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New Records of Gasteroid and Secotioid Fungi from Sand Dunes in Northwestern South Dakota

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New Records of Gasteroid and Secotioid Fungi from Sand Dunes in Northwestern South Dakota Proceedings of the South Dakota Academy of Science, Vol. 90 (2011) 125 NEW RECORDS OF GASTEROID AND SECOTIOID FUNGI FROM SAND DUNES IN NORTHWESTERN SOUTH DAKOTA A. C. Gabel and M. L. Gabel* Biology Department Black Hills State University 1200 University Spearfish, South Dakota 57799 *Corresponding author email: [email protected] ABSTRACT Sand dunes in northwestern South Dakota were surveyed for fungi from 2002–2006 in Harding County and from 2003–2005 in Perkins County. In total, eight taxa of gasteroid fungi, Calvatia cyathiformis, C. fragilis, Disciseda bovista, Geastrum campestre, G. pectinatum, G. xerophilum, Tulostoma brumale, and T. fimbriatum var. campestre, and one secotioid fungal species, Montagnea arenaria, were recovered. Calvatia fragilis, Geastrum xerophilum, Montagnea are- naria, and Tulostoma brumale are new records for South Dakota, and M. arenaria is reported east of the Rocky Mountains for the first time. By contributing to the inventory of fungal biota in South Dakota and extending the known range for the taxa reported here, we highlight the importance of basic surveys of fungal diversity in the United States. Keywords distribution, gasteroid and secotioid fungi, northwestern South Dakota, sand dunes INTRODUCTION Fungal organisms with fruiting bodies consisting of a mass of spores (glebum) enclosed by an outer layer (peridium) have been termed gasteroid fungi. Once considered a growth form of a single class of fungi, the Gasteromycetes, it is now known that gasteroid fungi in the Basidiomycota have evolved several times from their agaricoid (mushroom) ancestors (Hibbett et al. 1997). Intermediate between the agaricoid and gasteroid forms are the secotioid fungi that resemble mushrooms with unexpanded caps and poorly developed gills (Hopple and Vilgalys 1999). Gasteroid and secotioid fungi can be found in a wide range of habitats, from high altitude forests to sea shores, and are particularly well repre- sented in arid areas throughout the world (Coker and Couch 1928; Bates 2004; Bates 2006; Bates et al. 2009). 126 Proceedings of the South Dakota Academy of Science, Vol. 90 (2011) Orders of epigeous (emerging above ground) gasteroid fungi include the Lyco- perdales and Tulostomatales (Alexopoulos et al.1996; Kendrick 2000). Species in the Lycoperdales are known to inhabit xeric pastures, prairies, and deserts in North America (Coker and Couch 1928; Smith 1951; Miller and Miller 1988; Bates et al. 2009). The gasteroid order Tulostomatales includes species that are cosmopolitan in arid areas such as desert habitats and sand dunes, which occur primarily in the western and mid-continental United States (Wright 1987). Fruitings of saprophytic, secotioid fungi, such as Podaxis pistillaris and Agaricus texensis (= Longula texensis), are frequently reported from sandy, arid areas in- cluding the southwestern and western United States and Mexico (Zeller 1943; Bowerman and Groves 1962; Miller 1983; Miller and Miller 1988). Sand dune areas in northwestern South Dakota have not been well surveyed for fungi and the only gasteroid species reported previously from the area (Hard- ing County) were Disciseda subterranea (Lycoperdaceae), Geastrum drummondii (Geastraceae), and Tulostoma campestre (Tulostomataceae) (Gabel et al. 2004). There are no reports of secotioid fungi, however, from this area. The objective of this study was to collect and document more thoroughly gasteroid and secotioid fungi that occur in sand dunes of northwestern South Dakota. The results of our survey are reported from frequent visits to a sand dune complex in Hard- ing county from 2002–2006, and to a sand dune area in Perkins County from 2003–2005. METHODS Collecting sites were in Harding and Perkins Counties. The Harding County site was on the Hunsucker Ranch, T18N, R5E, in the north central portion of Section 3, latitude N45o 33.439’, longitude W103o 29.630’, ranging in altitude from 875 to 900 m. The Perkins County site was on the McKenna Ranch, T14 N, R10E, SW 1/4 of Section 24, latitude N45o 9.349’, longitude W102o 51.007’, at an altitude of approximately 800 m. Average annual precipitation recorded from 1948–2005 at the Antelope Station, the nearest recording station to Hunsucker Ranch, was 37.6 cm. The Perkins County (Zeona) Station, the nearest to McKenna Ranch, recorded an average annual precipitation of 38.4 cm from 1949–2002. Average wind velocities for these same time periods were 19.0 kph and 20.1 kph, for the Antelope and Zeona stations respectively. The sandy prairies of these areas included swales (depressions), dunes (elevated areas), and blowouts. Gabel and Simonson (2008) found vegetation in swales