Gene flow pattern and mating system in a small population of Quercus semiserrata Roxb. (Fagaceae)

Pollen flow from external sources is important for the conservation of tree species in fragmented forests or small populations, because it can be sufficient to prevent differentiation among them, and appears to be able to prevent the loss of their genetic diversity through genetic drift. In this study, we examined the genetic heterogeneity of pollen pools accepted by each Quercus semiserrata seed parent at the Khun Wang Royal Agriculture Research Center, Thailand, both within and among two mast fruiting years (2005 and 2007), using paternity analysis and analysis of molecular variance (AMOVA). The mating systems of the trees were also examined using the multilocus mating system model (MLTR), after determining the genotypes at eight microsatellite loci of 26 seed-trees and 435 seeds from 8 seed-trees in the 2 mast fruiting years. The average distance of effective pollen flow within the plot was estimated to be 52.4 m, and 95% of effective pollen was dispersed within 200 m, indicating that effective pollen flow is highly localized and that most effective pollen is contributed by near-neighbor trees. The proportion of effective pollen that immigrated from external sources was estimated to be 26.2%. The AMOVA analysis based on the pollen haplotypes showed that the pollen pools, both total and for each reproductive year, significantly genetically differed among the seed parents. Using a mixed mating model, the estimate of biparental inbreeding for the total population (t_m − t_s) was 0.013, indicating that a low proportion of mating occurred among close relatives. The effective number of pollen donors (N_ep) was estimated to be 9.987 using the TwoGener model, or 10.989 using the mixed mating model. The effective number of pollen donors of seeds was higher in the mast fruiting year 2005 than in the other examined year, 2007. Consequently the allelic richness and genetic diversity of seeds produced in 2005 were higher than those produced in 2007. Overall, the results show that high outcrossing rates, high levels of gene flow from other populations and heterogeneity in the pollen received by an individual may enhance the ability of populations to maintain effective population sizes. Therefore, these processes may be sufficient to prevent loss of genetic diversity through genetic drift of Q. semiserrata at this study site.     

Seed rain,soil seed bank,seed loss and regeneration of Castanopsis fargesii (Fagaceae) in a subtropical evergreen broad-leaved forest

Understanding the seed rain and seed loss dynamics in the natural condition has important significance for revealing the natural regeneration mechanisms. We conducted a 3-year field observation on seed rain, seed loss and natural regeneration of Castanopsis fargesii Franch., a dominant tree species in evergreen broad-leaved forests in Dujiangyan, southwestern China. The results showed that: (1) there were marked differences in (mature) seed production between mast (733,700 seeds in 2001) and regular (51,200 and 195,600 seeds in 2002 and 2003, respectively) years for C. fargesii. (2) Most seeds were dispersed in leaf litter, humus and 0–2 cm depth soil in seed bank. (3) Frequency distributions of both DBH and height indicated that C. fargesii had a relatively stable population. (4) Seed rain, seed ground density, seed loss, and leaf fall were highly dynamic and certain quantity of seeds were preserved on the ground for a prolonged time due to predator satiation in both the mast and regular years so that the continuous presence of seed bank and seedling recruitments in situ became possible. Both longer time observations and manipulative experiments should be carried out to better understand the roles of seed dispersal and regeneration process in the ecosystem performance.     

The relationship between seed weight, germination and biochemical reserves of maritime pine (Pinus pinaster Ait.) in Morocco

Seed germination and biochemical reserves of maritime pine (Pinus pinaster Ait.) were studied with the aim of providing germination information for reforestation and conservation programs. Ten natural populations were used to assess variation in seed weight, germination characteristics and biochemical reserves and to examine the relationship between these characteristics. The analysis of variance showed highly significant population effects for seed weight, germination characteristics and protein content in both seeds and the female gametophyte. The mobilization of protein content in female gametophyte during seed germination differed more among populations than sugar content, suggesting that protein content was more sensitive to environment effects than sugar content. A strong positive correlation between germination capacity and the protein content in both seeds and female gametophyte indicated that the best populations in term of germination capacity may also be the richest in protein content. Seeds that were heavier and had a lower speed of protein content mobilization in the female gametophyte appeared to be better adapted to drought conditions. The results also suggested that as much as possible of the potentially valuable genetic variation among populations of this species should be preserved for reforestation and conservation purposes.     

