Are Introduced Species Better Dispersers Than Native Species? A Global Comparative Study of Seed Dispersal Distance

We provide the first global test of the idea that introduced species have greater seed dispersal distances than do native species, using data for 51 introduced and 360 native species from the global literature. Counter to our expectations, there was no significant difference in mean or maximum dispersal distance between introduced and native species. Next, we asked whether differences in dispersal distance might have been obscured by differences in seed mass, plant height and dispersal syndrome, all traits that affect dispersal distance and which can differ between native and introduced species. When we included all three variables in the model, there was no clear difference in dispersal distance between introduced and native species. These results remained consistent when we performed analyses including a random effect for site. Analyses also showed that the lack of a significant difference in dispersal distance was not due to differences in biome, taxonomic composition, growth form, nitrogen fixation, our inclusion of non-invasive introduced species, or our exclusion of species with human-assisted dispersal. Thus, if introduced species do have higher spread rates, it seems likely that these are driven by differences in post-dispersal processes such as germination, seedling survival, and survival to reproduction.     

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.     

Seed dispersal of the non-native invasive tree <Emphasis Type="Italic">Ailanthus altissima</Emphasis> into contrasting environments

Ailanthus altissima has a long history of invasion in urban areas and is currently spreading into suburban and rural areas in the eastern U.S. The objectives of our study were to (1) determine whether A. altissima seed dispersal distance differed between populations on the edges of open fields and intact deciduous forest, and (2) determine whether dispersal differed for north and south winds. We also assessed the relationship between seed characteristics and distance from source populations in fields and forests, and whether seeds disperse at different rates throughout the dispersal season. Using two fields, two intact forest stands, and one partially harvested stand, we sampled the seed rain at 10 m intervals 100 m into each site from October to April 2002–2003. We compared seed density in field and intact forests using a three-way ANOVA with distance from source, wind direction, and environmental structure as independent variables. To assess the accuracy of common empirical dispersal models, mean seed density data at each site were fitted with alternative regression models. We found that mean seed dispersal distance depended on environmental structure and wind direction, a result driven in large part by dispersal at a single site where seed density did not decline with distance. The two alternative regression models fit each site’s dispersal curve equally well. More seeds were dispersed early than in mid- or late-season. Large, heavy seeds traveled as far as small light seeds. Turbulent winds appear to be necessary for seed release, as indicated by a wind tunnel experiment. A. altissima is able to disperse long distances into fields and into mature forests, and can reach canopy gaps and other suitable habitats at least 100 m from the forest edge. It is an effective disperser and can spread rapidly in fragmented landscapes where edges and other high light environments occur. These conditions are increasingly common throughout the eastern U.S. and in other temperate regions worldwide.     

Seed dispersal in Erythronium grandiflorum (Liliaceae)

Primary and secondary seed dispersal was investigated for the glacier lily Erythronium grandiflorum in the Colorado Rocky Mountains. These heavy seeds have no obvious adaptations for biotic or abiotic dispersal, but can be thrown short distances when the dehiscent fruits are shaken by wind. We used sticky traps to measure primary transport of seeds up to 1 m away from individual plants. A seed cafeteria experiment examined the role of ants and rodents in secondary seed transport. Primary dispersal by wind was positively skewed and median transport distances were influenced by variation in plant height. Secondary dispersal was negligible compared to Viola nuttallii, an elaiosome-bearing species. Thus, seed dispersal was highly restricted in E. grandiflorum, and a 1 m radius encompassed the modal section of the seed dispersal curve. The seed dispersal component of gene flow was quantified and combined with previous measurements of pollen flow to yield a more complete estimate of Wright's neighborhood size, N _e, for E. grandiflorum. The lack of a special seed dispersal mechanism in E. grandiflorum is discussed in terms of a source-sink model for seedling establishment with respect to distance from the parental plants.     

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.     

