Insights into the tussock growth form with model-data fusion.

Some rhizomatous grass and sedge species form tussocks that impact ecosystem structure and function. Despite their importance, tussock development and size controls are poorly understood due to the decadal to centennial timescales over which tussocks form. We explored mechanisms regulating tussock development and size in a ubiquitous arctic tussock sedge (Eriophorum vaginatum) using field observations and a mass balance model coupled with a tiller population model. Model-data fusion was used to quantify parameter and prediction uncertainty, determine model sensitivity, and test hypotheses on the factors regulating tussock size. The model accurately captured the dynamics of tussock development, characteristics, and size observed in the field. Tussock growth approached maximal size within several decades, which was determined by feedbacks between the mass balance of tussock root necromass and density-dependent tillering. The model also predicted that maximal tussock size was primarily regulated by tiller root productivity and necromass bulk density and less so by tiller demography. These predictions were corroborated by field observations of tussock biomass and root characteristics. The study highlights the importance of belowground processes in regulating tussock development and size and enhances our understanding of the influence of tussocks on arctic ecosystem structure and function.

Landscape Genomics Provides Evidence of Ecotypic Adaptation and a Barrier to Gene Flow at Treeline for the Arctic Foundation Species Eriophorum vaginatum.

Global climate change has resulted in geographic range shifts of flora and fauna at a global scale. Extreme environments, like the Arctic, are seeing some of the most pronounced changes. This region covers 14% of the Earth's land area, and while many arctic species are widespread, understanding ecotypic variation at the genomic level will be important for elucidating how range shifts will affect ecological processes. Tussock cottongrass (Eriophorum vaginatum L.) is a foundation species of the moist acidic tundra, whose potential decline due to competition from shrubs may affect ecosystem stability in the Arctic. We used double-digest Restriction Site-Associated DNA sequencing to identify genomic variation in 273 individuals of E. vaginatum from 17 sites along a latitudinal gradient in north central Alaska. These sites have been part of 30 + years of ecological research and are inclusive of a region that was part of the Beringian refugium. The data analyses included genomic population structure, demographic models, and genotype by environment association. Genome-wide SNP investigation revealed environmentally associated variation and population structure across the sampled range of E. vaginatum, including a genetic break between populations north and south of treeline. This structure is likely the result of subrefugial isolation, contemporary isolation by resistance, and adaptation. Forty-five candidate loci were identified with genotype-environment association (GEA) analyses, with most identified genes related to abiotic stress. Our results support a hypothesis of limited gene flow based on spatial and environmental factors for E. vaginatum, which in combination with life history traits could limit range expansion of southern ecotypes northward as the tundra warms. This has implications for lower competitive attributes of northern plants of this foundation species likely resulting in changes in ecosystem productivity.

Responses of root phenology in ecotypes of Eriophorum vaginatum to transplantation and warming in the Arctic.

The effect of climate change on phenology and growth is less understood for belowground plant tissues than for aboveground plant tissues, particularly in high-latitude regions. Ecotypes within a species adapted to a locality may display different responses to climate change. We established two common garden plots in the Arctic tundra north of the Brooks Range in northern Alaska. Three ecotypes of Eriophorum vaginatum along a latitudinal gradient were transplanted into common gardens, and half of the transplants were warmed using open-top chambers (OTCs). Minirhizotrons were used to track the root phenology during the growing seasons of 2016 and 2017. Warming with OTCs (approximately +1 °C in air) did not affect the root biomass, root production or root phenology. The southern ecotype (from 67°16'N) of Eriophorum vaginatum transplanted northward experienced delayed startup and root production compared to two northern ecotypes (from 68°38'N and 69°25'N), although significant differences were not observed in the three ecotypes in terms of root production, root biomass and growth duration at the two sites. Our results suggest that as the climate warms, ecotypes of Eriophorum vaginatum may be able to adjust their duration of root growth and root productivity by phenotypic plasticity, although the degree of plasticity controlling the root startup time may vary between southern and northern ecotypes.

Comparative transcriptomics of an arctic foundation species, tussock cottongrass (Eriophorum vaginatum), during an extreme heat event.

