Woody plant taxonomic, functional, and phylogenetic diversity decrease along elevational gradients in Andean tropical montane forests: Environmental filtering and arrival of temperate taxa.

Mountains are paramount for exploring biodiversity patterns due to the mosaic of topographies and climates encompassed over short distances. Biodiversity research has traditionally focused on taxonomic diversity when investigating changes along elevational gradients, but other facets should be considered. For first time, we simultaneously assessed elevational trends in taxonomic, functional, and phylogenetic diversity of woody plants in Andean tropical montane forests and explored their underlying ecological and evolutionary causes. This investigation covered four transects (traversing ca. 2200 m a.s.l.) encompassing 114 plots of 0.1 ha across a broad latitudinal range (ca. 10°). Using Hill numbers to quantify abundance-based diversity among 37,869 individuals we observed a consistent decrease in taxonomic, functional, and phylogenetic diversity as elevation increased, although the decrease was less pronounced for higher Hill orders. The exception was a slight increase in phylogenetic diversity when dominant species were over-weighted. The decrease in taxonomic and functional diversity might be attributed to an environmental filtering process towards highlands, where the increasingly harsher conditions exclude species and functional strategies. Besides, the differences in steepness decrease between Hill orders suggest that rare species disproportionately contribute to functional diversity. For phylogenetic diversity the shifting elevational trend between Hill orders indicates a greater than previously considered influence in central Andean highlands of tropical lowlands originated species with strong niche conservatism relative to distantly related temperate lineages. This could be explained by a decreasing presence and abundance of temperate, extratropical taxa towards the central Andes relative to northern or southern Andes, where they are more prevalent.

Phylogenomics and Systematics of Overlooked Mesoamerican and South American Polyploid Broad-Leaved Festuca Grasses Differentiate F. sects. Glabricarpae and Ruprechtia and F. subgen. Asperifolia, Erosiflorae, Mallopetalon and Coironhuecu (subgen. nov.).

Allopolyploidy is considered a driver of diversity in subtribe Loliinae. We investigate the evolution and systematics of the poorly studied Mesoamerican and South American polyploid broad-leaved Festuca L. species of uncertain origin and unclear taxonomy. A taxonomic study of seven diagnostic morphological traits was conducted on a representation of 22 species. Phylogenomic analyses were performed on a representation of these supraspecific taxa and all other Loliinae lineages using separate data from the entire plastome, nuclear rDNA 45S and 5S genes, and repetitive DNA elements. F. subgen. Mallopetalon falls within the fine-leaved (FL) Loliinae clade, whereas the remaining taxa are nested within the broad-leaved (BL) Loliinae clade forming two separate Mexico-Central-South American (MCSAI, MCSAII) lineages. MCSAI includes representatives of F. sect. Glabricarpae and F. subgen. Asperifolia plus F. superba, and MCSAII of F. subgen. Erosiflorae and F. sect. Ruprechtia plus F. argentina. MCSAII likely had a BL Leucopoa paternal ancestor, MCSAI and MCSAII a BL Meso-South American maternal ancestor, and Mallopetalon FL, American I-II ancestors. Plastome vs. nuclear topological discordances corroborated the hybrid allopolyploid origins of these taxa, some of which probably originated from Northern Hemisphere ancestors. The observed data indicate rapid reticulate radiations in the Central-South American subcontinent. Our systematic study supports the reclassification of some studied taxa in different supraspecific Festuca ranks.

Evolutionary Dynamics of the Repeatome Explains Contrasting Differences in Genome Sizes and Hybrid and Polyploid Origins of Grass Loliinae Lineages.

