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.     

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.     

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.     

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.     

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.     

Reproductive traits of Pinus halepensis in the light of fire – a critical review

Fire is known to be a major factor in shaping plants and vegetation worldwide. Many plant traits have been described as adaptations for surviving fire, or regenerating after it. However, many of the traits are also advantageous for overcoming other disturbances. The fact that fire in the Mediterranean Basin has been almost exclusively of anthropogenic origin, and thus is of short duration in an evolutionary time scale, cast doubt on the possibility that fire can act as a selective force in the Mediterranean Basin. Our aim here is to review the ecological advantages of Pinus halepensis traits and their possibility to be selected by fire. The non-self pruning of cones and branches, and the high resin content increase the probability of canopy fires and consequent death of P. halepensis trees. Post-fire regeneration of P. halepensis depends totally upon its canopy-stored seed bank. The seedlings grow quickly and they first reproduce at an early age. Young reproductive trees function first as females with a high percentage of serotinous cones. Thus, young P. halepensistrees allocate many resources to seed production, reducing their `immaturity risk' in a case of an early successive fire. The proportion of serotinous cones is higher in post-fire naturally regenerating stands than in unburned stands, and seeds from serotinous cones germinate better under simulated post-fire conditions. The extremely high pH of the ash-bed under the burned canopies creates the post-fire regeneration niche of P. halepensis exactly under their parent trees. All these traits are advantageous for post-fire regeneration, but could they also be selected during the time scale of anthropogenic fires in the Mediterranean Basin? Pinus halepensis is a relatively short living tree with almost no recruitment under forest canopy. The longest estimated fire-return interval and generation length are about 125 years. The earliest solid evidence for the first hominid-controlled fire in the Mediterranean basin is 780,000 years ago, and thus the estimated number of post-fire generations is 6240. We suggest that such a number of generations is sufficient for the selection and radiation of fire adaptive traits in P. halepensis.

     

Seed colour and post-fire bird predation in a Mediterranean pine forest

In a Pinus halepensis Mill. forest, a field experiment was designed to evaluate post-fire seed predation as affected by combinations of seed colour and soil substrates: light grey and black seeds combined with light grey ash, dark grey ash and pale brown sand. A survey of bird species inhabiting the area was also carried out and polyphenolic content of seed coat was assessed in seed lots of different colour. Light grey seeds were observed to be less predated on light grey ash, suggesting eucrypsis as a protective strategy against bird predation. On the contrary, no clear pattern was observed for the predation of black seeds on different substrates. In the study area both bird species breaking the seed coat and eating the endosperm and bird species swallowing the whole seed were monitored. We have estimated that more seeds were swallowed than broken, in all colour categories. Light grey seeds, which were found to have a higher content of polyphenols, were predated more than black seeds when exposed on the same substrate. Thus, no evidence was produced that the amount of polyphenols in seed coat could protect seeds from predation.     

Fire‐induced population reduction and landscape opening increases gene flow via pollen dispersal in Pinus halepensis

Population reduction and disturbances may alter dispersal, mating patterns and gene flow. Rather than taking the common approach of comparing different populations or sites, here we studied gene flow via wind‐mediated effective pollen dispersal on the same plant individuals before and after a fire‐induced population drop, in a natural stand of Pinus halepensis. The fire killed 96% of the pine trees in the stand and cleared the vegetation in the area. Thirteen trees survived in two groups separated by ~80 m, and seven of these trees had serotinous (closed) prefire cones that did not open despite the fire. We analysed pollen from closed pre and postfire cones using microsatellites. The two groups of surviving trees were highly genetically differentiated, and the pollen they produced also showed strong among‐group differentiation and very high kinship both before and after the fire, indicating limited and very local pollen dispersal. The pollen not produced by the survivors also showed significant prefire spatial genetic structure and high kinship, indicating mainly within‐population origin and limited gene flow from outside, but became spatially homogeneous with random kinship after the fire. We suggest that postfire gene flow via wind‐mediated pollen dispersal increased by two putative mechanisms: (i) a drastic reduction in local pollen production due to population thinning, effectively increasing pollen immigration through reduced dilution effect; (ii) an increase in wind speeds in the vegetation‐free postfire landscape. This research shows that dispersal can alleviate negative genetic effects of population size reduction and that disturbances might enhance gene flow, rather than reduce it.     

