Seed predation and selection exerted by a seed predator influence subalpine tree densities

Strongly interacting species often have pronounced direct and indirect effects on other species. Here we focus of the effects of pine squirrels (Tamiasciurus spp.), which are a dominant pre-dispersal seed predator of many conifers including limber pines (Pinus flexilis) and whitebark pines (P. albicaulis). Pine squirrels depress seed abundance by harvesting most limber and whitebark pine cones on their territories. Pine squirrels further reduce seed availability for Clark's Nutcrackers (Nucifraga columbiana), the primary seed disperser of these pines, because selection exerted by pine squirrels has reduced the number of seeds per cone and causes seeds to be less accessible. We predicted that, if fewer seeds were available for dispersal by nutcrackers, pine recruitment should be suppressed in areas with pine squirrels. In support of this prediction, stand densities were about two times greater in areas where pine squirrels are absent than in areas where they are present. Alternative explanations that we considered do not account for these differences; however, precipitation may limit stand densities in the absence of seed limitation by pine squirrels. In sum, pine squirrels apparently depress limber and whitebark pine stand densities, with the potential for ecosystem impacts because these pines are foundation species within Western subalpine ecosystems.     

Soil vs. canopy seed storage and plant species coexistence in species-rich Australian shrublands

The fire-prone shrublands of southwestern Australia are renowned for their high plant species diversity and prominence of canopy seed storage (serotiny). We compared species richness, abundance, and life history attributes for soil and canopy seed banks in relation to extant vegetation among four sites with different substrate conditions and high species turnover (50-80%) to identify whether this unusual community-level organization of seed storage might contribute to maintenance of high species richness. Soil seed bank (SSB) densities were low to moderate (233-1435 seeds/m2) compared with densities for other Mediterranean-type vegetation and were lowest for sites with highest canopy seed bank (CSB) species richness and lowest nutrient availability, but not richness or abundance of resprouters. Annuals were infrequent in the lowest nutrient sites, but there was no evidence that small SSB size was due to low seed inputs or a trade-off between seed production/storage and seed size in response to low nutrient availability. Sorensen's similarity between SSB and extant vegetation was 26-43% but increased to 54-57% when the CSB was included, representing levels higher than reported for most other ecosystems. Resprouting species were well represented in both the SSB and CSB, and there was no evidence for lower seed production in resprouters than in non-sprouters overall. The SSB and CSB held no species in common and were characterized by markedly different seed dispersal attributes, with winged or small seeds in the CSB and seeds dispersed by ants, birds, and wind (though none with wings) in the SSB. There was no evidence of spatial differentiation in the distribution of seeds of SSB species between vegetated and open microsites that might facilitate species coexistence, but most woody non-sprouters showed aggregation at scales of 1-2 m, implying limited seed dispersal. High similarity between overall seed bank (SSB + CSB) and extant species composition, high number of resprouting species, and seed dispersal processes before (SSB) and after fire (CSB) leading to differential spatial aggregation of post-fire recruits from the two seed bank types may buffer species composition against rapid change and provide a mechanism for maintaining species coexistence at the local scale.     

Hunting Increases Dispersal Limitation in the Tree Carapa procera, a Nontimber Forest Product

Abstract:  The sustainability of seed extraction from natural populations has been questioned recently. Increased recruitment failure under intense seed harvesting suggests that seed extraction intensifies source limitation. Nevertheless, areas where more seeds are collected tend to also have more intense hunting of seed-dispersing animals. We studied whether such hunting, by limiting disperser activity, could cause quantitative dispersal limitation, especially for large crops and for crops in years of high seed abundance. In each of four Carapa procera (Meliaceae) populations in French Guiana and Surinam, two with hunting and two without, we compared seed fate for individual trees varying in crop size in years of high and low population-level seed abundance. Carapa seeds are a nontimber forest product and depend on dispersal by scatter-hoarding rodents for survival and seedling establishment. Hunting negatively affected the proportion of seeds dispersed and caused greater numbers of seeds to germinate or be infested by moths below parent trees, where they would likely die. Hunting of seed-dispersing animals disproportionally affected large seed crops, but we found no additional effect of population-level seed abundance on dispersal rates. Consistently lower rates of seed dispersal, especially for large seed crops, may translate to lower levels of seedling recruitment under hunting. Our results therefore suggest that the subsistence hunting that usually accompanies seed collection is at the cost of seed dispersal and may contribute to recruitment failure of these nontimber forest products. Seed extraction from natural populations may affect seedling recruitment less if accompanied by measures adequately incorporating and protecting seed dispersers.     

