Anne Cochrane

Predicting the impact of increasing temperatures on seed germination among populations of Western Australian Banksia (Proteaceae)

Temperature is a significant factor influencing seed germination and for many species temperature-mediated germination cues are vital for plant persistence. Rising temperatures forecast as a result of anthropogenic climate change may have a substantial influence on the population and range dynamics of plant species. Here, we report on the thermal constraints on seed germination in natural populations of four congeneric Banksia species collected from a longitudinal climate gradient in Western Australia. We investigated whether germination niche: (1) varied between species; (2) varied among populations of each species; and (3) varied in a consistent manner reflecting the climatic gradients of seed origin. We hypothesized that species would differ and that populations from warmer sites would have a broader temperature window for germination than populations from cooler sites. Species differed in the breadth of their germination niche, but temperatures that stimulated the most rapid and complete germination were similar across all species. A sharp reduction in germination percentage occurred above the optimum temperature, which coincided with significant delays in germination relative to the optimum. The temperatures causing these declines varied among populations. Across the species, there was a significant correlation between optimum germination temperature and mean annual temperature at seed source; however, there was no relationship at the population level for individual species. These data provide insight into the vulnerability of Banksia species to climate change, with those populations that require lower temperatures for germination, or have narrower optimal ranges for germination, likely to be most vulnerable to a warming climate.

The phenotypic response of co-occurring Banksia species to warming and drying

Projected warmer and drier climates are expected to impact heavily on plant diversity in Mediterranean-climate ecosystems, but experimental investigations of sensitivity and adaptive capacity are needed to better understand species responses. Here, we examine the effects of warming and drying on growth and allocation in seedlings from populations of two co-occurring Banksia shrub species from south-west Western Australia. We hypothesised that the species would show ecological divergence in functional traits reflecting niche differentiation. We expected to see tolerance to warming and drought correlated with position of the population on a climate gradient. We predicted that populations at the warmer/drier end of the gradient would show greater homeostasis of growth and allocation patterns in response to experimental treatment. Seedlings of the two species differed in leaf and allocation traits and in responses to experimental warming and drying. B. coccinea had smaller leaves with higher specific leaf area, and accumulated less overall biomass compared to B. baxteri when grown under cooler conditions. Under warmer conditions, B. coccinea could maintain growth, whereas B. baxteri suffered significant decline in biomass accumulation. Under water deficit conditions, both species showed significant reductions in leaf mass and area. Under combined warmer/drier conditions B. baxteri forfeited height growth and biomass and increased leaf allocation. The results support our hypothesis that seedlings of B. baxteri and B. coccinea are divergent in key functional traits and their sensitivity to warming and drying. However, we found no evidence for inter-population variation in traits being associated with position on a climate gradient.

Can sensitivity to temperature during germination help predict global warming vulnerability

Seed germination is vital for persistence in species that rely on seeds for post-disturbance regeneration. It is a high-risk phase and vulnerable to environmental parameters. Here, I assessed temperature sensitivity for germination in Banksia L.f. (Proteaceae) from south-western Australia, screening all 38 endemic obligate seeder species. A bi-directional temperature gradient plate with 49 temperature combinations (constant and fluctuating) between 5 and 40°C was used to profile germination temperature requirements and identify upper and lower temperature thresholds for germination. Using these data the impact of increasing temperatures on germination in these species was modelled under high and low greenhouse gas scenarios for 2050 and 2070. The results suggest that many Banksia species from the region have wide physiological tolerance for high germination temperatures, although a number of common, but geographically restricted species, such as B. praemorsa, B. oreophila and B. quercifolia , have more narrow temperature windows for germination than at least one of the rarer species ( B. verticillata ). Only B. dryandroides is expected to decline in germination in the future; however, the optimal germination timing for many species is predicted to occur later under climate warming. In conjunction with declining rainfall, this germination delay will place seedlings closer to the summer dry in this seasonal Mediterranean-climate ecosystem and thus they will be more vulnerable to desiccation. The framework developed here can be used to identify vulnerable species for monitoring of early population decline.

Modelling seed germination response to temperature in Eucalyptus L'Her. (Myrtaceae) species in the context of global warming

Seed germination is vital for persistence of many plant species, and is linked to local environmental conditions. Small increases in temperature during this critical life history transition may threaten species by altering germination timing and success. Such changes in turn may influence population dynamics, community composition and the geographic distributions of species. In this investigation, a bi-directional temperature gradient plate was used to profile thermal constraints for germination in 26 common, threatened and geographically restricted Eucalyptus species (Myrtaceae) from southern Western Australia. These observed data were used to populate models to predict optimum germination responses (mean time to germination, germination timing and success) under current (1950–2000 averages) and future (2070 high greenhouse gas emission climate scenario) mean monthly minimum and maximum temperatures. Many species demonstrated wide physiological tolerance for high germination temperatures and an ability to germinate outside current and forecast future autumn–winter wet season temperatures, suggesting that climatic distribution is a poor proxy for thermal tolerance for Eucalyptus seed germination. Germination for some species is predicted to decline under forecast conditions, but the majority will maintain or improve germination particularly during the cooler winter months of the year. Although thermal tolerance may benefit persistence of many Eucalyptus species in southern Western Australia as warming becomes more severe, large rainfall declines are also forecast which may prove more detrimental to plant survival. Nonetheless, this framework has the potential to identify seed resilience to heat stress in an early life history phase and hence species vulnerability to one characteristic of forecast environmental change.

Differences in seedling water-stress response of two co-occurring Banksia species

In the South-west Australian Floristic Region, timing of rainfall is critical for successful seedling establishment, as is surviving the first year’s summer drought for population persistence. Predictions of a warmer, drier future, therefore, threaten the persistence of obligate seeding species. Here, we investigate the drought tolerance of two co-occurring Banksia (Proteaceae) species by withholding water in pots to different extents of soil drying. Seed was collected from high- and low-rainfall populations, to test for niche differentiation in water-use strategies at the species level, as well as population differentiation. On the basis of a more negative leaf water potential at minimal levels of stomatal conductance and quantum yield, B. coccinea was considered to be more drought tolerant than B. baxteri. This was supported at the anatomical level according to xylem-vessel attributes, with a higher estimated collapse pressure suggesting that B. coccinea is less vulnerable to xylem cavitation. Population contrasts were observed mainly for B. baxteri, with a lower leaf-expansion increment rate in the low-rainfall population providing for drought avoidance, which was reflected in a higher rate of survival than with the high-rainfall population in which 87.5% of plants showed complete leaf senescence. The implications of species differences in water-use strategies are that community dynamics may start to shift as the climate changes. Importantly, this shift may be population dependent. A systematic understanding of adaptive capacity will help inform the choice of population for use in revegetation programs, which may lead to increased resilience and persistence in the face of environmental change. The results of the present study suggest that should declines in B. baxteri populations be noted, revegetating with seed collected from the low-rainfall population may help improve the chances of this species surviving into the future.

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.