Research Article |
Corresponding author: Gregory H. Adler ( adler@uwosh.edu ) Academic editor: Cássio Cardoso Pereira
© 2023 Autumn B. Phillips-Lewis, Thomas D. Lambert, Gregory H. Adler.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Phillips-Lewis AB, Lambert TD, Adler GH (2023) Influence of tree-fall gaps on directional seed dispersal by small mammals in Central Panama. Neotropical Biology and Conservation 18(1): 73-82. https://doi.org/10.3897/neotropical.18.e97653
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Small mammals, particularly rodents, are often important seed-dispersal agents in Neotropical forests. Directional seed dispersal into tree-fall gaps may enhance seedling survival of light-demanding species and thus influence forest regeneration. To examine this proposition, we tracked seeds of a light-demanding palm (Attalea butyracea), with a focus on spiny rats (Proechimys semispinosus), the most-likely seed-removal agents. We established seed-removal stations at three distances relative to 28 gaps (gap center, gap edge, and intact forest 10 m from a gap edge) in a lowland forest in Central Panama. We placed five fresh fruits (with their seed) in semi-permeable exclosures to exclude larger mammals at each station and tracked the directions in which seeds were moved and deposited intact. More seeds were moved toward or into gaps when removed from gap center or edge stations; however, seeds dispersed from intact forest stations showed no such directionality. Small mammals may have dispersed seeds into and within tree-fall gaps because they favored caching seeds in areas that offered increased cover, which is typical of gaps, and consequently protection from predation. The lack of directional dispersal from intact forest stations may have been because spiny rats were able to find sufficient cover in the young intact forest that was closer than the gaps. In older forest, the contrast between intact forest and gaps may be greater, resulting in directed dispersal into gaps.
Attalea butyracea, directed dispersal, forest regeneration, Proechimys semispinosus, rodents, spiny rat
Seed dispersal increases the fitness of plants by enabling escape from species-specific enemies and by allowing seeds to be moved into favorable sites (
Numerous studies have examined the role of larger mammals, such as agoutis and monkeys, in seed dispersal in Neotropical forests; however, few studies have examined seed dispersal by smaller mammals whose relative abundance and biomass may surpass those of larger species, especially in disturbed areas (
This study was conducted in Soberania National Park in Central Panama (9.1413, -79.7192, Fig.
Attalea butyracea is a common palm within lowland forests of the northern Neotropics and is particularly abundant in disturbed areas (
Proechimys semispinosus (Fig.
Second-growth forest along the first 2 km at the southeast end of Pipeline Road was thoroughly searched for the presence of tree-fall gaps (Fig.
Seeds from A. butyracea with intact fresh mesocarp and ectocarp were collected, and holes were drilled in one end of each seed using a 1.2-mm drill bit. Such fruits range from 4.5–8.5 cm long and 3–4.5 cm in diameter and rely almost exclusively on animal dispersal (
We used χ2 analysis to determine if the number of seeds dispersed differed from random dispersal expectations. To determine the expected number for each direction, we divided the area around a seed station into four hypothetical quadrants. Therefore, it was expected that 25% of the total removals would be dispersed toward the tree-fall gap, 25% would be dispersed away from the tree-fall gap, and 50% would be dispersed in directions parallel to the gap if the presence of the tree-fall gap was not affecting dispersal direction. This method was not applicable to analyze the direction of seeds dispersed from gap-center seed stations; therefore, the results for those stations are reported as percentages only. We used a mixed-effects analysis of variance (ANOVA) to determine if there was a difference in the distance that seeds were dispersed among tree-fall gaps or stations or between years.
Over the span of two data-collection seasons, we tracked 282 removed and deposited seeds (71 in 2012 and 211 in 2013) at the 28 gaps. Of those, 116 were deposited less than 0.30 m from seed exclosures and therefore were not considered “dispersals” in this study. Additionally, 11 removals were taken into trees and were not included in the analysis. Dispersed seeds were typically deposited under leaf litter.
Of the 155 dispersed seeds, 114 were moved into or toward the tree-fall gap, 25 were moved in directions parallel to the gap, and 16 were moved away from the gap (Table
The number and percentage of seeds that were dispersed in each direction from each seed station location.
Station | Toward or into gap | Parallel to gap | Away from or into gap | |||
---|---|---|---|---|---|---|
Number | % | Number | % | Number | % | |
Center | 72 | 96.0 | - | - | 3 | 4.0 |
Edge | 33 | 57.9 | 13 | 22.8 | 11 | 19.3 |
10 m | 9 | 39.1 | 12 | 52.2 | 2 | 8.7 |
The mixed-effects ANOVA showed that there was no difference in dispersal distance among tree-fall gaps (P = 0.143, d.f. = 13) or between data collection years (P = 0.745, d.f. = 1). There was also no difference in dispersal distance between station locations (P = 0.051, d.f. = 151). Finally, there was no difference in dispersal distance between intact forest and edge stations (P = 0.176, d.f. = 151), nor between gap and edge stations (P = 0.227, d.f. = 151).
Directed dispersal occurs when an animal safely transports a seed and deposits it in a nonrandom location that coincides with more favorable germination conditions. We hypothesized that small mammals such as P. semispinosus, because of their associations with tree-fall gaps and other disturbed areas of forest (
The presence of a tree-fall gap did indeed influence seed-dispersal direction by removal agents at gap center and edge locations. Proechimys semispinosus, in particular, may have directionally-dispersed seeds from center and edge locations within and into tree-fall gaps, respectively, due to its association with tree-fall gaps (
We found no such directional effect with seeds located in intact forest 10 m from a gap edge. This study was conducted in young, second-growth forest where the understory consisted of many structures, including patches of dense understory vegetation that could potentially offer protection from predators of small mammals, including P. semispinosus, while they consumed fruits or seeds or cached seeds. Center and edge stations were located at a tree-fall gap; therefore, individuals likely preferentially dispersed seeds in this direction because the closest structure offering safety was within a gap. However, as the distance from the gap increased, individuals needed to move seeds greater distances to find structures that provided safety. These structures may have been within the tree-fall gap but also could have included other structures within the surrounding second-growth forest if they were closer than the tree-fall gap. According to optimal foraging theory, small mammals should carry the seed to the closest structure that provided safety, whether it was within the tree-fall gap or in a different direction, which resulted in non-directional dispersal.
Our results suggest possibilities for future research to further understand the driving forces underlying directional seed dispersal into tree-fall gaps in Neotropical forests of various ages. While this study was conducted in a young, second-growth forest, the results may have differed if the study were conducted in old-growth forest where there is less cover near the ground in intact forest, and the structural contrast inside and outside of tree-fall gaps would be greater.
Our results demonstrate that small mammals such as spiny rats nonrandomly disperse seeds into tree-fall gaps if such seeds are already within or close to gaps. This directed dispersal may promote recruitment of light-demanding forest trees and consequently influence forest regeneration.
We thank Scott Mangan, Frostburg State University, and the Smithsonian Tropical Research Institute for logistical support, Frank Ammer and Richard Raesly for comments on earlier drafts of this manuscript, Andrew Bretscher and Dan Jarvis for field assistance, Mamadou Coulibaly for assistance with the map of the study area, and an anonymous reviewer for valuable comments.