Tussock cottongrass (Eriophorum vaginatum) is a foundation species for much of the arctic moist acidic tundra, which is currently experiencing extreme effects of climate change. The Arctic is facing higher summer temperatures and extreme weather events are becoming more common. We used Illumina RNA-Seq to analyse cDNA libraries for differential expression of genes from leaves of ecologically well-characterized ecotypes of tussock cottongrass found along a latitudinal gradient in the Alaskan Arctic and transplanted into a common garden. Plant sampling was performed on a typical summer day and during an extreme heat event. We obtained a de novo assembly that contained 423,353 unigenes. There were 363 unigenes up-regulated and 1,117 down-regulated among all ecotypes examined during the extreme heat event. Of these, 26 HSP unigenes had >log2-fold up-regulation. Several TFs associated with heat stress in previous studies were identified that had >log2-fold up- or down-regulation during the extreme heat event (e.g., DREB, NAC). There was consistent variation in DEGs among ecotypes, but not specifically related to whether plants originated from taiga or tundra ecosystems. As the climate changes it is essential to determine ecotypic diversity at the genomic level, especially for widespread species that impact ecosystem function.

Differential responses of ecotypes to climate in a ubiquitous Arctic sedge: implications for future ecosystem C cycling.

The response of vegetation to climate change has implications for the carbon cycle and global climate. It is frequently assumed that a species responds uniformly across its range to climate change. However, ecotypes - locally adapted populations within a species - display differences in traits that may affect their gross primary productivity (GPP) and response to climate change. To determine if ecotypes are important for understanding the response of ecosystem productivity to climate we measured and modeled growing season GPP in reciprocally transplanted and experimentally warmed ecotypes of the abundant Arctic sedge Eriophorum vaginatum. Transplanted northern ecotypes displayed home-site advantage in GPP that was associated with differences in leaf area index. Southern ecotypes exhibited a greater response in GPP when transplanted. The results demonstrate that ecotypic differentiation can impact the morphology and function of vegetation with implications for carbon cycling. Moreover they suggest that ecotypic control of GPP may limit the response of ecosystem productivity to climate change. This investigation shows that ecotypes play a substantial role in determining GPP and its response to climate. These results have implications for understanding annual to decadal carbon cycling where ecotypes could influence ecosystem function and vegetation feedbacks to climate change.

Effect of growth temperature on photosynthetic capacity and respiration in three ecotypes of Eriophorum vaginatum.

Ecotypic differentiation in the tussock-forming sedge Eriophorum vaginatum has led to the development of populations that are locally adapted to climate in Alaska's moist tussock tundra. As a foundation species, E. vaginatum plays a central role in providing topographic and microclimatic variation essential to these ecosystems, but a changing climate could diminish the importance of this species. As Arctic temperatures have increased, there is evidence of adaptational lag in E. vaginatum, as locally adapted ecotypes now exhibit reduced population growth rates. Whether there is a physiological underpinning to adaptational lag is unknown. Accordingly, this possibility was investigated in reciprocal transplant gardens. Tussocks of E. vaginatum from sites separated by ~1° latitude (Coldfoot: 67°15'N, Toolik Lake: 68°37', Sagwon: 69°25') were transplanted into the Toolik Lake and Sagwon sites and exposed to either an ambient or an experimental warming treatment. Five tussocks pertreatment combination were measured at each garden to determine photosynthetic capacity (i.e., Vcmax and Jmax) and dark respiration rate (Rd) at measurement temperatures of 15, 20, and 25°C. Photosynthetic enhancements or homeostasis were observed for all ecotypes at both gardens under increased growth temperature, indicating no negative effect of elevated temperature on photosynthetic capacity. Further, no evidence of thermal acclimation in Rd was observed for any ecotype, and there was little evidence of ecotypic variation in Rd. As such, no physiological contribution to adaptational lag was observed given the increase in growth temperature (up to ~2°C) provided by this study. Despite neutral to positive effects of increased growth temperature on photosynthesis in E. vaginatum, it appears to confer no lasting advantage to the species.

Ecotypic differences in the phenology of the tundra species Eriophorum vaginatum reflect sites of origin.