Germination and early seedling growth of <Emphasis Type="Italic">Pinus densata</Emphasis> Mast. provenances

We studied seed germination and early seedling growth of Pinus densata to explore the range of variability within the species and to inform afforestation practices. Phenotypes were evaluated at a forest tree nursery under conditions that support Pinus yunnanensis, one of the presumed parental species of P. densata. Seeds were collected from 20 open-pollinated trees within each of eight autochthonous populations representing the natural distribution of P. densata in China to assess variation in germination traits and early seedling growth, and to examine the relationships among these traits. Results showed that seeds from all populations germinated and seedlings established successfully. There were significant differences among populations in 13 of 14 traits evaluated. Seed germination and early seedling growth were strongly related to seed size and seed weight. Bigger seeds germinated earlier and faster than small seeds, and seedling size was positively correlated with seed size. Some germination traits were strongly and significantly correlated with climatic variables associated with the provenance of the studied populations. Based on these observations, we conclude there were large, significant, and biologically important differences among P. densata populations in seed germination and seedling growth traits. The observed variability probably reflects a high degree of adaptive differentiation among populations that is likely to be relevant for future afforestation.     

Assessment of diversity among populations of Rauvolfia serpentina Benth. Ex. Kurtz. from Southern Western Ghats of India,based on chemical profiling,horticultural traits and RAPD analysis

Genetic, morphological and chemical variations of ten natural populations of Rauvolfia serpentina Benth. Ex. Kurtz. from Southern Western Ghats of India were assessed using RAPD markers reserpine content and morphological traits. An estimate of genetic diversity and differentiation between genotypes of breeding germplasm is of key importance for its improvement. Populations were collected from different geographical regions. Data obtained through three different methods were compared and the correlation among them was estimated. Statistical analysis showed significant differences for all horticultural characteristics among the accessions suggesting that selection for relevant characteristics could be possible. Variation in the content of Reserpine ranges from 0.192 g/100 g (population from Tusharagiri) to 1.312 g/100 g (population from Aryankavu). A high diversity within population and high genetic differentiation among them based on RAPDs were revealed caused both by habitat fragmentation of the low size of most populations and the low level of gene flow among them. The UPGMA dendrogram and PCA analysis based on reserpine content yielded higher separation among populations indicated specific adaptation of populations into clusters each of them including populations closed to their geographical origin. Genetic, chemical and morphological data were correlated based on Mantel test. Given the high differentiation among populations conservation strategies should take into account genetic diversity and chemical variation levels in relation to bioclimatic and geographic location of populations. Our results also indicate that RAPD approach along with horticultural analysis seemed to be best suited for assessing with high accuracy the genetic relationships among distinct R. serpentina accessions.     

Quantitative variation among Kenyan populations of Acacia senegal (L.) Willd. for gum production, seed and growth traits

Acacia senegal (L.) Willd. is an African leguminous multipurpose tree species belonging to subfamily Mimosoideae and subgenus Aculeiferum, highly valued for gum arabic production. This study estimated the genetic diversity within and among Kenyan populations of the species in the Bulla Sambul, Kutulo, Wamba and Meisori populations based on 12 quantitative traits, which included; tree height, diameter at breast height (DBH), number of branches, bark thickness, gum weight and various pod and seed traits on 20 randomly selected trees in every population. Duncan’s multiple range test (DMRT) and the dendrogram distinctively divided the populations into two groups; Wamba and Meisori in one group while Bulla Sambul and Kutulo in the other, indicating geographical structuring of the genetic variability. The trees were larger, taller with greater pod and seed traits in the Wamba and Meisori than the Bulla Sambul and Kutulo populations. Contrarily, trees had lighter seeds, more branches with higher gum weight in the Kutulo and Bulla Sambul than the Wamba and Meisori populations. The Shannon–Weaver diversity index (H), depicted 1,000-seed weight as the most varied trait across the populations. These findings provide essential information on the genetic diversity of the species, necessary for delineation of particular zones for seed sources for germplasm conservation, selection for domestication and improved gum arabic production.     

