Dispersión Primaria de Semillas por Primates y Dispersión Secundaria por Escarabajos Coprófagos en Tikal,Guatemala1

We linked primary dispersal by spider monkeys (Ateles geoffroyi) and howler monkeys (Alouatta pigra) to post‐dispersal seed fate by studying the effects of dung type and defecation pattern on secondary seed dispersal by dung beetles. First, we described the defecation patterns for both primate species. Howler monkeys generally defecated in groups (88% of observed defecations), with each individual producing on average 31 g of dung, resulting in a large area of the forest floor (31 m²) covered by large amounts of dung (clumped spatial pattern). Spider monkeys generally (96% of observed defecations) defecated individually, each individual producing an average of 11 g of dung, resulting in a small area of the forest floor (2 m²) covered by small amounts of dung (scattered spatial pattern). Secondly, we captured dung beetles using as bait the dung of both primate species, to detect differences in the assemblages of these secondary seed dispersers attracted to the dung of both primates. More individual dung beetles, but not more species, were attracted to howler monkey dung than to spider monkey dung. Finally, we assessed experimentally (using plastic beads as seed mimics) how dung type (Ateles vs. Alouatta) and defecation pattern (scattered vs. clumped) affect secondary seed dispersal by dung beetles. We found that post‐dispersal seed fate was affected by dung type, with more seeds being buried when present in howler monkey dung, than in spider monkey dung, but was not affected by defecation pattern. It is important to consider post‐dispersal processes, such as secondary seed dispersal by dung beetles, when comparing species of primary dispersers.     

Sleeping Sites and Latrines of Spider Monkeys in Continuous and Fragmented Rainforests: Implications for Seed Dispersal and Forest Regeneration

Spider monkeys (Ateles geoffroyi) use sites composed of one or more trees for sleeping (sleeping sites and sleeping trees, respectively). Beneath these sites/trees they deposit copious amounts of dung in latrines. This behavior results in a clumped deposition pattern of seeds and nutrients that directly impacts the regeneration of tropical forests. Therefore, information on the density and spatial distribution of sleeping sites and latrines, and the characteristics (i.e., composition and structure) of sleeping trees are needed to improve our understanding of the ecological significance of spider monkeys in influencing forest composition. Moreover, since primate populations are increasingly forced to inhabit fragmented landscapes, it is important to assess if these characteristics differ between continuous and fragmented forests. We assessed this novel information from eight independent spider monkey communities in the Lacandona rainforest, Mexico: four continuous forest sites and four forest fragments. Both the density of sleeping sites and latrines did not differ between forest conditions. Latrines were uniformly distributed across sleeping sites, but the spatial distribution of sleeping sites within the areas was highly variable, being particularly clumped in forest fragments. In fact, the average inter-latrine distances were almost double in continuous forest than in fragments. Latrines were located beneath only a few tree species, and these trees were larger in diameter in continuous than fragmented forests. Because latrines may represent hotspots of seedling recruitment, our results have important ecological and conservation implications. The variation in the spatial distribution of sleeping sites across the forest indicates that spider monkeys likely create a complex seed deposition pattern in space and time. However, the use of a very few tree species for sleeping could contribute to the establishment of specific vegetation associations typical of the southeastern Mexican rainforest, such as Terminalia-Dialium, and Brosimum-Dialium.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

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.     

Moms are better nurses than dads: gender biased self‐facilitation in a dioecious Juniperus tree

