Effects of liana load,tree diameter and distances between conspecifics on seed production in tropical timber trees

Seed production in tropical timber trees is limited by abiotic resources, pollination and pre-dispersal seed predation. Resource availability is influenced by the number of competing trees and by lianas that often reach high densities in disturbed parts of tropical forests. The distance between conspecific trees affects pollination efficiency and seed predation intensity, and may therefore indirectly affect the long-term sustainability of selective logging. Here we investigate how reproductive status and the number of seeds dispersed per tree are affected by liana load, distance to the nearest conspecifics, number of competing neighbours and tree diameter in the timber trees Cariniana ianeirensis and Terminalia oblonga. The study is based on a large-scale silvicultural experiment in lowland Bolivia. We found that the reproductive status of the two species was negatively correlated with liana cover and positively with tree diameter. In C. ianeirensis the most liana-infested trees dispersed fewer seeds. In T. oblonga the intensity of pre-dispersal seed predation decreased with distance to the nearest conspecifics. There was no evidence that seed viability or seed production decreased with distance to nearest conspecifics in either species as would be expected if isolation resulted in increased self-pollination. Our results indicate that reproduction can be severely reduced in timber trees if the largest, most healthy and least liana-covered trees are logged, but that liana cutting on the remaining seed trees can considerably improve seed production. In some species seed production may be further improved by ensuring that seed trees are located far apart.     

Effects of seed quality and seed location on the removal of acorns and beechnuts

We examine whether different guilds of foragers remove seeds differentially according to seed quality (seed size and insect infestation) and seed location (habitat and microhabitat) in a mixed oak-beech forest. Video recordings indicated that the wood mouse (Apodemus sylvaticus) was first to encounter seeds. Foragers preferred acorns to beechnuts, large to small size and sound to infested. Nevertheless, infested seeds were removed by rodents even when sound seeds were present. Seeds that were not preferred by scatter-hoarding rodents remained longer on the ground and were more vulnerable to predation and desiccation (4% moisture loss per day). However, seeds that were removed by scatter-hoarders were moved away from their mother trees (96%) and cached individually (32%), increasing their moisture content (3% per day). Buried seeds, simulating scatter-hoarding behavior, experienced only a 17% removal after 4 months. Seed removal differences among habitats were not due to habitat attributes but to the spatial distribution of rodent-preferred microhabitats. Thus, a significant lower seed removal was observed under the tree canopy with no shrubs. However, seed removal in forest gaps with deadwood cover was not significantly different from the preferred microhabitat (under shrub cover). In pure beech forests, seed removal by rodents only occurred under Ilex aquifolium (the only perennial cover) and under woody debris. This study concludes that seed quality and seed location determine the contribution of different removers (predators vs. dispersers), their seed selection and their removal speed, leading to different seed fates which will eventually affect tree regeneration.     

Management of burnt wood after fire affects post-dispersal acorn predation

The management of burnt wood after a fire may affect seed predation by vertebrates due to the change produced in habitat structure. We analyze the effect of burnt wood management on post-dispersal seed predation in the Holm oak. Three plots were established in a burnt forest, with three treatments per plot: (1) non-intervention (NI, all trees left standing), (2) “partial cut plus lopping” (PCL, felling 90% of trees, cutting their main branches, leaving all the biomass in situ), and (3) “salvage logging” (SL, felling the logs for their removal and masticating the woody debris). Acorns were buried to mimic dispersal by jays or rodents two and three years after fire, with two trials per year (7200 monitored acorns), and the predation rate was evaluated until the time of seedling emergence. The spatial patterns of acorn predation were assessed by computing a transformed-Ripley's K function and Moran's I correlograms. There was a large spatial and temporal variability in acorn predation, with differences among trials, plots, and replicates within treatments and plots. Overall, PCL showed the lowest predation values (83.0% versus 87.4 in NI and 88.0 in SL). Predator species (mice versus wild boar) also differed among treatments, wild boar having a negligible effect in PCL, presumably due to the physical barrier of felled logs and branches. The results support that: (1) salvage logging offers no advantage against predators and (2) that post-fire burnt wood management alters the guild of acorn predators and may reshape the pattern of seedling establishment.     

