Mika Bendiksby

The Plant Cell Wall–Decomposing Machinery Underlies the Functional Diversity of Forest Fungi

Comparative genomic analysis of “dry rot” fungus shows both convergent evolution and divergence among fungal decomposers. Brown rot decay removes cellulose and hemicellulose from wood—residual lignin contributing up to 30% of forest soil carbon—and is derived from an ancestral white rot saprotrophy in which both lignin and cellulose are decomposed. Comparative and functional genomics of the “dry rot” fungus Serpula lacrymans, derived from forest ancestors, demonstrated that the evolution of both ectomycorrhizal biotrophy and brown rot saprotrophy were accompanied by reductions and losses in specific protein families, suggesting adaptation to an intercellular interaction with plant tissue. Transcriptome and proteome analysis also identified differences in wood decomposition in S. lacrymans relative to the brown rot Postia placenta. Furthermore, fungal nutritional mode diversification suggests that the boreal forest biome originated via genetic coevolution of above- and below-ground biota.

MOLECULAR PHYLOGENETICS, CHARACTER EVOLUTION, AND SUPRAGENERIC CLASSIFICATION OF LAMIOIDEAE (LAMIACEAE)

Abstract This paper presents a phylogenetic analysis of Lamiaceae subfam. Lamioideae (including subfamily Pogostemonoideae) based on sequences of the trnL intron, trnL-trnF intergenic spacer, and rps16 intron of the plastid genome. It is the first analysis that includes all major lamioid and pogostemonoid genera. Monophyly of Lamioideae s.l. (i.e., including Pogostemonoideae) is strongly supported, with Cymaria Benth. as its sister group, and Pogostemonoideae, which sometimes has been recognized as a subfamily, is subsumed in Lamioideae. On the basis of the phylogenetic hypothesis, Lamioideae is divided into nine tribes. Three new tribes are established: Gomphostemmateae Scheen & Lindqvist, Phlomideae Mathiesen, and Leucadeae Scheen & Ryding. The other six tribes are: Pogostemoneae Briq., Synandreae Raf., Stachydeae Dumort., Leonureae Dumort., Lamieae Coss. & Germ., and Marrubieae Vis. The genus Betonica L. is reestablished. The results also strongly suggest that the genera Stachys L., Sideritis L., Ballota L., and Leucas R. Br. are polyphyletic or paraphyletic. The results were used to examine evolution of non-molecular characters.

Evolutionary history of Serpulaceae (Basidiomycota): molecular phylogeny, historical biogeography and evidence for a single transition of nutritional mode

BackgroundThe fungal genus Serpula (Serpulaceae, Boletales) comprises several saprotrophic (brown rot) taxa, including the aggressive house-infecting dry rot fungus Serpula lacrymans. Recent phylogenetic analyses have indicated that the ectomycorrhiza forming genera Austropaxillus and Gymnopaxillus cluster within Serpula. In this study we use DNA sequence data to investigate phylogenetic relationships, historical biogeography of, and nutritional mode transitions in Serpulaceae.ResultsOur results corroborate that the two ectomycorrhiza-forming genera, Austropaxillus and Gymnopaxillus, form a monophyletic group nested within the saprotrophic genus Serpula, and that the Serpula species S. lacrymans and S. himantioides constitute the sister group to the Austropaxillus-Gymnopaxillus clade. We found that both vicariance (Beringian) and long distance dispersal events are needed to explain the phylogeny and current distributions of taxa within Serpulaceae. Our results also show that the transition from brown rot to mycorrhiza has happened only once in a monophyletic Serpulaceae, probably between 50 and 22 million years before present.ConclusionsThis study supports the growing understanding that the same geographical barriers that limit plant- and animal dispersal also limit the spread of fungi, as a combination of vicariance and long distance dispersal events are needed to explain the present patterns of distribution in Serpulaceae. Our results verify the transition from brown rot to ECM within Serpulaceae between 50 and 22 MyBP.

Elucidating the evolutionary history of the Southeast Asian, holoparasitic, giant-flowered Rafflesiaceae: pliocene vicariance, morphological convergence and character displacement.