Seed dispersal distance,seed morphology,and recruitment in the Chilean sclerophyllous tree Quillaja saponaria: implications for passive restoration in a semiarid ecosystem

Recolonization of wind-dispersed tree species in degraded areas may decline with distance from remnant forest fragments because seed rain frequently decreases with distance from the seed source. However, regeneration of these species may be even more limited to sites close to the seed source if dispersal distance is negatively affected by seed mass, and germination probability is positively affected by seed mass. We evaluated these hypotheses in a Mediterranean-type ecosystem of central Chile, using the wind-dispersed tree species Quillaja saponaria. We assessed the seed rain curve in a degraded open area adjacent to a remnant forest fragment of this species, and related seed mass with dispersal distance from the seed source. Then, we evaluated the relationship between seed mass, germination, and seedling growth, and if seeds that fall nearer the seed source have greater germination probability. We found a decreasing seed rain with the distance from the seed source. Seed mass was not related to dispersal distance, although seeds with higher wing area dispersed further. Germination probability was significantly and positively related to the seed mass. We observed no significant relationship between distance and germination probability. We conclude that germination probability of this species does not vary along the seed rain curve, and that the recruitment density would be greater near the seed source only due to decreasing seed rain with distance. Our results suggest that this species has the potential to be passively restored in degraded areas, especially within the first 70 m from the remnant forest fragments.

     

Seed dispersal in a cycad,Macrozamia riedlei

The influence of animals on patterns of seed dispersal has not been well explored, despite the importance of seed dispersal in plant population biology. Furthermore, the exploitation of animals as dispersal agents has been considered one reason for the replacement of gymnosperms by angiosperms in the Cretaceous. We studied seed dispersal in a cycad, Macrozamia riedlei, and showed that in the absence of animals, ripe seeds drop directly beneath the parent not more than 40 cm from the base. Several animals eat the fleshy outer integument of the seeds, and possums (Trichosurus vulpecula), transport seeds up to 24 m from the parent. The postdispersal positioning of seeds gave a log density-log distance relationship of y = 7.20 – 2.85x (r²= 0.96, N = 17). Effective local dispersal may have contributed to the persistence of cycads in the world flora for the last 200 million years.     

Seed dispersal characteristics of Brassavola nodosa (Orchidaceae)

We used mathematical models for wind-dispersed seeds and wind-tunnel experiments to predict modal seed dispersal distance of the Neotropical orchid Brassavola nodosa under conditions approximating those found in its natural habitat: mangrove islands in Belize, Central America. Key variables in a simple ballistic model for predicting modal dispersal distance (xm) of an individual seed include: height of release (h); free-stream velocity (Uc); and terminal velocity of the seed (Ut): xm = h Uc/Ut. Modal dispersal distance of dust-like orchid seeds were predicted adequately by this ballistic model at low wind velocities and low release heights, but it underestimated the increasing importance of turbulence at higher wind velocities and greater release heights. We estimated the magnitude and relative importance of one measure of turbulence, vertical mixing velocity (W*), on xm in wind tunnel experiments. Our estimates of W* were within the same order of magnitude as those found for other small dust-like seeds and pollen. In high turbulence conditions, incorporation of vertical mixing velocity effects into the ballistic model of seed dispersal overestimated modal seed dispersal distances.     

Seed dispersal effectiveness by a large‐bodied invasive species in defaunated landscapes

Animal‐dispersed plants are increasingly reliant on effective seed dispersal provided by small‐bodied frugivores in defaunated habitats. In the Neotropical region, the non‐native wild pig (Sus scrofa) is expanding its distribution and we hypothesized that they can be a surrogate for seed dispersal services lost by defaunation. We performed a thorough analysis of their interaction patterns, interaction frequencies, seed viability, and characteristics of the seed shadows they produce. We found 15,087 intact seeds in 56% of the stomachs and 5,186 intact seeds in 90% of the scats analyzed, 95% of which were smaller than 10 mm in diameter. Wild pigs were the third most effective disperser among 21 extant frugivore species in a feeding trail experiment in terms of quantity of seeds removed. Gut retention time was 70 ± 23 hr, indicating wild pigs can promote long‐distance seed dispersal. Seed survival after seed handling and gut passage by wild pigs was positively related with seed size, but large seeds were spat out and only smaller seeds were defecated intact, for which we observed a positive or neutral effect on germination relative to manually de‐pulped seeds. Finally, deposition of seeds was four times more frequent in unsuitable than suitable sites for seedling recruitment and establishment. Seed dispersal effectiveness by wild pigs is high in terms of the quantity of seeds dispersed but variable in terms of the quality of the service provided. Our study highlights that negative and positive effects delivered by non‐native species should be examined in a case by case scenario. Abstract in Portuguese is available with online material.     

Seed dispersal distance classes and dispersal modes for the European flora

Motivation

Although dispersal ability is one of the key features determining the spatial dynamics of plant populations and the structure of plant communities, it is also one of the traits for which we still lack data for most species. We compiled a comprehensive dataset of seed dispersal distance classes and predominant dispersal modes for most European vascular plants. Our seed dispersal dataset can be used in functional biogeography, dynamic vegetation modelling and ecological studies at local to continental scales.

