COPRINUS Pers.
The species of Coprinus are readily recognised from the black spores in addition to the fact that the gills, at maturity, dissolve into a black or inky fluid. The larger species especially form in this way an abundance of the black fluid, so that it drops from the pileus and blackens the grass, etc., underneath the plant. In some of the smaller species the gills do not wholly deliquesce, but the cap splits on top along the line of the longer gills, this split passing down through the gill, dividing it into two thin laminæ, which, however, remain united at the lower edge. This gives a fluted appearance to the margin of the pileus, which is very thin and membranaceous.
Figure 31.—Coprinus comatus, "shaggy-mane," in lawn.
The plants vary in size, from tiny ones to those which are several inches high and more than an inch broad. Their habitat (that is, the place where they grow) is peculiar. A number of the species grow on dung or recently manured ground. From this peculiarity the genus received the name Coprinus from the Greek word κοπρὁς, meaning dung. Some of the species, however, grow on decaying logs, on the ground, on leaves, etc.
Coprinus comatus Fr. Edible.—One of the finest species in this genus is the shaggy-mane, or horse-tail mushroom, as it is popularly called. It occurs in lawns and other grassy places, especially in richly manured ground. The plants sometimes occur singly, or a few together, but often quite large numbers of them appear in a small area. They occur most abundantly during quite wet weather, or after heavy rains, in late spring or during the autumn, and also in the summer. From the rapid growth of many of the mushrooms we are apt to be taken by surprise to see them all up some day, when the day before there were none. The shaggy-mane often furnishes a surprise of this kind. In our lawns we are accustomed to a pretty bit of greensward with clumps of shrubbery, and here and there the overhanging branches of some shade tree. On some fine morning when we find a whole flock of these shaggy-manes, which have sprung up during the night, we can imagine that some such kind of a surprise must have come to Browning when he wrote these words:
"By the rose flesh mushroom undivulged
Last evening. Nay, in to-day's first dew
Yon sudden coral nipple bulged,
Where a freaked, fawn colored, flaky crew
Of toadstools peep indulged."
Figure 32.—Coprinus comatus. "Buttons," some in section showing gill slits and hollow stem; colors white and black. (Natural size.)
The plant is called shaggy-mane because of the very shaggy appearance of the cap, due to the surface being torn up into long locks. The illustrations of the shaggy mane shown here represent the different stages of development, and the account here given is largely taken from the account written by me in Bulletin 168 of the Cornell University Agr. Exp. Station.
Figure 33.—Coprinus comatus (natural size).]
In Fig. [32] are shown two buttons of the size when they are just ready to break through the soil. They appear mottled with dark and white, for the outer layer of fungus threads, which are dark brown, is torn and separated into patches or scales, showing between the delicate meshes of white threads which lie beneath. The upper part of the button is already forming the cap, and the slight constriction about midway shows the lower boundary or margin of the pileus where it is still connected with the undeveloped stem.
At the right of each of these buttons in the figure is shown a section of a plant of the same age. Here the parts of the plant, though still undeveloped, are quite well marked out. Just underneath the pileus layer are the gills. In the section one gill is exposed to view on either side. In the section of the larger button the free edge of the gill is still closely applied to the stem, while in the small one the gills are separated a short distance from the stems showing "gill slits." Here, too, the connection of the margin of the pileus with the stem is still shown, and forms the veil. This kind of a veil is a marginal veil.
Figure 34.—Coprinus comatus (natural size). This one entirely white, none of the scales black tipped.
The stem is hollow even at this young stage, and a slender cord of mycelium extends down the center of the tube thus formed, as is shown in the sections.
The plants are nearly all white when full grown. The brown scales, so close together on the buttons, are widely separated except at the top or center of the pileus, where they remain close together and form a broad cap.
A study of the different stages, which appear from the button stage to the mature plant, reveals the cause of this change in color and the wide separation of the dark brown scales. The threads of the outer layer of the pileus, and especially those in the brown patches seen on the buttons, soon cease to grow, though they are firmly entangled with the inner layers. Now the threads underneath and all through the plant, in the gills and in the upper part of the stem, grow and elongate rapidly. This pulls on the outer layer, tearing it in the first place into small patches, and causing them later to be more widely separated on the mature plant. Some of these scales remain quite large, while others are torn up into quite small tufts.
Figure 35.—Coprinus comatus, sections of the plants in Fig. [33] (natural size).
