Rust, Smut, Mildew, & Mould: An Introduction to the Study of Microscopic Fungi

CHAPTER IV.

Chapter 55,252 wordsPublic domain

_MILDEW AND BRAND._

DR. WITHERING’S “Arrangement of British Plants” in 1818 reached its sixth edition. This is less than half a century ago, and yet the whole number of species of Fungi described in that edition was only 564, of which three hundred were included under the old genus _Agaricus_. Less than eighty of the more minute species of Fungi, but few of which deserve the name of microscopic, were supposed to contain all then known of these wonderful organisms. Since that period, microscopes have become very different instruments, and one result has been the increase of Withering’s 564 species of British Fungi to the 2,479 enumerated in the “Index Fungorum Britannicorum.” By far the greater number of species thus added depend for their specific, and often generic characters, upon microscopical examination. The proportion which the cryptogamic section bears to the phanerogamic in our local Floras before 1818, now almost involuntarily causes a smile. Even such authors as were supposed to pay the greatest possible respect to the lower orders of plants could never present an equal number of pages devoted to them, as to the higher orders. Relhan, for instance, only occupies one-fifth of his “Flora Cantabrigiensis,” and Hudson one-fourth of his “Flora Anglica,” with the Cryptogamia. At the present time, it will be seen that, with a liberal allowance for “hair-splitting,” the number of British species of flowering plants scarcely exceeds three-fourths of the number of Fungi alone, not to mention ferns, mosses, algæ and lichens, and yet we have no “Flora” which contains them, and but a minority of our botanists know anything about them. If we need excuse for directing attention to some of the most interesting of these plants, let the above remarks suffice in lieu of formal apology.

“Mildew” is just one of those loose terms which represent no definite idea, or a very different one to different individuals. Talk of _mildew_ to a farmer, and instantly he scampers mentally over his fields of standing corn in search of the brown lines or irregular spots which indicate the unwelcome presence of _Puccinia graminis_, known to him, and to generations of farmers before him, as “mildew.” Try to convince a Norfolk farmer that anything else is “mildew,” and he will consider you insane for your pains. Speak of _mildew_ in your own domestic circle, and inquire of wives, or daughters, or servants, what it means, and without hesitation another, and even more minute species of fungus, which attacks damp linen, will be indicated as the true mildew, to the exclusion of all others; and with equal claims to antiquity. Go to Farnham, or any other hop-growing district, and repeat there your question,—What is _mildew_?—and there is every probability that you will be told that it is a kind of mould which attacks the hop plant, but which differs as much from both the mildew of the farmer and the laundry-maid as they differ from each other. The vine-grower has his mildew, the gardener his mildewed onions, the stationer his mildewed paper from damp cellars, the plasterer his mildewed walls, and in almost every calling, or sphere in life, wherever a minute fungus commits its ravages upon stock, crop, or chattels, to that individual owner it becomes a bug-bear under the name of “mildew.” Reluctantly this vague term has been employed as a portion of the title to this chapter, but it must be limited in its application to the “mildew of corn,” known to botanists as _Puccinia graminis_, and _not_ to include the numerous other microscopic Fungi to which the name of _mildew_ is often applied.

The origin of this term and its true application may undoubtedly be traced to _mehl-thau_, “meal dew.” A singular proof of the ignorance which prevails in regard to all the fungal diseases of corn, may be found in the fact that at least one of our best etymological dictionaries states that the _mildew_ in corn is the same as the _ergot_ of the French. Had the writer ever been a farmer, he would have known the difference; had he ever seen the two, he could scarcely have made such a mistake. It is barely possible for him ever to have heard the ergot of grain called by the name of _mildew_.

How long this disease has been known, is an unsolved problem. About the middle of the last century a tract was published on this subject in Italy, but this was probably not even the first intimation of its fungoid character. Before such conclusion had been arrived at, men may have struggled in the dark, through many generations, to account for a phenomenon with which they were doubtless familiar in its effects. In 1805, Sir Joseph Banks published his “Short Account,” illustrated by engravings from the inimitable drawings of Bauer, whereby many in this country learnt, for the first time, the true nature of _mildew_.

