Anatomic basis and insulation of serotinous cones in Pinus halepensis Mill

Pinus halepensis Mill., a widespread, low elevation conifer common in Mediterranean Basin, shows a dual reproductive strategy: post-fire obligate seeder (from serotinous cones) and an early coloniser (from non-serotinous cones). Release of seeds encased in serotinous cones is induced either by fire (pyriscence, serotiny or bradychory) or by drying (xeriscence). Morphological differences in serotinous and non-serotinous cones in natural populations of P. halepensis in Southeastern Italy were analyzed. Relationships between tree size (diameter class) and serotiny were checked by counting and sampling serotinous and non-serotinous cones. The macro and microscopic characteristics that could affect cones’ opening were measured in sampled cones. Protection against high temperatures offered by wood scales was also evaluated by applying different temperatures and time exposures, and following the inner thermal raise. Results showed that non-serotinous cones had bigger resin ducts and more separate scales. Also it was highlighted that ovuliferous scales of serotinous cones were bigger and thicker. These scales had more lamellated (multilayered) sclereid cells, and were significantly thinner with a shorter lumen diameter. Continuous temperature-monitoring heat tests inside cones showed that temperatures close to the cone axis were rather low, so seed germination was not influenced. Results confirm that serotinous cones are more compact, rigid and consistent than non-serotinous cones. These characteristics explain the lower insulation, seed protection and the ease opening of non-serotinous cones as well. In conclusion, opening mechanism of pinecone scales under the effect of fire or dry conditions seem related to anatomic differences and it provides seeds with an efficient protection against heat.     

Germination response to fire-related factors of seeds from non-serotinous and serotinous cones

Pinus halepensis, a Mediterranean pine tree, is a partially serotinous species: individual trees of this species carry both non-serotinous and serotinous cones. Serotinous cones open mainly after fire, whereas non-serotinous cones open in absence of fire. In this study we addressed the question, whether or not this cone response is linked with the germination response of seeds to fires. Two main factors associated with fire are heating of seeds and soil pH. A combination of high heat and high pH simulates a scenario with fire, whereas low heat and low pH simulates a scenario without fire. We assessed the separate and combined effects of heat and pH on the germination rate and the percentage of germination of seeds from non-serotinous cones and two age classes of serotinous cones of P. halepensis. Heat had no effect on the percentage of germination of seeds from any of the cone types, but did positively affect the germination rates of seeds from both age-classes of serotinous cones. High pH negatively affected the germination rate of seeds from all cone types as well as the percentage of germination of seeds from non-serotinous cones. The combinations of heat and pH had different effects on the three cone types: percentage of germination and rate of germination of seeds from non-serotinous cones was higher in the combination high heat-high pH than in the combination low heat-low pH. In the combination high heat-high pH, seeds from serotinous cones germinated better than seeds from non-serotinous cones. The different germination responses of seeds from non-serotinous and serotinous cones could not be attributed to differences in cone age. Our results indicate that the cone response is linked to the germination response of the seeds in P. halepensis, with seeds from serotinous cones being more tolerant to fire related factors. This revised version was published online in June 2006 with corrections to the Cover Date.     

Canopy seed bank structure in relation to: fire,tree size and density

To assess the canopy seed bank structure of Pinus halepensis, we measured the level of serotiny and the seed bank size and density of trees in unburned stands and post-fire regenerated stands in Israel. We analysed the effects of tree size, tree density and fire history on the level of serotiny. The level of serotiny decreased with an increase in tree height. The high level of serotiny in short trees could be explained by selection to increase regeneration chances after burning at pre-mature age. Also, limitation of long-distance seed dispersal opportunities in short trees may favour high serotiny levels. The level of serotiny was higher in post-fire stands than in unburned stands, suggesting a fast selection for serotiny by fire. Unburned stands had a higher total stand seed density than post-fire regenerated stands, but the proportion of seeds in serotinous cones of the total stand seed density was higher in post-fire regenerated stands. The fact that P. halepensis bears simultaneously serotinous and non-serotinous cones reflects its dual strategy as both a post-fire obligate seeder, mainly from serotinous cones and an early coloniser during fire-free periods, mainly from non-serotinous cones. The relative investment in these strategies is dependent on fire history and varies with tree height. Furthermore, mature brown cones can contribute to post-fire regeneration in case of spring fires, and serotinous cones are known to open partially also in dry spell events. Thus, post-fire regeneration and invasion are strategies, which seem to complement each other.     