Cones structure and seed traits of four species of large‐seeded pines: Adaptation to animal‐mediated dispersal

Seed dispersal selection pressures may cause morphological differences in cone structure and seed traits of large‐seeded pine trees. We investigated the cone, seed, and scale traits of four species of animal‐dispersed pine trees to explore the adaptations of morphological structures to different dispersers. The four focal pines analyzed in this study were Chinese white pine (Pinus armandi), Korean pine (P. koraiensis), Siberian dwarf pine (P. pumila), and Dabieshan white pine (P. dabeshanensis). There are significant differences in the traits of the cones and seeds of these four animal‐dispersed pines. The scales of Korean pine and Siberian dwarf pine are somewhat opened after cone maturity, the seeds are closely combined with scales, and the seed coat and scales are thick. The cones of Chinese white pine and Dabieshan white pine are open after ripening, the seeds fall easily from the cones, and the seed coat and seed scales are relatively thin. The results showed that the cone structure of Chinese white pine is similar to that of Dabieshan white pine, whereas Korean pine and Siberian dwarf pine are significantly different from the other two pines and vary significantly from each other. This suggests that species with similar seed dispersal strategies exhibit similar morphological adaptions. Accordingly, we predicted three possible seed dispersal paradigms for animal‐dispersed pines: the first, as represented by Chinese white pine and Dabieshan white pine, relies upon small forest rodents for seed dispersal; the second, represented by Korean pine, relies primarily on birds and squirrels to disperse the seeds; and the third, represented by Siberian dwarf pine, relies primarily on birds for seed dispersal. Our study highlights the significance of animal seed dispersal in shaping cone morphology, and our predictions provide a theoretical framework for research investigating the coevolution of large‐seeded pines and their seed dispersers.     

Post-dispersal predation on Pinus sylvestris seeds by Fringilla spp: ground substrate affects selection for seed color

Background matching might lower the risk of seeds being eaten by seed predators that search visually. In aviary experiments, we analyzed the selection of diff erent-colored seeds by ground-feeding finches (Fringillacoelebs and F.montifringilla) against four naturally occurring forest soil substrates. The substrates were fresh burn (black), 6-year-old burn (brown), mineral soil (pale yellow) and Pleuroziumschreberi feather moss (green). We used color-sorted seeds of Pinussylvestris, a species with a large natural variation in seed color, ranging from pale yellow to black. Although seeds were scattered on the substrates at a density of only 91 seeds m⁻², birds removed seeds effectively. Both bird species found more pale than dark seeds on the fresh burn substrate. F. montifringilla also recovered more pale than dark seeds on the old burn, and more dark than pale seeds on mineral soil. In moss, the birds found very few seeds compared to the other substrates, and there was no color discrimination. P.sylvestris is frequently regenerating after fire, suggesting that dark seeds would be favored under selection from visually searching predators. Fire-adapted conifers with serotinous cones, e.g., Pinuscontorta ssp. latifolia, which spread their seeds primarily on freshly burnt surfaces, produce uniformly black or dark brown seeds. However, regeneration of the non-serotinous P.sylvestris is often extended for several years after a fire, during which substrate color and structure change. This may have helped to maintain variation in seed color. When regeneration of a plant species occurs on a substrate of uniform color, we believe that selection by visually searching seed predators will result in the evolution of cryptic seed color. Received: 16 August 1996 / Accepted: 30 November 1996     

Early postfire seed dispersal, seedling establishment and seedling mortality of Pinus coulteri (D. Don) in central coastal California, USA

Seedling recruitment in many highly serotinous populations of Pinus coulteri on California's central coast depends almost entirely on periodic, stand-replacing fire. Compared to serotinous pines of the Mediterranean Basin, little detailed information is available on the postfire demography of California closed-cone pines, including P. coulteri. In September 1996 a wildfire burned the 760-ha American Canyon Research Natural Area (RNA). Using aerial photography, we mapped burn severity of P. coulteri-chaparral woodlands and forests within the RNA. From May to September of 1997, we also quantified seedling establishment and mortality in relation to biophysical site characteristics including fire severity. Seventy-six percent of P. coulteri forests and woodlands experienced high-severity burns, 9% moderate-severity burns, and 15% low-severity or unburned. Of the 53 plots used for seedling counts, 70% were high-severity, 26% moderate-severity, and 4% low-severity. Seedling densities 13 months postfire were low (0.21 m^–2), but seedling mortality also was low (8.4%). Aerial seed bank size increased from north-facing to south-facing slopes and from high-severity to low-severity burns. Seedling recruitment was unrelated to burn severity and increased with the size of the canopy seed bank (cone density). Many seedlings established from rodent seed caches; 23% of the seedlings established in clumps from seeds cached by Dipodomys agilis, Chaetodipus californicus and Peromyscus maniculatus. Pinus coulteri seeds have low potential for dispersal by wind, but secondary dispersal by rodents moves seeds away from source trees and into neighboring chaparral. We discuss the potential importance of rodent seed caching to postfire demography of California and Mediterranean serotinous pines.     

