A Survey of Myxomycetes from the Black Hills of South Dakota and the Bear Lodge Mountains of Wyoming
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Proceedings of the South Dakota Academy of Science, Vol. 89 (2010) 45 A SURVEY OF MYXOMYCETES FROM THE BLACK HILLS OF SOUTH DAKOTA AND THE BEAR LODGE MOUNTAINS OF WYOMING A. Gabel1*, E. Ebbert2, M. Gabel1 and L. Zierer2 1Biology Department Black Hills State University 1200 University Spearfish, South Dakota 57799 2Deceased *[email protected] ABSTRACT Fruiting bodies of slime molds were collected intermittently from the field from 1990-2009. During the summers of 2007-2009 slime molds also were cultured in moist chambers from downed, decaying wood, dried deciduous leaf litter, and dried grass litter that was gathered three times each summer from four areas at each of the following six sites. Alabaugh Canyon and Boles/Redbird Canyons were dry, prairie/woodlands in Fall River and Custer Counties in the southern Black Hills. Pony Gulch and Spring Creek, in the central hills were moist, broad gulches in Pennington County, and the northern hills sites were Botany Bay and Grigg’s Gulch, narrow, moist canyons in Lawrence County. Contents of a total of 648 cultures were examined. Presence of a species in a culture was considered to be a collection and some cultures contained more than one species. Eighty-two species from both field and culture surveys were identified, and 62 are considered as new records for the Black Hills, and 61 for western South Dakota. Six hundred fifty-seven collections representing 65 spe- cies developed in culture included species from all orders of Myxomycetes. Only five species occurred at a relative abundance > 5% and 20 occurred only once. Four hundred eighty-three collections representing 54 species occurred on wood, 115 collections representing 29 species occurred on leaf litter, and 59 collections representing 10 species were on grass litter. Several species occurred on more than one substratum. The highest number, 178 collections and 39 species, oc- curred on substrata from the Boles/Redbird Canyon site, the southwest dry, open prairie/woodland. The lowest number, 74 collections and 19 species, occurred on substrata from Grigg’s Gulch, a northern, moist site with a high canopy cover. Fuligo megaspora was frequently collected from southern, dry sites. Keywords Black Hills, Fuligo megaspora, Myxomycetes, South Dakota 46 Proceedings of the South Dakota Academy of Science, Vol. 89 (2010) INTRODUCTION Plasmodial slime molds (Myxomycetes) are unique organisms with two distinctly different growth forms. The free-living, multinucleate plasmodium resembles an amoeboid-organism which feeds on bacteria and other microor- ganisms. When environmental conditions are appropriate, a reproductive phase develops and fruiting bodies that resemble fungi are formed which contain spores for dispersal and survival. In addition to spores, microcysts (encysted myxamoebae and swarm cells) and sclerotia (encysted plasmodia) serve as rest- ing stages and are capable of surviving unfavorable environmental conditions. Slime molds occur on a variety of substrata including the bark of living trees and woody vines, decayed logs on the ground, herbaceous plants and grasses, and litter of herbaceous plants and grasses. Herbivore dung and soils also support growth (Kilgore et al. 2009, Keller and Braun 1999, Martin et al. 1983, Martin and Alexopolous 1969). In the past slime molds were classified as fungi and studied by mycologists, but because of the amoeboid stage most taxonomists currently place them with the protists (Keller and Everhart 2010, Kilgore et al. 2009, Spiegel et al. 2004, Ing 1994, Frederick 1990) accommodating both the amoeboid stage and the fruiting bodies. Regardless of their confusing taxonomy, many myxomycete species are dis- tributed worldwide and more are reported as new regions are explored. Martin and Alexopolous (1969) reported species from North America, Central America, South America, Europe and Asia. More recently Chen (1999) reported species from China, Harkonen (1977) from Finland, Lado (1993) from the Mediter- ranean, Rojas and Stephenson (2007) from Costa Rica, Schnittler (2001) from Kazakhstan, Schnittler et al. (2006) from Germany, Stephenson (2009) from Ascension Island, Stephenson and Johnston (2001) from New Zealand, Stephen- son et al. (2000) from Russia, Greenland and Iceland, Stephenson et al. (2004) from Ecuador, and Tran et al. (2008) from Thailand. Worldwide approximately 1000 species have been described (Keller and Everhart 2010, Schnittler et al. 2006). Numerous surveys have been conducted in the United States of America. Alexopoulos (1965) and Alexopoulos and Henney (1971) reported collections from Texas. Ndiritu et al. (2009) and Tiffany and Knaphus (2001) reported collections from Big Bend National Park in Texas. Blackwell and Gilbertson (1980) and Evenson (1961) reported species from Arizona, Blackwell and Gilb- ertson (1984) from the Saguaro National Monument, Eliasson et al. (1988) from Arkansas, Graff (1928) from western Montana, Keller and Braun (1999) from Ohio, and Scarborough et al. (2009) from Missouri. Scarborough