Questions: Can gender of nurse plants affect regeneration patterns and spatial population structure? Is there a seed‐seedling conflict in the regeneration process? What factors are responsible for the clumped spatial population structure observed for adult trees? Location: Mediterranean cold semi‐arid high mountains in Spain. Methods: The spatial pattern of adult Juniperus thurifera trees was studied by means of Ripley's K‐analysis. χ² analyses were used to test for natural seedling frequency in each of three main microhabitats: (1) under female and (2) male tree canopies and (3) in open interspaces. The observed pattern was explained experimentally by studying seed and seedling survival for two years. Survival probabilities were calculated across life stages for each of three main microhabitats. Results: Adult J. thurifera trees were aggregated in space. Most seedlings were found underneath female J. thurifera trees. Experimental studies demonstrated that from seed dispersal to seedling survival all life stages showed the same positive or negative trend within a given microhabitat, indicating stage coupling and no seed‐seedling conflicts. Attraction of frugivo‐rous birds by reproductive female junipers and improvement of environmental conditions beneath tree canopies were the main factors responsible for the variation in seedling density among microhabitats; highest underneath female trees and lowest in open interspaces. Conclusions: In dioecious species, the gender of nurse plants can significantly determine the spatial population structure. In J. thurifera forests, facilitation beneath female trees occurs among all life stages without any sign of seed‐seedling conflict. The most critical factors shaping the spatial population structure were directed seed dispersal and environmental amelioration beneath female conspecific trees.     

The relative contributions of seed and pollen dispersal to gene flow and genetic diversity in seedlings of a tropical palm

Seed and pollen dispersal shape patterns of gene flow and genetic diversity in plants. Pollen is generally thought to travel longer distances than seeds, but seeds determine the ultimate location of gametes. Resolving how interactions between these two dispersal processes shape microevolutionary processes is a long‐standing research priority. We unambiguously isolated the separate and combined contributions of these two dispersal processes in seedlings of the animal‐dispersed palm Oenocarpus bataua to address two questions. First, what is the spatial extent of pollen versus seed movement in a system characterized by long‐distance seed dispersal? Second, how does seed dispersal mediate seedling genetic diversity? Despite evidence of frequent long‐distance seed dispersal, we found that pollen moves much further than seeds. Nonetheless, seed dispersal ultimately mediates genetic diversity and fine‐scale spatial genetic structure. Compared to undispersed seedlings, seedlings dispersed by vertebrates were characterized by higher female gametic and diploid seedling diversity and weaker fine‐scale spatial genetic structure for female gametes, male gametes and diploid seedlings. Interestingly, the diversity of maternal seed sources at seed deposition sites (N _em) was associated with higher effective number of pollen sources (N _ep), higher effective number of parents (N _e) and weaker spatial genetic structure, whereas seed dispersal distance had little impact on these or other parameters we measured. These findings highlight the importance maternal seed source diversity (N _em) at frugivore seed deposition sites in driving emergent patterns of fine‐scale genetic diversity and structure.     

Spatial Patterns of Seed Dispersal by White-Faced Capuchins in Costa Rica: Evaluating Distant-Dependent Seed Mortality

Spatial patterns of seed dispersal are the focus of numerous theoretical examinations of endozoochory. Here, we examine the spatial pattern of seed dispersal by white-faced capuchin monkeys Cebus capucinus in Santa Rosa National Park, Costa Rica, and the neighborhood characteristics and distance variables most closely associated with seed survival and germination, and seedling survival and growth in various locations. Overall, distance to the nearest fruiting conspecific tree has the most positive, consistent effect on growth and survival variables, which supports a variation of the Janzen–Connell seed escape hypothesis.     

Seed Dispersal of a High Quality Fruit by Specialized Frugivores: High Quality Dispersal?1

Dispersal quality, as estimated by the cumulative effects of dispersal, germination, seed predation, and seedling survival, was examined for Beilschmiedia pendula (Lauraceae) in Monteverde, Costa Rica. I determined the pattern of dispersal by finding seeds deposited by birds, protected the seeds from seed predators with cages to assess germination and seedling survival, and examined seed predation rates with marked seeds. Seed predation, germination, and seedling survival were compared between seeds naturally dispersed by birds and seeds placed at randomly located sites. Approximately 70 percent of seeds dispersed by birds (N= 244) were deposited <10 m from crown edges of fruiting B. pendula trees, although some seeds were dispersed at least 70 m away. Larger seeds were more likely to be dispersed under or close to the parent trees, and larger seeds produced larger seedlings. Seed size was not correlated directly with seedling survival, but larger seedlings at three months were most likely to survive one year. Seed predation by mammals and insects and seedling mortality due to fungal pathogens were concentrated beneath the crowns of parent trees. Seedlings and saplings were more abundant beneath fruiting B. pendula trees, but individuals farther away were taller on average. Thus, dispersal is beneficial for B. pendula, but such benefits appear most pronounced at a small spatial scale; seeds dispersed >30 m from the crown edges actually had a lower probability of survival than those dispersed 10–20 m. Only 10 percent of B. pendula. seeds received high‐quality dispersal in terms of landing in the zone with the highest per seed probability of seedling survival 10–20 m from parental crowns.     

Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

R.M. Cowling,D.M. Richardson & S.M. Pierce (eds.) 1977. Vegetation of Southern Africa.

We determined the role of bird dispersal in seed and seedling dynamics of the tree Kalopanax pictus from 7 years of observing seed rain and seedling emergence in a broad-leaved deciduous forest in central Japan. We also performed an experiment on the influence of seed pulp on germination of seeds of K. pictus. Seeds of this species can lie dormant for several years, and this causes rather constant yearly seedling emergence in spite of irregular seed production. The spatial distribution of the seedlings that emerged each year (maximum distance from nearest conspecific seed-bearing tree of 90 m) was wider than that of gravity-dispersed seeds (max. distance of 37 m), suggesting seed dispersal by birds in winter. Emerged seedling densities at sites over 20 m from the nearest conspecific seed-bearing tree were highest in the spring of 1991, about half a year after the largest seed fall of the observation period. However, emerged seedling densities within 20 m from seed-bearing trees were highest in 1992, 1.5 years after the largest seed fall. These field observations may be explained by the experimental results on the effects of seed pulp on germination. Intact seeds germinate slowly at low germination rates, while seeds without seed pulp germinate quickly at high germination rates. Fallen seeds with seed pulp thus appear to form a seed bank near seed sources (temporal dispersal), while seeds scattered by birds appear to increase the possibility of reaching the present safe sites in distant areas with quick germination (spatial dispersal).     

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.     

Low plant density enhances gene dispersal in the Amazonian understory herb Heliconia acuminata

In theory, conservation genetics predicts that forest fragmentation will reduce gene dispersal, but in practice, genetic and ecological processes are also dependent on other population characteristics. We used Bayesian genetic analyses to characterize parentage and propagule dispersal in Heliconia acuminata L. C. Richard (Heliconiaceae), a common Amazonian understory plant that is pollinated and dispersed by birds. We studied these processes in two continuous forest sites and three 1‐ha fragments in Brazil's Biological Dynamics of Forest Fragments Project. These sites showed variation in the density of H. acuminata. Ten microsatellite markers were used to genotype flowering adults and seedling recruits and to quantify realized pollen and seed dispersal distances, immigration of propagules from outside populations, and reproductive dominance among parents. We tested whether gene dispersal is more dependent on fragmentation or density of reproductive plants. Low plant densities were associated with elevated immigration rates and greater propagule dispersal distances. Reproductive dominance among inside‐plot parents was higher for low‐density than for high‐density populations. Elevated local flower and fruit availability is probably leading to spatially more proximal bird foraging and propagule dispersal in areas with high density of reproductive plants. Nevertheless, genetic diversity, inbreeding coefficients and fine‐scale spatial genetic structure were similar across populations, despite differences in gene dispersal. This result may indicate that the opposing processes of longer dispersal events in low‐density populations vs. higher diversity of contributing parents in high‐density populations balance the resulting genetic outcomes and prevent genetic erosion in small populations and fragments.     