As the plant ages, the next inner layers of the pileus grow less rapidly, so that the white layer beneath the brown is torn up into an intricate tangle of locks and tufts, or is frazzled into a delicate pile which exists here and there between well formed tufts. While all present the same general characters there is considerable individual variation, as one can see by comparing a number of different plants. Figure [34] shows one of the interesting conditions. There is little of the brown color, and the outer portion of the pileus is torn into long locks, quite evenly distributed and curled up at the ends in an interesting fashion which merits well the term "shaggy." In others the threads are looped up quite regularly into triangular tresses which appear to be knotted at the ends where the tangle of brown threads holds them together.
Figure 36.—Coprinus comatus, early stages of deliquescence; the ring is lying on the sod (natural size).
There is one curious feature about the expansion of the pileus of the shaggy-mane which could not escape our attention. The pileus has become very long while comparatively little lateral expansion has taken place. The pileus has remained cylindrical or barrel-shaped, while in the case of the common mushroom the pileus expands into the form of an umbrella.
Figure 37.—Coprinus comatus, later stage of deliquescence, pileus becoming more expanded (natural size).
The cylindrical or barrel-shaped pileus is characteristic of the shaggy-mane mushroom. As the pileus elongates the stem does also, but more rapidly. This tears apart the connection of the margin of the pileus with the base of the stem, as is plainly shown in Fig. [33]. In breaking away, the connecting portion or veil is freed both from the stem and from the margin of the pileus, and is left as a free, or loose, ring around the stem. In the shaggy-mane the veil does not form a thin, expanded curtain. It is really an annular outer layer of the button lying between the margin of the cap and the base of the stem. It becomes free from the stem. As the stem elongates more rapidly than the cap, the latter is lifted up away from the base of the stem. Sometimes the free ring is left as a collar around the base of the stem, still loosely adherent to the superficial layer of the same, or it remains for a time more or less adherent to the margin of the pileus as shown in the plant at the left hand in Fig. [33]. It is often lifted higher up on the stem before it becomes free from the cap, and is then left dangling somewhere on the stem, or it may break and fall down on the sod. In other instances it may remain quite firmly adherent to the margin of the pileus so that it breaks apart as the pileus in age expands somewhat. In such cases one often searches for some time to discover it clinging as a sterile margin of the cap. It is interesting to observe a section of the plants at this stage. These sections can be made by splitting the pileus and stem lengthwise through the middle line with a sharp knife, as shown in Fig. [35]. Here, in the plant at the right hand, the "cord" of mycelium is plainly seen running through the hollow stem. The gills form a large portion of the plant, for they are very broad and lie closely packed side by side. They are nowhere attached to the stem, but at the upper end round off to the cap, leaving a well defined space between their ends and the stem. The cap, while it is rather thick at the center, i. e., where it joins the stem, becomes comparatively thin where it spreads out over the gills. At this age of the plant the gills are of a rich salmon color, i. e., before the spores are ripe, and the taste when raw is a pleasant nutty flavor, reminding one of the meat of fresh green hickory nuts. In a somewhat earlier stage the edges of all the gills are closely applied to the stem which they surround. So closely are they applied to the stem in most cases that threads of mycelium pass from the stem to the edge of the gills. As the cap expands slightly in ageing, these threads are torn asunder and the stem is covered with a very delicate down or with flocculent particles which easily disappear on handling or by the washing of the rains. The edges of the gills are also left in a frazzled condition, as one can see by examining them with a good hand lens.
The spores now begin to ripen and as they become black the color of the gills changes. At the same time the gills and the cap begin to dissolve into an inky fluid, first becoming dark and then melting into a black liquid. As this accumulates it forms into drops which dangle from the cap until they fall away. This change takes place on the margin of the cap first, and advances toward the center, and the contrast of color, as the blackening invades the rich salmon, is very striking. The cap now begins to expand outward more, so that it becomes somewhat umbrella shaped. The extreme outer surface does not dissolve so freely, and the thin remnant curls upward and becomes enrolled on the upper side as the cap with wasted gills becomes nearly flat.
