Heritability and quantitative genetic divergence of serotiny,a fire-persistence plant trait

Background and Aims

Although it is well known that fire acts as a selective pressure shaping plant phenotypes, there are no quantitative estimates of the heritability of any trait related to plant persistence under recurrent fires, such as serotiny. In this study, the heritability of serotiny in Pinus halepensis is calculated, and an evaluation is made as to whether fire has left a selection signature on the level of serotiny among populations by comparing the genetic divergence of serotiny with the expected divergence of neutral molecular markers (Q_ST–F_ST comparison).

Methods

A common garden of P. halepensis was used, located in inland Spain and composed of 145 open-pollinated families from 29 provenances covering the entire natural range of P. halepensis in the Iberian Peninsula and Balearic Islands. Narrow-sense heritability (h²) and quantitative genetic differentiation among populations for serotiny (Q_ST) were estimated by means of an ‘animal model’ fitted by Bayesian inference. In order to determine whether genetic differentiation for serotiny is the result of differential natural selection, Q_ST estimates for serotiny were compared with F_ST estimates obtained from allozyme data. Finally, a test was made of whether levels of serotiny in the different provenances were related to different fire regimes, using summer rainfall as a proxy for fire regime in each provenance.

Key Results

Serotiny showed a significant narrow-sense heritability (h²) of 0·20 (credible interval 0·09–0·40). Quantitative genetic differentiation among provenances for serotiny (Q_ST = 0·44) was significantly higher than expected under a neutral process (F_ST = 0·12), suggesting adaptive differentiation. A significant negative relationship was found between the serotiny level of trees in the common garden and summer rainfall of their provenance sites.

Conclusions

Serotiny is a heritable trait in P. halepensis, and selection acts on it, giving rise to contrasting serotiny levels among populations depending on the fire regime, and supporting the role of fire in generating genetic divergence for adaptive traits.     

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.     

The influence of cone age on the relative longevity of Banksia seeds

Background and Aims

Serotiny is common in the genus Banksia, so any seed collection is likely to be comprised of seeds that were produced in many different years. This study aimed to determine the impact of cone age and degree of serotiny on longevity in ex situ storage.

Methods

Cones of identifiable age classes were collected from three species of Banksia. Seeds were extracted from cones and the degree of serotiny calculated. An estimate of initial viability (K_i), the time for viability to fall by one probit (σ) and the relative longevity of seeds (p₅₀) for each species and cone age class was determined using a comparative longevity test (50 °C, 63 % relative humidity).

Key Results

The degree of serotiny ranged from moderate (7·9) for Banksia attenuata to strong (40·4) for B. hookeriana. Survival curves for all seed age classes within each species could be described by regressions with a common slope (1/σ), but with different values for K_i. The time taken for viability to fall by one probit (σ) could be described by a common value (29·1 d) for all three species.

Conclusions

Differences in seed longevity between cone age classes and species was related to variation in initial viability (K_i) rather than to differences in σ. While targeting the youngest mature seed cohort on a plant will maximize the viability of seeds collected, a wide range of age classes should be collected (but stored as separate cohorts if possible) for quality conservation/restoration seed collections where genetic diversity is important.     

Field heritability of a plant adaptation to fire in heterogeneous landscapes

The strong association observed between fire regimes and variation in plant adaptations to fire suggests a rapid response to fire as an agent of selection. It also suggests that fire‐related traits are heritable, a precondition for evolutionary change. One example is serotiny, the accumulation of seeds in unopened fruits or cones until the next fire, an important strategy for plant population persistence in fire‐prone ecosystems. Here, we evaluate the potential of this trait to respond to natural selection in its natural setting. For this, we use a SNP marker approach to estimate genetic variance and heritability of serotiny directly in the field for two Mediterranean pine species. Study populations were large and heterogeneous in climatic conditions and fire regime. We first estimated the realized relatedness among trees from genotypes, and then partitioned the phenotypic variance in serotiny using Bayesian animal models that incorporated environmental predictors. As expected, field heritability was smaller (around 0.10 for both species) than previous estimates under common garden conditions (0.20). An estimate on a subset of stands with more homogeneous environmental conditions was not different from that in the complete set of stands, suggesting that our models correctly captured the environmental variation at the spatial scale of the study. Our results highlight the importance of measuring quantitative genetic parameters in natural populations, where environmental heterogeneity is a critical aspect. The heritability of serotiny, although not high, combined with high phenotypic variance within populations, confirms the potential of this fire‐related trait for evolutionary change in the wild.     