With a view to the clearer understanding of these parasites in the phases of their development, let us select one, and we cannot do better than adhere to that of the wheat and other graminaceous plants. A fine day in May or June dawns upon our preparations for a stroll, far enough into the country to find a wheat-field. Even now, with the area of the metropolis constantly widening, and banishing farmers and wheat fields farther and farther from the sound of Bow-bells, a corn field may be reached by a good stiff walk from Charing-Cross, or a six-penny ride at the most, in nearly any direction. Having reached the field, it may be premised that a walk into it of less than twenty yards will be sure to reward you with the fungus we are in quest of. Look down at the green leaves, especially the lower ones, and you will soon find one apparently grown rusty. The surface seems to be sprinkled with powdered red ochre, and grown sickly under the operation. Pluck it carefully, and examine it with a pocket lens. Already the structure of a healthy leaf is familiar to you, but in the present instance the cuticle is traversed with numerous longitudinal cracks or fissures, within which, and about their margins, you discern an orange powder, to which the rusty appearance of the leaf is due. Further examination reveals also portions in which the cuticle is distended into yellowish elongated pustules, not yet ruptured, and which is an earlier stage of the same disease. This is the “rust” of the agriculturist, the _Trichobasis rubigo-vera_ of botanists, the first phase of the corn mildew.

To know more of this parasite, we must have recourse to the microscope; having therefore collected a few leaves for this purpose, we return homewards to follow up the investigation. We will not stay to detail the processes of manipulation, since these will not offer any deviation from the ordinary modes of preparation and examination of delicate vegetable tissues.

The vegetative system of the “rust,” and similar fungi, consists of a number of delicate, simple, or branched threads, often intertwining and anastomosing, or uniting one to the other by means of lateral branchlets. These threads, termed the mycelium, penetrate the intercellular spaces, and insinuate themselves in a complete network, amongst the cells of which the leaf, or other diseased portion of the plant, is composed. High powers of the microscope, and equally high powers of patience and perseverance, are necessary to make out this part of the structure. We may regard the whole mycelium of one pustule, or spore-spot, as the vegetative system of one fungal plant. At first this mycelium might have originated in a number of individuals, which afterwards became confluent and combined into one for the production of fruit, that is to say, an indefinite number of points in the vicinity of the future mycelium developed threads; and these, in the process of growth, interlaced each other, and ultimately, by means of transverse processes, became united into one vegetative system, in which the individuality of each of the elementary threads became absorbed, and by one combined effort a spore-spot, or cluster of fruit, was produced. In the first instance a number of minute, transparent, colourless cellules are developed from the mycelium: these enlarge, become filled with an orange-coloured endochrome, and appear beneath the cuticle of the leaf as yellowish spots. As a consequence of this increase in bulk, the cuticle becomes distended in the form of a pustule over the yellow cellules, and at length, unable longer to withstand the pressure from beneath, ruptures in irregular, more or less elongated fissures (Plate VII. fig. 141), and the yellow bodies, now termed _spores_ (whether correctly so, we do not at present inquire), break from their short pedicels and escape, to the naked eye presenting the appearance of an orange or rust-coloured powder. In this stage the spores are globose, or nearly so, and consist of but one cell Plate VII. figs. 142, 144). It will afford much instructive amusement to examine one of these ruptured pustules as an opaque object under a low power, and afterwards the spores may be viewed with a higher power as a transparent object. The difference in depth of tint, the nearly colourless and smaller immature spores, and the tendency in some of the fully matured ones to elongate, are all facts worthy of notice, as will be seen hereafter.

A month or two later in the season, and we will make another trip to the cornfield. Rusty leaves, and leaf-sheaths, have become even more common than before. A little careful examination, and, here and there, we shall find a leaf or two with decidedly brown pustules intermixed with the rusty ones, or, as we have observed several times during the past autumn, the pustules towards the base of the leaf orange, and those towards the apex reddish-brown. If we remove from the browner spots a little of the powder, by means of a sharp-pointed knife, and place it in a drop of water or alcohol on a glass slide, and after covering with a square of thin glass, submit it to examination under a quarter-inch objective, a different series of forms will be observed. There will still be a proportion of subglobose, one-celled, yellow spores; but the majority will be elongated, most with pedicels or stalks, if they have been carefully removed from the leaf, and either decidedly two-celled, or with an evident tendency to become so. The two cells are separated by a partition or dissepiment, which divides the original cell transversely into an upper and lower cell, with an external constriction in the plane of the dissepiment (Plate IV. fig. 59). These bilocular or two-celled spores are those of the “corn mildew” (_Puccinia graminis_), which may be produced in the same pustules, and from the same mycelium, as the “corn rust,” but which some mycologists consider to be a distinct fungus, others only a modification or stage of the same fungus. After an examination of the different forms in the allied genera to which these chapters are devoted, we shall be able with less of explanation and circumlocution to canvass these two conflicting opinions.