Coprinus atramentarius (Bull.) Fr. Edible.—The ink-cap (Coprinus atramentarius) occurs under much the same conditions as the shaggy-mane, and is sometimes found accompanying it. It is usually more common and more abundant. It springs up in old or newly made lawns which have been richly manured, or it occurs in other grassy places. Sometimes the plants are scattered, sometimes two or three in a cluster, but usually large clusters are formed where ten to twenty or more are crowded closely together (Fig. [39]). The stems are shorter than those of the shaggy-mane and the cap is different in shape and color. The cap is egg-shaped or oval. It varies in color from a silvery grey, in some forms, to a dark ashen grey, or smoky brown color in others. Sometimes the cap is entirely smooth, as I have seen it in some of the silvery grey forms, where the delicate fibres coursing down in lines on the outer surface cast a beautiful silvery sheen in the light. Other forms present numerous small scales on the top or center of the cap which are formed by the cleavage of the outer surface here into large numbers of pointed tufts. In others, the delicate tufts cover more or less the entire surface, giving the plant a coarsely granular aspect. This is perhaps the more common appearance, at least so far as my observation goes. But not infrequently one finds forms which have the entire outer surface of the cap torn into quite a large number of coarse scales, and these are often more prominent over the upper portion. Fine lines or striations mark also the entire surface of all the forms, especially toward the margin, where the scales are not so prominent. The marginal half of the cap is also frequently furrowed more or less irregularly, and this forms a crenate or uneven edge.
Plate 9, Figure 38. Coprinus comatus, drops of inky fluid about to fall from wasted pileus (natural size).
Plate 10, Figure 39.—Coprinus atramentarius, nearly smooth form, gray color (natural size)
Figure 40.—Coprinus atramentarius, scaly form (natural size).
The annulus or ring on the stem of the ink-cap is very different from that of the shaggy-mane. It forms an irregularly zigzag elevated line of threads which extend around the stem near the base. It is well shown in Fig. [41] as a border line between the lower scaly end of the stem and the smooth white upper part. It is formed at the time of the separation of the margin of the cap from the stem, the connecting fibres being pulled outward and left to mark the line of junction, while others below give the scaly appearance. It is easily effaced by rough handling or by the washing of the rains. A section of a plant is illustrated by a photograph in Fig. [42]. On either side of the stem is shown the layer of fibres which form the annulus, and this layer is of a different texture from that of the stem. The stem is hollow as seen here also. In this figure one can see the change in color of the gills just at the time when they begin to deliquesce. This deliquescence proceeds much in the same way as in the shaggy-mane, and sometimes the thin remnant of the cap expands and the margin is enrolled over the top.
Figure 41.—Coprinus atramentarius, showing annulus as border line between scaly and smooth part of the stem (natural size).
Coprinus micaceus (Bull.) Fr. Edible.—The glistening coprinus received its name because of the very delicate scales which often cover the surface of the cap, and glisten in the light like particles of mica. This plant is very common during the spring and early summer, though it does appear during the autumn. It occurs about the bases of stumps or trees or in grassy or denuded places, from dead roots, etc., buried in the soil. It occurs in dense tufts of ten to thirty or more individuals; sometimes as many as several hundred spring up from the roots of a dead tree or stump along the streets or in lawns, forming large masses. More rarely it occurs on logs in the woods, and sometimes the plants are scattered in lawns. From the different habits of the plant it is sometimes difficult to determine, especially where the individuals are more or less scattered. However, the color, and the markings on the cap, especially the presence of the small shining scales when not effaced, characterize the plant so that little difficulty is experienced in determining it when one has once carefully noted these peculiarities.
Figure 42.—Coprinus atramentarius, section of one of the plants in Fig. [41] (natural size).
Figure [43] is from a group of three young individuals photographed just as the margin of the pileus is breaking away from the lower part of the stem, showing the delicate fibrous ring which is formed in the same way as in Coprinus atramentarius. The ring is much more delicate and is rarely seen except in very young specimens which are carefully collected and which have not been washed by rains. The mature plants are 8–10 cm. high (3–4 inches), and the cap varies from 2–4 cm. in diameter. The stem is quite slender and the cap and gills quite thin as compared with the shaggy-mane and ink-cap. The gills are not nearly so crowded as they are in the two other species. The cap is tan color, or light buff, or yellowish brown. Except near the center it is marked with quite prominent striations which radiate to the margin. These striations are minute furrows or depressed lines, and form one of the characters of the species, being much more prominent than on the cap of the ink-cap.
Figure 43.—Coprinus micaceus, young stage showing annulus, on the cap the "mica" particles (natural size).
Figure 44.—Coprinus micaceus, plants natural size, from floor of coal mine at Wilkesbarre. Caps tan color. Copyright.
In wet weather this coprinus melts down into an inky fluid also, but in quite dry weather it remains more or less firm, and sometimes it does not deliquesce at all, but dries with all parts well preserved, though much shrunken of course, as is the case with all the very fleshy fungi.
Plate 11, Figure 45.—Panæolus retirugis, group of plants from lawn along street, showing veil in young plants at the left, which breaks into V-shaped loops and clings to margin of the cap. Cap dark smoky color at first, becoming grayish in age (natural size). Copyright.
