Long-distance dispersal helps germinating mahogany seedlings escape defoliation by a specialist caterpillar

Herbivores and pathogens with acute host specificity may promote high tree diversity in tropical forests by causing distance- and density-dependent mortality of seedlings, but evidence is scarce. Although Lepidoptera larvae are the most abundant and host-specific guild of herbivores in these forests, their impact upon seedling distributions remains largely unknown. A firm test of the mechanism underpinning the Janzen–Connell hypothesis is difficult, even for a single tree species, because it requires more than just manipulating seeds and seedlings and recording their fates. Experimental tests require: (1) an insect herbivore that is identified and highly specialised, (2) linkage to an in situ measure (or prevention) of herbivory, and (3) evaluation and confirmation among many conspecific adult trees across years. Here we present experimental evidence for a spatially explicit interaction between newly germinating seedlings of a Neotropical emergent tree, big-leaf mahogany (Swietenia macrophylla, Meliaceae), and caterpillars of a noctuid moth (Steniscadia poliophaea). In the understory of a southeastern Amazon forest, the proportion of attacks, leaf area lost, and seedling mortality due to this specialised herbivore peaked near Swietenia trees, but declined significantly with increasing distance from mature fruiting trees, as predicted by the Janzen–Connell hypothesis. We conclude that long-distance dispersal events (>50 m) provided an early survival advantage for Swietenia seedlings, and propose that the role of larval Lepidoptera as Janzen–Connell vectors may be underappreciated in tropical forests.     

Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities

Animals aid population growth and fitness in tropical forest communities through dispersal and negatively impact populations through seed predation. The interaction between dispersal and seed predation can produce distance- or density-dependence; powerful mechanisms for maintaining species diversity incorporated in the Janzen–Connell model. Large mammals, the highest biomass seed predators of intact Amazonian communities and at risk due to human disturbance, are potentially central to these interactions. This study tests the Janzen–Connell model and investigates the impact of mammalian seed predators on seedling recruitment and maintenance of tree diversity. Patterns of both vertebrate and invertebrate seed predation and seedling recruitment were studied in the two most abundant canopy tree species in western Amazonia (Arecaceae: Astrocaryum murumuru and Iriartea deltoidea). We specifically examined effects of both spatial and temporal variation of the highest biomass seed predator in southwest Amazonian forests, the white-lipped peccary (Tayassu pecari), on recruitment through disturbed and undisturbed sites and through a fortuitous 12 year natural extinction and recolonization event of T. pecari. Distance-dependent seedling recruitment was found in Astrocaryum and Iriartea at both sites. However, the median distance of seedlings was ~1.5× farther from reproductive adults in both palms at the undisturbed site. The number of Iriartea seeds escaping predation increased 6,000% in both space and time due to the decline of T. pecari abundance. The results demonstrate that Janzen–Connell effects are stronger in intact ecosystems and tie these mechanistically to changes in seed predator abundance. This study shows that anthropogenic changes in mammal communities decrease the magnitude of Janzen–Connell effects in Amazonian forests and may result in decreases in tree diversity.     

Herbivores limit the population size of big‐leaf mahogany trees in an Amazonian forest

The Janzen–Connell hypothesis proposes that specialized herbivores maintain high numbers of tree species in tropical forests by restricting adult recruitment so that host populations remain at low densities. We tested this prediction for the large timber tree species, Swietenia macrophylla, whose seeds and seedlings are preyed upon by small mammals and a host‐specific moth caterpillar Steniscadia poliophaea, respectively. At a primary forest site, experimental seed additions to gaps – canopy‐disturbed areas that enhance seedling growth into saplings – over three years revealed lower survival and seedling recruitment closer to conspecific trees and in higher basal area neighborhoods, as well as reduced subsequent seedling survival and height growth. When we included these Janzen–Connell effects in a spatially explicit individual‐based population model, the caterpillar's impact was critical to limiting Swietenia's adult tree density, with a > 10‐fold reduction estimated at 300 years. Our research demonstrates the crucial but oft‐ignored linkage between Janzen–Connell effects on offspring and population‐level consequences for a long‐lived, potentially dominant tree species.     