Birds as Suppliers of Seed Dispersal in Temperate Ecosystems: Conservation Guidelines from Real‐World Landscapes

Abstract: Seed dispersal by animals is considered a pivotal ecosystem function that drives plant‐community dynamics in natural habitats and vegetation recovery in human‐altered landscapes. Nevertheless, there is a lack of suitable ecological knowledge to develop basic conservation and management guidelines for this ecosystem service. Essential questions, such as how well the abundance of frugivorous animals predicts seeding function in different ecosystems and how anthropogenic landscape heterogeneity conditions the role of dispersers, remain poorly answered. In three temperate ecosystems, we studied seed dispersal by frugivorous birds in landscape mosaics shaped by human disturbance. By applying a standardized design across systems, we related the frequency of occurrence of bird‐dispersed seeds throughout the landscape to the abundance of birds, the habitat features, and the abundance of fleshy fruits. Abundance of frugivorous birds in itself predicted the occurrence of dispersed seeds throughout the landscape in all ecosystems studied. Even those landscape patches impoverished due to anthropogenic disturbance received some dispersed seeds when visited intensively by birds. Nonetheless, human‐caused landscape degradation largely affected seed‐deposition patterns by decreasing cover of woody vegetation or availability of fruit resources that attracted birds and promoted seed dispersal. The relative role of woody cover and fruit availability in seed dispersal by birds differed among ecosystems. Our results suggest that to manage seed dispersal for temperate ecosystem preservation or restoration one should consider abundance of frugivorous birds as a surrogate of landscape‐scale seed dispersal and an indicator of patch quality for the dispersal function; woody cover and fruit resource availability as key landscape features that drive seedfall patterns; and birds as mobile links that connect landscape patches of different degrees of degradation and habitat quality via seed deposition.     

Incorporating animal behavior into seed dispersal models: implications for seed shadows

Seed dispersal fundamentally influences plant population and community dynamics but is difficult to quantify directly. Consequently, models are frequently used to describe the seed shadow (the seed deposition pattern of a plant population). For vertebrate-dispersed plants, animal behavior is known to influence seed shadows but is poorly integrated in seed dispersal models. Here, we illustrate a modeling approach that incorporates animal behavior and develop a stochastic, spatially explicit simulation model that predicts the seed shadow for a primate-dispersed tree species (Virola calophylla, Myristicaceae) at the forest stand scale. The model was parameterized from field-collected data on fruit production and seed dispersal, behaviors and movement patterns of the key disperser, the spider monkey (Ateles paniscus), densities of dispersed and non-dispersed seeds, and direct estimates of seed dispersal distances. Our model demonstrated that the spatial scale of dispersal for this V. calophylla population was large, as spider monkeys routinely dispersed seeds >100 m, a commonly used threshold for long-distance dispersal. The simulated seed shadow was heterogeneous, with high spatial variance in seed density resulting largely from behaviors and movement patterns of spider monkeys that aggregated seeds (dispersal at their sleeping sites) and that scattered seeds (dispersal during diurnal foraging and resting). The single-distribution dispersal kernels frequently used to model dispersal substantially underestimated this variance and poorly fit the simulated seed-dispersal curve, primarily because of its multimodality, and a mixture distribution always fit the simulated dispersal curve better. Both seed shadow heterogeneity and dispersal curve multimodality arose directly from these different dispersal processes generated by spider monkeys. Compared to models that did not account for disperser behavior, our modeling approach improved prediction of the seed shadow of this V. calophylla population. An important function of seed dispersal models is to use the seed shadows they predict to estimate components of plant demography, particularly seedling population dynamics and distributions. Our model demonstrated that improved seed shadow prediction for animal-dispersed plants can be accomplished by incorporating spatially explicit information on disperser behavior and movements, using scales large enough to capture routine long-distance dispersal, and using dispersal kernels, such as mixture distributions, that account for spatially aggregated dispersal.     