The aim of the present study is to elucidate the evolutionary history of the enigmatic holoparasitic Rafflesiaceae. More specifically, floral morphological evolution is interpreted in a molecular phylogenetic context, the biogeographic history of the family is investigated, and the possibility of character displacement to have been operating in this family is assessed. Parsimony and Bayesian methods are used to estimate phylogeny and divergence times among Rafflesiaceae species based on nuclear and mitochondrial DNA sequence data from Barkman et al. (2008) as well as new sequence data from additional samples and an additional genetic marker, the plastid 16S. Ancestral areas are inferred using dispersal-vicariance analysis (DIVA) as well a more recently developed parametric likelihood method (LAGRANGE), now including an update that allows for estimation over the posterior distribution of dated trees. Our extended molecular phylogeny of Rafflesiaceae implies a general lack of morphological synapomorphies as well as a high level of morphological homoplasy. In particular, a high level of floral morphological homoplasy is detected among Rafflesia species suggestive of similar patterns of pollinator-based selection in different geographic areas, and multiple instances of divergent floral size evolution is consistent with a model of character displacement. Initial diversification of Rafflesiaceae during the Late Cretaceous was followed by a long period of no-net diversification, likely due to extinctions caused by a Late Eocene to Miocene dramatic reduction in rainforest cover. A Late Miocene to Early Pliocene rise in sea-level probably caused the vicariant diversification observed between areas of endemism. The most recent species divergences are concordant with Pleistocene changes in climate and sea-levels, but apparently with no successful inter-area migrations, supportive of savannah, rather than rainforest, covered landbridges. An explosive increase in net diversification rate, most pronounced in Rafflesia, may be explained by Mid-Miocene to Pliocene rainforest-favorable conditions as well as natural selection promoting character displacement for Rafflesia flower size.

Accelerated Rates of Floral Evolution at the Upper Size Limit for Flowers

Evolutionary theory explains phenotypic change as the result of natural selection, with constraint limiting the direction, magnitude, and rate of response [1]. Constraint is particularly likely to govern evolutionary change when a trait is at perceived upper or lower limits. Macroevolutionary rates of floral-size change are unknown for any angiosperm family, but it is predicted that rates should be diminished near the upper size limit of flowers, as has been shown for mammal body mass [2]. Our molecular results show that rates of floral-size evolution have been extremely rapid in the endoholoparasite Rafflesia, which contains the worlds largest flowers [3]. These data provide the first estimates of macroevolutionary rates of floral-size change and indicate that in this lineage, floral diameter increased by an average of 20 cm (and up to 90 cm)/million years. In contrast to our expectations, it appears that the magnitude and rate of floral-size increase is greater for lineages with larger flowered ancestors. This study suggests that constraints on rates of floral-size evolution may not be limiting in Rafflesia, reinforcing results of artificial- and natural-selection studies in other plants that demonstrated the potential for rapid size changes [4-6].

Molecular phylogeny of tribe Stachydeae (Lamiaceae subfamily Lamioideae)

Although tribe Stachydeae (Lamiaceae) is considered monophyletic, relationships within the tribe are still poorly understood. The complexity of Stachydeae includes paraphyletic genera, considerable morphological plasticity, a range of ploidy levels, and presumably frequent natural hybridization. We performed parsimony and Bayesian phylogenetic analyses of nuclear (ribosomal ITS) and plastid (trnL intron, trnL-trnF spacer, rps16 intron) DNA sequence data from a taxonomically and geographically broad sampling of the tribe to identify major evolutionary lineages and to test taxonomic hypotheses within this largest of all lamioid tribes. We included 143 accessions corresponding to 121 species, representing both Old and New World species, and all 12 recognized genera of tribe Stachydeae. Both nuclear and plastid data corroborate monophyly of the tribe, with Melittis as sister to all remaining Stachydeae. For the latter well-supported clade, we suggest the phylogenetic name Eurystachys. Within Eurystachys, although monophyly is supported by both nuclear and plastid data for several named and unnamed groups, the majority of recognized taxa appear to be para- or polyphyletic. The taxon compositions of most subclades are congruent between the plastid and nuclear tree topologies, whereas their relative phylogenetic placements are often not. This level of plastid-nuclear incongruence suggests considerable impact of hybridization in the evolution of Stachydeae.