Post‐fire seed predation: Does distance to unburnt vegetation matter?

Human‐induced changes to fire regimes result in smaller, more patchy fires in many peri‐urban areas, with a concomitant increase in potential edge effects. In sclerophyll vegetation, many structurally dominant serotinous plants rely on the immediate post‐fire environment for recruitment. However, there is little information about how fire attributes affect seed predation or recruitment for these species. We examined the influence of distance to unburnt vegetation on post‐dispersal seed predation for five serotinous species from sclerophyll vegetation in the Sydney region, south‐eastern Australia; Banksia serrata L.f., Banksia spinulosa Sm. var. spinulosa, Hakea gibbosa (Sm.) Cav., Hakea teretifolia (Salisb.) Britten (all Proteaceae) and Allocasuarina distyla (Vent.) L. Johnson (Casuarinaceae). We used cafeteria trials and differential exclusion of vertebrates and invertebrates to test whether rates of seed removal for these five species differed among (i) unburnt, (ii) burnt‐edge (approx. 10 m from unburnt vegetation) and (iii) burnt‐interior (approx. 100 m from unburnt vegetation) locations. When all animals had access to seeds, seeds were removed at lower rates from burnt‐interior areas than from other locations. Vertebrates (small mammals) showed this pattern markedly the first time the experiment was run, but in a repeat trial this effect disappeared. Rate of seed removal by invertebrates differed among plant species but we did not detect any such differences for removal by vertebrates. Overall rates of seed removal also differed significantly between the two fires studied. Our results indicate that small mammal seed predation can be substantial for large‐seeded serotinous shrubs, and that differences in the perimeter: area ratio, severity or size of a fire are likely to affect seed predation.     

Non‐serotinous woody plants behave as aerial seed bank species when a late‐summer wildfire coincides with a mast year

Abstract

1. Trees which lack obvious fire-adaptive traits such as serotinous seed-bearing structures or vegetative resprouting are assumed to be at a dramatic disadvantage in recolonization via sexual recruitment after fire, because seed dispersal is invariably quite constrained. We propose an alternative strategy in masting tree species with woody cones or cone-like structures: that the large clusters of woody tissue in a mast year will sufficiently impede heat transfer that a small fraction of seeds can survive the flaming front passage; in a mast year, this small fraction would be a very large absolute number.
2. In Kootenay National Park in British Columbia, we examined regeneration by Engelmann spruce (Picea engelmannii), a non-serotinous conifer, after two fires, both of which coincided with mast years. Coupling models of seed survivorship within cones and seed maturation schedule to a spatially realistic recruitment model, we show that (1) the spatial pattern of seedlings on a 630 m transect from the forest edge into the burn was best explained if there was in situ seed dissemination by burnt trees; (2) in areas several hundred meters from any living trees, recruitment density was well correlated with local prefire cone density; and (3) spruce was responding exactly like its serotinous codominant, lodgepole pine (Pinus contorta).
3. We conclude that non-serotinous species can indeed behave like aerial seed bank species in mast years if the fire takes place late in the seed maturation period. Using the example of the circumpolar boreal forest, while the joint probability of a mast year and a late-season fire will make this type of event rare (we estimate P = 0.1), nonetheless, it would permit a species lacking obvious fire-adapted traits to occasionally establish a widespread and abundant cohort on a large part of the landscape.

