout, usually climbing. Fronds 12-18 in., very much like the large form of Z. cultrata, but the rhizome and scales differing —This is a very rare fern in North India, having apparently been collected but on three occasions. The locality, Nilgherries, given for it by Mr. Baker, is objected to by Col. Beddome; and I can find no Nilgherry example at Kew. Lindsaya pectinata and Davallia repens of Hk.& Baker, Syn. Fil., are so identical that I imagine the two are only retained in the text because stereotyped. Both species are attributed to North India on the faith of Griffith’s three sheets, two of which (collected from the same plant?) are arranged one in the Davallia repens bundle, the other in the Lindsaya pectinata bundle. Baker doubts whether L. scandens, Hk., is distinct ; but Mr. Baker has marked one sheet L. pectinata, which I consider typical L. scandens, and this, of course, spoils all. There remains the question whether the species should be called Lindsaya or Davallia. I should resolve this by transferring the whole section Odontoloma to Lindsaya. e 3. L. FLABELLULATA, Dryand. in Trans. Linn. Soc. iii. 41, t. 8. Frond bipinnate or simply pinnate; veins free; pinnæ without a distinct midrib, flabellulate-veined, curved or . excised on the lower margin.—Hk. & Grev. Ic. Fil. t. 75; Hook. Sp. Fil. i. 211; Bedd. Ferns South. Ind. t. 216; Hk. & Baker, Syn. Fil. 107; Benth. Fl. Austral. vii. 720. . L. tenera, Dryand. in Trans. Linn. Soc. iii. 42, t. 10; Wall. Cat. 146; Hook. Sp. Fil. i. 211; Bedd. Ferns South. Ind. t. 24. L. striata, Blume, Enum. Pl Jav. Fil. 220. D. polymorpha, Wall. Cat. 147; Hk. & Grev. Ic. Fil. t. 75. L. interrupta, Wall. Cat. 2195. Vittaria interrupta, Roxb. in Cale. Journ. Nat. Hist. iv. 511. Davallia trichomanoides, Bedd. Ferns Brit. Ind. t. 178, not of Blume. D. schizophylla, Hk. & Baker, Syn. Fil. 468.. 4.52 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Khasia and Assam, Griffith. Jaintea; Jarain, alt. 4000 feet, C. B. Clarke.—Distrib. Mts. of South India and Ceylon, Malay Peninsula and Islands, South China, "— Australia. d Stipes tufted, sometimes strongly dimorphic; the barren fronds 2-3 in., simply piiniat; on stipes 1 in. ; the fertile fronds 6-8 in., more or less compound, on stipes 4-6 in. The simply pinnate form may be passed in the field for L. cultrata, and is perhaps not rare in North India; but it appears to have been collected there on three occasions only. 4. L. ENsIFOLIA, Swartz, Syn. Fil. 118. Frond usually pinnate, with entire narrow lanceolate pinna; veinlets reticulating —Hk. & Grev. Ie. Fil. t. 111; Hook. Sp. Fil. i.. 220; Hook. Garden Ferns, t. 62 ; Hk. & Baker, Syn. Fil. 112; Luerssen, Fil. Graeff. 226; Benth. Fl. Austral. vii. 721. Z. lanceolata, Labill. Pl. Nov. Holl. ii. 98, t. 248. LZ. Griffithiana, Hook. Sp. Fil. i. 219, t. 68 B. LZ. pentaphylla, Hook. Sp. Fil. i. 219, t. 67. . L. pteroides, Wall. Cat. 2198. L. longipinna, Wall. Cat. 2194. Schizoloma ensifolium, Bedd. Ferns South. Ind. t. 25. S. Griffithianum, Bedd. Ferns Brit. Ind. t. 20. Pteris angustata, Wall. Cat. 93. From Sikkim to Muneypoor and Chittagong, alt. 500-4000 feet, frequent.— Distrib. South Deccan and Ceylon, Malay Peninsula and Islands to Polynesia and North Australia. Tropical and Southern Africa with its eastern islands. Rhizome creeping somewhat widely. Pinnz often few, sometimes one only, rarely with long lobes. There is very little of this at Kew from North India; but it is by no means rare. Collectors have perhaps often passed it supposing it to be the common Pteris cretica.—As to the name, Mettenius keeps up Labillardiere's L. lanceolata, which was published in the same year as Swartz's. I cannot discover which has the right of priority, and therefore follow Mr. Baker. | [Lindsaya Finlaysoniana, Wall. (i. e. L. heterophylla, Hk. & Baker, Syn. Fil. 112, partly), is stated by Beddome (Ferns Brit. Ind. Suppl. p. 6) to grow in North India; buf there is no example further north than the Nilgherries in the Kew Herbarium, and I suspect some error. | 15. ADIANTUM, Linn. ~ * Simply pinnate. 1. A. LUNULATUM, Burm. Fl. Ind. 235. Pinnæ glabrous, their petioles usually 4-} in- or more.—Hk. & Grev. Ic. Fil. t. 104; Wall. Cat. 77; Blume, Enum. Pl. Jav. Fil. 215; Hook. Fil. Sp. ii. 11; Carr. in Fl. Viti. 346; Bedd. Ferns South. Ind. t. 15 Milde, Fil. Europ. 28; Hk. & Baker, Syn. Fil. 114; Benth. Fl. Austral. vii. 723. A. dolabriforme, Hook. Ie. Pl.t. 191. A. filiforme, Hook. Ic. PL t. 503. Pteris lunulata, Retz. Obs. ii. t. 4; Roxb. in Cale. Journ. Nat. Hist. iv. 506. Throughout North India, in moist places, alt. 0-4500 feet; very common.—Distrib. South India and Ceylon, Malay Peninsula, the tropics of nearly the whole world. One of the most generally diffused of Indian Ferns, plentiful in ditches in Calcutta. In the hills it has sometimes large pinne 1 in., sometimes small } in. which are occa- MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 458 sionally nearly orbicular.—A. soboliferum, Wall. Cat. 74 (from Ava), referred here by Baker /. e does not belong; it is nearer 4. caudatum. 2. A. CAUDATUM, Linn. Mant. 308. Pinne hairy, their petiole rarely exceeding 4 in.— Roxb. in Calc. Journ. Nat. Hist. iv. 512; Hook. Exot. Flora, t. 104; Hook. Sp. Fil. ii. 18; Bedd. Ferns South. Ind. t. 2; Milde, Fil. Europ. 29; Hk. & Baker, Sp. Fil. 115. A. vestitum, Wall. Cat. 75. A. flagelliferum, Wall. Cat. 76. A. proliferum, Roxb. in Calc. Journ. Nat. Hist. iv. 512. A. hirsutum, Wall. Cat. 2176. A. cili- atum, Blume, Enum. Pl. Jav. Fil. 215. Throughout North India, alt. 0-3000 feet, very common.—Distrib. South India and Ceylon, Malay Peninsula and Islands to South China, Tropical Africa to the Verdes. Var. rhizophorum, Wall. Cat. 82. Pinnæ and stipe glabrous, or nearly so.— Hook. Sp. Fil. ii. 12, t. 80 A. 4. Edgworthii, Hook. Sp. Fil. ii. 14, t. 81 B; Bedd. Ferns Brit. Ind. t. 17. Gurwhal, Edgeworth, A. Hume, Hope. Nepaul, Wallich. Mooltan, Edgeworth. ** Frond usually 2- (or 8—4-) pinnate. 9. A. CAPILLUS-VENERIS, Linn. Sp. Pl. 1558. Frond usually 2-pinnate; segments very thin.—Engl. Bot. t. 1564; Wall. Cat. 73; Hk. & Bauer, Gen. Fil. t. 66 5; Hook. Sp. Fil ii. 36, t. 74 5; Hook. Brit. Ferns, t. 41; Bedd. Ferns South. Ind. t. 4; Milde, Fil. Europ. 30; Hk. & Baker, Syn. Fil. 123; Benth. Fl. Austral. vii. 728. A. ca- pillus, Wall. Cat. 73. A. tenerum, Roxb. in Cale. Journ. Nat. Hist. iv. 513, not of Swartz. Chittagong; Barobkoondo and Seetakoondo, J. Scott, C. B. Clarke. Khasia, Assam, Sikkim, rare. From Nepaul to Kafaristan, plentiful.— Distrib. Malabaria, from Bombay to Ceylon (rare) ; from Cabul to England and Morocco; in tropical and temperate Africa and America; Queensland. 4. A. vENUsTUM, Don, Prodr. Fl Nep. 17. Frond 3-4-pinnate; segments somewhat rigid.—Wall Cat. 81; Hook. Sp. Fil. ii. 40, t. 768; Bedd, Ferns Brit. Ind. t. 20; Hk. & Baker, Sp. Fil. 125. A. microphyllum, Roxb. in Cale. Journ. Nat. Hist. iv. 513. : . Nepaul, Wallich, Gurwhal to Kashmir, alt. 3000-10,000 feet, abundant.—Distrib. Cabul. One of the commonest ferns of the North-east Himalaya. Sir W. J. Hooker gives also Khasia as a locality; but I find no example thence, and Mr. Baker confines the species (I believe correctly) to the Himalaya. *** Fronds dichotomous, 5. A. PEDATUM, Linn. Sp. Pl. 1557. Rhachis glabrous, once-forked, the pinns placed scorpioid-like on each fork.—Schkuhr, Crypt. t. 115; Hook. Sp. Fil. ii. 28; Bedd. Ferns Brit. Ind. t. 167; Milde, Fil, Europ. 31; Hk. & Baker, Syn. Fil. 126. ^ 454 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. North-west Himalaya, alt. 6000-9000 feet, scattered, not plentiful; from Gurwhal (Strachey & Winterbottom) to Sikkim (Sir J. D. Hooker).—Distrib. Japan, North America (nearly the whole). 6. A. FLABELLULATUM, Linn. Sp. Pl. 1557. Scales on the rhizome long, linear, lax, chestnut-coloured ; rhachis hairy, repeatedly dichotomous; segments glabrous; sori often 45; in. broad.— Hook. Sp. Fil. ii. 30; Bedd. Ferns South. Ind. t. 218 ; Hk. & Baker, Syn. Fil. 126, not of Wall. 4. amenum, Wall. Cat..78; Hk. & Grev. Ic. Fil. t. 103. Nepaul, Wallich. Assam, Simons. Khasia, Hk. f. & T., T. Lobb. Sylhet, Hk. f & T. —Distrib. Ceylon, Malay Peninsula and Islands, South China, and Japan. In A. hispidulum, Swartz (A. flabellulatum, Wall. Cat. 2177, not of Linn.), the scales on the rhizome are lanceolate, shining black, rigid, adpressed; the sori scarcely Ze in. broad. 16. CHEILANTHES. * Fronds not powdered beneath ; stipe and main rhachis scaly and hairy. 1. CH. FRAGRANS, Swartz, Syn. Fil. 127, 325, 326, t. iii. fig. 6. Frond bipinnate, tripin- natifid, or tripinnate; ultimate segments: $ in. in diam., ovate or round, without hairs beneath.— Webb & Berth. Phyt. Canar. iii. 452; Mett. Ferngatt. Cheil. 38; Hook. Sp. Fil.ii. 81; Milde, Fil. Europ. 34; Bedd. Ferns Brit. Ind. t. 338; Hk. & Baker, Syn. Fil. 134. Ch. odora, Swartz, Syn. Fil. 127, 327 ; Schkuhr, Crypt. t. 123. Ch. sua- veolens, Swartz, Syn. Fil. 127; Sibth. Fl. Greec. t. 966. Polypodium fragrans; Linn. Mant. 307, not Sp. Pl. 1550. Mts. S. of Kashmir; Murree, alt. 4000-5000 feet, H. QC. Levinge. Kishtwar, alt. 3500-5000 feet, C. B. Clarke.—Distrib. Cabul, all round the Mediterranean, Ma- caronesia. 3 The partial rhachises are often scaly and hairy with chestnut submoniliform hairs; but the surface of the frond beneath is entirely devoid of the plentiful white curled woolly hairs so abundant in the next species.—Mr. Baker carefully follows Sir W. J. ` Hooker in stating that the present species is not Swartz's Cheilanthes fragrans, while they both say it is Polypodium fragrans of Linnzeus's * Mantissa., Swartz (Syn. Fil. 326) states not only that his Ch. fragrans is Linnsus's Polypodium fragrans, but that to prevent confusion he obtained, through Sir J. E. Smith, from Linneus’s Herbarium the ipsissimum exemplum collected by Koenig on which Linnzus founded the species. 1 E. suspect that the whole blunder arose because Webb and Berth. supposed Kcenig’s ex- ample to have come from India, and that therefore their own Macaronesian fern would — be something different. Sir W. J. Hooker (Sp. Fil. ii. 94) says that Swartz’s Ch. fra- = grans was of Indian origin, gathered by Koenig probably in the Madras Peninsula; ` ` but on turning to Linn. Mant. 307 it will be seen that Kcenig’s specimen was oci | not in India, but on the walls of Funchal, m it grows now. Ty ue 2. Cn. SzovrrZu, Fisch. & Meyer, in Bull. Soc. Mose. 1838, 241. Frond bipinnate, tri- pinnatifid or — ultimate segments } in. in diam., round, with miang crisped À MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 455 white hairs beneath.—Hook. Sp. Fil. ii. 98, t. O4 B ; Mett. Farngatt. Cheil. 37; Milde, Fil. Europ. 33; Bedd. Ferns Brit. Ind. t. 145; Hk. & Baker, Syn. Fil. 139. Ch. fimbriata, Vis. Fl. Dalm. t. 1. fig. 1. Notholena persica, Bory in Bélang. Voy. 23. Kashmir and Baltistan; alt. 5000-7500 feet, frequent. Kulu, ZEdgeworth.—Distrib. Cabul, West Asia, South Europe. Exceedingly like Ch. fragrans, and only to be distinguished by the indusial hairs. The hairs are really confined to the sori, which occupy a large portion of the very small segments, so that the lower surface of the frond appears densely matted. Ch. Szovitzii is the oldest name. Notholena persica is 4 years later than the full description of Oh. Szovitzii by Fisch. & Meyer. ** Fronds not powdered beneath ; stipe not hairy, sometimes slightly scaly. 9. CH. VARIANS, Hook. Sp. Fil. ii. 89, t. 103 a. Glabrous; frond long-lanceolate, pin- nate; pinnz subentire, pinnatifid or with a few secondary pinnæ ; involucres attain- ing §-+ in.—Bedd. Ferns South. Ind. t. 189; Hk. & Baker, Syn. Fil. 127. Pteris varians, Wall. Cat. 86. - Pt. cespitosa, Wall. Cat. 90. - East Bengal Plain from Assam to Chittagong, general on red Terai soil: ascending the Khasi range to 2000 feet alt.—Distrib. Moulmein, Ava, South China, Luzon. Ana- mallays (Beddome). This is a fern of the plains; common at Dacca. Mr. Baker says its locality is the Himalaya ; but I never saw it there, nor is there any example thence at Kew. Mr. Baker also does not admit Col. Beddome's locality in Malabaria; but Col. Beddome's figure above quoted is so exactly the plant, that I suppose Mr. Baker suspected that Col. Bed- dome did not prepare it from his Anamallay plant. 4. CH. TENUIFOLIA, Swartz, Syn. Fil. 129, 332. Glabrous; frond triangular-lanceolate, 2-3-pinnate, secondary pinnz numerous; involucres rarely exceeding 3 in. (though often confluent in age).—Schkuhr, Crypt. t. 125; Blume, Enum. Pl. Jav. Fil. 137; Hook. Sp. Fil. ii. 82, t. 87 c; Carr. in Fl. Viti. 347; Mett. Farngatt. Cheil. 27; Bedd. Ferns South. Ind. t. 188; Hk. & Baker, Syn. Fil. 138; Benth. Fl. Austral. vii. 726. Ch. micrantha, Wall. Cat. 68, as to type sheet. Ch. rupestris, Wall. Cat. 07. Ch. Sieberi, Kunze; Hook. Sp. Fil. ii. 83, t. 97. Pellea nudiuscula, Hook. Sp. Fil. ii. 151. Pteris gracilis, Roxb. in Cale. Journ. Nat. Hist. iv. 508. Tricho- manes tenuifolia, Burm. Fl. Ind. 237. : Bengal Plain; alt. 0-2000 feet, common on red soil, as iñ Assam, Chittagong, Dacca, and throughout Chota Nagpore: ascends the Khasi range to 3500 feet alt. Sikkim, alt. 1000 feet.—Distrib. Deccan and Ceylon, Malay Peninsula and Islands, extending to China, Australia, New Zealand, Polynesia, Uruguay. This fern is also said by Mr. Baker to be confined in North India to the dire, there is no example from the Himalaya in the Kew Herbarium, but I have collected it in Sikkim. ‘Wall. Cat. 68 has Pellea nitidula, Baker, mixed with it. There isa marked SECOND SERIES.—BOTANY, VOL. I. 3n 456 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. difference in cutting between the Decean and East Bengal examples; the figure of Sir W. J. Hooker represents the East Bengal type; the figure of Col. Beddome represents the Chota Nagpore type. Chota Nagpore is, phytographically as well as geographically and geologically, the north-east extremity of the Deccan Plateau. SE. *** Fronds with some powder or hair beneath (but Ch. farinosa, var. Dalhousie, often denudate when old). | 5. CH. SUBVILLOSA, Hook. Sp. Fil. ii. 87, t. 98B. Frond elongate-lanceolate; the lowest pair of pinnz distant, narrower than the pair above; stipe shorter than the frond, glabrous, with a few broad-lanceolate uniform-coloured scales near the base ; rhachis of primary pinne with crisped woolly salmon-coloured hairs beneath, but no scales; involucre continuous, slightly crenulate, not fimbriate on the margin.—Mett. Farn- gatt. Cheil. 48; Bedd. Ferns Brit. Ind. t. 142; Hk. & Baker, Syn. Fil. 137. North-west Himalaya; Pabur Valley and Kotghur, near Simla, Zdgeworth. Scales of the rhachis light-chestnut-coloured or scarious. Main rhachis glabrous beneath, the partial rhachises woolly. Pinne often 8-10 pairs, separate, the lower 2 in. apart. Involucre as in Pellea, to which this species might be referred. In this and the neighbouring species of Cheilanthes the margin of the frond is at first reflexed, con- tinuous, becoming in the age of the fruit beaded or broken up; in Pellæa the margin is continuous in age; the distinction is very small.—Col. Beddome expresses an opinion that this species is only a form of Ch. farinosa; but he has perhaps never seen it, his figure being taken from Sir W. J. Hooker’s. There is no powder on the examples, of which, however, there are but 2 sheets at Kew. 6. CH. ALBO-MARGINATA, C. B. Clarke. Frond lanceolate; the lowest pair of pinn® usually more developed than any superior pair; stipe shorter than the frond, glabrous, with many lanceolate white-margined scales; pinnz, when young, densely scaly beneath, often with yellow powder also, but not hairy, when old usually scaly beneath on the rhachises; involucres peltate, usually lacerate on their margins. (Pl. LII.) North-west Himalaya, Falconer. Kashmir; Basaoli, alt. 5000 feet, C. B. Clarke. Dalhousie, alt. 6000 feet, C. B. Clarke. Simla, alt. 7000 feet, T. Thomson. Gurwhal, alt. 2000-9000 feet, H. C. Levinge.—Distrib. Nilgherries, fide Mojor F. Henderson. Scales on the lower part of the stipe numerous, lanceolate-linear, secund, ascending, somewhat rigid, dark chestnut, nearly black in the centre, their margins glistening white; tufts of hair-pointed scales on the rhizome. Young fronds usually entirely thickly covered beneath with lanceolate chestnut coloured scales, and generally with ` yellow powder, in which state it has been confused with Ch. rufa; but it has none of — the woolly hairs in which Ch. rufa always abounds. This is probably what Col. Beddome means when when he says (Ferns, Suppl. p. 7) that “ he has lately obtained De traces of it are sometimes permanent —1 haye collected large quantities of this Ka as P has H. C. Levinge. Bee MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 457 7. Cu. RUFA, Don, Prodr. Fl. Nep. 16. Frond lanceolate; lowest pair of pinne usually smaller than the next superior pair; stipe and whole frond beneath woolly with crisped hairs; margin of involucre much fimbriate.—Hook. Sp. Fil. ii. 79, t. 994; Mett. Farngatt. Cheil. 47; Bedd. Ferns Brit. Ind. t. 144; Hk. & as Syn. Fil. 141. Ch. tomentosa, Herb. Wall. ` Khasia; alt. 4000 feet, plentiful wherever there is limestone. Sikkim; alt. 5000 feet, rare (as is limestone). Gurwhal, alt. 2000 feet, H. C. Levinge.—Distrib. Mergui. I have collected much of this fern, but only on limestone ; it is generally closely pro- cumbent, curling up on the rock, and easily recognized by its woolly hairiness. Scales often none, or undistinguishable from the hairs; scales, when present on the stipe mixed with the hairs, are narrow-linear uniform-coloured. Fronds above laxly flocculose or woolly, or almost tomentose.—Col. Beddome remarks (Ferns, Suppl. p. 7) that Wallich was right in calling this plant Ch. farinosa, var. vestita; but Wallich calls it Ch. tomen- tosa in his Herb., and some other hand (apparently) has directed it to be put * with 71," i.e. Ch. farinosa.. 8. Cu. FARINOSA, Kaulf, Enum. Fil 212. Frond triangular-lanceolate or long-lanceo- late; lowest pair of pinnæ often distant, as much developed as any of the superior pinne ; stipe shorter than the frond, glabrous, with some lanceolate-linear uniform- coloured scales near the base; pinnse without hairs beneath, rarely with a few scales, always more or less powdered ; involucre usually toothed, sometimes lacerate. —Hk. & Grev. Ic. Fil. t. 134; Hook. Sp. Fil. ii. 77; Bot. Mag. t. 4765; Mett. Farngatt. Cheil. 46; Bedd. Ferns South. Ind. t. 191; Hk. & Baker, Syn. Fil. 142. Ch. dealbata, Don, Prodr. Fl. Nep. 16; Wall. Cat. 71, as to type sheet. Ch. rigi- dula, Wall, Cat. 2175. Ch. bullosa, Bedd. Ferns South. Ind. t. 199. Pteris farinosa, Forskh. Fl. ZEgypt. Arab. 187. Pt. bicolor, Roxb. in. Calc. Journ. Nat. Hist. iv. 507. Throughout Northern India, in the hills, alt. 0—5000 feet from Kashmir to Assam gnd Chittagong; also in Chota Nagpore.—Distrib. Whole Deccan and Ceylon; Eastern Africa with its islands and Arabia; Java and the Philippines; Tropical America. This species is easily separable from Ch. rufa and Ch. albomarginata above. I cannot separate it satisfactorily from Ch. argentea, var. chrysophylla. All the India examples are white- or pale-yellow-powdered below, except a very large dark-green form sent by Mr. Batcock from Ootacamund, marked var. concolor. | Var. Dalhousie (sp.), Hook. Sp. Fil. ii. 10. Pinnæ (even when young) without hairs, scales, or powder beneath; involucres deeply crenulate, toothed or lacerate on the margin.—Hk. & Baker, Syn. Fil. 137. (Pl. LI) Western Himalaya, from Kashmir to Kumaon, alt. 6000-9000 feet, frequent. Sikkim; Lachen, alt. 10,000 feet, Sir J.. D. Hooker.—Hook. Sp. Fil. ii. t. 78 B, is named Ch. de- albata, but quoted at page 80 for Ch. Dalhousie; but Mr. Baker has withdrawn it, as it is taken from a sheet of examples containing a mixture of forms.—Bedd. Ferns South. Ind. t. 193, does not show the rhizome and lower scales; nor is the cutting that of Ch. Dathousia ; all Col. Beddome’s examples of Ch. Dathousie EE from the 3R 458 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Deccan exhibit more or less powder, and I should call them Ch. farinosa type. Sorting the specimens my own way, I fancy that I can keep Ch. Dalhousie distinct; but I bow to the opinion of Baker and F. Henderson, and rank it as a variety. Var. chrysophylla, Hook. Fil. Exot. t. 95. fig. 1. Frond triangular-lanceolate, with sometimes 6-8 pairs of free pinnze; powder beneath golden, never white, sometimes scanty.— Ch. chrysophylla (sp.), Hook. Sp. Fil. ii. 113; Hook. Ic. Pl. t. 901; Mett. Farngatt. Cheil. 47. Khasia; alt. 5500 feet, not rare. Sikkim, W. S. Atkinson.— This is a most doubtful plant, that has been confused with Ch. argentea, var. sulphurea, to which it bears little resemblance except in having yellow powder beneath. Both W. 8. Atkinson and Major F. Henderson think it should be appended to Ch. farinosa rather than to Ch, argentea. 9. CH. ARGENTEA, Kunze in Linnea, 1850, 242. Frond triangular; lowest pair of pinne most developed, approximate; remainder of the frond pinnatifid or with one other free pair of pinne; stipe glabrous, often longer than the frond, with sublinear scales near the base; pinne glabrous beneath, but with white powder; involucre crenate, rarely much toothed.—Hook. Sp. Fil. ii. 76; Mett. Farngatt. Cheil. 45; Bedd. Ferns Brit. Ind. t. 143; Milde, Fil. Europ. 37; Hk. & Baker, Syn. Fil. 142. Pieris argentea, Gmel.; Langsd. & Fisch. Pl. Voy. Russes, t. 22. Khasia; alt. 3000-5500 feet, frequent.—Distrib. Mergui, Northern Asia to Siberia, Japan, and Kamtschatka. : In my typical example the stipe is 84 in., the frond 31; free pinne one pair only, very white beneath. Var. sulphurea, Hook. Fil. Exot. t. 95. fig. 3. Exactly as in the type, but the powder often yellow or golden. Khasia and Mergui; growing sometimes on the same rhizome with white-powdered fronds. 17. Onycuium, Kaulf. 1. O. AvRATUM, Kaulf. Enum. Fil. 144. Coriaceous, shining on both surfaces; barren ` frond sub-4-pinnate, fertile frond 3-pinnate; some of the involucres often exceeding ` ` i in.; ripe capsules golden.—Hook. Sp. Fil. ii. 121; Fée, Gen. Fil. 131, t. 7. fig. C; Bedd. Ferns South. Ind. t. 30; Hk. & Baker, Syn. Fil. 143. Lomaria aurea, Wall. — Cat. 38. ZL. caruifolia, Wall. Cat. 39. L. decomposita, Don, Prodr. Fl. Nepal. 14. Pteris chrysocarpa, Hk. & Grev. Ic. Fil. t. 107. From Nepaul to Assam and Chittagong, alt. 0-4000 feet, common: sometimes found ` far from the hills in East Bengal.—Distrib. Malay Peninsula and Islands, New Guinea. ` In large tufts. Scales numerous at the base of the stipe, linear, chestnut-coloured. ` Involucres on the terminal segments much longer than the others.—The form L. carui- Jolia, with shorter sori, less golden capsules (from Amherst) is marked by Kurz as 9. ` var. of O. japonicum, to which view Capt. F. Henderson inclines. Mr. Baker agrees with — me that it is better placed with O. auratum. t aa MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 459 2. O. JAPONICUM, Kunze, in Schkuhr, Fil. Suppl. 11. Coriaceous, shining on both surfaces, especially the upper; fertile frond often 4-pinnate, as much divided as the barren ; no involucre attaining 4 in. ; ripe capsules deep brown, numerous, broader than the segment, and forcing the involucre completely open.—Hook. Sp. Fil. ii: 122; Hk. & Baker, Syn. Fil. 143. O. lucidum, Spreng.; Hk. & Bauer, Gen. Fil t. 11; Hook. Sp. Fil. ii. 121. O. capense, Kaulf. Enum. Fil. 145, t. 1. fig. 8. Cheilanthes lucida, Wall. Cat. 69. Trichomanes japonicum, Thunb. Fl. Jap. 340. From Gurwhal to Mishmee and Khasia, alt. 3000—6000 feet; common from Nepaul eastward.—Distrib. Ava, China, Japan. Tufted. Scales at the base of the stipe numerous, linear, pale brown. Var. multisecta, (sp.) F. Henderson, MS. Fertile frond very finely cut, often 5-pinnate ` ripe capsules straw-coloured, not numerous; involucre remaining closed over the ripe capsules. Cheilanthes contigua, Wall. Cat. 72. Leptostegia lucida, Don, Prodr. Fl. Nepal. 14. From Kumaon to Kashmir, alt. 6000-8000 feet, frequent.— Frond herbaceous, hardly shining, not coriaceous.—This is more easily separated from O.japonicum type than is O. auratum, and has been estimated a species both by Wallich and F. Henderson, to which opinion the area lends support. But if it is estimated a species, I do not know to which the next variety should be attached. Var. intermedia. Frond lax, more coarsely cut ; involucres often 1 in.— O. lucidum, Bedd. Ferns Brit. Ind. t. 21. Kumaon ; alt. 7500 feet. Sikkim; Lachen, alt. 9000—1000 feet, Sir J. D. Hooker.— This form, exactly figured by Col. Beddome, seems halfway between O. japonicum, Kunze, and O. multisectum, F. Henderson. [After going through the Kew material with me, F. Henderson would still prefer to make O. multisectum a species.] 18. CnyPTOGRAMME, R. Br. 1. C. crispa, R. Br. in Richardson's Append. to Franklin's First Journal, 54.—Hk. & Bauer, Gen. Fil. t. 115 5 ; Hook. Sp. Fil. ii. 128-130; Hook. Brit. Ferns, t. 39; Hk. & Baker, Syn. Fil. 144.. C. Brunoniana, Wall. Cat. 396; Hk. & Grev. Ic. Fil t.158 ; Bedd. Ferns Brit. Ind. t. 164. C. acrostichoides, Hk. & Grev. Ic. Fil.t.29. Allosorus crispus, Bernh. in Schrad. neu. Journ. Bot. ii. 36; Milde, Fil. Europ. 28. Phoro- lobus crispus, Desy.; Fée, Gen. Fil. 130, t. TD. Pteris crispa, Linn. MS.; Engl. Bot. t. 1160. Osmunda crispa, Linn. Sp. Pl. 1522. : From Kumaon to Kashmir and Baltistan ; alt. 10,000—15,000 feet, plentiful. Sikkim ; alt. 10,000-14,000 feet (head of Lachen valley), Sir J. D. Hooker.—Distrib. Arctic and Alpine Europe, Asia and North America. Glabrous, tufted ; scales at the base of the stipe lanceolate acute, pale brown ; often a few ovate-lanceolate acute similar scales scattered on the lower part of the stipe. Fertile frond wholly fertile, i. e. not with the lowest pinnæ barren.—1 can see no difference between the Himalayan and European plants, nor can I distinguish any Himalayan variety. Milde says the Himalayan form has the barren fronds with the 460 MR. C. B. CLARKE ON THE FERNS OF NORTHERN. INDIA. ultimate segments more acutely serrate; but I suspect Milde's stock of Himalayan material on which he ventured this distinction was small. I have collected the plant more than twenty times between Dhurmsala and the Karakorum. None resemble the American var. acrostichoides—The name of this fern is a queest. vex. Most writers prefer Allosorus crispus, Bernh., who first separated the species from Pteris; but Allosorus has been so variously understood that the genus has been altogether dropped by Sir W. J. Hooker. 19. PELLÆA, Link. 1. P. eracus, Hook. Sp. Fil ii. 138, t. 133 B. Glabrous; rhizome wiry, creeping; . stipes not tufted; fronds 1-2-pinnate, often with the lowest pinne barren, the upper fertile ; young sori subterminal on the veins, clustered, not forming a marginal line till old—Hk. & Baker, Syn. Fil. 145. Sect. IV. Doryopteris. Veins copiously anastomosing, without free included veinlets. Fronds lobate pinnatifid, or scarcely 1-pinnate. 17. P. LUDENS, Wall. Cat. 88. Rhizome creeping, with linear adpressed scales, dark chestnut with whitened margins; stipes solitary, distant.— Hook. Sp. Fil. ii. 210; Hk. & Baker, Syn. Fil. 166. P. multifida, Roxb. in Cale. Journ. Nat. Hist. iv. 567, not of Poir. Litobrochia pedata and L. ludens, Bedd. Ferns Brit. Ind. tt. 26, 27, excluding synonyms. Chittagong Hills, alt. 0—1000 feet, Roxburgh, C. B. Clarke. Orissa; Balasore Hin Blanford.—Distrib. Malay Peninsula, Ava. P. Stipe often with dusky subtomentose pubescence at benc and apex, sometimes with a & few scales. Frond glabrous beneath, with shining ribs.—Col. Beddome (in Brit. Ferns, |. Suppl. p. 8) proposes, as I understand him, to unite P. ludens and P. palmata undet ` P. pedata, Linn. ; but, as Sir W. J. Hooker pointed out (Syn. Fil. ii. 211), all the Burmese ` - P. ludens has a long creeping rhizome, whereas (in the large Kew collection of P. pal- mata and pedata) the stipes are in every case closely tufted. As to Col. Beddome's fig- — of Litobrochia pedata, it does not show the rhizome, but was taken from a Moulmein. Le specimen, where P. ludens is common, P. pedata unknown. E Wallich’s example from Dindighul marked Pteris mysurensis, referred MH Hooker x .MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 411 P. pedata, by Baker to P. palmata, consists of a fertile frond without the lower part of the stipe. It might be P. ludens; but the cutting is that of P. palmata, Hook. Garden Ferns, t. 22, or P. pedata, Hook. Fil. Exot. t. 34. There is in Wight's Peninsula Ind. Or. Herbarium a complete example, marked Doryopteris sagittifolia, which has the fronds closely tufted, and resembling in form Pteris sagittifolia, Raddi, Hook. Fil. Exot. t. 39; but the stipe is plentifully covered with long bright chestnut hairs, which extend to the ribs of the frond, and even over its surface beneath. In this scarcity of material from the Deccan the P. pedata of South India must remain obscure: P. ludens; of Chittagong, Burma, and Ava is completely known. Sect. V. Litobrochia. Veins copiously anastomosing, with some free included veinlets. Fronds 2-pinnate or still further divided. 