Coker WC, Couch JN. 1928. The Gasteromycetes of the Eastern United States and Canada. The University of North Carolina Press, Chapel Hill.
Lycoperdon marginatum Vitt.
L. papillatum Schaeff . in sense of Hollos L. cruciatum Rostk. L. separans Peck L. caivescens B. & C.
Plates 55, 56 and 113
Plants 1.2-4.5 cm. broad and nearly always broader than tall; narrowing below to a short stem and root. Cortex pure white, then a little discolored or pinkish, becoming clay colored or deep brown in age, composed of densely set, sharp, erect warts which are short, thick and simple or more slender and fascicled with their tips cohering, at maturity breaking up into large or small flakes (see below), usually beginning in a median line between the stem and top, but often rupturing at every point or at many points at once (see below for woods form). The lower third, covering the stem and lower part of the body, does not fall away. Inner peridium strong to pale olivaceous brown when first exposed, even, or in one of our collections (Warrenton, Va.) obviously pitted, much as in gemmatum. It is peculiar in having beneath the spines a covering of a minute brown furfurescence which disappears rather slowly and leaves the surface paler and shining. This furfurescence is different from that of any other plant we know. It is composed of little flat scales, the prevailing shape of which is spherical to oblong, vary- ing to irregular and angled, the majority elongated and up to 60 x 100u. Subgleba ample, composed of distinct, rather large compartments, occupying the stem and lower part of the body and persistent after the top has disappeared, remaining as a dark brown pedicel with a thin, collapsed margin.
Spores (of No. 482) spherical, smooth, 3.6-4.1 u thick, with one oil drop. The spores have mixed with them about an equal number of small rods like bacilli, that average about twice the length of the spore's width. Basidia (of No. 5225) 6.6-7.4 u thick, usually 4-spored. Capillitium composed of yellowish brown fibers very little branched, about as thick as the spores as a rule, but up to 5 u in diameter, attenuated at the ends.
The spores and capillitium of our plants have been compared with Peck's collections of L. separans and are the same. One of the most characteristic qualities of the plant is its strong odor when ripening or fading like that of horse urine, a distinction it shares with L. gemmatum. It is a very common plant with us in open places and cultivated ground, and less often in woods. The spines are very variable, in some cases very thick and short and not split up into fascicles, in others slender and moderately short or long, and in groups of several with their tips united. In Chapel Hill there are two rather distinct forms; in one (typical) of cultivated or open ground the cortex falls off in larger flakes and more irregularly and the spores have a little mucro left after the pedicel breaks off; in the other one of woods each group of spines with its underlying tissue is apt to fall off singly or with a small group of others, and the denudation is more regular, proceeding from the top downward. In this form the spores have no mucro. The scales on the denuded surface are just like those of the typical form. This character alone is sufficient to distinguish the species. For general discussion of the species see Lloyd (Myc. Notes, pp. 214 and 231).
It seems certain that L. cruciatum is L. marginatum, as Rostkovius's good figure could not be referred to anything else. Spores of three specimens labeled L. cruciatum in Bresadola's herbarium (sent by Romell) are not those of the present species. This is true also of a specimen labeled L. marginatum in the same herbarium. We have seen European material of true L. marginatum, as L. cruciatum (Herb. Hasskarl & Herb. Klotzsch), received from the Ryks Herbarium in Leiden. They show the same spores and peridial scales as in our specimens (spores 3.6-4.5 u). The capillitium is not colorless but of about the same tint as in ours. We have compared capillitium of plants just maturing in our No. 482 with other mature ones of the same collection and find that the capillitium does not change after the spores are mature.
A bit of the inner peridium and spores and capillitium of L. calvescens B. & C. from the Curtis Herbarium (Wright, No. 6366, co-type), kindly sent us by Dr. Thaxter, shows it to be L. marginatum. The peridial scales and the spores are the same as in the typical form of L. marginatum. Lycoperdon pratense Pers. is also considered a synonym by Hollos, but Lloyd thinks that the typical L. pratense does not occur in the United States. A specimen from New Zealand determined as L. pratense by Lloyd does not look like our L. marginatum, as the spines are much shorter and the denuded peridium shows no obvious furfurescence. The spores are slightly smaller and both they and the capillitium are paler than in our plants.
From the description and illustration, L. abscissum R. E. Fries from Bolivia and Argentina is nearest L. marginatum. It is remarkable in the comparatively large size of the subgleba which occupies most of the fruit body.
Hard. Mushrooms, fig. 467. Hollos. 1. c, pl. 20. figs. 17-22; pl. 29, fig. 6 (as L. papillatum). Lloyd. Myc. Works, pl. 51 (as L. cruciatum). 1905. Lloyd. Photogravure of Am. Fungi, No. 3 (as L. separans). Morgan. 1. c, 14: pl. 2. fig. 1. Murrill. Mycologia 1: pl. 15, fig. 7 (as L. Wrightii). 1909. The three plants on right may be true L. Wrightii. Rostkovius in Sturm. Deutsch. Fl. Pilze 1: pl. 8 (as L. cruciatum). 1813. Vittadini. Monog. Lycoperd., pl. 1, fig. 11. 1843.
99. On ground in cemetery, October 27, 1911. Spores spherical, smooth, 3.6-4u thick, with a very distinct oil drop. 482. Glen Burnie meadow and on campus, October 3, 1912. 922. In mixed woods, September 13, 1913. 1762b. In pine woods, September 12, 1915. 5225. In a lowground pasture, June 23, 1922. 7194. In mixed woods, October 14, 1923. Spores 3.7-4.2u thick, with pedicels entirely broken off.
Haywood County. In mold in semi-open place by Pigeon River, Aug. 9, 1926. Coker, coll. (U. N.C. Herb., No. 8131).
South Carolina. Ravenel. Fungi Car. Exs. No. 73. (Phil. Acad. Herb, and N. Y. B. G. Herb., as L. gemmaatum).
Florida. Couch, coll. (U. N. C. Herb., Nos. 7266-7272).
Alabama. Auburn. Earle, coll. (N. Y. B. G. Herb.).
Virginia. Mountain Lake. In moist woods. Murrill, coll. (N. Y. B. G. Herb.). This is exactly like our woods form No. 7194. Warrenton. Coker, coll. (U. N. C. Herb.). Spores faintly rough, 3. 5-4.2u. Woods form.
District of Columbia. Cook, coll. (U. S. Nat'l. Herb., as L. cruciatum).
New Jersey. Newfield. Ellis, coll. (N. Y. B. G. Herb., as L. Wrightii and as L. pedicellatum).
New York. Sand Lake. (N. Y. B. G. Herb., as L. separans). Newcomb. House, coll. (Herb. N. Y. St. Mus. and U. N. C. Herb.). Spores smooth, 3.7-4.2u. Scales on inner peridium as usual.
Connecticut. Miss White, coll. (N. Y. B. G. Herb.).
Ohio. Lloyd, coll. No. 2655. (Bres. Herb., as L. separans, and U. N. C. Herb.). Cleveland. Beardslee, coll. (Bres. Herb., as L. separans, and U. N. C. Herb.). Spores 3.6-4.2u.
Wisconsin. La Crosse. (Univ. Wis. Herb, and U. N. C. Herb.). Spores smooth, 3.5-4u, sometimes with a mucro.