Changes in seed dispersal processes and the potential for between-patch connectivity for an arid land daisy

Dispersal is a major and critical process in population biology that has been particularly challenging to study. Animals can have major roles in seed dispersal even in species that do not appear specifically adapted to animal-aided dispersal. This can occur by two processes: direct movement of diaspores by animals and modification of landscape characteristics by animals in ways that greatly influence dispersal. We exploited the production of large, persistent dispersal structures (seed heads, henceforth) by Erodiophyllum elderi (Asteraceae), a daisy from arid Australia, to further understand secondary dispersal. Seed head dispersal on and off animal tracks in eight E. elderi patches was monitored for 9.5 months by periodically recording the location of marked seed heads. Sites were located inside a reserve that excludes sheep but not kangaroos, and in a nearby area with both kangaroos and sheep. The distance moved and likelihood of seed head movement was higher in areas with sheep, and especially along animal tracks. There was clear evidence that seed heads were channeled down animal tracks during large rainfall events. Seed head dispersal away from patches occurred to a limited extent via their physical contact with sheep and potentially via wind dispersal. Thus, the advantages of this study system allowed us to demonstrate the two postulated effects of herbivores on dispersal via direct movement of seed heads, and two distinct indirect effects through landscape modification by herbivores from the creation of animal tracks and the denudation of vegetation.     

Flying foxes cease to function as seed dispersers long before they become rare

Rare species play limited ecological roles, but particular behavioral traits may predispose species to become functionally extinct before becoming rare. Flying foxes (Pteropodid fruit bats) are important dispersers of large seeds, but their effectiveness is hypothesized to depend on high population density that induces aggressive interactions. In a Pacific archipelago, we quantified the proportion of seeds that flying foxes dispersed beyond the fruiting canopy, across a range of sites that differed in flying fox abundance. We found the relationship between ecological function (seed dispersal) and flying fox abundance was nonlinear and consistent with the hypothesis. For most trees in sites below a threshold abundance of flying foxes, flying foxes dispersed < 1% of the seeds they handled. Above the threshold, dispersal away from trees increased to 58% as animal abundance approximately doubled. Hence, flying foxes may cease to be effective seed dispersers long before becoming rare. As many species' populations decline worldwide, identifying those with threshold relationships is an important precursor to preservation of ecologically effective densities.     

Neighborhood effects on seed dispersal by frugivores: testing theory with a mistletoe-marsupial system in Patagonia

The outcome of the dispersal process in zoochorous plants is largely determined by the behavior of frugivorous animals. Recent simulation studies have found that fruit removal rates and mean dispersal distances are strongly affected by fruiting plant neighborhoods. We empirically tested the effects of conspecific fruiting plant neighborhoods, crop sizes, and plant accessibility on fruit removal rates and seed dispersal distances of a mistletoe species exclusively dispersed by an arboreal marsupial in Northern Patagonia. Moreover, in this study, we overcome technical limitations in the empirical estimation of seed dispersal by using a novel 15N stable isotope enrichment technique together with Bayesian mixing models that allowed us to identify dispersed seeds from focal plants without the need of extensive genotyping. We found that, as predicted by theory, plants in denser neighborhoods had greater fruit removal and shorter mean dispersal distances than more isolated plants. Furthermore, the probability of dispersing seeds farther away decreased with neighborhood density. Larger crop sizes resulted in larger fruit removal rates and smaller probabilities of longer distance dispersal. The interplay between frugivore behavioral decisions and the spatial distribution of plants could have important consequences for plant spatial dynamics.     

Behavior rather than diet mediates seasonal differences in seed dispersal by Asian elephants

Digestive physiology and movement patterns of animal dispersers determine deposition patterns for endozoochorously dispersed seeds. We combined data from feeding trials, germination tests, and GPS telemetry of Asian elephants (Elephas maximus) to (1) describe the spatial scale at which Asian elephants disperse seeds; (2) assess whether seasonal differences in diet composition and ranging behavior translate into differences in seed shadows; and (3) evaluate whether scale and seasonal patterns vary between two ecologically distinct areas: Sri Lanka's dry monsoon forests and Myanmar's (Burma) mixed-deciduous forests. The combination of seed retention times (mean 39.5 h, maximum 114 h) and elephant displacement rates (average 1988 m in 116 hours) resulted in 50% of seeds dispersed over 1.2 km (mean 1222-2105 m, maximum 5772 m). Shifts in diet composition did not affect gut retention time and germination of ingested seeds. Elephant displacements were slightly longer, with stronger seasonal variation in Myanmar. As a consequence, seed dispersal curves varied seasonally with longer distances during the dry season in Myanmar but not in Sri Lanka. Seasonal and geographic variation in seed dispersal curves was the result of variation in elephant movement patterns, rather than the effect of diet changes on the fate of ingested seeds.     