18. P. Gerten, Thunb. Prodr. Fl. Capens. 171; Blume, Enum. Pl. Jav. Fil. 212; Hook. Sp. Fil. ii. 280; Hk. & Baker, Syn. Fil. 172; Benth. Fl. Austral. vii. 732. P. ves- pertilionis, Labill. Nov. Holl. t. 245. P. aurita, Blume, Enum. Pl. Jav. Fil. 213; Mett. Fil. Hort. Bot. Lips. t. 14; Hook. Sp. Fil. ii. 281. P. sinuata, Wall. Cat. 84; Brack. Fil of U.S. Explor. Voy. t. 14; Hook. Sp. Fil. ii. 282. — Litobrochia aurita, Bedd. Ferns South. Ind. t. 221. Sikkim, Bhotan, and Khasia; alt. 3000-6500 feet, frequent.—Distrib. Ceylon, Malaya; wide scattered in the tropics and southern subtropics of Asia, Australia, Polynesia, America, Africa. (P. marginata, Bory, Hk. & Baker, Syn. Fil. 172, has attributed to it Syllet as a locality, on the authority of Wallich, both by Hooker and Baker. There is no North- Indian specimen ; and I believe the fern has not been collected north of Malacea and Ceylon. Wallich mixed his collections from remote localities under the same number, frequently before distribution; and he has in so many known instances thus mixed different species, that I am not at all disposed to accept the locality of Sylhet on his sole authority. He not improbably confused the species P. marginata and P. Wallichiana altogether. Wallich’s example, supposed to be from Sylhet, is perhaps that on which Agardh founded his P. revolvens (Recens. Gen. Pter. 74).] 21. CERATOPTERIS, Brongn. 1. C. THALICTROIDES, Brongn. ; Hk. & Bauer, Gen. Fil. t. 12; Hook. Sp. Fil. ii. 235 ; Wall. Cat. 83; Blume, Enum. Pl. Jav. Fil. 240; Bedd. Ferns South. Ind. t. 75; Hk. & Baker, Syn. Fil. 174; Benth. Fl. Austral. vii. 695. Parkeria pteridioides, Hook. Exot. Flora, tt. 147, 231; Hk. & Grev. Ie. Fil. t. 97; Hk. & Bauer, Gen. Fil. t. 50. Pteris succulenta, Roxb. in Cale. Journ. Nat. Hist. iv. 508. Acrostichum thalic- troides, Linn. Sp. Pl. 1527. . From the Punjab to Bhotan and Chittagong; alt. 0-3000 feet, common.—Distrib. Deccan and Ceylon; Malay Peninsula; in the tropics of the whole world. In rice-swamps, floating ; but much more commonly erect, tufted, in ditches, or even SECOND SERIES.—BOTANY, VOL. I. 9T 412 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. in dry spots during the rains. The floating and the erect forms both produce their barren and fertile fronds. In ditches the rhizome is somewhat creeping and stout. 22. Lomarta, Willd. [Beddome (Brit. Ferns, Suppl. p. 9) states that Lomaria Patersoni, Spreng., Hk. & Baker, Syn. Fil. 174, is generally distributed in India; but I never heard of its being collected in North India, and in the Kew Herbarium there is no specimen north of the Nilgherries. | Sect. Plagiogyria. Base of stipe dilated, triquetrous. Capsules with an oblique ring. 1. L. ADNATA, Blume, Enum. Pl. Jav. Fil. 205. More than half the pinnze of the barren frond sessile adnate.—Hook. Sp. Fil. iii. 19, t. 147; Hk. & Baker, Syn. Fil. 182. L. euphlebia, Hook. 2nd Cent. Ferns, t. 89, not of Kunze. Plagiogyria adnata, Bedd. Ferns Brit. Ind. t. 51. P. scandens ?, Mett. Farn. Plagiog. 9. Khasia ; alt. 4000-5000 feet, plentiful.—Distrib. Java, Japan. Pinnæ of the barren frond usually approximate, often all adnate, so that the frond is ` pinnatifid, scarcely pinnate; pinne often faleate, simply serrate, or nearly entire in their lower portion, varying from scarcely acute to caudate.—The Indian L. adnata is easily separated from L. euphlebia, but the Japan critical examples unite the two species, 80 that I cannot name them. The typical L. adnata is sent from Japan; but Maximowicz names the whole series Z. euphlebia, Mett.—As to the name L. adnata, Blume, the only example from Java so named is a portion of a fertile frond quite impossible of deter- mination. The Khasi plant may yet turn out something quite different from L. adnata. 2. L. etauca, Blume, Enum. Pl. Jav. Fil 204. Lower pinns of the barren frond white-glaucous beneath, minutely serrate in their lower half, suddenly truncate to a very short stalk.—Kunze, 2nd Suppl. to Schkuhr, t. 138; Hook. Sp. Fil. iii. 22; Hk. & Baker, Syn. Fil. 182, Plagiogyria glauca, Mett, Farn. Plagiog. 9; Bedd. Ferns Brit. Ind. t. 90. Khasia, alt. 4000-5000 feet; Moflong Wood, Hk. f. & T.; Vale of Rocks, Upper Kalapani, Sohra Reen (Surareem), O.B. Clarke.—Distrib. Java. I think only a var. of L. pycnophylla, powdered beneath. There is no difference between the two in the degree of serration and acumination of the points of the pinnt, — or in the frequency of the presence of a gland at their base. F. Henderson observé — that the lower pinnæ, not the mere auricles, are distant. Some of the Java Z. glauca | are much larger than the Khasia, s 3. L. rxcoPHYLLA, Kunze, in Bot. Zeit. 1848, 143. Lower pinns of the barren frond - green beneath, minutely serrate in their lower half, suddenly truncate to à vey — _ short stalk.—Hook. Sp. Fil. iii, 21, t. 148; Hk. Baker, Syn. Fil. 183. D. callosa : MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 473 _ Fée, Gen. Fil. 70. Plagiogyria pycnophylia, Mett. Farn. Plagiog. 8; Bedd. Ferns Brit. Ind. t. 52. A x & Trans.Linn. Soc. Sxn 2. Bor. Vor. ].Pr.50. DAVALLIA UROPHYLLA, Hook (wot Walt.) ; : Fitch imp ; Trans. Linn. Soc. SER. 2.Bor.VoL.1.PL. 51. "SÉ dal ot keh, CHEILANTHES FARINOSA, Aaud Var DALHOUSIA, (sp. Hook) Pitch imp Trans. Linn.S oc.SeR.2.Bor.Vou.1.Pu. 52. CHEILANTHES ALBO-MARGINATA, C2C~arée. = AM Cockerill del et lith. TRANS. LINN. Soc. SER. 2.Bor.VoL.lPr.53. Fitch imp. PTERIS QUADRIAURITA, Zesz. Ver KHASIANA. E SE MEC aU iln 0 p Trans. Linn. Soc. SER.2. Bor. Von Pu.54 : PTERIS GREVILLEANA, Wad oa Trans. Linn. Soc. Sen? Boa Vor! Pr.55 Fitch amp PTERIS QUADRI-AURITA, Zæz. Ver BLUMEANA, Agardh. ~ ` RN CBClarke. Trans. Linn. Soc. BER.Z.Bor.Vor I. Dr, EE H L So 5 et cue D 1 PTERIS SUBINDIVISA, £ZZZaz4e. 2. ASPLENIUM PEKINENSE, zzz. Fitch imp. XN oi ve TEP A n UN ee oh > : Y Ñ > We IE BS T NAUN oa 5 LENE Vs y SSO We CFT VINE Me V NS A LE SOON, OY TATE Die vá = GIVI) PS x UNS EEN, c n 9888 V d : N d antis f) andes Trans Low Boc SEr.2.Bor. VoL 1. P1 57 Eu SU Q Ser " xU QUSS 2 Ü e ry en PI S e cy \Y) ^ AX be A di Q yy Do de 7 "d OV Gre, S À a CBClarke DC esch WO i i FS N Vd D SËCH Lë, 5 ep NOD - a E MONA TA ES SOUL UES Ben e EC Ae 1 E \ EU GE ot SN a ëch NIMM STOUR Sé DEM, ! ec A | ANUS inn nouo AN MW att) | : D i} SPO 70) 000 X eu h Y d Nd | SC? KT SEU, Dn, NET Y AA GE sg OW EEN AAS GG, n N K UN ^ d^ ; fa". v A 20 y 3 RR NS SQ o 2 A re matt br. z e Ke D ES Q = a $ N T 3 3 SS o uU) a bd = < = E Z Ej = EN Ge < — E. Fitch imp ci M QUSS BAM 2 eto Ne 3 Se 5. » A Trans.Linn.Soc.SeR2Z Bor Vor I.Pr.58. 2 U e AN xx. 4 Y SS qw fer 2-37] 2 Ve aE EE ZU STC | MeV TUE Wis A N iu D RF nk S ASPLENIUM FILIX-F(E MINA, ZerzZ. Var PECTINATA. $^ AN à f à 4 ? zs A d | "zd SO e BY T EN Ge AE TEM GE Ai EE, e? BA ` NS VN 2 GE GE U ` 4 a : d AY A oar ki 2 Nat. size. % - KE Fitch imp. o t£» 3 3 LI H N o Ñ pa e : : » less \ $ Kei i ha | 8 d o Div Dei | 2 K 3 S ; ES g^ oO d E à M : RS z < «D e S 2272777 "e nm E^ INe N 7 | Ke s a UU E A S = N a SE Ü eg? ` Sc? St Res 00, mv d L Sz as pee” ^ D d yp’ SA) j s ei o 7795 GET, | e AA : H 7 T "a | S =) E EA E «m des Sas EK e LASPLENIUM FILIX-F(EMINA D ei e e m HE AM Cockerill Ad a kith 2. CBClarke. Traws.Linn.Soc. Szr.2. Bor. Vor, I.Pr.60. AMCockerill Lej ASPLENIUM FILIX-F(EMINA, Bern’. Var FLABELLULATA, CA are Fitch imp C.B.Clarke. Trans. Linn. Soc. Ser.2.Bor. Vor I Pr. 8l. ASPLENIUM FILIX-F @ MINA, Bernd. Var POLYSPORA, CBClarke Zernh. Var PARASNATHENSIS, C2Clarke. Fitch imp. i is AN Cockerill del. et lith. KN MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 495 Himalaya, from East Kashmir to Bhotan, alt. 5000-12,000 feet, very common. . Here are collected all the Himalayan forms that are at least 3-pinnate, except the low-level A. Filix-feemina, var. pectinatum. The form figured by Beddome is the least divided form, which, I fear, is only arbitrarily separated from the most divided form of A. macrocarpum, called A. decipiens by Mett. Wallich’s typical plant is 4-pinnate sub- 5-pinnatifid, the ultimate segments narrow, sharply toothed. Var. foliosa, Wall. Cat. 359, partly, not as to type sheet. Rhachis often very red and waved, glandular in the axis of the pinnze; ultimate segments broader than in the type; sori small. (Pl. LXII. fig. 2.) Frequent at high levels in Sikkim and Nepaul.—Oblong or triangular in outline. Var. spheropteroides. Involucre small, early disappearing ; sori both lateral and ter- minal on the veins, becoming soon globose, often appearing elevated from the frond. (Pl. LXII. fig. 1.) : Throughout the Himalaya, alt. 9000—13,000 feet, very common.—I include here a number of forms which I have distributed under the names spAeropteroides, rubricaulis, and Andersoni. A. spheropteroides type is often 6-8 feet high, triangular in outline, the lowest pinnz 2 feet, the quaternary pinne oblong, hardly lobed or pinnatifid. Sir W. J. Hooker had a specimen of this in over-ripe fruit, which he marked “ Davallia an Spheropteris sp. 2?” There are small 3-pinnate examples which, having gathered from the same tufts, I know to be varieties of this. There are other forms with the sori large, ultimately covering the whole lower surface of the frond; on one of these Col. Beddome has noted “ vix 4. fimbriatum.” Ihave another very large compound plant, finely cut, with smaller sori and more persistent involucres, which connects A. sphero- pteroides with the type. Pl. LXII. fig. 1 is taken from a very small example of the var. spheropteroides. isis | | Subgenus V. Pseud-Allantodia. Veins free. Involucres oblong or subquadrate, not curved in age, rarely placed back to back, not dehiscing from the outer edge, but breaking up from the middle of the back irregularly. (Differs from genus Allan- todia by the free veins.) | 38. A. PROCERUM, Wall. Cat. 2203. Rhizome very shortly creeping; stipe muricate, eri yellowish ; frond large, 3-4-pinnate, herbaceous, green; primary pinne 1-2 feet; ultimate pinne pinnatifid 1 or 3 to the midrib into oblong crenate, scarcely serrate lobes; veins in the lobes often forked; sori short, oblong, mostly in two rows, near the midrib of the ultimate pinne and parallel thereto; involucre delicate.— 4. um- brosum, var: procerum, Hk. & Baker, Syn. Fil. (2nd ed.). (Pl. LXIII. fig. 1.) Himalaya, from Kumaon to Bhotan, alt. 4000-8000 feet, very common. Assam, Khasia, alt. 3000-6000 feet, very common. A most abundant Sikkim and Khasia fern, remarkably constant in character. It is ho use quoting synonyms for it, as I arrange the material in the Kew Herbarium bundles differently from Baker. Col. Beddome has marked it “ Athyrium aspidioides ? ; SECOND SERIES.—BOTANY, VOL. I. Lag i A 3Y ; 496 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. but the involucre is totally different as from all forms of 4. australe, Brack. A. pro- cerum is perhaps nearest 4. multicaudatum, in which the involucres are not allantodioid, and which has a wide-creeping root; the sori are differently placed also. "A Madeira form of A. umbrosum, A. Smith, has the sori allantodioid, but the cutting of tho frond is totally unlike that of 4. procerum. ` Var. Mishmica. Ultimate segments more approximate than in the typical A. procerum ; sori small, in two rows near the midrib of the segments. Mishmee, Griffith. Khasia and Sikkim, alt. 1000 feet, C. B. Clarke.—Involucre allantodioid, not rarely diplazioid; veins in the ultimate segments often forked. 39. A. BELLUM, C. B. Clarke. Stipes tufted, from a caudex sometimes standing a foot out of the ground; stipe muricate, reddish, with long lanceolate chestnut scales at the base; frond large, sub-3-pinnate; primary pinne 1-2 feet; tertiary pinnæ (seg- ments) oblong crenate serrate, hardly pinnatifid; veins rarely forked; sori short, oblong, not very close to the midrib.— Aspidium marginatum, Wall. Cat. 391, partly. (Pl. LXIII. fig. 2.) Sikkim and Bhotan, alt. 1000-5000 feet, frequently collected and very many times seen, C. B. Clarke. Khasia, alt. 4000 feet, Cherra, Sir J. D. Hooker. Sir J. D. Hooker marked his specimen Gymnosphera gigantea; it has been removed to Asplenium by Baker without providing any name for it. The cutting and venation greatly resemble that of Alsophila glabra, and the sori are subquadrate ; but the invo- lucre bursting at the back the sori as they ripen appear in a Cyatheoid hyposorous involucre, which, however, is boat-shaped instead of hemispherical. The tertiary pinnze (though not quite free themselves) are sometimes subpinnatifid when the veins in them are subpinnate (as in similar cases in all ferns): these examples look different from the type, but I have cut them from the same caudex. | Subgenus VI. Diplazium. Veins free. Involueres linear or oblong, not or very ` slightly eurved, some placed back to back, dehiseing from their outer edge. * Fronds simple, entire or nearly so. 40. A. LANCEUM, Thunb. Fl. Japon. 333; Ic. Pl. Japon. Dec. ii. t. 8. -Rootstock wiry, creeping, with small, linear, black scales; stipes manifest, not crowded; frond narrow-lanceolate or linear entire.—Hook. Sp. Fil. iii. 235; Mett Farngatt. Aspl. 161; Benth. Fl. Hongk. 451; Hk. & Baker, Syn. Fil. 229. A. subsinuatum, Wall. Cat. 199; Hk. & Grev. Ic. Fil. t. 27. Scolopendrium dubium, Don, Prodr. Fl. Nep. 9. Diplazium lanceum, Presl; Bedd. Ferns South. Ind. t. 227, not of Boryjin Bélanger Voy. Bot. Crypt. 37, t. iv. . Assam and North-east Bengal, alt. 500-5000 feet, not very common ; NODE to Chittagong, C. B. Clarke, and to East Nepanl, Wallich; —Distrib. agin (hints, For- mosa, Japan, Frond variable in length, but not much in breadth—from 3 by Zi in. to 12 by tis in MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 497 ** Fronds 1-pinnate, not 2-pinnate except in A. japonicum, var. 41. A. BANTAMENSE, Hk. & Baker, Syn. Fil. 231. Rootstock creeping, as thick as a goose-quill; stipe with lanceolate caducous scales; pinne on each side the main rhachis 3-6, usually all free, alternate or not rarely subopposite, rhomboid rounded or acute at base, entire or crenate-serrate towards the apex.— A. alternifolium, Mett. . Fil. Hort. Lips. 75, t. 12. figs. 1, 2, and in Ann. Mus. Lugd. Bat. ii. 237; Hook. Fil. Exot. t. 17, Sp. Fil. iii. 239. 4. fraxinifolium, Wall. Cat. 194; Hook. Sp. Fil. iii. 240, 2nd Cent. Ferns, t. 19; Benth. Fl. Hongk. 454, partly. A. Donianwm, Mett. Farngatt. Aspl 177. .4. Hookerianum, Wall. Cat. 7090, not of Wall. Cat. 2682, nor of Colenso. Diplazium alternifolium and bantamense, Blume, Enum. Pl. Jay. Fil 190,191. D. fraxinifolium, Bedd. Ferns Brit. Indi t. 69; the syn. A. elegans to be excluded everywhere. Khasia; alt. 4000 feet, frequent. Cachar, alt. 500 feet, Sir J. D. dicken R. L. Keenan. Chittagong, alt. 200 feet, C. B. Clarke.—Distrib. Tinnevelly; Malay Peninsula and Islands ; South China. The Aneiteum examples have the pinnæ meine and Gerten a bud on the rhachis at the base of the penultimate pinna: this is Wallich’s 4. fraxinifolium. The pinnz are often subopposite in Indian examples; the penultimate bud I have only once found.—Diplazium falcatum, Don, Prodr. Fl. Nep. 13, from the description seems the present fern, but there is no authentic example; and 4. dantamense is not otherwise known from the Himalayas, so that it is not likely that Wallich collected it so far west as i This is, I believe, the only authority for the locality Himalayas. 42. À. Tveim, Hook. 2nd Cent. Ferns, t. 17, Sp. Fil iii. 244. Coriaceous, upper- most pinnz coalescing, or the terminal pinnz more or less pinnatifid ; lateral pinnze 5-12, on either side the main rhachis free, serrate or subentire.—A. pinnatifido- pinnatum, Hook. Sp. Fil. iii. 238; Hk. & Baker, Syn. Fil. 281. Diplazium pinna- tifido-pinnatum, Bedd. Ferns Brit. Ind. t. 244. Mishmee, Griffith; four fronds apparently collected on one occasion.—~Distrib. Hong- kong, Java. The Java frond is 18 in., with 12 free pinne on either side the main rhachis. The Hongkong fronds are 4-6 in., with 5-6 free pinnae on either side the main rhachis. Griffith’s examples are halfway between the two; and the same fern beyond doubt as mine, As to the long pinnatifid terminal pinna insisted on by Sir W. J. Hooker, two of Griffith’s fronds do not show it.—This is very near 4. sylvaticum, which has more numerous herbaceous pinns, crenate or very slightly pinnatifid into obtuse lobes, rather than serrate. Bentham’s description of 4. sylvaticum, in Fl. Hongk. 452, seems intended to cover the examples since referred to 4. Lobbianum. 43. A. SYLVATICUM, Presl, Rel. Haenk. i. 42, excl. var. B. Stipes nearly naked; frond a eg pinnz numerous, horizontal, the upper scarcely pinnatifid, the lowest pinnatifid } rarely 3 the distance to the midrib ; sori linear, many diplazioid.— 3Y 2 498 MR. €. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Mett. Farngatt. Aspl. 179, partly; Hook. Sp. Fil. iii. 248; Benth. Fl. Austral. vii. 750; Hk. & Baker, Syn. Fil. 232. A. acuminatum, Wall. Cat. 205; cf. Mett. in Ann. Mus. Lugd. Bat. ii. 238. A. elatum, Mett, Farngatt. Aspl. 180. Diplazium sylvaticum, Schk. Krypt. Gew. 80, t. 85 b; Bedd. Ferns South. Ind. t. 161, Ferns Brit. Ind. t. 243. Anisogonium sylvaticum, Hk. & Bauer, Gen. Fil. t. 56 B. Sylhet Station, alt. 100 feet, Jaintea, Jarain, alt. 4000 feet, C. B. Clarke.—Distrib. Nilgherries and Ceylon; Malay Peninsula and Islands and South China, Mauritius (Central Africa, America, Polynesia ?). My fronds are typical specimens, i. e. they resemble exactly single pinne of 4. latifo- lium, Don; they are attached to the caudex, and are very fully fruiting. Wallich’s ex- ample, marked by his hand Æ. soboliferum, is exactly the same thing, from Chappedong Hill, in Birma; but the number on the ticket (viz. 1670) is wrong, and the reference of the specimen to A. porrectum, Wall. (Hook. Sp. Fil. iii. 251), seems unusually wide of the mark. This fern appears to me only to be fronds from the young caudex (not young fronds) of A. latifolium; G. W. Cat. Ferns Ceylon, 5, appears to distinguish between such 1-pinnate fronds and 4. sylvaticum; but I do not know how he does it. As to Diplazium bulbiferum, Brack. U.S. Explor. Ferns, t. 18. fig. 1, it may he either this or A. japonicum, if these two can be kept distinct. 44. A. TOMENTOSUM, Hook. Sp. Fil. iii. 249, not of Mett. Caudex 1-2 in., erect from the ground, sending down very wiry black roots; main rhachis pubescent, not tomentose ` pinnze narrow-oblong, falcate, usually auricled at base on the upper margin, entire, or in large fronds pinnatifid halfway to the midrib; sori long, often reaching nearly to the margin —Hk. & Baker, Sp. Fil. 234. A. soboliferum, Wall. Cat. 201, type sheet. Diplazium tomentosum, Bedd. Ferns Brit. Ind. t. 195. Khasia, Griffith (one example).—Distrib. Burma, Malay Peninsula. This species here: stands, as Mr. Baker has left it, on its non-creeping rhizome. Bed- dome’s figure is excellent. On the same caudex are many short-stiped fronds, with sub- entire pinnæ, and often a few long-stiped fronds with pinne pinnatifid halfway down. The Indian type is not exactly that of Diplazium tomentosum, Blume, Enum. Pl. Jav. Fil. 192; the authentic specimen of which (does not show the rhizome, and may therefore be another species, and) has the pinne pinnatifid to the rhachis nearly, and the main rhachis very villous, subtomentose.—The fern of Griffith’s referred by Fée to his Asplenium argutans (Gen. Fil. 194, See Mém. 53) is 4. tomentosum, Hook.; but it probably has nothing to do with the Bourbon plant, which is Fée's type of 4. argutans.—The type specimens of Wallich have a rooting penultimate bud. 45. A. JAPONICUM, Thunb. Fl. Jap. 334. Rhizome creeping; main rhachis glabrous or pubescent; pinne alternate or subopposite, pinnatifid halfway or nearly the whole way to the rhachis into obtuse lobes; veins all simple, or some forked; sori linear, commencing near the midrib, falling short of the margin.—Hk. & Baker, Syn. Fil. — 294. A. Schkurii, Hook. Sp. Fil. iii. 251, not of Mett., nor of Thwaites. A.ambi- — MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 499 guum, Schk. Krypt. Gew. t. 75, not t. 75 b. A. decussatum, Wall. Cat. 2208. A. Thwaitesii, A. Braun; Mett. Farngatt. Aspl. 183; Hook. 2nd Cent. Ferns, t. 45, Sp. Fil. iii. 250; Hk. & Baker, Syn. Fil. 235. .4. tomentosum, Mett. Farngatt. Aspl. 182, not of Hook. A. lasiopteris, Hk. & Baker, Syn. Fil. 235. Diplazium Schkuhrii, Bedd. Ferns South. Ind. t. 230 (not quite Asplenium Schkuhrii of Thwaites and Baker). D. Thwaitesii, Bedd. Ferns Brit. Ind. t. 291. D lasiopteris, Kunze ; Bedd. Ferns South. Ind. t. 160. D. bulbiferum, Brack. U.S. Explor. Ferns, t. 18. fig. 2 (but the names under the two ferns figured in this plate have perhaps been transposed). North-west Himalaya; Nepaul, Wallich; Nynee Tal, Edgeworth, H. C. Levinge; Kangra, Edgeworth; Kumaon, Strachey and Winterbottom. Khasia; alt. 3000 feet, abundant, C. B. Clarke. Chittagong; alt. 200 feet, common, C. B. Clarke.— Distrib. South India and Ceylon, Burma, Malaya, China, Japan, Polynesia. The distribution and varieties of this fern are puzzling. The North-west Himalayan plant is the same as the typical South-Indian; no other North-Indian examples except these three sheets exist at Kew; but the fern is abundant in East Bengal, whence I have brought in large quantity two Khasi and two Chittagong forms. All these North- Indian forms have a distinctly creeping rhizome, and are (in my judgment) trifling varieties (except var. chattagramica). They vary much in size and hairiness: the ordinary Khasi and Chittagong plant is very glabrous; the 2nd Khasi form has the rhachis pubescent, and is altogether more hairy than 4. tomentosum, Hook. ; it has been distributed by me as A. tomentosum, Hook., erroneously. Small forms of this much resemble A. grammitoides, Hk. & Baker, Syn. Fil. (ed. 2), 491. Var. chattagramica. Frond 2-pinnate; some of the secondary pinnz subpetioled ; sori reaching to the edge, but not commencing from the midrib. (Pl. LXIV. fig. 1.) Kasalong, Chittagong Hills, €. B. Clarke.—Mr. Baker is not willing to place this under A. japonicum; but I feel sure, from its variation, that it belongs here. As to the nearly allied forms I may mention, in passing :— (Var. ?) Schkuhrii, (sp.) Thwaites; Hk. & Baker Syn. Fil. (ed. 2), 491, excl. the figure of Bedd. quoted. Large, compound ; the secondary pinnze 3 in., deeply pinnatifid. —D. decurrens, Bedd. Ferns South. Ind. t. 229. : Cachar, R. L. Keenan. | | A. polyrrhizon, Hk. & Baker, Sp. Fil. (ed. 2), 490, only differs from A. japonicum in having a non-creeping rhizome. Beddome (Ferns Brit. Ind. Suppl. p. 12) doubts if this distinction can be relied on: I share his doubts. (A. polyrrhizon does not approach 4. tomentosum, Hook.) Diplazium malaccense, Presl; Fée, Gen. Fil. 213, t. 176. The authentic specimen of this is the tip of a very large pinna of D. latifolium, Don. Asplenium Schkuhrii, Mett. Farngatt. Aspl. 182. The authentic specimen marked by Mettenius’s hand is 4. sylvaticum. ^^ Diplazium speciosum, Bedd. Ferns Brit. Ind. t. 290, is said by Beddome to be equal to * 500 -MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Aspl. acwminatum, Wall., and the figure appears made therefrom. This Aspl. acuminatum, Wall., I cannot separate from A. sylvaticum. But the authentic example of Diplazium speciosum, Blume, differs considerably from 4. acuminatum, Wallich, and appears a good species. At all events A. acuminatum, Wall., should be attached to A. sylvaticum rather than to A. mme? | 46. A. Sronrczk m, C. B. Clarke. Caudex very stout, erect, stipes below densely clothed with linear scales 1-1 in.; pinne 2-6 by 1-1 in., caudate, regularly pinnatifid nearly to the rhachis; sori ohai or linika riots not produced to the margin of the frond.— 4. marginatum, Wall. Cat. 209, type sheet partly, not of Linn. A.sorzo- gonense, Hook. Sp. Fil. iii. 252, partly; Hk. & Baker, Syn. Fil. 236, partly, not of Presl and Mett. . Diplazium sorzogonense, Bedd. Ferns Brit. Ind. t. 246. D. Sto- licske, Bedd. Ferns Brit. Ind. Suppl. 13, t. 361. Himalaya, alt. 7000—10,000 feet, from Nepaul to Bhotan, in large quantity and gre- garious around Darjeeling. My examples show a greater Biflécéhos in width of pinna than do Beddome's figures of A. sorzogonense and Stolicehe.—A. sorzogonense, Presl, is founded on Cuming’s No. 301 (see also Mett. Farngatt. Aspl. 185), a Luzon plant; it is exactly =A. parallelum, Wall. Cat. 228, from Penang and Malacea. This is a thoroughly tropical fern, the pinnæ not caudate, the segments obtuse, the sori long, reaching quite to the margin. There is an example of the true A. sorzogonense marked * Khasia, Griffith," but without any original ticket. -I suspect Griffith collected this at Malaeca.—The Javan A. sorzo- gonense, referred to the Luzon plant by Mettenius, differs again somewhat, but may belong; it certainly does not approach the Sikkim A. Stoliczhe.—A. sorzogonense B. majus, Hook. (Sp. Fil. iii. 252), Hk. & Baker (Syn. Fil. 236), differs totally.— A. polymorphum, Wall. Cat, 230,.is greatly mixed in the mounting; the Kew Herb. sheet contains a frond of 4. Stoliceke, and another not A. polymorphum. T 3 Fonds RAEREIS or more compound. 47. A. TORRENTIUM, C. B. Clarke. Catdex erect; stipes nearly glabrous; frond 6-18 in., 2-pinnate very irregularly cut ; secondary pinne 1-2 in., pinnatifid 3-3 the way to the midrib ; sori long ; involucre early disappearing. (Pl. LXIV. figs. 2, pe Sikkim, alt. 1500-5000 feet, ©. B. Clarke. | Collected on several oceasions, always on the margins of torrents, goe the caudex stands, from sand between rocks, often several inches out of the ground. The most marked feature in this fern is the very fugacious involucre; I long supposed it a Gymnogramme. It is possible that it is a form of A. latifolium, Don, growing irregu- larly in an abnormal situation. The secondary pinnæ vary greatly in size, and in the degree to which they are developed. - 48. A. SIKKIMENSE, C. B. Clarke. Primary pinne 2 feet, resembling tes ing A. = : MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 501. polypodioides, but the ultimate segments longer; rhachis of the primary pinna rough with small short scales; secondary pinne eut down nearly to the rhachis into close, oblong, regular obtuse segments, their rhachises pubescent not squamose; segments 4-2 in., crenate-serrate not pinnatifid; veins mostly forked, microscopi- cally pilose ; involucres very short, in two oblique rows quite close to the rhachis of the segment. (Pl. LXV. fig. 1.) Sikkim, Sir J. D. Hooker; alt. 500 feet near the Teesta, C. B. Clarke. This fern has been placed with Xx a ES Sc? AN Sg ZEE eR Trans. Linn. Soc. Gen? Bor. Vor.I.Pr.62. Hoot. Var FOLIOLOSA, (Sp) Wade. » BRIATUM, Zook Var SPHEROPTEROIDES. l.ASPLENIUM FIM e PUTAS wë y 3 ge, EN SEN y PA DA SS ` Ce ea pec anil Sh SSS MÀ à CUTER DONT Pare" Cet pu Fe) AMCockerill del et Hth. E D d Lem) e el c bn Ern So Me kon PILLS ay We SS, wl SAS 117 2t L^ br WIZE 2 al Wes e UP UM am. EN PE 17an m 4 LL PERSE BS ON Vë aoe ANS “ZS Ke [^v AUS Aes CAS Urs VANS Gë V E PN Ng AM (UNS Wes N $ fA GS AD (m Uh ZA A (| R N ANS. MAN JAD IN S & Š A Be TAN ü ES S IRS DS He y ls (R AS RS oy ONSEN ee e, AM Cockerill del.et hth. lASPLENIUM PROCERUM, Wed. vy N i | € S Ww) LIN MA A ER a kA: ST sg - AT (f ri AN, EE RTE, SAN ot Gen M, 4, y Se Le Lé (| (4 EFL EE j^ ee) 273 A Zëss E ST Ee | (7 d dee A AAN Sl) Si A rdi AN CAN GV £y) Zee CUN su 7 AY AN Ww ^ ` wéi ™ ` - ap, A $ X ën n d DË E EE, ie e 2 Eeer 2. A BELLUM, ZZ Clarke. Fitch imp. Trans. Linn. Soc. SerZ Bor. Vor I. Pr..64. ; l.ASPLENIUM JAPONICUM, ZZzz:J. "an CHATTAGRAMICA. (0 AMCockerill delet lith. 23ATORRENTIUM CZZ/arze. +A. SUCCULENTUM, CZ Clarke, Fitch imp. Trans. LINN. Soc. SER. Bar VoL 5 COORD Cees a WK FN ER Eh? | | AR S 2253 W t S AAAS (Zs ^ JD {ibe £ - SS Wee du SS Zo. wi Aad i HA ats 7 Mrz TC VR £ Lay} Zi 7 Ds DBOD E AP e Ze "re E AMCockerill delet lith, MISPLENIUM SIKKIMENSE, GR@¥ente 2 NEPHRODIUM GRACILESCENS, Zoa. Var DECIPIENS. Fitch imp. saga ABPIDIUM PRESCOTTIANUM, Wad. Var BAKERIANA, (sp) WS Adkinson. = Ta i e e wie Vi X i d f 2 E | MN i z ra P A. [7 i S LL a A 1. an A d if p - e" x P 3 i e i À - V : £ K f / TY 1 ARE aig x d SS fel ae SSW ^ J P . x ; e = on rs - GC i Z * ~ - s ———— +, > s T E = " ies á ^ , A ~ el H Së b E. d f S è . S * " ~ | IN à - d Dës d e Ki on atur y Lech à à K ge } Trans. Liny. Soc. Szr2. Bor. Vol Pu 66. < =< ~ Se .. Trans. Linn. Soc. Zen 2 Bag Von] Pie? I ator £s hh AW Bed OE lNEPHRODIUM GRACILESCENS, Hoos. Yer. HIRSUTIPES. | 2 is : Var DIDYMOCHL(ENOIDES. CBClarke. : Trans.Linn.Soc. Ser 2.Bor VoL I Pri 868. l.NEPHRODIUM FILIX-MAS, Zicha. Var. PANDA. Bec : » Var NORMALIS. AM Cockerill delet bth Trans. Linn. Soc. Sen? Boe Vo Prb) ge dd Deen NV v 7 2^ Zee MS AM.Cockerill del.et lith. us SE Fitch imp. E | CBClarke. pes TBAus.Liww.Soc. Sek 2.Bo T. VoL I.Pr.70. CBClarke. Trans. Linn. Soc. SER.2 BoT. VoLIPL 71. go P (0 AM Coekerill Ai lith NEPHRODIUM FILIX-MAS, ichd. Var MARGINATA, [Sp] WaZ : Inaxs.Limm.Soc.S8mn2Bor VoL IP. 72. M RHODOLEPIS, CBClarke. = 8 A S Y ; CBClarke. Trans. Linn.Soc. SER2 Bot. Vou I Pu.73. , WSAtkinson. Ka (i z SI z Z = 2 A Q O o DC A, D a, ; ST E WH Lat ech VA ee ON TST Rs dy NNUS (SND ma Ss C.B.Clarke. MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 567 Var. niphoboloides. Surface beneath densely brown tomentose, the sori buried in the tomentum. Bhotan, Griffith.