Within-species Variation Research Articles

"And if you gaze long into an abyss, the abyss gazes also into thee": four morphs of Arctic charr adapting to a depth gradient in Lake Tinnsjøen.

The origin of species is a central topic in biology. Ecological speciation might be a driver in adaptive radiation, providing a framework for understanding mechanisms, level, and rate of diversification. The Arctic charr Salvelinus alpinus L. is a polymorphic species with huge morphological and life‐history diversity in Holarctic water systems. We studied adaptive radiation of Arctic charr in the 460‐m‐deep Lake Tinnsjøen to (a) document eco‐morphology and life‐history traits of morphs, (b) estimate reproductive isolation of morphs, and (c) illuminate Holarctic phylogeography and lineages colonizing Lake Tinnsjøen. We compared Lake Tinnsjøen with four Norwegian outgroup populations. Four field‐assigned morphs were identified in Lake Tinnsjøen: the planktivore morph in all habitats except deep profundal, the dwarf morph in shallow‐moderate profundal, the piscivore morph mainly in shallow‐moderate profundal, and a new undescribed abyssal morph in the deep profundal. Morphs displayed extensive life‐history variation in age and size. A moderate‐to‐high concordance was observed among morphs and four genetic clusters from microsatellites. mtDNA suggested two minor endemic clades in Lake Tinnsjøen originating from one widespread colonizing clade in the Holarctic. All morphs were genetically differentiated at microsatellites (F ST: 0.12–0.20), associated with different mtDNA clade frequencies. Analyses of outgroup lakes implied colonization from a river below Lake Tinnsjøen. Our findings suggest postglacial adaptive radiation of one colonizing mtDNA lineage with niche specialization along a depth–temperature–productivity–pressure gradient. Concordance between reproductive isolation and habitats of morphs implies ecological speciation as a mechanism. Particularly novel is the extensive morph diversification with depth into the often unexplored deepwater profundal habitat, suggesting we may have systematically underestimated biodiversity in lakes. In a biological conservation framework, it is imperative to protect endemic below‐species‐level biodiversity, particularly so since within‐species variation comprises an extremely important component of the generally low total biodiversity observed in the northern freshwater systems.

Does masting scale with plant size? High reproductive variability and low synchrony in small and unproductive individuals

In a range of plant species, the distribution of individual mean fecundity is skewed and dominated by a few highly fecund individuals. Larger plants produce greater seed crops, but the exact nature of the relationship between size and reproductive patterns is poorly understood. This is especially clear in plants that reproduce by exhibiting synchronized quasi-periodic variation in fruit production, a process called masting. We investigated covariation of plant size and fecundity with individual-plant-level masting patterns and seed predation in 12 mast-seeding species: Pinus pinea, Astragalus scaphoides, Sorbus aucuparia, Quercus ilex, Q. humilis, Q. rubra, Q. alba, Q. montana, Chionochloa pallens, C. macra, Celmisia lyallii and Phormium tenax. Fecundity was non-linearly related to masting patterns. Small and unproductive plants frequently failed to produce any seeds, which elevated their annual variation and decreased synchrony. Above a low fecundity threshold, plants had similar variability and synchrony, regardless of their size and productivity. Our study shows that within-species variation in masting patterns is correlated with variation in fecundity, which in turn is related to plant size. Low synchrony of low-fertility plants shows that the failure years were idiosyncratic to each small plant, which in turn implies that the small plants fail to reproduce because of plant-specific factors (e.g. internal resource limits). Thus, the behaviour of these sub-producers is apparently the result of trade-offs in resource allocation and environmental limits with which the small plants cannot cope. Plant size and especially fecundity and propensity for mast failure years play a major role in determining the variability and synchrony of reproduction in plants.

Diversity within species: interpreting strains in microbiomes.

Studying within-species variation has traditionally been limited to culturable bacterial isolates and low-resolution microbial community fingerprinting. Metagenomic sequencing and technical advances have enabled culture-free, high-resolution strain and subspecies analyses at high throughput and in complex environments. This holds great scientific promise but has also led to an overwhelming number of methods and terms to describe infraspecific variation. This Review aims to clarify these advances by focusing on the diversity within bacterial and archaeal species in the context of microbiomics. We cover foundational microevolutionary concepts relevant to population genetics and summarize how within-species variation can be studied and stratified directly within microbial communities with a focus on metagenomics. Finally, we describe how common applications of within-species variation can be achieved using metagenomic data. We aim to guide the selection of appropriate terms and analytical approaches to facilitate researchers in benefiting from the increasing availability of large, high-resolution microbiome genetic sequencing data.