     

Bet‐hedging and germination in the Australian arid zone shrub Acacia ligulata

The diaspore of the Australian arid zone shrub Acacia ligulata is dispersed by birds and ants. To investigate the benefits of providing a dispersal structure attractive to both groups, we compared the germination response and viability of seeds eaten by birds, handled by ants or collected from trees to simulated precursors of germination: scarification, fire and rainfall were simulated. Seed germination and viability were related to the degree of preheating disturbance to the seed coat. Heating increased the germinability of seeds not scarified or eaten by birds. In the absence of heating, ingestion by birds increased germinability. Heating increased the mortality of seeds. Our results suggest that ingestion of seeds by birds may break seed dormancy and hence enable some seeds to germinate soon after dispersal. Alternatively, seeds not eaten by birds are likely to remain dormant until sufficiently scarified by soil or stimulated by fire. Consequently, in areas such as the Simpson Desert, A. ligulata may be able to use a range of seedling establishment ‘windows’ provided by monsoon rains, post‐fire environments and unseasonal winter rains, and also spread the risk of unsuccessful seedling establishment by retaining dormant seeds in the seedbank.     

Role of heat tolerance and cone protection of seeds in the response of three pine species to wildfires

The post-fire regenerative ability of Pinus halepensis, Pinus nigra and Pinus sylvestris, three of the most important pine species present in the West Mediterranean basin, has been analyzed in the light of seed tolerance to different temperatures and times of exposure, and of seed position during the fire event (seeds inside cones versus free seeds). The combination of different fire intensities and degrees of seed protection allows us to draw different scenarios during the fire event: canopy scenarios (seeds inside cones), surface scenarios (seeds on the ground surface), and soil scenarios (seeds in the top soil layers). There were interspecific differences in the pattern of cone opening under the different heat treatments: cones of P. nigra and P. sylvestris showed similar percentages of opening, but considerably higher than those of P. halepensis. In the three species, seeds inside cones showed higher percentages of germination than those that were free, emphasizing the important role of cones in the protection of pine seeds from high temperatures. The percentage of germination decreased when both the temperature and the time of exposure increased, and there was also a significant species effect: P. halepensis showed higher germination rates than P. nigra, and both were higher than P. sylvestris. The overall scores of seed germination of these three pine species under the conditions tested suggests that their regeneration after fire should come either from the soil bank, or from the canopy bank, but rarely from the ground surface. As the existence of a permanent seed bank in Mediterranean pines is probably limited or nil, pine recruitment after fire appears to be mainly controlled by the existence of a canopy seed bank. The contribution of this canopy bank to the differences in postfire regeneration success of the three pine species is discussed in the light of their seeding phenology and the effects of fire severity on cone opening. The results obtained in this study contribute to explain the successful regeneration of P. halepensis, and the failure of P. nigra and P. sylvestris after fire.     

Long‐distance dispersal in a fire‐ and livestock‐protected savanna

Savannas are highly diverse and dynamic environments that can shift to forest formations due to protection policies. Long‐distance dispersal may shape the genetic structure of these new closed forest formations. We analyzed eight microsatellite loci using a single‐time approach to understand contemporary pollen and effective seed dispersal of the tropical tree, Copaifera langsdorffii Desf. (Fabaceae), occurring in a Brazilian fire‐ and livestock‐protected savanna. We sampled all adult trees found within a 10.24 ha permanent plot, young trees within a subplot of 1.44 ha and open‐pollinated seeds. We detected a very high level of genetic diversity among the three generations in the studied plot. Parentage analysis revealed high pollen immigration rate (0.64) and a mean contemporary pollen dispersal distance of 74 m. In addition, half‐sib production was 1.8 times higher than full‐sibs in significant higher distances, indicating foraging activity preference for different trees at long distances. There was a significant and negative correlation between diameter at breast height (DBH) of the pollen donor with the number of seeds (r = ?0.640, P‐value = 0.032), suggesting that pollen donor trees with a higher DBH produce less seeds. The mean distance of realized seed dispersal (recruitment kernel) was 135 m due to the large home range dispersers (birds and mammals) in the area. The small magnitude of spatial genetic structure found in young trees may be a consequence of overlapping seed shadows and increased tree density. Our results show the positive side of closed canopy expansion, where animal activities regarding pollination and seed dispersal are extremely high.     

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.     