Seed Dispersal by Avian Frugivores: Non‐random Heterogeneity at Fine Scales

Seed dispersal studies have primarily examined dispersal as a function of distance from the parent tree and/or heterogeneity in dispersal due to animal use of nesting, roosting and sleeping sites. However, non‐random heterogeneity in seed dispersal is also likely to result from the post‐foraging behavior and movement of frugivores which prefer certain trees. To characterize variation in seed rain at fine scales, we studied the dispersal curve of Prunus ceylanica, a primarily bird‐dispersed species. We compared seed rain at conspecifics, heterospecific fruiting trees with similar frugivore assemblages, emergent trees, and the landscape surrounding these trees. Seed rain of P. ceylanica was found to peak globally under the canopy of conspecifics but to peak locally under the canopy and immediate neighborhood of heterospecific fruiting trees. Our results demonstrate that seed rain is highly clumped even at fine spatial scales. A large proportion of seeds are dispersed in specific, localized regions. This variation can have important implications for plant population dynamics and might significantly alter the impact of post‐dispersal processes. Seed dispersal models may need to incorporate this heterogeneity to explain manifestations of spatially explicit dynamics like mixed species ‘orchards’.     

Spit it out: Monkeys disperse the unorthodox and toxic seeds of Clivia miniata (Amaryllidaceae)

Seeds of many Amaryllidaceae are unorthodox (recalcitrant) and toxic, and cannot survive ingestion, yet are packaged in brightly colored fruits suggestive of zoochory. Seed dispersal and germination of the understory amaryllid, Clivia miniata, were investigated in KwaZulu‐Natal, South Africa. Motion‐activated cameras revealed that samango monkeys (Cercopithecus mitis labiatus) are the primary disperser of C. miniata seeds. They eat the mesocarp and, to a lesser extent, the exocarp, and spit the large (13 mm diameter) seeds whole and cleaned onto the forest floor. Most seeds were dispersed farther than 1 m from the parent. Experimental removal of the fruit pulp had a small positive effect on the rate of seed germination, but did not affect subsequent seedling growth rates. The main advantages of monkey dispersal of Clivia seeds appear to be short‐distance dispersal away from the dense foliage of clumped parent plants and occasional long‐distance dispersal through cheek‐pouching behavior.     

Monsoon rain forest seedling dynamics, northern Australia: contrasts with regeneration in eucalypt-dominated savannas

Aim To explore: (1) the relative influences of site conditions, especially moisture relations, on pathways and rates of monsoon rain forest seedling and sapling regeneration, especially of canopy dominants, in northern Australia; and (2) contrasts between regeneration syndromes of dominant woody taxa in savannas and monsoon rain forest. Location Four monsoon rain forest sites, representative of regional major habitat and vegetation types, in Kakadu National Park, northern Australia. Methods A decadal study involved: (1) initial assessment over 2.5 years to explore within‐year variability in seed rain, dormant seed banks and seedling (< 50 cm height) dynamics; and (2) thereafter, monitoring of seedling and sapling (50 cm height to 5 cm d.b.h.) dynamics undertaken annually in the late dry season. On the basis of observations from this and other studies, regeneration syndromes of dominant monsoon rain forest taxa are contrasted with comparable information for dominant woody savanna taxa, Eucalyptus and Corymbia especially. Results Key observations from the monsoon rain forest regeneration dynamics study component are that: (1) peak seed rain inputs of rain forest taxa were observed in the wet season at perennially moist sites, whereas inputs at seasonally dry sites extended into, or peaked in, the dry season; (2) dormant soil seed banks of woody rain forest taxa were dominated by pioneer taxa, especially figs; (3) longevity of dormant seed banks of woody monsoon rain forest taxa, including figs, was expended within 3 years; (4) seedling recruitment of monsoon rain forest woody taxa was derived mostly from wet season seed rain with limited inputs from soil seed banks; (5) at all sites rain forest seedling mortality occurred mostly in the dry season; (6) rain forest seedling and sapling densities were consistently greater at moist sites; (7) recruitment from clonal reproduction was negligible, even following unplanned low intensity fires. Main conclusions By comparison with dominant savanna eucalypts, dominant monsoon rain forest taxa recruit substantially greater stocks of seedlings, but exhibit slower aerial growth and development of resprouting capacity in early years, lack lignotubers in mesic species, and lack capacity for clonal reproduction. The reliance on sexual as opposed to vegetative reproduction places monsoon rain forest taxa at significant disadvantage, especially slower growing species on seasonally dry sites, given annual–biennial fires in many north Australian savannas.     

