Mycological Society of America Agaricicolous Species of Hypomyces Author(s): Clark T. Rogerson and Gary J. Samuels Source: Mycologia, Vol. 86, No. 6 (Nov. - Dec., 1994), pp. 839-866 Published by: Mycological Society of America Stable URL: http://www.jstor.org/stable/3760597 Accessed: 15-12-2015 18:49 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Mycological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Mycologia. http://www.jstor.org This content downloaded from 147.8.31.43 on Tue, 15 Dec 2015 18:49:01 UTC All use subject to JSTOR Terms and Conditions Mycologia, 86(6), 1994, pp. 839-866. ? 1994, by The New York Botanical Garden, Bronx, NY 10458-5126 of Agaricicolous species Hypomyces Clark T. Rogerson phoraceous hosts are less restricted. The Russulaceae The New York Botanical Garden, Bronx, New York J. P. Lotsy is the host family of preference for the 10458-5126 agaricicolous Hypomyces species, with teleomorphs of eight species found only on species of Lactarius or GaryJ. Samuels Russula. Hypomyces hyalinus is found only on Amanita USDA-ARS, Systematic Botany and Mycology Lab., Rm. Two are on viz. 304, B-011A, BARC-W, Beltsville, Maryland 20705- species. species brown-spored agarics, 2350 H. succineus on Pholiota sp., and H. tremellicola on Cre- pidotus spp.; and H. porphyreus is on the pink-spored Leptonia strigosissima. Hypomyces odoratus is the least Abstract: Thirteen species of Hypomyces occur on gilled specialized ofthe agaricicolous species. Its teleomorph fungi. Most are found on members ofthe Russulaceae; was previously known only in cultures of paired Eu? other hosts include Amanita spp., Crepidotus spp., Lep- ropean isolates of its anamorph, Cladobotryum myco- tonia strigosissima, and Pholiota sp. Anamorphs have philum, that were isolated from a variety of fungi. We been proven only for the four species, H. armeniacus have found perithecia in Puerto Rico and the United (Cladobotryum verticillatum), H. odoratus (C. mycophil? States on a polyporaceous basidiomycete. Hypomyces um), H. succineus (Verticillium succineum comb. nov.), armeniacus, better known in Europe as H. ochraceous and H. tremellicola (Verticillium sp.). Anamorphs have (but see below), begins developing on basidiomata of been putatively linked to H. lateritius (C. tulasnei), H. Russula or Lactarius, completely destroys the host and lithuanicus (C. arnoldii), andH. petchii (Verticillium sp.). grows away from it, then appears to be growing on Key Words: Agaricales, Cladobotryum, Hypocrea- the substratum of the actual host. Hypomyces lateritius ceae, Hypomyces, systematics, Verticillium and H. lithuanicus appear restricted to species of Lac? tarius. Hypomyces lithuanicus may be the most special? ized of the parasites, occurring only on species of the INTRODUCTION Lactarius torminosus (Schaeff.: Fr.) S. F. Gray complex, while other species may be restricted to either Lactarius This is the fourth in a series of publications of mono- or Russula but not to any species complex. In large graphic studies of Hypomyces (Fr.) L.-R. Tulasne (Hy? populations of Hypomyces lactifluorum where the host In earlier papers we have described the pocreales). can be determined with some confidence, the host has species that occur on discomycetes (Rogerson and proven to be Russula brevipes; but associated nonpar- Samuels, 1985), on boletes (Rogerson and Samuels, asitized hosts often belong to the Lactarius piperatus 1989), and on Aphyllophorales (Rogerson and Sam? complex. In our experience Hypomyces luteovirens oc? uels, 1993). As in these papers, we continue to place curs only on species of Russula. Peckiella (Sacc.) Sacc. and Apiocrea H. Sydow in syn? Basidiomata of agaric species that have been trans- onymy with Hypomyces. formed by their parasites, including H. hyalinus, H. Species of Hypomyces can be distributed along the lactifluorum, and H. macrosporus, are considered by some lines of their substrata, with most found on boletes, to be edible (Herrera and Guzman, 1961; Lincoff, 1981), agarics, or Aphyllophorales (including polypores and bearing in mind the possibility that the host itself may thelephores) and with little overlap among the broad be poisonous. Hypomyces lactifluorum and H. macros? substratum groups. Thirteen species are recognized porus are available in Mexican markets, where they may as occurring primarily on members of the Agaricales, be found mixed (Herrera and Guzman, 1961). including H lactifluorum, the type species ofthe genus. Species of Hypomyces that parasitize boletes (Rogerson