Mark K. J. Ooi

Seed bank persistence and climate change

The strong mechanistic relationship between climatic factors and seed dormancy and germination suggests that forecast climatic changes will significantly affect seed bank persistence. This review focuses on the potential impact of changing temperature, rainfall and fire regimes on the longevity of long-term persistent seed-banks. Currently, there are few studies investigating the mechanistic responses of demographic processes, such as seed-bank dynamics, to forecast climate change. However, from the work that has been published, several key points have been highlighted. First, increased air temperatures will produce significantly higher soil temperatures in open and sparsely vegetated habitats. Some evidence shows that this could accelerate the decline of seed viability and compromise bet-hedging strategies of species in dryland regions. Second, changes to rainfall season may determine the relative success of recruitment, with lower levels of success producing net losses to seed bank longevity. Finally, higher temperatures are likely to produce increased fire frequency, compromising the persistence of plant populations dependent on long-lived seed banks. Improving our understanding of both the mechanistic response and adaptive capacity of seed banks to climate change will provide a solid basis for improved predictions of future species distributions and risk of extinction, particularly in ecosystems subjected to temporally stochastic disturbances. It is necessary to develop functional groups based on key life-history trait responses to changing environmental conditions, to enable broader-scale predictions of distribution and persistence in the future.

Persistence of obligate-seeding species at the population scale: effects of fire intensity, fire patchiness and long fire-free intervals

Understanding how a species persists under a particular fire regime requires knowledge of the response to fire of individual plants. However, categorising the fire response of a species solely based on known responses of individual plants can be misleading when predicting a population response. In the present study, we sought to determine the fire responses of several Leucopogon species at the population level, including the threatened L. exolasius. We found that, whilst all species studied were obligate seeders, the population responses of species to fire were dependent upon fire intensity and patchiness. Results showed first that low intensity fires were significantly patchier than higher intensity fires. Second, the proportion of plants killed within a population decreased with increased fire patchiness. We also assessed how populations were structured and found that stands were multi-aged at most sites, and did not have a single-aged structure, which is often assumed for obligate seeders. Both spatial complexity within the fire regime leading to adult plant persistence, and inter-fire recruitment, contributed to the multi-aged structure. It is possible that these Leucopogon species are gap recruiters, and may tolerate fire rather than be specifically adapted to it. Inter-fire recruitment may enable L. exolasius populations to persist for a much longer fire-free period than many other species in the region.

Projected soil temperature increase and seed dormancy response along an altitudinal gradient: implications for seed bank persistence under climate change

Background and aimsUnderstanding the mechanistic effects of climate change on species key life-history stages is essential for predicting ecological responses. In fire-prone regions, long-term seed banks allow post-fire recovery and persistence of plant populations. For physically dormant species, seed bank longevity depends on the maintenance of dormancy which is controlled primarily by temperature. Successful inter-fire recruitment is rare and dormancy loss between fires produces a net loss to the seed bank. We assessed whether temperature increases related to climate change can affect seed dormancy and, potentially, seed bank longevity.MethodsWe quantified the relationship between air temperatures and soil temperatures. Seeds of two shrub species, from four populations along an altitudinal gradient, were then exposed to a range of soil temperatures calculated to occur at the end of the 21st century, using projected mean and heat wave scenarios. Alterations to dormancy were assessed via germination.ResultsFor every 1°C increase in air temperature, associated soil temperature increased by 1.5°C. Mean temperature increase had no affect on seed dormancy. However, future heat wave conditions produced soil temperatures that significantly increased dormancy loss. This impact was greatest in seeds from cooler, high elevation populations.ConclusionsProjected heat wave events produce conditions that provide a mechanism for seed bank compromise. Dormancy-breaking temperatures for each population were positively related to parental environment temperatures, indicating local adaptation. Whilst heat from fire may govern post-fire recruitment response, we suggest that parental climate is the key selective force determining dormancy-breaking threshold temperatures, ensuring inter-fire seed bank persistence.

Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change

Variation in dormancy thresholds among species is rarely studied but may provide a basis to better understand the mechanisms controlling population persistence. Incorporating dormancy-breaking temperature thresholds into existing trait frameworks could improve predictions regarding seed bank persistence, and subsequently species resilience in response to fire, climate change and anthropogenic management. A key ecological strategy for many species from fire-prone ecosystems is the possession of a long-lived seed bank, ensuring recovery after fire. Physical dormancy is dominant in these ecosystems and maintaining this dormancy is directly linked to seed bank persistence. We identified a suite of seed-related factors relevant to maintaining populations in fire-prone regions for 14 co-occurring physically dormant species. We measured variation in initial levels of dormancy and then applied experimental heating treatments, based on current seasonal temperatures and those occurring during fires, to seeds of all study species. Additionally, higher seasonal temperature treatments were applied to assess response of seeds to temperatures projected under future climate scenarios. Levels of germination response and mortality were determined to assess how tightly germination response was bound to either fire or seasonal cues. Six species were found to have dormancy cues bound to temperatures that only occur during fires (80°C and above) and were grouped as having obligate pyrogenic dormancy release. The remaining species, classified as having facultative pyrogenic dormancy, had lower temperature dormancy thresholds and committed at least 30% of seeds to germinate after summer-temperature treatments. Evidence from this study supports including dormancy-breaking temperature thresholds as an attribute for identifying functional types. High temperature thresholds for breaking dormancy, found in our obligate pyrogenic group, appear to be a fire-adapted trait, while we predict that species in the facultative group are most at risk to increased seed bank decay resulting from elevated soil temperatures under projected climate change.

To germinate or not to germinate: more than just a question of dormancy

Consider the following four quotations concerning the distinction between breaking dormancy and stimulating germination.

Delayed post-fire seedling emergence linked to season: a case study with Leucopogon species (Epacridaceae)

In many fire-prone habitats around the world, natural fire regimes have shaped the evolution of the associated flora. Critical life history stages are often linked to fire in species that occur in these fire-prone habitats but many species are unstudied. We investigated seedling emergence patterns over time, after several fires in south-eastern Australia, for three obligate seeders in the Genus Leucopogon (Family Epacridaceae): L. setiger, L. esquamatus and L. exolasius. Fixed quadrats were monitored for 12 to 30 months for newly emerged seedlings, both after fire and in unburnt L. exolasius and L. esquamatus habitats. There was a flush of seedling emergence in the first year after fire for all three Leucopogon species, with smaller pulses recorded in subsequent years. Time elapsed between fire and the onset of emergence differed between fires, but not between Leucopogon species. Whatever the timing of the fire, seedling emergence was restricted to the late autumn and winter periods, coinciding with emergence in unburnt habitat. This contrasts with patterns of emergence previously reported for other taxa, and also in this study, where emergence of the Leucopogon species after fire was delayed compared with co-occurring species in other genera. Our results suggest that seasonal factors are important to the germination ecology of these species and that combinations of fire-related and seasonal factors are necessary to maximise germination. Rainfall has a non-seasonal pattern in the study region and seasonal emergence post-fire has not previously been recorded. The magnitude of delay to emergence of species with seasonal emergence patterns will be determined by the season of fire but not by a seasonal pattern of rainfall in the post-fire year. A shift of the peak fire season could increase this delay, possibly affecting plant population recovery.

Sensitivity analyses of decision rules in World Conservation Union (IUCN) Red List criteria using Australian plants

Abstract The World Conservation Union (IUCN) has developed quantitative criteria for assessing the conservation status of species. The criteria comprise five decision rules that specify threshold values in rates of decline, distributional range size, population size and risk of extinction. The criteria are currently under review to identify and resolve difficulties in their application. Four issues raised in the review include (i) the effect of skewed size distributions on the assessment of the number of populations or locations, (ii) the effect of varying scales on measurement and assessment of distributional range, (iii) structural parity between the range and population criteria (B and C), and (iv) the treatment of missing data. Unless resolved, these issues may result in biases in the outcomes of Red List assessments. We applied sensitivity analyses to explore the nature of these biases and examined how they might be overcome with relatively minor modifications to the criteria. We applied these analyses to a data set comprising 135 rare or threatened vascular plant species from south-eastern Australia. The analyses suggest that biases may be reduced by: excluding very small locations and subpopulations from consideration; adopting a scale for area of occupancy thresholds that is commensurate with a biologically appropriate scale for measurement; implementing a common structure of decision rules in range and population criteria to minimise differences in the way continuing declines, fragmentation and fluctuations are assessed; and adopting a precautionary protocol for invoking ‘Data Deficient’ status when assessments based on only one or few criteria fail to identify threatened status.

