Filtering effect of temporal niche fluctuation and amplitude of environmental variations on the trait-related flowering patterns: lesson from sub-Mediterranean grasslands.

Timing of flowering is a critical component of community assembly, but how plant traits respond to heterogeneity of resources has been identified mostly through observations of spatial variations. Thus, we performed a trait-based phenological study in sub-Mediterranean grasslands to assess the importance of temporal variation of resources in the species assemblage processes. We found that early flowering species have traits allowing for slow resource acquisition and storage but rapid growth rate. Instead, mid- and late-flowering species exhibited sets of strategies devoted to minimizing water loss by evapotranspiration or aimed at maximizing the species' competitive ability, thanks to slow growth rate and more efficient resource acquisition, conservation and use. Our findings were consistent with the fluctuation niche theory. We observed that the amplitude of the environmental fluctuations influences the type and number of strategies positively filtered by the system. In fact, in the most productive grasslands, we observed the highest number of indicator trait states reflecting strategies devoted to the storage of resources and competition for light. Results seem also indicate that temporal variation of resources plays a role in trait differentiation and richness within a plant community, filtering traits composition of grasslands in the same direction, as formerly proved for spatial heterogeneity of resources.

The evolution of defense mechanisms correlate with the explosive diversification of autodigesting Coprinellus mushrooms (Agaricales, Fungi).

Bursts of diversification are known to have contributed significantly to the extant morphological and species diversity, but evidence for many of the theoretical predictions about adaptive radiations have remained contentious. Despite their tremendous diversity, patterns of evolutionary diversification and the contribution of explosive episodes in fungi are largely unknown. Here, using the genus Coprinellus (Psathyrellaceae, Agaricales) as a model, we report the first explosive fungal radiation and infer that the onset of the radiation correlates with a change from a multilayered to a much simpler defense structure on the fruiting bodies. We hypothesize that this change constitutes a key innovation, probably relaxing constraints on diversification imposed by nutritional investment into the development of protective tissues of fruiting bodies. Fossil calibration suggests that Coprinellus mushrooms radiated during the Miocene coinciding with global radiation of large grazing mammals following expansion of dry open grasslands. In addition to diversification rate-based methods, we test the hard polytomy hypothesis, by analyzing the resolvability of internal nodes of the backbone of the putative radiation using Reversible-Jump MCMC. We discuss potential applications and pitfalls of this approach as well as how biologically meaningful polytomies can be distinguished from alignment shortcomings. Our data provide insights into the nature of adaptive radiations in general by revealing a deceleration of morphological diversification through time. The dynamics of morphological diversification was approximated by obtaining the temporal distribution of state changes in discrete traits along the trees and comparing it with the tempo of lineage accumulation. We found that the number of state changes correlate with the number of lineages, even in parts of the tree with short internal branches, and peaks around the onset of the explosive radiation followed by a slowdown, most likely because of the decrease in available niches.

Phylogeny and species delimitation in the genus Coprinellus with special emphasis on the haired species.

We inferred the phylogenetic structure and species limits within Coprinellus by using a newly generated multigene alignment of LSU, ITS and ß-tubulin sequences. We sampled 154 specimens of 71 species out of ca. 80 known taxa in Coprinellus and inferred phylogenetic relationships by Bayesian MCMC and ML bootstrapping. Morphological and phylogenetic analyses revealed 17 new species, five of which are described herein. Coprinellus was split into three large clades, one comprising species with an initially continuous sheath of veil (Domestici/Micacei clade) and two consisting of primarily setulose (haired) species (eurysporoid and Core Setulosi clades). The separation of veiled species and the clade structure therein corresponds well to the morphology-based sectional classification, although the inclusion of certain setulose taxa (e.g. C. disseminatus, C. verrucispermus, C. curtus etc.) in these clades necessitates a new morphological definition for the sections. Morphological traits are discussed for all clades of Coprinellus. Species limits in the eurysporoid and Core Setulosi clades are scrutinized in detail. Future directions of species-rank research and an artificial key to the recovered setulose species of Coprinellus is presented.

Understanding the evolutionary processes of fungal fruiting bodies: correlated evolution and divergence times in the Psathyrellaceae.

Fruiting body evolution is one of the central topics in fungal evolutionary biology. A number of hypotheses have been developed to explain the contemporary diversity of fruiting body forms, but their evaluation has been hampered by the lack of well-sampled data sets and suitable statistical methods. Phylogenetic evidence of the physiological changes that accompany switches in fruiting body type is lacking, and very little is known about the age of major events of fruiting body evolution. Based on a new multigene phylogeny, by using Bayesian methods, we demonstrate the existence of correlation between a number of morphological features and switches from nondeliquescent to deliquescent (autodigesting) fruiting bodies in the mushroom family Psathyrellaceae. Our results show that switches in the anatomy of two types of spacer cells (cystidia and pseudoparaphyses) and basidia (bimorphic or monomorphic) as well as the structure of the mushroom cap follow the evolution of deliquescent fruiting bodies, which suggests strong functional linkage between these traits. We performed Bayes factor-based tests, referred hereafter to as evolutionary pathway test (EPT), to decide which of the correlated characters were gained first during evolution. The EPTs strongly suggest that deliquescence was gained first, followed after short waiting times by the other morphological features. Bayesian relaxed molecular clock analyses suggest that the various events of switching between fruiting body types occurred independently at various ages during the history of the family. The utility of two mushroom fossils (Archaemarasmius and Protomycena), the only ones with unambiguous taxonomic positions, for the calibration of agaric trees were also examined. Based on our results, we suggest that the evolutionary benefit of deliquescence may be prevention against desiccation via accelerated ontogeny of the fruiting body. Hypotheses regarding the functional significance of the correlated evolution are presented and discussed. Further, we argue that the changes in fruiting body types in mushrooms in general can be attributed to independent events (e.g., dispersal and adaptation) and not to particular geologic ages.