Plant density can increase invertebrate postdispersal seed predation in an experimental grassland community

Janzen–Connell effects are negative effects on the survival of a plant's progeny at high conspecific densities or close to its conspecifics. Although the role of Janzen–Connell effects on the maintenance of plant diversity was frequently studied, only few studies targeted Janzen–Connell effects via postdispersal seed predation in temperate grassland systems. We examined effects of conspecific density (abundance of conspecific adult plants) on postdispersal seed predation by invertebrates of three grassland species (Centaurea jacea, Geranium pratense, and Knautia arvensis) in experimental plant communities. Additionally, we examined the impact of plant species richness and different seed predator communities on total and relative seed predation (= seed predation of one plant species relative to others). We offered seeds in an exclusion experiment, where treatments allowed access for (1) arthropods and slugs, (2) arthropods only, (3) small arthropods only, and (4) slugs only. Treatments were placed in plots covering a gradient of abundance of conspecific adults at different levels of plant species richness (1, 2, 3, 4, 8 species). Two of the plant species (C. jacea and K. arvensis) experienced higher rates of seed predation and relative predation with increasing abundance of conspecific adults. For C. jacea, this effect was mitigated with increasing plant species richness. Differences in seed predator communities shifted seed predation between the plant species and changed the magnitude of seed predation of one plant species relative to the others. We exemplify density‐dependent increase in seed predation via invertebrates in grassland communities shaping both the total magnitude of species‐specific seed predation and seed predation of one species relative to others. Further differences in seed predator groups shift the magnitude of seed predation between different plant species. This highlights the importance of invertebrate seed predation to structure grasslands via density‐dependent effects and differing preferences of consumer groups.     

Seed shadow, seedling recruitment, and spatial distribution of Buchenavia capitata (Combretaceae) in a fragment of the Brazilian Atlantic forest.

Here we describe the seed shadow, seedling recruitment, ontogenetic structure and spatial distribution of Buchenavia capitata (an emergent canopy tree) in a 380-ha fragment of the Atlantic forest in northeast Brazil. In particular, we examine seed distribution around 10 parental trees and both seedling recruitment and mortality, during an 18 month period beneath and around parental trees. Moreover, we describe: (1) B. capitata occurrence within treefall gaps; (2) population structure in terms of ontogenetic stages for the whole site; and (3) spatial distribution of adults within an area of 51 hectares. 99% of seeds were found beneath parent crowns (n = 4,236) and seed density reached 14.6 +/- 29.9 seeds/m2 (0-140 seeds/m2). 49% of all seeds germinated but seedling mortality reached 100% after an 18 month period. In addition, saplings of B. capitata were not found in forest understory and within 30 treefall gaps (94-2,350 m2). The adults showed an average DBH of 69.3 +/- 22.1 cm, were 19.2 +/- 2.9 m tall and presented a clumped spatial distribution. B. capitata matched some of the features presented by shade intolerant trees or large-gap specialists, and we hypothesize that low rates or even lack of long distance seed dispersal events may be reducing the probability of B. capitata seeds reaching suitable habitats for successful seedling recruitment and growth. Because of that (1) seedlings face high levels of early mortality; (2) there is no sapling recruitment at the study site; and (3) local population faces senility and it is threatened by local extinction.     

Distance- and density-dependent seedling mortality caused by several diseases in eight tree species co-occurring in a temperate forest

To examine whether the Janzen–Connell mechanism applies to temperate forests, seedling survival and causes of mortality were investigated at two distances (beneath, far) from conspecific adults and at two densities (high, low) at each distance for seedlings (n = 7935) of eight tree species co-occurring in a hardwood forest. Six of the eight species showed distance- and/or density-dependent seedling mortality mainly caused by diseases and rodents. In four of the five species primarily killed by disease (i.e. damping-off, blight, rot, powdery mildew), the infectivity (probability of infection by the disease) and/or the virulence (proportion of seedlings killed to those infected by the disease) were higher beneath than far from conspecific adults. These findings suggest that host specificity and/or spatially heterogeneous activity of natural enemies play an important role in the reciprocal replacement of tree species, maintaining species diversity in temperate forests.     