— But there is a series of examples connecting this var. with the type. —As to the name of this species, I cannot find where P. venustum, Desv., was described ; it has, at all events, so completely slipped out of sight that it no way interferes with P. venustum, Wall. 67. P. LEIoRHIzON, Wall. Cat. 308, chiefly. Rhizome creeping shortly, with lax slaty- brown, ovate, not acute scales; pinnze narrow, lanceolate-linear, coriaceous ; primary parallel veins hardly more prominent than the secondary venation, with one sorus between each pair of such parallel obscure veins.—Mett. Fil. Hort. Lips. 37, t. 25. fig. 17; Farngatt. Polypod. 103; Hook. Fil. Exot. t. 25, Sp. Fil. v. 92; Hk. & Baker, Syn. Fil. 369. P. lucidum, Roxb. in Cale. Journ. Nat. Hist. iv. 486. Pleopeltis leiorhizon, Bedd. Ferns South. Ind. t. 174. North India, alt. 04000 feet, very common in the lower hills, extending west to Nepaul.—Distrib. South India. 33. NOTHOLENA, R. Br. 1. N. Maranra, R. Br. Prodr. Fl. Nov. Holl. 145. Nothochlena Marante, Hook. Sp. Fil. v. 120; Bedd. Ferns Brit. Ind. t. 1; Hk. & Baker, Syn. Fil. v. 120. 4crosti- chum Marante, Linn. Sp. Pl. 1527; Schk. Krypt. Gew. t. 4; Sibth. Fl. Gree. .. 1.964. Gymnogramme Morante, Mett: ; Milde, Fil. Europ. 21. Alpine Himalaya, from Kashmir to Kumaon; rare. Sikkim, alt. 9000-15,000 feet, Lachen Valley, Sir J. D. Hooker.—Distrib. South Europe and the Mediterranean Region from Maearonesia to the Caucasus and Abyssinia. The scales on -the back of the fronds may be called “ pseudo-wool"' (Añvoc); I do not see how the fern can be said to have a **pseudo-indumentum ;" it has a real indu- mentum. Rob. Brown called it Notholena ; and I am convinced that he meant Notholena, and that Desveaux blundered. 34. GYMNOGRAMME, Desy. Series I. Veins free. | Subgenus I. Leptogramme. Sori oblong, not forked. 1. G. Torra, Schlecht. Adumbr. 15, t. 6. ` Rhizome shortly creeping ; stipe and frond on both surfaces pilose; frond l-pinnate; pinnz pinnatifid about halfway to the midrib ; veins in the lobes simple.—Blume, Fl. Jav. Fil. 90, t. 388; Hook. Sp. Fil. v. 138; Hk. & Baker, Syn. Fil. 376. G. aspidioides, Blume, Enum. Pl. Jav. Fil. 112. G. Loveii, Mk. & Grev. Ie. Fil. t. 89. Polypodium Tottum, Willd. Sp. Pl. v. 201. Grammitis Totta, Presl; Hk. & Bauer, Gen. Fil. t. 728; Bedd. Ferns South. Ind, t. 49. Phegopteris Totta, Mett. Farngatt. Pheg. & Asp. 19; Milde, Fil. Europ. 101. North-west Himalaya, Kashmir to Kumaon, alt. 6000 feet, rare. Khasia; alt. 3000- SECOND SERIES.—BOTANY, VOL. I. _ | 4H 568 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 5000 feet, common.—Distrib. South India, Ceylon, Java, China, Japan; Africa (almost the whole) with its islands. . G. AvRITA, Hook. Sp. Fil. v. 141, 2nd Cent. Ferns, tt. 74, 89. Rhizome extensively creeping; base of the stipe curved, with deflexed ovate-acute scales; frond 1-pin- nate; pinnæ pinnatifid nearly to the midrib; veins in the lobes many of them ` forked.—Hk. & Baker, Syn. Fil. 377. Grammitis aurita, Bedd. Ferns Brit. Ind. 51. .152. Sikkim and Bhotan, alt. 3000-6000 feet; common. Assam and Khasia, alt. 2000- 5000 feet, very common. Beddome (Ferns Brit. Ind. Suppl. p. 24) doubts if this is more than a Ss of Poly- podium distans: the rhizome is very different, and is constantly accompanied by a differ- ence in the sori. The fern is glabrous beneath, or the rhachises glandular-puberulous or subpubescent. Var. Levingii. Frond weak; pinnz not auriculate; lobes broadly oblong (not ge pilose on both surfaces; sori marginal. Kashmir, Jhelum, and Chittapani Valleys, alt. 4000-7000 feet; .H. C. Levinge. —This has the texture and hairiness of G. Totta, but the rhizome and venation of G. aurita ; while the cutting is deeper than that of C. Totta, less auriculate than that of G. aurita. 3. G. OPACA, Spreng. Syst. Veg. iv. 89. Stipes tufted, with ovate-acute scales near the foot; frond 2-pinnate, texture thin, succulent, drying very black, margin crenate, scarcely serrate; lowest pinnz little shorter than those above.—Bedd. Ferns Brit. - Ind. t. 238; Hk. & Baker, Syn. Fil. 378. G. obtusata, Blume, Enum, Pl. Jav. Fil. 119, Fl. Jav. Fil 97, t. 43; Hook. Sp. Fil. v. 143. Phegopteris opaca, Mett. Farngatt, Pheg. & Asp. 15. Hemionitis opaca, Don, Prodr. Fl. Nep. 13. From Nepaulto Bhotan, alt. 4000-7000 feet; frequent. Khasia, alt. 3000-5000 feet ; : common.—Distrib. Java. wi G. arborescens, De Vriese, reduced here by Sir W. J. Hooker, does not perhaps belong: ` the examples are firm stout green, with an acutely toothed margin. Subgenus II. Syngramme. Sori elongate, often forked or confluent. | AG vestra, Hook. Sp. Fil. v. 143, Ic. Pl. t. 115. Frond linear, l-pinnate; pinn® ovate, stalked, entire, densely rufous-villous beneath.—Hk. & Baker, Syn. Fil. 379. : Grammitis ? vestita, Wall. Cat. 19. Syngramma vestita, Bedd. Ferns Brit. Ind. : t. 154. From Chumba to Nepaul ; alt. 6000-9000 feet, common. Sipe Griffith. sch ; South and North China, E 5. G. ANDERSONL, Bedd. Ferns Brit. Ind. t. 190. Frond l-pinnate ; pinnæ . " : We MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 569 ‘lobed, densely clothed beneath with soft golden hair—Hk. & Baker, Syn. Fil. 380. . Kumaon, 7. Anderson. Nothing is known about this at Kew but the "MR and description of Beddome. 6. G. FRAXINEA, Bedd. Ferns Brit. Ind. Suppl. p. 24. Frond large, glabrous, or very .. nearly so, 1-2-pinnate ; ultimate pinne large, stalked, linear-lanceolate-acuminate, . entire or minutely serrulate.— G. javanica, Blume, Enum. Pl. Jav. Fil. 112, Fl. Jav. Fil. 95, t. 41; Hook. Sp. Fil.v. 145 ; Bedd. Ferns Brit. Ind. t. 57, Ferns South. Ind. t. 232; Hk. & Baker, Syn. Fil. 382. ` OG. serrulata, Blume, Enum. Pl. Jav. Fil. 113, Fl. Jav, Fil. 96, t. 42. — Diplasium fraxineum and falcatum, Don, Prodr. Fl. Nep. 12, 18. Grammitis procera, Wall. Cat. 3. G.caudata, Wall. Cat. A OG. affinis, Wall. Cat. 11. p Himalaya, from Chumba to Bhotan ; alt. 1000-8000. feet, very common. Khasia, alt: ` 1000-5000 feet; common.—Distrib. Ceylon, Malaya, Japan, Polynesia, Tropical Africa. Var. pilosa, (sp.) Brack. U. S. Explor. Ferns, 22, t. 4. Pubescent beneath. . Sikkim, alt. 6000 feet, C. B. Clarke.—Distrib. Sandwich Isles. 7. G. MICROPHYLLA, Hook. Sp. Fil v. 129, 2nd Cent. Ferns, t. 16. Fronds tufted, 2—4 in. high, delicately green, 3-4-pinnatifid ; ultimate segments 3—$ in., obovate- oblong.—Hk. & Baker, Syn. Fil. 383. Grammitis microphylla, Bedd. Ferns Brit. Ind. t. 148. . Khasia; Surareen, alt. 5000 feet, Griffith. Sikkim; Tonglo, alt. 7000—10,000 feet, Si; J. D. Hooker, H. C. Levinge, Col. Oakes ; Sinchul, alt. 8000 feet, Gamble; Dikeeling, alt. 7500 feet, C. B. Clarke. Series II. Veins inarching. | Subgenus III. Steguogramme. Stipes continuous with the rhizome. Frond 1-pinnate. 8. G. AsPIDIOIDES, Hk. & Bauer, Gen. Fil. t. 120 B. Rhachises beneath pilose; pinne pinnatifid 3-1 the way to the midrib, 2 pairs of veinlets uniting; sori curved.— Hook. Sp. Fil. v. 150, 2nd Cent. Ferns, t. 50; Hk. & Baker, Syn. Fil. 378. G. steg- nogramme, Blume, Fl. Jav. Fil. 98, t. 44. Stegnogramme aspidioides, Blume, Enum. Pl. Jav, Fil 173; Presl, Tent. Pter. t. 9. fig. 5; Bedd. Ferns Brit. Ind. t. 149. Ca hegopteris stegnogramme, Mett. Farngatt. Pheg. & Asp. 26. Khaia; alt. 4000-6000 feet, Griffith, Hook. f. $ T. Thoms. — Distrib. Ceylon, Java. The Jus examples have the hairs denser, needle-like, the pinne less lobed, with 3-6 pairs of veinlets uniting, the sori straight, hence much shorter than in the Khasi plant, Wherein the veinlets curve much upwards to the sinus. It is this last character that _ Makes the Khasi possibly a distinct species from the Javan; but much more material > required from both localities to found a species on such a character.—De Vriese and P. No. 612, referred here by Sir W. J. Hooker, has the frond EE below into auricles, and is T auriculatum, Wall. (vel prox.). 402 570 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Subgenus IV. Selliguiea. Stipes articulated on the rhizome. Fronds undivided or scarcely 1-pinnate. 9. G. LANCEOLATA, Hook. Sp. Fil. v. 156. Rhizome creeping, not very stout; frond 1-$ in. wide, simple, linear, entire, coriaceous, all the veins obscure; sori often nearly parallel with the main rhachis.—Hk. & Baker, Syn. Fil. 387. Grammitis lanceolata, Swartz, Syn. Fil. 22, 212, t. 1. fig. 4; Blume, Enum. Pl. Jav. Fil. 117; Hk. & Grev. Ic. Fil. t. 48. Antrophyum lanceolatum, Blume, Fl. Jav. Fil. 84, i. 36. Loxogramme lanceolata, Presl; Hk. & Bauer, Gen. Fil. t. 73 B; Bead. Ferns South. Ind. t. 51. Polypodium Loxogramme, Mett, Farngatt. Polypod. 112, t. 3. fig. 25. Selliguiea lanceolata, Carr. in Fl. Viti. 371. Khasia, alt. 4000-5000 feet, frequent. —Distrib. South India, Ceylon, Malaya, China, Japan, Africa with its eastern islands. 10. G. rNvonvTA, Hook. Sp. Fil. v. 155. Rhizome stout, shortly creeping; frond 3-23 in. wide, simple, linear, entire, coriaceous, all the veins obscure; sori usually oblique to the main rhachis.—Hk. & Baker, Syn. Fil. 387. Grammitis involuta, Don, Prodr. Fl. Nep. 14; Hk. & Grev. Ic. Fil. t. 53; Blume, Enum. Pl. Jav. Fil. 87. G. cuspidata, Zenk. Pl. Ind. i. £. 2. OG scolopendrina, Bory in Voy. Coq. t. 30. fig. 1. OG flavescens, Wall. Cat. 6. G. acuminata, Wall. Cat. Y. G. macrophylla, Wall. Cat. 10. Antrophyum involutum, Blume, Fl. Jav. Fil. 87. A. coriaceum, Blume, Fl. Jav. Fil. t. 37. fig. 1. Loxogramme involuta, Presl; Bedd. Ferns South. Ind. t. 50. Polypodium involutum, Mett. Fil. Hort. Lips. 97, t. 25. figs. 26, 27. P. Blumei, Mett. Farngatt. Polypod. 118, t. 3. fig. 27. Selliguiea Wallichana, Hook. Ic. Pl. t. 204. Himalaya, alt. 2000-7000 feet; from Gurwhal to Bhotan, very common. Khasia, alt. 1000-5000 feet, very common.—Distrib. South India, Ceylon, Malaya, Polynesia. I do not know how this fern differs from G. lanceolata, except by its larger size; the ` scales on the rhizome, the venation, and the texture are (so far as I can make out) exactly the same. 11. G. Haminronrana, Hook. Sp. Fil. v. 160. Rhizome creeping; fronds linear, simple, | with conspicuous venation, the fertile smaller on much longer stipes.—Hk. & Baker, E Syn. Fil. 389. Grammitis Hamiltoniana, Wall. Cat. 9. Ceterach pedunculata, ` Hk. & Grev. Ic. Fil t. 5. Selliguiea Hamiltoni, Presl, Tent. Pterid. 216, t. WC fig. 16; Bedd. Ferns Brit. Ind. t. 239. Polypodium pedunculatum, Mett. in Herb. North and East Bengal, alt. 0-4000 feet ; from Nepaul to Mishmee and Chittagong, common; extending also some way into the plain, as at Chattuck, C. B. Olarke— Distrib. Yunan. : 12. G. znurPriCA, Hk. & Baker, Syn. Fil. 389. Rhizome creeping; fronds deeply pin: . natifid (sometimes 1-pinnate) or deeply 3-5-lobed ; main veins of the lobes distinct- — 4G. decurrens, Hook. Sp. Fil. v. 161; Benth. Fl. Hongk. 457. Poigpodiem 6 eli s r6 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 571 cum, Thunb. Fl. Jap. 335; Mig. in Ann. Mus. Lugd. Bat. ii. 225. P. pothifolium, Mett. Farngatt. Polypod. 108, t. 3. figs. 22-24. , Hemionitis pothifolia, Don, Prodr. Fl. Nep. 18. Grammitis decurrens, Wall. Cat. 5. G. Finlaysoniana, Wall. Cat. 776. Selliguiea decurrens, Presl; Bedd. Ferns Brit. Ind. t. 150. From Nepaul to Bhotan and Khasia, alt. 2000-5000 feet, very common.-—Distrib. Burma, China, Philippines, Japan. . Very variable in development ; large specimens are 1-pinnate, with the pinne distant, 113 by ŝ in.; other specimens are subpalmately divided into 3-5 short lobes ; and I have an example (in full fruit) quite simple. I believe it is all one.—The Hongkong example is dimorphie; the fertile frond reduced, with narrow pinne. 35. BRAINEA, Hook. I. B. insignis, Hook. Kew Gard. Misc. ix. 354, Sp. Fil. v. 162; Benth. Fl. Hongk. 460; Bedd. Ferns Brit. Ind. t. 189; Hk. & Baker, Syn. Fil. 390; Kurz, For. Fl. Brit. Burma, ii. 574. Bowringia insignis, Hook. Kew Gard. Misc. v. 287, t. 2, Fil. Exot. t. 38. - Khasia, alt. 3000-4000 feet ; Pomrang, Hook. f. $ T. Thoms; Jainka, Col. Godwin- Austen.—Distrib. Hongkong. 36. Meniscium, Schreb. l. M. TRIPAYLLUM, Swartz, Syn. Fil. 19, 206. Rhizome creeping, with chestnut lanceo- late-linear scales at the extremities; frond with 1-9 pinne; pinne narrowed at the base, villose beneath; secondary venation not prominent.— Wall. Cat. 61; Hk. & Grev. Ic. Fil. t. 120; Kunze, Farnkr. Schk. Suppl. t. 52 ; Bedd. Ferns South. Ind. t. 56; Hk. & Baker, Syn. Fil. 391. M. erosum, Wall. Cat. 62. M. Cumingii, Fée, . Gen. Fil. 222. JM. Parishii, Bedd. Ferns Brit. Ind. t. 184. a - Eastern Bengal, alt. 0-500 feet, from Mishmee and Cachar to Chittagong, where it is plentiful. Sikkim Terai; Dulkajhar, Gamble.—Distrib. Ceylon, Burma, China, Philip- pines, Malacca. . M. simplex, Hook., has the fronds less divided, more cordate at the base, the secondary venation raised beneath, prominent, but seems very near.—Jf. Parishit, Bedd., is referred in Hk. & Baker, Syn. Fil. 392, to M. cuspidatum, Blume; but it is exactly our common form of M. triphyllum in the Chittagong Hills. An example collected in Mishmee by Griffith has 13 pinne in all, and is marked M. villosum, J. Smith in Herb. 2. M. CUSPIDATUM, Blume, Enum. Pl. Jav. Fil. 114, Fl Jay. Fil. 102, t. 45. Frond l-pinnate ; pinns 3-6 in., pubescent or nearly glabrous except the mg e narrowly oblong, acuminate caudate, drying blackish, margin often crenulate.— Bedd. Ferns Brit. Ind. t. 309; Hk. & Baker, Syn. Fil. 392. Jaintea; Jowye, alt. 4000 feet, C. B. Clarke. Sylhet, alt. 250 feet, C. B. Clarke— Distrib. Java, Philippines. tae - l ipe no difficulty in distinguishing this from P. urophyllum, Wall. (as seen under * 572 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA.. that species above); but it is very difficult to separate from M. triphyllum; indeed Mr. Baker has sorted our M. triphyllum, var. Parishii, under M. cuspidatum, Blume. Var. longifrons, (sp. JWall. Cat.60. Pinnæ 12-15 in., glabrous, entire, with 15-20 pairs of uniting veinlets, suddenly narrowed into a subulate apex 1-2 in. long. Himalaya and Khasia, alt. 500-4000 feet, very common.—This is probably a good species, but I can fix on no good diagnostic character; none of the Javan material approaches it, nor does it run into P. wrophyllum, with which it is often confounded. It is marked M. reticulatum in the Kew bundles. 3. M. DELTIGERUM, Wall. Cat. 59. Fronds 1-pinnate, glabrous; fertile frond longer than the barren, the pinnse little narrower than those of the barren, crenated; sori ultimately confluent, forming deltoid masses between the very distinct main veins beneath.—Acrostichum virens, var., Hk. & Baker, Syn. Fil. 420. A. costatum, var., Hook. Sp. Fil. v. 262; Bedd. Ferns Brit. Ind. t. 114. Himalaya, alt. 500-2000 feet, from Nepaul to Bhotan, common. Chardooar and Roopraee Chang Hills, in Assam, Simons. Chittagong Hills, C. B. Clarke. If the position of the sori and the nature of the venation be adjudged matters of no weight in ferns, this species may be got into the same genus with Aerostichum virens; it will then be very difficult to unite it specifically, as there are no intermediate forms, and the fertile pinnz are only slightly dimorphous. Col. Beddome's figure unfortunately does not show the main veins beneath the fertile pinna, which are very strong and a diagnostic mark. 37. ANTROPHYUM, Kaulf. 1. A.comrAcEUM, Wall. Cat. 43. Stipe 0 or short; frond linear lanceolate acuminate, 3-12 by 4-1 in., plicate, owing to the sori being much sunk in the frond, their lines distinctly raised on the upper surface.—Hook. Sp. Fil. v. 169; Hk. & Baker, Syn. Fil. 398. A. plicatum, Fée, Ze Mém. Foug. 44, t. 5. fig. 1. Hemionitis coriacea, Don, Prodr. Fl. Nep. 18. : Nepaul, Wallich. Sikkim; alt. 2000-5000 feet, C. B. Clarke. Mishmee, Griffith. Khasia, Griffith, Simons, alt. 250 feet at Shaila, C. B. Clarke.—Distrib. Mergui. There are both trigonous and ovoid spore-cases in this species; perhaps in all. Beddome doubts (with reason), Ferns Brit. Ind. Suppl. p . 25, whether the species is mg tinet from A. reticulatum. 2. A. PLANTAGINEUM, Kaulf.; Bory in Voy. de la Coq. Bot. Crypt. t. 28. fig. 1. Stipes — many of them long ; frond oblong, acute, not plicate ; sori immersed.—Fée, 3" Mém. — Foug. 45; Carr. in Fl. Viti. 371. 4. reticulatum, Bedd. Ferns South. Ind. t. 52. : Sikkim; Tehonpong, alt. 4000 feet, Dikeeling, alt. 5000 feet, C. B. Geng? Assam, 7 Simons. Khasia, Griffith.—Distrib. Ceylon, Malaya, Polynesia. | Tr Tem This differs from A. reticulatum by the long stipes, from A. dalifolium by eng deeg .MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 573 not obovate nor lobed at the summit, and by the sori more sunk in the substance of the frond. | 3, A. RETICULATUM, Kaulf.; Fée, 9" Mém, Foug. 44. Stipe short or 0; frond linear or linear-lanceolate, 38-10 by i-1i in., not plicate; sori immersed.— Wall. Cat. 40; Hook. Sp. Fil. v. 169; Bedd. Ferns South. Ind. t. 231; Hk. & Baker, Syn. Fil. 393; Benth. Fl, Austral. vii. 777, in part. .4. falcatum, Blume, Fl. Jav. Fil. 76, t. 32. Hemionitis reticulata, Forst. Prodr. 79. = Sikkim, Dr. Jerdon.—Distrib. Ceylon, Malacca, Polynesia, Queensland. There is only one North-Indian specimen now left in the Kew bundle, and that I suspect is our common A. coriaceum, dried under much pressure by Dr, Jerdon, so as to show the plication less than usual. Wallich’s A. reticulatum is very large, 3-4 in. broad.— Fée says his 4. reticulatum has a long stipe, and quotes as a picture Blume's Fl. Jav, Fil. t. 32, which shows the fronds attenuate to the rhizome. Var. parvula, Fronds 1-4 by 4-4 in., sides very parallel; sori appearing as though forming parallel lines the whole length of the frond, but really reticulating. ~ Sikkim, W. S. Atkinson; Yoksun, alt. 4500 feet, C. B. Clarke.—l know no reason why this should not really be the same as 4. parvulum, Blume, Fl. Jav. Fil. 78, t. 34. fig. 3. That figure represents the fronds as being rather larger, with less parallel sides, the sori in less continuous lines. 4. A. LATIFOLIUM, Blume, Fl. Jav. Fil. 75. Stipe long; frond large, broadly obovate, often acuminate or acutely lobed at the apex ; sori superficial, ultimately elevated beneath.—Fée, 3™ Mém. Foug. 48; Hook. Sp. Fil. v. 172; Bedd. Ferns Brit. Ind. t. 176; Hk. & Baker, Syn, Fil 394. A. Boryanum, Blume, Fl. Jav. Fil. 75, t. 31, not of Hk. & Grey. Hemionitis Boryana, Blume, Enum. Pl. Jav. Fil. 111, syn, excl. Sikkim and Bhotan, alt. 2000-6000 feet, frequent. Assam and Khasia, alt. 1000- 4000 feet, frequent.—Distrib. Java. My large Sikkim examples are 15 by A8 in., the anterior margin acutely lobed, and are easily called A. Jatifolivm. But my smallest Sikkim examples are 32 by 4 in.; and, except that they are obovate, often lobed or notched at the vertex, I do not rom how they are to be separated from .4. plantagineum: I see no tangible difference in the elevation of the sori beneath. 38. VITTARIA, Smith. LV. ELONGATA, Swartz, Syn. Fil. 109, 302. Rhizome somewhat long; scales many, with long, very black hair-like points; fronds 4-26 by i-i = midrib cma beneath, even in the upper half of the frond; outer lip of the involucre not tin- . guishable from the margin of the frond; sori ultimately elevated.— Wall. Cat, 144; — Fée, 3™ Mém. Foug. 22, t. 3. fig. 5; Bedd. Ferns South. Ind. t. 21; Hk. & Baker, ‘Syn. Fil. 395; Luerssen, Fil. Graeff. 90, cum syn. V. rigida, Kaulf.; Wall. Cat. 574 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 143 ; Hook. Sp. Fil. v. 184; Carr. in Fl. Viti. 372. V. plantaginea, Hk. & Grev. Ic. Fil. t. 187. V. zosterefolia, Bory ; Fée, 3"* Mém. Foug. t. 2. fig. 2; Hook, Sp. Fil. v. 183. Pteris angustifolia and graminifolia, Roxb. in Cale. Journ. Nat. Hist. iv. 502, 508, t. 33. . Plain of North India, abundant; from the sea-face of the Soonderbun to the inner Himalayan valleys, but not often found above 4000 feet alt.—Distrib. South India, Ceylon, Burma, Malaya, Queensland, Polynesia, Mauritius, Tropical Africa. A very uniform fern; the large examples reach 24 in. often, and are V. elongata of Wallich, Pteris graminifolia, Roxb. ; the shorter fronds of 6-10 in. are V. rigida, Wall, Pteris angustifolia, Roxb.—Without examination of the involucre this species (as to the Indian material) is easily sorted from the next by the rhizome and scales. 2, V. FLEXUOSA, Fée, 3" Mém. Foug. 16. Rhizome very shortly creeping; stipes tufted; scales acuminate, with slaty-brown or fuscous caudate points; fronds 4-18 by 4-4 in., midrib distinct beneath even in the upper half of the frond; outer lip of the involucre easily distinguishable from the margin of the frond ; sori sunk in a groove, ultimately more or less exsert.—Hook. Sp. Fil. v. 178; Luerssen, Fil. Graeff. 84, 92. H lineata, Roxb. in Cale. Journ. Nat. Hist. 509, t. 33; Hook. Sp. Fil. v. 180, as to the Indian material; Hk. & Baker, Syn. Fil. 396, as to the Indian mate- rial Teniopsis lineata, Bedd. Ferns South. Ind. t. 54. Himalaya, alt. 2000-12,000 feet; from Gurwhal to Bhotan, very common. Khasia, alt. 1000-6000 feet ; very common.—Distrib. South India, Ceylon ?. V. lineata, Swartz, with which this is united by Mr. Baker, has the stipes less tufted, the soral groove more marginal (as in the next species). V. fleruosa is very doubtfully distinct ; but I have retained the name because all our North-Indian V. lineata belongs to this one form. The geographical distribution is of course as doubtful as the synonymy. Miquel's V. japoniea seems the Himalayan plant. 3, V. SIKKIMENSIS, Kuhn in Linnea, xxxvi. 66, as to the Sikkim plant only. Rhizome very shortly sai cain with slaty-rufous hair-pointed scales; stipes densely tufted ; frond 3-4 by 4; in., subobtuse, midrib beneath obscure or slightly depressed; soral lines large, marginal, the outer lip undistinguishable from the edge of the frond; sori, when young, deeply sunk in a groove; sporangiasters few, intestiniform, clavate; sporothecze ovoid.— V. minor, Fée, var. minima; Bedd. Ferns Brit. Ind. t. r (Pl. LXXXIV. fig. 3.) Sikkim, alt. 2000-6000 feet ; common. Khasia, alt. 2500 feet, Mowlong, C. B. Clarke. — Kuhn put together this Sikkim tender grass-green species and the Moulmein and ` Malay plant Vittaria minor, Fée (Tenitis pusilla, Mett. in Ann. Mus. Lugd. Bat. w ^ 172), which has a coriaceous broader frond, the midrib distinctly elevated beneath, | E stout wiry roots, the lines of sori four times as large, &c. This V. minor may be, aS — Mr. Baker notes, V. falcata, var.—Kuhn and Beddome had, however, only some 4 scraps of the Sikkim plant to work with. This Sikkim species seems to me closely e allied to the American V. lineata; except that it is small, I see no parnou E resem- MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 575 blance between it and V. minor, Fée. Fully developed Sikkim plants are much longer, but narrower than the form figured by Beddome. 39. Tanitis, Swartz. 1, T. BLECHNOIDES, Swartz, Syn. Fil. 24, 220. Fertile frond 1-pinnate; pinns linear- lanceolate or linear entire; sori in a continuous line about midway between the midrib and the margin.—Wall. Cat. 141; Blume, Fl. Jav. Fil. t. 28. fig. 2 & t. 29; Fée, 3"° Mém. Foug. 26; Hook. Sp. Fil. v. 187 ; Bedd. Ferns Brit. Ind. t. 54; Hk. & Baker, Syn. Fil. 397. T. pteroides, Schk. Krypt. Gew. t. 6b. T. interrupta, Wall. Cat. 142; Hk. & Grev. Ic. Fil. t. 63. Sylhet; Wallich.—Distrib. Malay Peninsula and Islands, Ceylon. The form T. interrupta, Wall, is hardly worthy the rank of a variety ; it only differs from the type in having the lines of fruit more or less broken irregularly.—I do not doubt the locality of this plant, as Wallich has noted that his single specimen was for- warded him from Sylhet by his collector Da Silva, who collected there. 40. DRYMOGLOSSUM, Presl. l. D. carnosum, Hook. Sp. Fil. v. 189. Rhizome wide-creeping, bearing scattered lanceolate-linear spreading blackish scales; soral lines when young not marginal, but in age often covering nearly the whole frond.—Hk. & Bauer, Gen. Fil. t. 784 ; Fée, 3™ Mém. Foug. 29; Bedd. Ferns Brit. Ind, t. 55; Hk. & Baker, Syn. Fil. 397. Notholena carnosa, Wall. Cat. 138. Nepaul, Sikkim, Bhotan, Khasia, alt. 2000-5000 feet, frequent. em Fertile fronds sometimes linear, sometimeselliptic. The species is easily distinguished from the next by the scales of the rhizome: Hooker both figured and described the scales of D. carnosum from a rhizome of D. piloselloides, and Beddome's figure of the scales is rather suggestive of a compromise than characteristic. D. subcordatum, Fée (not an Indian form), has the rhizome densely clothed with spreading linear scales, and may be à variety of D. carnosum. 2. D. PILOSELLOIDES, Pres], Tent. Pterid. 227, t. 10. figs. 5, 6. Rhizome e . €elothed completely with adpressed, peltate, diamond-shaped, fulvous-red 18. » decurrens (not from North India). ` 19. » alcicorne (not from North India). 20. ae . fuciforme (not from North India). 596 Ec MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 21. Aerostichum punctulatum (not from North India). . 22. ag contaminans = A. crispatulum, Wall., var. 23. E: triquetrum = Lomaria euphlebia, Kunze. 24. de crispatulum, Wall. Good. ' 25. m flagelliferum, Wall. Good. 26. va costatum, Wall. Good. 27. is rigidum (not from North India). 28. » viviparum = A. appendiculatum, Willd. 29. ps Hamiltonianum = A. appendiculatum, Willd. 30. » setosum = A. appendiculatum, Willd. 3l. y aureum, Linn. Good. 32. Lomaria serpens = Acrostichum axillare, Cav. . CS attenuata (not from North India). ER. y secunda (not from North India). 98. y limonifolia (not from North India). a. . re scandens = Acrostichum scandens. Mir er spondizfolia (not from North India). 38 5 aurea = Onychium auratum, Kaulf. 39 caruifolia = Onychium auratum, Kaulf. 40. "ST og reticulatum, Kaulf. Good? 4l. » Boryanum (not from North India). 42. $ pumilum (not from North India). 43 5 coriaceum, Wall. Good. 44. Hemionitis cordifolia = H. arifolia, Bedd. 45. Psilotum flaccidum (not from North India). 46. » triquetrum = Ps, nudum, Griseb. 47. Ophioglossum cordifolium — O. reticulatum, Linn. 48. Botrychium lanuginosum, Wall., = B. virginianum, Swartz, var. 49. » subcarnosum, Wall., = B. daucifolium, Hook. 50. Osmunda speciosa = O. regalis, Linn. » Leschenaultii = O. regalis, Linn. 52. » monticola = O. Claytoniana, Linn. 53. Anemia flexuosa (not from North India). 54. Helminthostachys dulcis = H. zeylanica, Hk. & Bauer. - s 55. Pleopeltis incana (not from North India). 96. Blechnum glandulosum (not from North India). 57. y orientale, Linn. Good. 58. Woodwardia radicans, Swartz. Good. . Meniscium deltigerum, Wall. Good. $5 longifrons — M. cuspidatum, Blume, var. 5 triphyllum, Swartz. Good. » erosum — M. triphyllum, gegen 5 salicifolium (not from North India). - Dicksonia arborescens (not from North India). » appendiculata, Wall. Good. - Cheilanthes mysurensis, Wall. (not from North India). Good. j rupestris, Wall., = Ch. tenuifolia, Swartz. is micrantha, Tall, = Ch. tenuifolia, Swartz. _ » lucida, Wall., = Onychium j tages Kunze. » pallens (not from North India). die . dealbata — Ek mes Rm. x MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 597 Cheilanthes tomentosa — Ch. rufa, Don. (This sheet is without number, but named by 25 Wallich's hand.) contigua — Onychium multisectum, F. Henderson. 73. Adiantum capillus-veneris, Linn. Good. 3) 25 25 33 25 25 33 3) A3 soboliferum (not from North India). Good? vestitum = A. caudatum, Linn. flagelliferum = A. caudatum, Linn. lunulatum, Burm. Good. amcenum = A. flabellulatum, Linn. scabrum (not from North India). reniforme (not from North India), venustum, Don. Good. rhizophorum = A. caudatum, Linn. var. 83. Ceratopteris thalictroides, Brongn. Good. 84. Pteris sinuata = Pt. incisa, Thunb. 