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Restoration of tropical forest depended in large part on seed dispersal by fruit-eating animals that transported seeds into planted forest patches. We tested effectiveness of dispersal agents as revealed by established recruits of tree and shrub species that bore seeds dispersed by birds, bats, or both. We documented restoration of dispersal processes over the first 76 months of experimental restoration in southern Mexico. Mixed-model repeated-measures randomized-block ANOVAs of seedlings recruited into experimental controls and mixed-species plantings from late-secondary and mature forest indicated that bats and birds played different roles in the first years of a restoration process. Bats dispersed pioneer tree and shrub species to slowly regenerating grassy areas, while birds mediated recruitment of later-successional species into planted stands of trees and to a lesser extent into controls. Of species of pioneer trees and shrubs established in plots, seven were primarily dispersed by birds, three by bats and four by both birds and bats. Of later-successional species recruited past the seedling stage, 13 were of species primarily dispersed by birds, and six were of species dispersed by both birds and bats. No later-successional species primarily dispersed by bats established in control or planted plots. Establishment of recruited seedlings was ten-fold higher under cover of planted trees than in grassy controls. Even pre-reproductive trees drew fruit-eating birds and the seeds that they carried from nearby forest, and provided conditions for establishment of shade-tolerant tree species. Overall, after 76 months of cattle exclusion, 94% of the recruited shrubs and trees in experimental plots were of species that we did not plant.

Theory predicts widespread loss of genetic diversity from drift and inbreeding in trees subjected to habitat fragmentation, yet empirical support of this theory is scarce. We argue that population genetics theory may be misapplied in light of ecological realities that, when recognized, require scrutiny of underlying evolutionary assumptions. One ecological reality is that fragment boundaries often do not represent boundaries for mating populations of trees that benefit from long-distance pollination, sometimes abetted by long-distance seed dispersal. Where fragments do not delineate populations, genetic theory of small populations does not apply. Even in spatially isolated populations, where genetic theory may eventually apply, evolutionary arguments assume that samples from fragmented populations represent trees that have had sufficient time to experience drift, inbreeding, and ultimately inbreeding depression, an unwarranted assumption where stands in fragments are living relicts of largely unrelated predisturbance populations. Genetic degradation may not be as important as ecological degradation for many decades following habitat fragmentation. /// La teoría predice una gran pérdida de diversidad genética por deriva y endogamia de árboles sujetos a la fragmentación de hábitat; sin embargo, el soporte empírico de esta teoría es escaso. Argumentamos que la teoría de la genética de poblaciones puede ser mal aplicada a la luz de realidades ecológicas que, al ser reconocidas, requieren del escrutinio de los supuestos evolutivos subyacentes. Una realidad ecológica es que los límites de los fragmentos a menudo no representan los límites para las poblaciones de árboles que se benefician con la polinización a larga distancia, a veces favorecida por la dispersión de semillas a larga distancia. Donde los fragmentos no delinean poblaciones, la teoría genética de las poblaciones pequeñas no aplica. Aun en poblaciones espacialmente aisladas, donde la teoría genética eventualmente puede aplicar, los argumentos evolutivos asumen que las muestras de poblaciones fragmentadas representan árboles que han tenido suficiente tiempo para experimentar deriva, endogamia y, finalmente, depresión por endogamia, una suposición que carece de base y donde los individuos de los fragmentos son relictos vivientes de poblaciones no emparentadas antes de la perturbación. La degradación genética puede no ser tan importante como la degradación ecológica durante muchas décadas después de la fragmentación del hábitat.

