The evolutionary ecology of nut dispersal

A variety of nut-producing plants have mutualistic seed-dispersal interactions with animals (rodents and corvids) that scatter hoard their nuts in the soil. The goals of this review are to summarize the widespread horticultural, botanical, and ecological literature pertaining to nut dispersal inJuglans, Carya, Quercus, Fagus, Castanae, Castanopsis, Lithocarpus, Corylus, Aesculus, andPrunus; to examine the evolutionary histories of these mutualistic interactions; and to identify the traits of nut-bearing plants and nut-dispersing rodents and jays that influence the success of the mutualism. These interactions appear to have originated as early as the Paleocene, about 60 million years ago. Most nuts appear to have evolved from ancestors with wind-dispersed seeds, but the ancestral form of dispersal in almonds (Prunus spp.) was by frugivorous animals that ingested fruit. Nut-producing species have evolved a number of traits that facilitate nut dispersal by certain rodents and corvids while serving to exclude other animals that act as parasites of the mutualism. Nuts are nutritious food sources, often with high levels of lipids or proteins and a caloric value ranging from 5.7 to 153.5 kJ per propagule, 10–1000 times greater than most wind-dispersed seeds. These traits make nuts highly attractive food items for dispersers and nut predators. The course of nut development tends to reduce losses of nuts to insects, microbes, and nondispersing animals, but despite these measures predispersal and postdispersal nut mortality is generally high. Chemical defenses (e.g., tannins) in the cotyledons or the husk surrounding the nut discourage some nut predators. Masting of nuts (periodic, synchronous production of large nut crops) appears to reduce losses to insects and to increase the number of nuts dispersed by animals, and it may increase cross-pollination. Scatter hoarding by rodents and corvids removes nuts from other sources of nut predation, moves nuts away from source trees where density-dependent mortality is high (sometimes to habitats or microhabitats that favor seedling establishment), and buries nuts in the soil (which reduces rates of predation and helps to maintain nut viability). The large nutrient reserves of nuts not only attract animal dispersers but also permit seedlings to establish a large photosynthetic surface or extensive root system, making them especially competitive in low-light environments (e.g., deciduous forest) and semi-arid environments (e.g., dry mountains, Mediterranean climates). The most important postestablishment causes of seedling failure are drought, insufficient light, browsing by vertebrate herbivores, and competition with forbs and grasses. Because of the nutritional qualities of nuts and the synchronous production of large nut crops by a species throughout a region, nut trees can have pervasive impacts on other members of ecological communities. Nut-bearing trees have undergone dramatic changes in distribution during the last 16,000 years, following the glacial retreat from northern North America and Europe, and the current dispersers of nuts (i.e., squirrels, jays, and their relatives) appear to have been responsible for these movements.     

花楸树种子散布、萌发与种群天然更新的关系

花楸树是我国东北林区重要的非木质资源树种,其实生天然更新不良.本文通过研究花楸树种子散布、土壤种子库及种子萌发出土过程,分析花楸树实生天然更新的影响因素.结果表明：自然散落的花楸树果实96.1%分布于母株2 m范围内,凋落物层和土壤表层（0～2 cm）的种子数占土壤种子库总数的97.0%;不同季节花楸树土壤种子库种子数量差别很大,当年11月上旬种子数量最多,达(257.7±69.2)粒·m^-2;翌年7月下旬种子数量最少,仅为(2.9±2.9)粒·m^-2;温度不是花楸树种子萌发出土过程的限制因子,0 ℃～5 ℃时幼苗出苗率达(67.5±6.6)%,但对其出苗速率影响显著.土壤含水量为50%时,花楸树出苗率最高,达(74.7±4.2)%;含水量为60%时,幼苗死亡率最低,为(32.6±0.6)%.花楸树种子的散布格局和土壤种子库的时空分布格局影响种子的萌发出土过程,进而影响其种群的天然更新.     