Enhanced seed defenses potentially relax selection by seed predators against serotiny in lodgepole pine

Serotiny, the retention of seeds in a canopy seed bank until high temperatures cause seeds to be released, is an important life history trait for many woody plants in fire‐prone habitats. Serotiny provides a competitive advantage after fire but increases vulnerability to predispersal seed predation, due to the seeds being retained in clusters in predictable locations for extended periods. This creates opposing selection pressures. Serotiny is favored in areas of high fire frequency, but is selected against by predispersal seed predators. However, predation also selects for cone traits associated with seed defense that could reduce predation on serotinous cones and thereby relax selection against serotiny. This helps explain the elevated defenses in highly serotinous species. However, whether such interactions drive variation in seed defenses within variably serotinous populations has been studied rarely. We investigated the effects of phenotypic selection exerted by red squirrel (Tamiasciurus hudsonicus) predation on Rocky Mountain lodgepole pine (Pinus contorta latifolia) seeds. Squirrels preferentially harvested cones with more and larger seeds, indicating a preference for a higher food reward. We found evidence for stronger selection on trees with serotinous cones, which presumably accounts for the elevated defenses of and lower predation on serotinous compared to non‐serotinous cones. Lower levels of predation on serotinous cones in turn lessen selection against serotiny by squirrels. This has important implications because the frequency of serotiny in lodgepole pine has profound consequences for post‐fire communities and ecosystems widespread in the Rocky Mountains.     

Fruit-seed relations in Hakea: serotinous species invest more dry matter in predispersal seed protection

Abstract The ability to maintain a canopy stored seedbank (serotiny) is characteristic of many woody genera inhabiting fire-prone environments. The relationship between level of serotiny for 94 Hakea (Proteaceae) species in southwestern Australia and follicle mass, density, three wall thicknesses and seed mass was investigated. Two species were non-serotinous (fruits open at maturity), 12 were weakly serotinous (majority of fruits open at maturity), 9 were moderately serotinous (fruits open within five years of maturity) and 71 were strongly serotinous (fruits still closed at least five years after reaching maturity). A positive relationship existed between the level of serotiny and follicle morphology. Strongly serotinous species were more likely to have heavier, woodier and thicker-walled follicles than non- and weakly serotinous species. Moderately/strongly serotinous species invested more energy (six times higher follicle:seed mass ratio) than non weakly serotinous species, consistent with increased protection of the seeds from granivores, pathogens, desiccation and/or heat. Recent work has shown that thicker fruit walls (strongly serotinous species) provide better insulation to seeds from heat, although the need to survive fire is just as critical for thin-walled, weakly serotinous species. Greater protection from granivores may provide a better explanation for the adaptive significance of dense, thick-walled serotinous follicles, as the opportunities for predispersal granivore damage are low among weakly serotinous species.     

Non‐fire induced seed release ina weakly serotinous pine: climatic factors,maintenance costs or both?