Implications of Behavior and Gut Passage for Seed Dispersal Quality: The Case of Black and Gold Howler Monkeys

The most generalized mechanism of seed dispersal in tropical and subtropical forests is dispersal by vertebrates. The pattern of seed dispersal and germination capacity of dispersed seeds can determine the seed disperser quality because both affect the postdispersal fate of dispersed seeds. Animals' behavior is an important determinant of seed deposition pattern. One of most abundant frugivores of Neotropical forests are howler monkeys Alouatta spp., so my aim was to evaluate the potential quality of Alouatta caraya as a seed disperser, obtaining details of: (1) seed deposition pattern, (2) behavioral context of defecations, and (3) germination capacity of dispersed seeds. During 2 years the seed shadow created by howlers and the behavioral context of depositions was examined in the Paraná flooded forest, and germination tests were conducted in the laboratory. Black and gold howler monkeys consumed fruits of five canopy trees, three understory trees, one shrub, and two vines. Howlers dispersed seeds in a complex pattern: 35 percent (337 scats) of scats were deposited individually, and 65 percent were deposited in big latrines associated with the main sleeping trees and in small latrines associated with secondary sleeping trees and confrontations at territory edges. From 261 dispersal distances recorded, 20 and 40 m were the mode, but 57 percent of distances were >50 m and 31 percent were >100 m. Germination speed increased significantly in ingested seeds of Banara arguta, Ocotea diospyrifolia and Eugenia punicifolia . The seed deposition pattern generated by A. caraya may produce a diversification of environmental conditions for dispersed seeds that should be considered in future evaluations.     

Spatiotemporal patterns of seed dispersal in a wind-dispersed Mediterranean tree (Acer opalus subsp. granatense): implications for regeneration

Seed dispersal can severely limit the quantity of plant recruits and their spatial distribution. However, our understanding of the role of dispersal in regeneration dynamics is limited by the lack of knowledge of seed deposition patterns in space and time. In this paper, we analyse the spatiotemporal variability of seed dispersal patterns in the Mediterranean maple, Acer opalus subsp. granatense, by monitoring seed rain along two years at a broad spatial scale (2 mountain ranges, 2 populations per range, 4 microhabitats per population). We quantified seed limitation and its components (source and dispersal limitation), and explored dispersal limitation in space by analysing dispersal distances, seed aggregation, and microhabitat seed distribution. Acer opalus subsp. granatense was strongly seed‐limited throughout the gradients explored, being always dispersal limitation much higher than source limitation. The distribution of seeds with distance from adult individuals was leptokurtic and right‐skewed in all populations, being both kurtosis and skewness higher the year of the highest seed production. Dispersal distances were shorter than expected by random in the four populations, which suggests distance‐limited dispersal. Dispersal patterns were highly aggregated and showed a preferential direction around adults. At the microhabitat scale, most seeds accumulated under adult maples. However, there were no more seeds under trees and shrubs other than maple than in open interspaces, implying that established vegetation does not disrupt patterns of seed deposition by physically trapping seeds. When compared with patterns of seedling establishment, limited dispersal ability and inter‐annual spatial concordance in seed rain patterns suggest that several potentially safe sites for recruitment have a very low probability of receiving seeds in most maple populations. These findings are especially relevant for rare species such as Acer opalus subsp. granatense, and illustrate how dispersal studies are not only crucial for our understanding of plant population dynamics but also to provide conservation directions.     

Toucans (Ramphastos ambiguus) facilitate resilience against seed dispersal limitation to a large‐seeded tree (Virola surinamensis) in a human‐modified landscape