Roles of seed and pollen dispersal in natural regeneration of Castanopsis fargesii (Fagaceae): Implications for forest management

Natural regeneration is an important process to reverse the loss of forests. Understanding the process of natural regeneration is crucial for achieving sustainable forest management. In this study, we examined the effects of seed and pollen dispersal in naturally regenerated populations of Castanopsis fargesii. Genetic variation in six populations along two successional series (three successional stages in each series: early, pre-climax, and climax) was assayed using RAPD (random amplified polymorphic DNA) markers. High genetic variability was observed as measured with Shannon's information index. A majority of genetic variation was distributed within populations (Φ_st = 0.1271) and significant isolation by distance existed among populations. A contrasting pattern of genetic variation along these two series was observed, representing different scenarios of natural regeneration processes. The ratio of the number of migrants between the mature populations to the number of migrants from the mature to immature populations was estimated as 1.146 ± 0.099 to 1.981 ± 0.164, implying that comparable seed and pollen dispersal might exist at a fine spatial scale (∼2 km²). The results suggest the critical role of seed dispersal in shaping genetic composition and diversity in the second-growth forests. Barriers to seed dispersal from a variety of genetic sources could result in low genetic diversity in naturally regenerated populations. Management that facilitates the connectivity of newly regenerated stands with mature forests could be effective for natural regeneration given the predominant role of short-distance dispersal of seeds in this species.     

云杉天然群体表型多样性研究

以系统揭示云杉全分布区天然群体表型变异程度和变异规律为目的 ,以其全分布区 10个有代表性天然群体 ,每个群体 30个个体为试材 ,对针叶、球果、种鳞、种翅和种子等 17个表型性状进行系统比较分析 ,采用方差分析、多重比较、相关分析和聚类分析等统计分析方法 ,讨论了群体间和群体内的表型多样性。结果表明 :云杉种内表型性状在群体间和群体内存在极其丰富的遗传差异。云杉球果、针叶、种鳞、种翅和种子 5个表型性状的变异系数分别为 19 14 %、2 6 4 6 %、13 5 8%、19 78%和 174 0 % ,种鳞性状稳定性较其他性状高。表型分化系数 (VST)的变幅为 2 0 9%～ 4 0 6 2 % ,群体间表型分化系数均值为 30 99%。群体间变异 (30 99% )小于群体内变异 (6 9 0 1% )。 17个表型性状间多数呈极显著或显著的正相关 ,球果长度、球果径、种鳞长度、种鳞宽度和种子千粒重为云杉易测定和重要的表型性状。云杉种内群体表型变异在空间分布上呈现以纬度为主的单向变异模式 ,利用群体间欧氏距离进行UPGMA聚类分析表明 ,云杉群体可以划分为 5类     

Germination and early seedling growth of Pinus densata Mast. provenances

We studied seed germination and early seedling growth of Pinus densata to explore the range of variability within the species and to inform afforestation practices.Phenotypes were evaluated at a forest tree nursery under conditions that support Pinus yunnanensis, one of the presumed parental species of P. densata. Seeds were collected from 20 open-pollinated trees within each of eight autochthonous populations representing the natural distribution of P. densata in China to assess variation in germination traits and early seedling growth, and to examine the relationships among these traits. Results showed that seeds from all populations germinated and seedlings established successfully. There were significant differences among populations in 13 of 14 traits evaluated. Seed germination and early seedling growth were strongly related to seed size and seed weight. Bigger seeds germinated earlier and faster than small seeds, and seedling size was positively correlated with seed size. Some germination traits were strongly and significantly correlated with climatic variables associated with the provenance of the studied populations. Based on these observations, we conclude there were large, significant, and biologically important differences among P. densata populations in seed germination and seedling growth traits. The observed variability probably reflects a high degree of adaptive differentiation among populations that is likely to be relevant for future afforestation.     

