Ready for change: seed traits contribute to the high adaptability of mudflat species to their unpredictable habitat

Question A better understanding of species distribution and establishment requires in-depth information on their seed ecology. We hypothesised that seed traits of mudflat species may indicate a strong environmental adaptation in their highly specialised habitat. Furthermore, we asked the question, do seeds of mudflat species have a specific trait value to contribute high adaptability to small-scale variation in their unpredictable habitat? Location Central Europe. Methods Seeds of 30 typical mudflat species were used to measure 15 traits that govern seed dispersal (buoyancy and production), persistence (seed desiccation, mass and persistence in soil), and germination and establishment (germination response to different light, temperature and oxygen conditions). Cluster analysis and phylogenetic principal components analysis (pPCA) were conducted to define potential mudflat species functional groups as per their ecological optima. Results Seed production and seed mass displayed extremely high variation while seed buoyancy, desiccation and persistence in soil showed almost no variation. All study species produced buoyant, desiccation-tolerant and long-term persistent seeds. Germination and establishment traits also displayed similarity in their responses to different germination treatments as the majority (73%) of species has a moderate seed germination niche width. They germinated well under light/aerobic conditions irrespective to temperature fluctuations. The cluster analysis and pPCA separated species into three potential plant functional groups as 'true', 'flood-resistant' and 'facultative', mudflat species. Conclusion Moderate variation in the seed traits of mudflat plants suggests they employ different ecological strategies that seem highly predictive to the peculiarity of their specific micro-habitats, which are largely controlled by the hydroperiod gradient. It implies that seed trait information, which further needs to be tested for their adaptability, can advance our understanding of how community composition at the micro-habitat level depends on trait values of participating species

Inferring community assembly processes from functional seed trait variation along elevation gradient

Assembly of plant communities has long been scrutinized through the lens of trait-based ecology. Studies generally analyse functional traits related to the vegetative growth, survival and resource acquisition and thus ignore how assembly rules may affect plants at other stages of their life cycle, particularly when seeds disperse, persist in soil and germinate. Here, we analysed an extensive dataset of 16 traits for 167 species measured in-situ in 36 grasslands located along an elevation gradient and studied the impact of abiotic filtering, biotic interactions and dispersal on traits reflecting different trait categories: plant vegetative growth, germination, dispersal and seed morphology. For each community, we quantified community-weighted means (CWMs) and functional diversity (FD) for all traits and established their relationships to mean annual temperature. The seed traits were weakly correlated with vegetative traits. Therefore, these traits constituted independent axes of plant phenotypical variation that could be affected differently by community assembly rules. Abiotic filtering impacted mostly vegetative traits and to a lesser extent seed germination and morphological traits. Increasing low-temperature stress in upland sites selected for short-stature, slow-growing and frost-tolerant species that produce small quantities of small seeds with high degree of dormancy, high temperature requirements for germination and low germination speed. Biotic interactions, specifically competition in the lowlands and facilitation in uplands, also filtered some functional traits in the studied communities. The benign climate in lowlands seems to promote plant with competitive strategies that include fast growth and resource acquisition (vegetative growth traits) and early and fast germination (germination traits), whereas the effects of facilitation on the vegetative and germination traits were cancelled out by the strong abiotic filtering. The changes in the main dispersal vector from zoochory to anemochory along the elevation gradient strongly affected the dispersal and the seed morphological trait structure of the communities. This may be explained by stronger vertical turbulence and moderate warm upwinds and low grazing intensity in the uplands that select for light and non-round shaped seeds with lower terminal velocity and endozoochorous potential. Synthesis. We demonstrate that, in addition to vegetative traits, seed traits can substantially contribute to functional structuring of plant communities along environmental gradients. Thus, the ‘hard’ seed traits related to germination and dispersal are critical to detect multiple, complex community assembly rules. Consequently, such traits should be included in core lists of plant traits and, when applicable, be incorporated into the analysis of community assembly

Seed Germination Strategies of Mediterranean Halophytes Under Saline Condition

The study of the ecological strategies adopted by seed plants to ensure their success in different environments is closely related to germination ecology. This implies a careful knowledge of ecophysiology of seeds and, therefore, also of interaction between plants and the complexity of external factors. In particular, the environmental conditions of the area where a plant grows and produces seeds represent the main factors that influence successful seedling establishment. The physical-chemical features of habitats, and therefore their heterogeneity, affect the behavior of seeds in different ways. In addition to the timing of seed production, they can induce or terminate dormancy and/or germination and influence the germination pattern of different seeds in the same plant and so the composition and dispersal of soil seed banks. Salinity is a major abiotic stress affecting growth and plant productivity worldwide, constituting one of the main topics of study in the field of plant physiology. Halophytes are the plants that have the availability to survive and develop in different types of saline habitats. In this chapter, we consider some examples to illustrate the main adaptive strategies used by the seeds of halophytes on ecophysiological perspectives to survive in habitats affected by high levels of salinity. The focus is on the species that live in the brackish or salt coastal areas of the Mediterranean Basin. On these environments, the salt stress may act synergistically with intense anthropic pressure, generating profound alterations in the ecosystem and threatening the survival of the plant species very sensitive to the effects of climate change also. The results show the main diverse strategies, such as dormancy cycling, seed heteromorphism, and recovery capacity, from saline shock, favoring the chances of seed survival. The interaction between temperature and salinity during germination was also discussed assessing its crucial role as an ecological strategy