Let us proceed, for the third and last time, to our cornfield, when the corn is nearly or fully ripe, or let us look over any bundle of straw, and we shall find blackish spots, from the size of a pin’s head to an inch in length, mostly on the sheaths of the leaves, often on the culm itself. This is the fully developed _mildew_, and when once seen is not likely afterwards to be confounded with any other parasite on straw (fig. 57). The drawings of Bauer have already been alluded to. Bauer was botanical draughtsman to George III., and his exquisite drawings, both of the germination of wheat and the fungi which infest it, are marvels of artistic skill. A reduced figure from part of one of his drawings is given (Plate IV. fig. 58), exhibiting a tuft of the bilocular spores of _Puccinia graminis_ bursting through a piece of wheat straw. These closely-packed tufts or masses of spores, when examined with a common lens, seem, at first, to resemble the minute sorus of some species of fern; but when seen with higher powers, the apparent resemblance gives place to something very different. The tufts consist of multitudes of stalked bodies, termed spores, which are constricted in the middle and narrowed towards either extremity. The partition, or septum, thrown across the spore at the constriction, separates it into two portions, each of which consists of a cell-wall enclosing an inner vesicle filled with the endochrome (fig. 59) or granular contents, in which a nucleus may often be made out. This species of _Puccinia_ is very common on all the cereals cultivated in this country, and on many of the grasses. A variety found on the reed was at one time considered a distinct species; but the difference does not seem sufficient to warrant a separation. However near some other of the recognized species may seem to approximate in the form of the spores, a very embryo botanist will not fail to observe the distinctive features in the spores of the corn mildew, and speedily recognize them amongst a host of others; subject, as they may be, to slight deviations in form, resulting either from external pressure, checks in development, or other accidental circumstances, or the variations of age.

There is no doubt in the minds of agriculturists, botanists, _savans_, or farm-labourers, that the mildew is very injurious to the corn crop. Different opinions may exist as to how the plants become inoculated, or how infection may be prevented or cured. Some have professed to believe that the spores, such as we have seen produced in clusters on wheat straw, enter by the stomata, or pores, of the growing plant, “and at the bottom of the hollows to which they lead they germinate and push their minute roots into the cellular texture.” Such an explanation, however plausible at first sight, fails on examination, from the fact that the spores are too large to find ingress by such minute openings. It is improbable that the _spores_ enter the growing plant at all. The granular contents of the spores may effect an entrance either through the roots or by the stomata, or the globose bodies produced upon the germination of the spores may be the primary cause of infection. We are not aware that this question has been satisfactorily determined. It is worthy of remembrance by all persons interested in the growth of corn, that the mildew is most common upon plants growing on the site of an old dunghill, or on very rich soil. As the same _Puccinia_ is also to be found on numerous grasses, no prudent farmer will permit these to luxuriate around the borders of his fields, lest they should serve to introduce or increase the pest he so much dreads.

The germination of the spores of the corn mildew is a very interesting and instructive process, which may be observed with a very little trouble. If the spores be scraped from the sori of the preceding year (we are not sure that those of the current year will succeed), and kept for a short time in a damp atmosphere under a glass receiver, minute colourless threads will be seen to issue both from the upper and lower divisions of the spores. These will attain a length several times that of the spores from whence they spring. The extremities of these threads ultimately thicken, and two or three septæ are formed across each, dividing it into cells, in which a little orange-coloured endochrome accumulates. From the walls of each of these cells, or joints, a small pedicel, or spicule, is produced outwards, the tip of which gradually swells until a spherical head is formed, into which the orange-coloured fluid passes from the extremities of the threads.[4] A quantity of such threads, bearing at their summits from one to four of these orange-coloured, spherical, secondary fruits, supply a beautiful as well as interesting object for the microscope. When matured, these globose bodies, which Tulasne has called _sporidia_, fall from the threads, and commence germinating on their own account. It is not impossible that the sporidia, in this and allied genera, may themselves produce a third and still more minute fruit, capable of diffusion through the tissues of growing plants, or gaining admission by their stomata. Nothing of the kind, however, has yet been of certainty discovered.