Effects of silvicultural disturbance on acorn infestation and removal

Oak (Quercus spp.) regeneration is an important forest management goal in the central hardwood forest region of the United States. Silvicultural methods that target oak regeneration, such as the creation of canopy openings and removal of shade-tolerant midstory trees, are complicated by the impact of pre- and post-dispersal predators of oak acorns like acorn weevils (Curculio spp.) and granivorous small mammals. Understanding the effects of forest disturbance created by silviculture on these predators is important to promote successful oak regeneration. We conducted a 6-year study of acorn production, weevil infestation, and acorn removal at 113 black (Q. velutina) and white oak (Q. alba) trees impacted by three types of silvicultural treatments: adjacent to harvest openings, midstory removal (first-stage shelterwood harvest), and control (no harvest) in south-central Indiana, USA. We observed highly variable acorn production across years, but minimal impact of silvicultural treatments. Levels of weevil infestation and acorn removal by small mammal seed predators also varied greatly, and predation pressure was highest in years when acorns were scarce. Weevil infestation was reduced following midstory removal in shelterwood harvests, but probability of acorn removal by small mammals was unchanged following harvest. Damaged, germinated, and weevil-infested acorns were less likely to be removed by seed predators, suggesting additive effects of pre- and post-dispersal predators. This study emphasizes the importance of considering acorn predators in oak regeneration silviculture, and timing harvests to follow large acorn crops in order to reduce predation pressure and generate a high number of seedlings.     

Spatial and temporal patterns in seed predation as revealed by reciprocal experiments and video surveillance in neighbouring beech and spruce forests

European beech and Norway spruce are late successional and competitive species meeting each other in southern Norway. Beech is predicted to expand northwards with increased temperature, implying increased competition between the two tree species. Seed survival is a key process in this competition and here we estimate detection times and predation rates of beech and spruce seeds in adjacent beech and spruce forests by combining quantitative and qualitative approaches based on reciprocal seed predation experiments and video surveillance. These experiments were repeated over two growing seasons, and by using a video-surveillance technique, we were able to precisely reveal seed detection times and high seed predation rates, up to 15 seeds?min^?1, with bank vole and chaffinch as main predators. Moreover, the two main predators were different in their seed preferences, that is, beech seeds were exclusively removed or predated by the vole, while the finch consumed spruce seeds only. As we show high predation rates, even small variations in seed predator population densities would impact the regeneration success of beech and spruce. Because the highest predation rates were recorded for spruce seeds, it is likely that this will alter the competitive balance between the two tree species in favour of beech.     

Evaluation of regeneration potential of Pinus koraiensis in mixed pine-hardwood forests in the Xiao Xing’an Mountains, China

Large scale harvest of Korean pine(Pinus koraiensis) seeds as a food product in the mixed Korean pine-hardwood forest of northeastern China poses a serious threat to the sustainability and restoration of this endangered regional ecosystem.Seed collection over past decades greatly reduced the seed bank and subsequent seedling and sapling recruitment,and impacting a wide array of granivorous animals that rely on the pine seeds.We surveyed Korean pine seeds,including solid seeds(SS),insect consumed seeds(ICS) and other(animal) consumed(OCS) kernels,of the seed bank(forest floor and the top 10 cm of mineral soil),the seedlings and saplings from 1 m 2 sample plots in five forest types in Liangshui Nature Reserve(LNR) of the southern Xiao Xing’an Mountains in northeastern China to provide accurate information for assessing the Korean pine regeneration potential.The average number of pine seeds in the seed bank were 11.2 seeds/m 2,9.1 seeds/m 2,4.6 seeds/m 2,1.1 seeds/m 2,and 0.2 seeds/m 2 in Korean pine-basswood forest,mixed Korean pine-hardwood forest,mixed conifer-hardwood forest,white birch forests,and oak forests,respectively.In the first three forest types,percentages of SS(potentially viable seeds) were 11.2%,3.5% and 27.8%,respectively.The percentages of ICS(not viable seeds) were consistent at around 35%.The higher but variable percentages of OCS(not viable seeds) indicated high seed predation in these forests.Compared with other studies,we recorded higher percentages of seed damage,probably due to our survey approach and the increased depth of seed bank sampled in our study.Depletion of pine seeds in the seed bank greatly reduced seedling and sapling recruitment.Densities of pine seedlings varied from about 180 trees/ha in the mixed Korean pine-hardwood forest to about 5,400 trees/ha in the mixed conifer-hardwood forests and showed a high degree of spatial variation.Saplings were rare in the mixed Korean pine-hardwood forest,but ranged in the thousands in other forests.Large scale pine seed harvest has seriously threatened the sustainability of the mixed Korean pine-hardwood forest ecosystem.Scaling down the seed harvest or supplemental planting of pine saplings are urgently needed to maintain the health of the existing Korean pine forests and to restore this endangered ecosystem.     