and and Samuels, 1989; Helfer, 1991) agarics (Helfer, TELEOMORPH CHARACTERS 1991; and herewith) tend to be specific to host family All here have the essential features or genus, whereas most species found on aphyllo- species reported of Hypomyces, the Hypocreaceae and the Hypocreales (Barr, 1990; Hanlin, 1963,1964; Rogerson, 1970; Rog? Accepted for publication July 27, 1994. erson and Samuels, 1989; Samuels, 1973). Hypomyces is 839 This content downloaded from 147.8.31.43 on Tue, 15 Dec 2015 18:49:01 UTC All use subject to JSTOR Terms and Conditions 840 Mycologia most easily recognized by its basidiomycetous habitat, tegrity and the subiculum assumes a pseudoparenchy- fusiform, apiculate ascospores, and anamorphs. Per? matous, stromatic aspect. ithecia typically form in a light or brightly colored Among the agaricicolous species perithecial ontog? subiculum on the hymenium of agarics, boletes, or the eny has been described for H. lactifluorum (Hanlin, Aphyllophorales. 1963) and is consistent for the genus (see references The verrucose and apiculate ascospores (Figs. 1- in Rogerson and Samuels, 1993). There are variations 13) are unusual among the pyrenomycetes and are in the anatomy of the perithecial apex of the agari? thus highly diagnostic for Hypomyces. Four agaricico- cicolous species. In Hypomyces in general the perithe? lous species have unicellular ascospores (H. bannin- cial apex is formed of globose to somewhat elongated giae, H. lateritius, H. luteovirens, H. lithuanicus), and cells that are arranged in files (e.g., H. macrosporus, the ascospores of H. hyalinus are apiosporous with the Fig. 72; H. petchii, Fig. 98), the terminal cells of which septum in the lower third ofthe ascospore. Ascospores are more or less conspicuously clavate; the perithecial of H. banningiae, H. porphyreus, and H. tremellicola lack apex is not confluent with the surrounding subiculum. a conspicuous apiculus; rather their ends have a blunt However, in the perithecial apex of H. lateritius (Figs. appendage that is probably homologous with the apic? 48, 49), H. lithuanicus (Figs. 57, 58), and//. luteovirens ulus of other species. Ascospores of H. succineus are (Fig. 63) the arrangement of papillar cells into files is merely acute although the spines may appear as apiculi not obvious, and hyphae merge the papilla with the when in a terminal position. Ascospores of H. tremel? subiculum. Cells of the papilla of H. luteovirens are licola are also spinulose rather than warted and are arranged in files that separate from each other. neither apiculate nor acute. While the of are in agaricicolous species Hypomyces ANAMORPH CHARACTERS general morphologically, anatomically, and biologi- cally homogeneous, there are variations. Reaction of Only four of the agaricicolous species of Hypomyces subiculum and perithecia with 3% (aq.) potassium hy- have been linked unequivocally through ascospore cul? droxide (KOH) is characteristic of a species. In some tures to anamorphs, viz. H. aremniacus (Figs. 20-29), a pronounced violet to purple color forms (KOH+). H. odoratus (Figs. 76-83), H. succineus (Figs. 118- The subiculum and perithecial apex of H. lithuanicus 121), and H. tremellicola (Figs. 129-131). The ana? become reddish in KOH, but the color reaction could morphs of H. armeniacus and H. odoratus can be clas? be overlooked easily. In others no color change is sified in Cladobotryum sensu Rogerson and Samuels discernible (KOH?); but the natural color of their (1993). The conidiogenous cells of the Cladobotryum perithecia may intensify or fade. anamorphs lack periclinal thickening at the conidi? The subiculum of most species is initially of cottony ogenous locus suggesting holoblastic development; co? hyphae and remains cottony throughout development. nidia are held singly or in groups of a few at the tip Hyphae at the surface of the subiculum of H. luteo? of the condiogenous cell but are not aggregated in a virens, H. lateritius (Fig. 50), H. lithuanicus (FiG. 59), drop of liquid. The anamorphs of H. succineus and H. and H banningiae have many free ends; this feature tremellicola are Verticillium species. These Verticillium is conspicuous in microtome sections but also is easily anamorphs produce phialides that have periclinal seen in crush mounts. Cells of these hyphae often swell thickening at the tip, denoting enteroblastic conidial and the hyphae may appear to be somewhat monili? development, and a multiplicity of conidia held in a form. Distinctive moniliform hyphae of the subiculum drop of liquid