The advantages of canopy seed retention (serotiny) for plants inhabiting fire‐prone ecosystems are well documented. However, very few species are completely serotinous and non‐fire induced opening of serotinous fruits is commonly observed (weak serotiny). Two non‐mutually exclusive causes are envisaged to contribute to this process: mechanical changes in serotinous fruits mediated by climatic conditions (e.g. drought) or the costs of maintenance for the plant of these long‐lasting structures. However, their relative contribution to the spontaneous opening of serotinous fruits remains elusive as well as the consequences for the build‐up of the canopy seed bank and inter‐individual differences in serotiny. In this study we monitored the dynamics of cone production and cone opening in the weakly serotinous Pinus halepensis for five years (2004–2008), including two severe drought episodes (2005, 2006). Drought decreased the production of conelets, increased the abortion of immature cones, reduced the seed quality in the cohorts of cones produced during these years, and increased the opening of serotinous cones. During the first drought episode, a higher proportion of serotinous cones opened in those pines bearing a larger crop of younger cones. This suggests that not only passive changes induced by drought but also competition among cones for resources (e.g. water) might be involved in this process. The opening of serotinous cones in pines bearing more cones made inter‐individual differences in the size of the canopy cone bank to narrow or even to reverse from 2004 to 2008. These results may help to understand the decrease in serotiny when pines grow and accumulate more cones and the large inter‐individual variability in the degree of serotiny observed in P. halepensis forests. In addition, the negative effects of drought episodes for the size of the canopy cone bank and the seeds contained can be an unexplored cause of post‐fire regeneration constraint.     

Evidence for physiological seed dormancy cycling in the woody shrub Asterolasia buxifolia and its ecological significance in fire‐prone systems

• Dormancy cycling is a key mechanism that contributes to the maintenance of long‐term persistent soil seed banks, but has not been recorded in long‐lived woody shrub species from fire‐prone environments. Such species rely on seed banks and dormancy break as important processes for post‐fire recruitment and recovery.
• We used germination experiments with smoke treatments on fresh seeds and those buried for 1 year (retrieved in spring) and 1.5 years (retrieved the following late autumn) to investigate whether Asterolasia buxifolia, a shrub from fire‐prone south‐eastern Australia with physiologically dormant seeds, exhibited dormancy cycling.
• All seeds had an obligation for winter seasonal temperatures and smoke to promote germination, even after ageing in the soil. A high proportion of germination was recorded from fresh seeds. but germination after the first retrieval was significantly lower, despite high seed viability. After the second retrieval, germination returned to the initial level. This indicates a pattern of annual dormancy cycling; one of the few observations, to our knowledge, for a perennial species. Additionally, A. buxifolia’s winter temperature and smoke requirements did not change over time, highlighting the potential for seeds to remain conditionally dormant (i.e. restricted to a narrow range of germination conditions) for long periods.
• For physiologically dormant species, such as A. buxifolia, we conclude that dormancy cycling is an important driver of successful regeneration, allowing seed bank persistence, sometimes for decades, during fire‐free periods unsuitable for successful recruitment, while ensuring that a large proportion of seeds are available for recruitment when a fire occurs.

     

Long‐term livestock exclusion facilitates native woody plant encroachment in a sandy semiarid rangeland

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Life histories of Mediterranean pines

The life history of Spanish pines and their relation to fire as the main disturbance factor in their ecosystems was analysed. The primary ecological attributes studied were the canopy seed bank (onset of cone production, percentage and persistence of serotinous cones), seed and cone morphology, sprouting and bark thickness. Four ecological groups were separated using multivariate cluster analysis and their life-history characteristics are discussed. Serotiny and early flowering in Pinus halepensis and P. pinaster reflect their evader strategy in relation to fire as this character is advantageous to survive frequent crown fires and to attain successful post-fire recruitment. Late flowering and absence of serotinous cones in P. nigra, P. sylvestris and P. uncinata indicate that their natural forest did not evolve under frequent crown fires. P. canariensis and P. pinea appeared in two single groups because of their sprouting capability and their seed size respectively. Intraspecific variation in P. pinaster was also analysed using the same criteria and high variability was found in its life history traits. A group of P. pinaster populations showed high levels of serotiny and thin bark as a possible adaptation to frequent stand-replacing crown fires. In contrast, a group of non- or weakly-serotinous populations seems to have evolved under a low-intensity fire regime where the best fitness corresponds to thick-barked individuals capable of surviving ground fires. Intermediate strategies were also evident in this species and were discussed in relation to the effect of different fire regimes caused by the understorey vegetation.     