Experimental manipulation of seed shadows of an Afrotropical tree determines drivers of recruitment

The loss of animals in tropical forests may alter seed dispersal patterns and reduce seedling recruitment of tree species, but direct experimental evidence is scarce. We manipulated dispersal patterns of Manilkara mabokeensis, a monkey-dispersed tree, to assess the extent to which spatial distributions of seeds drive seedling recruitment. Based on the natural seed shadow, we created seed distributions with seeds deposited under the canopy ("no dispersal"), with declining density from the tree ("natural dispersal"), and at uniform densities ("good dispersal"). These distributions mimicked dispersal patterns that could occur with the extirpation of monkeys, low levels of hunting, and high rates of seed dispersal. We monitored seedling emergence and survival for 18 months and recorded the number of leaves and damage to leaves. "Good dispersal" increased seedling survival by 26%, and "no dispersal" decreased survival by 78%, relative to "natural dispersal." Using a mixed-effects survival model, we decoupled the distance and density components of the seed shadow: seedling survival depended on the seed density, but not on the distance from the tree. Although community seedling diversity tended to decrease with longer dispersal distances, we found no conclusive evidence that patterns of seed dispersal influence the diversity of the seedling community. Local seed dispersal does affect seedling recruitment and survival, with better dispersal resulting in higher seedling recruitment; hence the loss of dispersal services that comes with the reduction or extirpation of seed dispersers will decrease regeneration of some tree species.     

Antennal responses of the western pine beetle, <Emphasis Type="Italic">Dendroctonus brevicomis</Emphasis> (Coleoptera: Curculionidae), to stem volatiles of its primary host, <Emphasis Type="Italic">Pinus ponderosa</Emphasis>, and nine sympatric nonhost angiosperms and conifers

Summary. Stem volatile extracts from ten trees that are sympatric with the western pine beetle, Dendroctonus brevicomis LeConte (Coleoptera: Curculionidae) were assayed by gas chromatographic-electroantennographic detection analysis (GC-EAD). The extracts were from the primary host, ponderosa pine, Pinus ponderosa Dougl. ex Laws. (Pinaceae); two nonhost angiosperms, California black oak, Quercus kelloggii Newb. (Fagaceae), and quaking aspen, Populus tremuloides Michx. (Salicaceae); and seven nonhost conifers, white fir, Abies concolor (Gord. & Glend.) Lindl. ex Hildebr. (Pinaceae), incense cedar, Calocedrus decurrens (Torr.) Florin (Cupressaceae), Sierra lodgepole pine, P. contorta murrayana Grev. & Balf. (Pinaceae), Jeffrey pine, P. jeffreyi Grev. & Balf. (Pinaceae), sugar pine, P. lambertiana Dougl. (Pinaceae), Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco (Pinaceae), and mountain hemlock, Tsuga mertensiana (Bong.) Carr. (Pinaceae). Sixty-four compounds were identified from the ten trees, 42 of which elicited antennal responses in D. brevicomis, usually in both sexes. In addition, several synthetic compounds, including a number of the antennally-active compounds from the extracted trees and some bark beetle pheromone components, elicited antennal responses in a manner similar to that observed with the extracts. Of the antennally-active compounds known to be present in trees sympatric with D. brevicomis, only geraniol was unique to its host. Four antennally-active compounds were found in the host and in other conifers; five compounds were found only in nonhost conifers; eight compounds were found in either or both of the nonhost angiosperms; eight compounds were found in either or both of the angiosperms and in nonhost conifers, but not in the host; and 19 were found in both the host and in angiosperms and/or nonhost conifers. Several bark beetle pheromone components were found in the stem volatile extracts. Conophthorin was identified from both nonhost angiosperms; exo-brevicomin was identified in A. concolor; verbenone was identified from a number of nonhost conifers; and chalcogran was identified from P. tremuloides. The number of nonhost volatile chemicals that D. brevicomis encounters and is capable of detecting, and the diversity of sources from which they emanate, highlight the complexity of the olfactory environment in which D. brevicomis forages. This provides a basis for further work related to chemically-mediated aspects of foraging in this insect and perhaps other coniferophagous bark beetles, and highlights the need to consider foraging context in the design and implementation of semiochemical-based management tactics for tree protection.     

Introduced Birds and the Fate of Hawaiian Rainforests

Abstract:  The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.     