啮齿动物对米心水青冈种子命运的影响

有性生殖(实生更新)和无性生殖(萌生更新)是植物繁殖的两种关键方式。自然界中,个别物种同时具备两种更新方式,米心水青冈(Fagus engleriana Seem.)就是典型的物种。已有研究表明萌生更新在米心水青冈种群生活史中普遍存在,可使其占据原有生态位(生态位占据假说),但对米心水青冈实生更新的研究却未见报道。为了探索啮齿动物对米心水青冈实生更新的作用机制,笔者在神农架米心水青冈林中通过坚果摆放实验,研究啮齿动物对米心水青冈坚果传播机制的影响。结果显示,鼠类对坚果的原地捕食率高达81.22%;坚果被扩散率低,仅18.56%;坚果在原地平均留存时间为(5.25±5.95)d;坚果扩散距离为(2.63±1.60)m。研究发现,米心水青冈坚果被啮齿动物捕食率极高,贮藏率极低,且坚果被传播距离在同属植物中较小,不利于该物种更新繁殖。萌生更新在很大程度上弥补了米心水青冈实生更新繁殖方式的不足。本研究结果有助于对水青冈属植物繁殖策略的理解。     

A telemetric thread tag for tracking seed dispersal by scatter-hoarding rodents

The seeds of many tree species are dispersed more than once, and this secondary seed dispersal is believed to enhance seedling recruitment. However, the effectiveness of secondary seed dispersal has rarely been assessed because it is difficult to track seeds until they die or germinate. We describe a new technique that uses thread tags attached to radio transmitters (telemetric thread tags) to track long-distance multistep seed dispersal by scatter-hoarding rodents. These telemetric thread tags can be turned off with a magnet and are reactivated when the seed moves. This method allows for seed tracking with minimal cache disturbance or distance bias, over long time spans, multiple seed movements, and with few effects on animal behavior. We used telemetric thread tags to track seed dispersal of the palm tree Astrocaryum standleyanum in a Neotropical forest, and achieved near-complete recovery of dispersed seeds tracked over distances as far as 241?m. We were also able to record the recovery time and fate of cached seeds without disturbing caches. Neither the removal rate nor the dispersal distance differed between seeds with telemetric thread tags and thread-tagged seeds. We conclude that telemetric thread tags can be used to document secondary seed dispersal by scatter-hoarding animals with unprecedented efficacy and precision. Given the size of these tags relative to the size of seeds and their dispersers, this method is applicable to the majority of tree species that are secondarily dispersed by scatter-hoarding mammals.     

Impact of site‐induced mouse caching and transport behaviour on regeneration in Castanea crenata

Abstract. Dispersal and retrieval site selection by mice, transport distance, cache depth, and emergence and survival of seedlings of Castanea crenata (Japanese chestnut) were investigated by a magnet‐locating experiment in two habitat conditions (gap vs. forest understorey). Magnets were inserted into nuts (n= 450) and the nuts placed in the edge of forest gaps. Although wood mice (Apodemus speciosus and A. argenteus) initially buried nuts singly in shallow surface caches near the nut source, by the following spring these cached nuts were retrieved and re‐cached in larger, deeper caches farther from the source, particularly in forest understories, probably to reduce the threat of pilferage. All the nuts cached in the forest understories were consumed, but 4 seedlings emerged in gaps, apparently because of lower foraging activity in the gaps by the mice. Seed size was not correlated with cache depth or cache site selection. With increasing seed size, transport distance increased, particularly in gaps, possibly due to a greater potential energy gain (relative to handling cost to the cacher), or to attempts to prevent density‐ or mass‐dependent loss of caches by other foragers. Variable seed dispersal behaviour based on variation in seed size may influence the chances of colonization and distribution of the light‐demanding Castanea trees in mosaic landscapes and may play an important role in community organization and dynamics.     