Footnote 4:

Similar in all essential particulars to the germination of _Aregma_ (Plate III. fig. 45).

Forty other species of _Puccinia_ have been recorded as occurring in Great Britain, to all of which many of the foregoing remarks will also apply—viz., such as relate to their two-celled spores being found associated with, and springing from, the same mycelium as certain orange-coloured one-celled spores; and also the main features of the germinating process.

A very singular and interesting species is not uncommon on the more delicate grasses, being found chiefly confined to the leaves, and produced in smaller and more rounded, or but slightly elongated, patches (Plate IV. fig. 60). We have met with it plentifully amongst the turf laid down in the grounds of the Crystal Palace at Sydenham, and also on hedge-banks and in pastures. The spores are rather smaller than those of _Puccinia graminis_, but, like them, much elongated, slightly constricted, and borne on persistent peduncles. The most prominent distinction may be found in the apices of the spores, which, in this instance, are not attenuated, but crowned with a series of little spicules, or teeth, whence the specific name of _coronata_ has been derived (Plate IV. fig. 62).

The Labiate family of plants and its ally the Scrophulariaceæ are also subject to the attacks of several kinds of Brand, a name, by the bye, often applied locally to the corn mildew and other similar parasites, and which may have originated in the scorched or _burnt_ appearance which the infected parts generally assume. In the former natural order the different kinds of mint, the ground-ivy, the wood-sage, and the betony, and in the latter, the water figwort and several species of veronica, or speedwell, are peculiarly susceptible; and on most a distinct species of _Puccinia_ is found. To provide against doubt which the less botanical of our readers may possess of the meaning or value of the term _Puccinia_, which has already occurred two or three times in this chapter, a brief explanation may be necessary, which more scientific readers will excuse.

In botany, as in kindred sciences, acknowledged species have their trivial, or specific name, generally derived from the Latin. In the last species referred to, this was _coronata_, meaning _crowned_, in reference to the coronated apex of the fruit. Any indefinite number of species with some features in common are associated together in a group, which is termed a _genus_, and the term prefixed to the specific name of each species constituting that genus is its generic name, also commonly derived from the Latin or Greek. In this instance it is _Puccinia_, derived from the Greek _puka_, meaning _closely packed_, singularly applicable to the manner in which the spores are packed together in the pustules. The common features, or generic distinctions, of this genus, are uniseptate spores borne on a distinct peduncle.

In returning to the species found on Labiate plants, let us suppose ourselves to have strolled towards Hampstead Heath, and south of the road leading from Hampstead to Highgate, near certain conspicuous and well-known arches, built for a purpose not yet attained, are two or three muddy ponds nearly choked up with vegetation. Some fine autumnal afternoon, we must imagine ourselves to have reached the margin of the most northern of these ponds, and amidst a thick growth of reeds, sedges, and other water-loving plants, to have found the water-mint in profusion and luxuriance, with every leaf more or less occupied, on its under surface, with the yellow spores of a species of rust (_Trichobasis_) mixed with the browner septate spores of the mint brand (_Puccinia Menthæ_). This is common also on the horse-mint and corn-mint; we have found it on the wild basil and wild thyme, and once only on marjoram. Having collected as many leaves as we desire, and returned to home and the microscope, we proceed to examine them in the same manner as we have already examined the mildew, and as a result of such proceeding arrive at the following conclusions:—The pustules are small and round, never elongated as in the corn mildew, and generally confined to the under surface of the leaves (Plate IV. fig. 69). The spores are subglobose, slightly constricted, and the two cells nearly two hemispheres, with their flat surfaces turned towards each other (fig. 70). The form delineated in figure 75 is that of the sorus of many of the epiphytal brands, the centre being occupied by the closely-packed spores, surrounded to a greater or less extent by the remains of the ruptured epidermis.

Although the species of Puccinia (_P. glechomatis_) found on the leaves of the ground-ivy is said to be very common, we sought it in vain amongst every cluster of that plant met with during last summer and autumn, until, nearly despairing of finding it at all, we at last encountered a plot of ground-ivy covering the ground to the width of two or three yards, and in length eight or ten, nearly every plant being attacked by the _brand_. This was in the corner of a pasture, and the only time we found infected plants. The fungus, however, may be as common as the plant in other localities. The pustules on the leaves are larger than those of the mints, and also confined to the inferior surface (fig. 73). The spores are elliptic and but slightly constricted; the apex is often pointed, though not always so much as in our figure (fig. 74).