Significant effects of temperature on the reproductive output of the forest herb Anemone nemorosa L.

Climate warming is already influencing plant migration in different parts of the world. Numerous models have been developed to forecast future plant distributions. Few studies, however, have investigated the potential effect of warming on the reproductive output of plants. Understorey forest herbs in particular, have received little attention in the debate on climate change impacts.This study focuses on the effect of temperature on sexual reproductive output (number of seeds, seed mass, germination percentage and seedling mass) of Anemone nemorosa L., a model species for slow colonizing herbaceous forest plants. We sampled seeds of A. nemorosa in populations along a 2400 km latitudinal gradient from northern France to northern Sweden during three growing seasons (2005, 2006 and 2008). This study design allowed us to isolate the effects of accumulated temperature (Growing Degree Hours; GDH) from latitude and the local abiotic and biotic environment. Germination and seed sowing trials were performed in incubators, a greenhouse and under field conditions in a forest. Finally, we disentangled correlations between the different reproductive traits of A. nemorosa along the latitudinal gradient.We found a clear positive relationship between accumulated temperature and seed and seedling traits: reproductive output of A. nemorosa improved with increasing GDH along the latitudinal gradient. Seed mass and seedling mass, for instance, increased by 9.7% and 10.4%, respectively, for every 1000 °C h increase in GDH. We also derived strong correlations between several seed and seedling traits both under field conditions and in incubators. Our results indicate that seed mass, incubator-based germination percentage (Germ%_Inc) and the output of germinable seeds (product of number of seeds and Germ%_Inc divided by 100) from plants grown along a latitudinal gradient (i.e. at different temperature regimes) provide valuable proxies to parameterize key population processes in models.We conclude that (1) climate warming may have a pronounced positive impact on sexual reproduction of A. nemorosa and (2) climate models forecasting plant distributions would benefit from including the temperature sensitivity of key seed traits and population processes.     

庞泉沟自然保护区河岸林群落的土壤种子库特征

以人工拣种结合种子萌发法研究了庞泉沟自然保护区9种河岸林群落的土壤种子库特征,结果表明: 9种群落的土壤种子库密度间于684±108 - 2 696±266粒·m^-2,69.27% - 88.65%的种子留存于0 - 5 cm的层次,沙棘灌丛的种子库密度最大,乌柳灌丛最小;离河岸10 m和20 m处的种子库密度大于岸边、离河岸30 m和40 m处;种子库包含的40种植物以多年生草本为主,灌木和草本种多为耐干扰种,存在典型的湿地植物,但种子储量以先锋种白桦最丰;辽东栎无完整种子,白桦、华北落叶松、云杉和油松的种子活力保持期为1 - 2年,多数草本的种子活力保持期超过2年;按照群落土壤种子库的相似性,将群落划分为林分开始期、林分排除期、下层再现期和老熟期4个演替阶段。演替后期群落的种子库密度、丰度以及与地上植被的相似性都小于前期群落。土壤种子库是森林自然更新及种群恢复的重要基础,本文对于研究地区河岸林的保护和重建具有重要的理论意义。     

Effect of seed mass on emergence and seedling development in Pterocarpus marsupium Roxb.