Use of carbofuran for control of eastern white pine cone and seed insects

The major insects that destroy cones and seeds of eastern white pine, Pinus strobus L., were controlled with soil applications of the systematic insecticide carbofuran in orchards at Murphy and Morganton, NC, U.S.A. Carbofuran applied once a year at rates of 4.5, 9.0 or 13.5 g AI (active ingredient) per cm of diameter at breast height protected trees from the white pine cone beetle, Conophthorus coniperda (Schwarz), the leaffooted pine seed bug, Leptoglossus corculus (Say), the shieldbacked pine seed bug, Tetyra bipunctata (Herrich-Schäffer), and the white pine cone borer, Eucosma tocullionana Heinrich. Control was reflected by increased cone survival and increased yields of filled seeds per cone. Increasing the rate of application from 4.5 to 9.0 or 13.5 g generally improved control, but the improvement was not of statistical or practical significance. Carbofuran was ineffective against a seed chalcid, Megastigmus atedius (Walker). White pine clones varied in inherent susceptibility to insect attacks.     

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.     

Cone traits of jack pine and black spruce in young seedling seed orchards

The number of fertile and infertile scales, filled and empty seeds, cone volume, seed efficiency and the incidence of insect and disease damage to seed were evaluated for seven jack pine (Pinus banksiana Lamb) and six black spruce (Picea mariana [Mill.] B.S.P.) seedling seed orchards in northern Ontario, Canada. On average, the seed potential of jack pine and black spruce cones was 50 and 82 seeds, respectively. Cone volume and the number of fertile scales were under strong genetic control and well correlated with one another for both species. Seed efficiency values were high for jack pine (60%) but poor for black spruce (24%). The incidence of seed insect damage was less than 2.5% for both species and nil for seed diseases.     

马尾松种子园种实害虫、害鼠的研究

经观察马尾松种子园种实害虫和害鼠的种类有２１种，严重危害的有机松瘿纹，马尾松窃蠹，芽梢斑螟，松实小卷蛾、油松球果小卷蛾、微红梢斑螟和社鼠。根据马尾松生殖发育规律，观察了 主要种实害虫、害鼠的 危害顺序及其程度。     

Cone and seed production from stone pine (<Emphasis Type="Italic">Pinus pinea</Emphasis> L.) stands in Central Range (Spain)

Natural regeneration of stone pine (Pinus pinea L.) stands in the Central Range of Spain can be difficult to attain. The success of this regeneration is affected by factors such as the total amount of seed available, the short dispersal ability of the pinyon, the conditions required for germination, specific problems associated with the establishment, early survival of the seedlings due to severe summer heat and drought, competition for water and mineral resources and grazing damage. This study focuses on seed availability. The amount of available seeds depends on the number and size of the cones and the number of viable pinyons within the cones. In stone pine stands, both variables show great year to year variability. Both cone and seed production, for a given year, are also conditioned by the vigour and health of the tree, its size, the condition and attributes of the stand and the loss of seed through pests or predation. In this study, the main factors which influence cone and pinyon production are identified and a multivariate model to predict annual cone and viable seed production is developed. To consider the correlation among observations coming from the same tree, stand or year, random components are included in the model. The multivariate random structure allows for future calibration of the model for a given year from a small additional sample of observations. It is important to know the total amount of viable seed produced in a stand for a given year since regeneration cuttings for Stone pine should be concentrated in high crop years.     