33 22 22 25 22 25 23 22 33 2 23 3) EE) 25 25 22 3) 2 2 2 25 A3 23 2 2 25 » 23 hastata (not from North India). varians = Cheilanthes varians, Hook. mysurensis (not from North India). Pellæa geranisfolia. ludens, Wall. Good. nitidula = Pellea nitidula, Hk. & Baker. cæspitosa = Cheilanthes varians, Hook. digitata = P. pellucida, Presl, var. polita (not from North India). Lindsaya. angustata = Lindsaya ensifolia, Swartz, scabripes = Pt. pellucida, Presl, var. læta = Pt. cretica, Linn. nervosa = Pt. pellucida, Presi, var. semipinnata, Linn. Good. lanuginosa = Pt. aquilina, Linn. densa = Pt. aquilina, Linn. firma = Pt. aquilina, Linn. terminalis = Pt. excelsa, Gaud. semihastata = Pt. aquilina, Linn. lorigera = Pt. aquilina, Linn. subquinata = Pt. quadriaurita, Retz. var. linearis (not from North India). nemoralis = Pt. biaurita, Linn. (as to the type sheet). aspericaulis = Pt. quadriaurita, Retz. longipinnula, Wall. Good. umbrosa = Pt. Wallichiana, Ag. subpedata (not from North India). longifolia, Linn. Good. amplectens = Pt. longifolia, Linn. recurvata = Pt. aquilina, Linn. 114. Lycopodium subulifolium = L. setaceum, Ham. pulcherrimum = L. setaceum, Ham. var, Hookeri = L. squarrosum, Forst. urostachyum (not from North India). serratum, Thunb. Good. | 25 A3 2: 2 Se"? verticillatum = D. squarrosum, Forst, dis concinnum = Selaginella, sp. 998 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 122. Lycopodium Willdenovii — Selaginella, sp. 133. 134. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. - 171. 2 25 22 25 25 25 33 A3 25 25 2 35 3? 25 35 pubescens — Selaginella, sp. tetragonostachyum = Selaginella, sp. fulcratum = Selaginella, sp. semicordatum = Selaginella, sp. argenteum = Selaginella, sp. elegans = Selaginella. aloifolium = L. Hamiltonii, Spreng. (N.B. Blank sheet in type set.) - cernuum, Linn. . Good. divaricatum = L. clavatum, Linn. Heyneanum. Blank sheet. Phlegmaria, Linn. Good. obtusifolium = L. Hamiltonii, Spreng. gnidioides. Blank sheet. subdiaphanum = Selaginella, sp. caulescens = Selaginella, sp. Notholzna carnosa = Drymoglossum carnosum, Hook. A3 A3 piloselloides = Drymoglossum piloselloides, Pres/. (In the additional sheets there is also mixed Niphobolus adnascens and Drymoglossum Beddomei.) undulata = Acrostichum virens, Wall. var. Tænitis blechnoides, Swartz. . Good. interrupta = T. blechnoides, Swartz. Vittaria rigida, Wall., = V. elongata, Swartz. elongata, Swartz. Good. Lindsaya lucida — L. cultrata, Swartz. 33 22 A3 » 25 25 25 25 3) 2 dentata (not from North India). polymorpha — L. flabellulata, Dryand. pallens = L. cultrata, Swartz. . recurvata (not from North India). serpens (not from North India). attenuata — L. cultrata, Swartz. lobata (not from North India). decomposita (not from North India). lanuginosa = Nephrolepis volubilis, J. Smith. Gléüchéhis Hermanni = G. linearis, C. B. Clarke. A3 A8 attenuata (not from North India). gigantea = G. glauca, Hook. (The 2nd sheet is Polypodium erubescens.) Trichomanes setigerum = T. javanicum, Blume. A3 LÉI 22 3 3 A3 A3 alchemillezefolium (not from North India). undulatum (not from North India). rigidum = T. javanicum, Blume. pyramidalis (not from North India). villosula (not found among Wallich’s ferns: noted to be Salvinia natans). longisetum (not from North India). umbrosum = T. radicans, Swartz. anceps — T. radicans, Swartz, var.' EN iban Boryanum (not from North India). A3 22 A8 E Telfairianum (not from North India). crispatum — H. javanicum, Spreng. exsertum, Wall. Good. densum (as to half type sheet) = H. exsertum, Wall; ` | as to the other half = Trichomanes tipmotatun, Poir- ; 203. 204. 205. 206. 207. 208. 209. 210. 211. 212. 213. 214. 215. 216. 217. 218. 219. 220. MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 2 35 3) 33 35 A3 » 3) 3) A3 3) 22 32 32 33 » A3 35 . Hymenophyllum badium — H. polyanthos, Swartz. E serpens = H. javanicum, Spreng. . Lygodium microphyllum, R. Br. Good, except the first specimen of the type sheet, which is L. flexuosum, Swartz. longifolium = L. flexuosum, Swartz. dichotomum = L. circinatum, Swartz, polystachyum (not from North India), . Cyathea spinulosa, Wall. Good. Brunonis (not from North India). venulosa (not from North India), excelsa (not from North India). robusta (not from North India). . Spheropteris barbata = Peranema cyatheoides, Don. . Matonia pectinata (not from North India). . Lomaria Boryana (not from North India). . Marattia fraxinea (not from North India). . Angiopteris crassipes = A. evecta, Hoffm. . Asplenium reticulatum = Allantodia javanica, Bedd. planicaule = A. laciniatum, Don, var. decurrens = A. unilaterale, Lamk. Finlaysonianum = A. faleatum, Lamk. var. urophyllum (not from North India). Trichomanes, Linn. Good. fraxinifolium = A. bantamense, Baker. ovatum (not from North India). Penangianum (not from North India). radiatum = Actinopteris dichotoma, Bedd. Nidus, Linn. Good. subsinuatum = A. lanceum, Thunb. ensiforme, Wall. Good. soboliferum = A. tomentosum, Hook. proliferum = A. esculentum, Pres/ (in the 2nd sheet A. latifolium, Don, is mixed). diversifolium = A. latifolium, Don. porrectum (not from North India). acuminatum = A. sylvaticum, Presl. tenuifrons = A. Clarkei, Bedd. (but A. nigripes, Mett., is mixed on the type sheet). multijugum = À. normale, Don. multicaule = A. normale, Don. letum = A. unilaterale, Lamk. lobulosum = A. longifolium, Don. cristatum = A. unilaterale, Lamk. hirsutum (not from North India). . mysurense (not from North India). pulchellum = A. nitidum, Swartz. bullatum = A. bulbiferum, Forst. concinnum = A, tenuifolium, Don. cespitosum = A laciniatum, Don. heterocarpum, Wall. Good. ` . flagelliferum = A. longissimum, Blume, i brevisorum, Wall. Good. .: S.—BOTANY, VOL. I. (All the sheets.) AM 599 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 221. Asplenium alternans, Wall. Good. 222. » auritum (not from North India). 323. " oxyphyllum, Wail. (not from North India). 224, 5 porrectum (lege polyodon) (not from North India). 225. o falcatum, Lamk. . Good. 226. $ pellucidum (not from North India). 227. » nitens (not from. North India). 228. » parallelum = A. sorzogonense, Presi. 229. p caudatum, Wall. Good. . ; 230. j polymorphum = A. latifolium, Don (in the additional sheets Gymnogramme opaca is mixed). 331. a pectinatum = A. Filix-feemina, Bernh. (in the additional sheets A. nigripes is mixed). 382. a nitidum (not from North India). 233. si Ruta? — A. varians, Hk. & Grev. 334. » depauperatum = A. laciniatum, Don. 235. » Prescottianum (not from North India). 236. 5 proliferum (not from North India). 337. » spectabile = A.multicaudatum, Wall. (Second sheet is Nephrodium Boryanum, Baker, var.) ij 238. Cænopteris furcata (not from North India). 239. 5 vivipara (not from North India). 240. Davallia lonchitidea — D. platyphylla, Don. 241. o lobulosa (not from North India). 242. » angustata (not from North India). 243. — ,, flagellifera (not from North India). 244. » villosa = D. marginalis, Hk. & Baker. 245. » tenuifolia = D. chinensis, Swartz. 246. ,, ornata (not from North India). 247. » parvula (not from North India). 248. » achillezfolia (not from North India). : $ 249, » Serreformis (not from North India). 250. .,, pedata, Smith. Good. 251. » parallela (not from North India). 252. ` ,, coniifolia (not from North India). 253. — ,, elegans (not from North India). 954. Equisetum scirpioides, Herb. Ham., = E. elongatum, Willd. 255. ap membranulosa, Wall. Good. : 256. p immersa, Wail. Good. 257. js rhomboidea — D. polypodioides, Don, var. 258. » bullata, Wall. Good. 259. » cherophylla = D. pulchra, Don. ` 260. a stipellata = D. nodosa, Hook. 261. » pyramidata = D. flaecida, R. Br. var. T. 262. ^ ,, puberula = D. flaccida, R. Br. E: 263. » pilosula = D. flaccida, R. Br. var. SSC 264. E virens (not from North India). E 265. Polypodium costatum — P. stigmosum, Swartz. (Second sheet is mainly P. flocculosum, Don.) — 266. » porosum — P. fissum, Baker. 267. » ` pertusum = P. adnascens, Swartz. c dm. is adnascens, Swartz. Good. 969. =, mysurense = P. fissum, Baker, MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 270, Polypodium vittarioides — P. adnascens, Swartz. 971. j loriforme = P. lineare, Thunb. 272. » spherocephalum = P. angustatum, Swartz, 278. - polycephalum — P. punctatum, Swartz. 274. - longifrons — P. normale, Don. 275. » sesquipedale = P. lineare, Thunb. 276. » ovatum, Wall. Good. 277. » Mauritianum (not from North India). 278. d» furfuraceum (not from North India). 279. 5 gladiatum (not from North India). 280. ï zosteræforme (not from North India), 281. » glabrum — P. punctatum, Swartz. 282. » grandifolium = P. membranaceum, Don. 283, D hymenodes = Acrostichum axillare, Cav. 284. » hemionitideum, Wall. Good. 285. an eontiguum = P. lineare, Thunb. 286. » Horsfieldii (not from North India). Good. 287. ls Wallichii, R. Br. Good, 288. 5 coronans, Wall. Good. 601 289. » alternifolium = P, nigrescens, Blume. Var. polyphylla = ep, nae, Blume. 290. Se amoenum, Wall. Good. 291, ep quercifolium, Linn. Good, (Third sheet has P. Linnzi mixed. 292. » curvinervium (not from North India). 293. » propinquum, Wall. Good, (Third sheet is P. ebenipes, Hook. ; fourth sheet is P. hastatum, Thunb.) 294. » oxylobum = P. hastatum, Thunb, var. 295. » dilatatum, Wall. Good. 296. » verrucosum (not from North India), 297. $ glaucistipes (not from North India). 298. » caudigerum = Nephrodium cucullatum, Baker (at least as to type sheet). 299. » urophyllum, Wall. Good. 300. ^» lineatum, Colebr. Good. 301. » secundum — Nephrodium unitum, R. Br. 302. » scabridum — Nephrodium aridum, Baker. 303. » : leiorrhizon, Wall. Good. (Third sheet = P. juglandifolium, Don.) 304, 5 Lindleyanum (not from North India). 305. 5 venustum, Wail. Good. 9E S i capitellatum — P. juglandifolium, Don. 307. » meriifolium (not from North India). 308. J argutum, Wall. Good. 309. $ elongatum = Nephrodium cuspidatum, Baker. 310. » lachnopus, Wall. Good, i. e. as to the main part of the type sheet : the other sheets are largely P. microrrhizoma, C. B. Clarke. 311. » subpectinatum, Wall. (not from North India). 312. » proliferum, Rozb. Good. 313. e serra (not from North India). 314. » auriculatum, Wall. Good. ` 815. » tridactylon, Wall., = B Seen, Blume. lon Wall., = P. distans, e MR Wall., = Nephrodium parasiticum, C. B. Clarke. 318, » — Wall, (not from. North India). Alsophila, sp. 4x2 602 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 319. Polypodium comosum (not from North India). Alsophila, sp. 320. 321. 322. 323. 324. 325. 326. 327. A8 3 23 25 35 35 35 33 3 35 35 35 3» A3 35 3 3 33 LÉI 33 33 33 25 23 35 23 35 25 -marginatum = A, auriculatum, Swartz, var. . contaminans = Alsophila glauca, J. Smith. giganteum = Alsophila glabra, Hook. marginale. Type sheet contains Nephrodium pulvinuliferum, Baker, with a small piece of P. punctatum, Swartz: 2nd sheet is Nephrodium scabrosum, Baker, but is ticketed as from Nepaul. Leschenaultianum (not from North India), Alsophila. oxyphyllum = Asplenium oxyphyllum, Hook. confluens (not from North India). coniifolium = P. subdigitatum, Blume. ornatum, Wall. Good. adnatum = P. distans, Don. alternans (not from North India). Alsophila. erubescens, Wall. Good. phymatodes, Linn. (not from North India). Good. molliusculum = Nephrodium parasiticum, C. B. Clarke. brunneum = P. distans, Don, var. dentigerum = Asplenium Filix-feemina, Bernh. var. tenericaule = Nephrodium tenericaule, Hook. umbrosum (not from North India). Alsophila. ‘ Audios pilosulum — Nephrodium crenatum, C. B. Clarke. puberum = Nephrodium falcilobum, Hook. var. fimbriatum = Asplenium fimbriatum, Hook. var. patentissimum = Nephrodium Filix-Mas, Richd. var. coniifolium = A. aristatum, Swartz. eriocarpum = Nephrodium crenatum, C. B. Clarke. subdiaphanum = Nephrodium crenatum, C. B. Clarke. Brunonianum = Nephrodium Brunonianum, Hook. apiciflorum = Nephrodium apiciflorum, Hook. multidentatum = Davallia multidentata, Baker. glanduliferum = Nephrodium prolixum, Baker. ~ multijagum = Nephrodium multijugum, C. B. Clarke, not of Baker. appendiculatum = Nephrodium canum, Baker. (The other sheets have some Polypodium erubescens, Wall., intermixed.) i solutum = Nephrodium parasiticum, C. B. Clarke. ciliatum = Nephrodium ciliatum, C. B. Clarke. venulosum = Nephrodium aridum, Baker. multilineatum = Nephrodium truncatum, Presl. canescens = Nephrodium parasiticum, C. B. Clarke. prionophyllum = Nephrodium truncatum, Presi. squarrosum = Asplenium macrocarpum, Blume. crinitum (not from North India). , unitum = Nephrodium cucullatum, Baker. foliolosum = Davallia nodosa, Hook. ocellatum = A. auriculatum, Swartz, var. fuscipes, Wall., = Nephrodium sagenioides, Baker (type sheet), but nearly all the rest is Nephioainit membranifolium, Presi. riparium (not from North India). . | Prescottianum, Wall. Good. rhomboideum = A. amabile, Blume. ` sublanosum = Nephrolepis exaltata, Schott. MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 603 367. Aspidium cespitosum = A. auriculatum, Swartz, var. 368. » pungens — A. ilicifolium, Don. 369. á rufo-barbatum = A. aculeatum, Swartz, var. 370. » affine = A. aculeatum, Swartz, var. (In the 3rd sheet Nephrodium Filix-Mas, var., is mixed.) 371. » setosum = A. aculeatum, Swariz. 372. » spectabile — Nephrodium spectabile, C. B. Clarke. 373. » Wallichianum = Oleandra Wallichii, Hook. 374. js Singaporianum (not from North India). 975. y articulatum (not from North India). 376. » caryotideum = A. falcatum, Swartz. 377. » coadunatum — Nephrodium cicutarium, Baker. (In the later sheets Nephrodium membranifolium, Presl, is mixed.) [377. » multicaudatum = Nephrodium multicaudatum, C. B. Clarke. | 378. » alatum = Nephrodium vastum, Baker. 379. 5 variolosum — Nephrodium variolosum, Baker. 380. » atratum — Nephrodium hirtipes, Hook. var. 381. » caducum, Hall, Good. 382. e polymorphum = Nephrodium polymorphum, Baker. 383. js rostratum = Nephrodium polymorphum, Baker. 384. 5 aduncum. Blank sheet. 385. J Telfairianum (not from North India). 386. » terminans. Blank sheet. 387. » canum — Nephrodium ciliatum, C. B. Clarke. 388. » semi-bipinnatum. Blank sheet. 389. i eburneum = Nephrodium canum, Baker (as to type specimen ; the other spe- cimen on the first sheet is Asplenium oxyphyllum, Hook. ; the second sheet is Asplenium macrocarpum, Blume). 390. T densum = Nephrodium sparsum, Don, var. - 391. » marginatum, Wall., = Asplenium bellum, C. B. Clarke (as to type sheet). 392. j nitidulum = Nephrodium sparsum, Don, var. 393. ;, ^ divisum = Nephrodium Boryanum, Baker. 394. 55 tectum (not: from North India). 395. Arthrobotrys macrocarpa — Nephrodium cochleatum, Don. 396. Cryptogramme Brunoniana = C. crispa, R. Br. 397. Equisetum debile, Roxb. Good. 398. 5 scoparium — E. diffusum, Don. 775. Spheropteris Hookeriana = Diacalpe aspidioides, Blume. 776. Grammitis Finlaysoniana =Gymnogramme elliptica, Baker. 1031. Aspidium exaltatum = Nephrolepis exaltata, Schott. e Var. = Nephrolepis volubilis, J. Smith. 1032. i Tavoyanum — Nephrolepis cordifolia, Hk. & Baker. * 1033. Acrostichum virens, Wall., replaced by Equisetum panonieum = E. diffusum, Don. 1034. Arthrobotrysavana = Nephrodium cochleatum, Don. 1035. Asplenium Tavoyanum = A. falcatum, Lamk. 1036. is Grevilleanum (not from North India). . 1037. Equisetum pallens — Equisetum debile, Rozb. 2162. Acrostichum Finlaysonianum, Wall. Blank sheet. 2163. e Wightianum = A. appendiculatum, Willd. 2164. ý vestitum (not from North India). 2165. » fivulare. Blank sheet. 2166. H contractum. Blank sheet. 604 2167, 2168. 2169. 2170, 2171. 2172. 2173. 2174. 2175. 2176. 2177. 2178. 2179. 2180. 2181. 2182. 2183. 2184. 2185. 2186. 2187. 2188. 2189. 2190. 2191. 2192. 2193. 2194. 2195. 2196. 2197. 2198. 2199. 2200. 2201. 2202. 2203. 2204. 2205. 2206. 2207. 2208. MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. ` Acrostichum stelligerum. Blank sheet. S terminans = A. virens, Wail, Lomaria Wightiana, Wall. Blank sheet. Hemionitis hastata = H. arifolia, Bedd. Ophioglossum reticulatum, Herb. Ham. Not found. Blechnum Finlaysonianum, Wall, Blank sheet. Dicksonia scabra, Wall, Good. j moluecana,.Rozb. Blank sheet. Cheilanthes rigidula, Wall., = Ch. farinosa, Kaulf. Adiantum hirsutum, Willd. Blank sheet. » flabellulatum — A. hispidulum, Swartz. Pteris Wightiana — Pt. aquilina, Linn. » arguta, Ham. Blank sheet. » geminata = Pt. biaurita, Linn. var. » ensiformis, Ham. Blank sheet. » alternifolia (not from North India). Lycopodium rotundifolium. Blank sheet. » Wightianum (not from North India). P. levigatum. Blank sheet. » complanatum, Selaginella, sp. 5 ornithopodioides. Selaginella, sp. J bryopteris. Selaginella, sp. circinale. Selaginella, sp. Lonchitia hirsuta (not from N orth India). Vittaria divergens, Roxb. Blank sheet. Lindsaya attenuata, Wall., in Herb. Finlayson, therefore not from North India. (N.B. Not the same as L. attenuata, Wall. Cat. 151). » pteroides, Wall., = L. ensifolia, Swartz, » longipinna, Wall., = L. ensifolia, Swartz. » interrupta, Roxb., sub Vittaria. Blank sheet. » parasitica, Roch, sub Vittaria. Blank sheet. » Finlaysoniana, Wall. No locality (not from North India). Hymenophyllum leve, Ham. (not from North India). Trichomanes campanulatum, Rozó. No locality, Lygodium pubescens, Wall., = L, ae, Ze, 3 japonicum, Swartz. Good. » Finlaysonianum = L. japonicum, Swartz. Asplenium procerum, Wall. Good. » divaricatum, Wall. Blank sheet. 3 foliolosum, Wi all. Blank sheet, » furcatum (not from North India). » parvulum, Wail. Blank sheet. 2 decussatum, Wall., ses A. 3 japonicum, Thunb. e a » marginatum = A, Stoliezke, C. B. Clarke, the other part of the type sheet ` being A. porrectum, Wail, » contaminans, Wail. Blank sheet. » pulchrum (not from North India). ge puberulum = A. esculentum, Presl. » trapeziforme, Rozó, Blank sheet. 5 cultratum, Roxb, Blank sheet. 5 Wigktandm (not from North India). » rie en (not from North India). MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 605 2217. Asplenium obliquum, Swartz, Blank sheet. 2218. Davallia Roxburghii, Wall., = D. polypodioides, Don, var. 2219. 2 moluccana, Roxb. Blank sheet. ZA 5 caudata (not from North India). 2221. Polypodium Finlaysonianum, Wall. Blank sheet. 2222. 5 Wightianum, Wall., = P. lineare, Thunb. 2223. 5 adherens, Wall. Blank sheet. 2224. » inoides, Ham. Blank sheet. 2225. j scabrum, Roxb. Blank sheet. 2226. - arboreum, Roxb. Blank sheet. 2297. 5 subarboreum, Ham. Blank sheet. 3228. js venosum, Ham. Blank sheet. 2229. » semi-bipinnatum, Wall. (not from North India). 2230. "7 aycefolium (?), Wall. (not from North India). 3231. sinuosum, Wall. (not from North India). 9232. ptor Hamiltonianum, Wall. Blank sheet. 232988. 5» auriculatum = Nephrolepis cordifolia, Hk. & Baker (chiefly). 2234, » flagelliferum, Roxb. Blank sheet. 2235. e Finlaysonianum (not from North India). 2236. e trifoliatum (not from North India). 2297. » coriaceum (not from North India). 2238. » sophoroides (Roxé.). Blank sheet. 2239. » parasiticum, Herb. Madr. Blank sheet. 2240. 5 hirsutulum, Ham. Blank sheet. 3241. splendens, Ham. (not from North India). 2680. Pieris Grevilleana, Wall. Good. 2681. ,, multidentata = Pt. ensiformis, Burm. 2682. Asplenium Hookerianum = A. Finlaysonianum, Hook. (not of Wall. Cat. 191). 2683. Davallia urophylla = D. marginalis, Baker, var. 2684. » Hookeriana, Wall. Good. 2685. Acrostichum ludens = A. appendiculatum, Wi Ud. 4727. Polypodium acutissimum = P. lineare, Thunb. 5169. » Grevilleanum = P. lineare, Thunb. 7073. Alsophila Brunoniana, Wall. The blank sheet has been filled up by pieces of fronds “taken from the specimen on the staircase,” supposed (as I think erro- neously) to have been Wallich’s plant. 7074. b glaucescens — A. glauca, J. Smith. 7075. » Grevilleana = Davallia flaccida, R. Br. var. 7076. Cyathea bipinnata, Roxb. No locality (not from North India). 7077. Polypodium Russelianum = Nephrodium tenericaule, Hook. 7078. » pubigerum = Nephrodium Leuzeanum, Hook. 7079. ge polyodon = Aspidium caducum, Wail. 7080. A oppositum = Nephrodium sparsum, Don. 7081. » -sophoroides (not from North India). 7082. Aspidium iticum = Nephrodium parasiticum, C. B. Clarke. 7083. r amgang = Nephrodium falcilobum, Hook, (Another Mauritius fern is, however, mixed on the type sheet.) 7084. Hymenophyllum tenue, Ham. (not from North India). 7085. Pteris ensiformis, Ham., = Pt. ensiformis, Burm. 7086. ,, scabra (not from North India). 606 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA, 7087. Lycopodium pulvinatum. Selaginella, sp. 7088. p verticillatum = L. squarrosum, Forst. 7089. » pallidum. Selaginella, sp. 7090. Asplenium Hookerianum, Wall., = A. bantamense, Blume. 7091. m pellucidum (not from North India). DISTRIBUTION OF THE GENERA OF THE FERNS OF NORTH INDIA. dl 3 j : | s 4| dT 41" i ail A Fagl | 3 : E Hee] [d GESEIS SIESPIEFIBEPT LS D Ze SE, b sg Lä. al Sa o 3 modu 32/28/8388] = | ee EREECHEN: eGkEISEIpéëIé als és SEIREIES|Ssl a MA Gleichnis r.cs.. 2} l| 2| 2; 0| O0 |Nephrodium Orsi i oi ead i 0 0 1 Ord Pleocnemia ...... 91 01.8] 2T11 3 Homes 12.5.2. 9} 0} 21 21.0 1-1 Bagenis i29 Guni WA SI TUM] i Ap Tee 6| 0| 6| 5| 0| 3] Nephrolepis ........ 4| 1] 1] 440] U Deane ous 3i 0: 3i 1i OI EI Obis ..——— 3] 1| 9| 815919 DEN eee AS: 1| 0| 1| 1| 0| O | Polypodium WEE essel S] 3] 2| 01 OT*8 Phegopteris ...... 1$] 7]1.9| 061091 2 Peranema ...... subtle] 1i- 1| 01l.04-Goniopris ^ al 8| 2] Sidi t eweg VM 41.24 41.21.-01.2 Dictyopteris ...... 1| 01 0l E St Hymenophyllum `... 1 6} 3| 6| 5| 0| 2 Eupolypodium ....| 3|] 1| 2| 2] 01 2 Trichomanes ........ 7} Tt. 410] 0|-9 Goniophlebium ....| 7| 4| 6| 3] 0| 6 Diva ooa 18| 7|15|15| 2| 3]| Niphobolus........ ict | 7| T1! Cystopteris ........ $i d] Shat Or 01. DS I eui 11.01.04 t40] 9 AT ui ion 41 01 9| ft Ul Oy Dye Ee dE 9] 91921 aa ARR o xou ore! 6}. 5| 2] 9 Phymatodes ...... 23 | 6|17]|16]| 2| 8 Cheilanthes ........ 9. 0i 3] 5| 3: 2 Pleopeltis ........ £t 11.4] 81501. 1 éi ka 21 1} 2| 2| 1 jb: Od Notholens EE Li Eied4 01:501 9 Cryptogramme ...... 1| 1| 1| 0| 0|. 0] Gymnogramme ......|12| 6| 8|10| 0| 3 PME, Qus al e] I| 0 0] 21 Bamna x I Ur) | II 13 bas DU pe ue ue 17| 7|12|15| 6| 4| Meniscium .......... 1] 14 3| 9T 01 3 Ceratopteris ........ bI Ebgil| |. 0| Antfophyuh. utes 4|.0[-4] SHOV Lone. 2.5 ey n *1,90121 41 01 0] Vitteria om... ees Si 1| 9| 9| 312 Bléchtum Ee SI Ur at 21.0. +) ee |... pete 01 0| 91 11 9 7 Woodwardia ...... e| 1| 1| 1| 1| 0| Off Drymoglosum ...... $1 ol 11 3} 113 Asplenium Hemionitis.......... S| 04 0| Pa 9 TE 1| 0| 1| 1| 0| pl Acrostichum , ei 15} 1]-91184159]| * Euasplenium .. .... 24|14| 12 |31 | 2] 2 | Osmmda .........- 21.91.14 $4.90] 9 DE cr ll OL 1| f| 061] Gy PM, S ES ti Orel 11917 Athyrium ........ 12|] elit FS Poo P9 P Euedus (NOR 4| 2|. 3|:4|3| 0 Pseud-Allantodia ..| 2| 1| 2| 2| 0| 2| Angiopteris ........ 1| 0j 1|.23170| 91. Diplazium ........ 131.8] ak 8 01. 41 been e. 1l] 0] 0| elen oe Anisogonium ...... 2| 0| 1| 0| 1] O | Ophioglossum........ s|sgt 9| 112 e Demidieteum 2| 1| 1| 1| 0} 0} Helminthostachys....| 1| 0| 0| 1| 1 ol. Allantodia .......... 1| 0| 1| 1| o oy Botrychtam £s $i 9] 3|. en ý Actiniopteris ........ PPLE Ol 040 dt Od Pilotin scx os ovitele ds 1:01.09] es ol Aspuliüm oi ceci os 12| 9|11| 6| 1| 6| Lycopodium ........| 11| 3110] 9|-2 H Nephrodium Equisetum .......... ET 4-9] 317 Totas... 30 | 13/2% | 16| 2111 as LT Eunephrodium ISL St bad (84 4 Totál | send. ue 379 |149 |269 |258 | 52 KS. MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 607 EXPLANATION OF THE PLATES. PLATE X LIX. Fig. 1. Cyathea spinulosa, Wall., drawn from Wallich’s own example n. 178 of his Catalogue.—4A. Barren secondary pinna, nat. size. B. Fertile secondary pinna, nat. size. i Fig. 2. Hymenophyllum denticulatum, Swartz, drawn from a Khasi example.—B. Two entire fronds on one rhizome, nat. size, seen from above. A. One pinna, magn. 5 diam., seen from beneath. Fig. 3. Hymenophyllum Levingii, C. B. Clarke, drawn from a Yoksun example.—A. One frond, nat. size. B. Barren lobe of pinna, magn. 20 diam., seen from beneath. C. Fertile lobe of pinna, magn. 20 diam., seen from beneath. Fig. 4. Davallia pulchra, Don, var. pseudocystopteris, (sp. Kunze, from a North-west Himalayan example; extremity of an ultimate pinna, magn. 8 diam., showing the very thin indusia of large oblong cells. PLATE L. Davallia urophylla, Hook. (not Wall.), from Griffith’s Bhotan example; one lower pinna, nat. size. PLATE LI. Cheilanthes farinosa, Kaulf., var. Dalhousie, (sp.) Hook., from North-west Himalayan examples. A. an entire frond from a small specimen, and B, the lowest pinna of a frond from a larger ` specimen, both nat. size. C. Scale from the stipe, magn. 8 diam. Prate LII. Cheilanthes albo-marginata, C. B. Clarke, from a Kashmir example.—A. A whole plant, nat. size. B. Scale from the stipe, magn. 8 diam. Prare LIII. Pteris quadriaurita, Retz., var. Khasiana, from a Walong example.—A. The lowest pinna of a frond, nat. size. (The entire lanceolate-linear irregular reduced secondary pinna at the base of the pinna in this example has been faithfully added by the artist; it was accidentally present. here, but in none of the other examples.) B. A basal secondary pinna, nat. size. Prare LIV. Pteris Grevilleana, Wall., from Keenan's Cachar example; one barren and one fertile frond on the same- rhizome, nat. size. Prats LV. Pteris quadriaurita, Retz., var. Blumeana, (sp.) Agardh, from a Chittagong example ; one basal (bipartite), pinna, nat. size. | Puate LVI. : Fig. 1. Pteris subindivisa, C. B. Clarke, from a Teesta (Sikkim) example; a rhizome with one frond and part of another, nat size. (The basal pinnz are sometimes much shorter than in this fully- developed specimen, or obsolete.) : : : Fig. 2. Asplenium pekinense, Hance, from Levinge's Kashmir example; rhizome with one frond, nat. _ SECOND SERIES.— BOTANY, VOL. I. AN - 608 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. Prats LVII. Asplenium Atkinsoni, C. B. Clarke, var. Andersoni, from a Tonglo (Sikkim) example; one basal pinna of a frond, nat. size. Prag LVIII. Asplenium Filiz-femina, Bernh., var. pectinata, (sp. Wall.), from a Sikkim example; the middle of a frond (5 pairs of pinnz), nat. size. On the side is sketched (on a reduced scale), in outline, the frond from which the larger figure of the Plate was drawn. ; t Prag LIX. Fig. 1. Asplenium Filiz-femina, Bernh., var. attenuata, C. B. Clarke, from a Kashmir example; one frond, nat. size. Fig. 2. Asplenium Filiz-femina, Bernh., var. refusa, (sp.) Decne., subvar. rubricaulis, Edgw., from a North-west Himalayan example; the greater part of one frond, nat. size. Prats LX. Asplenium Filiz-femina, Bernh., var. flabellulata, C. B. Clarke, from a Sikkim example; the lower half of a frond (5 pairs of pinnze), nat. size. Puate LXI. Fig. 1. Asplenium Filiz-femina, Bernh., var. polyspora, C. B. Clarke, from a Chumba example; the greater part of a frond, nat. size. Fig. 2. Asplenium Filia-feemina, Bernh., var. Parasnathensis, C. B. Clarke, from a Parasnath example; a frond, nat. size. Prag LXII. ( Fig. 1. Asplenium fimbriatum, Hook., var. spheropteroides, C. B. Chas from a Sikkim example; the greater part of a frond, nat size. Fig. 2. Asplenium fimbriatum, Hook., var. foliolosa, (sp. Wall.), from a Sikkim example; nearly half a frond, nat. size. Prate LXIII. Fig. 1. Asplenium procerum, Wall., from a Sikkim example; basal part of one pinna, nat. size. Fig. 2. Asplenium bellum, C. B. Clarke, from a Sikkim example ; basal part of one pinna, nat. size. PraATE LXIV. ME te Fig. 1. Asplenium Japonicum, Thunb., var. chattagramica, C. B. Clarke, from a Chittagong example; ` basal pinna of the frond, nat. size. oe Fig. 2. Asplenium torrentium, C. B. Clarke, from a Sikkim large example; basal pinna of the frond, e nat. size. a Fig. 3. Asplenium torrentium, C. B. Clarke, from a Sikkim small example; basal pinna « the frond, nat. size. Fig. 4. Asplenium succulentum, C. B. Clarke, from e chung, basal portion of $ | Hager the whole of the lower pence seer nat. size. : MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 609 Prate LXV. Fig. 1. Asplenium sikkimense, C. B. Clarke, from Hooker’s Sikkim example; basal portion of the lowest pinna of the frond, nat. size. Fig. 2. Nephrodium gracilescens, Hook., var. decipiens, C. B. Clarke, from a Sikkim example; a frond, nat. size—A. One segment of a pinna, magn. 3 diam., showing the divided venules. Prate LXVI. Aspidium Prescottianum, Wall, var. Bakeriana, (sp. W. S. Atkinson), from a Sikkim example.