Broad Seed Germination Niche as an Adaptation to Heterogeneous Aquatic Habit—a Case Study of Four Potamogeton species

Aquatic habitats represent an extremely heterogeneous media for the inhabiting its plants, because key environmental factors vary there rapidly at relatively small scales. In this study, we explore the seed germination strategies of four pondweed species (Potamogeton compressus L., P. natans L., P. pectinatus L. and P.vaginatus Turcz.) in relation to the habitats they occupy. We hypothesised that, due to the high heterogeneity of aquatic environments, none of the environmental factors, such as light, temperature fluctuations or oxygen would play the role of a reliable trigger for seed germination of these species. The results of the germination experiments confirm the proposed hypothesis: under almost all conditions - fluctuating and constant temperatures, light and darkness, aerobic, flooded and hypoxia conditions - germination percentages ranged from 88.5 to 100%. An interspecific comparison of germination patterns also evidenced that all the investigated species showed the same response to all the studied environmental factors. We speculate that similar adaptations present in other aquatic macrophytes may point out the broader occurrence of this phenomenon in aquatic plants

Seed germination traits shape community assembly along a hydroperiod gradient

Background and Aims Hydroperiod drives plant community composition in wetlands, resulting in distinct zonation patterns. Here, we explored the role of seed germination traits in shaping wetland community assembly along a hydroperiod gradient. Specifically, we tested the hypothesis that seeds of reed, mudflat, swamp, shallow- and deep-water communities only germinate under a specific set of environmental factors characterized by the community-specific optimal conditions for seedling survival and growth. Methods In a three-factorial experiment, we tested the seed germination response of 50 species typical for temperate wetlands of Europe to temperature fluctuations (constant vs. fluctuating temperature), illumination (light vs. darkness) and oxygen availability (aerobic vs. hypoxia). Phylogenetic principal component analysis, cluster analysis and phylogenetic linear regressions were used to confirm the community-specific seed germination niches. Key Results Our study revealed the presence of five distinct, community-specific seed germination niches that reflect adaptations made by the study communities to decreasing light intensity, temperature fluctuations and oxygen availability along the hydroperiod gradient. Light as a germination trigger was found to be important in mudflats, swamps and shallow water, whereas the seeds of reed and deep-water species were able to germinate in darkness. A fluctuating temperature is only required for seed germination in mudflat species. Germination of species in the communities at the higher end of the hydroperiod gradient (reed and mudflat) demonstrated a strict requirement for oxygen, whereas swamp, shallow- and deep-water species also germinated under hypoxia. Conclusions Our study supports the recent argument that the inclusion of seed germination traits in community ecology adds significant insights to community response to the abiotic and biotic environment. Furthermore, the close relationship between seed germination adaptations and community assembly could help reach a better understanding of the existing patterns of wetland plant distribution at local scales and wetland vegetation dynamics, as well as facilitate nature conservation measures and aquatic habitat restoration

The sweet and musky scent of home: biogenic ethylene fine‐tunes seed germination in wetlands

Wetlands are known for comparatively high production of biogenic ethylene from decomposed sediment. Because the gas has various well-documented effects on seed physiology, we asked whether it can be a vital seed germination cue for wetland plants. Specifically, we explored whether ethylene plays an ecological role in (i) breaking/weakening seed dormancy, (ii) broadening the germination niche width, (iii) promoting germination speed or (iv) altering the germination requirements of six plant species with different occurrence along a hydroperiod gradient. In a factorial experiment, both ethylene-treated and untreated seeds were incubated in combinations of temperature (constant versus fluctuating), illumination (light versus darkness) and oxygen (aerobic versus hypoxia) with and without cold stratification. Our results revealed seed exposure to ethylene did not weaken or break dormancy without cold stratification treatment. However, ethylene helped to broaden the germination niche width, increased overall germination percentage and speed of cold-stratified (non-dormant) seeds. This indicates that ethylene helps those seeds that lost dormancy (non-dormant) to sense favourable water-saturated versus flooded substrate depending on their requirement for aerobic versus hypoxic conditions to trigger germination. We conclude that ethylene does not interfere directly with the dormancy-breaking process in autumn-dispersed seeds that are naturally cold-stratified in winter and germinate in spring/summer. However, ethylene plays a crucial ecological role as a ‘flood detector' for different wetland plant communities (reed, mudflat, swamp, shallow water) to synchronize germination of non-dormant seeds in the most suitable habitat at the right time