Directed seed dispersal towards areas with low conspecific tree density by a scatter‐hoarding rodent

Scatter‐hoarding animals spread out cached seeds to reduce density‐dependent theft of their food reserves. This behaviour could lead to directed dispersal into areas with lower densities of conspecific trees, where seed and seedling survival are higher, and could profoundly affect the spatial structure of plant communities. We tested this hypothesis with Central American agoutis and Astrocaryum standleyanum palm seeds on Barro Colorado Island, Panama. We radio‐tracked seeds as they were cached and re‐cached by agoutis, calculated the density of adult Astrocaryum trees surrounding each cache, and tested whether the observed number of trees around seed caches declined more than expected under random dispersal. Seedling establishment success was negatively dependent on seed density, and agoutis carried seeds towards locations with lower conspecific tree densities, thus facilitating the escape of seeds from natural enemies. This behaviour may be a widespread mechanism leading to highly effective seed dispersal by scatter‐hoarding animals.     

Edge Effects Act Differentially on Multiple Early Regeneration Stages of a Shade‐tolerant Tree Tapirira mexicana

Forest fragmentation is pervasive in tropical landscapes, and one pathway by which fragmentation may negatively impact populations is via edge effects. Early life‐stages are particularly important for species regeneration as they act as bottlenecks, but how edge effects may act differentially on different life‐stages is unknown. This study evaluated edge effects on multiple early life‐stages of a currently common animal‐dispersed, shade‐tolerant tree Tapirira mexicana (Anacardiaceae). The study was conducted in tropical premontane wet forest fragments in a highly deforested region of Costa Rica. The stages assessed were pre‐dispersal predation, primary dispersal, post‐dispersal predation, secondary dispersal, ex situ germination, in situ seed longevity, first and second year seedling abundance, second year seedling survivorship, and basal diameter growth. Results showed that impacts of edge effects were not equal across stages, but were limited to specific stages and times. One stage which may act as a bottleneck for species regeneration was pre‐dispersal predation. Over 60 percent of the seeds were predated by larvae, and predation was higher near the edge than interior habitat. Seeds lost viability within 10 d in the forest. Germination to first year seedling stage was also lower near edges, but such effect was eliminated within a year after that. Primary dispersal, seedling survivorship, and growth were not affected by proximity to edges, and both secondary dispersal and post‐dispersal predation were rare. This study demonstrates that current population abundance may not guarantee future species persistence and the importance of considering multiple life‐stages for a comprehensive assessment of forest fragmentation effects on species regeneration.     

Experimental field test of spatial variation in rodent predation of nuts relative to distance and seed density

The spatial context in which seed predation occurs may modify the spatial structure of recruitment generated by seed dispersal. The Janzen–Connell (J-C) model predicts that granivores will exert greater pressure on the parent plant or at those sites where the density of dispersed seeds is higher. We have investigated how the probability of post-dispersal survival of Juglans australis varies with nut density across a hierarchy of spatial scales. We experimentally evaluated the survival of 3,120 nuts at three spatial scales: meso-scale (≤1.5 ha), as forest sites with two densities of fruiting J. australis individuals; intermediate scale (<0.2 ha), as individual trees with two experimental crop sizes; small scale (<0.1 m²), as microsites with two factors (number of nuts and distance from source). Nut removal coincided with seed predation, a condition that allowed us to test the density-dependent seed predation hypothesis. We found that the probability of nut survival was greater at forest sites with higher J. australis density. Nut survival was not affected by nut density in the seed shadow of individual specimens: at sites where J. australis density was greater, the proportion of surviving nuts did not differ between microsites located at different distances from the parent plant, but it was greater at microsites with greater initial nut density. Nut survival depended on the scale at which rodents responded to nut density, being negatively density dependent at the meso-scale and spatially random at intermediate and small scales. At the meso-scale, excess nut supply increased the probability of nut survival, which is in agreement with a model of granivore satiation near the seed source. Rodent satiation at the meso-scale may favour maintenance of sites with high J. australis density, where individual trees may have greater probabilities of passing their genes onto the next stage of the dispersal cycle.     

