Ecological niches in the polyploid complex Linum suffruticosum s.l.

Introduction: The high frequency of polyploidy in the evolutionary history of many plant groups occurring in the Mediterranean region is likely a consequence of its dynamic paleogeographic and climatic history. Polyploids frequently have distinct characteristics that allow them to overcome the minority cytotype exclusion. Such traits may enable polyploid individuals to grow in habitats different from their parentals and/or expand to new areas, leading to spatial segregation. Therefore, the successful establishment of polyploid lineages has long been associated with niche divergence or niche partitioning and the ability of polyploids to cope with different, often more stressful, conditions. In this study, we aimed to explore the role of environmental variables associated with the current distribution patterns of cytotypes within the polyploid complex Linum suffruticosum s.l.. Methods: The distribution and environmental niches of the five main cytotypes of Linum suffruticosum s.l. (diploids, tetraploids, hexaploids, octoploids and decaploids) were studied across its distribution range. Realized environmental niche of each cytotype was determined using niche modelling tools, such as maximum entropy modelling and niche equivalency and similarity tests. Results: Differences in the environmental conditions of L. suffruticosum s.l. cytotypes were observed, with polyploids being associated with habitats of increased drought and soil pH, narrower temperature ranges and decreased soil water and cation exchange capacities. Diploids present the widest environmental niche, and polyploids occupy part of the diploid niche. Although some polyploids have equivalent potential ecological niches, cytotypes do not co-occur in nature. Additionally, the ecological niche of this polyploid complex is different between continents, with North African habitats being characterised by differences in soil texture, higher pH, and low cation exchange capacity, precipitation and soil water capacity and higher temperatures than habitats in southwest Europe. Discussion: The different ecological conditions played a role in the distribution of cytotypes, but the mosaic distribution could not be entirely explained by the environmental variables included in this study. Other factors, such as reproductive isolation and competitive interactions among cytotypes, could further explain the current diversity and distribution patterns in white flax. This study provides relevant data on the niche requirements of each cytotype for further competition and reciprocal transplant experiments. further competition and reciprocal transplant experiments.

Invasive Acacia Tree Species Affect Instream Litter Decomposition Through Changes in Water Nitrogen Concentration and Litter Characteristics.

Non-native nitrogen-fixing Acacia species have been invading riparian ecosystems worldwide, potentially threatening stream communities that strongly depend on allochthonous litter. We examined the effects of the invasion of native deciduous temperate forests by Acacia species on litter decomposition and associated fungal decomposers in streams. Litter of native (Alnus glutinosa and Quercus robur) and invasive (Acacia melanoxylon) species were enclosed in fine-mesh bags and immersed in three native and three invaded streams, for 14-98 days. Litter decomposition rates, fungal biomass, and aquatic hyphomycete sporulation rates were higher in invaded than in native streams, likely due to the higher water nitrogen concentration found in invaded streams. Alnus glutinosa litter had higher aquatic hyphomycete sporulation rates and species richness, and higher decomposition rates, probably because they were soft and nitrogen rich. Quercus robur litter also had high aquatic hyphomycete sporulation rates but lower decomposition rates than Al. glutinosa, probably due to high polyphenol concentration and carbon:nitrogen ratio. Acacia melanoxylon litter had lower aquatic hyphomycete sporulation rates and species richness, and lower decomposition rates, most likely because it was very tough. Thus, litter decomposition rates varied in the order: Al. glutinosa > Q. robur > Ac. melanoxylon. The aquatic hyphomycete community structure strongly differed between native and invaded streams, and among litter species, suggesting that microbes were sensitive to water nitrogen concentration and litter characteristics. Overall, increases in water nitrogen concentration and alterations in litter characteristics promoted by the invasion of native riparian forests by Acacia species may affect the activity and community structure of microbial decomposers, and instream litter decomposition, thus altering the functioning of stream ecosystems.

Invasion of temperate deciduous broadleaf forests by N-fixing tree species - consequences for stream ecosystems.