Predation by vertebrates and invertebrates on the seeds of five canopy tree species of an Amazonian forest

We studied the pre-germination loss of seeds to invertebrate and vertebrate seed predators of 5 species of Amazonian trees (Astrocaryum macrocalyx—Palmae; Bertholletia excelsa—Lecithydaceae; Calatola venezuelana—Icacinaceae; Dipteryx micrantha—Leguminosae (Papilionoidae); Hymenaea courbaril-Leguminosae (Caesalpinoidae)). These five species were selected from a large tree flora on several criteria. All possess large (3–10 cm) well-protected seeds that might plausibly be attractive to mammalian seed predators. The reproductive biology of three of the species, or close congeners, had been studied elsewhere in the Neotropics (Astrocaryum, Dipteryx, Hymenaea); one is important to the economy of southeastern Peru (Bertholletia); and one, despite large and apparently edible seeds, appeared to suffer no pre-germination loss to predators (Calatola). We conducted the research in mature forests in the Manu National Park of southeastern Peru where mammal densities are unperturbed by human activities. Densities of adult trees of the five species in our area range from very high (>30 per ha: Astrocaryum) to very low (1 per ha: Hymenaea).Loss of seeds to all causes, and to mammalian seed predators in particular, was determined for seeds placed in 2-square meter mammal exclosures and in open controls located at 10 m (near) and 50 m (far) from a large mature individual of the target species (with minor variations in the design for Astrocaryum and Calatola). The exclosures were of two types: impermeable—designed to exclude all mammals, but not invertebrate seed predators, and semipermeable—designed to admit small (<500 g), but not large mammals. Experimental and control plots were stocked with apparently viable seeds during the dry-wet transition period (October–November) and scored one year later.A significant distance effect (higher predation near vs far from a large conspecific adult) was found in only one of the species (Astrocaryum), the only one to be attacked with high frequency by invertebrate seed predators. The absence of any detectable distance effect attributable to mammals suggests that mammals, over the course of a year, thoroughly search the forest floor for seeds. Invertebrates may thus be responsible for most pre-germination distance (density) effects. With respect to the treatments, we found three qualitatively distinct results: seeds of three species (Astrocaryum, Bertholletia, Dipteryx) were significantly protected by the impermeable, but not semipermeable exclosures, implicating small mammals in seed loss; the seeds of one species (Hymenaea) were significantly protected by exclosures of both types, implicating large mammals; and the seeds of one species (Calatola) exhibited 100% survival, whether or not protected by exclosures.The importance of large mammals as seed predators is generally underestimated in these experiments because semipermeable exclosures may serve as foraging reserves for small mammals. Finally, we noted no relationship between the intensity of mammalian seed predation (as suggested by the survival of unprotected seeds) and the abundance of adults of the five species in the environment. The diversity of results obtained for the five species reveals that large-seeded tropical trees may display a wide range of demographic patterns, and points to the likely importance of post-germination bottlenecks in the population biology of many species, even those that may experience severe pre-germination seed loss.     

Central-place seed foraging and vegetation patterns

We investigate how central-place seed foragers with a nest in the proximity of one or more seed sources determine the formation of different vegetation patterns. In particular, we discuss the ecological conditions that lead to the formation of hump-shaped (Janzen–Connell) patterns in a two-dimensional landscape. Our analysis shows that central-place predation can generate Janzen–Connell patterns even if predators’ movement strategies are exclusively based on resource abundance, both in the single-plant/single-nest case and in a forest with several seed sources. We also show that social foraging may either promote or work against the formation of Janzen–Connell patterns, depending upon the way foragers take advantage of social interactions.     