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.     

Carbon and Nitrogen Dynamics of Boreal Jack Pine Stands With and Without a Green Alder Understory

We compared the species composition, structure and selected components of the carbon (C) and nitrogen (N) budgets of similar-aged, mature boreal jack pine (Pinus banksiana Lamb.) forests with and without green alder [Alnus crispa (Ait.) Pursh.] in two different boreal environments. The C and N content of the overstory biomass components (for example, stem, branch, and foliage), total vegetation, forest floor, and mineral soil were greater (P= 0.05 to P= 0.10) for jack pine with alder (JPA) stands than for jack pine without alder (JP) stands at both study areas. Jack pine foliage N isotopic discrimination (δ¹⁵N) and annual litterfall N content were significantly greater (P < 0.05) in the JPA than the JP stands at both study areas, suggesting that alder was fixing N and that N availability was greater in the JPA than the JP stands. The greater leaf area index (LAI) and overstory C accumulation in the JPA than the JP stands (P < 0.05) is likely because of the greater N availability in the JPA stands, but the effect of soil texture discontinuity on water availability in the JPA stands can not be dismissed. Percent ground cover by feathermoss varied among the jack pine communities and was positively correlated with overstory LAI (r ²= 0.83, P< 0.05). One index of N-use efficiency (NUE), defined as aboveground net primary productivity (ANPP) per litterfall N, was significantly greater (P < 0.05) for the JP than the JPA stands, but a second index of NUE, ANPP/N uptake, did not differ between the two jack pine communities. Jack pine trees growing without alder produced more organic matter per unit of N, but percent N retranslocation from senescing foliage and N mean residence time in the overstory did not differ between the JPA and the JP stands. A conceptual model is presented that illustrates the potential influence of alder on the species composition, structure, and function of boreal jack pine forests. Received 6 January 1998; accepted 15 April 1998.     

A review of factors affecting the population dynamics of jack pine budworm (Choristoneura pinus pinus Freeman)

Jack pine budworm (Choristoneura pinus pinus Free.) (Lepidoptera: Tortricidae) is a native insect that periodically defoliates areas of jack pine (Pinus banksiana Lamb.) in the subboreal forests of North America east of the Rocky Mountains. Outbreaks of jack pine budworm generally occur at 6- to 12-year intervals and collapse after 2–4 years. Periodicity of outbreaks varies and is associated with site-related factors. Survival of early-instar larvae during spring dispersal is tied to the abundance of pollen cones (microsporangiate strobili), which provide a refuge for larvae until current-year needles expand. Jack pine trees that have been heavily defoliated produce few pollen cones in the following year, often resulting in high mortality of early-stage larvae. A diverse guild of generalist parasitoids attack jack pine budworm, but only a few species account for most mortality in any area. Collapsing jack pine budworm populations are characterized by sharp declines in early instar survival, coupled with an increased rate of parasitism in the late larval and pupal stages. The reciprocal interaction between heavy defoliation and low pollen cone production, and increased parasitism of late-stage larvae or pupae, are consistent with second-order density dependence factors identified in analysis of a long-term population data set. Since the 1950s, several jack pine budworm outbreaks have been roughly synchronous over a large geographic area, suggesting that Moran effect processes, as well as moth dispersal or other factors, may be involved in jack pine budworm dynamics. Although the short duration of outbreaks enables most trees to recover, over time dead trees and top-killed trees accumulate in jack pine stands. Jack pine is well adapted to fire and when fires ignite, the accumulation of dead trees and woody debris often leads to intense wildfires followed by prolific regeneration. The three-way interaction of jack pine, jack pine budworm, and fire ultimately serves to maintain vigorous stands and ensures continued hosts for jack pine budworm. Received: October 1, 1999 / Accepted: September 22, 2000     

