Predicting effects of warming requires a whole-of-life cycle perspective: a case study in the alpine herb Oreomyrrhis eriopoda.

Global warming is affecting plant phenology, growth and reproduction in complex ways and is particularly apparent in vulnerable alpine environments. Warming affects reproductive and vegetative traits, as well as phenology, but seldom do studies assess these traits in concert and across the whole of a plant's life cycle, particularly in wild species. Thus, it is difficult to extrapolate from such effects to predictions about the persistence of species or their conservation and management. We assessed trait variation in response to warming in Oreomyrrhis eriopoda, an Australian native montane herb, in which populations vary in germination strategy (degree of dormancy) and growth characteristics as a function of ecological factors. Warming accelerated growth in the early stages of development, particularly for populations with non-dormant seed. The differences in growth disappeared at the transition to reproduction, when an accelerating effect on phenology emerged, to varying degrees depending on germination strategy. Overall, warming reduced flower and seed production and increased mortality, indicating a reduction in reproductive opportunities, particularly for populations with dormant seed. Developmental condition affected germination strategy of the next generation seed, leading to increased degree of dormancy and slowed germination rate. But there were no whole-scale shifts in strategy or total germination percent. Following through the life cycle reveals that warming will have some potentially positive effects (early growth rates) and some negative effects (reduced reproductive output). Ultimately, warming impacts will depend on how those effects play out in the field: early establishment and an accelerated trajectory to seed maturity may offset the tradeoff with overall seed production. Small differences among germination strategies likewise may cascade to larger effects, with important implications for persistence of species in the alpine landscape. Thus, to understand and manage the response of wild species to warming takes a whole-of-life perspective and attention to ecologically significant patterns of within-species variation.

The seed germination spectrum of alpine plants: a global meta-analysis.

Assumptions about the germination ecology of alpine plants are presently based on individual species and local studies. A current challenge is to synthesise, at the global level, the alpine seed ecological spectrum. We performed a meta-analysis of primary data from laboratory experiments conducted across four continents (excluding the tropics) and 661 species, to estimate the influence of six environmental cues on germination proportion, mean germination time and germination synchrony; accounting for seed morphology (mass, embryo : seed ratio) and phylogeny. Most alpine plants show physiological seed dormancy, a strong need for cold stratification, warm-cued germination and positive germination responses to light and alternating temperatures. Species restricted to the alpine belt have a higher preference for warm temperatures and a stronger response to cold stratification than species whose distribution extends also below the treeline. Seed mass, embryo size and phylogeny have strong constraining effects on germination responses to the environment. Globally, overwintering and warm temperatures are key drivers of germination in alpine habitats. The interplay between germination physiology and seed morphological traits further reflects pressures to avoid frost or drought stress. Our results indicate the convergence, at the global level, of the seed germination patterns of alpine species.

Temperature variability drives within-species variation in germination strategy and establishment characteristics of an alpine herb.

Plant establishment and subsequent persistence are strongly influenced by germination strategy, especially in temporally and spatially heterogeneous environments. Germination strategy determines the plant's ability to synchronise germination timing and seedling emergence to a favourable growing season and thus variation in germination strategy within species may be key to persistence under more extreme and variable future climates. However, the determinants of variation in germination strategy are not well resolved. To understand the variation of germination strategy and the climate drivers, we assessed seed traits, germination patterns, and seedling establishment traits of Oreomyrrhis eriopoda from 29 populations across its range. Germination patterns were then analysed against climate data to determine the strongest climate correlates influencing the germination strategy. Oreomyrrhis eriopoda exhibits a striking range of germination strategies among populations: varying from immediate to staggered, postponed, and postponed-deep. Seeds from regions with lower temperature variability were more likely to exhibit an immediate germination strategy; however, those patterns depended on the timescale of climatic assessment. In addition, we show that these strategy differences extend to seedling establishment traits: autumn seedlings (from populations with an immediate or staggered germination strategy) exhibited a higher leaf production rate than spring seedlings (of staggered or postponed strategy). Our results demonstrate not only substantial within-species variation in germination strategy across the species distribution range, but also that this variation correlates with environmental drivers. Given that these differences also extend to establishment traits, they may reflect a critical mechanism for persistence in changing climate.

A research agenda for seed-trait functional ecology.

Trait-based approaches have improved our understanding of plant evolution, community assembly and ecosystem functioning. A major challenge for the upcoming decades is to understand the functions and evolution of early life-history traits, across levels of organization and ecological strategies. Although a variety of seed traits are critical for dispersal, persistence, germination timing and seedling establishment, only seed mass has been considered systematically. Here we suggest broadening the range of morphological, physiological and biochemical seed traits to add new understanding on plant niches, population dynamics and community assembly. The diversity of seed traits and functions provides an important challenge that will require international collaboration in three areas of research. First, we present a conceptual framework for a seed ecological spectrum that builds upon current understanding of plant niches. We then lay the foundation for a seed-trait functional network, the establishment of which will underpin and facilitate trait-based inferences. Finally, we anticipate novel insights and challenges associated with incorporating diverse seed traits into predictive evolutionary ecology, community ecology and applied ecology. If the community invests in standardized seed-trait collection and the implementation of rigorous databases, major strides can be made at this exciting frontier of functional ecology.