Genetic relations among and within wild and cultivated Thymus species based on morphological and molecular markers.

In the present study, the diversity of 11 Thymus species was assessed using molecular and morphological markers. Essential oil content and morphological traits were also investigated during two agronomic years. The result of the analysis of variance showed considerable differences among morphological traits. In the first and second years, the essential oil content of the species varied from 0.63 to 1.94% and 0.86 to 2.34%, respectively. T. vulgaris and T. migricus showed the highest essential oil content in two agronomic years. In this research, nine ISSR primers were also used to amplify 151 polymorphic bands in 77 accessions belonging to 11 Thymus species. Cluster and principal component (PCA) analyses classified the species in three major groups. Among the species, T. vulgaris and T. fedtschenkoi presented relatively higher genetic distance in comparison with other species. Analysis of molecular variance (AMOVA) revealed that 72.34% of the total variation was belonged to within-species variation, while 27.66% was associated among the species. High gene flow (Nm = 1.11) and genetic differentiation (Gst = 0.31) were also observed among the species. T. transcaspicus exhibited the highest genetic variation (0.19), polymorphism % (57.69%), and Shannon index (0.29). The STRUCTURE analysis also showed a high admixture of Thymus species that might be originated from a high rate of natural hybridization. Finally, based on molecular and morphological information, T. vulgaris and T. carmanicus can be suggested as good candidate species for further breeding programs in Thymus species.

What Drives the Diversification of Eyespots in Morpho Butterflies? Disentangling Developmental and Selective Constraints From Neutral Evolution

Eyespots are wing colour pattern elements repeatedly observed in many butterflies, whose developmental genetics has been extensively documented. Nevertheless, the evolutionary forces shaping their diversification across species still remain largely unknown. Here we investigate the evolution of eyespots in the 30 species of the neotropical genus Morpho. Morpho butterflies display a series of eyespots located on the ventral side of their wings, highly variable among species, ranging from large, conspicuous structures to relictual spots. Applying geometric morphometrics to a large sample (N = 910) spanning all Morpho species and both sexes, we assess eyespots number, position, size and shape. We detect a divergence in eyespots position between understory and canopy species that opposes an L-shaped configuration in canopy species to a line pattern in understory species, where the eyespots tend to fuse and form a stripe in white and pale species. This effect is stronger than expected based on a Brownian motion model of phylogenetic divergence, suggesting an adaptation to the microhabitat and an influence of the wing colour on the evolution of pattern elements. Remarkably, this shift in colour pattern is strongly correlated with a shift in wing shape. However, using a thin-plate spline interpolation, we show that the shape change is insufficient to explain the evolution of eyespots position as a developmental side-effect of wing shape evolution, also pointing at an adaptive effect. Finally, we find a significant negative correlation between eyespot relative size and the within-species variation in eyespot number, position, size and shape, suggesting a relaxed or apostatic selection on small eyespots. We hypothesize that contrasted ecology may explain the observed differences between species: large and phenotypically stable eyespots might act as attack deflectors; small, variable faded eyespots might rather enhance crypsis; and pale species stripe pattern might disrupt the outline of the wing.

Genetic richness affects trait variation but not community productivity in a tree diversity experiment.

Biodiversity-ecosystem functioning experiments found that productivity generally increases with species richness, but less is known about effects of within-species genetic richness and potential interactions between the two. While functional differences between species can explain species richness effects, empirical evidence regarding functional differences between genotypes within species and potential consequences for productivity is largely lacking. We therefore measured within- and among-species variation in functional traits and growth and determined stand-level tree biomass in a large forest experiment factorially manipulating species and genetic richness in subtropical China. Within-species variation across genetic seed families, in addition to variation across species, explained a substantial amount of trait variation. Furthermore, trait responses to species and genetic richness varied significantly within and between species. Multivariate trait variation was larger among individuals from species mixtures than those from species monocultures, but similar among individuals from genetically diverse vs genetically uniform monocultures. Correspondingly, species but not genetic richness had a positive effect on stand-level tree biomass. We argue that identifying functional diversity within and among species in forest communities is necessary to separate effects of species and genetic diversity on tree growth and community productivity.

Components of leaf-trait variation along environmental gradients.