Discrimination among pinyon pine trees by Clark's Nutcrackers: effects of cone crop size and cone characters

Summary The influences of Colorado pinyon pine (Pinus edulis) cone crop size, cone and seed weight, cone length, number of seeds per cone, number of viable seeds, and percent viable seeds on the foraging behavior of avian seed dispersal agents were examined in field and laboratory settings. In the field, there was a significant positive relationship between cone number per tree and both the absolute number of cones and the percentage of the cone crop from which seeds were harvested. Cone weight and the number of viable seeds were also significantly related to seed harvest intensity. Laboratory experiments examined the relationship between crop size and cone characters on seed harvest by 18 Clark's Nutcrackers (Nucifraga columbiana). Nutcrackers were offered a choice of two tree types: one with 20 cones attached, and another with 10 cones attached. Significantly more birds chose to remove seeds first from the tree with 20 cones than the tree with 10 cones. In timed trials, they also harvested seeds from significantly more cones on the tree with the higher cone density. In the laboratory, cones chosen for seed removal by the nutcrackers had significantly more viable seeds, more seeds, and were longer compared to cones that were not chosen. Such discriminatory foraging behavior may increase avian foraging efficiency and result in differential reproductive success of pinyon pines. This behavior may therefore influence the evolution of pinyon pine reproductive traits.     

Fire season effects on the recruitment of non‐sprouting serotinous Proteaceae in the eastern (bimodal rainfall) fynbos biome,South Africa

Abstract Research in Mediterranean‐climate shrublands in both South Africa and Australia shows that recruitment of proteoid shrubs (non‐sprouting, serotinous Proteaceae) is best after warm‐season (summer and autumn) fires and worst after cool‐season (winter and spring) ones. This pattern has been attributed to post‐dispersal seed attrition as well as size of pre‐dispersal seed reserves. Here we investigate patterns of post‐fire recruitment for four proteoid species in the eastern part of South Africa's fynbos biome, which has a bimodal (spring and autumn) rainfall regime. Despite the lack of significant differences in recruitment between cool‐ and warm‐season burns, we find some evidence for favourable recruitment periods following fires in spring and autumn, immediately before, and coinciding with, the bimodal rainfall peaks. This suggests that enhanced recruitment is associated with conditions of high soil moisture immediately after the fire, and that rapid germination may minimize post‐dispersal seed attrition. In two of the species, we also find a shift from peak flowering in winter and spring in the Mediterranean‐climate part of the fynbos biome, to summer and autumn flowering in the eastern part. Because these two species are only weakly serotinous, warm‐season flowering would result in maximal seed banks in spring, which could explain the spring recruitment peak, but not the autumn one. We conclude that eastern recruitment patterns differ significantly from those observed in the western and central parts of the biome, and that fire management protocols for the east, which are currently based on data and experience from the winter‐rainfall fynbos biome, need to be adjusted accordingly. Fire managers in the eastern fynbos biome should be less constrained by requirements to burn within a narrow seasonal range, and should therefore be in a better position to apply the required management burns.     

Seed Dispersal of a High Quality Fruit by Specialized Frugivores: High Quality Dispersal?1

Dispersal quality, as estimated by the cumulative effects of dispersal, germination, seed predation, and seedling survival, was examined for Beilschmiedia pendula (Lauraceae) in Monteverde, Costa Rica. I determined the pattern of dispersal by finding seeds deposited by birds, protected the seeds from seed predators with cages to assess germination and seedling survival, and examined seed predation rates with marked seeds. Seed predation, germination, and seedling survival were compared between seeds naturally dispersed by birds and seeds placed at randomly located sites. Approximately 70 percent of seeds dispersed by birds (N= 244) were deposited <10 m from crown edges of fruiting B. pendula trees, although some seeds were dispersed at least 70 m away. Larger seeds were more likely to be dispersed under or close to the parent trees, and larger seeds produced larger seedlings. Seed size was not correlated directly with seedling survival, but larger seedlings at three months were most likely to survive one year. Seed predation by mammals and insects and seedling mortality due to fungal pathogens were concentrated beneath the crowns of parent trees. Seedlings and saplings were more abundant beneath fruiting B. pendula trees, but individuals farther away were taller on average. Thus, dispersal is beneficial for B. pendula, but such benefits appear most pronounced at a small spatial scale; seeds dispersed >30 m from the crown edges actually had a lower probability of survival than those dispersed 10–20 m. Only 10 percent of B. pendula. seeds received high‐quality dispersal in terms of landing in the zone with the highest per seed probability of seedling survival 10–20 m from parental crowns.