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10531 2007 9252 Article-Web 1..17 Biodivers Conserv (2008) 17:285–301 DOI 10.1007/s10531-007-9252-9 ORIGINAL PAPER Myxomycete diversity and distribution from the fossil record to the present Steven L. Stephenson · Martin Schnittler · Yuri K. Novozhilov Received: 8 February 2007 / Accepted in revised form: 10 April 2007 / Published online: 23 November 2007 © Springer Science+Business Media B.V. 2007 Abstract The myxomycetes (plasmodial slime molds or myxogastrids) are a group of eukaryotic microorganisms usually present and sometimes abundant in terrestrial ecosys- tems. Evidence from molecular studies suggests that the myxomycetes have a signiWcant evolutionary history. However, due to the fragile nature of the fruiting body, fossil records of the group are exceedingly rare. Although most myxomycetes are thought to have very large distributional ranges and many species appear to be cosmopolitan or nearly so, results from recent studies have provided evidence that spatial distribution patterns of these organ- isms can be successfully related to (1) diVerences in climate and/or vegetation on a global scale and (2) the ecological diVerences that exist for particular habitats on a local scale. A detailed examination of the global distribution of four examples (Barbeyella minutissima, Ceratiomyxa morchella, Leocarpus fragilis and Protophysarum phloiogenum) demon- strates that these species have recognizable distribution patterns in spite of the theoretical ability of their spores to bridge continents. Keywords Distribution patterns · Ecology · Long-distance dispersal · Microorganisms · Slime molds Special Issue: Protist diversity and geographic distribution. Guest editor: W. Foissner. S. L. Stephenson Department of Biological Sciences, University of Arkansas, Fayetteville, AR 72701, USA M. Schnittler (&) Institute of Botany and Landscape Ecology, Ernst Moritz Arndt University Greifswald, Grimmer Str. 88, 17487 Greifswald, Germany e-mail: [email protected] Y. K. Novozhilov V.L. Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov St. 2, 197376 St. Petersburg, Russia 1 C 286 Biodivers Conserv (2008) 17:285–301 Introduction The myxomycetes (also called plasmodial slime molds or myxogastrids) are a group of eukaryotic microorganisms usually present and sometimes abundant in terrestrial ecosys- tems. Myxomycetes have been known from their fruiting bodies since at least the middle of the seventeenth century, when the Wrst recognizable description of a member of the group (the very common species now known as Lycogala epidendrum) was provided by the Ger- man mycologist Thomas Panckow. Evidence from molecular studies (e.g., Baldauf and Doolittle 1997; Baldauf et al. 2000) indicates that the myxomycetes should be placed within the “crown” clade of eukaryotes, which would suggest that they have a signiWcant evolutionary history. However, due to the fragile nature of the fruiting body, fossil records of the group are exceedingly rare. Domke (1952) described a species of Stemonitis and Dörfelt et al. (2003) a species of Arcyria from Baltic amber dating from the Eocene, whereas Waggoner and Poinar (1992) reported the fossil of a myxomycete plasmodium in amber from Eocene-Oligocene deposits in the Dominican Republic. The maximum age that could be assigned to any of these fossils would not exceed about 50 million years, which is greater than that of the few records of fossil spores that appear to be those of myxomycetes, which date only from the Oligocene and Pleistocene (Graham 1971). Life cycle The myxomycete life cycle (Fig. 1) encompasses two very diVerent trophic stages, one consisting of uninucleate amoebae, with or without Xagella, and the other consisting of a distinctive multinucleate structure, the plasmodium (Martin et al. 1983). Under favorable conditions, the plasmodium gives rise to one or more fruiting bodies containing spores. IdentiWcation of myxomycetes is based almost exclusively upon features of the fruiting body (Martin and Alexopoulos 1969). The fruiting bodies produced by myxomycetes are somewhat suggestive of those produced by higher fungi, although they are considerably smaller (usually no more than 1–2 mm tall). The spores of the vast majority of myxomycetes range in size from 5 to 15 m in diameter, with most species producing spores 10 § 2 m in diameter. Presumably, the spores are wind-dispersed and complete the life cycle by germinating to produce the uninucleate amoeboXagellate cells. These feed and divide by binary Wssion to build up large populations in the various microhabitats in which these organisms occur. Ultimately, this stage in the life cycle gives rise to the plasmodium. This process can result from gametic fusion between compatible amoeboXagellates or it can be apomictic (Collins 1980, 