Main Types of Variables Contained

Species were classified into seven ordered classes with similar dispersal distances estimated based on the predominant dispersal mode, the morphology of dispersal units (diaspores or propagules), life form, plant height, seed mass, habitat and known dispersal by humans. We evaluated our results by comparing them with dispersal distances calculated using the ‘dispeRsal’ function in R.

Spatial Location

Europe.

Time Period

Present.

Major Taxa and Level of Measurement

The seed dispersal dataset contains information on dispersal distance classes and the predominant dispersal mode for 10,327 most frequent and locally dominant European vascular plant species.

Software Format

Data are available in .csv format.     

Seed dispersal and realized gene flow of two forest orchids in a fragmented landscape

• Species with vast production of dust‐like windborne seeds, such as orchids, should not be limited by seed dispersal. This paradigm, however, does not fit recent studies showing that many sites suitable for orchids are unoccupied and most seeds land close to their maternal plant. To explore this issue, we studied seed dispersal and gene flow of two forest orchid species, Epipactis atrorubens and Cephalanthera rubra, growing in a fragmented landscape of forested limestone hills in southwest Bohemia, Czech Republic.
• We used a combination of seed trapping and plant genotyping methods (microsatellite DNA markers) to quantify short‐ and long‐distance dispersal, respectively. In addition, seed production of both species was estimated.
• We found that most seeds landed very close to maternal plants (95% of captured seeds were within 7.2 m) in both species, and dispersal distance was influenced by forest type in E. atrorubens. In addition, C. rubra showed clonal reproduction (20% of plants were of clonal origin) and very low fruiting success (only 1.6% of plants were fruiting) in comparison with E. atrorubens (25.7%). Gene flow was frequent up to 2 km in C. rubra and up to 125 km in E. atrorubens, and we detected a relatively high dispersal rate among regions in both species.
• Although both species occupy similar habitats and have similar seed dispersal abilities, C. rubra is notably rarer in the study area. Considerably low fruiting success in this species likely limits its gene flow to longer distances and designates it more sensitive to habitat loss and fragmentation.

     

Directed vertebrate-mediated seed dispersal of a fleshy-fruited amaryllid in a heterogenous habitat

Most plants with fleshy fruits have seeds that are ingested by animals, but a less well-understood mode of seed dispersal involves fleshy fruits containing seeds that are discarded by frugivorous animals because they are too large or toxic to be ingested. We studied the seed dispersal biology of Haemanthus deformis, an amaryllid lily species found in a mosaic of bush clumps in a grassland matrix in South Africa. We asked whether seed dispersal is directed in and among bush clumps and whether germination and survival are greater for seeds dispersed to bush clumps than for those dispersed into grassland. Using camera trapping, we found that fruits are consumed mainly by birds and rodents. The pulp was removed from the seeds which were then discarded without ingestion. While many seeds were dispersed close to the parent plant, most (c. 78.5%) were dispersed further than 1 m away from the parent plant. Longer distance dispersal resulted mainly from birds flying off with fruits in their bill or from rodents engaging in scatter-hoarding behavior. Seedling survival was most successful within bush clumps as compared to grasslands and shade was identified as a primary requirement for seedling survival. Seeds from which the fruit pulp had been removed germinated faster than those in intact fruits. Haemanthus deformis deploys a system of directed seed dispersal, whereby both birds and rodents contribute to the dispersal of seeds within patchy bush clumps that are favorable for seedling survival.     

Seed Rain and Seed Dispersal Agents in Pure and Mixed Plantations of Native Trees and Abandoned Pastures at La Selva Biological Station, Costa Rica

Lack of seed dispersal can be an important obstacle to natural regeneration (NR) of degraded pastures in the humid tropics. Tree plantations can facilitate secondary forest succession by attracting seed dispersal agents from nearby forests. We studied seed rain and seed dispersal agents in 12–13 years old pure and mixed native tree plantations at La Selva Biological Station, Costa Rica from July to December 2004. Plantations of Balizia elegans (5,522), Dipteryx panamensis (2,263), and Jacaranda copaia (2,091) had the greatest total seed abundance; treatments with the least total seed abundance were Calophyllum brasiliense (56), nonplanted abandoned pasture control 2 (353), Mixed Species 2 (389), and control 1 (836). Plantations of J. copaia and Hyeronima alchorneoides had the greatest seed species richness density, whereas the lowest seed species richness was found in the control treatments. The NR plots had more seeds dispersed by wind, whereas in the plantations, the most important dispersal agents were birds and mammals. The most abundant seeds were those of Miconia spp. (14,492), Psychotria bracheata (2,252), and the Poaceae family (1,346), all species from early successional stages. Plantations of native species are effective in attracting seed dispersal agents and thus facilitating regeneration of degraded pasturelands in the region.     