et al. (2009), Snell and Keller (2003), Snell et al. (2003), and Stephenson et al. (2001) report- ed slime molds from the Great Smoky Mountains National Park, Stephenson (1988) from Virginia, and Stephenson and Laursen (1993) from Alaska. Morris (1954) collected from mountainous areas of Wyoming, and Sturgis (1913) and Bethel and Sturgis (1907) collected from the mountainous areas of Colorado. Huffman et al. (2008) included the Myxomycetes in their field guide of fungi of the United States of America. Proceedings of the South Dakota Academy of Science, Vol. 89 (2010) 47 Little attention has been directed toward surveying the Great Plains in the mid-continental United States. This area as described by Kaul (1986) includes northwestern Texas, eastern New Mexico, western Oklahoma, eastern Colorado, Kansas, Nebraska, eastern Wyoming, South Dakota, eastern Montana and North Dakota. Keller and Schoknecht (1989) reported Fuligo megaspora from Colo- rado and northwestern Nebraska. Keller and Brooks (1971), Kramer (1956), Brooks (1941, 1942), and Schaffner (1904) collected in Kansas, and Seaver (1908) collected in North Dakota. Rollins (2007) initiated surveys in mid-con- tinental grasslands and has reported collections from the Thunder Basin National Grassland in Wyoming. Even fewer reports focus on the Black Hills in South Dakota and Bear Lodge Mountains in Wyoming. In 1898, Macbride published a list in the Proceedings of the Iowa Academy of Science of 30 species he collected during one August visit in the Black Hills of South Dakota. Later, Macbride (1922) listed an ad- ditional 3 species from the Black Hills, and 6 more species were located by staff in collections at the National Fungal Collection at Beltsville, Maryland. The Black Hills is a unique area where several biomes meet and overlap (Froiland 1990). The Nature Conservancy’s Ecoregions of the United States of America (1999) identifies the Black Hills as a unique ecoregion (Fig. 1). An- nual average precipitation for the Black Hills area from 1961-1990 was from less than 40 cm near the periphery to 60 cm in the Lead/Deadwood area (Trimarchi et al. 1998). The area is located along the western border of South Dakota and northeastern Wyoming, extending approximately 166 km north to south and 108 km east to west. It is approximately 480 km from the Rocky Mountains and even more isolated from the boreal and eastern deciduous forests. However, the flora includes representatives from each of these biomes, and the mid-continent grasslands. Geologically, it is an area characterized by an uplifted dome with a Precam- brian core. Some of the older rocks date to 2.5 billion years before the present. Several episodes of uplift and erosion have occurred, accompanied by sedimenta- tion. Near the end of the Cretaceous the last of the epicontinental seas retreated and more uplifts occurred extending as late as the Eocene. A final minor uplift occurred in the Pleistocene, producing essentially the modern appearance of the area (Froiland 1990). The primary goal of this survey was to extend our knowledge of the occurrence of plasmodial slime molds in the Black Hills of South Dakota and adjacent Wyo- ming, and western South Dakota, and to contribute to the ongoing world survey of Myxomycetes. The present survey includes specimens collected randomly and intermittently from the field for several years and specimens fruiting in culture on substrata that were collected from six selected sites in the Black Hills from 2007-2009. For the culture studies we hypothesized that more species and speci- mens would occur from wetter sites with higher canopy covers, and high plant species diversity/hectare in the northern hills compared to drier, more open sites with less plant species diversity/hectare in the southern hills (Gabel and Gabel 2007). We also hypothesized that more species and specimens would be recov- ered from woody substrata than on deciduous leaf and grass litter substrata. 48 Proceedings of the South Dakota Academy of Science, Vol. 89 (2010) METHODS Field Collections—From 1990-2009 fruiting bodies of plasmodial slime molds were collected intermittently and randomly from western South Dakota and adjacent eastern Wyoming. Specimens were air dried, placed in boxes with label information and placed in the fungal collection located in the Black Hills State University Herbarium (BHSC). Labels included: order, species, habitat, elevation, substrate, date collected, location (latitude/longitude or township, range and section), county, collection number, accession number, collector and identifier. Latitudes and longitudes are listed as the centroid for the site unless a reading was taken for a particular collection. Moist Chamber Cultures—The 6 sites in the Black Hills from which substrata were collected are described in Table 1 and located in Figure 1. Sites included 2 in the southern hills (Alabaugh Canyon and Boles/Redbird Canyons), 2 in the central hills (Pony Gulch and Spring Creek), and 2 in the northern hills (Botany Bay and Grigg’s Gulch). Downed, decaying wood, and dead deciduous leaf and grass litter retrieved from the ground were collected during the summers of 2007-2009.