Leaf area (LA), mass per area (LMA), nitrogen per unit area (Narea ) and the leaf-internal to ambient CO2 ratio (χ) are fundamental traits for plant functional ecology and vegetation modelling. Here we aimed to assess how their variation, within and between species, tracks environmental gradients. Measurements were made on 705 species from 116 sites within a broad north-south transect from tropical to temperate Australia. Trait responses to environment were quantified using multiple regression; within- and between-species responses were compared using analysis of covariance and trait-gradient analysis. Leaf area, the leaf economics spectrum (indexed by LMA and Narea ) and χ (from stable carbon isotope ratios) varied almost independently among species. Across sites, however, χ and LA increased with mean growing-season temperature (mGDD0 ) and decreased with vapour pressure deficit (mVPD0 ) and soil pH. LMA and Narea showed the reverse pattern. Climate responses agreed with expectations based on optimality principles. Within-species variability contributed <10% to geographical variation in LA but >90% for χ, with LMA and Narea intermediate. These findings support the hypothesis that acclimation within individuals, adaptation within species and selection among species combine to create predictable relationships between traits and environment. However, the contribution of acclimation/adaptation vs species selection differs among traits.

Size-related effects and the influence of metabolic traits and morphology on swimming performance in fish.

Energy metabolism fuels swimming and other biological processes. We compared the swimming performance and energy metabolism within and across eight freshwater fish species. Using swim tunnel respirometers, we measured the standard metabolic rate (SMR) and maximum metabolic rate (MMR) and calculated the critical swimming speed (Ucrit). We accounted for body size, metabolic traits, and some morphometric ratios in an effort to understand the extent and underlying causes of variation. Body mass was largely the best predictor of swimming capacity and metabolic traits within species. Moreover, we found that predictive models using total length or SMR, in addition to body mass, significantly increased the explained variation of Ucrit and MMR in certain fish species. These predictive models also underlined that, once body mass has been accounted for, Ucrit can be independently affected by total length or MMR. This study exemplifies the utility of multiple regression models to assess within-species variability. At interspecific level, our results showed that variation in Ucrit can partly be explained by the variation in the interrelated traits of MMR, fineness, and muscle ratios. Among the species studied, bleak Alburnus alburnus performed best in terms of swimming performance and efficiency. By contrast, pumpkinseed Lepomis gibbosus showed very poor swimming performance, but attained lower mass-specific cost of transport (MCOT) than some rheophilic species, possibly reflecting a cost reduction strategy to compensate for hydrodynamic disadvantages. In conclusion, this study provides insight into the key factors influencing the swimming performance of fish at both intra- and interspecific levels.

Effects of urbanization on bird migration

Nearly one in five bird species is migratory, but not all individuals within a migratory species necessarily migrate: in partially migratory species, some do and some do not. Such within-species variability provides a natural experiment for investigating the mechanisms driving bird migration. Previous studies at the species level suggest that migrating provides a way to escape harsh winters, and to secure an increased access to resources, particularly important during the breeding season. Urbanization, by altering local temperatures (‘heat island’ effect) and resource availability (e.g. through garbage or garden feeders) can buffer the effects of winter harshness and modify breeding-season resource availability, potentially affecting individual migratory strategies. Here, we use ringing data from twelve North American partially migratory bird species to investigate the effects of natural environmental conditions (winter temperature, breeding season resource surplus) and urbanization on the propensity of individuals to migrate. We find strong support for the hypothesis that individuals migrate to avoid harsh winters, with, for eleven species, significantly higher probabilities of residency in areas with milder winters. We also found (significant for five species) that resource surplus in the breeding season reduces the propensity to migrate. Finally, urbanization increased the likelihood that individuals remain year-round in their ranges, avoiding to migrate away from their breeding range (four species) or their wintering areas (eight species), after controlling for climate and resources. Our results thus indicate that bird migratory strategies will respond to global change – in climate and land use – and indeed are already doing so.

Disentangling functional trait variation and covariation in epiphytic lichens along a continent-wide latitudinal gradient.

Characterizing functional trait variation and covariation, and its drivers, is critical to understand the response of species to changing environmental conditions. Evolutionary and environmental factors determine how traits vary among and within species at multiple scales. However, disentangling their relative contribution is challenging and a comprehensive trait-environment framework addressing such questions is missing in lichens. We investigated the variation in nine traits related to photosynthetic performance, water use and nutrient acquisition applying phylogenetic comparative analyses in lichen epiphytic communities on beech across Europe. These poikilohydric organisms offer a valuable model owing to their inherent limitations to buffer contrasting environmental conditions. Photobiont type and growth form captured differences in certain physiological traits whose variation was largely determined by evolutionary processes (i.e. phylogenetic history), although the intraspecific component was non-negligible. Seasonal temperature fluctuations also had an impact on trait variation, while nitrogen content depended on photobiont type rather than nitrogen deposition. The inconsistency of trait covariation among and within species prevented establishing major resource use strategies in lichens. However, we did identify a general pattern related to the water-use strategy. Thus, to robustly unveil lichen responses under different climatic scenarios, it is necessary to incorporate both among and within-species trait variation and covariation.