The northern limit of Pinus banksiana Lamb. in Canada: explaining the difference between the eastern and western distributions

Aim Present northern distribution limit of jack pine (Pinus banksiana Lamb.) follows the northern limit of continuous open boreal forest in western Canada, but not in eastern Canada where it is located further south. We tested the hypothesis that fire plays a more important role than climate in explaining the present position of the northern distribution limit of jack pine. Location An experimental jack pine plantation was set up in 1992, c. 300 km north of the present distribution limit of the species, in the Boniface river area of northern Québec (57°43′ N, 76°05′ W). Methods Climate and fire data were used to compare sites at and north of the present distribution limit of jack pine. In 2001, surviving individuals from the plantation were measured (total height, annual shoot elongation, basal diameter, and presence/absence of cones). Results Climate data from the ten weather stations used in this study did not show major differences. The northern limit of jack pine distribution is closely associated with the occurrence of fires larger than 200 ha. Survival of the planted jack pines was 31%. About 25% of the surviving pines qualified as normal, single‐stem individuals; the others were slightly uprooted and/or showed marks of erosion or foraging. Cones were produced, although no viable seeds were found. Main conclusions The low number of degree‐days above 5 °C at the plantation site could explain why the seeds were not viable. However, such climate conditions are not sufficient to prevent growth, as was shown by annual shoot elongation measurements. Most of the surviving jack pines from the Boniface river plantation are relatively healthy and follow a normal developmental programme. Low fire frequency and small fire size are amongst the main factors that prevented P. banksiana from migrating further north or east following deglaciation in northern Québec and Labrador.     

Host-plant influence on the population ecology of the jack pine budworm, Choristoneura pinus (Lepidoptera: Tortricidae)

Abstract.

• 1 Newly-emerged, second-instar jack pine budworm (Choristoneura pinus Freeman) establish spring feeding sites preferentially in the pollen cones of their host tree, Pinus banksiana Lamb.
• 2 Laboratory studies showed that the rate of establishment and survival of jack pine budworm on pollen cones was high throughout the entire spring emergence period of the insect.
• 3 In contrast, the rate of establishment and survival of jack pine budworm on vegetative buds was very poor early in the spring. Vegetative buds were only acceptable as feeding sites to the jack pine budworm for a relatively brief period in late spring.
• 4 Field studies showed that the change in population density of jack pine budworm during the spring emergence stage, as expressed by k-values, was a function of the abundance of pollen cones in the stand. Population reduction was greatest in those stands with the fewest pollen cones.
• 5 Direct measurement of spring dispersal by jack pine budworm showed that dispersal and consequent losses to the budworm population were greatest in stands with the fewest pollen cones.
• 6 We conclude that changes in the density of jack pine budworm are strongly influenced by production of pollen cones in the host stand. Because pollen cone production is related to previous years of defoliation by the jack pine budworm, we propose that pollen cones act as a density-dependent factor governing the density of early-stage jack pine budworm.
• 7 The resulting dynamics are compared to those of other budworm species and used to explain observed regional and temporal patterns of jack pine budworm outbreaks.