Walnut hoarding by the Japanese wood mouse,Apodemus speciosus Temminck

Walnuts,Juglans ailantifolia Carriere, are large and the shell is difficult to open. Thus, seed predators in Japan are restricted mainly to the Japanese squirrel (Sciurus lis Temminck) and the wood mouse (Apodemus speciosus Temminck), which can shave away the hard shells with their sharp incisors. Previous work indicated that the squirrels are not only predators, but also disperse walnuts. In this study, 95 walnuts in Period1 (September 1996–February 1997) and 80 walnuts in Period 2 (September 1999–February 2000) each attached to a miniature radio transmitter, were placed in a feeding box, and 93 and 53 of them were transported by free-living wood mice in the respective periods. Most of the walnuts provided from September to November were scatter-hoarded in the litter layer, while from December to February, the mice fed immediately and seldom scatter-hoarded on the ground surface. In all, 66 walnuts were hoarded and 80 were eaten soon after discovery, although walnuts were eaten more frequently in Period 1 than in Period 2, which may be related to annual differences in seed crop size in the study site. The mean transport distance of hoarded seeds was 6.2 ± 5.0 m (mean ± SD,n=66). The fate of hoarded walnuts was followed from September 1996 to May 1997; 21 (75%) of the scatter-hoarded walnuts were recovered, but 7 (25%) were left uneaten. Consequently, the Japanese wood mouse also plays a role in seed-dispersion of the walnut, but the transport distance for mice was shorter than that for squirrels.     

Estimating seed quantity and quality in shortleaf pine (Pinus echinata Mill.) cones from natural stands

The quantity and quality of seeds were studied in 886 shortleaf pine cones collected from 48 trees in each of two 15-ha natural stands in western Arkansas, USA. The number of seeds per cone ranged from 0 to 102, and averaged 34.6 and 38.0 in the two stands. Between 30 and 50 percent of the seeds were sound. Overall mean values for total number and total sound seeds compare with previous work. However, the wide variation between individual trees in a stand needs to be considered when evaluating cone abundance and estimating potential seed yields. For the stands observed in this study, a sample of 65 to 85 cones would be needed to estimate the number of sound seeds per cone (within ±3 seeds) with 95 percent probability. Sampling 1 or 2 cones from each of about 3 trees per ha would provide a reliable estimate of sound seed contents.     

The Role of the White-Winged Guan (Penelope albipennis) in Seed Dispersal and Predation in Tumbesian Dry Forest,Peru

The white-winged guan (Penelope albipennis, Cracidae), thought extinct until 1979, today numbers <300 individuals. We investigated the role of the guans in seed dispersal and predation dynamics by recording seed germination from fecal samples of wild birds during the dry-season (May–December 2010). This study was conducted at Chaparri Ecological Preserve, in the critically endangered Tumbesian ecosystem on the western slope of the Andes. Field observations show that this bird is a frugivore; Cordia lutea (Boraginaceae) was the primary fruit consumed of the eight species of seeds collected from droppings. Collected seeds were sown in pots under nursery conditions and monitored for germination. Germination rates of passed seeds for Cordia lutea were compared to those that had been collected from a control group of trees. Germination was significantly different between fecal and control seeds, suggesting the plant germinated better after passing through the guans alimentary tract. The identification of flower structures, and digested seeds in feces show that guans can be effective fruit and flower predators. This study provides further information regarding the diet for conservation of the white-winged guan and suggests the bird may aid seed germination in some instances and may act as a predator in others.     