Nut predation and dispersal of Harland Tanoak Lithocarpus harlandii by scatter-hoarding rodents

Plants that use the propagule to co-opt animals as dispersal agents must balance the costs of seed predation with the benefits of dispersal. Successful post-dispersal germination is a key metric that reflects these costs and benefits. By tracking individual nuts with coded tin-tags over 3 years (2000–2003), this study quantified nut predation and dispersal of harland tanoak (Lithocarpus harlandii) by seed-caching rodents in a subtropical evergreen broadleaved forest in the Duiangyan Region of Sichuan Province, Southwest China. We found that tanoak seedlings established from rodent-generated caches in the primary stands over a 12-month post-dispersal period. Our results indicate that seed-caching rodents are effective dispersers of tanoak nuts, but dispersal effectiveness varies among years and stands, probably due to mast seeding of harland tanoak or community-level seed availability according to the predator satiation hypothesis. Some nut traits in tanoak species, e.g. large seed size, hard nut husk, lower tannin and mast seeding, are important characteristics for seed dispersal by scatter-hoarding rodents, compared with oak species with higher tannin content.     

Re-caching of acorns by rodents: Cache management in eastern deciduous forests of North America

Scatter-hoarding rodents such as tree squirrels selectively cache seeds for subsequent use in widely-spaced caches placed below the ground surface. This behavior has important implications for seed dispersal, seedling establishment, and tree regeneration. Hoarders manage these caches by recovering and eating some seeds, and moving and re-caching others. This process of re-caching, however, is poorly understood. Here, we use radio-telemetry to evaluate re-caching behavior for the management of acorn caches by rodents in eastern deciduous forests. We also test the hypothesis that as seeds are re-cached, the distance from the source increases. Radio transmitters were implanted in Northern red oak (Quercus rubra) acorns and presented to rodents in a natural setting over 3 seasons. We used radio-telemetry to track and document evidence of recovery and re-caching. We tracked a total of 102 acorns. Of the 39 radio-tagged acorns initially cached, 19 (49%) were cached on two or more occasions; one acorn was cached four times. The hypothesis that rodents move seeds to progressively greater distances from the source is not well-supported, suggesting that acorns are being moved within an individual's home range. Given the species of rodents in the study area, gray squirrels (Sciurus carolinensis) are the most likely to be responsible for the caching and re-caching events. Gray squirrels appear to engage in extensive re-caching during periods of long-term food storage, which has important implications for understanding how caching behavior influences acorn dispersal and oak regeneration.     

Survival and Scatterhoarding of Frugivores-Dispersed Seeds as a Function of Forest Disturbance

The effect of forest disturbance on survival and secondary dispersal of an artificial seed shadow (N= 800) was studied at Brownsberg Natural Park, Suriname, South America. We scattered single seeds of the frugivore‐dispersed tree Virola kwatae (Myristicaceae), simulating loose dispersal by frugivores, in undisturbed and disturbed secondary forest habitats. Seed survival rate aboveground was high (69%) within 2 wk and was negatively correlated with scatterhoarding rate by rodents, the latter being significantly lower in the undisturbed forest (9%) than in the disturbed forest (20%). Postdispersal seed predation by vertebrates was low (3%) and infestation of seeds by invertebrates was almost zero in all instances. Therefore, secondary seed dispersal by rodents in forest is not as critical for recruitment as observed among other bruchid‐infested large‐seeded species. Secondary seed dispersal by rodents may, however, facilitate seedling recruitment whether cached seeds experience greater survival than seeds remaining above ground surface.     

Insufficient Evidence of <Emphasis Type="Italic">Jatropha curcas</Emphasis> L. Invasiveness: Experimental Observations in Burkina Faso,West Africa