Large‐seeded plants may suffer seed dispersal limitation in human‐modified landscapes if seed dispersers are absent or unable to disperse their seeds. We investigated dispersal limitation for the large‐seeded tree Virola surinamensis in a human‐modified landscape in southern Costa Rica. During two fruiting seasons, we monitored crop size, seed removal rates, the number of fruiting conspecifics within 100 m, and feeding visitation rates by frugivores at trees located in high and low forest disturbance conditions. Seed removal rates and the total number of seeds removed were high regardless of the disturbance level, but these parameters increased with tree crop size and decreased with the number of fruiting V. surinamensis trees within a 100 m radius. Trees at low disturbance levels were more likely to be visited by seed dispersers. Black mandibled toucans (Ramphastos ambiguus) and spider monkeys (Ateles geoffroyi) were the most important seed dispersers, based on visitation patterns and seed removal rates. Spider monkey feeding visits were more frequent at high disturbance levels, but the monkeys preferentially visited isolated trees with large yields and surrounded by a low number of fruiting Virola trees within 100 m. Toucan visitation patterns were not constrained by any of the predictors and they visited trees equally across the landscape. We suggest that isolated and highly fecund Virola trees are an important food resource for spider monkeys in human‐modified landscapes and that toucans can provide resilience against seed dispersal limitations for large‐seeded plants in human‐modified landscapes in the absence of hunting.     

Seed dispersal by lizards on a continental‐shelf island: predicting interspecific variation in seed rain based on plant distribution and lizard movement patterns

Aim We estimated the patterns of seed deposition provided by the eyed lizard, Timon lepidus, and evaluated whether these patterns can be generalized across plant species with different traits (fruit and seed size) and spatial distributions. Location Monteagudo Island, Atlantic Islands National Park (north‐western Spain). Methods We radio‐tracked seven lizards for 14 days and estimated their home ranges using fixed kernels. We also geo‐referenced all fruit‐bearing individuals of four plant species dispersed by eyed lizards in the study area (Corema album, Osyris alba, Rubus ulmifolius and Tamus communis), measured the passage time of their seeds through the lizard gut, and estimated seed predation in four habitats (bare sand, grassland, shrub and gorse). Seed dispersal kernels were estimated using a combination of these data and were combined with seed predation probability maps to incorporate post‐dispersal seed fate (‘seed survival kernels’). Results Median seed gut‐passage times were around 52–98 h, with maximum values up to 250 h. Lizards achieved maximum displacement in their home ranges within 24–48 h. Seed predation was high (80–100% of seeds in 2 months), particularly under Corema shrub and gorse. Seed dispersal kernels showed a common pattern, with two areas of preferential seed deposition, but the importance of these varied among plant species. Interspecific differences among dispersal kernels were strongly reduced by post‐dispersal seed predation; hence, seed survival kernels of the different plant species showed high auto‐ and pairwise‐correlations at small distances (< 50 m). As a result, survival to post‐dispersal seed predation increased with dispersal distance for O. alba and T. communis, but not for C. album. Main conclusions Seed dispersal by lizards was determined primarily by the interaction between the dispersers’ home ranges and the position of the fruit‐bearing plants. As a result, seed rain shared a common template, but showed considerable variation among species, determined by their specific spatial context. Seed predation increased the spatial coherence of the seed rain of the different species, but also resulted in contrasting relationships between seed survival and dispersal distance, which may be of importance for the demographic and evolutionary processes of the plants.     

Do spatial effects play a role in the spatial distribution of desert‐dwelling Acacia raddiana?

Abstract. We investigated the spatial pattern of A. raddiana in the Negev desert of Israel in order to gain insights into the factors and processes driving the dynamics of this species. Using a scale‐dependent measure, the ring statistic, we analysed both patterns observed in the field and time series of spatial tree distributions produced by a simulation model. In the field, random spacing was the predominant pattern observed. However seedlings were clumped on small scales. We ran the model under two contrasting scenarios representing hypotheses that explain the clumping of seedlings and the random distribution of trees. One hypothesis is that there is spatial heterogeneity in seed distribution, germination and seedling mortality, but that these heterogeneities are not correlated with each other in space. The second hypothesis assumes a correlation between these heterogeneities leading to areas suitable for establishment. However, the suitability of the sites is temporally variable. Furthermore, the second hypothesis assumes density‐dependent tree mortality due to competition. Both hypotheses lead to spatial distributions that are in qualitative agreement with the patterns observed in the field. Therefore, the classical view that a clumped seedling distribution and a random pattern of older trees is due to clumped regeneration and density‐dependent mortality may not hold for Acacia trees in the Negev.