— b. A portion of the centre of a frond, nat. size. a. The whole of the same frond, } nat. size, in outline, to show the proportion between length and breadth of the frond, and the non- attenuation of the base; the cutting of the pinn: not shown. Prate LXVII. Fig. 1. Nephrodium gracilescens, Hook., var. hirsutipes, from a Khasi example.—a. Base of a frond, nat. size. ò. Two segments of a pinna, magn. 4 diam., showing the hairy involucres. Fig. 2. Nephrodium gracilescens, Hook., var. didymochlænoides.—c. Base of a frond, nat, size. d. Two segments of a pinna, magn. 3 diam., showing the elliptic involucres. Dag LXVIII. Fig. 1. Nephrodium Filix-mas, Richd., var. panda, C. B. Clarke, from a Chumba example; basal por- tion of a frond, nat. size. Fig. 2. Nephrođium Filix-mas, Richd., var. normalis, C. B. Clarke, — a Khasi example ; a frond, nat. size. Dag LXIX. Fig. 1. Nephrodium Filix-mas, Richd., var. Khasiana, C. B. Clarke, from a Khasi example ; lower portion of a frond, nat. size. Fig. 2. Nephrodium Filix-mas, Richd., var. Schimperiana, C. B. Clarke, from a Sikkim example ; lower portion of a frond, nat. size. Nephrodium Filiz-mas, Richd., var. fibrillosa, C. B. Clarke, from the North-west Himalaya; a frond, nat. size. Prare LXXI. Nephrodium Filiz-mas, Richd., var. marginata, sp. Wall, from a Sikkim example; basal portion of the lowest pinna of a frond, nat. size. Nephrodium rhodolepis, C. B. Clarke, from a Sikkim example ; basal portion of the lowest pinna of a frond, nat. size. Prate LXXIII. Nephrodium i ingens, W. S. Atkinson, from a Sikkim example; fragment from the base of a frond, nat. size. The lowest pinna (A) is fertile; the next (B), and all the superior pinne, barren. 4N2 610 MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. PrATE LXXIV. Fig. 1. Nephrodium glandulosum, Hook., from Blume’s Java example of Aspidium glandulosum, Blume; basal portion of a frond, nat. size. Fig. 2. Nephrodium glandulosum, Hook., var. lete-strigosa, C. B. Clarke, from a Chittagong example; basal portion of a frond, nat. size. Prate LXXV. Nephrodium membranifolium, Presl.—A. Var. typica, from Chittagong; a fruiting (small) frond, nat. size. B,C. Var. dimorpha, from Sylhet; b, basal pinna of a barren frond, nat. size; c, basal portion of a fertile frond, nat. size. Dapp LX XVI. Nephrodium Wightii, C. B. Clarke, from Wight’s Deccan examples. The barren and fertile fronds depicted (nat. size) have been pasted down on one sheet, and I do not doubt were collected by Wight as of one fern. PuatTe LX XVII. Nephrodium mullicaudatum, C. B. Clarke, from a Jaintea Terai example; stipe together with the lowest secondary pinna of the lowest primary pinna of the frond, nat. size. Prag LXXVIII. Nephrolepis volubilis, J. Smith, from Hooker’s Sylhet example.—4A. Portion from the upper part of the scandent rhizome with barren fronds, nat. size. B. Fragment of a barren frond, from a lower part of the scandent rhizome, nat. size. Pratt LX XIX. Fig. 1. Polypodium distans, Don, var. minor, from a Sikkim example; a frond in fruit, nat. size. Fig. 2. Polypodium appendiculatum, Bedd., var. squamestipes, from a Sikkim example ; lower half of a frond, nat. size. Puate LXXX. Fig. 1. Polypodium subtripinnatum, C. B. Clarke, from a Neebay (Sikkim) example; basal portion e. one of the lower pinnz of the frond, nat. size. Fig. 2. Polypodium subdigitatum, Blume, from a Sikkim example ; ultimate pinna, magn. 5 diam. PrATE LXX XI. Polypodium chattagramicum, C. B. Clarke, from a Chittagong example; rhizome with one barren and ` one fertile frond, nat. size. | Prate LX XXII. Fig. 1. Polypodium clathratum, C. B. Clarke, from a Kashmir example.—A. Rhizome with three fronds, - nat. size. B. Clathrate scale from the sorus, magn. 8 diam. | Fig. 2. Polypodium subamenum, C. B. Clarke, from a Sikkim example.—A. Rhizome with one frond, 7 nat. size. B. Scale from the rhizome, magn. 8 diam. D Fig. 3. Polypodium membranaceum, Don, from a Sikkim example; fragment of a frond, nat. size (showing ` | the venation and arrangement of the sori). | SS Fig. 4. Polypodium Jaintense, C. B. Clarke, from a Jaintea example; rhizome with one fertile and one MR. C. B. CLARKE ON THE FERNS OF NORTHERN INDIA. 611 Prate LXXXIII. Polypodium cyrtolobum, J. Smith, from a Sikkim example ; rhizome with one frond, nat. size. Prare LXXXIV. Fig. 1. Lycopodium lucidulum, Michx., from a Sikkim example ; fruiting branch, nat. size. Fig. 2. Acrostichum crispatulum, Wall., from Sikkim examples.—B. Fragment from the middle of a barren frond of Wallich’s type, 6 pinnæ shown, nat. size. D. A part of one of the pinnæ of B, magn. 14 diam. A. Fragment from the middle of a barren frond of var. contaminans (sp. Wall.), 7 pinnæ shown, nat. size. C. A part of one of the pinnæ of A, magn. 1} diam. Fig. 3. Vittaria Sikkimensis, Kuhn, from a Sikkim example; transverse section of a fruiting frond, magn. 10 diam. ERRORS. On page 444, line 18 from top, dele “ Cystopteris dimidiata, Dene.” ; which is correctly referred to D. BULLATA on p. 445. On page 512, bottom line, for * F. cusrIpatUm " read N. cusprpatum. AM Cockerill dal. et Ré CBClarke SSC aN ae UE UM B 227,7) SSN y suey z pan S j J ge SE k 1 \ Pi d K wie ` : y À Mauer DEER, A x — Une ` eee a N e ea y cd - à C 7 ZU. s Beie: gees e me ) Ka ` pi rem e / / F ; j f A Z Gë ESA SS LA LA 1. NEPHRODIUM GLANDULOSUM, Ame. 2 Ui P MIA A Ve Var LÉTE STRIGOSA. TN : M Spe 3 2 aiana ania, EH, Paii aaar ananas a aaa aca D / j í € i i | ' Fitch imp. CBClarke. Trans. Linn. Soc. SeR.2.Bor. Vou J. Pap i -— j i ( Ai p Ze: K F (A / ere ^ j zs iN 4 d xe \ i E / jl = , oer Ls. J ie AM Cockerill dl ot keh NEPHRODIUM FUSCIPES , Wedd. 3 Fitch imp. d Y H Än V 7 N / P A e R ` Y X N 4 \ > ic A d G " KH N à Tana Liu Soc. Sax? "Bow Nout Pz.76. SBClarke. TRANS.LINN. Soc. SER.2.BorT.VoL.I. bm ^ TrRaws. Linn. Soc. SERZ Bor. Vou 1. Pr 78. 2 SA Smith. NEPHROLEPIS VOLUBILIS wf HV 2 dd N. ade A. T B 33. x >~ ALIE | AS o bk v, "e EI d | Z4" E v Gë E eh, CX m 27M heh DÉI pe A y Clarke. CBC = TRANS. Linn. Soc.SER2.BorT. Vor. IPr.78. | NI "y 9 LATVIA AQAA 277777771) 22272 Gë 22. ` YY, 7 N 77 Y Y P ^ Af "M POLYPODIUM DISTANS, on. - ~ | l. AM Cockerill del.et lith. g Y [UA p Wu APPENDICULATUM, Beda. Var SQUAMJEST IPES. E. E os "Se P ` CBClarke. | Trans.Linn. Soc. SERZ.Bor. Vor.LPr.80. D YT * NY ^ Nites, Ra D SU N Nal |e Sur AS \ SA SUED d ) Je AN H z5 4 ENS Ne MOS SR d SS NA Ga wie re ` Ge e E V À MN SNR qr NUN Salt f Ns € P R^ Sos. Gu, Spo M Ge | a SGN NU ho bo E E i UA. Reeg = er Cafe he e, ai a j oí p. 3 Fa es a t. P d 7*4 Le "TS" TINE Y 25 E ale, ME AA G A y. , r gi v D, qu Z4 g^ d "Lg x i Ul Zu. 2. | | LPOLYPODIUM SUPTRIPINNATUM, ZZZare AMCockerildelethth. ` ` 4i " SUBDIGITATUM, Zune. Trans. Linn. Soc. Sen? Bor Vo! PB CBClarke. POLYPODIUM CHATTAGRAMICUM, Zane. `~ C.B.Clarke. TRANS. LINN. Soc. SER2.Bor. Vou! Pu.83. ^ AM Cockerill del.et kth. -— : POLYPODIUM CYRTOLOBUM, Smith. Trans. Linn. Soc. SER.Z.Bor.Vor. I PL 84. LLYQOPODIUM LUCIDULUM, Miche. | ISPATULUM, Wadd. — 3. VITTARIA SIKKIMENSIS, Sein, This page number was omitted in the original pagination. No page or information was missed during scanning. [ 613 ] XXVI. On the Origin Ge so-called Scorpioid Cyme. By the Rev. GzoncE Hunsiew, M.A., F.L.S., F. G.8., F.O.P.S. (Plate LXXXV.) Read November 6th, 1879. Part I. (Descriptive). Ina paper in the Transactions of the Linnean Society (sec. ser. Botany, vol, i. p. 37) I have shown that when a stem or branch with opposite and decussate leaves below, bears alternate leaves above, the arrangement of the latter is very generally represented by the fraction 2, inasmuch as the order of the leaves at once assumes a spiral direction, indi- cated by drawing a line from leaf to leaf the nearest way round to each successively, and that three pairs of opposite and decussate leaves will thus form one cycle, the sixth leaf being (both originally and finally) over the first, in this alternate arrangement. There is, however, another method of arrangement, in which, instead of forming a continuous spiral line round and round the axis, a line drawn to each successive leaf returns after reaching every two successive leaves, so that it forms a curved zigzag line, the curvatures oscillating through three quarters of a circle (Pl. LX XXV. fig. 1). I first observed this method of change from opposite and decussate leaves to alternate in the foliage of a Lagerstremia in the Palm-stove at Kew, and recorded the observation in a note at the end of my paper referred to above. I have now found that a similar “ oscillating ” arrangement occurs sd the flowers and bracts of what are usually called scorpioid cymes, and which, as the sequel will show, are not definite in character, but may be more correctly termed scorpioid racemes. Fig. 1 represents a projected arrangement of the foliage of Lagerstremia, in which it will be seen that the first two pairs of opposite leaves that ** broke up " follow the usual spiral order; but from leaf number 4 the spiral returns and henceforth follows the * oscillating " method, as I propose to call it. Lathrea squamaria is another very illustrative plant. On the lower part of the stem the leaf-scales are opposite and decussate, while the bracts (though retaining the same relative positions as the scales, and therefore arranged in four vertical rows at angular distances of 90° apart) have become alternate by slight developments of the internodes. Every bract bears a flower in its axil; so that the inflorescence consists of a dense spike of four vertical rows of flowers, all, however, being twisted to one and the same side. | In this species I found several combinations of the above two methods of resolving opposite into alternate arrangements. The bracts usually commence on a spiral line for two or three coils, as in Lagerstremia, but then follow the oscillating method. In one case (fig. 2), commencing on a spiral line to the right as far as the 4th bract, the line then reversed the spiral direction to the 8th bract; the oscillating method was C14 REV. GEORGE HENSLOW ON THE ORIGIN OF then pursued to the 12th. The spiral method is once more resumed as far as the 16th braet, when it again changed and finally proceeded on the oscillating system to the apex- Such changes as described in the last example seem to be due to the fact that the flowers are so crowded that it would appear to be a mere accident, so to say, as to which of two opposite bracts became slightly elevated above its companion. The arrangement, however, is immediately affected by it. Whenever the internodes between the bracts are well pronounced in Lathrea, then the oscillating plan is for the most part adhered to with precision. Fig. 3 represents another example of Lathrea in which frequent changes were exhibited. It will be easily understood from what has been explained with reference to fig. 2. The next plant to be referred to is Silene pendula. De Candolle *, in describing the scorpioid cyme, refers the reader to species of Silene with *'spicate" or, rather, “racemose” inflorescences, as being reduced forms of the * dichotomous eyme " so common in other species of Silene as well as of the Caryophyllee generally. “In this case," he says, ** the flowers are generally arranged on one side "— i. e. more or less inclined to be secund—“ either by a tendency of the branches to abort on the same side, or by a torsion of the axis. The branches or stems in which this dis- position takes place are in general before their development rolled into a volute on the exterior side;" and he mentions Drosera, *spicate" species of Silene, Echium, and other Boraginaceous plants. De Candolle designates such unilateral inflorescences by the name cimes scorpioides T. Before I was aware that De Candolle had referred to Silene at all, my attention happened to be drawn to the subsecund arrangement of the flowers of Silene pendula; and it occurred to me that it might throw light on the origin of the scorpioid cyme, which I had always thought to be erroneously explained in text-books. The inflorescence of Silene pendula (fig. 9) consists apparently of opposite and decussate bracts; and if it be held before the observer so that one flower faces him, the next flower is in the axil of the bract (say, to the right hand); the next is over the first and also faces him, the fourth being over the second (again on the right hand), and so on. Hence the flowers are in two vertical rows at an angular distance of 90° apart. On the other hand there are two rows of bracts, also 90° apart, which have no flowers in their axils. Moreover, the bracts beneath the flowers are all much more reduced in size than those opposite to them, showing a strong tendency to be arrested. Lastly, there is a great propensity for all the flowers to turn to one side. It may be added that in Silene there is no tendency to produce any disarrangement in the opposite and decussate bracts +. * Organographie végétale, i. p. 413 seg. + He here uses the word “ volute;" but it does not appear to have been intentional to express the conical form of spiral indicated by the term, but rather the flat; “ helix.” + Silene pendula has all the appearance of being a monopodial raceme; and had we not other members of the o genus (or of the family Caryophylleæ), such as S. calabrica, which produce true dichotomous cymes below and ` similar “ racemose " branches above, it would be impossible to interpret the inflorescence of the species aright. - As 2 » it is, ho ue, I think we miser tae each flower as a terminal blossom, one ee ee ae veloped, THE SO-CALLED SCORPIOID CYME. 615 If we now pass to genera of the Boraginec, we find they furnish us with inflorescences both bracteate (as Borago) and ebracteate (as Myosotis). Ifin the former a line be drawn the shortest way from, say, right bract to flower, left bract to flower, and so on continu- ously, it will be found that, commencing with No. 4, fig. 1 will exactly represent the positions of the bracts and flowers. Hence, as far as the order in the arrangement of the bracts and flowers are concerned, it is completely interpreted by the oscillating method. The reader will now perceive the significance of the examples I have taken; for, regarding the scorpioid cyme as monopodial and not sympodial, as it is usually considered, I derive it from opposite and decussate bracts, which have passed into alternate by the oscillating method (the foliage of the stem having adopted the spiral method). The preliminary stage of this is represented by Lathrea, though it is often uncertain in its method of alternation, and, moreover, retains a flower in the axil of every bract. Silene pendula only aids us in showing how one of two opposite bracts may have a strong tendency to be reduced in size when a flower-bud is produced above it. Indeed, as observed in the note above, not only is the bract reduced, but the bud in its axis is suppressed altogether. Combining these effects in Borago, we find the floral bracts completely gone, while the alternation by oscillation is thoroughly established. Other facts remain which demand interpretation. From the preceding explanation of Lathrea it would be inferred that the bracts should be in two vertical rows at first exactly opposite to the two rows of flowers, respectively ; and assuming, on the separation of opposite bracts, that the new internodes would grow to about the same length, then one of the bracts should be on a level, about halfway between any two flowers, and on the opposite side of the stem. Such is not quite the case. The bracts have undergone a further change in position; they have been “spirally uplifted,” so that instead of being lower down on the opposite side of the axis, each is uplifted to a position a little lower than that of the flower, but very near to it, as shown by the dotted lines in the figure of the floral axis of Hyoscyamus (fig. 4, bb). In using the expression “ spirally uplifted,” I only mean that the bract is, on the one hand, raised vertically a little above its theoretical position (or exactly halfway between the heights of any two flowers), and, on the other hand, shifted horizontally towards the nearest flower, these two motions, so to say, making together a slight spiral turn. This will account for their otherwise anomalous position close beside the flowers. Indeed, so close are they in some cases, that the latter might even. be supposed to spring from their axils. This is especially deceptive when the bracts are inserted by broad bases ; but the flowers are never immediately over the midribs of the bracts. That bracts can be uplifted, either by an elongation of the axis or by adhesion to it, is the other, namely that between the floral pedicel and its adjacent bract, being arrested; so that the whole forms a definite, sympodial, and racemose inflorescence with opposite and decussate bracts. De Candolle refers to this as explaining thescorpioid eyme ; but it will be seen hereafter that he, Payer, and other writers have confounded such sympo nodial “ racemes " and “spikes” (adopting such terms for what are really definite inflorescences) with what I now propose to call the (truly indefinite) scorpioid raceme. | SECOND SERIES.—BOTANY, VOL. I. 40 616 REV. GEORGE HENSLOW ON THE ORIGIN OF a fact well known and often commented upon. Thus Guillard observes :—** La Feuille aisselióre est surhaussée par adhérence à son axillaire’’*. Bravais also alludes to les soudures des feuilles as of common occurrence f. The elevation of the bracts in the bracteate scorpioid inflorescences is very slight; while the shifting of their positions is quite in accordance with the normal change from the 2 spiral to the distichous arrangement in the Cherry-Laurel, Ze In that plant, whenever a shoot grows vertically the leaves are mostly to be represented by Z or $ ; but when (as is their usual habit) the branches grow horizontally, then the leaves are distichous. Similarly other changes are not at all unfrequent. Leaves may be 5 at the base, $ higher up, and 35; above, as Payer describes them in Echinocactus ; not to add the fact, on which the thesis of this paper is based, that the alternate arrangements have arisen out of opposite and decussate. ! This common fact, of a change of position of a leaf, proves that the point of emergence of a leaf-papilla can easily be shifted according to circumstances of growth; thus gravity is possibly an active eause in producing the distichous arrangement in dieotyledons with horizontal branches. In the case of the scorpioidal inflorescence, perhaps some other influence may determine the emergence of the bracts and flowers being all on one side of a diameter drawn across the flower-stalk; for it will be observed that the two rows of bracts, instead of being 90° apart, are now diametrically opposite to each other, 4. e. each bract has passed through an angle of 45° towards the side which bears the flowers. Similarly with regard to the flowers, instead of being in two vertical rows 90° apart, as in Lathrea, the rows have become drawn nearer to each other, having passed through an angular distance together of about 30°. This approximation is apparently fore- shadowed in the strong inclination to a secund arrangement in Lathrea. Thus, I believe, do we obtain the characteristic inflorescence of those genera Ss Boraginee and Solanee of which the scorpioidal inflorescence consists of two rows of flowers together with two rows of bracts—such, for example, as of Symphytum, Borago, Iycopsis, and Hyoscyamus. In Myosotis the bracts have all become completely arrested, so that there is eet | left but the two parallel rows of flowers; and as the pedunele or morphological axis elongates, the flowers become separated. There yet remains a further observation to be made. The pair of scorpioid racemes which terminate the axis of Myosotis and other species of the Boraginee often have the appearance of dichotomy; sometimes, indeed, again and again repeated, as in Heliotropium, and not infrequently with a flower situated in the fork, so that the two branches curl in opposite directions. This appearance has, I believe, been the source of an error in the usual interpretation. Ifa large number of specimens be examined, the above description will be accurately true for some of them; but more frequently the lowermost flower * Bull. Soc. Bot. de Fr. iv. p. 933. ie + Ann. des Sc. Nat. 2° sér. vii. p. 298. As further references to the uplifting of bracts the reader is — to: e um Paver, ‘Éléments de Botanique, p. 116. Wanwze, Botanisk Tidskrift, 1869, Bind 3, Tab.1; also, Viđensk. Belsk. Skr. 5 Række, Naturvidenskabelig., Afd. Bd. 10; French résumé, ‘Recherches sur la ramification Hes or p. xxii, Barrow, “ Sur les lois de l'entrainement dans les — eegener ` P THE SO-CALLED SCORPIOID CYME. 617 is on one or other of the branches; and not exactly between them. Secondly, two flowers may sometimes, but rarely, be found in the fork instead of one. Thirdly, two opposite leaves may sometimes be seen, one below each branch. Fourthly, axillary scorpioid racemes spring from the axils of several of the upper leaves of the stem. - . The interpretation I would offer is the following :—The inflorescence is really indefinite : of the two branches, one is terminal, the other is strictly axillary; but an adhesion, with occasional uplifting of the bracts as well, usually takes place between the terminal and last. axillary branch. Whenever this adhesion ceases exactly at the lowest flower of either branch, it gives rise to the appearance of dichotomy with a terminal flower in the fork*. Whenever the two uppermost leaves, which should be separated by an inter- node, are occasionally brought together, then they are only abnormally opposite. The inflorescence of Myosotis will illustrate this interpretation. A selection has been made and represented in the following figures out of some hundreds examined of several species of this genus :— Fig. 10 represents, diagrammatically, an unequal development between the terminal and the last axillary branch; both carry bracts. Fig. 11. The axillary branch is here nearly equal in strength to the terminal; the bracts are arrested. | Fig. 12. The bracts are likewise arrested on both the terminal and the axillary branches, which are still more nearly equal. Fig. 13. The terminal and axillary branches are of equal growth, and an adhesion has now taken place between them from a to b. Fig. 14 shows a greater degree of adhesion, in that it has extended to the lowest flower, which consequently appears to issue from the fork, giving the appearance of dichotomy. | E - Fig. 15 shows excessive adhesion beyond the lowest flower, so that four flowers rise from below the fork. : Fig. 16 illustrates a case where the leaf out of the axil of which the branch has sprung is “uplifted”? beyond the lowest flower on the axillary branch. ; | Fig. 17 exhibits excessive adhesion as in fig. 15, with the uplifting of the bract as well. . This condition of a leaf or bract being uplifted is not without frequent parallel else- where. In the Crassulacee, where modified cymes often occur, such uplifting as well as elevation by adhesion is far from being uncommon (see fig. 5). | It will not be unadvisable here to summarize the successive steps or processes through which, as I suppose, the inflorescence has passed im order to arrive at the condition presented in the so-called scorpioid cyme of Myosotis. -* Bravars, I find; had long ago anticipated this observation, for he observes:—“ dans la plupart des Borragindes qui offrent des cimes scorpioides axillaires, le premier pédoncule de chaque cisco emm Ss au moins PE: la tige centrale " (Ann. des Sc. Nat. 2° sér. vii. p. 298, 1837). He also notices how this union gives rise to * double" scorpioid . . eymes (7. c. p. 300). poe ps de qu : Lo SÉ 618 REV. GEORGE HENSLOW ON THE ORIGIN OF 1. Opposite and decussate bracts with axillary flowers have given rise to alternate by the oscillating system. This stage is seen in Lathrea. 2. The suppression of the flowers in the axils of two of the vertical rows of bracts with a diminution of the size of the floral bracts, as exemplified by Silene pendula. 3. In both Lathrea and Silene pendula the tendency of the flowers to be secund foreshadows the actual approximation of the bracts and flowers in Borago. 4. The four vertical rows have shifted their positions, passing to one side of the axis, all being situated within a semicircle (if projected). Two rows have lost their bracts, the flowers remaining; the other two rows have lost their flowers, the bracts remaining (as seen in Borago, &c.). 5. The bracts have become “ spirally uplifted ” and so take up a position close to, but at a slightly lower level than, that of the flowers (seen in fig. 4, Nadine 7a _ 6. The bracts may be all suppressed, as in Myosotis. 