Invasive plants usually experience population differentiation as they expand from their initial invasive range to the edge. Moreover, invasive plants usually encounter competitors which shared different co-evolutionary histories with them. These factors may lead to varying responses of invasive plant populations to elevated nitrogen deposition during expansion. However, this issue has received limited attention in prior research. To address these challenges, we conducted a greenhouse experiment to investigate how population differentiation of Galinsoga quadriradiata interacts with the presence of various competitors in response to increased nitrogen deposition. Competitor types (new or old that shared short or long co-evolutionary history with the invader, respectively) were set to compete with the invasive central and edge populations under different nitrogen addition treatments. Individuals from the central population of G. quadriradiata, originating from the initial invasion range, showed greater total mass, reproduction and interspecific competitiveness compared with the edge population. Nitrogen addition improved growth and reproductive performance in both populations, and the central population had a stronger response compared with the edge population. The performance of G. quadriradiata was inhibited more effectively by old competitors than new competitors. Our results indicate that population differentiation occurs in terms of growth and competitiveness during the range expansion of G. quadriradiata, with the central population exhibiting superior performance. Co-evolutionary history with competitors is considered unfavorable for invasive plants in this study. Our results highlight the combined effects of population differentiation in invasive species and their co-evolution history with competitors in the context of global change factors.

Climate refugia can serve as a remnant habitat or stepping stones for species dispersal under climate warming. The largest freshwater lake by surface area, Lake Superior, USA and Canada, serves as a model system for understanding cooling-mediated local refugia, as its cool water temperatures and wave action have maintained shoreline habitats suitable for southern disjunct populations of arctic–alpine plants since deglaciation. Here, we seek to explain spatial patterns and environmental drivers of arctic–alpine plant refugia along Lake Superior’s shores, and assess future risk to refugia under moderate (+3.5 °C) and warmest (+5.7 °C) climate warming scenarios. First, we examined how the interactive effects of summer surface water temperatures and wind affected onshore temperatures, resulting in areas of cooler refugia. Second, we developed an ecological niche model for the presence of disjunct arctic–alpine refugia (pooling 1253 occurrences from 58 species) along the lake’s shoreline. Third, we fit species distribution models for 20 of the most common arctic–alpine disjunct species and predicted presence to identify refugia hotspots. Finally, we used the two climate warming scenarios to predict changes in the presence of refugia and disjunct hotspots. Bedrock type, elevation above water, inland distance, July land surface temperature from MODIS/Terra satellite and near-shore depth of water were the best predictors of disjunct occurrences. Overall, we predicted 2236 km of the shoreline (51%) as disjunct refugia habitat for at least one species under current conditions, but this was reduced to 20% and 7% with moderate (894 km) and warmest (313 km) climate change projections.

Global climate change poses a severe threat to mountain biodiversity. Phenotypic plasticity and local adaptation are two common strategies for alpine plant to cope with such change. They may facilitate organismal adaptation to contrasting environments, depending on the influences of the environment or genotype or their interacted effects. In this study, we use an endemic alpine plant (Rorippa elata) in the Hengduan mountains (HDM) to unravel its phenotypic basis of adaptation strategy and evaluate the relative contributions of environment and genotype to its phenotype. We transplanted 37 genotypes of R. elata into two common gardens across low and high elevations (2800 vs. 3800 m) during 2021–2022. Nine fitness-related traits were measured, including flowering probability and glucosinolates (GS) content. We estimated the environmental or genotypic contributions to the phenotype and identified the main environmental components. Our results revealed that both environment and genotype-by-environment interactions contributed to the phenotypes of R. elata. Latitudinal heterogeneity was identified as a key factor that explained 24% of the total phenotypic variation. In particular, genotypes of the northern HDM showed significantly higher plasticity in flowering probability than those of the southern HDM. Furthermore, within the southern HDM, GS content indicated local adaptation to herbivory stresses for R. elata genotypes along elevations. In conclusion, our results suggest that R. elata may have adapted to the alpine environment through species-level plasticity or regional-level local adaptation. These processes were shaped by either complex topography or interactions between genotype and mountain environments. Our study provides empirical evidence on the adaptation of alpine plants.

