Effect of forest fragmentation on dung beetle communities and functional consequences for plant regeneration

I compared dung beetle communities and assessed some of their functional effects (dung removal, seed burial, seedling establishment) in continuous forest with those in 1-ha and 10-ha forest fragments in Central Amazonia. I followed the fate of seeds until seedling establishment for three native tree species, using clean seeds and seeds surrounded by dung. The 1-ha fragments had half the number of dung beetle species captured in continuous forest and in 10-ha fragments. The continuous forest sites and the 1-ha fragments had similar number of individuals, but in the 10-ha fragments dung beetles were twice as abundant. Mean beetle size increased with increasing forest area. Dung removal and seed burial rates were higher in continuous forest than in forest fragments. Seed predation rates were higher in the forest fragments. In all sites, the proportion of seedlings established from seeds surrounded by dung vs clean seeds was the same, and it was the same in continuous forest vs fragments. When comparing seeds that remained on the forest floor with seeds buried by dung beetles, a higher percentage of seedlings established from the latter. Conservation programs that aim to maintain the regeneration ability of forest fragments must incorporate all the important components involved in seedling establishment; in Central Amazonia these include dung beetles as secondary dispersers. It is important that studies start measuring directly not only the first-order effects of forest fragmentation on species, but also the higher-order functional effects.     

Decreased frugivory and seed germination rate do not reduce seedling recruitment rates of <Emphasis Type="Italic">Aristotelia chilensis</Emphasis> in a fragmented forest

Habitat fragmentation reduces frugivorous bird abundance. Such a reduction may lead to a reduction in seed dispersal, thereby compromising seedling recruitment rate with far reaching consequences for plant population persistence. We assessed frugivory, seed germination, and seedling recruitment rates in a fragmented forest of central Chile by comparing a continuous forest with four forest fragments surrounded by pine plantations. Frugivory was 2.4 times higher in continuous forest than in forest fragments. Seeds eaten by birds germinated 1.7 and 3.7 times higher than non-eaten seeds from continuous forest and fragments respectively. Non-eaten seeds from continuous forest germinated 2.2 times higher than those from forest fragments, suggesting inbreeding depression. However, seedling recruitment rates at forest fragments were far higher than in continuous forest where no seedling recruited in the five years analysed. Therefore, despite forest fragmentation negatively affected frugivory, it did not translate into a decreased fitness of plants, thus highlighting the importance of considering the overall processes leading the reproductive success of plants following anthropogenic disturbances.     

Comparative habitat susceptibility to invasion by Chinese ash (<Emphasis Type="Italic">Fraxinus chinensis</Emphasis>: Oleaceae) in a tropical Andean landscape

Chinese ash (Fraxinus chinensis) is an exotic tree species that has been used in non-commercial monospecific plantations for revegetation programs in the Central Andes of Colombia. At the Otún river watershed, these plantations occur in patches intermixed with old pastures, oak forests, and successional forests. In this heterogeneous landscape, the ash has been able to invade some of its surrounding habitats. This study evaluates the invasion patterns of ash to each of these habitats and experimentally quantifies seed and seedling survivorship and seedling growth as three processes that could determine ash establishment. Of the four habitats examined, old pastures were the most vulnerable to invasion, followed by oak plantations, and successional forest. Ash plantations exhibited recruitment levels intermediate between pastures and oak plantations. Abandoned pastures showed the highest seed germination, seedling survivorship, and seedling growth. In the ash plantations, recruitment seemed negatively affected by the low number of germinated seeds, high mortality of seedlings, and low growth. Invasion in oak plantations was constrained by high seed mortality and burial of seedlings by leaf litter, although the seedlings that did survive grew fast and produced a high number of leaves. Within the successional forest, Chinese ash seeds germinated but establishment was constrained by a reduced seedling survivorship and low growth. This research offers new evidence of how different processes affecting the establishment of an invader may differ among landscape elements, and it also yields important information for the management and control of ash in these Andean landscapes.     