7. Adhesions of various degrees may take place between the terminal and axillary racemes, giving rise to the apparently more or less dichotomous character of the in- florescence of the Boraginee, and which often contains a seemingly terminal flower at the fork. ` 8. The unequal power of growth between the upper and underside of the peduncle, induced by the flower-buds being all on one side, has caused the characteristic scorpioid form of the undeveloped apex. Part II. (Critical). I have remarked that the interpretation of the so-called scorpioid cyme as given in botanical text-books appears to me to be erroneous; it will be desirable to point out ` what I believe to be the errors involved in the usual explanations. As Payer ‘ Eléments de Botanique' gives a rather fuller account than is to be found in many other text- books, I will direct my remarks especially to his views. This author evidently confounds two totally different kinds of inflorescence under the common terms cyme wnipare scorpioide. Thus the inflorescence represented by Pl. LXXXV. fig. 5 t of Sedum album is of quite different origin to that of Hyoscyamus (fig. 4) 1, the former being a true sympode; the latter, as I have shown, is of a monopodial origin. If we compare fig. 5 with fig. 6 §, the origin of the former is obvious ; but, then, it is not a scorpioid eme but a degraded form of dichasium or “ dichotomous eme" and is, in fact, a *spicate" cyme. The dotted lines will indicate the branches which are sup- pressed alternately right and left at each dichotomy. Had they been uniformly and always suppressed in one and the same direction, the result would have been such another spicate cyme as is indicated by the right-hand portion of fig. 6, as shown by the letters a,b,c. This would be the correct result of the process usually but MOES M "s giving rise to the Boragineous scorpioid cyme. * Uplifting and adhesion of leaves may be often seen in many genera of the — E + Reproduced from Payre, J. c. fig. 191, p. 121. t See Payer, J. c. fig. 155, p. 98. o$ This fig. represents a portion of Paxxz's fig. 188, l. c. p. 118. THE SO-CALLED SCORPIOID CYME. 619 ‘The following is Payer’s definition of the scorpioid cyme (J. c. p. 100) :—* La cime unipare scorpioide peut donc être définie: Une inflorescence dont toutes les fleurs sont de génération différente, oppositifoliées et rangées sur deux séries seulement.” Payer regards this cyme as being due to an alternate and quincuncial arrangement of the bracts of successive sympodial axes, they being also heterodromous successively, so that any bract is either to the right or left of the preceding and at an angular distance of two fifths of the circumference, or 144^. Duchartre gives a similar description in his * Éléments de Botanique 2nd ed. p. 588, and is apparently followed by Sachs *, at least so I interpret the paragraph on p. 160 of the English edition of his Text-book. This description of Payer's is far from being in accordance with facts, as seen in the so-called scorpioid cyme of the Boraginee, &c. The plants selected by Payer for illus- tration are Hyoscyamus and Sedum album. In the first place, in the scorpioid cyme, as of Hyoscyamus and Borago, no bract is ever exactly opposite to any flower, though it is so in Sedwm. The bracts, as I have de- scribed them above, are situated alternately to the right and to the left of each flower successively, very near to them, but a little below the level of their insertions +. Secondly, they are not placed at an angular distance of 144°, or two fifths of the circum- ference, but form two rows diametrically opposite to each other, while the flowers form two rows also, but in vertical planes at an angular distance of about 60° apart. In Sachs's Text-book (p. 159) the cyme is treated somewhat abstractedly, and that author makes no reference to bracts, which are of very great importance in any explana- tion of the inflorescence. , The editor of the English edition has pointed out that fig. D on p. 159, which I have reproduced for convenience (Pl. LXXXV. fig. 7), though described by Sachs as helicoid or bostrychoid, corresponds with what is usually called scorpioid in several of our text- books. Sachs, however, is more nearly correct, while the latter term is decidedly wrong. Figs. A and B of Sachs correspond to Payers cyme unipare, as of Sedum album (P1. LXXXV. fig. 5), both being sympodes and presumably having the suecessive branches oppositifoliar, though Sachs makes no allusion to bracts. Sachs’s description, therefore, applies to the “spicate” cyme, though designated scorpioid eme both by him and Payer, and not to the true scorpioid cyme, or rather, as I would now propose to call it, “ scorpioid raceme.” With regard to fig. C (Sachs, p. 159), which represents the usual form of the so-called dichasiwm of Schimper, or dichotomous cyme of the text-books, it is not clear whether this figure is supposed by the author to represent accurately a natural state of things (which it does not, as all the branches are in one plane), or whether it is only a projection of the inflorescence on to the plane of the paper, as, indeed, I take Sachs to assume the reader to perceive. The inflorescence itself in nature is more or less corymbose, as may be seen in many of the Caryophyllee, Radiola, &c.; and it must be - * Sacus, while calling this degradation of a dichasium a scorpioid eyme, recognizes the inflorescence of the oragineæ as of a different character (* Text-book,’ p. 522, Engl. ed.). | d o dun. = + geo PL LXXXV. fig. 4, in which b indicates the position of the insertion of the bracts; Jf represents the he. . pedicels which bore the capsules now removed. | | 620 REV. GEORGE HENSLOW ON THE ORIGIN OF carefully remembered—for the neglect of this fact lies at the bottom of the common misinterpretation—that in reality the direction of each successive pair of bracts. (or a common line drawn along their midribs) is in a plane at right angles to that of the pre- ceding pair; so that by suppressing all the branches to the right or to the left succes- sively, we should not get a flat helix, but an open angular spiral, as shown in the projeetion, Pl. LXXXV. fig. 8. Fig. 7 (fig. Din Sachs's Text-book, p. 159) would seem to be intended by that author to illustrate this, if we suppose the members 2-3, 3—4, 4-5, 5-6, Ee to be in planes at right angles to each other successively, and only the members 1-2, 2-8 to bein the plane - ofthe paper. There is nothing to show that the author did not mean this to be the case. I am, in fact, inclined to infer that he did; but even then it will not correspond with the helicoid cyme or bostryx of Schimper, which is derived from a system of axes arising from alternate and not opposite bracts, as in the inflorescence of Alstreemeria—a true helieoid cyme being a sympode with an homodromous development of bracts and floral axes, each successive bract being supposed to be at an angular distance of two-fifths of the circum- ference from. the preceding ` whereas in Sachs’s description the supposition is that the successive axes are in planes at right angles to one another, since he bases the whole on the false dichotomous cyme. We might, however, include fig. as another and new form of bostrychoid, presumably derived from opposite bracts, while the usually described bostrychoid or helicoid is only ` obtained from alternate bracts. | If, however, we adopt the supposition that fig. D is derived from fig. C, and is identical with the scorpioid cyme of descriptive botany, we at once fall into the same error which all our text-books have perpetuated. It is true that fig. D can be derived from fig. €; but then, as stated above, it does not give rise to the true scorpioid cyme, but to one form of spicate cyme, such as of fig. 6, a, 5, c, this being characterized by having a single row of flowers only, while the seorpioid eme has a double row. If we regard Silene pendula as a sympode, then it would fall under the same kind of inflorescence, or * racemose " eme, Another theory has been offered by Kaufmann *, and supported by Warming 1, that the scorpioid inflorescence of the Boraginee and Solanee is due to a repeated dichotomy of the vegetative apex, one of the papille thus produced developing into a flower, the other continues the apical growth. As, however, both papille are at first exactly alike, the term dichotomy would seem only to introduce a misleading idea, for it is identical in every respect with budding. The description of the process given by Kaufmann *, as well as his figures for Asperugo, which is a bracteate form, correspond in the order of — development exactly with the interpretation given in this paper, as follows:—Left bract— _ flower to the right of it; right bract—flower to the left of it; left bract &c. as before ` (that is, when the convex side of the scorpioid inflorescence is held before the observer). ` ` CH Zem Zeit. 1869, p. 886; also Nouv. Mém: de la Soc. Imp. des Nat. de Moscou, xiii. p. 248, plate xxi t “ Recherches sur la ramification des Phan¢rogames,” Vid, Selsk. Skr; 5 R; Afd. Bd, 10, 1 (with French abstract THE SO-CALLED SCORPIOID CYME. 621 He correctly observes that the flowers are not in the axils of the bracts; for had they been, they would not be situate so close together; and he notices that if they had been axillary, the plane which divides them would then have been parallel to the bracts, whereas it lies obliquely and alternately to the right and to the left of the observer as above placed. | This last observation has probably suggested to his mind the theory of dichotomy, for it certainly would have been difficult to account for the two rows of flowers by any other method known. I trust the reader will now perceive that the development of the bracts and flowers by simply budding out in the usual way from the axis, in accordance with the oscillating method described in this paper, fully accounts for their relative positions, and does not in the least depart from Nature's normal method of procedure. Lastly, the circinate apex of the scorpioid inflorescences is probably due simply to the greater development of the upperside of the axis, in consequence of the buds and braets, when present, being all within the upper and longitudinal half of the stem *. DESCRIPTION OF PLATE LXXXV. Fig. 1. Projection of the alternate leaf-arrangement of Lagerstremia according to the “ oscillating ” method. ‘The first four leaves are spirally arranged, which follow opposite and decussate pairs ; but subsequently the line passing through the leaves successively up the stem oscillates through 270°, or three quarters of a circle. ; Figs. 2 and 3. Two projections of the line connecting successive bracts of inflorescence of Lathrea squamaria. Both systems of resolution, from opposite and decussate to alternate arrangements, are followed. When the internodes are well pronounced, then the oscillating system is followed continuously, as from numbers 16 to 26 in fig. 2. Fig. 4. The peduncles (A and B) of Hyoscyamus niger, the bracts (b) and flowers (fl) removed, the letters indicating their positions. The dotted spiral lines show how the bract has been “ uplifted” from its (theoretically) normal position, on a level halfway between the flowers. Fig. 5. A portion of the inflorescence of Sedum album (after Payer), showing, by dotted lines, (1) the suppression of alternate lateral axes, and (2) the uplifting of bracts above their, theoretically, normal positions opposite to the flowers. Fig. 6. A portion ofthe dichasium, or false dichotomous cyme, of Sedum oppositifolium (after Payer), to indicate the origin of spicate forms of sympodial cymes (e. g. a, 5, c). . Fig. 7. Projection of a “ racemose cyme " (after Sachs), called “ bostrychoid " by that author. Nos. 1, : 2, 3, 4, 5, 6, as referred to in text. Fig. 8. True projection of a racemose or spicate cyme, falsely called “ scorpioid cyme ” by Payer, &c., . helicoid by Sachs, as deduced from a dichasium or “ dichotomous cyme.” Fig. 9. Inflorescence or “racemose sympode ” of Silene pendula. Figs. 10 to 17. Diagrammatic illustrations of the inflorescence of Myosotis, to indicate the origin of the falsely dichotomous appearance sometimes seen. It consists of a terminal and axillary raceme, with their basal portions more or less adherent. See p. 617 for descriptions. * With regard to the descriptions of the scorpioid cymes by other writers, I need do no more than allude to the latest works on the subject, such as Eichler's * Bliithendiagramme,’ and to Prof. A. Gray’s ‘ Text-book,’ part i. p. 156 (1879), who reproduces the classification of cymal inflorescences by that author, such being regarded by them as This page number was omitted in the original pagination. No page or information was missed during scanning. GHenslow del. App ith. 15 79 2j 17 Trans.Linn. 50c. SER. 2. Bor. VoL ln. 85. 25 79\\23\ 27 29 Fig.12. Fig 16. | Fig IT. Pe DIAGRAMS ILLUSTRATING THE SCORPIOID RACEME. Aconiopteris obtusa, 578. Acrophorus assamicus, 443 ; Hookeri, 443; immersus, 443; nodosus, 444; pseudo-cystopteris, 444; pulcher, 444 ; Thomsoni, 443. Acropteris australis, 505; radiata, 505. Acrospira (char. gen.), 255; aspho- deloides, 255, Acrostichum aleieorne, 595; angu- latum, 576 ; appendiculatum, 577, 578, 596, 603, 604, var. Hamil- | toniana, 578 ; asplenifolium, 577 ; aureum, 582, 596; australe, 505; axillare, 579, 596, 601; Blume- anum, 580; conforme, 576, 577, 595; eontaminans, 596; con- tractum, 603; coriaceum, 595; costatum, 580, 581, (Wall.) 596, var. (Hk.) 572; crispatulum, 580, - 581, 596, var. Blumeana, 580, var. contaminans, 580, 596, var. pseud-undulata, 581; danesfo- INDEAS mulariefolium, 554; obovatum, 554; palustre, 577; proliferum, 581; punctatum, 561; punctula- tum, 596 ; radiatum, 505 ; repan- dum, 580; rigidum, 596; rivu- lare, 579, 603; scandens, 577, 596; septentrionale, 478; setosum, 577, 596; speciosum, 582; spi- catum, 582; stelligerum, 577, 604 > stigmatolepis, 577; subcrenatum, 581; terminans, 581, 604; tha- lictroides, 471; Thelypteris, 517; | trieuspe, 581; triquetrum, 473, 596; variabile, 578, 579, var. lanceolata, 579; vestitum, 603; . virens, ( Wall.) 581, 598, 603, 604, (Bk.) 580, (Hk. & Grev.) 581, var. (Hk.) 580, var. (Hk. & Bk.) 572, viscosum, 577, 581, 595; vivipa- | rum, 577, 578, 596 ; Wightianum, 577, 603. Actiniopteris dichotoma, 505; radi- ata, 505. lium, 582 ; decurrens, 595 ; dicho- tomum, 505; diversifolium, 580 ; emarginatum, 582; Finlaysonia- num, 603 ; flagelliferum, 579, 581, 596; fuciforme, 595; Gayanum, 576; glandulosum, 576 ; gorgo- neum, ( Kaulf.) 578, (Bl.), 576; Ha- miltonianum, 577,596; heterophyl- lum, 575; hybridum, 595; in- | æquale, 582; laurifolium, 576; | ludens, 577, 578, 605; Marante, | - 567; marginatum, 576,595 ; minus, | 579; neriifolium, 577, 595 ; num- | Actinostachys digitata, 583, 595. Adenotheca aphylla (Welw.), 255. Adiantum amenum, 454, 597; ca- pillus, 453 ; Capillus-Veneris, 453, 597; caudatum, 453, 597, var. rhizophorum, 453 ; ciliatum, 453 ; dolabriforme, 452; Edgworthii, 453; filiforme, 452; flabellatum, 454, 597, 604 ; flagelliferum, 453, 597; hirsutum, 453, 604; hispi- dulum, 454, 604; lunulatum, 452, 597; mierophyllum, 453; peda- tum, 453; proliferum, 453; reni- * The names of species adopted by the author precede synon SECOND SERIES.—BOTANY, VOL. I. yms; the latter are in alphabetical order. forme, 597; repens, 442; rhizo- phorum, 597; scabrum, 597; sobo- liferum, 453, 597; tenerum, 453; venustum, 453, 597 ; vestitum, 453. Adoxa, structure of, Rev. G. Hens- low on, 177. Æstivations, origin of floral, 177. Agaricini, 119 ; of Brisbane, 399. Agaricus fimicola, 119; Gardneri, 399; junceus, 119; semiorbicu- laris, 119; spadiceus, 119; syl- vaticus, 119. Agasta, 59; asiatica, 61 ; indica, 63; splendida, 60. Aglaia odorata, distrib. of, 233. Ahuta = Agasta splendida, 60. | Aira, awn of, 164. | Albinism in Balsams, Dr. Lowne's . exper. on, 328. Albuca angolensis, 251; chlorantha, .251; galeata, 251; monophylla, 251; myogaloides, 250; subspi- cata, 251. ; Alismaceæ, self-fertilization of, 378, 393. Allantodia denticulata, 490 ; incisa, 490; javanica, 505, 599; sylva- tica, 501; tenella, 491. Allium angolense, 262. Allosorus calomelanos, 461 ; crispus, 459, 460; gracilis, 460 ; Stelleri, 460. Aloe andongensis 263; angolensis, 263 ; littoralis, 263; palmiformis, 963; platyphylla, 264; zebrina, 264. H ÅP 624 Alsophila Andersoni, 433; Bruno- niana, 427, 430, 431, 605, ( Bedd.) 432; comosa, 433; contaminans, 432; decipiens, 430; gigantea, 433; glabra, 433, 496; glauca, 431, 433, 605; glaucescens, 432, 605 ; Grevilleana, 449, 605; Hel- feriana, 433; khasiana, 432; latebrosa, 429, ( Bedd.) 432, (Hk.) 431, var. hemitelioides (J. Scott) 430, var. Schmidiana (Kze.) 432; Metteniana, 433; Oldhami, 432, 433; ornata, 432; Scottiana, 433 ; Sollyana, 432 ; Wallichiana, 432. Ampelopteris elegans, 548; firma, 548. Amphicosmia decipiens, 430. Androscepia arundinacea, awn of, 164. Anemia flexuosa, 596. Anemone montana, burying mecha- nism of achene of, 164; torsion of awn of, 158. Angiopteris crassipes, 585, 599; evecta, 585, 599. Angolan Herbarium of Welwitsch, J. G. Baker on the Liliaceæ, Iri- dacee, Hypoxidaces, and Hæmo- doraces of, 245. Anisogonium (subgen.), 503; esculen- tum, 503; heterophlebium, 503; se- rampurense, 503; sylvaticum, 498, Anisostemon (sect. DI), 113. Anthericum andongense, 257; arena- rium, 259; benguellense, 257 ; calyptrocarpum, 258; colubrinum, (Welw.) 256; dissitilorum, 957; | drimiopsis, 259 ; faleatum, ( Welw.) 256; Jacquinianum, 259, var. affinis, 259; limosum, 257; molle, 259; orchideum, 258; pleiosta- chyum, ( Welw.) 255 ; pterocaulon, 258; pyreniearpum, 259; tenel- lum, 256 ; ustulatum, 258. Anthesteria ciliata, awn of, 164. Antholyza huillensis, 270. Antrophyum Boryanum, 573, 596; coriaceum, ( Wall.) 572, 596, (BL) 570; faleatum, 573; involutum, 570 ; lanceolatum, 570 ; latifolium, 572, 573; parvulum, 573; plan- tagineum, 572, 573; plicatum, 572; pumilum, 596; reticulatum, INDEX. (Kaulf.) 573, 596, (Bedd.) 572, (Fée) 572, (Wall.) 572, var. par- vula, 573. Apocynaces, plumes of seed of, 165. Appunia (Hook. f.), 169. Arcyria umbrina, 128. Aristea angolensis, 270. Arrhenatherum, awn of, 164. Arthobotrys Avana, 522, 603; macro- carpa, 522, 603. Arthronia punctilliformis, 146. Asparagus africanus, 254; angolen- sis, 254; benguellensis, 253; de- flexus, 254; drepanophyllus, 254 ; equisetoides, 253; psilurus, 253 ; pubescens, 254 ; racemosus, 254. Aspergillus glaucus, 406. Aspidium abruptum, 534; aculeatum, 508, 509, (Swiz.) var. 603, var. biaristata, 510, var. biaristatum f. Khasiana, 511, var. lobata, 509,var. mucronifolia, 509, var. rufo-barbata, 509, var. semifertilis, 509, var. se- tosa, 510 ; acutum, 541 ; adnatum, 518; aduncum, 603; affine, 509, 603; alatum, 536, 603; amabile, 510, 602; amboinense, 532; an- gustifrons, 514; anomophyllum, 512; apiciflorum, 518, 602; ap- pendiculatum, 516, 543, 602; ari- dum, 531; aristatum, 511, 529, 602, var. affinis, 511, var. assamica, 511; articulatum, 603; Athyrium, 491; Atkinsoni, 506; atratum, 513, 603; auriculatum, (Swtz.) 506, 602, 603, 605, (Don) 507, ( Wall.) 540, var. ceespitosa, 507, var. lenta, 507, var. marginata, 507, var. normalis, 507; biaristatum, 511; Brunonia- num, 522, 602; caducum, 512, 603, 605 ; catophoron, 524; cxspitosum, 603; canariense, 521; canescens, 533, 602; canum, 515, 533, 603 ; caruifolium, 511; caryotideum, 512, 603; cicutarium, 539; cilia- tum, 515, 602; coadunatum, 540, 603; coniifolium, 511, 602; co- nioneuron, 529 ; conjugatum, 535; cordifolium, 540; coriaceum, 605; cornu-cervi, 511; crinitum, 602; cristatum, 519; cucullatum, 530; cuspidatum, 513; decurrens, 539; densum, 523, 603 ; discretum, 510; divisum, 527, 546, 603 ; Donianum, 593, var. australe (Mett.) 52 520; drepanopterum, 493; ebur- neum, 486, 493, 523, 603; elon- gatum, 520; eriocarpum, 525, 602; eusorum, 534; exaltatum, 541, 603; extensum, 529; falcatum, 512, 603; Filix-femina, 491; Filix-mas, 519; fimbriatum, 494, 602; Finlaysonianum, 605; flac- cidum, 513; flagelliferum, 605; feeniculaceum, 427, 434; folio- losum, 434, 444, 602; fontanum, 484; fuscipes, 534, 602; glandu- liferum, 516, 602; glanduligerum, 513; gracilescens, 513; grandi- folium, 537; Hamiltonianum, 605; hirsutulum, (Wall. 515, 605, (Ham.) 605, (Schk.) 541; hirtipes, 513; ilicifolium, 508, 608; imbricatum, 540; inter- medium, 526; Lachenense, 506, 510; Leuzeanum, 535; Lonchitis, 505 ; macrocarpum, 489; macro- phyllum, 539; marginatum, 496, 521, 602, 603; Maximowiczianum, 511; megaphyllum, 532; membra- nifolium, 535; microcarpon, 561; molle, 533; multicaudatum, 540 ; multidentatum, 443, 602; multi- jugum, 529, 533, 602; multili- neatum, 534, 602; musefolium, 542; neriiforme, 542; nitidulum, 603; nodosum, 444; nymphale, 530 ; obliquum, 507; obtusifolium, 540 ; ocellatum, 507, 602; octhodes, 516; odoratum, 525; paleaceum, 520; palmipes, 511; paludosum, 544; parasiticum, (Herb. Madr.) 605, (Swtz.) 533, 605; patentis- simum, 520, 602; pennigerum, 532; pilosulum, 448, 525, 602; pilosum, 541; polyblepharon, 510; polymorphum, (Bedd.) 538, 603, ( Wall.) 537, Prescottianum, 508, 510, 602, var. Bakeriana, 510, var. castanea, 510 ; prionophyllum, | 529, 534, 602; procurrens, 530 ; ! prolixum, 516; pteropus, (Kze.) 539, ( Mett.) 539; puberum, 448, 515, 602; pungens, 508, 603; ` purpurascens, 523, 524; remotum, | 523; repandum, 537; rhom- boideum, 602; rigidum, (Swtz.) ` : riparium, 602; rostratum, 537, 603 rufo-barbatum, 603 ; Schkuhrii, 541; semibipinnatum, 603 ; setosum,603 ; siifolium, 538, 539; sikkimense, (Bk.) 427, (Hk. & Bk.) 525; Simonsii,537; singaporianum, 603; solutum, 533, 602; sophoroides, 605; speciosum, 511; spectabile, (BL) 518, (Wall.) 435, 526, 603, splendens, (Ham.) 605, (Willd.) 541; squarrosum, (Don) 509, ( Wall.) 489, 602 ; stimulans, 508; subconfluens, 536 ; subdiaphanum, 525,602; subdigitatum, 546 ; sub- lanosum, 541, 602; syrmaticum, 518; Tavoyanum, 540, 603; tec- tum, 533, 603; Teijsmannianum, 538, 539; Telfairianum, 603; tenella, 491; terminans, 603; Thelypteris, 514, 517; Thomsoni, 508, 510, var. gracilis, 508 ; trifo- liatum, 605; truncatum, 534; uliginosum, 520, 528; unitum, 529, 602; variolosum, 538, 603 ; vastum, 536 ; venulosum, 531, 602; vestitum, 510; Wallichianum, (Spreng.) 520, (Wall.) 542 ; Wal- lichii, 542; Weigleanum, 523; xylodes, 516; Zollingerianum, 538. Asplenium abscissum, 481; acrosti- choides, 488; acuminatum, 498, 500, 599; Adiantum-nigrum, 483, 485; alternans, 476, 600; alter- nifolium, 497; ambiguum, 498; amcenum, 481; anceps, 477; an- thriscifolium, 482; arborescens, 503; argutans, 498; arifolium, 576; asperum, 502; aspidioides, 494; Atkinson), 487, var. An- dersoni, 487; auritum, 600; aus- tralasicum, 475; bantamense, 497, 504, 599, 606; |. Beddomei, 479 ; bellum, 496, 502, 603 ; bipinnatum, 503; brevisorum, 494, 599; Brunoniana, 505; Bru- mnonianum, 505; bulbiferum, 485, 599; bullatum, 485, 599; cespi- tosum, 482, 599; castaneum, 477; . caudatum, 600; Ceterach, 504; cheilosorum, 481; Clarkei, 489, 599; concinnum, 485, 599; con- taminans, 604; contiguum, 479 ; | eoriacoum, (Fio) 470, (Roxb) 480; Gg 4905 nicl, 479, 90; | australe, 505; ; INDEX. cristatum, 481, 599 ; cultratum, 604; cuneatum, var. splendens, 484 ; Dal- housiæ, 476 ; decipiens, 489, 495 ; decurrens, (Bedd.)503,( Wall.) 481, 599; decussatum, 499, 604; den- sum, 477; dentigerum, 492; depau- peratum, 482, 600; dilatatum, 502; divaricatum, 604; diversi- folium, 502, 599; Donianum, 497 ; drepanophyllum, 487; eburneum, 493; elatum, 498; elegans, 497; emarginato-dentatum, 481; ensi- forme, 476, 595, 599, var, aus- tralis, 476; erectum, 478, var. trapeziforme, 479 ; eroso-dentatum, 481; erythrocaulon, 481; escu- lentum, 503, 599, 604; exiguum, 484; falcatum, 479, 480, (Zamk.) yar. 599, 600, 603, var. caudatum, 480, var. macrophyllum, 480; fallax, 489; Filix-femina, 489, 491, 600, 602, var. attenuata, 492, var. dentigera, 491, var. flabellu- lata, 493, var. Parasnathensis, 493, var, pectinata, 492, 495, var. poly- spora, 493, var. retusa, 491, 492, var. tenella, 492; fimbriatum, ( Hk.) 494, (Kze.) 485, 489, 491, (Hk.) var. 602, var. foliosa, 495, var. leptophyllum, 485, var. sphæro- pteroides, 495; Finlaysonianum, (Hk.), 504, 605, ( Wall.) 480, 599; flagelliferum, 478, 599; foliolosum, . 492, 604; fontanum, 484 ; fraxi- nifolium, 497, 599; frondosum, 503; furcatum, 484, 604; Gard- neri, 480; Goringianum, 489 ; grammitoides, 499; Grevilleanum, 603; Grevillei, 476; Griffithianum, 476, var. australis, 476; Grif- fithii, 502; gymnogrammoides, (Bedd.) 490, (Klotzsch) 490 ; Hal- leri, 484; Hancei, 479; heterocar- pum, 480, 481, 599; heterophle- bium, 503; hirsutum, 484, 599; Hookerianum, 497, 504, 605, 606 ; humile, 483; insigne, 483; in- tegerrimum, 504; japonicum, 498, 502, 604, var. chattagramica, 499, var.? Schkuhrii, 499; javanicum, 505; Jerdoni, 502; laciniatum, 481, 482, 599, 610, var. depauperata, 482, var. planicaule, 482; letum, . 481, 599; lanceolatum, 485; lan- : 625 ceum, 496, 599; laserpitiifolium, 484; lasiopteris, 499; latifolium, 498, 500, 502, 599, 600, var. decurrens, 503, var. dilatatum, 503, var. frondosum,503, var. japonicum, 503 ; Lobbianum, 497; lobulosum, 478, 599; longifolium, 427, 478, 479, 599; longissimum, 478, 599; macrocarpum, 486, 488, 602, 603, var. Atkinsoni, 489, var. pinnata, 489, var. squarrosum, 493; macro- phyllum, 480, 504; marginatum, 500, 604; maximum, (Don) 503, (Hk. & Bk.) 502; minus, 477; monanthemum, 477; Moritzii, 503; multicaudatum, 496, 502, 600, var. tristis, 502; multicaule, 477, 599 ; multijugum, 477, 599; multisec- tum, 480 ; musefolium, 475; my- surense, (Roth.) 483, ( Wall.) 484, 599 ; Nidus, 475, 599, var. Simonsi- ana, 475 ; nigripes, 490, 491, 494, 599, var. dissecta, 491, var. sele- nopteris, 491; niponicum, 493; nitens, 600 ; nitidum, 483, 599, 600; normale, 477, 599, var. minor, 477, var. opacum, 478 ; ob- liquum, 605 ; odontophyllum, 604 ; opacum, 477; ovatum, 599; oxy- phyllum, 489, 491, 493, 600, 602, 603, var. kulhaitense, 494, var. oxyphyllum, 494; pachyphyllum, 476; parallelum, 500, 600; par- vulum, 485, 604; pavoninum, 478; pectinatum, 491, 600; pekinense, 483; pellucidum, 600, 606 ; penan- gium, 599 ; phyllitidis, 475 ; pinna- tifido-pinnatum, 497; planicaule, 479, 482, 599 ; polymorphum, 500, 501, 502, 503, 600; polyodon, 479; polypodioides, 501, 503, var. effusior, 501, var. sublatifolia, 501, ar. typica, 501, var. vestita, 501 ; | polyrrhizon, 499 ; porphyrocaulon, 481; porrectum, 498, 599, 600, 604; Prescottianum, 600; proce- rum, 427, 495, 501, 604, var. mishmica, 496; proliferum, 599, 600; prolongatum, 486; pube- rulum, 604; pubescens, 503; pul- chellum, 483, 599; pulchrum, 604; pumilum, 482, var. hymenophyl- loides, 482; pusillum, 477; radia- tum, 505, 597; resectum, 481; 626 reticulatum, 505,599; Ruta, 485; rutefolium, 486; Ruta-muraria, 482; Schkuhrii, (Hk.) 498, 503, (Mett.) 499, (Thw.) 503; septen- trionale, 478; sepulehrale, 483; ser- reforme, 481; sikkimense, 427, 500, 501; simplex, 475; soboli- ferum, 498, 599; sorzogonense, (Hk.) 500, (Prest) 500, 600 ; spec- tabile, 490, 502, 600; spinulosum, 486, var. subtriangularis, 487; splendens, 483; squarrosum, 489 ; stans, 486; stenophyllum, 476; Stoliczke, 500, 604; stramoneum, 494 ; strigillosum, 490, 491; sub- sinuatum, 496, 599; succulentum, 427, 502; sylvaticum, (Pres?) 497, 999, (Bl) 500, 501; Tavoyanum, 480, 603; tenerrimum, 482; tenui- folium, 485, 599 ; tenuifrons, 490, 491, 599; thelypteroides, 488; Thwaitesii, 499 ; tingens, 476; to- mentosum, ( Hk.) 498, 599, ( Mett.) 499; torrentium, 427, 500; tra- peziforme, 481, 604 ; Trichomanes, 477, 599, var. anceps, 477; um- brosum, 496, 501, var. proeerum, 495; unilaterale, 481, 599, var. udum, 481 ; urophyllum, 480, 599 ; varians, 483, 485, 486, 600; viride, . 477; Wichuræ, 479; Wightianum, 604. Asteranthos, 1, 17 ; analysis of, 21 ; brasiliensis, 18, Astrothelium ochrocleistum, 278 ; prostratum, 278; pyrenastroides, 278. Athyrium allantodioides, 488 ; aspi- dioides, 495 ; Atkinsoni, 487 ; aus- trale, 496, 502; brevisorum, 494 ; Clarkei, 490; faleatum, 487 $ fimbriatum, 494 ; Griffithii, 502 ; gymnogrammoides, 490, var. ery- . throrachis, 491;. Hookerianum, 486; macrocarpum, (Bedd.) 489, (Fée) 485; multicaudatum, 502; nigripes, 490; oxyphyllum, 493, 531 ; rubricaule, 492 ; stramineum, 493; thelypteroides, 488. Auricularini, 126 ; of Brisbane, 403. Australian fungi, 399. Auxemma, gen. char. of, 23; Gard- neriana, 24, Auxemme:e, description of, 23. _ INDEX. Avena elatior, fruit of, 163 ; sterilis, torsion of awn, 157, 162. Awn, on the bending of, at the knees, 159; hygroscopie action of, in Stipa, 150, in Avena, 162. Azadirachta, 234. Bactridium flavum, 406. iettoa ocellata, 279. Baker, J. G., Report on the Liliacex, Iridacee, Hypoxidacee, and Hæ- modoracez of Welwitsch’s Angolan Herbarium, 245. Ballia Brunonia, struct. of, 216 ; ealli- tricha, struct. and mode of growth, 211, cells of stems of, 215, 222, joints with stoppers, 212 ; Hombro- niana, 226; Mariana, 227; Rober- tiana, 227 ; scoparia, struet.of, 228. Barringtonia, 55; acuminata, 102; acutangula, 79, (Benth.) 86, (BL) 75, (Gaert.) 80,( Korth.) 87, (Roab.) 82, (Span.) 93, (Thw.) 81, (W. &. A.) 83 ; alata, 70, 72, 93:; alba, 65 ; caffra, 78; ceylanica, 77; conoi- dea, 65, 71, 72; costata, 88; cy- lindrostachys, 72 ; cylindrostachya (Griffith), 100; edulis, 76; elon- gata, 69; excelsa, (Benth.) 74, (BLY 74, 112, (Gray) 75; Hors- fieldia, 85; insignis, 75; inter- media, 73 ; macrocarpa, 109 ; ma- crophylla, 61; Montrouzieri, 89; neocaledoniea, 103; nitida, 88; obtusangula, 81; racemosa, (Bi.) 67, 69, (Gaud.), 75, (Griffith) 71, (Oliv.) 78, (Seem.) 73, (Thw.) 77, var. elongata ( Bl.) 109 ; Reinward- tii, 88 ; reticulata, 87; rosea, 102; rubra, 69; samoénsis, 74, 75; sarcostachys, 102; serrata, 87; speciosa, (Forst.) 56, ( Bl.) 55, 58, (Gaert.) 68, (Linn. fil.) 61, (W. & A.) 63; spicata, 85; sumatrana, 103; timoriensis, (BL) 109, var. alata (Miq.) 93; Vriesii, 106. Barringtoniaees, ealyx of, 47; fruit of, 48; embryo of, 48; geograph. distrib. of, 54 ; inflorescence of, 47 ; ovary of, 48; petals of, 47 ; posi- Belvisia (and position of), 1, 17. . Bennett, A. W., Prelim. Note no | tion of, 53; stamens of, 48 ; style | of, 48; synop. char. genera of, 54, | Growth of Female Flower.stalk of Vallisneria spiralis, 133. Berkeley, Rev. M. J., and C. E. Broome, on Fungi of Brisbane, Queensland, 399. Blechnidium melanopus, 475.. Blechnopsis Cumingiana, 474; elon- gata, 474; latifolia, 474 ; nitida, 474; pyrophila, 474; salicifolia, 474 ; stenophylla, 474. Blechnum brasiliense, 474 ; cartila- gineum, 473; elongatum, 474; Finlaysonianum, 474, 604; fla- bellatum, 505 ; glandulosum, 596 ; imbricatum, 474; melanopus, 474; moluceanum, 474; nitidum (Presl), 474, var. contractum ( Hook.) 474; orientale, 474, 596; pteridioides, 474 ; pyrophyllum, 474 ; radiatum, 505; serrulatum, 474; striatum, 474, Boraginex, peculiarities. inflorese. of, 615 ; self-fertilization sp. of, 375. Botrychium daucifolium,( Wall.) 587, (Hook.) 596; lanuginosum, 588, 596; Lunaria, 587; subbifolia- tum, 587 ; subcarnosum, 587, 588, 596 ; virginianum, 588, 596 ; zey- lanicum, 587. Botryopteris mexicana, 587. Botryoropsis luzonensis, 84. Bourreria clariuscula, 26 ; succulenta, 26. Bovisca lilacina, 406. | Bowringia insignis, 571. Bracts as adapted to self-fertilization, 380; uplifting of, 616. Brainea insignis, 571. Brisbane, Queensland, fungi of, 399. British Lichens, new, 237. Broome, C. E., and Rev. M. J. Berke- ley, on Fungi of Brisbane, Queens- land, 399. Bulbine asphodeloides, 261, _ Burdock, bur of, 165. | Butonica alata, 70; alba, 65; apicu- lata, 78; caffra, 78; calyptrata, 76 ; ceylanica, 77 ; edulis, 76; inelyta, 71; intermedia, 73; procera, 74; racemosa, (Juss.) 66, (BL) 67; rosata, 72; rubra, 70; Rumphiana, © L. 67, 68; samoénsis, 75; splendida, - Pe 60 ; terrestris, 69 ; terrestris rubra, Butonica, embryo of, 50, 52, 53 ; hist. and charac. of, 65. Butonicoides crenata, 90, 91. Byblis gigantea, striated cells of, 166. Cabralea, 236. Ceeomacei (of Brisbane), 407. Calicium kylemoriense, 242. Callipteris esculenta, 503. Calocera viscosa, 127. Calycanthemy of Primulaceæ, 286. Calyx as adapted to self: fertilization, 380. Camellia japonica, blight on leaves of, 301; minute structure of, 303. Campanulaces,' self-fertilization sp. of, 368. Campteria anamallayensis, 469 ; bi- aurita, 469; Rottleriana, 469; Wallichiana, 470. Campylandra dachystachys ( Welw.), 256. Carapa procera, 235. Cardiochlena alata, 539. Careya arborea, 97, var. australis, 91 ; crenata, 90, 91; herbacea, 96, 97; macrostachya, 85, 104; orbiculata, 98; pendula, 96, 995 spherica, 66, 97. Careya, charac. of, 96 ; history of, 95 ; seed of, 48. Caryocar, embryo of, 51. Caryophyllaceæ, self-fertilization sp. of, 353, 392. Cedrela Glaziovi, 236; tribution of, 233. Cenangium lichenoideum, 404. Cephalomanes aurieulatum, 441; ja- vanicum, 442 ; rhomboideum, 442; Zollingeri, 442. Ceratophyllez, self-fertilization sp. of, 393. Ceratopteris thalictroides, 471, 597. Ceterach aureum, 504; officinarum, 504; pedunculata, 570. ` Cheetocladium, fructification of, 420. Chasalia coffeoides, 176 ; divaricata, 176; Fontanesi, 175. — Cheilanthes albomarginata, 427, 456, 457 ; argentea, 458, var. sulphurea, . 