Variation in serotiny of three Banksia species along a climatic gradient

The degree of serotiny (i.e. the proportion of follicles remaining closed in each year's crop of cones since the last fire) was measured in Bank-sia attenuata, B. menziesii and B. prionotes at five sites along a climatic gradient extending 500 km north of Perth, Western Australia. The decrease in annual rainfall and increase in average temperature along the gradient paralleled a decrease in plant height and an increase in the degree of serotiny of all species. Extreme serotiny was recorded in the scrub-heath at the xeric end of the gradient whereas two species were essentially non-serotinous in the low woodland at the most mesic site. It is concluded that degree of serotiny is related to the fire characteristics of the site which depend on plant height. In xeric scrub-heath, the entire canopies of the Banksia spp. are consumed by fire which promotes massive release of seed. This facilitates recruitment in an otherwise unpredictable and unreliable seedbed. In mesic woodland, where cones rarely come into contact With flames, seeds are released spontaneously and site conditions are more conducive to recruitment in the inter-fire period.     

Fire severity drives variation in post-fire recruitment and residual seed bank size of <Emphasis Type="Italic">Acacia</Emphasis> species

Very high-severity fires are a component of many fire-prone ecosystems, yet are often viewed as detrimental to vegetation. However, species in such systems are likely to have adapted to persist under a fire regime that includes high-severity fires. We examined how fire severity affects post-fire recruitment and residual seed banks of Acacia species and whether severity may affect plant responses to fire intervals. Nine sites of either high or low burn severity were identified after a large-scale mixed-severity fire in Warrumbungle National Park, south-eastern Australia. Transects were used to sample above-ground woody plant density. Seed bank size was surveyed by soil extraction from two depths and manual searching for seeds. Residual soil seed bank and recruitment were compared across the two burn severities. Acacia seedling density was higher in areas burnt at high severity, indicating that increased severity triggers increased germination from the seed bank. Size of residual seed bank was smaller after high-severity fire, but varied between species, with few Acacia cheelii seeds remaining despite high above-ground abundance. In contrast, A. penninervis retained a small residual seed bank. There was little evidence of negative effects on populations of Acacia species after high-severity burns. However, we found that high fire severity may impact on the ability of a species to persist in response to a subsequent short fire interval. Fire management for maintaining biodiversity needs to consider other key aspects of the fire regime, including severity and season, rather than focusing solely on fire frequency.     

Increased canopy seed-storage in post-fire pine invaders suggests rapid selection mediated by fire

Exotic species storing seeds in the canopy (serotinous species) can experience a clear advantage in fire-prone communities that lack native taxa with such fire-resistant traits. In addition, selection in the new environment can potentially increase the frequency of fire-adapted characteristics such as serotiny. We studied the potential role of fire favoring the serotinous, non-native conifer Pinus radiata in NW Patagonia. We characterized the degree of serotiny (percentage of serotinous cones) and the size of the canopy seed bank in the unburned plantation and in stands of trees recruited after a fire 30 years ago as a proxy for invasion potential. Fire had a positive effect, increasing serotiny in post-fire P. radiata stands. Post-fire recruited cohorts showed higher serotiny levels and a larger canopy seed bank compared with plantations. Our study suggests that fire-linked traits like serotiny may be subjected to a rapid, fire-driven selection process in fire-adapted species such as P. radiata invading fire-prone ecosystems. Thus, increased serotiny can lead to higher postfire invasion densities, which in turn create a positive feedback loop in invaded areas under recurrent fires.     