Acorn defenses to herbivory from insects: Implications for the joint evolution of resistance,tolerance and escape

Acorn predation by insects and its effects on seedling establishment were investigated among three co-occurring oak species (Quercus variabilis, Q. serrata and Cyclobalanopsis glauca) in a subtropical evergreen broadleaved forest, Southwest China. All oak species had high tannin concentration (over 10%) but differed in acorn mass and germination schedule. We hypothesized that the defensive traits in acorns (e.g. seed mass, tannins and germination schedule) act together to reduce damage from insect seed predators. Q. variabilis had significantly lower predation (51.2%) than either Q. serrata (71.4%) or C. glauca (73.8%). Seedling establishment decreased with increasing injury for all oak species studied. As much as 43% of insect infested acorns germinated and established as viable seedlings despite extensive acorn mortality due to insect seed predators. The larger seed mass of Q. variabilis appears to tolerate insect infestation and retain enough stored reserves for seed germination and early growth of seedlings. Autumn germination also benefited Q. variabilis and Q. serrata, allowing escape from both direct and ancillary effects of insect feeding by fast reserve shifting to immediate germination of mature acorns. Our results indicate that large seed mass, tannin and autumn germination act together to ameliorate effects of insect seed predation, and the joint evolution of resistance and tolerance (as well as escape through autumn germination) in acorns may be the selective consequences driven in part by interactions with insect seed predators.     

红松种实害虫生物学特性的研究

红松种实害虫危害红松的球果及种子,造成种子产量下降、质量低劣,严重的可导致红松种子绝产。在黑龙江省牡丹江林区,直接危害红松种实的害虫主要有赤松梢斑螟、冷杉梢斑螟、松梢螟、红松球果种蝇、红松球果种小蜂等。其中冷杉梢斑螟为原始红松林果梢的重要害虫,赤松梢斑螟主要以幼虫危害红松幼树的球果、嫩梢及干部,是该地区危害红松种实的主要害虫。     

Flying vs. climbing: Factors controlling arboreal seed removal in oak-beech forests

Nuts are heavy and nutritious seeds that need animals to be successfully dispersed. Most studies address nut removal by a single animal species once seeds fall onto the ground. However, nuts are also accessible before the seed drop and usually to a wide guild of seed foragers. This study examines the factors controlling arboreal seed removal in oak-beech forests within the whole guild of nut foragers. We found that seed-dispersing rodents (Apodemus sylvaticus) were the main acorn removers in the oaks (up to 3.75 m height), with a rapid seed encounter and a high removal rate. However, rodents did not climb the beech trees, probably due to their smoother bark in comparison to oak bark and/or the lower nutritional value of beechnuts with regard to acorns. Jays (Garrulus glandarius) were more abundant in oak stands (both dense and scattered) and clearly preferred acorns to beechnuts whereas nuthatches (Sitta europaea) were more abundant in beech stands and preferred beechnuts to acorns. Non-storing birds such as great tits (Parus major) also removed acorns and beechnuts, especially in the stands where oaks are dominant. Jays and rodents preferred sound seeds over insect-infested seeds but such a preference was not found for nuthatches. This study highlights that pure beech stands showed a reduced guild of arboreal nut foragers in comparison to oak stands. This different guild could probably affect the spatial patterns of seed dispersal, with a proportionally higher number of long dispersal events for acorns (mostly jay-dispersed) than for beechnuts (mostly nuthatch-dispersed). Long-distance dispersal of beechnuts (by jays) is determined by the presence of other preferred species (oaks) and their frequency of non-mast years. Seed location in different habitats strongly determines the contribution of different arboreal removers (including climbing rodents) and their removal speed, leading to a differential seed fate that will eventually affect tree regeneration. As nuthatches are sedentary birds, it is important to maintain old and dead trees where they can breed (crevices), forage (arthropods) and store seeds in order to favor beechnut dispersal and gene flow. By maintaining or favoring oak trees within beech stands we will ensure a wider guild of arboreal nut dispersers.     