Inter-annual higher germination from smaller than medium-sized premontane wet forest fragments for an animal-dispersed tree species in Costa Rica

Forest fragmentation is pervasive in tropical landscapes, and animal-dispersed tree species are among the most threatened. Seed source is an important factor for active conservation and restoration efforts for such species but many studies show lower germination levels for seeds collected from small fragments compared to larger forests. However, the increasing rarity of large forests makes them difficult to be used as practical seed sources. We assessed the potential of small to medium-sized fragments (19–209 ha) to serve as seed sources for the conservation and restoration of an animal-dispersed tree Lacistema aggregatum (Lacistemataceae) in Costa Rican premontane wet forests. Germination, seedling survivorship, and growth for 2 years were quantified in a screen house environment. Two years later, more rigorous germination tests were conducted. Germination levels differed substantially among source trees. Seeds from some individuals had no germination inter-annually, suggesting that fecundity alone may be an incomplete indicator of individual fitness. Furthermore, in contrast to many previous studies, germination was better for seeds from smaller fragments for both study years. Subsequent seedling survivorship did not counteract the trends generated by germination, and the better performance of seeds from smaller fragments was retained for 2 years as ex situ seedlings. Higher seed quality in smaller fragments was associated with larger seed size in trees near forest edges, which might result from higher potential outcrossing rates. Our results suggest a previously unrecognized potential of small fragments as seed sources.     

Restoring Caribbean Dry Forests: Evaluation of Tree Propagation Techniques

Survival and height growth of tree seedlings and rooted cuttings introduced into artificially shaded and unshaded plots in a degraded dry forest were measured at intervals for nine months. Ten tree species were selected to represent a range of ecological characteristics of the dry–forest plant community on St. John, U.S. Virgin Islands. Of three propagule types – seeds, seedlings, and rooted cuttings – introduced to field plots, seedlings survived best (52%) over the initial nine-month period. Cuttings of six species rooted successfully in a shadehouse, but only two of these species survived the nine–month field experiment. Seed germination was low, under 11% for eight of ten species tested, and four species did not germinate. Subsequent mortality of seedling recruits was moderately high. Plumeria alba was the only species for which seedling height growth was not significantly greater than cutting height growth. Shading treatment (25% of full sun) significantly increased seedling survivorship (p= 0.03) but suppressed growth slightly for some species. Shading enhanced survival of seedlings produced from broadcast seeds, but not seed germination. Mortality occurred during dry periods, apparently from drought stress. Results suggest (1) that seedling introductions are the preferred propagule type (over seeding or rooted cuttings) for ecological restoration of degraded tropical dry forests, and (2) that some level of shading is required to increase the survivorship of many dry-forest species or to avert complete mortality of some species. This study suggests that early secondary dry forest may be best restored by underplanting within the existing vegetation. Sufficient shading suitable for growth of native dry-forest trees may be attained using a nurse crop of fast-growing leguminous trees.     

The effects of elevated CO2 on seed production and seedling recruitment in a sheep-grazed pasture

Seed production and seedling recruitment were measured over 2 years under ambient (360 ppm) and elevated (475 ppm) atmospheric CO₂ in a free air carbon dioxide enrichment (FACE) experiment, carried out in a sheep-grazed pasture on dry, sandy soil in New Zealand. In both years elevated CO₂ led to more dispersed seeds of the grasses Anthoxanthum odoratum, Lolium perenne and Poa pratensis, the legumes Trifolium repens and T. subterraneum and the herbs Hypochaeris radicata and Leontodon saxatilis. The increased seed dispersal in A. odoratum, H. radicata, Leontodon saxatilis and T. repens reflected both more inflorescences per unit area and more seeds per inflorescence under elevated CO₂. The increased seed dispersal in Lolium perenne, P. pratensis and T. subterraneum was due solely to more inflorescences per unit area. The number of seedlings that emerged and survived to at least 7 months of age was increased by elevated CO₂ for H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in both years and for A. odoratum and Lolium perenne in the first year. For species where increased seedling recruitment was noted, there was a significant positive correlation between seed production in summer and seedling emergence in the following autumn and winter, and sowing 200 extra seeds per species m^-2 resulted in more seedlings compared to unsown controls. Elevated CO₂ did not affect seedling survival in any species. There was no measurable effect of elevated CO₂ on canopy and soil surface conditions or soil moisture at the time of seedling emergence. The results suggest the dominant effect of elevated CO₂ on seedling recruitment in this pasture was an indirect one, reflecting effects on the number of seeds produced. The biomass of H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in the above-ground vegetation was greater under elevated than ambient CO₂. However, the size of individual seedlings and mature plants of these four species was unaffected by elevated CO₂. The results indicate an important way elevated CO₂ influenced plant species composition in this pasture was through changes in the pattern of seedling recruitment.     