Eriophorum vaginatum is a tussock-forming sedge that contributes significantly to the structure and primary productivity of moist acidic tussock tundra. Locally adapted populations (ecotypes) have been identified across the geographical distribution of E. vaginatum; however, little is known about how their growth and phenology differ over the course of a growing season. The growing season is short in the Arctic and therefore exerts a strong selection pressure on tundra species. This raises the hypothesis that the phenology of arctic species may be poorly adapted if the timing and length of the growing season change. Mature E. vaginatum tussocks from across a latitudinal gradient (65-70°N) were transplanted into a common garden at a central location (Toolik Lake, 68°38'N, 149°36'W) where half were warmed using open-top chambers. Over two growing seasons (2015 and 2016), leaf length was measured weekly to track growth rates, timing of senescence, and biomass accumulation. Growth rates were similar across ecotypes and between years and were not affected by warming. However, southern populations accumulated significantly more biomass, largely because they started to senesce later. In 2016, peak biomass and senescence of most populations occurred later than in 2015, probably induced by colder weather at the beginning of the growing season in 2016, which caused a delayed start to growth. The finish was delayed as well. Differences in phenology between populations were largely retained between years, suggesting that the amount of time that these ecotypes grow has been selected by the length of the growing seasons at their respective home sites. As potential growing seasons lengthen, E. vaginatum may be unable to respond appropriately as a result of genetic control and may have reduced fitness in the rapidly warming Arctic tundra.

Northward displacement of optimal climate conditions for ecotypes of Eriophorum vaginatum L. across a latitudinal gradient in Alaska.

Plants are often genetically specialized as ecotypes attuned to local environmental conditions. When conditions change, the optimal environment may be physically displaced from the local population, unless dispersal or in situ evolution keep pace, resulting in a phenomenon called adaptational lag. Using a 30-year-old reciprocal transplant study across a 475 km latitudinal gradient, we tested the adaptational lag hypothesis by measuring both short-term (tiller population growth rates) and long-term (17-year survival) fitness components of Eriophorum vaginatum ecotypes in Alaska, where climate change may have already displaced the optimum. Analyzing the transplant study as a climate transfer experiment, we showed that the climate optimum for plant performance was displaced ca. 140 km north of home sites, although plants were not generally declining in size at home sites. Adaptational lag is expected to be widespread globally for long-lived, ecotypically specialized plants, with disruptive consequences for communities and ecosystems.

Clinal variation in stomatal characteristics of an Arctic sedge, Eriophorum vaginatum (Cyperaceae).

PREMISE OF THE STUDY: In a large reciprocal transplant experiment, Eriophorum vaginatum tussocks transplanted along a latitudinal gradient in Alaska's interior exhibited genetic differentiation and phenotypic plasticity for vegetative traits. Using the same tussocks 30 yr later, we used estimates of growing season temperature at each site to ask whether there was a climatic cline for stomatal density, size, and conductance. METHODS: We created impressions of the abaxial leaf surfaces of the transplanted individuals for viewing under a microscope and measured stomatal density (SD) and length (SL) for 224 individuals. We used SD and SL to estimate stomatal conductance (C). Separate one-way analyses of variance were performed to quantify the effect of population genetic differences and latitudinal environmental variation on stomatal characteristics. KEY RESULTS: Our data suggest that stomatal size was influenced by both genetics and environment and that plasticity for stomatal density produced highest densities at the coolest sites. Stomatal conductance increased with decreasing temperature of site from which the populations originated. CONCLUSIONS: Our results demonstrate a cline in stomatal conductance in E. vaginatum, with some ability of populations to plastically produce an appropriate phenotypic response in a new environment. Because the species is a dominant species in many arctic plant communities, its ability to produce an appropriate stomatal phenotype and to optimize water use efficiency by decreasing stomatal conductance in warmer environments could affect both community composition and total primary productivity in future, warmer climates.

Acclimation of tropical tree species to hurricane disturbance: ontogenetic differences.