We investigated the effect of seed mass on emergence, seedling survival and growth ofPterocarpus marsupium Roxb., a medium to large, commercially valuable and deciduous tree species. Among the three size classes viz. small （10-12 mm）, medium （13-15 mm） and large （16-17 mm）, the maximum proportion, of seed by number （54.12%） and dry weight （51.87%） was recorded, in the medium size seed class. Seed length and seed width were greatest in the large seed class （16.50 mm, 7.33 mm） followed by medium （13.50 mm, 5.60 mm） and small （11.37 ram, 3.66 mm）. Similarly, hundred seed weight （100 sw） varied from a maximum of 12.92 g in the large seed class to intermediate 10.95 g in the medium seed class and minimum of 7.02 g in the small seed class. The large seed size showed maximum emergence and shoot length over the medium and small seed class. After six months of growth, significant variations due to seed size were also observed for the growth and dry weight of P. marsupium seedlings. Seedling vigour, expressed in terms of height, collar diameter, number of leaves and dry biomass, was sig- nificantly affected by seed class. Seedlings that emerged from large seeds showed better growth and produced heavier seedlings as compared to medium and small seeds.     

The utility of Trillium and Maianthemum as phyto-indicators of deer impact in northwestern Pennsylvania,USA

Trillium spp. and Maianthemum canadense Desf. are preferred deer forage throughout their range and have potential as indicators of deer impact throughout the eastern United States. This paper evaluates the use of these plants as indicators at a landscape level within the Kinzua Quality Deer Cooperative (KQDC, ∼30,000 ha) in McKean County, Pennsylvania where a high level of deer impact has been sustained over the last 60 years.We hypothesized that deer impacts to Maianthemum and Trillium spp. populations across the KQDC landscape would resemble those reported at much smaller scales in previous studies and would indicate spatial variation in deer impact within the KQDC area. We found that most of the Trillium and Maianthemum characteristics measured in the KQDC were below the “healthy” standards established in the literature and comparable to characteristics associated with high deer impacts: the percentage of Trillium flowering in the KQDC was 9.1%, compared to the standard of 21–34%; the average Trillium heights in 27% of the KQDC plots were below the 12–14 cm recommended for healthy Trillium populations; Maianthemum mean leaf length and percent flowering were 39 mm and 0.3%, compared to standards of 42 mm and 0.5% for plants in browse accessible areas and 55 mm and 20% flowering for plants in refugia. Despite these high levels of impact, no significant correlations were found between the indicator variables and a direct measure of deer impact, a browse index based on browsing measurements of preferred woody plant species.Based on these results we conclude that these two species do not track spatial variation in deer impact across the KQDC. Relationships between deer impact and indicator characteristics could be clouded by factors such as environmental variables not measured in our study and legacy effects of high deer abundance. Both species will bear further investigation in a long-term monitoring project within the KQDC and could serve as a measurement of recovery in areas that have experienced high deer impact when deer populations are lowered and monitoring sites are analyzed through time.     

不同种源盐肤木种子形态性状遗传变异研究

以21个种源的种子为材料,研究了盐肤木种子形态性状在种源水平上的遗传变异规律。研究结果表明:盐肤木种子形态性状在不同种源间存在极显著差异,但主要由环境因素引起,不能比较稳定的遗传;盐肤木种子大小表现出明显的经纬度地理变异模式,且降雨量越小,气温越高,其种子饱满度越高。盐肤木种子长、种子厚和种子千粒重之间显著正相关。     

Fruit,seed and seedling variation in Uapaca kirkiana from natural populations in Malawi

Uapaca kirkiana is a miombo fruit tree with a long history of human use. In Malawi, the species is rapidly gaining attention as an agroforestry tree for fruit production. Accordingly, a formal study to explore variations in fruit, seed and seedling traits within the species was undertaken at the Forestry Research Institute of Malawi. Fruits from 12 natural populations in Malawi were collected in November and December, 1994. Fresh fruit weight and dependent parameters (exocarp, mesocarp and seeds) and seed length, width, thickness and weight (following processing) varied significantly among the populations. When seedlings were raised at Zomba, there were significant differences in germination capacity and height, root collar diameter and biomass production at six months of age. None of these traits was correlated with source latitude, longitude or elevation. Variations in the traits reported emphasize the importance of selecting suitable seed sources before commencing domestication programmes. The levels of variation observed suggest useful gains might be obtained through a breeding programme following careful provenance selection.     