Plants typically experience great losses from their reproductive potential represented by ovule production to the post-dispersal crop of viable seed. We examined seed density and viability in a founder population of mountain birch (Betula pubescens ssp. tortuosa), aiming to quantify losses at different stages and examine potential selection forces on the reproduction success of the founder generation of an isolated population. At the time of the study (2017–2020), the population had recently reached reproductive maturity, following its colonization around 1990 through long-distance dispersal onto an early successional outwash plain in southeast Iceland. Seed densities were high, but 89% of apparently intact seeds did not contain an embryo, despite being visually indistinguishable from flled seeds. Externally evident losses amounted to about 45% of the total seed crop and were mostly due to predation by the gall midge Semudobia betulae. When all losses were accounted for, 2.7% of the seed crop remained viable and germinated. Pollen limitation may partially explain the high incidence of empty seeds. Excessive fower production is compatible with the predator satiation hypothesis but cannot explain pre-dispersal losses. Another adaptation to predation, masting, appears poorly developed in Iceland. Our results suggest the presence of constraints on the reproduction potential of the new island population, that are more limiting than in neighbouring populations, and we discuss their developmental, ecological, and environmental correlates.

The accuracy of the simulation of carbon and water processes largely relies on the selection of atmospheric forcing datasets when driving land surface models (LSM). Particularly in high-altitude regions, choosing appropriate atmospheric forcing datasets can effectively reduce uncertainties in the LSM simulations. Therefore, this study conducted four offline LSM simulations over the Tibetan Plateau (TP) using the Community Land Model version 4.5 (CLM4.5) driven by four state-of-the-art atmospheric forcing datasets. The performances of CRUNCEP (CLM4.5 model default) and three other reanalysis-based atmospheric forcing datasets (i.e. ITPCAS, GSWP3 and WFDEI) in simulating the net primary productivity (NPP) and actual evapotranspiration (ET) were evaluated based on in situ and gridded reference datasets. Compared with in situ observations, simulated results exhibited determination coefficients (R²) ranging from 0.58 to 0.84 and 0.59 to 0.87 for observed NPP and ET, respectively, among which GSWP3 and ITPCAS showed superior performance. At the plateau level, CRUNCEP-based simulations displayed the largest bias compared with the reference NPP and ET. GSWP3-based simulations demonstrated the best performance when comprehensively considering both the magnitudes and change trends of TP-averaged NPP and ET. The simulated ET increase over the TP during 1982–2010 based on ITPCAS was significantly greater than in the other three simulations and reference ET, suggesting that ITPCAS may not be appropriate for studying long-term ET changes over the TP. These results suggest that GSWP3 is recommended for driving CLM4.5 in conducting long-term carbon and water processes simulations over the TP. This study contributes to enhancing the accuracy of LSM in water–carbon simulations over alpine regions.

Focal plants are considerably affected by their neighbouring plants, especially when growing in heterogeneous soils. A previous study on grasses demonstrated that soil heterogeneity and species composition affected plant biomass and above- and belowground allocation patterns. We now tested whether these findings were similar for forbs. Three forb species (i.e. Spartina anglica, Limonium bicolor and Suaeda glauca) were grown in pots with three levels of soil heterogeneity, created by alternatively filling resource-rich and resource-poor substrates using small, medium or large patch sizes. Species compositions were created by growing these forbs either in monocultures or in mixtures. Results showed that patch size × species composition significantly impacted shoot biomass, root biomass and total biomass of forbs at different scales. Specifically, at the pot scale, shoot biomass, root biomass and total biomass increased with increasing patch size. At the substrate scale, shoot biomass and total biomass was higher at the large patch size than at the medium patch size, both in resource-rich and resource-poor substrates. Finally, at the community scale, monocultures had more shoot biomass, root biomass and total biomass than those in the two-species mixtures or three-species mixture. These results differ from earlier findings on the responses of grasses, where shoot biomass and total biomass decreased with patch size, and more shoot biomass and total biomass were found in resource-rich than resource-poor substrate. To further elucidate effects of soil heterogeneity on the interactions between neighbour plants, we advise to conduct longer-term experiments featuring a variety of functional groups.