The repeatome is composed of diverse families of repetitive DNA that keep signatures on the historical events that shaped the evolution of their hosting species. The cold seasonal Loliinae subtribe includes worldwide distributed taxa, some of which are the most important forage and lawn species (fescues and ray-grasses). The Loliinae are prone to hybridization and polyploidization. It has been observed a striking two-fold difference in genome size between the broad-leaved (BL) and fine-leaved (FL) Loliinae diploids and a general trend of genome reduction of some high polyploids. We have used genome skimming data to uncover the composition, abundance, and potential phylogenetic signal of repetitive elements across 47 representatives of the main Loliinae lineages. Independent and comparative analyses of repetitive sequences and of 5S rDNA loci were performed for all taxa under study and for four evolutionary Loliinae groups [Loliinae, Broad-leaved (BL), Fine-leaved (FL), and Schedonorus lineages]. Our data showed that the proportion of the genome covered by the repeatome in the Loliinae species was relatively high (average ∼ 51.8%), ranging from high percentages in some diploids (68.7%) to low percentages in some high-polyploids (30.7%), and that changes in their genome sizes were likely caused by gains or losses in their repeat elements. Ty3-gypsy Retand and Ty1-copia Angela retrotransposons were the most frequent repeat families in the Loliinae although the relatively more conservative Angela repeats presented the highest correlation of repeat content with genome size variation and the highest phylogenetic signal of the whole repeatome. By contrast, Athila retrotransposons presented evidence of recent proliferations almost exclusively in the Lolium clade. The repeatome evolutionary networks showed an overall topological congruence with the nuclear 35S rDNA phylogeny and a geographic-based structure for some lineages. The evolution of the Loliinae repeatome suggests a plausible scenario of recurrent allopolyploidizations followed by diploidizations that generated the large genome sizes of BL diploids as well as large genomic rearrangements in highly hybridogenous lineages that caused massive repeatome and genome contractions in the Schedonorus and Aulaxyper polyploids. Our study has contributed to disentangling the impact of the repeatome dynamics on the genome diversification and evolution of the Loliinae grasses.

Molecular Systematics of Valerianella Mill. (Caprifoliaceae): Challenging the Taxonomic Value of Genetically Controlled Carpological Traits.

Valerianella (cornsalad) is a taxonomically complex genus formed by 50-65 annual Holarctic species classified into at least four main sections. Carpological traits (sizes and shapes of achenes and calyx teeth) have been used to characterize its sections and species. However, the potential systematic value of these traits at different taxonomic ranks (from sections to species (and infraspecific taxa)) has not been tested phylogenetically yet. Here, we have assessed the evolutionary systematic value of Valerianella diagnostic carpological traits at different hierarchical ranks and have demonstrated their ability to separate taxa at the sectional level but not at species level for species of several species pairs. A total of 426 individuals (17 species, 4 sections) of Valerianella were analyzed using AFLP and plastid data. Genetic clusters, phylogenetic trees, and haplotype networks support the taxonomic classification of Valerianella at the four studied sectional levels (V. sects. Valerianella, Cornigerae, Coronatae, Platycoelae) but show admixture for ten taxa from five species pairs (V. locusta-V. carinata, V. coronata-V. pumila, V. multidentata-V. discoidea, V. dentata-V. rimosa, V. eriocarpa-V. microcarpa), which are not reciprocally monophyletic. Dating analyses indicate that the Valerianella sections are relatively old (mid-Miocene), while most species diverged in the Pliocene-Pleistocene. A new section Valerianella sect. Stipitae is described to accommodate the highly divergent and taxonomically distinct V. fusiformis type species. Taxonomic treatments that recognize the sectional ranks and that subsume the separate species of each species pair into single species represent a natural classification for Valerianella.

FunAndes - A functional trait database of Andean plants.

We introduce the FunAndes database, a compilation of functional trait data for the Andean flora spanning six countries. FunAndes contains data on 24 traits across 2,694 taxa, for a total of 105,466 entries. The database features plant-morphological attributes including growth form, and leaf, stem, and wood traits measured at the species or individual level, together with geographic metadata (i.e., coordinates and elevation). FunAndes follows the field names, trait descriptions and units of measurement of the TRY database. It is currently available in open access in the FIGSHARE data repository, and will be part of TRY's next release. Open access trait data from Andean plants will contribute to ecological research in the region, the most species rich terrestrial biodiversity hotspot.

Corrigendum: Museomics Unveil the Phylogeny and Biogeography of the Neglected Juan Fernandez Archipelago Megalachne and Podophorus Endemic Grasses and Their Connection With Relict Pampean-Ventanian Fescues.

[This corrects the article DOI: 10.3389/fpls.2020.00819.].