Seed germination ecology in Trapa natans L., a widely distributed freshwater macrophyte

Trapa natans is an aquatic annual plant that grows in stagnant water and occupies an extremely wide, yet discontinuous native range across temperate Europe, Asia, and Africa, and is naturalized in North America and Australia. Despite its wide distribution and invasion success, its seed germination ecology is poorly known. We hypothesized that, due to its wide distribution range, T. natans seeds should germinate under a broad spectrum of environmental conditions. In the present study, freshly matured seeds were cold stratified and tested for germination over a range of temperatures in light/dark, and dark under aerobic and hypoxia conditions. Seed and seedling tolerance to freezing and desiccation also were determined. Freshly matured seeds were physiologically dormant, requiring 9 weeks of cold stratification to overcome the dormancy. Seeds tolerated freezing down to -14 degrees C, whereas those with an emerged hypocotyl exhibited a high sensitivity to freezing down to -4 degrees C. The non-dormant seeds germinated (75-100%) under almost all environmental conditions, i.e. constant 22 degrees C and alternating temperature of 14/6 degrees C to 30/22 degrees C and 30/10 degrees C, light and darkness, oxic and hypoxia, thus supporting our proposed hypothesis. Finally, seeds were found to be highly desiccation-sensitive, even at high relative humidity. Our findings suggest that the wide germination niche could help explain its broad distribution and invasion success at local and global scales

Alpine plant communities differ in their seed germination requirements along a snowmelt gradient in the Caucasus

Mesoscale heterogeneity of alpine landscapes generates snowmelt gradients resulting in a distinct vegetation zonation from almost snow-free fellfields to long-lasting snowbeds. Although the vegetative trait variation along such gradients has been intensively studied, little is known about whether and how seed germination is adapted to the variable snowpack duration. Here, we compare the seed germination niches of 18 characteristic plant species occurring in four distinct plant communities (alpine lichen heath—Festuca varia grassland—Geranium-Hedysarum meadow—snowbed) along an alpine snowmelt gradient in the North Caucasus (Russia). In a fully factorial experiment, we tested seed germination responses to temperature (10/2, 14/6, 18/10, 22/14, 26/18 and 30/22 °C) and water potential (0, − 0.2, − 0.4, − 0.6 and − 0.8 MPa) gradients, reflecting the full spectrum of seedling establishment conditions in the study system. Community-specific germination niches were identified by generalised mixed model with Bayesian estimation. Our study revealed that the species from the four focal communities significantly differed in their temperature requirements for germination, whereas soil moisture was found to be a limiting factor for seedling establishment along the entire gradient. The snowbed germination was constrained to comparatively high temperatures above 10 °C, whereas seed germination in alpine lichen heaths occurred under a wide range of experimental conditions. The seed germination patterns of species from Festuca varia grasslands and Geranium-Hedysarum meadows reflected their intermediate position along the snowmelt gradient. We conclude that seed germination niche is affected by the environmental filtering along the snowmelt gradient thus departing from the general alpine germination syndrome

Seed dormancy and dormancy-breaking conditions of 12 West African woody species with high reforestation potential in the forest-savanna ecotone of Côte d'Ivoire

Information on the regeneration ecology of native woody species of the forest-savanna ecotone of West Africa is scarce, which is a major impediment to their optimal utilization in large-scale restoration programmes. The scattered information that is available for some of these species reveals that freshly matured seed are dormant. However, environmental heterogeneity among different habitats may results in inter-population seed dormancy variation. Thus, our objective was to re-examine the dormancy of 12 species from the forest-savanna ecotone that have been targeted for reforestation. Specifically, we aimed to examine the water-permeability of the seeds and explore the effectiveness of acid scarification and heat treatment to alleviate dormancy. Four species belonging to families other than Fabaceae and Malvaceae had water-permeable seeds. Two of them had nondormant (ND) seeds, and seeds of the other two species had a mixture of ND and other kinds of dormancy (possibly physiological dormancy, PD). Most species of Fabaceae and Malvaceae had water- impermeable seeds. All seeds of three species had physical dormant (PY), and some seeds of the remaining species had PY, while others were ND or had PD. Acid-scarification was effective in breaking PY and in augmenting imbibition and germination of non-PY seeds, while heat treatment was moderately effective in breaking dormancy. In general, acid scarification for 1-30 minutes and heat treatment for one hour at 55-75 degrees C were optimal to enhance seed germination, depending on species. The present study has wide practical implications for park conservationists and restoration ecologists interested in producing bulk quantities of high-quality planting stocks of native woody species for large-scale restoration programmes