Recruitment Limitation, Seedling Performance and Persistence of Exotic Tree Monocultures

Many native plant communities are replaced by exotic monocultures that may be successional stages or persistent community types. We surveyed a stand of Sapium sebiferum (Chinese Tallow Tree) that replaced tallgrass prairie in Texas and performed experiments with seeds and seedlings to determine the contributions of recruitment limitation and natural enemy release to allowing such a forest type to persist or to allowing native species to reduce Sapium dominance. The stand was dominated by Sapium, especially for mature trees (>99) and annual seed input (97) but less so for saplings (80). Field sown Sapium seeds had lower germination and survival rates than Celtis seeds. Together with the extreme dominance of Sapium in seed rain this suggests that native species are currently recruitment limited in this stand by seed supply but not by germination, early growth or survival. To investigate whether Sapium may benefit from low herbivory or diseases, we transplanted Sapium and Celtis seedlings into the forest and manipulated foliar fungal diseases and insect herbivores with sprays. As predicted, insect herbivores caused greater damage to Celtis seedlings than to Sapiumseedlings. However, suppression of insect herbivores caused significantly greater increases in survivorship of Sapium seedlings compared to Celtis seedlings. This suggests that herbivores in the understory of this Sapiumforest may significantly reduce Sapiumseedling success. Such a pattern of strong herbivore impact on seedlings growing near adult conspecifics was unexpected for this invasive species. However, even with insects and fungi suppressed, Sapium seedling performance was poor in this forest. Our results point towards Sapium as a successional species in a forest that will eventually be dominated by native trees that are currently recruitment limited but outperform Sapium in the understory.     

Establishment of chasmogamous and cleistogamous seedlings of an ant-dispersed UNDERSTORY HERB,CALATHEA MICANS (Marantaceae)

This study compares the establishment success of cleistogamous (CL, obligately selfed) and chasmogamous (CH, potentially outcrossed) seeds of the neotropical understory herb, Calathea micans (Marantaceae). In this species, CH and CL seeds are morphologically identical and are both dispersed by ants. I measured dispersal distances and analyzed seedling recruitment and seedling size for ant-planted CH and CL seeds, a protocol made possible by the fact that the seed coat of C. micans can be indelibly marked and remains attached to the base of the seedling for a few months. Seeds were taken by ants from experimental depots placed in natural conditions. In a second experiment, I planted CH and CL seeds in gap and understory sites to determine the effects of seed type and light on germination percentage, date of emergence, and seed and seedling survivorship. Ant-planted CH and CL seedlings did not differ in dispersal distance. CH and CL seeds did not differ significantly in recruitment probability and seedling size in either experiment. However, for the ant-planted seedlings, CH recruitment was higher in the understory than in the gap, while CL recruitment was uniform across light environments. In conclusion, I found a difference in the establishment success of CH and CL seeds in this understory herb, only after dispersal by ants in field conditions.     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

Forest Fragmentation Reduces Seed Dispersal of Duckeodendron cestroides, a Central Amazon Endemic

Fragmentation that alters mutualistic relationships between plants and frugivorous animals may reduce the seed dispersal of trees. We examined the effects of forest fragmentation on the distributions of seeds and seedlings of a Central Amazon endemic tree, Duckeodendron cestroides . In the dry seasons of 2002–2004, seeds and first-year seedlings were counted within wedge-shaped transects centered around Duckeodendron adults in fragments and nearby continuous forests at the Biological Dynamics of Forest Fragmentation Project. Analyses showed that fragmentation reduced seed dispersal quantity and quality. The percent and distance of dispersed seeds were both twice as great in continuous forest as in fragments. The distances of each tree's five furthest dispersed seeds were three times greater in continuous forest than fragments. Over the 3-yr study, 20 times more seeds were dispersed more than 10 m from parent crowns in continuous forest than fragments. A regression analysis showed more dispersed seeds at all distances in continuous forest than fragments. Dispersal differences were strong in 2002 and 2004, years of large fruit crops, but weak or absent in 2003, when fruit production was low. As distance from parent crowns increased, the number of seedlings declined more rapidly in fragments than continuous forest. Distance-dependent mortality between the seed and seedling stages appeared to be more important in continuous forest than fragments. This research provides ample, indirect evidence demonstrating that forest fragmentation can result in the breakdown of a seed dispersal mutualism, potentially jeopardizing the reproduction of a rare, tropical tree.     