PANAEOLUS Fr.
In Panæolus, the pileus is somewhat fleshy, or thin, the margin even, that is, not striate. The margin extends beyond the gills, and the gills are not uniform in color, being clouded or spotted with black and brown colors, the edge of the gills often white in contrast. The spores are black. The stem is usually smooth, sometimes floccose scaly, often long, firm, generally hollow. The veil is of interwoven threads, sometimes quite compact, especially when the plants are young. Peck, 23rd Report N. Y. State Mus., p. 10 et seq., gives a synopsis of five species.
Figure 46.—Panæolus retirugis, section of caps showing form and position of gills (natural size).
Panæolus retirugis Fr.—The color of this plant is not attractive, but it is one of the most beautiful species I have studied, if one regards form and the general features in its development. It is said to occur on dung. I have found it in lawns or grassy places, especially freshly made lawns or greenswards which have been heavily manured. The illustrations in Figs. 45–48 were made from photographs of plants which grew in a newly made boulevard along Buffalo street, Ithaca, N. Y. (No. 2356 C. U. herbarium). The plants are from 7–15 cm. high, the cap from 1–3 cm. in diameter, and the stem is 3–4 mm. in thickness. The size of the plants varies greatly according to the environment, being larger in moist soil and wet weather and smaller in dry soil and dry weather. It occurs in late spring and during the summer.
Figure 47.—Panæolus retirugis, showing rugose character of cap in left-hand plant (natural size). Copyright.
The pileus is oval to ovate and conic, and in some cases it becomes more or less expanded, but never, so far as I have observed, does it become depressed or even plane. In wet weather it is usually at first dark smoky in color, viscid, becoming grayish in age, and as the pileus dries it becomes shining. In lighter colored forms the pileus is at first light leather color to cream color. Toward the center of the pileus are irregular wrinkles or shallow pits, the wrinkles anastomosing more or less, and it is because of this character of the surface of the pileus that the plant receives its specific name. During dry weather there is a tendency for the pileus to crack, separating the dark color of the surface into patches showing the white flesh beneath. The pileus is often umbonate or gibbous, and the center is often darker than the margin. The pileus in rare cases is entirely white. The gills are adnate, broad in the middle, and in the more expanded forms as the gills separate more and more from the stem there is a tendency for them to become somewhat triangular. The spores are black in mass, are elliptical or short fusiform, and measure from 10–12 × 15–18 µ. The stem is cylindrical, sometimes tortuous, smoky gray, light reddish brown, or paler, sometimes entirely white, the lighter forms of the stem accompanying the light forms of the pileus; cartilaginous in texture, becoming hollow, always darker below and paler above, smooth, granulate with minute darker points, bulbous. The veil is very prominent and stout when the plant is young, and extends from the margin of the pileus to the stem when the plant is very young and the stem has not elongated. As the stipe elongates the veil separates from the stipe as a ring, and then, as the pileus expands, it is broken quite regularly into short segments which become arranged regularly around the margin of the pileus in the form of the letter V, which gives a beautiful appearance to this stage of the plant. It is only when the plants are fresh and moist that this condition of the veil can be seen, for on drying the veil collapses. Water is sometimes caught under the veil before the pileus separates far from the stem, and the spores falling thus float against the stem at this point and make a dark ring around the stem, which, however, should not be mistaken for the annulus. In no case was the veil observed to cling to the stem, and many plants have been observed to see if this variation might present itself.
Figure 48.—Panæolus retirugis, showing cracked surface of cap in the left-hand plant, also in same plant the ring mark of black spores which lodged before veil ruptured, in other plants showing well the V-shaped loops of veil on margin of cap (natural size). Copyright.
This peculiarity of the veil in clinging to the margin of the pileus has led Hennings to place the plant in Karsten's genus (Engler and Prantl, Pflanzenfamilien) Chalymotta, as Chalymotta retirugis. The plants have several times been eaten raw by me, and while they have a nutty flavor and odor, the taste is not entirely agreeable in this condition, because of the accompanying slimy sensation.
A number of smaller species, among them P. fimicola Fr., and P. papilionaceus Fr., occur in similar places. Panæolus solidipes Pk., is a large species with a long, solid stem, growing on dung. Psilocybe fœnisecii, abundant in lawns and grassy places during late spring and summer, resembles a Panæolus. The cap shows zones of light and dark color, due to different amounts of water, which disappear as the plant matures. It belongs to the purple-brown-spored agarics.