Hunting of hornbills by tribal communities is widespread in logged foothill forests of the Indian Eastern Himalaya. We investigated whether the decline of hornbills has affected the dispersal and recruitment of 3 large-seeded tree species. We hypothesized that 2 low-fecundity tree species, Chisocheton paniculatus and Dysoxylum binectariferum (Meliaceae) bearing arillate fruits, are more dispersal limited than a prolifically fruiting drupaceous tree Polyalthia simiarum (Annonaceae), which has potential dispersers other than hornbills. We estimated the abundance of large avian frugivores during the fruiting season along transects in 2 protected and 2 disturbed forests. We compared recruitment of the tree species near (<10 m) and far (10-40 m) from parent trees at protected and disturbed sites. Median abundance of Great (Buceros bicornis), Wreathed (Aceros undulatus), and Oriental Pied Hornbills (Anthracoceros albirostris) were significantly lower in disturbed forests, but sites did not differ in abundances of the Mountain Imperial Pigeon (Ducula badia). Overall, tree species showed more severely depressed recruitment of seedlings (77% fewer) and juveniles (69% fewer) in disturbed than in protected forests. In disturbed forests, 93% fewer seedlings of C. paniculatus were beyond parental crowns, and a high number of all seedlings (42%) accumulated directly under reproductive adults. In contrast, D. binectariferum and P. simiarum were recruitment rather than dispersal limited, with fewer dispersed seedlings surviving in disturbed than in protected forests. Results are consistent with the idea that disturbance disrupts mutualisms between hornbills and some large-seeded food plants, with the caveat that role redundancy within even small and specialized disperser assemblages renders other tree species less vulnerable to loss of regular dispersal agents.

Because bird species are lost when forests are fragmented into small parcels, trees that depend on fruit-eating birds for seed dispersal may fail to recruit seedlings if dispersal agents disappear. We tested this prediction in rainforest in the East Usambara Mountains of Tanzania, by using the endemic tree Leptonychia usambarensis (Sterculiaceae) and birds that disperse its seeds. We investigated bird abundance and Leptonychia dispersal ecology in fragments isolated for >70 yr, as compared with 3,500 ha of continuous forest. Birds that dispersed Leptonychia seeds in continuous forest were rare or absent in small fragments, where fewer seeds were removed from each tree, far fewer seedlings occurred >10 m from parent trees, and far more seedlings occurred in dense aggregations under parental crowns. Overall, our samples showed that fewer juvenile Leptonychia recruited in fragments than in continuous forest. We provide solid evidence that deficient dispersal due to habitat fragmentation seriously impacts the reproductive cycle of a tropical bird-dispersed tree.

Stepping‐stone restoration plantings can reconcile conservation goals and local land‐use needs in highly fragmented ecosystems. We explored how initial planting composition influences recruiting plant species density, diversity, abundance, and forest structure in a 13‐year‐old restoration experiment in Los Tuxtlas, Veracruz, Mexico. Treatments included eight fenced plantings with animal‐dispersed species, eight plantings with wind‐dispersed species, eight unplanted plots to favor natural succession, and eight plots in the primary forest as reference sites. We predicted that by attracting more seed dispersers, animal‐dispersed plantings would most closely resemble the primary forest. A census of trees taller than 2 m showed that while wind‐dispersed plantings had more recruits, the animal‐dispersed plantings most closely resembled the primary forest in pioneer abundance, species density and abundance of biotically dispersed and abiotically dispersed plants, individual tree basal area (in square meters per hectare), and vertical structure. The wind‐dispersed plantings more closely approximated the forest in non‐pioneer abundance and community composition. However, restoration treatments were more similar to each other than to the primary forest and did not differ in plant diversity. Animal‐dispersed and wind‐dispersed plantings did not differ in non‐pioneer species density and matched the primary forest in total plot basal area. Higher abundance of trees in wind‐plantings is explained by lower establishment limitations, seed legacy effects, and rapid reproduction of a few planted species. As the experiment continues, we expect treatment effects on seed dispersers will more strongly influence the recruiting plant community, leading the animal‐dispersed plantings to more closely resemble the primary forest in diversity and forest structure.

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed-dispersal limitation and seedling-establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (Eit ) were higher for animal-dispersed than wind-dispersed species, and for nonpioneer than pioneer species. Non-pioneers and animal-dispersed species had higher expected relative recruit abundance ( it ) in primary forest (median of 4 seeds recruit–1) whereas in secondary forest wind-dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind-dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.

