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Ozark Lichens PRELIMINARY DRAFT: OZARK LICHENS Enumerating the lichens of the Ozark Highlands of Arkansas, Illinois, Kansas, Missouri, and Oklahoma Prepared for the 14 th Tuckerman Lichen Workshop Eureka Springs, Arkansas October 2005 Corrected printing November 2005 Richard C. Harris New York Botanical Garden Douglas Ladd The Nature Conservancy Supported by the National Science Foundation grant 0206023 INTRODUCTION Well known as a biologically unique region North America, the Ozarks were long neglected from a lichenological standpoint. Systematic surveys and collecting work were initiated in the Missouri portion of the Ozarks in the early1980's, and were subsequently expanded to encompass the entire Ozark ecoregion, including small portions of Kansas and Illinois, and significant portions of Arkansas, Missouri and Oklahoma. These efforts have revealed a surprisingly rich diversity of lichens in the region, including a significant number of undescribed taxa. Despite considerable field work in every county in the region, new records continue to be found at a distressing rate, and we cannot yet state the total diversity of Ozark lichen biota. This draft is a tentative first attempt to provide a comprehensive treatment of the lichens of the Ozarks. Included here are general keys, brief synopses of genera, key to species within each genus with more than one Ozark taxon, and summaries of the Ozark distribution and ecology of each species, sometimes accompanied by more detailed taxonomic descriptions and other comments. As will be immediately evident to the reader, this draft is being rushed into preliminary distribution to be available for testing at the 2005 Tuckerman Lichen Workshop in the Ozarks. Hence a few disclaimers are stressed: this is an uneven treatment, in that some genera have been carefully studied, with detailed species descriptions and ecological profiles, while other groups are still problematical, with more cursory and provisional treatments. This project is an evolving work, and the user will note inconsistencies in style and approach from genus to genus. We have not included all unidentified species. Those we do include are treated inconsistently. Some are treated as new species, others as provisional (sp. provis. ), both as to their distinctness and as to the name supplied, and many known from only a single collection are distinguished simply by a collection number. In some cases, the keys will prove frustrating and require modification - it is with hope of meaningful criticism and input from users that this draft was rushed into production. Eventually, this draft will evolve into a unified comprehensive account of the lichen diversity of the Ozark highlands, complete with detailed distributional information, species descriptions and illustrations, and ecological information. ECOLOGICAL OVERVIEW OF THE OZARKS The Ozark region has long been recognized as a geologically, physiographically, ecologically, and culturally distinct area of North America (Figure 2). In conjunction with the Ouachita region to the south, the Ozarks comprise the only significant highland in midcontinental North America, and the only notable topographic relief between the Appalachians and the Rocky Mountains. This region is characterized by a diversity of terrestrial, aquatic, and karst habitats, ranging from extensive glades and tallgrass prairies to both coniferous and deciduous woodlands and cypress swamps, as well as fens, sinkholes, sloughs, and a suite of clear-flowing streams and rivers fed by an abundance of springs of all magnitudes, including some of the largest freshwater springs in North America. Encompassing 13.7 million hectares (34.3 million acres), the Ozarks includes portions of five states, with the majority of the region occurring within Missouri (67%) and Arkansas (24%) and smaller portions in Oklahoma (7%), Illinois (2%) and Kansas (0.1%). The Ozarks span a maximum of 270 miles (450 km) of north/south extent, and a maximum east/west extent 1 2 wide range of habitat types. This is a region of rugged uplands with copious exposed rocks and variable soil depths. The landscape in various terrestrial subsections of the Ozarks ranges from extensive areas of karst terrain on irregular plains, to highly dissected regions with steep hills and deeply entrenched valleys, as well as limited areas of ancient low mountains with elevations up to 925 meters (3000 feet). There are also smaller, linear areas of alluvial terrain and major riparian features. Bedrock geology of the Ozarks includes exposures of Precambrian igneous rocks in the eastern part of the Missouri Ozarks surrounded by alternating zones of Paleozoic sandstone and carbonate sedimentary rocks. Structurally, the Ozarks consist of a dome that has been slowly uplifted