1981). Bacteria apparently represent the main food resource for both trophic stages, but plasmodia are also known to feed upon yeasts, algae (including cyanobacteria), and fungal spores and hyphae (Stephenson and Stempen 1994). Under adverse conditions, such as drying out of the immediate environment or low temperatures, a plasmodium may convert into a hardened, resistant structure called a sclerotium, which is capable of reforming the plasmodium upon the return of favorable conditions. Moreover, amoeboXagellate cells can undergo a reversible transformation to dormant structures called microcysts. Both sclerotia and microcysts can remain viable for long periods of time and are probably very important in the continued survival of myxomycetes in some ecological situations and/or habitats, such as the bark surface of living trees and deserts. The fruiting bodies of many species of myxomycetes can achieve macroscopic dimen- sions and be collected and preserved for study in much the same way as the sporocarps of 1 C Biodivers Conserv (2008) 17:285–301 287 Fig. 1 Life cycle of a typical myxomycete. A, Spore. B, Germinating spore. C, Uninucleate amoeboid stage, with (right) or without (left) Xagella. D, Microcyst. E–F, Fusion of two compatible amoebae to produce a sin- gle cell. G, Zygote. H, Early plasmodium. I, Sclerotium. J, Portion of a mature plasmodium. K, Beginning of sporulation. L, Mature fruiting body with spores still enclosed. (Adapted from Stephenson and Stempen 1994) fungi or even specimens of bryophytes, lichens, and vascular plants. However, most species of myxomycetes tend to be rather inconspicuous or sporadic in their occurrence and are not always easy to detect in the Weld. Moreover, fruiting bodies of most species are relatively ephemeral and do not persist in nature for very long. Myxomycetes also spend a portion of their life cycle in a state where their very presence in a given habitat can be exceedingly diYcult if not impossible to determine. Because of their life history strategy and inconspicuous nature, these organisms provide an immense challenge in biodiversity assessments and, consequently, often have been neglected in such studies. Taxonomy Approximately 875 species of myxomycetes have been described (Lado 2001), and these have been placed in six diVerent taxonomic orders (Ceratiomyxales, Echinosteliales, 1 C 288 Biodivers Conserv (2008) 17:285–301 Liceales, Physarales, Stemonitales, and Trichiales). However, members of the Ceratiomyxales are distinctly diVerent from members of the other orders, and many modern workers have removed these organisms from the myxomycetes and reassigned them to the protostelids (Olive 1970, 1975; Olive and Stoianovitch 1979). The exact evolutionary aYnities of the myxomycetes are still debated, but these organisms constitute a well-deWned and homoge- nous group. Evidence from DNA sequence analysis (Baldauf and Doolittle 1997) suggests that even what appear to be closely related taxa on the basis of morphological similarity may have diverged from each other a long time ago (Clark 2000). In the Wrst phylogenetic study based on molecular data, Fiore-Donno et al. (2005) suggested that the Echinosteli- ales, which produce fruiting bodies with a simple structure, represented the most basal clade of myxomycetes, with two more advanced groups, the Wrst with light-colored spores and consisting of the Trichiales and the (presumably not monophyletic) Liceales, and the second having dark spores and made up of the Physarales and Stemonitales. Because of their small size and the limited array of morphological characters upon which their taxon- omy is based, determination of what constitutes a natural biological species, in the same sense that the concept is used for many of the more familiar groups of organisms (Mayr 1970), is sometimes rather problematic. It is now known that a number of the more common and widespread morphospecies actually consist of complexes of geographically restricted apomictic clonal lines (El Hage et al. 2000; Clark 2000; Clark and Stephenson 2000; Irawan et al. 2000). These genetically isolated lines are capable of independent evolution, which can lead to the accumulation of minor morphological diVerences that reXect speciWc adaptations to the particular set of environmental conditions in which they occur. For example, some of the forms found in special microhabitats (e.g., the inXores- cences of tropical herbs) diVer in some respects (e.g., color and size of the fruiting bodies) from specimens of the same species collected from more typical habitats. These almost certainly represent biotypes that are adapted to the microhabitat in question. Approximately 50% of all described species of myxomycetes are known only from the type locality or fewer
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