The importance of seed dispersal in the Alexandria Coastal Dunefield, South Africa

The endozoochorous dispersal of seeds by mammals and birds between distinct vegetation communities was assessed to determine the importance of these processes in coastal dune field management. Isolated pockets of thicket vegetation (bush-pockets) within a large coastal dune field provided the opportunity to study vertebrate seed dispersal and its contribution to their origin and maintenance. Mammalian and avian faeces were collected for the quantification of seeds dispersed via endozoochory. Birds and mammals showed considerable overlap, dispersing intact seeds of 17 and 29 plant species, respectively, but mammals dispersed a greater diversity and size range than birds. Extrapolation of mammalian faecal data indicates an annual input of 23 million intact seeds to the dune field. Significantly more seeds are deposited by mammals and birds in the bush-pockets than on open sand, and birds deposited greater numbers of seeds nearer the seed source. Zoochory appears to be critical for the maintenance of the bush-pocket habitats through the dispersal of climax woody plant species into the dune field. Directional dispersal by birds and mammals to the bush-pockets is considered to be responsible for the maintenance and possible origin of these bush-pockets. The high number of exotic plant propagules dispersed by both avian and mammalian zoochory highlights the importance of management of the Alexandria Coastal Dunefield (ACD) beyond the reserve boundaries. In a dynamic system such as the ACD which is within a declared nature reserve, the continued existence of the bush-pockets may depend on the maintenance, beyond the reserve boundaries, of a reservoir of not only plant material but vertebrate dispersers as well.     

Nonredundancy in the dispersal network of a generalist tropical forest tree

Plant species with generalized dispersal mutualisms are considered to be robust to local frugivore extinctions because of redundancy between dispersal agents. However, real redundancy can only occur if frugivores have similar foraging and ranging patterns and if fruit is a limiting resource. We evaluated the quantitative and qualitative contributions of seed dispersers for an endochorus mast-fruiting species, Prunus javanica (Rosaceae) in Khao Yai National Park, Thailand, to evaluate the potential redundancy of dispersers. Data were collected from tree watches, seed/fruit traps, and seed transects under and away from fruiting trees, feeding and seed deposition by gibbons (Hylobates lar), and evaluations of seed and first-year seedling survival. We identified three clusters of dispersers within the network. Most (>80%) frugivore species observed were small birds and squirrels that were not functional dispersers, dropping most seeds under or very near the tree crown, where seedling survival was ultimately nil. Monkeys (Macaca leonina) were low-quality, short-range dispersers, but they dispersed large numbers of seeds and were responsible for 67% of surviving first-year seedlings. Gibbons and Oriental Pied Hornbills (Anthracoceros albirostris) handled few fruits, but they provided the highest quality service by carrying most seeds away from the canopy to medium and long distances, respectively. Although there was overlap in the deposition patterns of the functional dispersers, they displayed complementary, rather than redundant, roles in seed dispersal. Satiation of all functional dispersers further limited their capacity to "replace" one another. Redundancy must be evaluated at the community level because each type of disperser may shift to different species in the non-masting years of P. javanica. Our results underscore the need for research on broader spatial and temporal scales, which combines studies of dispersal and plant recruitment, to better understand mechanisms that maintain network stability.     

Dispersal of seeds by the tropical sea breeze

Given the dependence of most wind-pollinated and wind-dispersed species on low relative humidity (RH) for abscission, and the minimization of RH in the early afternoon, there ought to be a marked directional bias in seed dispersal at sites with a strong local diurnal circulation. We filmed the abscission of seeds of five wind-dispersed tropical species near the coast of Mexico (Jalisco). We found that (1) most abscission occurred during the period from 10:00 to 17:00 hours; (2) there was a strong bias for landward dispersal due to the midday sea breeze; (3) the little nocturnal dispersal that occurred was toward the sea (due to the night land breeze); (4) there was no abscission in the absence of wind (i.e., indoors) except for one species; and (5) holding relative humidity constant, the proportion of seeds diurnally abscising is strongly correlated with horizontal wind speed. We predict that a similar bias (this time for upslope dispersal) for xerochastic dispersal will occur in complex terrain. We conclude that in coastal and mountainous terrain, dispersal models (and inverse modeling efforts) must include a directionality term.     