Effects of soil temperature regimes after fire on seed dormancy and germination in six Australian Fabaceae species

In addition to direct fire cues such as heat, smoke and charred wood, the passage of fire leads indirectly to changes in environmental conditions which may be able to break physical dormancy in hard-coated seeds. After a fire, the open canopy and the burnt material lying on the surface alter the thermal properties of the soil, resulting in elevated soil temperatures for long periods of time. We simulated daily temperature regimes experienced at different depths of soil profile after a summer fire. Our aim was to determine whether these temperature regimes and the duration of exposure (5, 15 and 30 days) play an important role breaking physical seed dormancy in six legumes from south-eastern Australia. Our results showed that simulated temperature regimes break seed dormancy. This effect is specially pronounced at temperatures that are expected to occur near the soil surface (0–2 cm depth). The duration of exposure interacts with temperature to break dormancy, with the highest germination rates reached after the longest duration and highest temperatures. However, the germination response varied among species. Therefore, this indirect post-fire cue could play a role in the regeneration of plant communities, and could stimulate seedling emergence independent of direct fire cues as well as in interaction with direct cues.

Distinguishing between persistence and dormancy in soil seed banks of three shrub species from fire-prone southeastern Australia

Question: Is primary dormancy required for seed bank persistence of Leucopogon species? How does the distinction between dormancy and persistence affect our understanding of seed bank dynamics in fire-prone regions? Location: Sclerophyllous plant communities, Sydney region, southeastern Australia. Methods: Seed bank longevity of three morphophysiologically dormant species was ascertained using seed burial trials. Seeds of each species were buried in situ in replicate mesh bags and retrieved annually over three years. Laboratory germination trials and embryo growth measurements over time were used to establish in which order each component of dormancy was overcome, and whether seed banks persisted after the loss of primary dormancy. Results: All species had long-term persistent seed banks, with estimated half-lives between 3.5 and 5.5 years. Physiological dormancy was broken by stratification prior to embryo growth, during the first year of burial. Seeds were able to germinate after retrieval, in light and at warm temperatures. Only low levels of germination occurred in situ over three years of burial, and did not increase over time. During germination trials, embryos remained underdeveloped unless seeds germinated, indicating that seeds were committed to germination once embryo growth was initiated. Conclusions: Long-lived seed banks are an important trait of plants from fire-prone communities, but cues observed to promote germination in situ are not necessarily the ones that break dormancy. A failure to distinguish between these two factors can divert attention away from mechanisms actually controlling dormancy and seed bank dynamics.

Physical dormancy in a changing climate

Species with physically dormant (PY) seeds make up over 25% of plant species in a number of ecologically important ecosystems around the globe, such as savannah and Mediterranean shrublands. Many of these ecosystems are subject to temporally stochastic events, such as fire and drought; but are in areas projected to experience some of the most extreme climatic changes in the future. Given the importance of PY in controlling germination timing for successful recruitment, we ask how plastic the PY trait is, and if changes to the maternal environment from climate change could alter recruitment. This review focuses on: (1) the evidence for inter- and intraspecific variation in PY; (2) the genetic, maternal and environmental controls involved; and (3) the ecological consequences of (1) and (2) above. Evidence for (within-community) interspecific variation in conditions required to break PY is strong, but for intraspecific variation evidence is contradictory and limited by a paucity of studies. Identifying controllers of variation in PY is complex, there is some suggestion that conditions of the maternal environment may be important, but no consensus on the nature of effects. The implications of PY plasticity for the persistence of seed banks, species and communities under climate change are discussed. We highlight a number of key knowledge gaps, such as a lack of research estimating the components of variation in non-agricultural species, and identify a suite of seed attributes relevant to understanding the potential impacts of climate change on the population dynamics of PY species in the future.