The Effects of Seed Ingestion by Livestock,Dung Fertilization,Trampling, Grass Competition and Fire on Seedling Establishment of Two Woody Plant Species

The increasing rate of woody plant encroachment in grasslands or savannas remains a challenge to livestock farmers. The causes and control measures of woody plant encroachment are of common interest, especially where it negatively affects the objectives of an agricultural enterprise. The objectives of this study were to determine the effects of gut passage (goats, cattle), dung (nutrients), fire, grass competition and trampling on establishment of A. nilotica and D. cinerea seedlings. Germination trials were subjected to the following treatments: 1) seed passage through the gut of cattle and goats and unpassed/ untreated seeds (i.e. not ingested), 2) dung and control (no dung), 3) grass and control (mowed grass), 4) fire and control (no fire), 5) trampling and control (no trampling). The interaction of animal species, grass and fire had an effect on seedling recruitment (P < 0.0052). Seeds retrieved from goats and planted with no grass and with fire (6.81% ± 0.33) had a significant effect on seedling recruitment than seeds retrieved from goats and planted with grass and no fire (2.98% ± 0.33). Significantly more D. cinerea and A. nilotica seeds germinated following seed ingestion by goats (3.59% ± 0.16) than cattle (1.93% ± 0.09) and control or untreated seeds (1.69% ± 0.11). Less dense grass cover, which resulted in reduced grass competition with tree seedlings for light, space and water, and improved seed scarification due to gut passage were vital for emergence and recruitment of Acacia seedlings. These results will contribute considerably to the understanding of the recruitment phase of woody plant encroachment.     

Variation in seed packaging of a fleshy‐fruited conifer provides insights into the ecology and evolution of multi‐seeded fruits

• The study of intraspecific seed packaging (i.e. seed size/number strategy) variation across different populations may allow better understanding of the ecological forces that drive seed evolution in plants. Juniperus thurifera (Cupressaceae) provides a good model to study this due to the existence of two subspecies differentiated by phenotypic traits, such as seed size and cone seediness (number of seeds inside a cone), across its range.
• The aim of this study was to analyse seed packaging (seed mass and cone seediness) variation at different scales (subspecies, populations and individuals) and the relationship between cone and seed traits in European and African J. thurifera populations.
• After opening more than 5300 cones and measuring 3600 seeds, we found that seed packaging traits followed different patterns of variation. Large‐scale effects (region and population) significantly contributed to cone seediness variance, while most of the seed mass variance occurred within individuals. Seed packaging differed between the two sides of the Mediterranean Sea, with African cones bearing fewer but larger seeds than the European ones. However, no differences in seed mass were found between populations when taking into account cone seediness. Larger cones contained more pulp and seeds and displayed a larger variation in individual seed mass.
• We validated previous reports on the intraspecific differences in J. thurifera seed packaging, although both subspecies followed the same seed size/number trade‐off. The higher seediness and variation in seed mass found in larger cones reveals that the positive relationship between seed and cone sizes may not be straightforward.We hypothesise that the large variation of seed size found within cones and individuals in J. thurifera, but also in other fleshy‐fruited species, could represent a bet‐hedging strategy for dispersal.

     