458 ; bullosa, 457; chrysophylla, 458 : contigua, 459, 597 ; Dal- housiæ, 457; dealbata, 457, 596; sinensis, dis- INDEX. farinosa, 456, 457, 604, var. chrysophylla, 458, var.. con- color, 457, var. Dalhousie, 456, 457, var. vestita, 457; fimbriata, 455; fragrans, 454; lucida, 459, 596; micrantha, 455, 596; mysurensis, 596; nitidula, 460; odora, 454 ; pallens, 596 ; pallida, 546; rigidula, 457, 604; rufa, 456, 457; rupestris, 455, 596; setigera, 528; Sieberi, 455; sua- veolens (Swtz.), 454; subvillosa, 456; Szovitzii, 454, 455; tenui- folia, 455, 596; tomentosa, 457, 597 ; varians, 455, 597. Cheiranthus Cheiri, sstivation of, 190. Chenopodiacez, self-fertilization sp. of, 377, 393. Chetobromus Dregeanus, awn of, 164. Chiodecton inconspicuum, 280 ; con- chyliatum, 280; moniliatum, 280 ; sinuosum, 280. Chisocheton paucijugum, 236. Chlorophytum andongense, 260 ; bracteosum, 260; debile, 260; filipendulum, 260; lancifolium, 260 ; leucolepis (Welw.), 259 ; ma- crophyllum, 259; - stenopetalum, 260. : Chnoophora glauca, 432. Choanephora, as illustrating conidial fructification in the by Dr. Cunningham, 409; Bre- feld's researches, 419 ; spores of, 428; conclusions from phenomena of develop., 419 ; coni- dia, devel. of, 414; exper. artificial cultivation of, 413; formation of zygospore of, 412; germinal tubes of, 414; history of, 409; minute structure of, 410; reproductive bodies differ, 418; Van Tieghem and Le Monnier's researches, 420 ; variations from nourishment of,416. Chrysodium aureum, 582; cayen- nense, 582 ; D’Urvillei, 582 ; fraxi- nifolium, 582; sculpturatum, 582. Chydenanthus, charac. of, 111; ex- celsus, 112, hist. of, 112. Cibotium assamicum, 435 ; Barometz, 435 ; glaucescens, 435. Circæa, 178. Circinella umbellata, 406. |. SECOND SERIES.— BOTANY, VOL. I. Mucorini, ` chlamydo- . 627 Cladosporium herbarum, 407. Clarke, C. B., Review of the Ferns of Northern India, 425. Clavaria argillacea, 403 ; 403 ; stricta, 403. Clavariei, 127 ; of Brisbane, 403. Clusiace:, embryo in, 50. Coenopteris appendiculata, 485 ; fur- cata, 600 ; vivipara, 600. Coffea, African species of, by W. P. Hiern, 169. Coffea Afzeli, 169, 174; arabica, (L. 169, 170, (Benth. 171, 172, (6, Willd.) 173, var. leuco- carpa, 171; bengalensis, 169 ; bre- vipes, 169 ; capitata, 176; cymosa, 175; divaricata, 176; hirsutus, 176; hypoglauca, 169, 173; jas- minoides, 169, 175 ; key to classi- fication, 170; Kraussiana, 176; laurifolia, 170; laurina, 175; li- berica, 169, 171; macrocarpa, 169, 173; mauritiana, 169, 173; me- lanocarpa, 169, 173; microcarpa, 164,174 ; mozambicana,175 ; Per- rotettii? (Steud.), 174; racemosa, ` 169, (Lour.) 175 ; ramosa (Roem. et Schult.), 175 ; rupestris, 169, 174 ; stenophylla, 169, 172 ; subcordata, 169, 174; sylvestris, 173; tenui- flora, 169 ; tetrandra, 169; zan- guebarica, 169, 172. Coleochete scutata, 311. Commersona, 68. Composite, plumes on seed of, 165; self-fertilization sp. of, 367, 392. Conidial fructification in the Muco- rini, by Dr. Cunningham, 409. Contabescence, 381. Contribution to the Lichenographia of New Zealand, by Charles Knight, 275. Convolvulaces, self-fertilization sp. of, 368, 392. Corallina monile, 204; officinalis, 203, histology, frond of, 204; opuntia, 202, Decaisne's obs. on, 198; tridens, 200, 202; tuna, 200, 202; vermicula, 200. Corallinacew, brilliant colours of, 200; colouring-matter of, 200; Duncan and Nelson on, 197, 200; of Ber- muda, struct. of, 200. Corolla as adapted to self-fertiliza- 4 Q rugosa, 628 tion, 380; irregular plants show- ing, 184; regular plants showing, 184. Corolla-lobes, develop. after stamens in Arenaria trinervis, 196; in Ceras- tium glomeratum, 196; in Lychnis dioica, 196; in Ranunculus acris, 196 ; in Stellaria Holostea, 196; in Veronica Chamedrys, 196. Cortieium leve, 403; levigatum, 127; Mougeotii, 127; violaceo- lividum, 127. Coussarea (Audl.), 169. Crassulacew, modification cymes of, 617. Craterispermum laurinum, 175. Cremaspora africana, 176. Cruciferæ, binary symmetry of, 193; marginal placentas of, 192; self. fertilization sp. of, 350, 391. Cruciferous flower, structure of, Rev. G. Henslow on, 177. ES Cryptogramme acrostichoides, 459; Brunoniana, 459, 603; crispa, 459, 460, 603, var. acrostichoides, 460. Cunningham, Dr. D. Douglas, on the occurrence of Conidial fructification in the Mucorini, illus. by Choane- phora, 409; on Mycoidea parasi- tica, and the part it plays in for- mation of certain Lichens, 301. Curculigo gallabatensis, 266. Currey, Fred., on a collection of Fungi made by Mr. Sulpiz Kurz, 119. Cyathea barbata, 435; bipinnata, 605 ; Brunonis, 599 ; excelsa, 599 ; robusta, 599; spinulosa, ( Wall.) 429, 430, 431, 599. ( Bedd.) 429, (J. Scott) 430; venulosa, 433, 599. Cyathus campanulatus, 405 ; fime- tarius, 405 ; intermedius, 128. Cyclamen, double petals, 287. Cyme, dichotomous, 618 ; scorpioid, origin of, by Rev. G. Henslow, 613, 618 ; spicate, 618. Cympolia barbata, 198. Cyrtomium (subgen.), 512 ; caducum, 512; caryotideum, 512 ; falcatum, 512. Cystopteris dimidiata, 444 ; fragilis (Bernh.) 450, f. Huteri (Milde), INDEX. 450; nodosa, 444; retusa, 450, 492; setosa, 450; spinulosa, 486 ; squamata, 444. Deedalea discolor, 125 ; Sprucei, 402 ; tenuis, 125 ; unicolor, 403 ; zonata, 125. Danthonia, awn of, 164. Darwin, Francis, on the Hygroscopic Mechanism by which certain Seeds are enabled to bury themselves in the ground, 149. Dasystachys, 255 ; campanulata, 256 ; colubrina, 256; falcata, 256 ; pleio- stachya, 255. Davallia achilleifolia, 600 ; affinis, 444 ; alpina, 442 ; angustata, 600 ; assamica, 443 ; Boryana, 451 ; bul- lata, 445; caudata, 605; chero- phylla, 444; chinensis, 449, 600; Clarkei (Hk. 4 Bk.), 443; conii- folia, 600 ; dareseformis, 427, 443 ; dimidiata, 445; divaricata, 445; elegans, 600; flaccida, 447, 448, 605, var. pilosula, 449, var. pubera, 449, var. pyramidata, 449; flagellifera, 600; Griffith- iana, 445; hemiptera, 451; hirta, 447 ; Hookeriana, 446, 447, 605; immersa, 443; jamai- censis, 448 ; khasiyana, 447; Kurzii, 446; ligulata, 444; lobu- losa, 600; lonchitidea, 446, 600; marginalis, 446, 600, 605, var. cal- vescens, 447 ; membranulosa, 442, 449; micans, 443; moluccana, 605 ; multidentata, 442, 443, 602; nodosa, 434, 443, 444, 602; ornata, 600 ; parallela, 600 ; parvula, 600 ; pedata, 442, 600 ; pilosa, 448 ; pi- losula, 447 ; platyphylla, 446, 600 ; polyantha, 445; polypodioides, 447, (Don) var., 605, var. hirta, 448, var. pilosula, 448, var. rhomboidea, 448, var. strigosa, 448; proxima, 447 ; puberula, 448 ; pulchra, 444, 445, var. pseudo-cystopteris, 444, 450; pyramidata, 448, 449; repens, 427, 451; rhomboidea, 447; Roxburghii, 447, 605; serrzfor- mis, 600; setosa, 450; scabra, 446, 447; schizophylla, 451; Spelunce, 448 ; sphseropteroides, 445 ; stipellata, 444 ; strigosa, (Swartz) 436, 447, (Hk. & Bk.) 447; subimbricata, 442; tenui- folia, 449, 600; trichomanoides, 451; trichosticha, 448 ; urophylla, 446, 447, 605; villosa, 446, 600. DeCandolle, Casimir, on the geogra- phical distribution of the Meliaces, 233. Dematiei (of Brisbane), 407. Dennstzedtia deltoidea, 436. Diacalpa aspidioides, 434, 445, 603 ; foeniculacea, 427, 434 ; Hookeriana, 494 ; sphzropteroides, 455. Dialysis, rare cases of, 193. Diatrype rugosa, 130. Dichasium of Schimper, 618, 619. Dicksonia appendiculata, 436, 596; — arborescens, 596; assamica, 435; Barometz, 435; deltoidea, 436; Elwesii, 496 ; Griffithiana, 435 ; moluccana, 604; polypodioides, 447; scabra, 436, 604. Dictyocline Griffithii, 576. Dictyopteris (subgen.), 548. Didymium farinaceum, 406. Didymoglossum euphlebium, 440; Filicula, 440; ]late-alatum, 440; . plieatum, 440, racemulosum, 440. Digrammaria ambigua, 503. Dipeadi comosum, 247; lateritium, 247 ; oxylobum, 246; Welwitschii, 246. ; Diplazium (subgen.), 496 ; alternifo- lium, 497 ; bantamense, 497 ; bul- biferum, 498, 499 ; dilatatum, 502 ; esculentum, 503; faleatum, 497, 569 ; fraxineum, 569; fraxinifo- lium, 497; Griffithii, 502; Jer- doni, 502; lanceum, 496; lasio- pteris, 499; lobulosum, 478 ; lon- gifolium, 478 ; malabarieum, 503; malaccense, 499 ; pinnatifido-pin- natum, 497; polypodioides, 501; Schkuhrii, 499, 503 ; sorzogonense, 500; speciosum, 499; Stoliczkee, 500; sylvaticum, 498; thelypte- — roides, 488 ; Thwaitesii, 499 ; to- mentosum, 498. ; Dipteris (subgen.), 555; Wallichii, 555. Doryopteris sagittifolia, 471. Double primroses, 287, 293. z Doxomma (n. gen. Miers), 98, 99; acuminatum, 102; angustatum, 105; cochinchinense, 101; cylin- drostachya, 100; macrostachyum, 104; magnificum, 105; neo- caledonica, 103; pendulum, 99; rigidum, 104; sarcostachys, 102, 103; sumatrana, 103; Vriesii, 106. Dracena acaulis, 252 ; fragrans, 252 ; interrupta, 252 ; ^ monostachya, 252 ; nitens, 252; parviflora, 252. Drymoglossum Beddomei, 576, 598 ; carnosum, 575, 598 ; piloselloides, 575, 598; subcordatum, 575. Drynaria (subgen.), 555; coronans, 557, 582; mollis, 556; propin- qua, 556; quercifolia, 556, 582. Dysoxylum spectabile, distrib. of, 233. Elaphoglossum conforme, 576 ; lauri- folium, 576; viscosum, 577. Ella-medella-gass—Stravadium acu- tangulum, 80. Embryo in Clusiaceæ, 50. Equisetum arvense, 594; bullata, 600; chærophylla, 600; debile, 594, 603; diffusum, 594, 600; elongatum, 595, 600; flaccida, 600, and var., 600; immersa, 600 ; laxum, 595 ; membranulosa, 600; nodosa, 600; pallens, 595, 603; palustre, 594; pilosa, 600; polypodioides (Don), var., 600; puberula, 600; pulchra, 600; pyramidata, 600; ramosis- simum, 595, ramosum, 595 ; rhom- boidea,600 ; robustum, 595; scir- pioides, 600 ; scoparium, 594, 603 ; stipellata, 600; virens, 600 ; vir- gatum ( BI.), 595. Eremobryoid series of Polypodium, 549. Erineum, tubes of, 166. à Eriospermum andongense, 261; flex- uosum, 261 ; ophioglossoides, 262 ; paludosum, 261; stenophyllum, 261. Erodium, manner in which seed of, is buried, 155. Eugenia acutangula, ( Linn.)70,79,80, (Lour.)101 ; erenata (Soland.), 91; racemosa, (Linn.) 66, (Seem.) 73, — "(Forst.) 73, (Roxb.), 82. INDEX. Eunephrodium (subgen.), 529, 547. Eupapaverz, 180. Euphorbiacese, self-fertilization sp. of, 978, 393. Eupolypodium (subgen.), 549. Feretia apodanthera, 169. Ferns, difficulties with regard to no- menclature, 426. Ferns of N. India, review of, 425. , change of names, 427; Col. Beddome’s tab. of distrib. of species, 426; new species, 427; Wallich’s collection, how to be quoted, 427, tabular reduction of, 595. Fissurina nove-Zelandie, 281. Flindersia, 235 ; Fournieri, 236. Floral estivations, diagrams of, 177 ; introductory remarks on, 177; origin of, Rev. G. Henslow on, HU Forms of sstivation, 178; causes which determine, 187, in irregular flowers, 188, in regular flowers, 187 ; cochlear, cochleate (footnote), 182; convolute and contorted, 179, 185; degrees of frequency, 179; distichous equitant, 178, 180; equitant, 180 ; half-equitant, 178, 180 ; half-imbricate, 178, 182; im- bricate proper, 178, 184 ; imbrica- tive,179; induplicative,179 ; nume- rical order of parts of whorls, 186 ; pentastichous, 178, 182 ; quincun- cial, 178, 182; reduplicative, 179 ; reversal of the spiral, 186; straight or open, 179; tristichous, 178, 181; valvate, 179, 185 ; valvative, 180; varieties in same species of diff. genera, 189. Fuchsia coccinea, sestivation of, 190. Fumago salicina, 130. Fungi, on a collection of, made by Mr. Sulpiz Kurz, 119. Galanthus nivalis, sstivation of, 190. Galium Aparine, bur of, 165. Geaster saccatus, 406; striatus, 406. Gentianaces, self-fertilization sp. of, 368. Geographical distribution of the Me- liacez, 233 ; map of (Pl. 31), 236 ; summary of, 236 ; tabular arrang. of (Pl. 30), 236. 629 Geraniaceæ, burying powers of, 165 ; self-fertilization sp. of, 357, 392. Gethyllis pilosa, 267. Geum, bur of, 165. Gladiolus andongensis, 269; ango- lensis, 269; benguellensis, 268; brevicaulis, 267 ; cærulescens, 267; gregarius, 268; laxiflorus, 268; luridus, 267; multiflorus, 269; quartinianus, 269; splendens ( Welw.), 268 ; Welwitschii, 268. Glaux, corolla wanting of, 287. Gleichenia arachnoides, 428; at- tenuata, 598; Bancroftii, 428; crassifolia, 428 ; dichotoma, 428 ; excelsa, 428; ferruginea, 428; gigantea, 428, 598; glauca, 428, 598; Hermanni, 428, 598; ja- ponica, 428; Klotzschii, 428; la- nigera, 428; linearis, 428, 598; longissima, 428 ; pteridifolia, 428. Gloriosa superba, 262 ; virescens,262. Goniophlebium (subgen.), 550 ; amce- num, 550; argutum, 551; ery- throcarpum, 565; Hendersoni, 550; lachnopus, 551; mierorhi- zoma, 551; serratifolium, 552; subaurieulatum, 552. Goniopteris (subgen.), 547; lineata, 547; multilineata, 548; penan- giana, 548 ; prolifera, 548. Gramineæ, self-fertilization sp. of, 979, 393. Grammitis acuminata, 570, 595; affinis, 569, 595; arborescens, 568 ; aurita, 568; caudata, 569, 595; Ceterach, 504; decurrens, 571, 595; diversifolia, 595; sco- lopendrina, 570; cuspidata, 570 ; Finlaysonia, 571, 603 ; flavescens, 570, 595; Hamiltoniana, 570, 595; involuta, 570; lanceolata, 570; macrophylla, 570, 595; mi- crophylla, 569 ; procera, 569, 595 ; Totta, 567, 568 ; vestita, 568, 595. Grandinia granulata, 126. Graphiola pheenicis, 130. Growth, curves of, A. W. Bennett on, 136; diagrams of, 137, 141; in Ballia callitricha, 211; in flower- stalk of Hyacinth, 139 ; in fl.-stalk of Vallisneria, 133. Guarea filiformis, 236 ; trichilioides, 236. 630 Guepinia ramosa, 127; spathularia, 403. Gustavia alata, 93; valida, 93, 94. Gymnogramme Andersoni, 568; as- pidioides, 567, 569; aurita, 544, 568, var. Levingii, 568; decur- rens, 570; elliptiea, 570, 595, 603; fraxinea, 569, 595, var. pilosa, 569; Hamiltoniana, 570, 595 ; involuta, 570, 595 ; javanica, 569; lanceolata, 570; Lovei, 567; Marante, 567; microphylla, 569; obtusata, 568; opaca, 568, 600; serrulata, 569; stegnogram- me, 569 ; Totta, 567 ; vestita, 568, 595. Gymnopteris axillaris, 579; decur- rens, 579; Féei, 579; lanceolata, 579; minor, 579; normalis, 579 ; spicata, 582; tricuspis, 581; va- riabilis, 579. Gymnosphiera gigantea, 496 ; glabra, 433. globosa, 86; Hæmodoraceæ, Angolan, 245. Haloragiacese, self-fertilization sp. of, 392. Harmena, 53. Haworthia angolensis, 263. Heliotrope, scorpioid racemes of, 616. Helminthosporium Ravellinii, 407. Helminthostachys dulcis, 595; zey- lanica, 587, 595. Helotium citrinum, 128. Helvella californica, 423. Helvellacei, 128; of Brisbane, 404. Hemidictyon Finlaysonianum, 504. Hemidictyum (subgen.), 504; Bru- nonis, 505 ; Ceterach, 504. Hemionitis arifolia, 576, 595, 604 ; Boryana, 573; cordata, 576; cor- difolia, 576, 595; coriacea, 572; esculenta, 503; Griffithii, 576 ; hastata, 575, 604 ; opaca, 568; po- thifolia, 571; reticulata, 573; sa- gittata, 575 ; Wilfordii, 575. Hemitelia Beddomei, 429; Bruno- niana, 427, 430, 431, var. ? Scottii, 431; decipiens, 430; latebrosa, 432, : Taer Rev. George, on Origin of systems of dee onem r INDEX. 37; on Origin of so-called Scor- pioid Cyme, 613; on the Origin of Floral Æstivations, 177; on the Self-fertilization of Plants, 317. Hermannia, whorls of, 177. Heteroneuron diversifolium, 580; heteroclitum, 580; proliferum, 581. Heteropogon contortus, awn of, 164 ; melanocarpus, awn of, 164. Hexagonia Kurzi, 126 ; polygramma, 125, 403; tenuis, 125. Heynea, 264. Hiern, W. P., on the African species of the genus Coffea, Linn., 169. Hinneola auricula Canis, 128; au- ricula Jude, 128, 404; polytricha, 404, Histology of Corallinee, 197; De- caisne on, 197 ; Quekett on, 197 ; Rosanoff on, 199. Holeus, awn of, 164. Humata pedata, 442. Hyacinth, on growth of flower-stalk of, 139, 143. Hydnei, 126; of Brisbane, 403. Hydnum auriculatum, 126; gelati- nosum, 126; udum, 126. Hydrocharis, whorl flower of, 15 Hydroglossum auriculatum, 584 ; cir- cinatum, 583; dichotomum, 583 ; flexuosum, 584; japonicum, 585; longifolium, 583; pedatum, 588; pinnatifidum, 584. Hygrophorus miniatus, 399. Hygroscope, how made, 155. Hygroseopie mechanism by which certain seeds are enabled to bury themselves in the ground, 149; hygroscopie torsion, 166. Hymenesthes, hist. and charac. of, 26; nitida, 26. Hymenochsete Cacao, 403; tenuissima, 403. Hymenolepis ophioglossoides, 582; revoluta, 582; spicata, 582. Hymenophyllum abietinum, 437 ; alatum, 440; badium, 437, 439, 599; Blumeanum, 437; Borya- num, 598; ciliatum, 437, 439; crispatum, 437, 598; dædaleum, 437; densum, 436, 440, 598; denticulatum, 438, 439, var. flac- - cidum, 438, 439; erosum, 437; hj exsertum, 436, 437, 598 ; Filicula, 440 ; fimbriatum, 437; flaccidum, 439; flexuosum, 437; himalai- num, 437; integrum, 437; java- nicum, 437, 439, 598, 599, var. badium, 438; khasianum, 438; leve, 604; Levingii, 427, 439; macroglossum, 436; micranthum, 437; microsorum, 437; osmun- dioides, 437; polyanthos, 437, 598, 599, var. fj. minor, 437 ; protru- sum, 437; pycnocarpum, 437; Reinwardti, 437; serpens, 437, 599; Simonsianum, 438 ; Smithii, 439; sphsrocarpum, 437; Tel- fairianum, 598 ; tenue, 605, Hyoscyamus, floral axis of, 615. Hypedomatium onustum, 525; Rup- pellianum, 525. Hypochsete debilis, 595. Hypoerea variabilis, 130. Hypolepis hostilis, 546; pteridioides, 468 ; tenuifolia, 546. Hypoxidacee, Report on, of Wel- witsch's Angolan herbarium, 245, 265. Hypoxis angolensis, 266; angusti- folia, 265; canaliculata, 265; cuanzensis, 265 ; monanthos, 266 ; polystachya, 266, var. andongensis, 266. Hypoxylon angolense, 405; cetra- rioides, 405; concentricum, 130, 405; cretaceum, 405; margina- tum, 130; suborbiculare, 130. Tleodictyon gracile, 405. Incomplete, self-fertilization sp. of, 377. India proper, Kurz’s divisions of, 425. India, Review of the Ferns of North- ern, 425. Inflorescenee, steps towards scorpioid cyme, 617. Tridacese (Angolan), 267; Report on, of Welwitsch's herbarium, 245. Iridopsis textilis, 270. Irish lichens, new, 241. Irpex flavus, 126, 403; palleseens, — 126; 403. sinuosus, 126; zonatus, Jambos sylvestris Saamstravadi, 66. Juncacee, 379, 393. self-fertilization sp. of, _ Kaulfussia æsculifolia, 585; assamica, 585; Korthalsii, 585; Lobbiana, 585. Knight, Chas., Contribution to the : Lichenographia of New Zealand, 275. Kniphofia andogensis, 246; benguel- lensis, 246. Kraussia floribunda, 176. Kurz, Mr. Sulpiz, on a collection of Fungi made by, 119. Labiatze, self-fertilization sp. of, 374, 909. ^. ; ; Lacaussadea appendiculata, 578 ; montana, 578; rhizophylla, 578. Lachnocladium furcellatum, 403. Lagerströmia, oscillating arrang. leaves of, (ftnote) 45, 613. Lagurus ovatus, awn of, 164. Lansium, 235. Lapeyrousia abyssinica, 272; cyanes- cens, 272; fragrans, 272; littoralis, 272 ; odoratissima, 273. Laschia Thwaitesii, 403. Lastrea (subgen.), 512; angustifrons, 527; apiciflora, 518; aristata, 511; Atkinsoni, 526; barbigera, 592; bella, 525; Bergiana, 515; Blumei, 526 ; Boryana, 527 ; Bru- noniana, 522; cana, 516; coch- leata, 522, 582; coniifolia, 511; crenata, 525; eburnea, 493; elon- gata, 513, 521, 523; Elwesii, 516; eriocarpa, 525; Fairbanki, . 517; falciloba, 515; Falconeri, 522; Filix-Mas, 582, var. Clarkei, 518, var. odontoloma, 521; flac- cida, 514, 528 ; foeniculacea, 434 ; gracilescens, 514 ; gracilis, 508 ; Hendersoni, 526; hirtipes, 513; immersa, 514; intermedia, 520; macrocarpa, 489; nigrescens, 515; octhodes, 516 ; patentissima, 520 ; platypus, 528 ; pulvinulifera, 435, 525 ; recedens, 526; sericea, 515; setigera, 528; setosa, 450; sparsa, 494, 524, var. obtusissima, 524; spectabilis, 519; spinulosa, 523; splendens, 527; Thelypteris, 517; tylodes, 516. INDEX. Lathreea Squamaria, arrang. inflores. of, 613. Leaf, peculiar change of position of, 616. Lecanora jejuna, Nyl., =L. subdiluta, Leight., 146. Lecidea abietina, 278 ; advenula, 146 ; antrophila, 242 ; atro-morio, 276 ; Dilleniana, 278 ; endomelæna, 239 ; formosa, 278; glaucomaria, 238 ; grumosa, 242; imponens, 238; impressula, 237; littoralis, 275; meiospora, 275; nigrescens, 276; parellaria, 238 ; petræa (F'lot.), var. neo-zelandica, 277, var. violacea, 277; polospora, 241; premnea, 278; prosechoides, 146; ryssolea, 237; schistacea, 276; stellulata, 277 ; subargillacea, 275; subbadio- atra, 276 ; subcoarctata, 276 ; sub- diluta, 145; subfarinosa, 277; subglobulata, 275; sublapicida, 276 ; subtubulata, 277 ; tephrizans, 237; iubulata, 277; Whakatipe, 277. Leguminose, self-fertilization sp. of, 360, 392. Leighton, Rev. W. A., new British Lichens, 145, 237; new Irish Lichens, 241. Lentinus esspitosus, 120; caprona- tus, 119; cyathus, 399; descen- dens, 119, 400; Dunalii, 400; exilis, 120, 400; fasciatus, 399 ; furfurosus, 120; glabratus, 120; inquinans, 120; irregularis, 121 ; Kurzianus, 120; Lecomtei, 400 ; Sajor Caju, 120; subdulcis, 400 ; velutinus, 120. Lenzites albida, 121 ; betulina, 400 ; deplanata, 400; Palisoti, 121. Lepiota, 119. Leptochilus axillaris, 579 ; decurrens, 579 ; lanceolatus, 579 ; minor, 579. Leptostegia lucida, 459. Leucostegia immersa, 443. Lichenographia of New Zealand, con- | tributions to the, 275. Lichens, new British, 145, 237; new Irish, 241. Liliacee, Report on, of Welwitsch’s Angolan herbarium, 245. self-fertilization sp. of, ——, , 357. 631 Lindsaya attenuata, 450, 451, 598, 604; chinensis, 449; cultrata, 450, 451, 452, 598; decomposita, 593; dentata, 598; ensifolia, 452, 604; Finlaysonia, 452, 604; fla- bellulata, 451, 598 ; gracilis, 450 ; Griffithiana, 452; heterophylla, 452; interrupta, 451, 604; lan- eeolata, 452; lanuginosa, 541, 598 ; lobata, 598 ; Lobbiana, 450; longipinna, 452, 634; lucida, 450, 598; neriiforme, 542; ob- longifolia, 451; odorata, 450; pallens, 450, 598 ; parasitica, 604 ; pectinata, 451; pentaphylla, 452; polymorpha, 451, 598; pteroides, 452, 604 ; recurvata, 598; repens, 427, 451; scandens, 451; serpens, 598; striata, 451; tenera, 451; tenuifolia, 449. List of Fungi from Brisbane, Queens- land, with descrip. of new Species, by the Rev. M. J. Berkeley and C. E. Broome, 399. Litobrochia aurita, 471 ; ludens, 470 ; pedata, 470. Lomaria adnata, 472; articulata, 473; attenuata, 596; aurea, 458 , 596 ; Boryana, 599 ; callosa, 472; caruifolia, 458,596 ; euphlebia,472, 473, 596; glauca, 472; limoni- folia, 577, 596; Patersoni, 472; pyenophylla, 472, var. 472, 474; scandens, 577, 596 ; secunda, 596; semicordata, 472, 473; serpens, 579, 596; spondisefolia, 596; Wightiana, 604. Lonchitis hirsuta, 604. Loxogramme involuta, 570; lanceo- lata, 570. Lycoperdacei, 128. Lyeoperdon brasiliense, 128; gem- matum, 128; pusillum, 128, 406. Lyeopodium aloifolium, 590, 598; annotinum, 592, 593; argenteum, 598; atroviride, 597; bryopteris, 604; esmspitosum, 594; casuari- noides, 593 ; caulescens, 598 ; cer- nuum, 591, 598, var. sikkimensis, 592; ceylanicum, 590; circinale, 604 ; clavatum, 592, 598; comans, 593; complanatum, 593, 604; concinnum, 597 ; curvatum, ( Dalz.) ÅR 632 594, (Swiz.) 591; 598 ; elegans, 598; empetrifolium, 590; epicesfolium, 591; filicaule, 593 ; flexuosum, 604; fulcratum, 598 ; gnidioides, 598; gramineum, 590; Hamiltonii, 590, 598, var. petiolata, 590; Heyneanum, 592, 598; Hookeri, 591, 597; java- nicum, 591; levigatum, 604; lucidulum, 590; miniatosporum, 594; mirabile, 593; myrtifolia, 593; obtusifolium, 590, 598; or- nithopodioides, 604; pallidum, 606; Phlegmaria, 592, 598, var. nummularifolia, 593; proliferum, 591; protensum, 591; pubescens, 598; pulcherrimum, 590, 597; pulvinatum, 606 ; rotundifolium, 593, 604 ; rubellum, 593 ; sabinæ- folium, 593; semicordatum, 598; serratum, 591, 597; setaceum, 590, 591, 597, var. pulcherrima, 590, var. subulifolia, 590; Squar- - rosum, 591, 597, 606; subdiapha- num, 598 ; subulifolium, 590, 597 ; suleinervium, 591; tetragonosta- chyum, 598; thyoides, 593 ; in- chiatum, 592; ulcifolium, 591; urostachyum,597 ; vernicosum,589, 590; verticillatum, 591, 597, 606 ; vulcanicum, 592 ; Wightianum, 593, 604; Willdenovii, 598. Lygodium circinatum, 583, 599 ; di- , chotomum, 583, 599; Finlaysoni- anum, 585, 604; flexuosum, 584, 585, 599; japonicum, 584, 604; longifolium, 583, 584, 599 ; micro- phyllum, 583, 584, 599; pedatum, 583; pinnatifidum, 584; polysta- chyum, 584, 599 ; pubescens, 584, 604 ; scandens, 584; semibipinna- SS 585 ; semihastatum, 584 ; ser- rulatum, 584; tenue, 585. Lysimachia nummularia, 287. Lythraces, self-fertilization sp. of, 363, 392. Macrochloa arenaria, awn of, 164. Malvaces, self-fertilization sp. of, ... 956, 392. Mammea asiatica, 58, 61. in 119; ramealis, 119. divaricatum, (Hk. & Grev.) 594, (Wall.) 592, INDEX. Masters, M. T., on some points in the Morphology of the Primulacesz, 285. Matonia pectinata, 599. Mechanism of torsion, 157. Medella = Stravadium acutangulum, 80. Megadendron, 109 ; ambiguum, 110 ; macrocarpum, 109 ; pallidum, 110. Melaspilea amphorodes, 281; meta- bola, 281; vermifera, 146. Melia Azedarach, distrib. of, 233, 235. Meliacez, on the geographical distri- bution of, 233. Melobesiz, filamentous processes of, 207; Rosanoff on histology of, 199, 207. Menichea rosata, 72. Meniscium Cumingii, 571; cuspida- tum, 547, 571, 572, 596, var. lon- gifrons, 572, 596; deltigerum,572, 580, 596 ; erosum, 571, 596; lon- gifrons, 596 ; Parishii, 571; proli- ferum, 548; reticulatum, 572; salicifolium, 596; simplex, 571; triphyllum, 571, 596; villosum, 571. Mertensia dichotoma, 428; emargi- nata,428 ; glabra, 428 ; glauca, 428. Mesocarpus nummuloides, (ftnote) structure of, 212. Meteorus coccineus, 83. Mierogonium bimarginatum, 440 ; sublimbatum, 439. Microlepia caudigera, 446; hirta, 447; platyphylla, 446; pteropus, 443; scabra, 447; spelunce, 448 ; strigosa, 436, 447 ; urophylla, ` 446. Mieropeltis applanata, 130. Mierosorium irregulare, 561. Miers; J., on Napoleona, Omphalocar- pum, and Asteranthos, 1; on the Auxemmes, a new Tribe of the Cordiacese, 23; on the Barringto- niaces, 47. Mitraria Commersoni, 68. Mode of growth, of Ballia callitricha, 211; of Corallinaces, 203. Monachosorum davallioides, 546. Monocotyledons, self-fertilization m. of, 378. D Montbretia abyssinica, 272, | Morsa andongensis, 271; candela- brum, 271; glutinosa, 271; gra- cilis, 272 ; Eege 271; textilis, 270; Welwitschii, 270. Morphology of Primulacez, Dr. M. T. Masters on, 285. Mortierella, fructification of, 421. Mucedines (of Brisbane), 406. Mycelia (of Brisbane), 407. Mycenastrum corium, 406. Mycoidea parasitica, Dr. D. Douglas Cunningham on, 301 ; alliances of, 311; descrip. of, 301; disk and filaments, 304, 305, 310; gen. charac., 312 ; life-history of, 301 ; oogonium and zoospores of, 308; relations when attacked by lichens and fungi, 312; sexual fructifica- tion of, 307 ; spores of, 306 ; zoo- spores of, 309. Myosotis, convolute corolla of, 177 ; scorpioid racemes of, 616. Myrtus alata, 93. Myxogastres, 128; of Brisbane, 406. Napoleona, analysis of, 2; calyx of, 9; corona of, 2, 5; fruit of, 6; ovary of, 4; seeds of, 4. angolensis, 6, 12; cuspidata, 10 ; Heudelotii, 4, 8, 21; imperia- lis, 6, 7, var. purpurea, 8; Mann, 11; Vogelii, 6, 9 ; Whitfieldii, 6, 9. Napoleonex, position and history of, 1. Nectria Eugenie, 130. Nephrodium abruptum, 532, 534; affine, 520 ; amboinense, 532 ; an- gustifrons, 527; apiciflorum, 517, 602, var. nidus, 518; appendicu- latum, 516; arbuscula, 532; ari- dum, 531, 601, 602; artinexum, 536; barbigerum, 522 ; Beddomei, 517; Boryanum, 527, 528, 600, 603, var. microstegioides, 528, 544 ; Brunonianum, 522, 602; Bucha- nani, 525; calcarata, 515 ; canum, 515, 543, 602, 603; cicutarium, 539, 540, 603, var. coadunata, 540 ; ciliatum, 514, 602, 603; Clarkei, ` .. 518,520, 536 ; coadunatum, 540 ; cochleatum, 520, 521, 603; cre- ` natum, 524, 602 ; — 533, currens, 539 ; delicatulum, 540 ; didymosorum, 530; dissectum, 526, 527, 534, 535; divisum, 527 ; elatum, 531, var. procerum, 531; elongatum, (Swiz.) 519, (Hk. A Grev.) 520; Elwesii, 516; erio- carpum, 525; eusorum, 534; ex- altatum, 541; extensum, 516, 529, 530, 533, var. lete-repens, 530, var. microsora, 530, var. 9. minor, (Bedd.) 532; Fairbankii, 517; falcilobium, 515, 517, 602, 605, var. pubera, 515; Falconeri, 522; ferox, 534; fibrillosa, 518 ; Filix- Mas, 519, 527, var. (Bedd.), 518, var. cristata, 519, ( Rich.) var. 602, var. fibrillosa, 520, var. interme- dia (Bedd.), 522, var. khasiana, 519, var. marginata, 520, 521, var. normalis, 519, var. panda, 519, var. patentissima, 520, var. Schim- periana, 520, ( Hochst.) 