Seasonality and facilitation drive tree establishment in a semi-arid floodplain savanna.

A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree-grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C4 grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C4 grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C4 grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.

Relative contribution of photodegradation to litter breakdown in Australian grasslands.

Grassy ecosystems cover ~40% of the global land surface and are an integral component of the global carbon (C) cycle. Grass litter decomposes via a combination of photodegradation (which returns C to the atmosphere rapidly) and biological decomposition (a slower C pathway). As such, decomposition and C storage in grasslands may vary with climate and exposure to solar radiation. We investigated rates of grass litter decomposition in Australian temperate grasslands along a climate gradient to uncouple the relative importance of photodegradation and climate on decomposition. Litterbags containing leaf litter from two common native grass species (Poa labillardierei, Themeda triandra) were deployed at six grassland sites across a precipitation gradient (380-890 mm) in south-eastern Australia. Bags were retrieved over 39 weeks to measure mass loss from decomposition. We used shade treatments on the litter of one species (T. triandra) to partition photodegradation from biological decomposition. The shade treatment reduced the rate of decomposition of T. triandra relative to the full-sun treatment at all sites, by an average of 38% at 39 weeks; the effect size of the shade treatment was not correlated with site productivity. The rate of decomposition in both species was positively correlated with rainfall midway through the experiment, but there were no significant differences in total decomposition among sites after 39 weeks. By week 39, total decomposition of T. triandra was significantly greater than for P. labillardierei. In general, we observed relatively linear decomposition rather than the strong negative exponential decay observed in many global litter decomposition studies. Synthesis: We found that solar radiation exposure was a strong contributor to litter decomposition in temperate Australian grasslands across a broad climate gradient, which may be related to a period of photopriming prior to further biotic decomposition. This study highlights the importance of litter composition and solar radiation exposure in our understanding of how decomposition patterns contribute to global C cycling.

Lower dormancy with rapid germination is an important strategy for seeds in an arid zone with unpredictable rainfall.

Seed germination traits are key drivers of population dynamics, yet they are under-represented in community ecology studies, which have predominately focussed on adult plant and seed morphological traits. We studied the seed traits and germination strategy of eight woody plant species to investigate regeneration strategies in the arid zone of eastern Australia. To cope with stochastic and minimal rainfall, we predict that arid seeds will either have rapid germination across a wide range of temperatures, improved germination under cooler temperatures, or dormancy and/or longevity traits to delay or stagger germination across time. To understand how temperature affects germination responses, seeds of eight keystone arid species were germinated under laboratory conditions, and under three diurnal temperatures (30/20°C, 25/15°C and 17/7°C) for 30 days. We also tested for decline in seed viability across 24 months in a dry-aging treatment (~20°C). Six of the eight arid species studied had non-dormant, rapidly germinating seeds, and only two species had physiological dormancy traits. Seed longevity differed widely between species, from one recalcitrant species surviving only months in aging (P50 = <3 months) and one serotinous species surviving for many years (P50 = 84 months). Our results highlight the importance of understanding the reproductive strategies of plant species in arid environments. Rapid germination, the dominant seed trait of species included in this study, allows arid species to capitalise on sporadic rainfall. However, some species also exhibit dormancy and delayed germination; this an alternative strategy which spreads the risk of germination failure over time.

The risk-takers and -avoiders: germination sensitivity to water stress in an arid zone with unpredictable rainfall.

Water availability is a critical driver of population dynamics in arid zones, and plant recruitment is typically episodic in response to rainfall. Understanding species' germination thresholds is key for conservation and restoration initiatives. Thus, we investigated the role of water availability in the germination traits of keystone species in an arid ecosystem with stochastic rainfall. We measured seed germination responses of five arid species, along gradients of temperature and water potential under controlled laboratory conditions. We then identified the cardinal temperatures and base water potentials for seed germination, and applied the hydrotime model to assess germination responses to water stress. Optimum temperatures for germination ranged from 15 to 31 °C under saturated conditions (0 MPa), and three species had low minimum temperatures for germination (<3 °C). A small proportion of seeds of all species germinated under dry conditions (Ψ ≤ -1 MPa), although base water potential for germination (Ψ b50) ranged from -0.61 to -0.79 MPa. Species adhered to one of two germination traits: (i) the risk-takers which require less moisture availability for germination, and which can germinate over a wider range of temperatures irrespective of water availability (Casuarina pauper and Maireana pyramidata), and (ii) the risk-avoiders which have greater moisture requirements, a preference for cold climate germination, and narrower temperature ranges for germination when water availability is low (Atriplex rhagodioides, Maireana sedifolia and Hakea leucoptera). High seed longevity under physiological stress in H. leucoptera, combined with a risk-avoiding strategy, allows bet-hedging. The hydrotime model predicted lower base water potentials for germination than observed by the data, further supporting our assertion that these species have particular adaptations to avoid germination during drought. This study provides insights into the complex physiological responses of seeds to environmental stress, and relates seed germination traits to community dynamics and restoration in arid zones.