Tanned or Burned: The Role of Fire in Shaping Physical Seed Dormancy

Plant species with physical seed dormancy are common in mediterranean fire-prone ecosystems. Because fire breaks seed dormancy and enhances the recruitment of many species, this trait might be considered adaptive in fire-prone environments. However, to what extent the temperature thresholds that break physical seed dormancy have been shaped by fire (i.e., for post-fire recruitment) or by summer temperatures in the bare soil (i.e., for recruitment in fire-independent gaps) remains unknown. Our hypothesis is that the temperature thresholds that break physical seed dormancy have been shaped by fire and thus we predict higher dormancy lost in response to fire than in response to summer temperatures. We tested this hypothesis in six woody species with physical seed dormancy occurring in fire-prone areas across the Mediterranean Basin. Seeds from different populations of each species were subject to heat treatments simulating fire (i.e., a single high temperature peak of 100°C, 120°C or 150°C for 5 minutes) and heat treatments simulating summer (i.e., temperature fluctuations; 30 daily cycles of 3 hours at 31°C, 4 hours at 43°C, 3 hours at 33°C and 14 hours at 18°C). Fire treatments broke dormancy and stimulated germination in all populations of all species. In contrast, summer treatments had no effect over the seed dormancy for most species and only enhanced the germination in Ulex parviflorus, although less than the fire treatments. Our results suggest that in Mediterranean species with physical dormancy, the temperature thresholds necessary to trigger seed germination are better explained as a response to fire than as a response to summer temperatures. The high level of dormancy release by the heat produced by fire might enforce most recruitment to be capitalized into a single post-fire pulse when the most favorable conditions occur. This supports the important role of fire in shaping seed traits.     

Demography of a non-sprouting and resprouting Hakea species (Proteaceae) in fire-prone Eucalyptus woodlands of southeastern Australia in relation to stand age,drought and disease

Population size-structures, seed production, canopy seed storage (serotiny), and recruitment were investigated in relation to fire, drought and disease for a pair of co-occurring resprouting and non-sprouting shrub species from the genus Hakea (Proteaceae) in fire-prone Eucalyptus woodlands in western Victoria, Australia. The non-sprouter species, Hakea decurrens, showed faster height growth, higher seed production and higher seed viability than the resprouter, Hakea rostrata. Population size structures in stands up to 24 years since last fire showed no evidence of inter-fire recruitment for either species. Following a fire in 1990 in a mixed species stand 15–20 years old, the estimated number of viable seeds released from canopy-stored seed banks was approximately equal for both species. However, the rate of seedling establishment in the first year was about 10 times higher, and seedling suvivorship over the first 5 years was seven times higher, for the non-sprouter. Seedlings of Hakea decurrens and resprouts of Hakea rostrata began to produce seeds within three years of the last fire, while the few surviving seedlings of Hakea rostrata showed no evidence of reproductive maturity after six years. Inter-fire recruitment was recorded for the non-sprouter, Hakea decurrens, in the oldest stand (burned in 1967) between 24 and 28 years since last fire. This was associated with an increased rate of seed release from serotinous fruits due to the onset of high rates of adult plant mortality. High adult mortality and increased seed release correlated with increasing stand age, the occurrence of severe drought, and the likely presence of Phytophthora cinnamomi, a fungal pathogen which damages the root system, reducing water and nutrient uptake. There were no new recruits for the resprouter Hakea rostrata in this stand, but old plants continued to resprout from basal lignotubers and no mortality was observed. While recruitment of strongly serotinous shrub species is commonly described as being restricted to the immediate post-fire period, the present study illustrates that other events (e.g., senescence, drought, disease) can lead to recruitment of serotinous non-sprouters and may be important in the maintenance of populations during unusually long periods without fire.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

Predator satiation and recruitment in a mast fruiting monocarpic forest herb

Background and Aims

Cross-pollination and satiation of seed predators are often invoked to explain synchronous mast reproduction in long-lived plants. However, explanations for the synchronous death of parent plants are elusive. The roles of synchronous seeding and post-reproductive mortality were investigated in a perennial monocarpic herb (Isoglossa woodii) in coastal dune forest in South Africa.

Methods

Pre-dispersal seed predation and seed production were assessed by measuring fruit and seed set of inflorescences sprayed with insecticide or water and with no spray treatments. Seed predation was measured at different densities of I. woodii plants by monitoring removal rates of seed from the forest floor. The influence of adult plants on establishment of I. woodii seedlings was assessed by monitoring growth and survivorship of seedlings in caged and uncaged 1 × 1 m plots in understorey gaps and thickets.