Factors affecting post-dispersal seed survival in a tropical forest

Summary Using the subcanopy tree Faramea occidentalis in Panama, I studied post-dispersal seed survival as a function of five characteristics describing seed locations. By simultaneously considering distance from a conspecific adult, size of the nearest conspecific adult, leaf litter quantity, proximity to logs or tree trunks, and whether or not the seed was in a gap, I was able to analyze the influences of individual factors, as well as the interactions among factors. Seed survival was significantly less in treefall gaps than in the forest understory. Seed survival was also influenced by the size of the nearest adult but in a complex interaction with distance to an adult. For seeds beneath adults, survival decreased with increasing tree size, while for seeds away from adults, survival was independent of the size of the nearest conspecific adult. Distance did not directly affect seed survival, nor did the quantity of leaf litter or the proximity to a tree trunk or a log. In a separate analysis, the relationship between distance and seed survival was consistent over four years, suggesting that single cohort studies may provide accurate insights into the consequences of dispersal. In contrast, the spatial locations of surviving seeds were not consistent over the four-year period. Transects with high survival one year did not tend to have high survival in other years, and the locations of surviving seeds in any particular year could not be predicted from the knowledge of where seeds survived in other years. While survival is patchy within a year, the locations of patches shift from year to year.     

Herbivore-induced expansion of generalist species as a driver of homogenization in post-disturbance plant communities

We used a controlled experiment to investigate how disturbance scale (canopy gap area) and herbivory influence post-disturbance plant community dynamics. Twenty canopy gaps were installed in a temperate hemlock-hardwood forest during the winter of 2002–2003: seven small gaps (50–150 m²), seven medium gaps (151–250 m²), and six large gaps (251–450 m²). Within each gap, we established 4–12 sample plots (depending on gap size); 1–3 of which were enclosed with wire mesh white-tailed deer (Odocoileus virginianus) exclosures in 2005. Gaps were revisited and intensively sampled in 2007. After five growing seasons, ground-layer plant communities in non-exclosed plots were more similar compositionally than exclosure plots. Non-exclosed plots in small and medium gaps were more similar to non-exclosed plots in large gaps than they were to exclosed plots in their respective gap size class. Shade-tolerant forbs and trees were less common outside exclosures, while generalist species associated with higher understory light levels and exotics were more prevalent outside exclosures. Our results suggest that even in forests with relatively low deer densities (6.5–9.3 deer km⁻²), white-tailed deer herbivory may influence the developmental trajectory of post-disturbance plant communities and be a mechanism for decreasing β-diversity along environmental gradients.     

格氏栲天然林林窗和林下种子散布及幼苗更新研究

以格氏栲(Castanopsis kawakamii)天然林为研究对象,探讨了林窗和林下格氏栲种子雨、种子库的分布特征及幼苗更新状况。结果表明:林窗种子雨总量和完好种子密度高于林下,未成熟种子比例低于林下;林窗和林下种子雨高峰期掉落数量分别占种子雨总量的77.13%和74.5%;林窗种子库储量低于林下,种子库中以全食或捡拾种子比例最高,其中种子库储量垂直分布表现为枯落物层(约占2/3)>腐殖质层(0～5 cm)(约占1/3)>心土层(5～10 cm)(小于1%),以格氏栲种子占绝对优势;格氏栲从种子到幼苗的转化率低,林窗中格氏栲完好种子密度与幼苗密度均高于林下。林窗微生境提高了种子散布过程中格氏栲成熟种子的密度和比例,有利于促进格氏栲的幼苗更新,表明林窗在格氏栲种群恢复过程中扮演着重要角色。     

Seed dispersal, plant recruitment and spatial distribution of Bactris acanthocarpa Martius (Arecaceae) in a remnant of Atlantic forest in northeast Brazil