Of other species found on allied plants we have not considered it necessary to give figures, or write much. The betony brand (_P. Betonicæ_, DC.) does not seem to be common enough to be readily found by any one desiring to examine it for himself; and the same may be said of the figwort brand (_P. Scrophulariæ_, Lib.), the wood-sage brand (_P. Scorodoniæ_, Lk.), and the speedwell brand (_P. Veronicarum_, DC.); all of these are, however, characterized by a distinct feature, or features, which have been considered of sufficient importance to constitute a separate species.

We have had occasion to refer incidentally to the brand found on the under surface of the leaves of the wood-anemone (_P. Anemones_, P.). This is one of the earliest and commonest species. Go wherever the wood-anemone abounds, in any of the woods lying immediately to the north of the metropolis, or any of the woods in Kent, and from March to May it will not be difficult to find attenuated, sickly-looking leaves, with the under surface covered with the pustules of this brand, looking so like the sori of some fern (fig. 65) that it _has_ been, and still _is_, sometimes considered as such. In Ray’s “Synopsis” (3rd edition, 1724), it is described in company with the maidenhair and wall-rue ferns; a figure is given of it in the same work (t. iii. fig. 1), and it is stated,—“this capillary was gathered by the Conjuror of Chalgrave.” When, afterwards, it was better understood, and the spots came to be regarded as true parasitic fungi, it still for a long time continued to bear the name, not even yet quite forgotten, of the Conjuror of Chalgrave’s fern.

An examination of the spores, both collectively in the pustules, and separately under a high power, will not fail to convince any one who has examined only the species we have already alluded to, that this parasite on the anemone (_P. Anemones_) is a true _Puccinia_, and a most interesting one. The two cells of the spores are nearly spherical, and the constriction is deeper and more positive than in any of the preceding. Moreover, the surface of the spore is minutely and beautifully echinulate, or covered with erect spines (Plate IV. fig. 66). Some few other of the species found in Britain have echinulate spores, but those are not common like the present. One word of caution to the amateur in search of the _Puccinia_ on the anemone. It will be fruitless looking for it on the large foliaceous bracts of the flower-stalk, since these may be turned up carefully, till the back aches with stooping, ere a solitary pustule will be found; but the true leaves, proceeding from the rhizomes, are certain soon to afford you specimens.

Everybody knows the dandelion, but it is not every one who has noticed the fungi found upon its leaves. These are most commonly of two kinds, or probably the unilocular and bilocular forms of the same species: the latter we have found in the month of May, and the former in August and September. The lower leaves of young seedlings have generally rewarded us with the best specimens of the septate-fruited brand (_Puccinia variabilis_, Grev.). The pustules occur on both sides of the leaf, and are very small and scattered (fig. 82). The spores are singularly variable in form: sometimes both divisions are nearly equal in size; sometimes the upper, and sometimes the lower, division is the smallest; occasionally the septum will be absent altogether; and more rarely, the spores will contain three cells. From the very variable character of the spores (fig. 83), the specific name has been derived.

No species in the entire genus makes so prominent an appearance as the one found on the radical leaves of the spear thistle (_Carduus lanceolatus_). This latter plant is exceedingly abundant, and so is its parasite (_Puccinia syngenesiarum_, Lk.). From the month of July till the frosts set in we may be almost certain of finding specimens in any wood. The leaves have a paler roundish spot, from one-twelfth to one-fourth of an inch in diameter, on the upper surface, and a corresponding dark brown raised spot on the under surface, caused by an aggregation of pustules, forming a large compound pustule, often partly covered with the epidermis. The individual pustules are small, but this aggregate mode of growth gives the clusters great prominence, and therefore they are not easily overlooked (Plate IV. fig. 63). Although not confined to this species of thistle, we have not yet found this _Puccinia_ on any other plant. The spores are elliptical, rather elongated, constricted, and without spines (fig. 64).