and eroded, resulting in a distinct landscape pattern. The oldest igneous rocks are exposed at the center of the uplift in southeast Missouri and surrounded by regions of Cambrian- and Ordovician-aged shallow water carbonates and beach sandstone strata. Further from the center are areas of younger Mississippian sedimentary rocks, including limestones and limited areas of riparian-derived freshwater sandstones (Nigh and Schroeder 2002). Dominant soils consist of Alfisols and Ultisols. The Alfisols, predominant in the less dissected terrestrial subsections, are thin loams with a clay component in the subsurface, and are generally thought to have formed under timber and some prairie vegetation types. Ultisols, predominant in the more rugged and dissected terrestrial subsections of the Ozarks, can in many ways be considered a more leached, weathered version of alfisols, with a much lower component of basic cations. Average precipitation in the Ozarks ranges from 39-52 inches (99-132 cm), with mean annual temperatures ranging from 54-63 °F (12-17 °C). The average frost free growing season ranges from 180-208 days. A major contributing factor to the region’s extreme biological diversity is that parts of the Ozarks have been continuously available for plant and animal life since the late Paleozoic some 230 million years ago, constituting perhaps the oldest continuously exposed land mass in North America, and one of the oldest on earth. Plants have presumably inhabited these rugged uplands since the origin of the modern angiosperms some 100 million years ago. Because of their central location within the continent, the Ozarks have on multiple occasions served as a refugium for organisms buffeted by climatic shifts associated with glacial and geologic events. The high levels of microhabitat diversity, influx of biota from divergent regions, and extreme antiquity of the landscape have combined to both sustain relictual populations and allow the development of new species, making the Ozarks a center of endemism in North America (i.e. Zollner et al. 2005). None of the four major continental glaciation events of the past two million years extended into the Ozarks. At the maximal extent of Wisconsin glaciation some 15,000 years ago, the climatic effects of a massive ice lobe extending into what is now central Iowa resulted in a boreal climate through much of midcontinental North America. At that time, the vegetation of the Ozarks was a combination of spruce-fir forests and jack pine parklands (Delcourt et al. 1986). Coincident with or preceding the glacial retreat, there has been a more or less continuous inhabitancy of the region by human cultures. These people had to secure all the necessities of survival from the local environment on a year round basis without significant trade or resource input from areas outside the Ozarks. The fact that such cultures not merely survived, but developed art, mythology, ceremony, religion, and other accoutrements of highly developed societies, testifies to their superb abilities to manage and interact with the Ozark environment. One of the most pervasive and effective tools available to early human populations in the region was wildland fire. An irrefutable body of evidence exists that the biological landscape of the Ozarks reflects the effects of millennia of frequent, low intensity, dormant season fires set by humans (e.g., Ladd 1991, Guyette & Cutter 1991). At the initiation of European settlement of the region, 3 predominate Native Americans in the Ozarks were the Osage. Parts of the eastern and southeastern Ozarks were home to the Quapaw. Thus, the pre-Eurosettlement vegetation in the Ozarks had been influenced since the end of the glacial period by an ongoing aboriginal fire regime. This vegetation consisted of a mosaic of matrix communities dominated by open woodland types, with various combinations of oaks and shortleaf pine as the principle overstory dominants in the uplands. Although Ozark woodlands differ significantly from the extensive deciduous woodlands extending eastward to the Atlantic coast, the Ozarks represent the westernmost extension of this eastern deciduous woodland formation that dominated much of eastern North America prior to European settlement. Extensive areas of tallgrass prairie occurred in the Ozarks, especially in the western terrestrial subsections (Schroeder 1981). Embedded within these matrix vegetation types was a diverse assemblage of small and large patch natural communities, including various types of fens, forests, wetlands, fluvial features and both carbonate and siliceous glades. The Ozarks ecoregion contains the largest extent of glade communities in North America (Nelson and Ladd 1980), and the extensive landscape of dolomite glades in the White River
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