Seed dispersal ecology of Bactris acanthocarpa Mart. (Arecaceae), an Atlantic forest understory palm, was investigated during two years as an attempt to test the following predictions: (i) seeds of Bactris are dispersed by mammals and large-gaped birds; (ii) Bactris benefits from seed dispersal in terms of seed predation avoidance, improvement of seed germination and seedling survival; and (iii) spatial distribution of adults is related to patterns of seed dispersal. The study was conducted at Dois Irmãos Reserve, a 387.4-ha reserve of Atlantic forest in northeastern Brazil (8º S–35º W). Black–rumped agoutis (Dasyprocta prymnolopha) and Guianan squirrels (Sciurus aestuans) were identified as the seed dispersers/predators, moving seeds short distances (< 4 m from parents) and at low rates (0.04-0.05 diaspore/palm/day). Pyrene burial prevented seed predation by vertebrates and reduced by half seed infestation by Scolytidae beetles. Only buried pyrenes germinated. Pyrene predation was not correlated with distance from conspecific adults. In contrast, early seedling mortality was higher near conspecific adults. Most adults (64%) had their nearest conspecific adult neighbour > 4 m away in contrast to 96% of seedlings that occurred concentrated within 4 m from adults (77% under the palm crowns). Here, we present evidence that spatial distribution of B. acanthocarpa is partly due to low rates of seed removal, short-distance seed dispersal by agoutis and squirrels, and early seedling mortality associated with presence of seedlings under palm crowns.     

Indirect interactions among tropical tree species through shared rodent seed predators: a novel mechanism of tree species coexistence

The coexistence of numerous tree species in tropical forests is commonly explained by negative dependence of recruitment on the conspecific seed and tree density due to specialist natural enemies that attack seeds and seedlings (‘Janzen–Connell’ effects). Less known is whether guilds of shared seed predators can induce a negative dependence of recruitment on the density of different species of the same plant functional group. We studied 54 plots in tropical forest on Barro Colorado Island, Panama, with contrasting mature tree densities of three coexisting large seeded tree species with shared seed predators. Levels of seed predation were far better explained by incorporating seed densities of all three focal species than by conspecific seed density alone. Both positive and negative density dependencies were observed for different species combinations. Thus, indirect interactions via shared seed predators can either promote or reduce the coexistence of different plant functional groups in tropical forest.     

Tree distribution pattern and fate of juveniles in a lowland tropical rain forest – implications for regeneration and maintenance of species diversity

Three analyses of species diversity in a lowland dipterocarp forest were conducted to examine whether the nature of forest community dynamics are determined by density-dependent recruitment and mortality of saplings with a data set obtained in a 50 ha plot in Pasoh Forest Reserve. The first analysis examined whether sapling density varied as a function of distance from the nearest conspecific adult. The second analysis assessed the relationship between the spatial distribution patterns of saplings and adult trees. A third analysis examined sapling recruitment and mortality based on data from 2 censuses, taken in 1985 and 1990. Four hundred forty-four species (each with more than 100 individuals) out of the total of 814 species recorded in the plot, were chosen for the analyses. Of these selected species, 56 species showed significant reduction in sapling densities close to the conspecific adults. Within this group, 11 species were in the emergent layer (29.0% of the total species in this layer), 17 were in the canopy layer (10.5%), 18 were in the understory layer (11.3%), and 10 were in treelet and shrub layer (11.8%). In contrast, the sapling densities of 53 species decreased with increasing distance from conspecific adults; 2 of these species were in the emergent layer (5.2% of the total species in this layer), 14 were in the canopy layer (8.6%), 21 were in the understory layer (13.2%), and 16 were in the treelet and shrub layer (18.8%). The saplings of 35 of the 444 total selected species were clumped, while adults were regularly or randomly distributed. Of the remaining species, in 183 species (41.2%), the distributions of both adults and saplings were clumped. Thus, these 2 analyses do not support the prediction that most of the species of lowland tropical forests fail to produce new adults in their vicinity and as a result of this, adult trees are more regularly distributed than their conspecific juveniles (Janzen 1970). In the third analysis, the recruitment of saplings of species in the emergent and canopy layers increased significantly and in proportion with mortality, suggesting that the dominant species suffer higher mortality than do less common species. This trend is not so apparent in the understory, and the treelet and shrub layers. The results imply that a dynamic equilibrium process, which prevents competitive exclusion and maintains space for minor species, may be active among the species in the upper layers (particularly the emergent layer); however, such a dynamic equilibrium condition is not due exclusively to the reduced recruitment of saplings near conspecific adults, and the dynamic equilibrium condition is not prevalent among the lower story species.     