Maintenance of biodiversity in tropical agrarian landscapes is challenging in the face of anthropomorphic simplification of habitats. As part of an experiment testing influences of planting treatment on tree recruitment in southern Mexico, counts of bird species were made over 10 years in twenty-four 30 × 30-m fenced plots in over-grazed pasture. Plots were planted with native tree species or left as unplanted controls in 2006. Annual censuses of birds in the plots from 2007–2016 indicated statistically significant increases in the number of fruit-eating species and individuals as vegetation matured, but increases in non-frugivorous species and individuals over the decade were not significant. Among four species of planted animal-dispersed trees that bore fruit during this time, Cecropia obtusifolia consistently produced substantial crops after 2009. In 2015, all 53 planted or passively recruited female trees of mature size of this species bore fruit. The summed body masses of fruit-eating birds in each of 24 plots were significantly correlated with rank order of available fruit per plot. Differential use of habitat patches in an agrarian landscape suggested substantial value to frugivores, but less to non-frugivorous birds than expected.

A profusion of fruit forms implies that seed dispersal plays a central role in plant ecology, yet the chance that an individual seed will ultimately produce a reproductive adult is low to infinitesimal. Extremely high variance in survival implies that variations in fruit production or transitions from seed to seedling will contribute little to population growth. The key issue is that variance in survival of plant life-history stages, and therefore the importance of dispersal, differs greatly among and within plant communities. In stable communities of a few species of long-lived plants, variances in seed and seedling survival are immense, so seed-to-seedling transitions have little influence on overall population dynamics. However, when seedlings in different circumstances have very different chances of survival—in ecological succession, for example, or when dispersed seeds escape density-dependent mortality near parent trees—the biased survival of dispersed seeds or seedlings in some places rather than others results in pervasive demographic impacts.

Many large-seeded Neotropical trees depend on a limited guild of animals for seed dispersal. Fragmented landscapes reduce animal abundance and movement, limiting seed dispersal between distant forest remnants. In 2006, experimental plantings were established in pasture to determine whether plantings enhance seed dispersal and, ultimately, seedling recruitment. We examined patterns of naturally recruited seedlings of Ocotea uxpanapana, a large-seeded bird-dispersed tree endemic to southern Mexico that occurs in the surrounding landscape. We used GIS and least-cost path analysis to ask: (1) Do restoration efforts alter recruitment patterns? (2) What is the importance of canopy cover and likely dispersal pathways to establishment? Patterns of seedling establishment indicated that dispersal agents crossed open pastures to wooded plots. Recruitment was greatest under woody canopies. Also, by reducing movement cost or risk for seed dispersers, wooded canopies increased influx of large, animal-dispersed seeds, thereby restoring a degree of functional connectivity to the landscape. Together, canopy openness and path distance from potential parent trees in the surrounding landscape explained 73% of the variance in O. uxpanapana seedling distribution. Preliminary results suggest that strategic fenced plantings in pastures increase dispersal and establishment of large-seeded trees, thereby accelerating forest succession in restorations and contributing to greater connectivity among forest fragments.

Grasses are recognized as a critical regeneration barrier in tropical pastures, yet the effects of rodents and rodent–grass interactions are not well understood. As selective foragers, rodents could shape tree communities, moderating biodiversity in regenerating tropical landscapes. We utilized a fully crossed two-way factorial design to examine the effect that grasses, rodents, and their interaction had on tree seedling establishment in pasture habitat. We followed two separate tree cohorts for 1 year each within the experimental framework. Multiple cohorts were used to better represent successional tree species variation and responses. Trees species were characterized by a gradient of seed masses and as pioneer or persistent successional type. Both cohort seedlings were altered when rodents were present compared to control treatments. In Cohort 1, rodents adversely affected seedlings of persistent tree species only in the absence of grass. In Cohort 2, seedlings of persistent tree species were decimated by rodents in the absence or presence of grass. In both cohorts, seedlings of persistent species established better in grass treatments, while seedlings of pioneer tree species were strongly suppressed. Tree species seed mass positively correlated with seedling establishment across all treatments except no grass–rodent treatments. Strong suppression of tree seedlings by rodents (Sigmodon toltecus) is a novel result in tropical land recently released from agriculture. One implication is that selective foraging by rodents on large-seeded persistent tree species may be facilitated by the removal of grass. Another implication is that temporary rodent control in pastures may permit higher establishment of deep-forest persistent species.