Polyploidy affects the seed, dormancy and seedling characteristics of a perennial grass, conferring an advantage in stressful climates.

Polyploidy (the state of having more than two genome copies) is widely distributed in flowering plants and can vary within species, with polyploid races often associated with broad ecological tolerances. Polyploidy may influence within-species variation in seed development, germination and establishment. We hypothesized that interactions between polyploidy and the seed developmental environment would affect subsequent dormancy, germination and early growth traits, particularly in stressful environments. Using seeds developed in a common garden under ambient and warmed conditions, we conducted germination trials under drought and temperature stress, and monitored the subsequent growth of seedlings. The study species, Themeda triandra, is a widespread, keystone, Australian native grass and a known polyploid complex. Tetraploid plants produced heavier, more viable seeds than diploids. Tetraploids were significantly more dormant than diploids, regardless of seed developmental environment. Non-dormant tetraploids were more sensitive to germination stress compared to non-dormant diploids. Finally, tetraploid seedlings were larger and grew faster than diploids, usually when maternal plants were exposed to developmental temperatures atypical to the source environment. Seed and seedling traits suggest tetraploids are generally better adapted to stressful environments than diploids. Because tetraploid seeds of T.triandra are more dormant they are less likely to germinate under stress, and when they do germinate, seedling growth is rapid and independent of seed developmental environment. These novel results demonstrate that polyploidy, sometimes in interaction with developmental environment and possibly also asexuality, can have within-species variation in seed and seedling traits that increase fitness in stressful environments.

Antennomere numbers in fireflies (Coleoptera: Lampyridae): unique patterns and tentative explanations

The plesiomorphic antenna with eleven articles is stunningly stable across the over 400.000 beetle species, despite the astonishing diversity of morphologies and ecological settings in which these organisms live. However, a few beetle lineages evolved different antennomere numbers, and these offer an interesting opportunity to understand how an otherwise fixed phenotype may eventually change, and to study what factors shape their patterns of variation. Here, we review for the first time, based on original and literature data, the unique patterns of variation in antennomere numbers in the firefly family (Coleoptera: Lampyridae), which has the greatest range (7–62) in antennomere numbers among beetles despite a relatively low species diversity (∼2000 species). We also tested the hypothesis that antennomere numbers are positively related to body size and explored patterns of antennal asymmetry across Amydetes spp, which have the greatest range in antennomere numbers (24–62). Out of the ∼100 firefly genera, nine show antennomere numbers lower or higher than 11. Among those nine genera with unusual antennomere numbers, five show intraspecific variation as well. In taxa whose antennae have less than 11 antennomeres, males seem to follow mainly light cues to find mates. In Amydetes, we found that antennomere numbers are positively related and negatively allometric to body size, suggesting that in this group the mechanism shaping antennal segmentation is sensitive to body size. Moreover, fluctuating asymmetry was found to be widespread in Amydetes spp., with a difference of up to six antennomeres between left and right antennae. The combined occurrence of unusual antennomere numbers and within-species variation suggests that in several firefly taxa a destabilization of the mechanism driving antennal segmentation has occurred.

Quantifying the primary biotic resource use by fisheries: A global assessment

In this paper, the specific primary production required (SPPR expressed as kg-NPP/kg-fish in wet weight) of more than 1700 marine species were calculated directly from 96 published food web models using the newly developed SPPR calculation framework. The relationship between SPPR and other ecological factors were then statistically analyzed. Among- and within-species variability of SPPR were found to be both explained by trophic level (TL), suggesting similar mechanisms underpinning both sources of variability. Among species, we found that harvesting species at higher mean trophic levels (MTL) increases the mean SPPR by a factor of 19 per 1 unit increase in MTL. Based on our empirical relationship, the mean SPPR of more than 9000 marine species were predicted and subsequently used to assess the primary production required (PPR) to support fisheries in five major fishing countries in Europe. The results indicated that conventional approach to estimating PPR, which neglects food web ecology, can underestimate PPR by up to a factor of 5. Within species, we found that harvesting populations occupying a higher TL leads to a higher SPPR. For example, the SPPR of Atlantic cod in the Celtic Sea (TL = 4.75) was 5 times higher than in the Gilbert Bay (TL = 3.3). Our results, which are based on large amounts of field data, highlight the importance of properly accounting for ecological factors during the impact assessment of fisheries.