Limits to tree species invasion in pampean grassland and forest plant communities

Factors limiting tree invasion in the Inland Pampas of Argentina were studied by monitoring the establishment of four alien tree species in remnant grassland and cultivated forest stands. We tested whether disturbances facilitated tree seedling recruitment and survival once seeds of invaders were made available by hand sowing. Seed addition to grassland failed to produce seedlings of two study species, Ligustrum lucidum and Ulmus pumila, but did result in abundant recruitment of Gleditsia triacanthos and Prosopis caldenia. While emergence was sparse in intact grassland, seedling densities were significantly increased by canopy and soil disturbances. Longer-term surveys showed that only Gleditsia became successfully established in disturbed grassland. These results support the hypothesis that interference from herbaceous vegetation may play a significant role in slowing down tree invasion, whereas disturbances create microsites that can be exploited by invasive woody plants. Seed sowing in a Ligustrum forest promoted the emergence of all four study species in understorey and treefall gap conditions. Litter removal had species-specific effects on emergence and early seedling growth, but had little impact on survivorship. Seedlings emerging under the closed forest canopy died within a few months. In the treefall gap, recruits of Gleditsia and Prosopis survived the first year, but did not survive in the longer term after natural gap closure. The forest community thus appeared less susceptible to colonization by alien trees than the grassland. We conclude that tree invasion in this system is strongly limited by the availability of recruitment microsites and biotic interactions, as well as by dispersal from existing propagule sources.     

Effects of Dispersal Pattern and Mammalian Herbivores on Seedling Recruitment for Virola michelii (Myristicaceae) in French Guiana1

The effects of dispersal pattern (seeds in small clumps vs. seeds scattered in pairs) and distance to the nearest Carapa procera (Meliaceae; a tree that produces seeds preferred by terrestrial vertebrates) on survival of seeds and seedlings were examined for the animal–dispersed tree species Virola michelii (Myristicaceae) in a mature forest‘at Paracou, French Guiana, in 1992 and 1993. We assessed the putative role of ground–dwelling mammalian herbivores, rodents, and ungulates that filter the seed shadow, acting either as dispersers or predators and thus modifying the original pattern of seed dispersal made by frugivores. We measured the effects of simulated seed burial by rodents using marked seeds and quantified the effect of protecting seeds and seedlings from ground–dwelling vertebrates on seedling germination and survival with fence exclosures in 1992. Dispersal pattern had short–term but no long–term effects on the proportion of V. michelii seeds that survived one year later as seedlings. In the short term, within six weeks, clumped seeds survived better than scattered seeds in both years. Marked seeds that were removed from their site of dispersal were eaten; rodents only rarely buried seeds of V. michelii, and seed burial reduced seed and seedling survivorship. The combined effect of the factors year and Carapa proximity significantly affected seed survival within six weeks. Although six–week seed survival was greater in 1993 than in 1992, seedling establishment was lower in 1993 than in 1992 following a lower rainfall regime during the key period of seed germination (February). One–year seed and seedling survivorship was similar between treatments and years. Seed survival and seedling establishment in V. michelii was dependent on vertebrates in the short term and on climate in the long term. Overall, seed and seedling survivorship depended on a combination of these factors.     

Burning Influences on Wiregrass (Aristida beyrichiana) Restoration Plantings: Natural Seedling Recruitment and Survival

Regeneration and expansion of Aristida beyrichiana and Aristida stricta (wiregrass) populations in remaining fire‐maintained Pinus palustris (longleaf pine) stands of the southeastern United States has become an objective of land managers. Although growing‐season fire is required for successful wiregrass seed production, studies examining naturally occurring wiregrass seedling dynamics are few. This study investigates how seedling survivorship is affected by season of burn, seedling size, time since germination, and proximity to adult plants. Restoration at this research site was begun in 1992 with the planting of containerized longleaf pine and wiregrass seedlings. Study plots were established in November 1997 after a growing‐season prescribed fire (June 1996) that resulted in successful seed production and seedling recruitment. Burn treatment plots included (1) no burn (control), (2) fire in the dormant season of the first year after germination (March 1998), (3) fire in the growing season of the first year after germination (August 1998), and (4) fire in the growing season of the second year after germination (July 1999). Seedling mortality increased with growing season burning and close proximity to planted adults. Natural seedling recruitment continued into the second year after initial seed‐drop in all plots, which verifies that wiregrass seed banking occurs for a minimum of 2 years after seed drop. Where wiregrass management objectives include population expansion, seedling recruits should be allowed 1 to 2 years post‐germination without growing season fire for successful establishment.     