Biofuel plants such as Jatropha curcas L. have potential to support the livelihoods of rural communities and contribute to sustainable rural development in Africa, if risks and uncertainties are minimized. Yet, recent papers have warned of the risk of biological invasions in such tropical regions as a consequence of the introduction of exotic biofuel crops. We investigated the seed dispersal risk and invasiveness potential of both J. curcas monoculture plantations and live fences into adjacent cultivated and uncultivated land use systems in Sissili province, Burkina Faso. Invasiveness potential was assessed through (i) detecting evidence of natural regeneration in perimeters around J. curcas plantations and live fences, (ii) assessing seed dispersal mechanisms, and (iii) assessing seedling establishment potential through in situ direct seed sowing. Spontaneous regeneration around the plantation perimeters of the three sites was very low. Individual seedling density around J. curcas live fences was less than 0.01 m^?2 in all sites. Seventy percent of the seedlings were found close to the live fence and most of them derived from the same year (96 %), which indicates low seed-bank longevity and seedling survival. J. curcas can be dispersed by small mammals and arthropods, particularly rodents and ants. In some sites, such as in Onliassan, high secondary seed dispersal by animals (up to 98 %) was recorded. There were highly significant differences in germination rates between seeds at the soil surface (11 %) and those buried artificially at 1–2-cm depth (64 %). In conclusion, we failed to find convincing evidence of the spreading of J. curcas or any significant impact on the surrounding environment.     

Artificial Bird Perches for the Regeneration of Degraded Tropical Peat Swamp Forest: A Restoration Tool with Limited Potential

Forest rehabilitation activities have been initiated on degraded peatland at several sites in Southeast Asia. In order to achieve rehabilitation efficiently and on the largest possible scale, cost‐effective, transferable methods need to be established. One potential method, which has previously proven successful in both temperate and neotropical forest ecosystems, is the construction of artificial bird perches outside the forest edge. These provide resting perches for frugivorous birds, encouraging them to fly out of the forest, thus increasing seed dispersal and subsequent seedling recruitment into the degraded area. This method was trialled for the first time in degraded tropical peat swamp forest in Indonesia. The results show that the perches were used by frugivorous birds, leading to a significant increase in seed dispersal; however, seedling recruitment was not increased. The frugivorous birds using the perches were degraded zone species and dispersed mainly tree species from the degraded area. Furthermore, while some seeds of forest‐area tree species were dispersed, largely only the degraded area tree species survived to seedling stage. Neither seasonality nor distance from forest edge proved to be significant factors influencing seed dispersal or seedling recruitment, the latter highlighting that seeds were principally being dispersed from within the degraded area rather than from the forest. Although artificial perches did increase seed dispersal, their use as an actual restoration tool in the process of forest regeneration on degraded tropical peatland is limited. Furthermore, when the cost‐ and effort‐to‐area factors are considered, this method is shown to be inefficient.     

Effectiveness of rodents as local seed dispersers of Holm oaks

In this study we assessed the effectiveness of rodents as dispersers of Quercus ilex in a patchy landscape in southeastern Spain. We experimentally followed the fates of 3,200 marked and weighed acorns from dispersal through the time of seedling emergence over three years. Rodents handled about 99% of acorns, and dispersed 67% and cached 7.4% of the dispersed acorns. Most caches were recovered and consumed, and only 1.3% of the original experimental acorns were found alive in caches the following spring. Dispersal distances were short (mean = 356.2 cm, median = 157 cm) and strongly right-skewed. Heavier acorns were dispersed further and were more likely to be cached and survive than lighter acorns. All caches were in litter or soil, and each contained a single acorn. Rodents moved acorns nonrandomly, mostly to oaks and pines. Most surviving acorns were either in oaks, a poor microhabitat for oak recruitment, or shrubs, a suitable microhabitat for oak recruitment. Our results suggest that rodents, by burying a relatively high proportion of acorns singly in shrubs and pines, act as moderately effective dispersers of Q. ilex. Nonetheless, this dispersal comes at a very heavy cost.     