Light availability and rhizobium variation interactively mediate the outcomes of legume-rhizobium symbiosis.

Nutrients, light, water, and temperature are key factors limiting the growth of individual plants in nature. Mutualistic interactions between plants and microbes often mediate resource limitation for both partners. In the mutualism between legumes and rhizobia, plants provide rhizobia with carbon in exchange for fixed nitrogen. Because partner quality in mutualisms is genotype-dependent, within-species genetic variation is expected to alter the responses of mutualists to changes in the resource environment. Here we ask whether partner quality variation in rhizobia mediates the response of host plants to changing light availability, and conversely, whether light alters the expression of partner quality variation. We inoculated clover hosts with 11 strains of Rhizobium leguminosarum that differed in partner quality, grew plants under either ambient or low light conditions in the greenhouse, and measured plant growth, nodule traits, and foliar nutrient composition. Light availability and rhizobium inoculum interactively determined plant growth, and variation in rhizobium partner quality was more apparent in ambient light. Our results suggest that variation in the costs and benefits of rhizobium symbionts mediate host responses to light availability and that rhizobium strain variation might more important in higher-light environments. Our work adds to a growing appreciation for the role of microbial intraspecific and interspecific diversity in mediating extended phenotypes in their hosts and suggests an important role for light availability in the ecology and evolution of legume-rhizobium symbiosis.

My niche: individual spatial niche specialization affects within- and between-species interactions.

Intraspecific trait variation is an important determinant of fundamental ecological interactions. Many of these interactions are mediated by behaviour. Therefore, interindividual differences in behaviour should contribute to individual niche specialization. Comparable with variation in morphological traits, behavioural differentiation between individuals should limit similarity among competitors and thus act as a mechanism maintaining within-species variation in ecological niches and facilitating species coexistence. Here, we aimed to test whether interindividual differences in boldness covary with spatial interactions within and between two ecologically similar, co-occurring rodent species (Myodes glareolus, Apodemus agrarius). In five subpopulations in northeast Germany, we quantified individual differences in boldness via repeated standardized tests and spatial interaction patterns via capture-mark-recapture (n = 126) and automated VHF telemetry (n = 36). We found that boldness varied with space use in both species. Individuals of the same population occupied different spatial niches, which resulted in non-random patterns of within- and between-species spatial interactions. Behavioural types mainly differed in the relative importance of intra- versus interspecific competition. Within-species variation along this competition gradient could contribute to maintaining individual niche specialization. Moreover, behavioural differentiation between individuals limits similarity among competitors, which might facilitate the coexistence of functionally equivalent species and, thus, affect community dynamics and local biodiversity.

A rapid insect species identification system using mini-barcode pyrosequencing.

Rapid and accurate species identification is not only important for biodiversity studies and pest quarantine and management, but in some cases may also influence the results of international trade negotiations. In this study, we developed a rapid species identification system for insects. A universal DNA mini-barcode primer pair was designed to target ∼ 120 bp of the mitochondrial 16S rDNA gene. This primer set can amplify the targeted region from all 300 species of 26 insect orders tested as well as other classes of Arthropoda. Although we found no within-species variation in this region, it provided enough information to separate closely related species or species complexes, in particular Thrips spp. and Bemisia spp. By combining a quick DNA extraction method with pyrosequencing, we were able to generate DNA sequences and complete species identification within 5h. Mini-barcode pyrosequencing of 16S rDNA coupled with the GenBank database provides a rapid, accurate, and efficient species identification system. This system is therefore useful for biodiversity discovery, forensic identification, and quarantine control and management. © 2019 Society of Chemical Industry.

Stem length, not climate, controls vessel diameter in two trees species across a sharp precipitation gradient.