Seed mass effects on performance of Pinus halepensis Mill. seedlings sown after fire

Pinus halepensis is a widely distributed species in the Mediterranean basin. It is generally well adapted to regenerate after wildfire, except when fire intervals are too short (≤15 years). In these latter situations, direct seeding could be a good alternative for restoring pine woodland after fire. Under dry climate, low seedling growth and survival can greatly limit the use of seeding. Early studies have shown that seedlings grown from large seeds have higher seedling establishment, growth and survival. Seed size grading may however reduce the genetic diversity of a seed lot by eliminating part or all of the families with relatively small seeds. An alternative to improve seed lot quality without losing genetic variability could be collecting and sieving seeds from each family separately. In order to explore the influence of seed mass on P. halepensis seedling performance, seeds from five half-sib families differing in mean seed mass were sown under greenhouse and field conditions. Final seedling emergence was unrelated to seed mass and half-sib family. The time of emergence was also unrelated to seed mass but it varied among families. Seed mass showed a positive effect on seedling height and diameter, both at population level and within family throughout the study period (9 months under greenhouse conditions and 20 months under field conditions). We also observed a negative relationship between seed mass and relative growth rate for seedling diameter, but it was not high enough to fully compensate the initial differences due to seed mass at the end of the 20-month study period. Seedling predation had a considerable impact on seedling survival, and it was not related to seed mass. When seedlings killed by predation were excluded from the survival analysis, larger seedlings, coming from larger seeds, showed slightly better survival, but only during the first growing period. The small advantages obtained from large seed mass in seedling development do not seem to justify the increased operational costs derived from seed mass selection for each family.     

Factors influencing levels of genetic diversity in woody plant species

The plant allozyme literature was reviewed to: (1) compare genetic diversity in long-lived woody species with species representing other life forms, and (2) to investigate whether the levels and distribution of genetic diversity in woody species are related to life history and ecological characteristics. Data from 322 woody taxa were used to measure genetic diversity within species, and within and among populations of species. Woody species maintain more variation within species and within populations than species with other life forms but have less variation among populations. Woody species with large geographic ranges, outcrossing breeding systems, and wind or animal-ingested seed dispersal have more genetic diversity within species and populations but less variation among populations than woody species with other combinations of traits. Although life history and ecological traits explain a significant proportion (34%) of the variation among species for the genetic parameters measured, a large proportion of the interspecific variation is unexplained. The specific evolutionary history of each species must play an important role in determining the level and distribution of genetic diversity.     

Distribution,colonization and diversity of arbuscular mycorrhizal fungi associated with central Himalayan rhododendrons

The objective of the present study was to investigate arbuscular mycorrhizal status of five species of rhododendrons distributed in Kumaun region of the Indian Central Himalaya. Root and rhizosphere soil samples of all the five species of rhododendrons, namely, Rhododendron anthopogon, R. arboreum, R. campanulatum, R. barbatum and R. lepidotum were collected from temperate, sub-alpine to alpine location in altitudinal range from 1500 to 4500 m amsl. The arbuscular mycorrhizal colonization in root samples ranged from 28 to 42%; and maximum and minimum colonization was observed in R. arboreum and R. lepidotum, respectively. The highest number of intraradical vesicles (12.5 ± 2.8 cm⁻¹ root segment) was recorded in R. arboreum and the lowest (7.0 ± 1.7 cm⁻¹ root segment) in R. barbatum; vesicles were not observed in R. lepidotum. Spores were extracted from the rhizosphere soil by wet sieving to perform microscopic identification of the species. The maximum and minimum populations of spores were detected in the rhizosphere soil samples of R. anthopogon (52.0 ± 1.5 spores 25 g⁻¹ soil) and R. lepidotum (32.0 ± 2.5 spore 25 g⁻¹ soil), respectively. Spore populations were found to belong to five genera—Acaulospora, Glomus, Gigaspora, Sclerocystis and Scutellispora; genus Glomus was found to be dominant in the rhizosphere soil samples of all the rhododendron species. The most frequent and abundant species was G. fasciculatum, however, this species was not isolated from the rhizosphere soil of R. barbatum. Finger millet (Eleucine coracana) was used to cultivate the trap culture of arbuscular mycorrhizal fungi to confirm the species identity. Spores of Glomus pustulatum, not detected in the rhizosphere soil, were recovered from the trap culture. Contrary to this, genus Gigaspora, which was present in the rhizosphere soil, did not sporulate in the trap culture. Shannon and Wiener index of diversity and Simpson's index of dominance indicated that the species richness, dominance and diversity of arbuscular mycorrhizal fungi decrease with increasing altitude. In two species of rhododendrons, namely R. campanulatum and R. anthopogon, dark septate mycelium was also observed.     