     

From the ground up: biotic and abiotic features that set the course from genes to ecosystems

Spatial variation in cone serotiny in Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia) across Yellowstone National Park influences initial pine recruitment after stand‐replacing fire with tremendous population, community, and ecosystem consequences. A previous study showed that much of the spatial variation in serotiny results from the balance of selection arising from high frequencies of fire favoring serotiny countered by opposing selection exerted by American red squirrels (Tamiasciurus hudsonicus) as seed predators. This earlier study, however, assumed stable local red squirrel densities over multiple generations of pines. Here, we examine environmental properties that might contribute to long‐term stability in the densities of red squirrels among sites. We found that the amount of clay in the soil, an indicator of plant and fungal growth—the latter an important food resource for red squirrels—and the coefficient of variation (CV) in diameter at breast height (DBH) of forest trees together account for a substantial amount of variation in red squirrel density. Soil development occurs over very long time scales, and thus, intersite variation in the amount of clay is unlikely to shift across pine generations. However, CV of DBH and squirrel density increase with stand age, which acts to amplify selection against serotiny with increasing interfire interval. Regardless, much of the variation in the CV of DBH is accounted for by soil bulk density, mean annual temperature, and surface curvature, which are unlikely to vary in their relative differences among sites over time. Consequently, these soil and abiotic attributes could contribute to consistent spatial patterns of red squirrel densities from one pine generation to the next, resulting in consistent local and spatial variation in selection exerted by red squirrels against serotiny.     

Forest floor properties across sharp compositional boundaries separating trembling aspen and jack pine stands in the southern boreal forest

Jack pine and trembling aspen are two early-seral boreal tree species with contrasting nutrient cycling strategies. Both species may form adjacent mono-specific stands separated by sharp compositional boundaries. We hypothesized that such boundaries result in wider functional ecotones. Spatial transitions in humus forms, forest floor chemistry and microbial communities were assessed across 32 m long transects set perpendicular to sharp compositional boundaries separating four jack pine and aspen stands. Split moving window analysis (SMWA) and moving window regression analysis (MWRA) were used to locate functional boundaries and ecotones. We found a gradual transition from moder (aspen) to mor (jack pine) humus spanning 16 m across the compositional boundary. An abrupt increase in forest floor water content at 3 m within jack pine stands was possibly due to aspen roots foraging for water beyond the boundary. The functional boundary and associated ecotone for forest floor pH, C:N ratio, Mg and ammonification were skewed toward jack pine stands, likely the result of aspen leaf dispersal. Low nitrification rates throughout jack pine stands and up to 11 m into aspen stands suggested that jack pine roots might extend far within aspen stands and produce metabolites that suppress nitrification. SMWA performed on the multivariate dataset of microbial fatty acids (FAs) revealed three distinct forest floor microbial communities that were skewed toward jack pine stands. Pine-type communities were associated to fungal FAs, pine-type and transition-type communities to non-fungal eukaryotic FAs, and aspen-type communities to bacterial FAs. Taken collectively, our data delimit a 24 m wide functional ecotone straddling sharp compositional boundaries separating trembling aspen and jack pine stands. We conclude that the functional diversity of boreal landscapes, where adjacent mono-specific stands are prevalent, is related to the patchiness of the landscape.     

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.     

Effectiveness of soil N availability indices in predicting site productivity in the oil sands region of Alberta

Background and aims

Quantitative relationships between soil N availability indices and tree growth are lacking in the oil sands region of Alberta and this can hinder the development of guidelines for the reclamation of the disturbed landscape after oil sands extraction. The aim of this paper was to establish quantitative relationships between soil N availability indices and tree growth in the oil sands region of Alberta.

Methods

In situ N mineralization rates, in situ N availability measured in the field using Plant Root Simulators (PRS? probes), laboratory aerobic and anaerobic soil N mineralization rates, and soil C/N and N content were determined for both the forest floor and the 0–20?cm mineral soil in eight jack pine (Pinus banksiana Lamb.) stands in the oil sands region in northern Alberta. Tree growth rates were determined based on changes in tree ring width in the last 6?years and as mean annual aboveground biomass increment.

