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Abstract
<p id="d9835915e181">
<b>Background and Aims</b>
<i>Arachnitis uniflora</i> is a mycoheterotrophic plant that exploits arbuscular mycorrhizal
fungi of neighbouring
plants. We tested
<i>A. uniflora</i>'s specificity towards fungi across its large latitudinal range,
as well as the role
of historical events and current environmental, geographical and altitudinal variables
on fungal genetic diversity.
</p><p id="d9835915e192">
<b>Methods</b>
<i>Arachnitis uniflora</i> mycorrhizas were sampled at 25 sites. Fungal phylogenetic
relationships were reconstructed,
genetic diversity was calculated and the main divergent lineages were dated. Phylogeographical
analysis was performed with the main fungal clade. Fungal diversity correlations with
environmental factors were investigated.
</p><p id="d9835915e200">
<b>Key Results</b> Glomeraceae fungi dominated, with a main clade that likely originated
in the Upper
Cretaceous and diversified in the Miocene. Two other arbuscular mycorrhizal fungal
families not previously known to be targeted by
<i>A. uniflora</i> were detected rarely and appear to be facultative associations.
High genetic diversity,
found in Bolivia and both northern and southern Patagonia, was correlated with temperature,
rainfall and soil features.
</p><p id="d9835915e208">
<b>Conclusions</b> Fungal genetic diversity and its distribution can be explained
by the ancient evolutionary
history of the target fungi and by micro-scale environmental conditions with a geographical
mosaic pattern.
</p>
• Here, we describe a new database, MaarjAM, that summarizes publicly available Glomeromycota DNA sequence data and associated metadata. The goal of the database is to facilitate the description of distribution and richness patterns in this group of fungi. • Small subunit (SSU) rRNA gene sequences and available metadata were collated from all suitable taxonomic and ecological publications. These data have been made accessible in an open-access database (http://maarjam.botany.ut.ee). • Two hundred and eighty-two SSU rRNA gene virtual taxa (VT) were described based on a comprehensive phylogenetic analysis of all collated Glomeromycota sequences. Two-thirds of VT showed limited distribution ranges, occurring in single current or historic continents or climatic zones. Those VT that associated with a taxonomically wide range of host plants also tended to have a wide geographical distribution, and vice versa. No relationships were detected between VT richness and latitude, elevation or vascular plant richness. • The collated Glomeromycota molecular diversity data suggest limited distribution ranges in most Glomeromycota taxa and a positive relationship between the width of a taxon's geographical range and its host taxonomic range. Inconsistencies between molecular and traditional taxonomy of Glomeromycota, and shortage of data from major continents and ecosystems, are highlighted.
More than 400 species of vascular plants, in 87 genera, are acholophyllous and heterotrophic, but not directly parasitic upon autotrophs. They are usually, but incorrectly, described as 'saprophytes'since they are in fact nourished by means of specialized mycorrhizal associations. Although distributed world-wide, they are most abundant and show the greatest species-richness in the Neotropics and Palaeotropical regions. Their aerial parts range in size from a few centimetres to extensive liane types up to 40 m long. With few exceptions, their habitats are dense moist forests in which there is a surface accumulation of leaf litter, often in situations which are too shaded for autotrophic growth. Although the achlorophyllous mycorrhizal mode of life has evolved independently many times and in widely disparate taxonomic groups, such plants show strong convergent evolution in particular adaptations to their peculiar mode of life. Most prominant amongst these are reductions in the size of seed and embryo, and the lack of differentiation of the embryo at maturity. The number of seeds produced by each flower is typically very large and the shape, structure and surface features of seeds involving adaptation for wind dispersal show remarkable parallels in many species. Specific adaptations for zoochory are rare but well developed in a small number of genera, some of which produce scents like fungal fruit bodies or floral parts which mimic fungal sporocarps. Vegetative parts are often even more conspicuously reduced. Most myco-heterotrophs are entirely subterranean for most of their lives and these stages exhibit adaptations consistent with a change in function from organs of absorption to organs of storage, shown by the almost universal loss of root hairs, decrease in surface area as exhibited in short cylindric'vermiform'and tuberous roots or, in extreme cases, the