Other species of _Puccinia_ are found on Composite plants, but with none of these is the present fungus likely to be confounded, if regard be had to its peculiar habit. The leaves, for instance, of the common knapweed (_Centaurea nigra_) are often sprinkled with the small pustules of the centaury brand (_Puccinia compositarum_, Sch.); these generally occupy the under surface of the lower radical leaves (fig. 67); occasionally a few of the pustules appear on the upper surface. We have not often found this fungus in the neighbourhood of London on the leaves of the knapweed, but, on the other hand, we have encountered it very commonly on those of the saw-wort (_Serratula tinctoria_). The spores are oval, scarcely constricted, and not attenuated in either direction (fig. 68). Other Composite plants than those above named are liable to attacks from this parasite.

In our school-days we remember to have spent many a stray half-hour digging for “earthnuts,” under which name we, as well as our elders and betters, knew the tubers of _Bunium flexuosum_. Not then, nor for many years after, did we notice, or regard if we did notice, the distorted radical leaves and leaf-stalks, and the blackish-brown spots, which reveal the cause in the presence of a brand, or parasitic fungus, of this genus (_Puccinia Umbelliferarum_, DC.), which is extremely common on this, as well as some other allied plants. If any spot is searched where this plant grows in any profusion, before the flowering stalks have made their appearance above the surrounding grass, this _Puccinia_ will be readily found by the twisted, contorted, sickly appearance of the infested leaves (fig. 71), the petioles of which are often swollen and gouty in consequence. The sporidia are shortly stalked, and generally very much constricted (fig. 72). The species found on the stems of the hemlock, and also that on Alexanders (_Smyrnium Olusatrum_), are distinct; the spores of the latter being covered with tubercles or warts (figs. 55, 56). During a botanical ramble through Darenth Wood in April of the year just passed away, in some parts of which the sanicle abounds, we found the bright glossy leaves of this singular and interesting plant freely sprinkled with the pustules of a _Puccinia_ (_P. Saniculæ_, Grev.), which is not at all uncommon on this, but has not hitherto been found on any other plant. Dr. Greville, of Edinburgh, was the first to describe this, as well as many other of our indigenous minute Fungi. For many years he has toiled earnestly and vigorously at the lower cryptogams, as evidenced by his “Scottish Cryptogamic Flora,” published in 1823; and yet his continual additions to the records of science show him to be earnest and vigorous still.

We have by no means exhausted the catalogue of Fungi belonging to this genus found in Britain, nor even those commonly to be met with; but the fear of prolixity, and the desire to introduce a description of other forms into the space still remaining to us, prompt us to dismiss these two-celled brands with but a brief allusion to such as we cannot describe. Box-leaves are the habitat of one species, and those of the periwinkle (Plate VI. fig. 132) of another. One vegetates freely on the leaves of violets through the months of July and August, and another less frequently on the enchanter’s nightshade. Several species of willow-herb (_Epilobium_) are attacked by one _Puccinia_ (Plate IV. figs. 78, 79), and a single species by another. Plum-tree leaves, bean-leaves, primrose leaves, and the half-dead stems of asparagus, have their separate and distinct species, and others less commonly attack the woodruff, bedstraw (Plate VIII. figs. 172, 173), knotgrass, ragwort, and other plants less common, more local, or, to the generality of the non-botanical, but imperfectly known.

We have found, not uncommonly in the autumn, the scattered pustules of a brand on the stems and leaves of the goat’s-beard, occupying the places which were scarred with the remains of cluster-cups that had flourished on the same spots a month or two previously (Plate IV. fig. 76). The pustules are by no means minute, but elongated and bullate; the spores beautifully studded with warts (Plate IV. fig. 77). This species cannot certainly be identical with _Puccinia compositarum_ (Schlecht), _P. syngenesiarum_ (Lk.), or _P. tragopogonis_ (Corda). In none of these do the spores appear to be warted, and the habits of both the latter are different. Its nearest associate appears to be _P. centauriæ_ (Corda), at least in the fruit, and whilst the form and character of these organs are considered of any value in the determination of species, smooth spores cannot be associated, we think, with tuberculate or echinulate spores under the same name.

In the spores of the species to which attention has been more specially directed we have types of the principal forms. In the “corn-mildew” they are elongated, and tapering towards either end; in the “coronated brand” the apex is crowned with spicular processes; in the “wind-flower brand” the entire spores are echinulate; in the “mint brand” they are globose; in the “composite brand” elliptic; in the “earth-nut brand,” nearly cut in two at the septum; and in the “dandelion brand,” so variable in form that no two are precisely alike. On the other hand, all are characterized by a transverse septum dividing each spore into two cells.