Spatial vs. temporal effects on demographic and genetic structures: the roles of dispersal, masting and differential mortality on patterns of recruitment in Fagus sylvatica

Trees' long lifespan, long-distance dispersal abilities and high year-to-year variability in fecundity are thought to have pervasive consequences for the demographic and genetic structure of recruited seedlings. However, we still lack experimental studies quantifying the respective roles of spatial processes such as restricted seed and pollen dispersal and temporal processes such as mast seeding on patterns of regeneration. Dynamics of European beech (Fagus sylvatica) seedling recruitment was monitored in three plots from 2004 to 2006. Six polymorphic microsatellite genetic markers were used to characterize seedlings and their potential parents in a 7.2-ha stand. These seedlings were shown to result from 12 years of recruitment, with one predominant year of seedling recruitment in 2002 and several years without significant recruitment. Using a spatially explicit mating model based on parentage assignment, short average dispersal distances for seed (δ(s) = 10.9 m) and pollen (43.7 m < δ(p) <57.3 m) were found, but there was also a non-negligible immigration rate from outside the plot (m(s) = 20.5%; 71.6% < m(p) < 77.9%). Hierarchical analyses of seedling genetic structure showed that (i) most of the genetic variation was within plots; (ii) the genetic differentiation among seedling plots was significant (F(ST) = 2.6%) while (iii) there was no effect of year-to-year seed rain variation on genetic structure. In addition, no significant effect of genetic structure on mortality was detected. The consequences of these results for the prediction of population dynamics at ecological timescales are discussed.     

Effects of seedling size, El Niño drought, seedling density, and distance to nearest conspecific adult on 6-year survival of Ocotea whitei seedlings in Panamá

We present an analysis of the long-term survival of two cohorts of seedlings of the tropical canopy tree Ocotea whitei (Lauraceae) on a 1-ha plot of mature, lowland moist forest on Barro Colorado Island, Panamá. In 1980, we counted an even-aged cohort of seedlings that germinated in 1979, then measured and tagged survivors in 1981. We also measured and tagged a second, smaller cohort of seedlings that germinated in 1981. We followed the subsequent survival of all seedlings through 1985. Seedling mortality was phenotypically, temporally, and spatially non-random. Important correlates of non-random mortality included: (1) seedling size and age, (2) an El Niño drought, and (3) biotic neighborhood. Larger and older seedlings survived better than smaller and younger seedlings, respectively, and the El Niño-related drought of 1982-1983 was associated with elevated mortality rates. Seedling density, which was strongly correlated with the proximity to the nearest conspecific adult, increased mortality. The observed mortality patterns suggest that processes consistent with the Janzen-Connell hypothesis operate during the recruitment phase of O. whitei population dynamics. However, the processes causing the observed density- and distance-dependent mortality may vary with factors such as total seed number, seedling size, and climatic variation, making it difficult to determine whether time-integrated seedling-to-adult spacing mechanisms other than self-thinning operate on a given plant population. After 6 years in the hectare studied, survivors remained densest and most numerous underneath the adult trees. We conclude that only long-term demographic data, collected at a variety of scales on a variety of species, will ultimately answer the question: do Janzen-Connell effects contribute substantially to structuring tropical forests?     

Adaptive traits of wild barley plants of Mediterranean and desert origin

Reciprocal introduction of seeds and seedlings was used to test for local adaptation and to identify a set of co-adapted traits of Mediterranean and desert ecotypes of wild barley Hordeum spontaneum. Evidence for local adaptation was found in seedling introductions into intact environments and from ecotype colonization success in the first generation after seed dispersal. Estimates of fitness were obtained at particular stages of the life cycle (seed, seedling and adult). Experiments that manipulated the environment (vegetation removal, different plant density) demonstrated the intensity and direction of natural selection in different life history episodes, but there was no strong evidence for local adaptation under these circumstances. The observed genetically determined differences between Mediterranean and desert ecotypes can be summarized as the following: reproductive output was higher in desert plants, with smaller seeds than in Mediterranean plants. There was a higher competitive ability of Mediterranean than desert plants. Plants of desert origin had significant reductions in yield when grown in mixed stands with Mediterranean plants; no such effect was observed for plants of Mediterranean origin. Seed germination and seedling survival was lower in seeds of desert origin. This was due to both: genetically determined higher dormancy of desert seeds and a trade-off between no. of seeds and their size (directly related to seed/seedling vigour).     