Key Results

Fruit and seed set were similar between spray treatments. An I. woodii stem produced 767·8 ± 160·8 seeds (mean ± s.e.) on dune crests and 1359·0 ± 234·4 seeds on the foredune. Seed rain was greater on the foredune than in other topographic locations. Seed predation rates were 32 and 54 % on dune crests and in dune slacks, respectively, and decreased with seed abundance, number of inflorescences per stem and plant height. Seedling recruitment was greater beneath synchronously dying adult plants than in natural understorey gaps (no I. woodii). However, seedling growth rate beneath I. woodii mid-way through its life-cycle was less than in gaps, although survivorship was similar.

Conclusions

The selective advantage of masting in I. woodii derives from satiation of both pre- and post-dispersal seed predators. In addition, post-seeding mortality of adult plants facilitates seedling establishment. Satiation of seed predators and the benefits of seedling establishment are strong drivers of the evolution of synchronous monocarpy in I. woodii.     

Fire and competition in Australian heath: a conceptual model and field investigations

Abstract. We describe a model of heath vegetation, in which species were classified into five functional groups based on characteristics of their propagule pools, post-fire growth, timing and mode of reproduction and competitive status. The model assumes no recruitment without fire and a simple competitive hierarchy based on vertical stature. A critical feature of the model is an initial post-fire window of 5–6 yr in which competition from overstorey species on understorey species is reduced. Understorey functional groups differ in their ability to exploit this window. In the field, we tested five predictions derived from the model: (a) overall species richness of understorey varies inversely with overstorey density as a result of a trend in richness of woody species, but not in herbaceous species; (b) where an overstorey was present in the previous fire interval, post-fire population density is reduced in a functional group of understorey serotinous resprouting shrubs, but not in a group of understorey obligate-seeding shrubs with soil seed banks; (c) in understorey serotinous resprouting shrubs, post-fire regrowth in resprouting individuals is adversely affected by the presence of an overstorey in the preceding fire interval; (d) in understorey serotinous resprouting shrubs, levels of pre-fire propagules are lower in the presence of an overstorey, reducing the density of post-fire recruits; and (e) in understorey serotinous resprouting shrubs, recruitment relative to the pre-fire population is unaffected by overstorey species within the window of reduced competition. Of these, three tests (a,b,d) supported the model, one (e) may support the model, but the results were inconclusive and one (c) did not support the model. Limitations and further applications of the model are discussed. Our results suggest that maintenance of high densities of overstorey populations is in conflict with conservation of some understorey species. Models of the type we propose will help identify and resolve such conflicts and promote the judicious use of fire to maintain full species diversity of plant communities.     

Bimodal colour pattern of individual Pinus halepensis Mill. seeds: a new type of crypsis

Variation in seed traits is a well‐known phenomenon affecting plant ecology and evolution. Here we describe, for the first time, a bimodal colour pattern of individual seeds, proposing an adaptive explanation, using Pinus halepensis as a model. Pinus halepensis disperses its seeds either by wind on hot dry days, from regular cones, or after fires, mainly from serotinous cones. Post‐dispersal seeds are exposed to strong predation by passerine birds, making crypsis important for seed survival. Individual seeds from non‐serotinous cones have a bimodal colour pattern: one side is light brown and the other black, exposing only one colour when lying on the ground. Serotinous cones from most trees have seeds with similar bimodal colour patterns, whereas seeds from serotinous cones of some trees are light brown on both sides. The dark side provides the seed with better crypsis on dark soils, whereas the light‐brown side is better adapted to light‐coloured soils, and mainly to light‐grey ash‐covered soil, which is the natural post‐fire regeneration niche of P. halepensis. The relative reflection curves of the black and brown seed colours differ, and their calculated relative chromatic distance is 5: meaning that seed‐predating passerine birds see them differently, and probably prefer seeds that present a higher contrast against the soil background. We propose that such a bimodal colour pattern of individual seeds is probably an overlooked general phenomenon mainly linked to seed dispersal in post‐fire and other heterogeneous environments. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 271–278.