complete suppression of roots and the formation of a swollen tuber or rhizome. Increased width of the root cortex often accommodates mycorrhizal infection and stores of carbohydrates and other materials obtained from the fungal symbiont. Mycorrhizal infection is confined to the below-ground parts of the plants but may be found there in modified stems as well as in roots. In many genera, stems are exceptionally slender and thread-like and their vascular tissues are either reduced to a single narrow cylinder of bicollateral bundles or, minimally, to four or six narrow bundles in the cortex. Secondary thickening is poorly developed in all but a tiny minority of species, lignification being confined to annular or, rarely, a few scalariform xylem vessels. Phloem is present in very small amounts and then mainly as parenchyma with sieve tubes frequently recorded as narrow and possibly with abherent sieve plates. Leaves are typically reduced to widely spaced achlorophyllous scales on the inflorescence axis. Occasionally, they are present only on underground rhizomes or tubers. The vascular supply to the leaf-scales, normally reduced to a single trace, may be absent. Vestigial stomata are sometimes found on leaves and, in a few species which retain traces of chlorophyll, on shoots but, in most fully heterotrophic species, stomata are absent from aerial parts. Since their seeds are very small and contain minimal reserve carbohydrates, the germination of myco-heterotrophs in nature would appear to depend upon infection by an appropriate symbiotic fungus at an early stage. The nature of the carbohydrates transferred from the fungus to the plants has not been determined. Once acquired from The fungal partner, most plants store carbon in a variety of forms, the most common of which is starch, although other compounds including glucomannan, fructan and calcium oxalate art important in some specks. Asexual reproduction is frequently important with root tubers, tubercles and rhizomes providing the means of vegetative spread. Nonetheless, all the angiospermous species recorded to date also reproduce sexually. Floral structures show varying degrees of reduction concomitant with myco-heterotrophy. Inflorescences are typically small, often with a single terminal flower, and the floral parts often show extreme simplification, with the production of unilocular or, more rarely, bilocular and trilocular ovaries. In some of the most highly adapted species, there is reduction of integumentary layers surrounding each ovule from the normal bitegmic condition to unitegmic or, occasionally, ategmic. With the principal exception of the Monotropaceae, placentation is typically trimerous and parietal. Flowers normally appear to be cross-pollinated and are brightly coloured. nectiferous, occasionally scented, and can demonstrate extreme morphological adaptations which attract insects as in the production of lone caudate tepals or fungus-mimicking structures. Much is still to be learned about the adaptive features and especially about the physiology of these plants and of their early developmental stages during which the essential associations with fungi are established. Similarly, studies of the taxonomy and physiology of most of their fungal partners are still in their infancy. Contents Summary 171 I. Introduction 172 II. Taxonomic and phylogenetic relationships of myco-heterotrophic plants 174 III. Distribution patterns 180 IV. Habitats 183 V. Embryology 185 VI. Characteristics of seeds 186 VII. Mycorrhizal infection 192 VIII. Morphologies of roots 196 IX. Characteristics of shoots 199 X. Carbon assimilation and storage by mycoheterotrophic plants 202 XI. Reproduction 208 XII. Mutualism or parasitistn? 210 XIII. Future directions for research in mycoheterotrophic plants 210 XIV. Conclusions 211 Acknowledgements 211 References 211.
The publication of a large number of taxon names at all levels within the arbuscular mycorrhizal fungi (Glomeromycota) has resulted in conflicting systematic schemes and generated considerable confusion among biologists working with these important plant symbionts. A group of biologists with more than a century of collective experience in the systematics of Glomeromycota examined all available molecular-phylogenetic evidence within the framework of phylogenetic hypotheses, incorporating morphological characters when they were congruent. This study is the outcome, wherein the classification of Glomeromycota is revised by rejecting some new names on the grounds that they are founded in error and by synonymizing others that, while validly published, are not evidence-based. The proposed "consensus" will provide a framework for additional original research aimed at clarifying the evolutionary history of this important group of symbiotic fungi.
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