Site-related influences on cone-borne inoculum and asymptomatic persistence of Diplodia shoot blight fungi on or in mature red pines

The shoot blight and canker pathogens Diplodia pinea and D. scrobiculata sporulate abundantly on cones of many pine hosts. Variation in incidence and abundance of potential inoculum from cones and frequency of asymptomatic persistence on or in shoots was examined for mature red pines in sites differing in dominant presettlement vegetation and soil type in Bayfield and Douglas counties in northern Wisconsin. Collections were made in each county from 6 plantations, 3 each in areas historically vegetated with jack pine and soils mapped as sands and three in areas historically vegetated with red pine with soils mapped as loamy sands. At each site, 5 cones were collected from each of 5 red pines and 10 shoots were collected from up to 5 red pines. Conidia from cones were quantified with a water wash and filtration technique. Diplodia species were cultured from surface-disinfested asymptomatic shoots. A species-specific PCR assay was used to identify the Diplodia species from cones and shoots. Although cones and asymptomatic shoots from each county yielded D. pinea and D. scrobiculata, D. pinea was detected more frequently. More conidia were obtained from cones from Douglas Co., where there is a history of severe shoot blight damage, than cones from Bayfield Co. In Douglas Co., more conidia were obtained from cones from plantations in areas of more sandy soil and presettlement jack pine dominance than cones from plantations in areas of less sandy soil and presettlement red pine dominance. The numbers of conidia and frequencies of cultural detection of Diplodia species from asymptomatic shoots at a site were positively correlated. These results provide evidence for site-related influences on abundance of pathogen inoculum and asymptomatic persistence on or in red pine crowns that may contribute to differences in frequency and severity of damage from Diplodia shoot blight.     

Timing of shedding seeds and cones, and production in different stands of Scots pines at Abernethy Forest, Scotland

A study was carried out over 11 seed years on the timing ofshedding of seeds and cones, and annual seed fall and cone productionin three stands of native Scots pinewood and a Scots pine plantationin Abernethy Forest, Scotland. Peaks in seed fall took placemainly in May, and cones were shed mainly between June and August.There were few residual seeds remaining in shed cones. Synchronizedpeaks in seed fall and cone production (mast years) took placeat 3-year intervals across the different stands. The differencebetween cohorts of high and low cone production ranged fromfactors of 5 to 20 among sites. Coefficients of variation forcone production ranged from 62 to 84 per cent among sites. Therewere no significant differences in cone production among sites,but there were site-related differences in seed fall. The largercanopy cover in the plantation probably accounted for the higherseed fall per square metre there, though variations in the amountof seed eaten by birds and mammals may also have been important.Canopy cover needs to be considered when converting cone densitiesunder crowns to cone density per unit of woodland area. A similarcalculation is difficult for seeds because they are lighterthan cones and many fall outside the area under the crowns.The results are discussed in relation to the potential for treeregeneration and the availability of food for birds and mammalsprior to seed dispersal.     

Tree density and site quality influence on Pinus halepensis Mill. reproductive characteristics after large fires

In Spain, many Pinus halepensis Mill. forests have been seriously affected by significant forest fires in the past decade, in 1994 alone, more than 100 000 ha were burned in Eastern Spain. In order to study the reproductive characteristics of P. halepensis after forest fires, six locations were selected in four areas affected by serious fires in the summer of 1994, and the following different precipitation zones were studied: dry-subhumid, dry and semi-arid. Ten years after the fires, data relevant to the production of pine cones: serotinous (grey), mature (brown), immature (green) and opened cones, was collected from areas with natural post-fire regeneration. Various cone and seed characteristics, such as pine cone seed number and weight, and germination percentage, were measured. The results showed greater production of cones and strobili in high-density sites. The biggest cone sizes (volume) and seed number per cone were related to site quality (dry-subhumid precipitation zone). Also, viability and germination percentages were higher with better site quality, with significant differences in values for serotinous and non-serotinous cones. Despite these differences, the canopy seed bank was large enough to ensure regeneration in this area for this age stand.