Biological invasions are a major threat to biodiversity and ecosystem functioning. Forest invasion by alien woody species can have cross-ecosystem effects. This is especially relevant in the case of stream-riparian forest meta-ecosystems as forest streams depend strongly on riparian vegetation for carbon, nutrients and energy. Forest invasion by woody species with dissimilar characteristics from native species may be particularly troublesome. The invasion of temperate deciduous broadleaf forests with low representation of nitrogen (N)-fixing species by N-fixers has the potential to induce ecosystem changes at the stream level. Although effects of tree invasion on stream ecosystems have been under assessed, knowledge of native and invasive tree characteristics allows prediction of invasion effects on streams. Here we present a conceptual model to predict the effects of forest invasion by alien N-fixing species on streams, using as a background the invasion of temperate deciduous broadleaf forests by leguminous Acacia species, which are among the most aggressive invaders worldwide. Effects are discussed using a trait-based approach to allow the model to be applied to other pairs of invaded ecosystem-invasive species, taking into account differences in species traits and environmental conditions. Anticipated effects of N-fixing species invasions include changes in water quality (increase in N concentration) and quantity (decrease in flow) and changes in litter input characteristics (altered diversity, seasonality, typology, quantity and quality). The magnitude of these changes will depend on the magnitude of differences in species traits, the extent and duration of the invasion and stream characteristics (e.g. basal nutrient concentration). The extensive literature on effects of nutrient enrichment of stream water, water scarcity and changes in litter input characteristics on aquatic communities and processes allows prediction of invasion effects on stream structure and function. The magnitude of invasion effects on aquatic communities and processes may, however, depend on interactions among different pathways (e.g. effects mediated by increases in stream nutrient concentration may contrast with those mediated by decreases in water availability or by decreases in litter nutritional quality). A review of the literature addressing effects of increasing cover of N-fixing species on streams suggests a wide application of the model, while it highlights the need to consider differences in the type of system and species when making generalizations. Changes induced by N-fixing species invasion on streams can jeopardize multiple ecosystem services (e.g. good quality water, hydroelectricity, leisure activities), with relevant social and economic consequences.

Corrigendum: Different Patterns of Ecological Divergence Between Two Tetraploids and Their Diploid Counterpart in a Parapatric Linear Coastal Distribution Polyploid Complex.

[This corrects the article DOI: 10.3389/fpls.2020.00315.].

Different Patterns of Ecological Divergence Between Two Tetraploids and Their Diploid Counterpart in a Parapatric Linear Coastal Distribution Polyploid Complex.

Polyploidization is a widespread mechanism of evolutionary divergence in flowering plants. Ecological divergence of polyploid lineages has been proposed as a key process shaping the distribution of cytotypes in nature (niche shift hypothesis); however, evidence for the role of niche separation in replicated diploid-polyploid species pairs is still needed. This study aimed to assess the role of abiotic factors shaping current cytotype distributions. For that, we examined the distribution and environmental niches of two varieties recognized in diploid-tetraploid Jasione maritima across the species range and within a putative contact zone on the Iberian Peninsula. We counted chromosomes, screened for ploidy across Iberian Peninsula and characterized environmental requirements using niche modeling tools. We found that J. maritima var. maritima is composed by diploids with disjunct distribution in the west coast of France and northwest Iberian Peninsula, and by tetraploids in Iberian Peninsula, while var. sabularia is tetraploid. In the Iberian Peninsula, two parapatric contact zones along a linear coastal distribution were detected, one between diploid and tetraploid var. maritima, and the other between tetraploids of the two varieties. Environmental variables of diploid populations from France are distinct from those of southern diploid populations, which are more similar to tetraploids. In general, niche modeling results are congruent with the observed distribution patterns, although the results suggest a wider contact zone between varieties and cytotypes. Tetraploids of both varieties revealed different degrees of environmental divergence in comparison with their diploid counterpart. Tetraploid var. sabularia differed environmentally from diploids suggesting niche divergence. In contrast, tetraploid var. maritima overlapped with diploid environmental niche and currently occupies its entire predicted range, whereas diploids are restricted to northern areas of their suitable environment. Differences in ecological envelopes facilitate the recognition of functional units of biodiversity within polyploid groups, allowing the study of factors related to post-polyploidization divergence. Thus, whereas changes in environmental requirements may have allowed tetraploid var. sabularia to spread in habitats not favorable to diploids, other factors are involved with the distribution of diploid and tetraploid var. maritima.

Complex cytogeographical patterns reveal a dynamic tetraploid-octoploid contact zone.

The distribution of cytotypes in mixed-ploidy species is crucial for evaluating ecological processes involved in the establishment and evolution of polyploid taxa. Here, we use flow cytometry and chromosome counts to explore cytotype diversity and distributions within a tetraploid-octoploid contact zone. We then use niche modelling and ploidy seed screening to assess the roles of niche differentiation among cytotypes and reproductive interactions, respectively, in promoting cytotype coexistence. Two cytotypes, tetraploids and octoploids, were dominant within the contact zone. They were most often distributed parapatrically or allopatrically, resulting in high geographic isolation. Still, 16.7 % of localities comprised two or more cytotypes, including the intermediate hexaploid cytotype. Tetraploids and octoploids had high environmental niche overlap and associated with similar climatic environments, suggesting they have similar ecological requirements. Given the geographical separation and habitat similarity among cytotypes, mixed-ploidy populations may be transitional and subject to the forces of minority cytotype exclusion which lead to pure-ploidy populations. However, seed ploidy analysis suggests that strong reproductive barriers may enforce assortative mating which favours stable cytotype coexistence. High cytogenetic diversity detected in the field suggests that unreduced gamete formation and hybridization events seem frequent in the studied polyploid complex and might be involved with the recurrent polyploid formation, governing, as well, the gene flow between cytogenetic entities.