Is farther seed dispersal better? Spatial patterns of offspring mortality in three rainforest tree species with different dispersal abilities

The paradigm that tropical trees with farther seed dispersal experience lower offspring mortality is currently based on within-species studies documenting higher survival of offspring located farther from conspecific adults and/or closer to light gaps. We determined whether the paradigm also holds among species by comparing spatial patterns of offspring mortality among three sympatric Neotropical rainforest tree species with the same seed dispersers but with different dispersal abilities. First, we assessed spatially non-random mortality for each species by measuring spatial shifts of the population recruitment curve (PRC) with respect to conspecific adults and light gaps across three early life stages: dispersed seeds, young seedlings and old seedlings. Then, we determined whether PRC shifts were greater for the species with short dispersal distances than for the species with greater dispersal distances. We found that the PRC shifted away from conspecific adults consistently across life stages, but we found no consistent PRC shifts towards gaps, suggesting that mortality was related more to the proximity of conspecifics than to absence of light gaps. PRC shifts away from adults were greatest in the species with the lowest dispersal ability, supporting the paradigm. Differential PRC shifts caused the spatial distribution of offspring to become almost independent with respect to adult trees and gaps in all three species, despite large differences in seed dispersal distance among these species. Our results provide direct empirical support for the paradigm that among tropical trees, species with farther seed dispersal are less impacted by spatially non-random mortality than are similar species with shorter dispersal distances. Thus, greater dispersal effectiveness merits extra investments of trees in seed dispersal ability, even at the cost of fecundity.     

Spatial patterns of acorn dispersal by rodents: do acorn crop size and ungulate presence matter?

Seed dispersal is qualitatively effective when it increases recruitment probability. A poorly studied factor likely affecting recruitment is the spatial distribution of dispersed seeds. Seed-caching animals are thought to disperse seeds in a way that reduces clumping and density to impede cache pilfering. Furthermore, dispersal might differ depending on whether the seed is immediately consumed versus cached for later consumption, and might differ depending on the ecological context. The main objectives of this study were to determine: 1) the spatial pattern of seed dispersal by rodents in a heterogeneous environment; 2) whether the patterns differ among years and among acorn competitor exclosure treatments, and 3) whether rodents create different spatial patterns of dispersal for acorns that are cached versus consumed immediately following dispersal. We studied the degree of spatial aggregation of acorn dispersal by rodents using two different estimators derived from the Ripley K and the Diggle G functions. We also analyzed various metrics of dispersal distances. For both analyses we used observed acorn dispersal patterns in two years differing in crop size and inside versus outside exclosures restricting access to acorn-consuming ungulates. During 2003, a year with a larger crop size, maximum seed dispersal distances were less, and the pattern of dispersed seeds was more clumped, than in 2004, a year with a smaller crop size. Median dispersal distances did not differ between years. In the presence of ungulates, seed dispersal was marginally sparser than in their absence. Cached acorns were dispersed more sparsely than acorns eaten immediately. These results have important implications for the quality of seed dispersal for oak recruitment that are likely relevant to other systems as well.     

When do localized natural enemies increase species richness?

The Janzen–Connell hypothesis states that local species‐specific density dependence, mediated through specialist enemies of offspring such as fungal pathogens and insect seed predators, can facilitate coexistence of species by preventing recruitment near conspecific adults. We use spatially explicit simulation models and analytical approximations to evaluate how spatial scales of offspring and enemy dispersal affect species richness. In comparison with model communities in which both offspring and enemies disperse long distances, species richness is substantially decreased when offspring disperse long distances and enemies disperse short distances. In contrast, when both offspring and enemies disperse short distances species richness more than doubles and adults of each species are highly spatially clumped. For the range of conditions typical of tropical forests, locally dispersing specialist enemies may decrease species richness relative to enemies that disperse long distances. In communities where dispersal distances of both offspring and enemies are short, local effects may enhance species richness.     