Results

Soil N availability indices across those forest stands varied and for each stand it was several times higher in the forest floor than in the mineral soil. The in situ and laboratory aerobic and anaerobic soil N mineralization rates, soil mineralized N, in situ N availability measured using PRS probes, soil C/N ratio and N content in both the forest floor and mineral soil, as well as stand age were linearly correlated with tree ring width of jack pine trees across the selected forest stands, consistent with patterns seen in other published studies and suggesting that N availability could be a limiting factor in the range of jack pine stands studied.

Conclusions

In situ and laboratory aerobic and anaerobic N mineralization rates and soil C/N ratio and N content can be used for predicting tree growth in jack pine forests in the oil sand region. Laboratory based measurements such as aerobic and anaerobic N mineralization rates and soil C/N ratio and N content would be preferable as they are more cost effective and equally effective for predicting jack pine growth.     

Explaining the distribution of Pinus spp. in a Canadian boreal insular landscape

Abstract. To evaluate the respective contributions of habitat, fire regime and colonization-extinction processes to the distribution of northern Pinus species, we investigated the distribution of P. banksiana (jack pine), P. resinosa (red pine) and P. strobus (white pine) on 117 islands of Lake Duparquet in northwestern Québec. Stepwise logistic regressions indicated that the extent of xeric areas on the islands was the sole factor predicting jack pine distribution. The distribution of white pine was predicted primarily by the combined effects of distance to the shoreline and elevation, with a smaller effect of area of xeric habitat. The distribution of red pine was predicted by other populations of red pine nearby, with a slightly smaller effect of the combined effects of distance to shoreline and elevation. None of the species completely saturates all available islands nor is any restricted to specific, very exposed aspects. The results suggest that pine is more frequent on islands with characteristics that promote lightning strikes and thus higher fire occurrence. However, absence of pine in several islands may not be explained by abiotic characteristics or recent fire history. The presence of very small populations, together with low invasion potential, suggests that the observed distribution is mainly driven by the process of random extinction. A disequilibrium between present and past fire regimes may explain why northern pines have discontinuous distributions inside their range limits.     

Seed and cone diversity and seed germination of Pinus pinea in Strofylia Site of the Natura 2000 Network

Variation in cone size, seed number per cone, seed potential, seed efficiency, seed morphology and seed germination behavior of Pinus pinea and its relation to stand conditions was analyzed. Data were collected from P. pinea forest in Strofylia, southern Greece, a forest that belongs to the Natura 2000 European network and the RAMSAR convention and is characterized by the absence of regeneration for many decades. The pine stands found in the area were distinguished into five categories according to a previous study and our observations, regarding stand age, canopy cover and the degree of stress by human pressure. The categories are: (I) young artificial, (II) closed-mature, (III) open-mature, (IV) over-mature stands and (V) highly degraded stands. Cones were collected from all stand types and their morphological characteristics as well as their seed production were measured. Seeds were extracted from the collected cones, measured and their germination behavior was tested. The findings showed that the over-mature and the high degraded stands and to a lesser extent, the closed-mature stands, produced significantly smaller cones with a lower seed potential, a lower number of filled seeds per cone, a greater number of not fully developed seeds and reduced seed morphological characteristics than the young and open-mature stands. However, the seed germination behavior of fully developed seeds was only slightly affected by the stand type.     

Geographical variation in population demography and life history traits of Tecate cypress (<Emphasis Type="Italic">Hesperocyparis forbesii</Emphasis>) suggests a fire regime gradient across the USA–Mexico border