Re-establishing the mid-storey tree <Emphasis Type="Italic">Persoonia longifolia</Emphasis> (Proteaceae) in restored forest following bauxite mining in southern Western Australia

Persoonia longifolia is a common mid-storey species that is difficult to return to post-mining environments. This study aimed to quantify in situ emergence of P. longifolia seeds on restored areas, investigate seed cueing prior to use in restoration and assess different tree guards for increasing seedling survival and health. Initial investigations found that <1 % of seeds buried or scattered on restored areas produced seedlings. However, if seeds were cued through burial in surrounding forest, retrieved and sown on restored areas, seedling emergence increased to 24 %. Significantly more seeds emerged as seedlings when buried (14.6 %) compared to those scattered on the soil surface (2.7 %). There was no significant difference in survival between seedlings planted at 2-3 weeks of age compared with those planted at 12 months of age after 20 months in situ growth. Additionally, those seedlings planted when younger were significantly taller (29.0 ± 2.9 cm) than those that were planted at 12 months of age (4.7 ± 0.3 cm). Use of “onion bag” guards improved survival from 58.1 ± 4.0 % (no guard) to 70.8 ± 3.4 % with an onion bag guard. The use of shade cloth guards did not significantly improve survival, however plant height did increase substantially after 32 months growth (22 cm compared with 7.2 cm for no guard). These data demonstrate that consideration needs to be given to specific species requirements to improve seedling emergence and survival when attempting to return difficult to germinate species to the post-mining environment.     

啮齿动物对印度栲种子扩散的效率

从2007 年11 月到2009 年11 月,在西双版纳热带雨林中选取三棵母树,在每棵树下每年释放标记印度栲种子200 粒(三年共计1 800 粒),并追踪其命运。通过调查啮齿动物搬运和分散贮藏印度栲种子的比例,以及
查贮藏种子的微生境、贮藏点大小和扩散距离,分析贮藏种子的存活情况,进而评估啮齿动物对印度栲种子扩散的效率。结果表明,啮齿动物搬运了69.3% 的印度栲种子,被搬运的种子中18% 被分散贮藏。所有被分散贮藏的种子均被埋于落叶下或埋于土壤表层,并且大部分贮藏点仅含一粒种子。种子的扩散距离从0.5 m 到43.8 m,平均距离为7.1 m,扩散距离在年间没有显著差异。2007 年(种子密度低,啮齿动物密度高) 没有种能最终存活到实验结束,而2008 和2009 年(种子密度高,啮齿动物密度低)分别有0.3% 和1.5% 的种子存活。研究表明,啮齿动物是印度栲有效的种子扩散者,但其扩散效率很大程度上取决于森林中种子的密度和啮齿动物的丰富度。     

Incorporating insect infestation into rodent seed dispersal: better if the larva is still inside

Many nutritious seeds are commonly attacked by insects which feed on the seed reserves. However, studies have not fully explored the ecological implications of insect infestation in animal seed dispersal and subsequent plant regeneration. Our question is whether the fact that an infested seed still contains the larva or not might increase/decrease the probability of being successfully dispersed by animals. This study examines the effects of weevil-infested seeds on the natural regeneration of a rodent-dispersed oak species. Rodents showed a high ability to discriminate between sound and infested seeds, even when the larva was still inside. As a result, rodents caused differential seed dispersal for sound and infested seeds by modifying multiple aspects of the dispersal process. We found that, for the same seed weight, infested acorns with a larva still inside can contribute to natural regeneration (0.7?% of seedlings in next summer), although in comparison to sound acorns they suffered higher predation rates by rodents (both partial and complete), were removed later from the ground (less preferred), cached less frequently, and dispersed to shorter distances, which reduced their potential to colonize new environments. However, infested seeds with exit holes are notably less preferred by rodents and, when dispersed, they are mostly deposited on the litter (uncached) with shorter dispersal distances and lower emergence success. Thus, the probability that larval-holed acorns will produce viable seedlings is extremely low (null in this study). Whether infested seeds still contain a larva or not clearly determines the probability of being successfully dispersed. Premature seed drop prolongs the presence of the larva inside the acorn after seed drop, and could be a possible mechanism to allow dispersal of infested seeds.     