In the early 20th century, numerous western botanists, often referred to as “plant hunters”, embarked on ambitious expeditions to China, playing a crucial role in the study of botany and botanical diversity. Despite their contributions, comprehensive assessments of their explorations are lacking. To bridge this gap, this article focuses on the work of Joseph Charles Francis Rock, a notable figure in that era. Our work revisits Rock’s botanical expeditions within the broader context of botanical diversity conservation. It outlines his historical experiences in collecting plants in China and enumerates the species composition and phenotypic traits of the plants he collected. Additionally, it also analyzes the spatial distribution of the species, the completeness of his collection, and the α-diversity and β-diversity of the plants he collected. Our findings reveal that Rock led four major botanical expeditions in China between 1922 and 1933, amassing a total of 28,184 sheets and 16,608 numbers across 204 families, 1,081 genera, and 4,231 species. His focus was predominantly on ornamental species, which exhibit a variety of flower colors and inflorescences. His collection work spanned 5 provinces, 35 cities, and 72 counties, with a notable concentration in the Hengduan Mountains, a current biodiversity hotspot. This study not only reconstructs Rock’s botanical legacy but also offers valuable historical data and fresh analytical insights for understanding contemporary plant diversity. It contributes to the ongoing discourse on the importance of preserving plant diversity as a cornerstone of environmental sustainability.

The dioecious plant, Hippophae rhamnoides, is a pioneer species in community succession on the Qinghai-Tibet Plateau (QTP), plays great roles in various ecosystem services. However, the males and females of the species differ both in their morphology and physiology, resulting in a change in the ratio of male to female plants depending on the environment. To further explore the functional traits critical to this sex-based distinctive response in the alpine grassland, we have surveyed the sex ratios, measured their photosynthetic parameters, height, leaf area and biomass allocation. The results showed that (i) The males had higher Pn, light saturation point, apparent quantum efficiency, A_max and lower water-use efficiency (WUE), which exhibited higher utilization efficiency or tolerance to strong light, while the females indicated higher utilization efficiency for low light and water. And it showed sex-specific biomass allocation patterns. (ii) H. rhamnoides populations across the successional stages all showed a male-biased sexual allocation, which was closely related to sex-specific WUE, Pn, root biomass/total biomass and root–crown ratio. (iii) The leaf traits of H. rhamnoides changed from higher N_area, P_area and leaf mass per area in the early and late to lower in the middle, which meant they moved their growth strategy from resource rapid acquisition to conservation as the succession progressed. (iv) The increasing soil total phosphorus mostly contributed to regulating the sex bias of populations and variations of traits during the succession. The results are vital for the management of grassland degradation and restoration due to shrub encroachment on the QTP.

Biological invasions by alien and range-expanding native plant species can suppress native plants through allelopathy. However, the homeland security hypothesis suggests that some native plants can resist invasion by producing allelopathic compounds that inhibit the growth of invasive plants. Most research has focused on allelopathic interactions between individual native and invasive plant species, with less emphasis on how allelopathy helps entire native communities resist invasions. Additionally, limited knowledge exists about allelopathic interactions between range-expanding native species and recipient native communities, and their influence on invasion success. To bridge this knowledge gap, we conducted two greenhouse competition experiments to test reciprocal allelopathic effects between a native woody range-expanding species, Betula fruticosa, and a community of four native herbaceous species (Sanguisorba officinalis, Gentiana manshurica, Sium suave, and Deyeuxia angustifolia) in China. We assessed whether B. fruticosa and the native community differed in their competitive effects and responses, and whether these were influenced by activated carbon, which neutralizes allelochemicals in the soil. Activated carbon reduced the suppressive effects of the native community on the above-ground biomass of B. fruticosa, which indicates that the native community exerted a strong allelopathic effect on B. fruticosa. In contrast, activated carbon only marginally enhanced the suppressive effects of B. fruticosa on the native community, which indicates that allelopathy is not the primary mechanism by which B. fruticosa exerts its suppression. Overall, these findings support the homeland security hypothesis and suggest that biotic resistance from the native herbaceous community may limit the invasion success of the woody range-expander B. fruticosa.

Drought can greatly impact the biodiversity of an ecosystem and play a crucial role in regulating its functioning. However, the specific mechanisms by which drought mediate the biodiversity effect (BE) on community biomass in above- and belowground through functional traits remain poorly understood. Here, we conducted a common garden experiment in a greenhouse, which included two plant species richness levels and two water addition levels, to analyze the effects of biodiversity on aboveground biomass (AGB), belowground biomass (BGB) and total biomass (TB), and to quantify the relationship between BEs and functional traits under drought conditions. Our analysis focused on partitioning BEs into above- and belowground complementarity effect (CE) and selection effect (SE) at the species level, which allowed us to better understand the impacts of biodiversity on community biomass and the underlying mechanisms. Our results showed that plant species richness stimulated AGB, BGB and TB through CEs. Drought decreased AGB, BGB and TB, simultaneously. In addition, the aboveground CE was positively associated with the variation in plant height. SEs in above- and belowground were negatively correlated with the community mean plant height and root length, respectively. Furthermore, drought weakened the aboveground CE by decreasing variation in plant height, resulting in a reduction in AGB and TB. Our findings demonstrate that the complementarity of species is an important regulator of community biomass in above- and belowground, the dynamics of biomass under environmental stress are associated with the response of sensitive compartments.