Seeds and seedlings of two neotropical montane understory shrubs respond differently to anthropogenic edges and treefall gaps

Edges resulting from forest clear-cutting and treefall gaps can affect plant populations and consequently the distribution of species across landscapes. These two types of disturbance might interact to exacerbate or ameliorate “edge effects”, a rarely tested possibility. We focused on the effects of distance from forest edge (0–10, 30–40, 60–70, and 190–200 m) and habitat within forest fragments (treefall gaps and intact forest) on the early stages of development of Palicourea gibbosa and Faramea affinis, two common shrubs of montane forests in southwest Colombia. Seed germination and seedling growth did not change with distance from forest edge. Within forest fragments, however, seed germination and seedling growth were higher in treefall gaps than in intact forest understory for both species. In contrast, seed predation was influenced by distance from forest edge and in P gibbosa it depended on habitat. Seed predation was highest in the forest interior (190–200 m from forest edge) and in P. gibbosa this was true only in treefall gap habitats. These results suggest that animal mediated processes such as post-dispersal seed predation are more likely than physiological processes to be affected by anthropogenic edges. Our results provide some evidence that treefall gaps may interact with “edge effects”, however, they are inconclusive as to whether they exacerbate or ameliorate them. Received: 31 August 1998 / Accepted: 18 February 1999     

Stochastically driven adult–recruit associations of tree species on Barro Colorado Island

The spatial placement of recruits around adult conspecifics represents the accumulated outcome of several pattern-forming processes and mechanisms such as primary and secondary seed dispersal, habitat associations or Janzen–Connell effects. Studying the adult–recruit relationship should therefore allow the derivation of specific hypotheses on the processes shaping population and community dynamics. We analysed adult–recruit associations for 65 tree species taken from six censuses of the 50 ha neotropical forest plot on Barro Colorado Island (BCI), Panama. We used point pattern analysis to test, at a range of neighbourhood scales, for spatial independence between recruits and adults, to assess the strength and type of departure from independence, and its relationship with species properties. Positive associations expected to prevail due to dispersal limitation occurred only in 16% of all cases; instead a majority of species showed spatial independence (≈73%). Independence described the placement of recruits around conspecific adults in good approximation, although we found weak and noisy signals of species properties related to seed dispersal. We hypothesize that spatial mechanisms with strong stochastic components such as animal seed dispersal overpower the pattern-forming effects of dispersal limitation, density dependence and habitat association, or that some of the pattern-forming processes cancel out each other.     

Seed predation patterns favor the regeneration of dominant species in forest gaps compared with the understory in an oak-pine mixed forest

Recent studies have demonstrated the higher likelihood of regeneration in forest gaps compared with the understory for the dominant species in pine-oak mixed forest. Here, we tested whether rodent seed predation or dispersal was beneficial for gap regeneration. We tracked the seed predation and dispersal of Quercus aliena var. acuteserrata and Pinus armandii using coded plastic tags in the forest understory close to gaps. Our results demonstrated that the proportions of initial buried seeds of both species were significantly more abundant in the forest understory compared with gaps. After seed caching, however, significantly lower proportions of the seeds of both species survived in the forest understory compared with gaps during the 30-day observation period. The final survival proportions of the seeds cached in the forest understory were lower than those cached in the gaps the next spring, which indicated that small rodents rarely retrieved scatter-hoarded seeds from forest gaps. Our findings suggest that rodent seed predation patterns contribute to the regeneration of the dominant species in gaps compared with the understory in a pine-oak mixed forest. In the study area, reforestation usually involves planting seedlings but direct sowing in forest gaps may be an alternative means of accelerating forest recovery and successional processes.