Modeling the Effect of Climate Change on the Distribution of Oak and Pine Species of Mexico

Abstract:  We examined the vulnerability of 34 species of oaks ( Quercus ) and pines ( Pinus ) to the effects of global climate change in Mexico. We regionalized the HadCM2 model of climate change with local climatic data (mean annual temperature and rainfall) and downscaled the model with the inverse distance-weighted method. Databases of herbaria specimens, genetic algorithms (GARP), and digital covers of biophysical variables that affect oaks and pines were used to project geographic distributions of the species under a severe and conservative scenario of climate change for the year 2050. Starting with the current average temperature of 20.2 °C and average precipitation of 793 mm, under the severe warming scenario mean temperature and precipitation changed to 22.7 °C and 660 mm, respectively, in 2050. For the conservative warming scenario, these variables shifted to 21.8 °C and 721 mm. Responses to the different scenarios of climate change were predicted to be species-specific and related to each species climate affinity. The current geographic distribution of oaks and pines decreased 7–48% and 0.2–64%, respectively. The more vulnerable pines were Pinus rudis , P. chihuahuana , P. oocarpa , and P. culminicola , and the most vulnerable oaks were Quercus crispipilis , Q. peduncularis , Q. acutifolia , and Q. sideroxyla . In addition to habitat conservation, we think sensitive pine and oak species should be looked at more closely to define ex situ strategies (i.e., seed preservation in germplasm banks) for their long-term conservation. Modeling climatic-change scenarios is important to the development of conservation strategies.     

Monoterpene chemodiversity of ponderosa pine in relation to herbivory and bark beetle colonization

Summary. Ponderosa pine, Pinus ponderosa Laws. (Pinaceae), forests in Arizona have suffered from a nine-year period of drought and bark beetle, Ips lecontei Swaine (Coleoptera: Scolytidae), outbreaks. Abiotic and biotic stress in ponderosa pine results in the induced synthesis of certain monoterpenes that may in turn affect bark beetle behavior and survival. In this study, we investigate whether induced monoterpene production could result in a different monoterpene composition that remains stored in the needles or the trunk resin of the tree. Needle and resin samples in addition to trunk cores were collected from ponderosa pines at three locations in Arizona. Ungulate browsing induced a significant increase in limonene (P=0.010) and in chemodiversity (P=0.009), a measure of the evenness of distribution among the monoterpenes present in needles. We compared the level of ‘stress’ of the trees by measuring the thickness of annual rings in living trees and those that were killed by bark beetles. Where drought occurred, the spacing of annual rings from the last 10 years of trees killed by bark beetles was significantly smaller (P=0.020) compared to living trees. There was no difference in the monoterpene composition between the core sections of closest spacing of annual rings (stressed years) compared to the sections of widest spacing, which indicates that monoterpenes are distributed evenly throughout the extended resin system. In the area where the degree of drought was less overall, none of the individual monoterpenes present in the resin was related to bark beetle killed trees. However, about half the living pines had resin in which one of the major monoterpenes (α-pinene, Δ³-carene, and limonene) was absent, and these trees had a lower monoterpene chemodiversity compared to trees killed by bark beetles. Trees with these three major monoterpenes, corresponding to the average relative proportion in living pines at that location, may sustain higher selection and colonization by bark beetles.     

Modeling seed dispersal distances: implications for transgenic Pinus taeda.

Predicting forest-tree seed dispersal across a landscape is useful for estimating gene flow from genetically engineered (GE) or transgenic trees. The question of biocontainment has yet to be resolved, although field-trial permits for transgenic forest trees are on the rise. Most current field trials in the United States occur in the Southeast where Pinus taeda L., an indigenous species, is the major timber commodity. Seed dispersal distances were simulated using a model where the major determinants were: (1) forest canopy height at seed release, (2) terminal velocity of the seeds, (3) absolute seed release, and (4) turbulent-flow statistics, all of which were measured or determined within a P. taeda plantation established from seeds collected from wild forest-tree stands at the Duke Forest near Durham, North Carolina, USA. In plantations aged 16 and 25 years our model results showed that most of the seeds fell within local-neighborhood dispersal distances, with estimates ranging from 0.05 to 0.14 km from the source. A fraction of seeds was uplifted above the forest canopy and moved via the long-distance dispersal (LDD) process as far as 11.9-33.7 km. Out of 10(5) seeds produced per hectare per year, roughly 440 seeds were predicted to be uplifted by vertical eddies above the forest canopy and transported via LDD. Of these, 70 seeds/ha traveled distances in excess of 1 km from the source, a distance too great to serve as a biocontainment zone. The probability of LDD occurrence of transgenic conifer seeds at distances exceeding 1 km approached 100%.