Variation in xylem conduit diameter traditionally has been explained by climate, whereas other evidence suggests that tree height is the main driver of conduit diameter. The effect of climate versus stem length on vessel diameter was tested in two tree species (Embothrium coccineum, Nothofagus antarctica) that both span an exceptionally wide precipitation gradient (2300-500mm). To see whether, when taking stem length into account, plants in wetter areas had wider vessels, not only the scaling of vessel diameter at the stem base across individuals of different heights, but also the tip-to-base scaling along individuals of similar heights across sites were examined. Within each species, plants of similar heights had similar mean vessel diameters and similar tip-to-base widening of vessel diameter, regardless of climate, with the slopes and intercepts of the vessel diameter-stem length relationship remaining invariant within species across climates. This study focusing on within-species variation--thus, avoiding noise associated with the great morphological variation across species--showed unequivocally that plant size, not climate, is the main driver of variation in vessel diameter. Therefore, to the extent that climate selects for differing vessel diameters, it will inevitably also affect plant height.

Floral Color, Anthocyanin Synthesis Gene Expression and Control in Cape Erica Species.

Introduction: The Cape Floristic Region (CFR) is a biodiversity hotspot, recognized globally for its unusually high levels of endemism. The origins of this biodiversity are a long-standing topic of research. The largest “Cape clade,” Erica, radiated dramatically in the CFR, its ca. 690 species arising within 10–15 Ma. Notable between- and within-species flower color variation in Erica may have contributed to the origins of species diversity through its impact on pollinator efficiency and specificity.Methods: We investigate the expression and function of the genes of the anthocyanin biosynthesis pathway that controls floral color in 12 Erica species groups using RT-qPCR and UPLC-MS/MS.Results: Shifts from ancestral pink- or red- to white- and/or yellow flowers were associated with independent losses of single pathway gene expression, abrogation of the entire pathway due to loss of the expression of a transcription factor or loss of function mutations in pathway genes.Discussion: Striking floral color shifts are prevalent amongst the numerous species of Cape Erica. These results show independent origins of a palette of mutations leading to such shifts, revealing the diverse genetic basis for potentially rapid evolution of a speciation-relevant trait.

Dramatic altitudinal variations in leaf mass per area of two plant growth forms at extreme heights

Knowledge of how leaf mass per area (LMA) and its components, leaf thickness (LT) and leaf density (LD), and related leaf traits may vary with altitude concerning different plant life forms at extremely high altitudes is scarce. We measured LMA, leaf nitrogen (Nmass), LT, LD, and the related anatomical characteristics of an evergreen shrub (Rhododendron aganniphum var. schizopeplum, RAS) and a wintergreen herb (Bergenia purpurascen, BP) along an altitudinal gradient above 4200 m a.s.l. to disclose how LMA for two different growth forms may vary with altitude and which leaf attribute determines the within-species variation of LMA. LMA significantly increased with rising altitude, at a pace of 40 g m−2/100 m for RAS and 12 g m−2/100 m for BP, respectively. LT was mainly responsible for the variability in LMA for RAS, and the variation of LT and LD contributed almost equally for BP. Nmass tended to decrease with rising altitude, and was negatively related to LMA and LT for both species. Foliar δ13C showed significantly rising trends with altitude, and was positively correlated with LMA and LT. The results indicated that plants at the extremely high altitudes are adapted to possessing especially higher LMA and steeper slopes of the relationships between LMA and altitude, as well as lower leaf nitrogen concentrations and higher δ13C to cope with the low-temperature environment. Compared with many other studies, LMA for both species are assumed to be more sensitive to the changing environment (especially warming). Also, Rhododendron shrub displayed steeper slope compared with dwarf herbs beneath the shrub layer indicating the former species should be more sensitive to warming climate. In all, exploring the plant strategies via leaf functional traits is of importance to understand the geography of evergreen species in alpine regions and its responses to future climate change.

Photosynthetic Responses of Turf-forming Red Macroalgae to High CO2 Conditions.

Seaweeds are important components of near-shore ecosystems as primary producers, foundation species, and biogeochemical engineers. Seaweed communities are likely to alter under predicted climate change scenarios. We tested the physiological responses of three perennial, turf-building, intertidal rhodophytes, Mastocarpus stellatus, Osmundea pinnatifida, and the calcified Ellisolandia elongata, to elevated pCO2 over 6weeks. Responses varied between these three species. E.elongata was strongly affected by high pCO2 , whereas non-calcified species were not. Elevated pCO2 did not induce consistent responses of photosynthesis and respiration across these three species. While baseline photophysiology differed significantly between species, we found few clear effects of elevated pCO2 on this aspect of macroalgal physiology. We found effects of within-species variation in elevated pCO2 response in M.stellatus, but not in the other species. Overall, our data confirm the sensitivity of calcified macroalgae to elevated pCO2 , but we found no evidence suggesting that elevated pCO2 conditions will have a strong positive or negative impact on photosynthetic parameters in non-calcified macroalgae.