Plant adaptations to fire often display spatial heterogeneity associated with geographical variation in fire regime. We examined whether populations of the Tecate cypress (Hesperocyparis forbesii Adams) in southern California and northern Baja, Mexico, exhibited spatial heterogeneity in cone serotiny, in other life history traits associated with fire-adaptation, and in population demographic structure, to assess a putative difference in fire regime across the USA–Mexico border. Demographic data, tree life history data, and tree ring series were used to compare the demographic structure and life history traits of three populations in southern California with three populations in northern Baja California. In Baja populations, a greater number of tree size classes were present (χ ² = 12,589; P < 0.05), cone serotiny was more facultative (Mann–Whitney U = 58, P < 0.05), and young adult trees had a higher reproductive output (Mann–Whitney U = 2.65, P < 0.05), suggesting that a difference in fire regime between southern California and northern Baja has existed long enough (ca 8000 years) to drive microevolutionary divergence between the two sets of populations, and is not solely the result of 20th century differences in fire management policies across the international border. The transitional area between the two different fire regimes does not appear to coincide with the border itself but may lie in a zone of ecological transition south of Ensenada. The range of phenotypic variation observed within the Tecate cypress metapopulation suggests this species has the capacity to adapt to future environmental changes.     

Modelling Variable Fire Severity in Boreal Forests: Effects of Fire Intensity and Stand Structure

It is becoming clear that fires in boreal forests are not uniformly stand-replacing. On the contrary, marked variation in fire severity, measured as tree mortality, has been found both within and among individual fires. It is important to understand the conditions under which this variation can arise. We integrated forest sample plot data, tree allometries and historical forest fire records within a diameter class-structured model of 1.0 ha patches of mono-specific black spruce and jack pine stands in northern Québec, Canada. The model accounts for crown fire initiation and vertical spread into the canopy. It uses empirical relations between fire intensity, scorch height, the percent of crown scorched and tree mortality to simulate fire severity, specifically the percent reduction in patch basal area due to fire-caused mortality. A random forest and a regression tree analysis of a large random sample of simulated fires were used to test for an effect of fireline intensity, stand structure, species composition and pyrogeographic regions on resultant severity. Severity increased with intensity and was lower for jack pine stands. The proportion of simulated fires that burned at high severity (e.g. >75% reduction in patch basal area) was 0.80 for black spruce and 0.11 for jack pine. We identified thresholds in intensity below which there was a marked sensitivity of simulated fire severity to stand structure, and to interactions between intensity and structure. We found no evidence for a residual effect of pyrogeographic region on simulated severity, after the effects of stand structure and species composition were accounted for. The model presented here was able to produce variation in fire severity under a range of fire intensity conditions. This suggests that variation in stand structure is one of the factors causing the observed variation in boreal fire severity.     

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.     

Net ecosystem productivity of boreal jack pine stands regenerating from clearcutting under current and future climates

Life cycle analysis of climate and disturbance effects on forest net ecosystem productivity (NEP) is necessary to assess changes in forest carbon (C) stocks under current or future climates. Ecosystem models used in such assessments need to undergo well-constrained tests of their hypotheses for climate and disturbance effects on the processes that determine CO₂ exchange between forests and the atmosphere. We tested the ability of the model ecosys to simulate diurnal changes in CO₂ fluxes under changing air temperatures (T_a) and soil water contents during forest regeneration with eddy covariance measurements over boreal jack pine (Pinus banksiana) stands along a postclearcut chronosequence. Model hypotheses for hydraulic and nutrient constraints on CO₂ fixation allowed ecosys to simulate the recovery of C cycling during the transition of boreal jack pine stands from C sources following clearcutting (NEP from −150 to −200 g C m⁻² yr⁻¹) to C sinks at maturity (NEP from 20 to 80 g C m⁻² yr⁻¹) with large interannual variability. Over a 126-year logging cycle, annualized NEP, C harvest, and net biome productivity (NBP=NEP–harvest removals) of boreal jack pine averaged 47, 33 and 14 g C m⁻² yr⁻¹. Under an IPCC SRES climate change scenario, rising T_a exacerbated hydraulic constraints that adversely affected NEP of boreal jack pine after 75 years. These adverse effects were avoided in the model by replacing the boreal jack pine ecotype with one adapted to warmer T_a. This replacement raised annualized NEP, C harvest, and NBP to 81, 56 and 25 g C m⁻² yr⁻¹ during a 126-year logging cycle under the same climate change scenario.