We investigated acclimation responses of seedlings and saplings of the pioneer species Cecropia schreberiana Miq. and three non-pioneer species, Dacryodes excelsa Vahl, Prestoea acuminata (Willdenow) H.E. Moore var. montana (Graham) Henderson and Galeano, and Sloanea berteriana Choisy ex DC, following a hurricane disturbance in a lower montane wet forest in Puerto Rico. Measurements were made, shortly after passage of the hurricane, on leaves expanded before the hurricane (pre-hurricane leaves) and, at a later time, on recently matured leaves that developed after the hurricane (post-hurricane leaves) from both seedlings and saplings at sites that were severely damaged by the hurricane (disturbed sites) and at sites with little disturbance (undisturbed sites). Pre-hurricane leaves of the non-pioneer species had relatively low light-saturated photosynthetic rates (A(max)) and stomatal conductance (g(s)); neither A(max) nor g(s) responded greatly to the increase in irradiance that resulted from the disturbance, and there were few significant differences between seedlings and saplings. Pre-hurricane leaves of plants at undisturbed sites had low dark respiration rates per unit area (R(d)) and light compensation points (LCP), whereas pre-hurricane leaves of plants at disturbed sites had significantly higher R(d) and LCP. Post-hurricane leaves of plants at disturbed sites had significantly higher A(max) and R(d) than plants at undisturbed sites. Compared with seedlings, saplings had higher A(max) and R(d) and showed greater acclimation to the increase in irradiance that followed the disturbance. Post-hurricane leaves of the non-pioneer species had significantly lower A(max) and were less responsive to changes in irradiance than the pioneer species C. schreberiana. Variation in A(max) across light environments and stages was strongly related to differences in leaf mass per unit area (LMA), especially in the non-pioneer species. As indicated by V(cmax) or J(max) per unit nitrogen, light acclimation of A(max) was determined by leaf morphology (LMA) for the non-pioneer species and by both leaf morphology and leaf biochemistry for C. schreberiana. Ontogenetic changes in A(max) were attributable to changes in leaf morphology. The ontogenetic component of variation in A(max) across light environments and stages differed among species, ranging from 36 to 59% for the non-pioneer species (D. excelsa, 59.3%; P. acuminata var. montana, 44.7%; and S. berteriana, 36.3%) compared with only 17% in the pioneer species C. schreberiana.

Effects of light regime on the growth, leaf morphology, and water relations of seedlings of two species of tropical trees.

An experiment was conducted with Heliocarpus appendiculatus, a pioneer or large gap species of tropical moist forest in Costa Rica, and Dipteryx panamensis, a small gap species. Seedlings were grown in full sun, partial (80%) shade, and full (98%) shade. After one month of growth they were switched between environments and grown for two more months.Growth in height of Heliocarpus was greatly affected by irradiance, being increased in response to full shade and decreased in full sun. Height of Dipteryx was unaffected by irradiance level. Survival of Heliocarpus seedlings was only 49% in full shade, whereas Dipteryx had 100% survival. Biomass of Heliocarpus was not significantly greater in full sun than in partial shade whereas it was for Dipteryx. The response of root: shoot ratio was similar for both species. They were lowest in full shade and highest in full sun. Heliocarpus exhibited greater changes in leaf thickness, specific leaf weight, and stomatal density than did Dipteryx. Stomatal conductance of both species was lower in full shade and full sun than in partial shade.The results of the experiment indicate that growth of Heliocarpus is more plastic than that of Dipteryx in response to changes in irradiance. Previous environment did not affect the response to the present environment in either species. Both species responded positively to increases in irradiance.

Water relations of five tropical tree species on Barro Colorado Island, Panama.

Diurnal curves of xylem pressure potential (P) and leaf conductance (C) were measured for five tree species of the lowland tropical forest on Barro Colorado Island, Panama. Measurements were taken just before and just after the beginning of the rainy season. The species studied were: Cordia alliodora, Faramea occidentalis, Heisteria concinna, Macquira costaricana, and Trichilia cipo.For all species, predawn (base P) and daytime values of P increased markedly after the beginning of the rainy season. Diurnal patterns of C were quite diverse. C. alliodora showed a decline in C during the day whereas F. occidentaus, H. concinna, and M. costaricana showed little change. Conductance of the last three species was lower before the first rains. T. cipo exhibited more complicated behavior for P and C. Before the rainy season, low values of base P and midday P were observed (-26 bars and-39 bars, respectively). Conductance was also low. After the first rains, base P increased to-4 bars. At this time, midday P and C appeared to respond to vapor pressure deficit (VPD), being less on a day with high VPD than on a day with low VPD.