Museomics Unveil the Phylogeny and Biogeography of the Neglected Juan Fernandez Archipelago Megalachne and Podophorus Endemic Grasses and Their Connection With Relict Pampean-Ventanian Fescues.

Oceanic islands constitute natural laboratories to study plant speciation and biogeographic patterns of island endemics. Juan Fernandez is a southern Pacific archipelago consisting of three small oceanic islands located 600-700 km west of the Chilean coastline. Exposed to current cold seasonal oceanic climate, these 5.8-1 Ma old islands harbor a remarkable endemic flora. All known Fernandezian endemic grass species belong to two genera, Megalachne and Podophorus, of uncertain taxonomic adscription. Classical and modern classifications have placed them either in Bromeae (Bromus), Duthieinae, Aveneae/Poeae, or Loliinae (fine-leaved Festuca); however, none of them have clarified their evolutionary relationships with respect to their closest Festuca relatives. Megalachne includes four species, which are endemic to Masatierra (Robinson Crusoe island) (M. berteroniana and M. robinsoniana) and to Masafuera (Alejandro Selkirk island) (M. masafuerana and M. dantonii). The monotypic Podophorus bromoides is a rare endemic species to Masatierra which is only known from its type locality and is currently considered extinct. We have used museomic approaches to uncover the challenging evolutionary history of these endemic grasses and to infer the divergence and dispersal patterns from their ancestors. Genome skimming data were produced from herbarium samples of M. berteroniana and M. masafuerana, and the 164 years old type specimen of P. bromoides, as well as for a collection of 33 species representing the main broad- and fine-leaved Loliinae lineages. Paired-end reads were successfully mapped to plastomes and nuclear ribosomal cistrons of reference Festuca species and used to reconstruct phylogenetic trees. Filtered ITS and trnTLF sequences from these genomes were further combined with our large Loliinae data sets for accurate biogeographic reconstruction. Nuclear and plastome data recovered a strongly supported fine-leaved Fernandezian clade where Podophorus was resolved as sister to Megalachne. Bayesian divergence dating and dispersal-extinction-cladogenesis range evolution analyses estimated the split of the Fernandezian clade from its ancestral southern American Pampas-Ventanian Loliinae lineage in the Miocene-Pliocene transition, following a long distance dispersal from the continent to the uplifted volcanic palaeo-island of Santa Clara-Masatierra. Consecutive Pliocene-Pleistocene splits and a Masatierra-to-Masafuera dispersal paved the way for in situ speciation of Podophorus and Megalachne taxa.

Linking patterns and processes of tree community assembly across spatial scales in tropical montane forests.

Many studies have tried to assess the role of both deterministic and stochastic processes in community assembly, yet a lack of consensus exists on which processes are more prevalent and at which spatial scales they operate. To shed light on this issue, we tested two nonmutually exclusive, scale-dependent hypotheses: (1) that competitive exclusion dominates at small spatial scales; and (2) that environmental filtering does so at larger ones. To accomplish this, we studied the functional patterns of tropical montane forest communities along two altitudinal gradients, in Ecuador and Peru, using floristic and functional data from 60 plots of 0.1 ha. We found no evidence of either functional overdispersion or clustering at small spatial scales, but we did find functional clustering at larger ones. The observed pattern of clustering, consistent with an environmental filtering process, was more evident when maximizing the environmental differences among any pair of plots. To strengthen the link between the observed community functional pattern and the underlying process of environmental filtering, we explored differences in the climatic preferences of the most abundant species found at lower and higher elevations and examined whether their abundances shifted along the elevation gradient. We found (1) that greater community functional differences (observed between lower and upper tropical montane forest assemblies) were mostly the result of strong climatic preferences, maintained across the Neotropics; and (2) that the abundances of such species shifted along the elevational gradient. Our findings support the conclusion that, at large spatial scales, environmental filtering is the overriding mechanism for community assembly, because the pattern of functional clustering was linked to species' similarities in their climatic preferences, which ultimately resulted in shifts in species abundances along the gradient. However, there was no evidence of competitive exclusion at more homogeneous, smaller spatial scales, where plant species effectively compete for resources.