Allopolyploid origins of the Galeopsis tetraploids – revisiting Müntzing’s classical textbook example using molecular tools

Whole-genome duplication coupled with hybridization is of prime importance in plant evolution. Here we reinvestigate Müntzings classical example of allopolyploid speciation; the first report of experimental synthesis of a naturally occurring allopolyploid species, Galeopsis tetrahit. Various molecular markers (cpDNA, NRPA2, amplified fragment length polymorphisms (AFLPs)) and flow cytometry were surveyed in population samples of subgenus Galeopsis, including two allopolyploid species and their potential diploid parents. The presence of two divergent copies of single-copy NRPA2 confirms the allopolyploid origins of G. tetrahit and Galeopsis bifida. However, the two allopolyploids do not share the same maternal genome, as originally suggested by Müntzing. The results support independent origins, but not recurrent formation, of the two allotetraploids. Data further indicate frequent gene flow and introgression within ploidy levels, but less so between ploidy levels. Our results confirm and elaborate on Müntzings classical conclusion about allopolyploid origins of G. tetrahit and G. bifida. We address questions of general interest within polyploidy research, such as recurrent formation, gene flow and introgression within and between ploidy levels.

Finding Evolutionary Processes Hidden in Cryptic Species

Cryptic species could represent a substantial fraction of biodiversity. However, inconsistent definitions and taxonomic treatment of cryptic species prevent informed estimates of their contribution to biodiversity and impede our understanding of their evolutionary and ecological significance. We propose a conceptual framework that recognizes cryptic species based on their low levels of phenotypic (morphological) disparity relative to their degree of genetic differentiation and divergence times as compared with non-cryptic species. We discuss how application of a more rigorous definition of cryptic species in taxonomic practice will lead to more accurate estimates of their prevalence in nature, better understanding of their distribution patterns on the tree of life, and increased abilities to resolve the processes underlying their evolution.

Molecular phylogenetics and taxonomy of the Calvitimela aglaea complex (Tephromelataceae, Lecanorales)

Contributing to the process of reassigning lecideoid lichens to natural taxa, we assessed phylogenetic relationships and species delimitation in the Calvitimela aglaea complex (Tephromelataceae) using DNA sequence data and morphological/anatomical and chemical characters. Phylogenetic analysis of nuclear (ITS, MCM7, TEF1-α) and mitochondrial (ribosomal SSU) DNA sequences revealed Mycoblastus as sister to a strongly supported clade comprising Calvitimela, Tephrolema and Violella. Species of these three genera fall into six strongly supported subclades with low backbone resolution. Two of these are represented by Tephromela and Violella, which are readily circumscribed morphologically. The remaining four subclades encompass lineages that have until now been assigned to Calvitimela. While Tephromela and Violella as currently circumscribed are recovered as monophyletic in our analyses, Calvitimela is paraphyletic, with four deeply divergent clades. We recognize these four clades as subgenera Calomela, Calvitimela, Paramela and Severidea. Our molecular results further support the recognition of two recently discovered sterile crusts as new species, Calvitimela cuprea and C. livida, distinguished from previously known species by their production of asexual diaspores and from each other by secondary metabolite chemistry. We also report Calvitimela perlata as new for continental North America.

Species delimitation, bioclimatic range, and conservation status of the threatened lichen Fuscopannaria confusa

Fuscopannaria confusa is a rare lichen restricted to very humid localities in boreal forests. Two Fuscopannaria species, F. ahlneri and F. mediterranea, and Parmeliella parvula are morphologi- cally problematic to distinguish from F. confusa. Our aim with the present study was to evaluate the taxonomic status of F. confusa and thereby clarify its conservation status in Norway. By phylogenetic analysis of multi-locus DNA sequences, we show that F. confusa is genetically well distinguished from F. ahlneri, F. mediterranea, and P. parvula. Fuscopannaria confusa should therefore be treated as a separate species. A species distribution modelling analysis indicates that F. confusa has a slightly continental but potentially wide geographic distribution in Norway. However, suitable localities are continuously being destroyed by clear-cut logging and hydroelectric power development. Because of the decline in suitable habitats, F. confusa should be regarded as highly threatened in Norway and listed as EN (endangered) at the national level.