Termites facilitate and ungulates limit savanna tree regeneration

Both large herbivores and termites are key functional groups in savanna ecosystems, and in many savanna areas, large termite mounds (termitaria) are associated with distinct woody clusters. Studies on the effect of large mammals on tree regeneration are few, and the results are conflicting. Large herbivores have been found to be important seedling predators in some areas, but facilitate tree regeneration by outcompeting small mammals and reducing grass cover in other areas. Through the use of the experimental fencing of termite mounds and adjacent savanna areas in this study, we investigated how termites and large herbivores influence tree regeneration. Termite mounds had a higher number of seedlings, more species richness, more alpha diversity (OD) and lower evenness (E) than savanna plots. Large herbivores did not significantly affect overall seedling density, species richness, OD or E. Beta diversity was higher in savanna areas than on termitaria, and beta diversity decreased in savanna areas when herbivores were excluded. Herbivore exclusion increased the density of the 12 (40 %) most common seedling species, representing 79 % of all seedlings, and fenced plots had relatively taller seedlings than open plots. Thus, termites were the main determinants of tree regeneration in our study area, but large mammals regulated the most common species. Although our study confirms previous work suggesting that large herbivores affect tree regeneration, we found that termites were an even more important determinant. Termite impacts on tree regeneration deserve increased attention by savanna ecologists.     

Effects of seed size of wind-dispersed pines (Pinus) on secondary seed dispersal and the caching behavior of rodents

Seeds and nuts dispersed by scatter-hoarding animals are relatively large compared to propagules dispersed by other means. Possible selective forces in the evolution of large seed size include the selectivity of foraging animals and the ways that food-storing animals treat seeds and nuts after harvest. Treatment by rodents, primarily yellow pine chipmunks ( Tamias amoenus ), of four species of pine seeds that vary in size was studied in the Carson Range of western Nevada. The pines, lodgepole pine ( Pinus contorta , 8.7 mg), ponderosa pine ( P. ponderosa , 55 mg), Jeffrey pine ( P. jeffreyi , 157 mg), and sugar pine ( P. lambertiana , 213 mg), produce winged seeds that are initially wind-dispersed but are gathered by rodents and cached in the soil. Radioactive scandium-46 was used to follow the fates of seeds of all fours species placed around three source trees during autumn 1998 to 2000. Rodents gathered the seeds of all four species, but they took fewer of the lodgepole pine seeds and only six lodgepole seed caches (n=2106 total caches) were found during the three years. Among the other three species, number of seeds per cache decreased with increasing seed mass. However, the product of number of seeds per cache and seed mass was similar for all species. Sugar pine seeds were cached slightly deeper than ponderosa and Jeffrey pine seeds. For the species examined, seed size appeared to have had little effect on several other attributes, including mean dispersal distance, substrate choice, and microhabitat choice. Large size decreases wind dispersibility of pine seeds, but secondary dispersal by scatter-hoarding rodents compensates for poor wind dispersal so that total dispersibility of large-seeded pines is not compromised.     

Predation and dispersal of large and small seeds of a tropical palm

Seed size may vary greatly among individuals within plant species. What effects the extremes of this variation have for seeds taken by small mammals are poorly understood. Not all seeds removed by small mammals are necessarily eaten. Small rodents are common seed predators, but they may disperse a significant proportion of seeds by scatter hoarding them via burial. Size-dependent predation and dispersal of seeds has not been directly tested within a plant species for tropical rodents. This study tested whether or not large and small nuts of Astrocaryum mexicanum (Palmae) differed in their fates due to handling by the spiny pocket mouse Heteromys desmarestianus (Heteromyidae). Exclosures were used to give small rodents exclusive access to A. mexicanum nuts. H. desmarestianus preferentially consumed large over small A. mexicanum nuts, but cached (in burrows and by scatter hoarding) similar proportions of these nuts by size. Small nuts tended to be buried farther away from exclosures than large nuts. Although sample sizes of buried nuts were small, the rodents retrieved all buried large nuts, but 30% of the small nuts remained buried long enough to germinate. I also examined predispersal predation by insects and found that insects appear to have no size preference for A. mexicanum nuts, but insect predation appears to hinder nut development. Thus, nuts attacked by insects develop to be significantly smaller, with a low proportion of undamaged endosperm, than uninfested nuts. It is hypothesized that the preferential predation of large A. mexicanum nuts by H. desmarestianus is a response by these rodents to insect predation.