Plants adapt to the limitation of soil phosphorus (P) induced by nitrogen (N) deposition through a complex interaction of various root and leaf functional traits. In this study, a pot experiment was conducted to explore the effects of different levels of N addition (control, low N [LN]: 25 kg N ha⁻¹ yr⁻¹, high N [HN]: 50 kg N ha⁻¹ yr⁻¹) on tree growth, leaf nutrient content, foliar P fractions and root characteristics of two dominant tree species, the pioneer species Salix rehderiana Schneid and the climax species Abies fabri (Mast.) Craib, in a subalpine forest in southwestern China. The results demonstrated that LN addition had a minimal impact on leaf N and P contents. Conversely, HN addition significantly decreased the leaf P content in both species. Salix rehderiana exhibited more pronounced increases in specific root length and specific root area under P deficiency triggered by HN addition when compared with A. fabri. In contrast, A. fabri showed weaker morphological responses to N addition but had a higher proportion of foliar P to metabolic P, as well as higher root exudates rate and root phosphatase activity in response to HN addition. Abies fabri employs a synergistic approach by allocating a greater amount of leaf P to metabolite P and extracting P from the soil through P-mobilizing exudates and root phosphatase activity, while S. rehderiana exhibits higher flexibility in modifying its root morphology in response to P limitation induced by HN addition. This study provides insights into subalpine tree species adaptation to N-induced P limitation, emphasizing its significance for guiding forest management and conservation in the context of global climate change.

Plants produce secondary chemicals that may vary along with latitude due to changing abiotic and biotic stress gradients and local environmental conditions. Teasing apart the individual and combined effects of these different abiotic, such as soil nutrients, and biotic factors, such as soil biota and herbivores, on secondary chemicals is critical for understanding plant responses to changing environments. We conducted an experiment at different latitudes in China, using tallow tree (Triadica sebifera) seedlings sourced from a population at 31° N. These seedlings were cultivated in gardens located at low, middle and high latitudes, with either local soil or soil from the original seed collection site (origin soil). The seedlings were exposed to natural levels of aboveground herbivores or had them excluded. Plant secondary chemicals (both foliar and root), aboveground herbivores and soil characteristics were measured. Results showed that most leaf and root secondary metabolites depended on the interaction of the experimental site and soil type. Leaf and root phenolic and tannin concentrations were higher at the middle latitude site, especially in the origin soil. Root and foliar flavonoid concentrations increased when aboveground herbivores were excluded. Microbial communities depended strongly on soil treatment. The different responses of tannins versus flavonoids suggest that these two chemical classes differ in their responses to the varying abiotic and biotic factors in these sites along latitudes. Taken together, our results emphasize the importance of considering the interactive effects of local environmental conditions, soil properties and herbivoryin regulating plant chemical defenses.

Examining the pollination biology of plant species is not only crucial for enhancing our understanding of their reproductive biology, but also essential for elucidating their adaptation and evolutionary history. Here, we investigated the breeding system and pollination biology of two closely related species in Actaea. The flower of A. purpurea is unique in the genus with purple and chartaceous (paper-like) sepals, fewer stamens with yellow anthers and purple filaments. Through three seasons of field observation and exclusion experiments, we determined that A. purpurea was primarily pollinated by a hornet species, Vespa bicolor, which also served as the most efficient pollinator. In contrast, A. japonica was primarily pollinated by large flies. Actaea purpurea exhibited a significantly higher cumulative nectar volume than A. japonica, which could be a crucial factor in attracting V. bicolor. A control experiment further demonstrated that removing the nectar leaf (petal) significantly decreased the visiting frequency of V. bicolor. Breeding system studies revealed that both species were self-compatible, yet they primarily underwent outcrossing in natural habitats. Our study presents a compelling case of possible pollinator shift in A. purpurea accompanied by morphological divergence. A more in-depth investigation of this system would offer crucial insights into the extent to which pollinators are involved in the plant speciation process and whether they contribute to reproductive isolation between closely related species.