Phenotypic variation in fruit and seed morphology of Adansonia digitata L. (baobab) in five selected wild populations in Malawi

Adansonia digitata L. (Baobab) is widely distributed throughout Sub-Saharan Africa stretching from Senegal to Sudan, and from Ethiopia to Natal. Information of phenotypic and genetic variation is a prerequisite for the domestication and improvement of baobab fruits from the wild. A study was done to determine within and between provenance variability in fruit and seed characteristics of five populations selected from four silvicultural zones and assess whether morphometric traits could delineate populations from different zones into land races. Fruits were characterized from 55 trees representing a wide geographical range. Six fruit traits and three individual seed traits were assessed. Results showed highly significant differences (P????0.001) in fruit, pulp, and seed weight, fruit length and width, number of seed, individual seed weight, seed length and width within and between populations. Mean fruit weight ranged from 125.8?±?3.25 to 162.9?±?3.25?g, seed weight ranged from 38.6?±?2.5 to 66?±?2.01?g and pulp weight ranged from 28.7?±?1.33 to 41.4?±?1.33?g. Single seed weight showed pronounced evidence of divergence of populations into ecotypes. The rich diversity found between and within populations is important for domestication purposes and tree improvement through selection and breeding. All populations could be used for seed source but distribution should be consciously done recognizing existence of races.     

Annual variation in predation and dispersal of Arolla pine (Pinus cembra L.) seeds by Eurasian red squirrels and other seed-eaters

Seed predation and dispersal are key processes in the survival and distribution of plant species. Many animals cache seeds for later consumption, and, failing to recover some of these seeds, act as seed dispersers, influencing post-dispersal seed and seedling survival. Both animal and plant benefit from scatterhoarding and natural selection of seed characteristics and adaptations of seed predators (and dispersers) is one of the most important examples of co-evolution and mutualism. We studied the producer–consumer Arolla pine (Pinus cembra)–red squirrel (Sciurus vulgaris) system in a subalpine forest in the Italian Alps. Arolla pine produced large seed-crops (masting) at irregular intervals, followed by years with poor or moderate seed production. Squirrel density fluctuated in synchrony with the food resource, eliminating the time-lag normally present when resources are produced in pulses. In all years except 2009 (a mast-crop year), all Arolla pine cones were harvested (their seeds consumed and/or cached) by September to late October by different species. Both squirrels and nutcrackers (Nucifraga caryocatactes) fed on seeds, and their relative pre-dispersal predation rates (on cones in the canopy) differed between years. Overall, nutcrackers consumed more seeds between July and October than squirrels, but in 1 year squirrels took the largest number of seeds. Pre-dispersal seed predation by squirrels tended to be lower in years with large seed-crop size and there was a positive correlation, over the entire study period, between density of recovered hoards and Arolla pine seed density of the previous year. We conclude that (i) squirrels and nutcrackers are important pre-dispersal seed predators and seeds dispersers; (ii) squirrels are also post-dispersal seed predators, and (iii) the proportion of cached seeds consumed by squirrels increased with the size of the Arolla pine seed-crop, suggesting that red squirrel is a conditional mutualistic scatterhoarder of Arolla pine seeds.