Association with mature plants protects seedlings from predation in an arid grassland shrub,Gutierrezia microcephala

Summary Survivorship of Gutierrezia microcephala (Compositae) seedlings was studied in an undisturbed arid grassland and in experimental plots where various components of the natural vegetation were removed following seed germination. The major causes of seedling mortality were herbivore damage from the specialist grasshopper, Hesperotettix viridis, and drought stress associated with competition from established plants. The relative intensity of these mortality factors varied strongly with seedling size. Large seedlings had higher overall survivorship but were most likely to be killed by defoliation; most small seedlings died of drought stress.In plots where all perennial grasses were removed (leaving established G. microcephala plants), seedling survivorship was 5 times greater than in undisturbed vegetation. Surviorship in plots where both grasses and mature G. microcephala were removed was slightly poorer than in undisturbed vegetation due to a large increase in mortality from defoliation. From May–August, when the herbivore H. viridis was abundant, seedling survivorship was better in the immediate vicinity of mature conspecific plants than in plots lacking mature G. microcephala, both in the presence and absence of perennial grasses. These results provide a counter example to theories predicting that the impact of specialist herbivores on seedling recruitment is greatest in the vicinity of parent plants.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

Effects of plant competition,seed predation,and nutrient limitation on seedling survivorship of spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis>)

We measured seed germination and seedling survivorship of spotted knapweed, Centaurea stoebe, in a series of laboratory and field experiments to evaluate the efficacy of seed limitation as a management focus. This work was initiated 6 years after introduction of several biological control agents. The soil seed bank of the site used in this study contained a mean density of 5,848 seeds/m² (ranging from 0 to 16,364 seeds/m²), and 92% of the seeds isolated from soils were shriveled, discolored, and/or partially decayed. Additionally, none of the intact seeds germinated, suggesting that the viable seed bank at our field study site has been exhausted. Centaurea stoebe seeds were planted into pots under a range of soil nitrogen (N) availability, with half of the pots containing a single density of previously established seedlings of a native cool-season grass, slender wheatgrass (Elymus trachycaulus). A watering regime mimicking local precipitation was applied. Spotted knapweed exhibited large biomass responses to N addition, but the presence of grasses suppressed the ability to exploit this N. Surprisingly, low soil N conditions improved knapweed survivorship in the presence of grasses. Nevertheless, recruitment and biomass were still far below the levels reached in the absence of competition. To evaluate the effect of density on successful recruitment, Centaurea stoebe seed was introduced into a meadow at three densities matching reduced levels of seed production under the constraints of seed predators. These densities were sown with or without a seed mixture of native species, into an existing plant community lacking C. stoebe, and seedling recruitment was recorded over 2.5 years. Across all plots and densities sown (568–2,272 seeds m⁻² year⁻¹), seedling recruitment was less than 1%. The invasion potential of spotted knapweed was greatly diminished when realistic levels of plant competition and biological control limit seed production. We therefore conclude that a combination of seed limitation and shortage of ‘safe sites’ within undisturbed vegetation can limit densities of C. stoebe.     

Scatter Hoarding of Seeds Confers Survival Advantages and Disadvantages to Large-Seeded Tropical Plants at Different Life Stages

Scatter hoarding of seeds by animals contributes significantly to forest-level processes, including plant recruitment and forest community composition. However, the potential positive and negative effects of caching on seed survival, germination success, and seedling survival have rarely been assessed through experimental studies. Here, I tested the hypothesis that seed burial mimicking caches made by scatter hoarding Central American agoutis (Dasyprocta punctate) enhances seed survival, germination, and growth by protecting seeds from seed predators and providing favorable microhabitats for germination. In a series of experiments, I used simulated agouti seed caches to assess how hoarding affects seed predation by ground-dwelling invertebrates and vertebrates for four plant species. I tracked germination and seedling growth of intact and beetle-infested seeds and, using exclosures, monitored the effects of mammals on seedling survival through time. All experiments were conducted over three years in a lowland wet forest in Costa Rica. The majority of hoarded palm seeds escaped predation by both invertebrates and vertebrates while exposed seeds suffered high levels of infestation and removal. Hoarding had no effect on infestation rates of D. panamensis, but burial negatively affected germination success by preventing endocarp dehiscence. Non-infested palm seeds had higher germination success and produced larger seedlings than infested seeds. Seedlings of A. alatum and I. deltoidea suffered high mortality by seed-eating mammals. Hoarding protected most seeds from predators and enhanced germination success (except for D. panamensis) and seedling growth, although mammals killed many seedlings of two plant species; all seedling deaths were due to seed removal from the plant base. Using experimental caches, this study shows that scatter hoarding is beneficial to most seeds and may positively affect plant propagation in tropical forests, although tradeoffs in seed survival do exist.