and dunes to be more dense and more diverse (an average of 46 species and 43 species per transect, respectively) than in blowouts (averaging at 18 species per transect). The same study found vegetation of these areas to be dominated by native species, which represented 86–100% of all species reported from the transects, and included forbs, sedges, and grasses. Common forbs of the area include Eriogonum annuum Nutt., Ladeania lanceolata (Pursh) Egan & Reveal (= Psoralidium lanceolatum), and Solidago missouriensis Nutt., as well as species of Ambrosia and Penstemon. Thirty percent of species reported by Gabel and Simonson (2008) were grasses, such as Andropogon hallii Hack., Bouteloua graci- Proceedings of the South Dakota Academy of Science, Vol. 90 (2011) 127 lis (Willd. ex Kunth) Lag. ex Griffiths, Calamovilfa longifolia (Hook.) Scribn., Hesperostipa comata (Trin. & Rupr.) Barkw., and Sporobolus cryptandrus (Torr.) A. Gray. Several fungi were collected from transects established for the floristic study of Gabel and Simonson (2008) and others were collected outside transects, result- ing in a repetition of collecting dates for some specimens. Collected specimens were brought to the laboratory and identified using standard microscopic proce- dures. As the identification of the Tulostoma species typically requires detailed ultrastructural information, spore and peristomes of collections from this genus were examined under scanning electron microscopy (SEM) using a JEOL 5600 LV. Date, site, location (latitude/longitude), collection number, habitat, and col- lector were recorded for each specimen, which are vouchered at the Black Hills State University Herbarium (BHSC), Spearfish, South Dakota. RESULTS Montagnea arenaria, a secotioid fungus, is newly reported from South Dakota, being the first published record of the species east of the Rocky Mountains. The nearest published records for M. arenaria are from southwest Wyoming and western Montana, approximately 600–650 km distant (Chen 1999). Calvatia fragilis, Geastrum xerophilum, and Tulostoma brumale are likewise new records for South Dakota. A detailed account of the specimens collected is given in Table 1. The following species are reported from Hunsucker Ranch sand dune area of Harding County: Calvatia cyathiformis, C. fragilis, Disciseda bovista (Fig. 1), Geastrum campestre (Fig. 2), G. pectinatum, G. xerophilum, Montagnea arenaria (Figs. 3–4), Tulostoma brumale (Figs. 5–7), and T. fimbriatum var. campestre (Figs. 8–10). The following species are reported from McKenna Ranch sand dune area in Perkins County: C. cyathiformis, C. fragilis, Disciseda bovista, G. pectinatum, G. xerophilum, M. arenaria, T. brumale, and T. fimbriatumvar. camp- estre. Where specimens were collected from transects, all species except one were collected from swales or dunes. A single specimen, D. bovista, was collected from a Harding County blowout. DISCUSSION Because some disagreement exists in the literature over the salient characters that define particular gasteroid species, we have included details on the morphol- ogy of the taxa collected over the course of our survey to clarify the taxonomic concepts used here. Calvatia cyathiformis and C. fragilis are distinguished by the morphology of the sterile portion of tissue below the gleba (termed a subglebum), as well as by their size, shape, and degree of spore ornamentation (Smith 1951; Bowerman and Groves 1962; Kuo 2008; Bates et al. 2009). Specimens in this study were identified as Calvatia fragilis if the gasterocarps were less than 6 × 6 cm, had a 128 Proceedings of the South Dakota Academy of Science, Vol. 90 (2011) tapered base, and the subgleba was absent or less than 1/3 to 1/2 of the gastero- carp height. Geastrum campestre specimens were small earthstars with a sulcate peristome (mouth area) that exhibited thick, fleshy, hygroscopic exoperidial rays (outer layer) that entirely enclosed the asperulate, sessile to nearly sessile endoperidium (spore containing area) when specimens were dry (Smith 1951; Cunningham 1979; Smith et al. 1981; Pegler et al. 1995). Geastrum pectinatum specimens exhibited a lead-purple endoperidium, a narrow, striate pedicel, and a sulcate peristome (Smith 1951; Cunningham 1979; Smith et al. 1981; Pegler et al. 1995; Miller and Miller 2006). Specimens of G. xerophilum had a sulcate peri- stome and a saccate endoperidium that is attached to a pedicel and surrounded by involute exoperidial rays (Ponce de Leon 1968). Scanning electron microscopy of spores from five specimens of each of the two Tulostoma species collected all exhibited distinct verrucae that coalesced into ridges or crests of different lengths (Figs. 7 and 10); therefore, other characteris- tic were important in distinguishing the species. Wright (1955) cites peristome morphology as an important character used to delimit species in the genus Tu- lostoma.
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