522 ; flac- cidum, 513, 514, 528; fuscipes, 526, 527, 535, 538; giganteum, 539 ; glandulosum, 531, 547, var. lete-strigosa, 532; gracilescens, 513, 517, 528, var. decipiens, 514, var. hirsutipes, 514, var. didymo- chlenoides, 514; 537; heterosorum, 537, 555 ; hir- sutum, 524; hirtipes, 513, ( HF.) var. 603; Hudsonianum, 534; immersum, 514 ; ingens, 526, 527, 585; lacerum, 522; latipinna, 532; Leuzeanum, 535, 536, 605; lineatum, 547; lingua, 553; ma- derense, 521; membranifolium, 534, 602, 603, var. typica, 535, : var. dimorphum, 535 ; meniscioides, . 532; microphyllum, 584 ; micro- stegium, 544; multicaudatum, 427, 540, 603; multijugum, 533, 602; nymphale, 533; octhodes (Hk.), 516; odontoloma, 521, var. (Baker), 518; odoratum, 525; pallidum, 593; parasiticum, 516, 532, 533, 534, 601, 602, 605, var. aurea, 533, var. multijuga, 533 ; Parishii, 525; patentissimum, 520; penni- gerum, 532, var. multilineata, 532 ; platypus, 528 ; polymorphum, 537, 603, var. Simonsii, 537 ; procur- — gens, 530; prolixum, 516, 517, 602; propinquum, 529; pteroides grandifolium, INDEX. (J. Sm.), 529 ; pteridioides (Griff.), 518 ; puberulum, 513, 514; pul- vinuliferum, 525, 526, 602; pur- puraseens, 523; remotum (Hk.), 521, 523, var. Chanterim, 523; repandum, 537; rhodolepis, 427, 526; rigidum, 523; rostratum, 538; sagenioides, 535, 602; sca- brosum, 527, 602 ; setigerum, 528, 546 ; siifolium, 538; sikkimense, 525 ; sparsum, 489, 519, 523, 605, (Don) var., 603, var. gracilis, 508, var. latisquama, 524, var. membra- naceum, 489, var. minor, 524, var. nitidula, 524, var. squamulosa, 524 ; spectabile, (C. B. C.) 526, (Hk.) 518, 608 ; spinulosum, 523 ; splen- dens, 527; subconfluens, 536; syrmaticum, 518; tenericaule, 598, 545, 602, 605; terminans, 529 ; Thelypteris, 514, 517, 544; truncatum, 534, 602; tuberosa, 540; undulatum, 524; unitum, 529, (R. Br.) 601, (Sieber) 530; variolosum, 538, 603; vastum, 536,603; venulosum, 534; Wightii, 427, 538. Nephrolepis acuta, 541 ; cordifolia, 540, 603, 605; exaltata, 541, 602, 603 ; volubilis, 541, 603. Neuronia asplenioides, 542. New British Lichens, by Rev. W. A. Leighton, 145, 237. New Zealand, contribution to the Lichenographia of, 275. Nidulariacei, 128 ; of Brisbane, 405. Niphobolus (subgen.), 552; adnascens, 552, 598; angustatus, 559; cau- datus, 552; costatus, 553; deter- gibilis, 555 ; elongatus, 552 ; fissus (B1.), 554 ; floccigerus, 554 ; levis, 552; macrocarpus, 559; nummu-— larisfolius, (Bedd.) 576, (J. Sm.) 554; porosus, 554, Schmidianus, 554; spherocephalus, 559; stic- ticus, 554; subfurfuraceus, 553; varius, 552; venosus, 553. Notholena carnosa, 575, 598; Ma- rante, 567; persica, 455; pilosel- loides, 575, 598; undulata, 598. Nullipores closely allied to Corallines, (footnote) 207. Nymphseacese, self-fertilization sp. of, 349. : 633 Odontoloma Boryanum, 451; repens, 451. Odontotrema longius, 239; majus, 239 ; minus, 239. (Ecidiacei, of Brisbane, 407. (Ecidium apocynatum, 407. Oidium lactis, filaments of, 421. Oleandra Cumingii, 542; mussfolia, 542; neriiformis, 541; Wallichii, 542, 603. Olfersia triquetra, 473. Omphalocarpum, 1; description and position of, 13; elatum, 16; pro- cerum, 15. Onagraceæ, plumes on seed of, 165; self-fertilization sp. of, 363, 392. On Adoxa Moschatellina, by Rev. G. Henslow, 194. On a Collection of Fungi made by Mr. Sulpiz Kurz, by F. Currey, 119. On a new Species of Helvella, by W. Phillips, 423. On Mycoidea parasitica, a new genus of parasitic Alge, and the part which it plays in the formation of certain Lichens, by Dr. D. D. Cun- ningham, 301. On Napoleona, Omphalocarpum, and Asteranthos, by J. Miers, 1. On origin of tertiary and quinary symmetry of Flowers with in- definite and spirally arranged mem- bers, by Rev. G. Henslow, 194. On some points in the Histology of certain species of Corallinacew, by Major-Gen. R. J. Nelson and Prof. Duncan, 197. On some points in the Morphology of the Primulaces, by Dr. Maxwell T. Masters, 285. On the African species of the genus Coffea, Linn., by W. P. Hiern, 169. On the Geographical Distribution of the Meliacee, by M. Casimir de Candolle, 233. On the Growth of the Flower-stalk of | the Hyacinth, by A. W. Bennett, 139. On the Hygroscopie Mechanism by which certain Seeds are enabled to bury themselves in the ground, by F. Darwin, 149. On the minute Structure and mode 634 of Growth of Ballia callitricha, by W. Archer, 211. On the Nature of the Corolla of Pri- mula, by Rev. G. Henslow, 195. On the occurrence of Conidial Fruc- tification in the Mucorini, illus- trated by Choanephora, by Dr. D. D. Cunningham, 409. On the origin of Floral ZEstivations, with Notes on the Structure of the Cruciferousflower, on that of Adoxa, and on the Corolla of Primula, by the Rev. G. Henslow, 177. On the Origin of the prevailing sys- tems of Phyllotaxis, by the Rev. G. Henslow, 37. On the Origin of the so-called Scor- pioid Cyme, by the Rev. G. Hens- low, 613. On the Self-fertilization of Plants, by the Rev. G. Henslow, 317. On the Symmetry of a Cruciferous flower, by Rev. G. Henslow, 191. Onoelea orientalis, 434. Onyehium auratum, 458, 596; ca- pense, 459; japonicum, 459, 596, var. intermedia, 459, var. (Kurz) 458, 459, var. multisecta, 459 ; lu- cidum, 459; multisectum, 459,597. Opegraphia saxicola, 280. Ophioglossum Aitchisoni, 586; cir- cinatum, 583; cordifolium, 586, 596; filiforme, 584; flexuosum, 583, 584; japonicum, 585; laci- niatum, 587; pedatum, 583; pen- dulum, 586, 587; polyphyllum, 586 ; reticulatum, 586, 596; scan- dens, 584; vulgatum, 586, var. . Aitehisoni, 586. Orchidaces, self-fertilization sp. of, 378. Ornithogalum benguellense, 248 ; ce- pefolium, 248. Nonne. self-fertilization sp. of, Loge Claytoniana, 582, 596; crispa, 459 ; interrupta, 582, 583; japonica, 583; lanceolata, 579; Leschenaultiana, 583, 596; Lu- naria, 587; monticola, 582, 596; regalis, 583, 596; speciosa, 583, 996; virginiana, 582; bebe Si DN. e INDEX. Palicourea (Aubl.), 169. Papaver, 178, 180 ; orientale, 180. Papaveraceæ, self-fertilization sp. of, 349. Paradigma, history and generic cha- racter of, 30, 31; Galeottiana, 31. Parasitic alga (Mycoidea parasitica), Dr. Cunningham on, 301. Parkeria pteridioides, 471. Passifloraces, self-fertilization sp. of, 366. Patagonula, 28; history of, 27. americana, 28; bahiensis, 30 ; glabra, 29; Tweediana, 29. Patania Elwesii, 436. Pavetta (Linn.), 169. Pelargonium, growth of flower-stalk of, 143. Pellæa calomelanos, 461; geraniæ- folia, 597 ; gracilis, 460 ; nitidula, 455, 460 ; nudiuscula, 455; Tam- burii, 460. Peranema aspidioides, 434 ; oides, 435, 599. Perigara globosa, 86 ; valida, 94. Pertusaria graphica, 275 ; incarnata, 241. Petersia, 107; africana, 108; ipit of, 48. Peziza aurantia, 128; cinereonigra, 404 ; rutilans, 128 ; scutellata, 404; vinoso-brunnea, 404. Phalloidei, of Brisbane, 405. Phegopteris (subgen.) 542; Dryo- pteris, 545 ; luxurians, 548 ; opaca, 568; polypodioides, 544; Rober- tiana, 545; rugulosa, 546; stegno- gramme, 569; Totta, 567; uro- phylla, 547 ; vulgaris, 544. cyathe- Phillips, W., new species of Helvella, | 423. Phlyctis, neo-zelandie, 281; ocel- lata, 281; sordida, 281. Phorolobus crispus, 459. Phycopeltis, 315. Phyllody of Primulaces, 286. Phyllotaxis, Airy’s condensation- theory of leaf-arrangement criti- cized, (footnote) 38; correlation between internodes and angular divergences, 40 ; di- and tristichous and pseudo-distichous arrangement, | 43; miss cg zr Dee pri- | mary series, 41; law of angular distance, 38; of Lepidodendron (fossil), 40 ; opposite leaves fail to produce 4 and 4 arrangement, 42; origin of cycles, 39; origin of pre- vailing system of, Rev. G. Henslow on, 37; reduction from whorled leaves, (footnote) 44 ; relative ages, di- and monocotyledons, 44; re- sults of arrest of development, 44 ; scheme of ordinary phyllotaxis, 44 ; spiral arrangement deduced from opposite and decussate leaves, 37 ; theoretical conditions, 41. Phymatodes (subgen.), 557 ; bum, 564; polycarpa, 561. Pinguiculacee self-fertilization sp. of, 377. Piptocephalis, fructification of, 420. Pistil, as adapted to self-fertilization, 380. Placodium argillaceum, 282; illitum, 282; lecanorinum, 282; Rhuderi, 282. Plagiogyria adnata, 472; glauca, 472 ; pycnophylla, 473 ; scandens, 472; triquetra, 473. Planchonia, charac. and history of, 90. crenata, 91; elliptica, 93; lit- toralis, 94 ; nitida, ( Bl.) 93, (Miq.) 88; sundiaca, 81, 92; tetra- ptera, 93; undulata, 92; valida, 81, 94. Plants, self-fertilization of, by the Rev. G. Henslow, 317. cyrtolo- Pleoenemis (subgen.), 534; Clarkei, 536 ; javanica, 535 ; Luzeana, 535. Pleopeltis (subgen.), 566 ; capitellata, 566; dilatata, 566 ; ebenipes, 564 ; Griffithiana, 560 ; hemionitidea, 562; heterocarpa, 561; himalay- - ensis, 566 ; incana, 596; irioides, 561; Lehmanni, 566; leiorhizon, 567; longissima, 565; malacodon, 563, 564; normalis, 558; nuda, 558 ; ovata, 560; oxyloba, 563; Parishii, 564 ; punctata, 561 ; rhyn- cophylla, 560; Stewartii, 563; ` superficialis, 557 ; tridactyla, 562. Plethostephia, 32; angiocarpa, 32. Peecilopteris costata, 580; flagel- lifera, 580; terminans, 581. . Polybotrya appendiculata, 571; i termedia, "on mal neglecta, 578; virens, 581. Polygales, self-fertilization sp. of, 352. Polygonacee, self-fertilization sp. of, 378, 393. Polypodium acrostichoides, 513, 552; aculeatum, 509 ; acutissimum, 558, 605; adherens, 605; adnascens, 552, 600, 601; adnatum, 602; alternans, 602; alternifolium, 565, 601; altissimum, 433; amoenum, 550, 601, var.? tonglensis, 550; angustatum, 559, 661, var. de- pauperata, 559; appendiculatum, (Bedd.) 543, (Bk.) 516, var. squa- mestipes, 543; arboreum, 605; argutum, 551, 552, 601; arista- tum, 511; atropunctatum, 558; auriculatum, 433, 543, (L.) 507, ( Wall.) 569, 601; Barometz, 435 ; Blumei, 570 ; Boothii, 555 ; brun- neum, 544, 602; caleareum, 545; capitellatum, 566, 601; eaudatum, 552; caudigerum, 530, 601; chat- tagrammicum, 427, 548, 549 ; chi- nense, 558; cicutarium, 539; cla- thratum, 427, 559; comosum, 602; confluens, 602; coniifolium, 546, 601 ; conjugatum, (Hk. & Bk.) 557, (Lamk.) 557; contaminans, 432, 602; contiguum, 557, 558, 601 ; cordifolium, 540; coriaceum, 559; coronans, 556, 557, 601; costatum, 553, 555, 600; cre- natum, 525; curvinervium, 601; eyrtolobum, 563, 564 ; dareseforme, 497, 443 ; davallioides, 546 ; den- tigerum, 491, 602; detergibile, 555; dichotomum, 428 ; dilatatum, 536, 565, 601; distans, 544, 568, 601, var. adnata, 544, var. glabrata, 544, var. minor, 545; Dodgsoni, 563; drepanopterum, 493, 494; | Dryopteris, 545; ebenipes, 563, 564, 601, var. Oakesii, 564, var. Parishii, 564; ellipticum, 570; elongatum, 513, 601; erubescens, 516, 543, 598, 602; erythrocar- pum, 427, 565; exaltatum, 541; ` excavatum, 558, 559, 565; falca- tum, 512; Fieldingianum, 551; Filix-Mas, 519; Finlaysonianum, 605 ; fissum, 554, 600, var. flocci- nodiflora, 577 ; | INDEX. gerum, 554; floeculosum, 554, 555, 600 ; furfuraceum, 601; fra- grans, 454 ; giganteum, 433, 601 ; glabrum, 561, 601; gladiatum, 558, 601 ; glaucistipes, 601 ; glau- eum, 428 ; grandifolium, 560, 601; Grevilleanum, 558, 605 ; Griffithi- anum, 560 ; Griffithii, 544 ; gibbo- sum, 549 ; hastatum, 562, 564, 601, var. oxyloba, 563, var. Thunbergii, 563; hemionitideum, 561, 601; Hendersoni, 550; heteractis, 553, 554; heterocarpum, 560, (BL) 561, 562, (Hk. & Bk.) 561; hi- malayense, 566 ; Hippocrepis, 539 ; Horsfieldii, 601; hymenodes, 557, 579, 601; hyperboreum, 434 ; in- volutum, 570 ; irioides, 603; jain- tense, 427, 552; javanica, 535; juglandifolium, 566, 601, var. bi- serialis, 566; khasyanum, 549; Kulhaitense, 493, 531; lachnopus, 550, 551, 601; lanceolatum, 560 ; latebrosum, 432 ; Lehmanni, 566 ; leiopteris, 558; leiorhizon, 566, 567, 601; Leschenaultianum, 602 ; Leuzeanum, 535; Linnei, 556, 601; Lindleyanum, 601; lineare, (Burm.) 428, 558, 559, (Thunb.) 601, 605, var. polymorpha, 559, var. steniste, 559 ; lineatum, 548, -` 601, var. penangiana, 548 ; lingua, 555, var. (Hk. & Bk.), 553; longi- folium, (.BI.) 601 ; longifrons, 558, 601 ; longipes, 544, 601; longissi- mum, 556, ( Mett.) 565 ; lonchites, 506; loriforme, 558, 559, 601; loxogramme, 570; lucidum, 567; luxurians, 548; macrodon, 536; malacodon, 564; marginale, 447, 602, ( Wall.) 546, 527, 601 ; mau- ritianum, 601; melanopus, 564; membranaceum, 560, 601; me- niscioides, 547; microrrhizoma, 551, 601; molle, 533; mollius- eulum, 533, 602; moulmeinsis, 566; multilineatum, 494, 547, 548 ; mysurense, 554, 600 ; nemo- rale, 533, 601; neriifolium, 601; nigrescens, 565, 601, var. poly- phylla, 601; normale, 557, 558, 601; nummulariefolium, 553, var. obovatum, 554; obliqua- | tum, 549; oppositum, 524, 605; SECOND SERIES.— BOTANY, VOL. I. | 635 ornatum, 528, 545, 546, 601 ; ova- tum, 560, 601; oxylobum, 601; oxyphyllum, ( Wall.) 486, 493, 602, (Kze.) 561; pallidum, 545, 546; paludosum, 544; palustre, 577; parasiticum, 533; Parishii, 564; parvulum, (Bedd.) 549, (Bory) 549; pedunculatum, 570; pertu- sum, 552, 600; phegopteris, 544 ; phlebodes, 558, 559 ; phymatodes, 557, 602; polycephalum, 561, 601; polyodon, 512, 605; polyphylla, 565 ; polypodioides, 544 ; porosum, 554, 600; pothifolium, 571; pro- liferum, 548, 601; propinquum, 556, 563, 600, var.? (Wall.), 563; pteropus, 562, 601; pubigerum, 535, 605 ; punctatum, (Swtz.) 601, (Thunb.) 528, 546, 561; quercifolium, 555, 556, 557, 601; rhyncophyllum, 560 ; rivale, 556 ; Robertianum,545 ; rostratum, 557 ; rubrinerve, 547 ; rugulosum, 545, var. hirsutum, 546; Russelli- anum, 528, 605; scabridum, 531 ; scabrum, 605 ; Scottii, 542 ; secun- dum, 529, 601; semibipinnatum, 605; serra, 601; serricula, 549 ; sesquipedale, 558, 601; siifolium, 538 ; sinuosum, 605 ; sophoroides, 605 ; Speluncee, 448 ; spheerocepha- lum, 559, 601 ; stenophyllum, 559 ; Stewartii, (C. B. C.) 563, (Bedd.) 563; stipitum, 544; stigmosum, 553, 600 ; subamenum, 427, 550; subarboreum, 605 ; subauriculatum, 551; subdigitatum, 546 ; subfalca- tum, 549; subfurfuraceum, 553, 555; subpectinatum, 601; subtri- pinnatum, 427, 545; superficiale, 557, var. semilinearis, 558 ; teneri- caule, 546, 602, (Hk.) 566, ( Wall.) 528; tenerifrons, 549 ; Thelypte- ris, 544; tomentosum, 555; Tot- tum, 567 ; trichodes, 546; tricho- manoides, 549, 550; tridactylon, 562, 601; trifidum, 563; triplum, 563; truncicola, 549; umbro- sum, 433, 632; unitum, 529; urophyllum, 547, 571, 601, var. khasiana, 547; varium, 552; ve- nosum, 605; venustum, ( Wall.) 566, (Desv.) 567, 601, var. nipho- boloides, 567; verrucosum, 552, 4g 636 601 ; viscosum, (Roxb.) 546, (Zipp.) 546; vittarioides, 552, 601 ; Wal- lichii, 555, 601 ; Wightianum, 558, 605; Willdenovii, 556 ; Zippellii, 561, (BL) 562, (Mett.) 562; zos- tereforme, 601; Zollingerianum, 562. Polyporei, 121; of Brisbane, 400. Polyporus affinis, 122, 401; amboi- nensis, 122; anebus, 124; appla- natus, 123; Broomei, 402; bru- malis, 400 ; brunneo-pictus, 122 ; ealeeus, 402; cichoraceus, 402; cinerascens, 124; cinereo-fuscus, 124, 402; cinnabarinus, 124; crassipes, 122; Feei, 124; ferreus, 124; flabelliformis, 122, 401; florideus, 121; fruticum, 401; funalis, 123, 402; fusco- lineatus, 401; hirsutus, 403 ; holosclerus, 123; hypoplastus, 121 ; ignarius, 401; incertus, 123 ; luci- dus, 122, 401 ; luteo-nitidus, 400 ; luteo-olivaceus, 402; marginatus, 123 ; modestus, 122 ; nilgheriensis, 122; perennis, 121; Persoonii, 123; picipes, 122, 401; pinsitus, 124; platotis, 401; quadrans, 400; rhipidium, 401; rubidus, 123, 402; sanguineus, 122, 401; scruposus, 124 ; senex, 401 ; Split- gerberi, 122 ; versicolor, 124 ; xan- thopus, 121, 400; xerophyllaceus, 124, Polysaecum olivaceum, 406 ; pisocar- pium, 406, Polystichum (subgen.), 585 ; aculea- tum, 509 ; aristatum, 511 ; Atkin- soni, 506; auriculatum, 489; ilicifolium, 506, var. minimum, 506, var. minus, 506 ; Lachenense, 506; Lonchitis, 506 ; Prescottia- num, 510; radicans, 507; sikki- mense, 525; stimulans, 508; . Thomsoni, 508, 509; truncatum, 2 UM Porina endochrysa, 280. Portulaces, self-fertilization sp. of, . 992. Preliminary Note on Growth of Female flower-stalk of Valisne- ria spiralis, by A. W. Bennett, ET T ; Primroses, double, 287, 293, INDEX. Primula, corolla of, Rev. G. Henslow on, 177. , wstivation and diagrams of, 177, 189 ; nature of the corolla of, 195; eupped petals of, 289. Primula cortusoides, 286 ; japonica, 286 ; preenitens, 286, sinensis, 286 ; vulgaris, 189. Primulaces, androscium of, 289 ; bib- liography of, 297; calyx of, 285 ; chorisis of, 288; corolla of, 286; develop. - parts of flower, 287; enation of, 288, 289; gynecium of, 292 ; morphology of, Dr. Masters on, 285 ; ovules of, 295 ; pistillody, stamens of, 290 ; placentation of, 292, Trécul’s view of, (ftnote) 293 ; self-fertilization sp. of, 376, 393 ; some gen. and sp. subject to changes, 727; summary of mor- phology, 296 ; tubular petals in, 289; variations in calyx of, 286. Prismatomeris (Thw.), 169. Prunus, calycine lobes of, 196. Psathyra, 119. Psathyrella, 119. Pseud-Allantodia (subgen.), 495. Psilosiphon cyanescens, 272; fra- grans, 272; odoratissimus, 273. Psilotum flaccidum, 596; nudum, 589, 596 ; triquetrum, 589,596. Psychotria ( Linn.), 169. Ptéris acuminatissima, 461; alterni- folia, 604 ; amplectens, 461, 597 ; amplexicaulis, 461; anamallayen- sis, 469 ; angustata, 597 ; angus- tifolia, 574; aquilina, 468, 597, 604, var. esculenta, 468, var. lanu- ginosa, 468; argentea, 458; ar- guta, 604; aspericaulis, 465, 467, 470, 597; aurita, 471; aus- tralis, 505; brevisora, 468; biau- rita, 469, 470, 597, var., 604, var. geminata, 469; bicolor, 457; Blumeana, 466; brevisora, 467, 468 ; cxespitosa, 455, 597 ; calome- lanos, 461; caudata, 468; ehry- socarpa, 458; costata, 461; cre- nata, 462, 463, 467 ; cretica, 452, 462, 463, 464, 597; crispa, 459; dactylina, 463 ; Dalhousie, 465; densa, 468, 597 ; digitata, 463, 597; dimidiata, 464; diversifolia, 461; ensiformis, (Burm.) 463, 464, 604, 605, (Ham.) 605, var. Grevilleana, 464; esculenta, 468; excelsa, 467, 597; farinosa, 457; firma, 468, 597; flabellata, 464 ; gemi- nata, 604; gracilis, 455, 460; graminifolia, 574; ^ Grevilleana, 464, 466, 605; Griffithii, 464; hastata, 461, 597; heterodactyla, 464; incisa, 471, 597 ; læta, 462, 597; lanigera, 468; lanuginosa, 468, 597 ; linearis, 597 ; longifolia, 461, 597; longipes, 465, 468; longipinnula, 466, 467, 469, 597 ; lorigera, 467, 468, 597; ludens, 470, 597; lunulata, 452; mar- ginata, 471; multidentata, 463, 605; multifida, 470; mysurensis, 470, 597; nemoralis, (Hk. 4 Bauer) 465, (Willd.) 469, 597; nervosa, 462, 597; nitidula, 460, 597 ; palmata, 470 ; pectinata, 465, 470; pedata, 470 ; pellucens, 468, 470; pellucida, 462, var. steno- _ phyla, 463, var., 597; piloselloides, 579 ; polita, 597 ; pyrophylla, 465 ; quadriaurita, 465, 466, 467, ( Hk.) 467, (Jietz.) 467, 469, 470, 597, var. Blumeana, 466, var. khasiana, 466, var. major, 466 ; radiata, 505 ; recurvata, 468, 597; repandula, 466; revoluta, 468; revolvens, 471; sagittifolia, 471; scabra, 605 ; scabripes, 463, 597; scandens, 577; semihastata, 467, 468, 597; semipinnata, 464, 597; setigera (or var.), 465 ; sinuata, 471, 597 ; spinescens, 465; Stelleri, 460; subquinata, 465, 467; succulenta, 471; subindivisa, 427, 467 ; sub- pedata, 597 ; terminalis, 467, 597 ; tripartita, 470; umbrosa, 470, 597 ; varians, 455, 597 ; venulosa, 462; vespertilionis, 471; vittata, (Z.) 461, (Roxb.) 461; Wallichiana, — 468, 469, 470, 597, var. quadri- - pinnata, 470; Wightiana, 468, ` 604 ; Zollingeri, 468. Queensland, fungi of, 399. Randia genipeflora (DC.), 169. ` Ranuneulaces, self-fertilization sp. awn of, 164. E Se SE Ranunculus bulbosus, sestivation of, 189. Report on the Liliacew, lridaces, Hypoxidaces, and Hæmodoraceæ of Welwitsch's Angolan Herbarium, by J. G. Baker, 245. Review of the Ferns of Northern India, by C. B. Clarke, 425, Rhizophora, embryo of, 52. Rhizormorpha Harrimanni, 407. Rhizophoraces, remarks on, 54. Rhododendron Griffithianum, maximum, 19. Ribes coccineum, sstivation of, 190. Romneyee, 180. Rosaceze, self-fertilization sp. of, 362, 392. Rubiacese, self-fertilization sp. of, 366, 392. Rudgea (Salisb.), 169. 19; Sacellium, gen. charac. of, 25 ; lanceo- latum, 25. Sagenia alata, 536; cicutaria, 535 ; coadunata, 540; gigantea, 539 ; heterocarpa, 537; hippocrepis, 539 ; macrodonta, 539 ; pteropus, 539; silhetensis, 540 ; variolosa, 540. Salvinia natans, 598. Samolus Valerandi, 286; variation corolla of, 287. Sandersonia littonioides, 262. Sandorieum indieum, 236; (Main- gayia) marginatum, 236. Sanseviera bracteata, 253. Saxifragaces, self-fertilization sp. of, 365. Schizwa dichotoma, 595; digitata, 583, 595. Schizobasis angolensis, 255. .— Schizoloma ensifolium, 452 ; Griffithi- anum, 452. . Schizophyllum commune, 121, 400. Scilla arenaria, 249; benguellensis, 249; congesta, 250; flaccidula, 249 ; hispidula, 248 ; lancesfolia, 250; laxiflora, 250; platyphylla, 250; polyphylla, 249 ; simiarum, 249. Sclerodesma Bovista, 406. Scolopendrium Ceterach,504 ; dubium, 496. Scorpioid cyme, Rev. G. Henslow on, 613. INDEX. Scorpioid cyme as exemplified in La- gerstroemia, 613 ; critical remarks on, 617; description of, 613 ; Hens- low's views, 613; in Silene pen- dula, 613; in Lathrea Squama- ria, 613; Kaufmann and War- ming’s views, 620; oscillating ar- rangement, 613; Payer’s defini- tion, 619 ; Sachs’s view, 619 ; sum- mary of successive steps to, 618. Serophulariacesm, self-fertilization sp. of, 370, 393. Seeds burying themselves, hypothesis on, 166; hygroscopic mechanism of, 149. Self-fertilization of Plants, Rev. G. Henslow on, 317, , advantages derived from, 319 ; are fittest to survive struggle for life, 396 ; are perfectly healthy, 388; chief facts tabulated, 322; cleistogamous flowers self-fert., 347; conservation of energy in reduct. stamens and pollen, 380; Darwin’s views on, 320; fertility and health does not decrease, 382 ; fertility may increase, 383; few flowers (physiolog.) self-sterile, 323; flowers may become self- fertilizing in absence of insects for intercrossing, 331 ; free from com- petition self-fertiliz. may equal intercrossed, 383; grafting effects of, 333 ; highly self-fert. var. arise under cultivation, 339; inconspi- cuous flowers self-fert. or else ane- mophilous, 342; introductory re- marks on, 317 ; loss of colour cor- related with self-fertilization, 327; origin of conspieuous flowers and causes inducing reversion to selí- fertilization, 333; majority of flowers self-fertile, 323; many plants morphol. self-sterile, 324 ; Meehan’s views on, 320; more productive than flowers depen- dent on insects, 390; naturalized abroad, replace native vegeta- tion, 395; partial and total arrest devel. of corolla, 327; partial ex- cision of corolla and stamens, 326 ; physiol. and morphol. self-sterile plants may become self-fertile, 325; plants may become self- 637 fertilizing on reduction of tempe- rature, 332; relative fertility may surpass crossed plants, 382; secured by flowers closing after expansion, 329; some derive no benefit from crossing with same on distinct stock, 385 ; some flower-buds never open, 330; special adap. for self- fertil. of Brit. plants, 347; sum- mary of adaptations to secure self- fertil., 380; value of intercrossing, 318; withering of corolla, 325; world-wide distrib. of self-fert. Brit. plants, 391. - Selliguiea decurrens, 571 ; Hamiltoni, 570; lanceolata, 570; hemioni- tidea, 562; Wallichiana, 570. Shuta, native name of Agasta splen- dens, 60. Silene pendula, scorpioid cyme of, 614. Sitolobium strigosum, 436. Smilaceze (Angolan), 264. Smilax Kraussiana, 264. Solanaces, self-fertilization sp. of, 370, 392. Solanew, scorpioidal infloresc. of, 616. Sphacelaria callitricha, struct. of, 220. Sphzria phaselina, 130 ; sublimbata, 130. Sphzriacei, 129; of Brisbane, 405. Spheropteris barbata, 435, 526, 599 ; Hookeriana, 434, 603. Sporocybacei, 129. Stamens as adapted to self-fertiliza- tion, 380. Stegnogramme aspidioides, 569. Stemonitis fusca, 406. Stenochlena pycnophylla, 473 ; scan- dens, 577 ; triquetra, 473. Stereum adustum, 127; cyathiforme, 127; elegans, 126; lobatum, 126, 403 ; medicum, 127 ; Ostrea, 126 ; papyrinum, 127; scytale, 127. Sticta canaliculata, 282. Stilbacei, of Brisbane, 406. Stilbum erythrocephalum, 129; in- conspicuum, 129. Stipa-awn, 155, hygroscope, 156. Stipa pennata, descrip. of seed, 149; hygroscopic action of awn of, 150 ; mechanism of torsion in awn of, 157. 638 Stravadium, 79, 80; acuminatum, 102; acutangulum, 80; album, 65; angustatum, 105; excelsum, 112; coccineum, 83; cochinchi- nense, 101 ; costatum, 88; demis- sum, 81, 83; denticulatum, 88; globosum, $86; gracile, 86; Horsfieldii, 85; insigne, 75; in- tegrifolium, 89 ; lucidum, 88; lu- zonense, 84; macrophyllum, 61 ; obtusangulum, 77, 81; pubescens, 83 ; Reinwardtii, 188 ; reticulatum, 87; Rheedii, 82; rubrum, 68, 69 ; Sarcostachys, 102; semisutum, 89 ; serratum, 87; spicatum, 85. Structure of Ballia callitricha, 211. Struthiopteris orientalis, 434. Suppression of 5th part cycle of flowers, 192. Swietenia Mahagoni, 236. Symplocines, 1. Syngramma vestita, 568. Teeniopsis lineata, 574. Tænitis blechnoides, 575; interrup- tus, 575; piloselloides, 575; pte- roides, 575; pusilla, 574. Thamnopteris Grevillei, 476; nidus, 475; phyllitidis, 475; Simonsiana, 476. Thecacophora globuligera, 407. Thelenella Wellingtonii, 280. Thelephora palmata, 126 ; pusilla,126. Thelotrema monosporum, var. pa- tulum, 278 ; saxatile, 278. Tigellum, term defined, 53. Torsion in Avena, 163 ; of cells, 161; of seeds, 164; on mechanism in Stipa-awn, 157 ; power of, 161. Torulacei, of Brisbane, 406. Trametes cingulatus, 195; lobatus, 124; occidentalis, 125; perennis, 402; rigida, 402; umbrinus, 124, Tremella foliacea, 127. Tremellini, 127, of Brisbane, 404. _ Tremellodon gelatinosum, 126. Trichia pyriformis, 128. Trichilia, 235; havanensis, 236 ; pe- ruviana, distrib. of, 233. INDEX. Trichobasis rubigo-vera, 407. - Trichócoma paradoxum, 128. Trichodermacei, 126, 128. .Trichogastres, of Brisbane, 406. Tricholoma, 119. $ Trichomanes alchemillefolium, 598 ; anceps, 598; auriculatum, 441 ; Belangeri, 441; bipunctatum, 440 ; 598; campanulatum, 604; chi- nense, 449 ; corticola, 440 ; den- ticulatum, 438 ; digitatum, 440 ; dissectum, 441; Filicula, 440; gracile, 440; insigne, 440; java- nicum, 442, 598; Kunzeanum, 441 ; Kurzii, 440 ; limbatum, 441; longisetum, 598; muscoides, 439, var. sublimbatum, 439; nanum, 440; parvulum, 440; plicatum, 440; proliferum, 441; pyrami- dalis, 598; pyxidiferum, 440, var. limbatum, 441; radicans, 441, 442, 598, var. anceps, 441 ; rigi- dum, 442, 598; Schmidianum, 440 ; setigerum, 442, 598 ; strigo- sum, 447 ; sublimbatum, 439, 440 ; umbrosum, 441, 598; undulatum, 598 ; villosa, 598. Trypethelium astroidea, 278; Cu- mingii, 278 ; pyrenuloides, 278. Tsjeria Saamstravadi, 82. Tulbaghia sequinoctialis, 246. Typhula fuscipes, 127. Ugena microphylla, 584; polymor- pha, 584; semihastata, 584. Umbelliferee, self-fertilization sp. of, 365, 392. Urceolaria novæ-zelandiæ, 275. Urginea psilostachya, 247; ehloran- tha, 248 ; comosa, 247. Ustilago axicola, 407 ; carbo, 407. Valerianaces, self-fertilization sp. of, 367. | Vallisneria spiralis, sd of female . flower-stalk of, 133. Vavea Amicorum, 235. Vellosia eapillaris( Welw.),264 ; squar- rosa (Welw.) 264; stenophylla ` (Welw.), 265; velutina: (Welw.), 265. Verbenaces, self-fertilization sp. of, 393. Verrucaria aspistea, 280 ; astata, 280; circumpressa, 278; dealbata, 280 ; epidermidis, var. gemellipara, 279 ; Eschweileri, 280 ; fumosaria, 239 ; gemellipara, 278; Larbalestierii, 242; micromma, 279; minutis- sima, 270; myriospora, 145; ne- ottizans, 239; nitida, 280; oc- culta, 279; pertenuis, 239; pru- ino-grisea, 279; pyrenastroides, 278; saxicola, 280; subbiformis, 279; succina, 145 ; Thwaitesii, 278. Viburnum Tinus, sestivation of, 189. Violacem, self-fertilization sp. of, 351. Vittaria divergens, 604; elongata, 573, (Swtz.) 598, (Wall.) 574; falcata, var., 574; flexuosa, 574; interrupta, 451; japonica, 574; lineata, 574; minor, 574, var. minima, 574; plantaginea, 574; rigida, (Kaulf.) 573, ( Wall.) 574, 578 ; sikkimensis, 574; zostere- folia, 574. Volvaria, 119. Vutu-dina (a Barringtonia ?), 57. Walleria angolensis, 262. Welwitsch’s Angolan Herbarium, re- port on Liliaceæ &c. of, 215. Woodsia elongata, 435; hyperborea, 434, 435; lanosa, 435; mollis- sima, 435, f. subcordata (Milde), 434, Woodwardia radicans, 475, 595. Xerophyta capillaris, 264; squarrosa, 264; stenophylla, 265; velutina, - ow Xylaria digitata, 129 ; flagelliformis, . 129 ; guyanensis, 129 ; hypoxylon, 129; Kurziana, 129; mutabilis, 129; pileiformis, 405 ; polymorpha, 129, 405; rhytidophlea, 405; tabacina, 129. PRINTED BY TAYLOR AND FRANCIS, LION COURT, FLEET STREET. _