Seed dispersal effectiveness in a plant–lizard interaction and its consequences for plant regeneration after disperser loss

Mutualistic disruptions, such as those promoted by the loss of seed dispersers, can have negative effects on the plant regeneration of those species that strongly depend upon them. In order to adequately assess how plant communities are affected by such disruptions, we need to know the importance of the dispersal phase, both in its quantitative and qualitative components. We examined this in the narrow interaction between the shrub Daphne rodriguezii and its (only) disperser, the lizard Podarcis lilfordi. We quantified fruit removal and the effect of fruit/seed-size selection, seed treatment in the disperser’s guts and seed deposition patterns on seedling emergence and survival. In the only locality in which lizards persist, they removed most fruits and showed preference for larger ones in one of the two study years. Seed treatment in lizard’s guts had no effect on germination, although it tended to reduce the effect of seed size on germination (differences between large vs. small seeds in seed germination were higher for non-ingested seeds). Probability of seedling emergence, but not survival, was higher in the locality with lizards. Dispersed seeds under heterospecific shrubs showed higher seedling survival than those under conspecifics in all localities, especially the year with higher rainfall. Our findings support that the movement of seeds to nurse shrubs by lizards is the most important component of the seed dispersal process in the only remaining locality where both species coexist.     

How important is long‐distance seed dispersal for the regional survival of plant species?

Long‐distance seed dispersal is generally assumed to be important for the regional survival of plant species. In this study, we quantified the importance of long‐distance seed dispersal for regional survival of plant species using wind dispersal as an example. We did this using a new approach, by first relating plant species’ dispersal traits to seed dispersal kernels and then relating the kernels to regional survival of the species. We used a recently developed and tested mechanistic seed dispersal model to calculate dispersal kernels from dispersal traits. We used data on 190 plant species and calculated their regional survival in two ways, using species distribution data from 36,800 1 km²‐grid cells and 10,754 small plots covering the Netherlands during the largest part of the 20th century. We carried out correlation and stepwise multiple regression analyses to quantify the importance of long‐distance dispersal, expressed as the 99‐percentile dispersal distance of the dispersal kernels, relative to the importance of median‐distance dispersal and other plant traits that are likely to contribute to the explanation of regional survival: plant longevity (annual, biennial, perennial), seed longevity, and plant nutrient requirement. Results show that long‐distance dispersal plays a role in determining regional survival, and is more important than median‐distance dispersal and plant longevity. However, long‐distance dispersal by wind explains only 1–3% of the variation in regional survival between species and is equally important as seed longevity and much less important than nutrient requirement. In changing landscapes such as in the Netherlands, where large‐scale eutrophication and habitat destruction took place in the 20th century, plant traits indicating ability to grow under the changed, increasingly nutrient‐rich conditions turn out to be much more important for regional survival than seed dispersal.     

金沙江干热河谷滇榄仁种子扩散与种子库特征研究

采用随机取样和跟踪调查的方法对滇榄仁(Terminalia franchetii)种子扩散规律、空间分布特征以及种子库动态进行了系统研究。结果表明,滇榄仁种子扩散受到较强的风力作用影响,具有明显的方向性,扩散距离是其树高的1.6倍以上。不同群落类型中,滇榄仁的土壤种子库存在较大差异,但不同时间段的变化趋势基本一致,即在雨季(6月、8月和10月)数量相对较少,12月到翌年4月,数量逐渐增加。滇榄仁植冠种子库从当年10月开始缓慢脱落,到翌年3月达到脱落高峰(15%以上种子此时脱落),然后再次进入缓慢脱落期,翌年6、7月的种子脱落速率最低,但直到翌年10月仍有部分种子(约15%左右)存在于植株冠层。总体来说,不同类型群落中,滇榄仁幼苗更新不良的原因不应是缺乏种源,可能与种子质量以及群落环境等影响群落天然更新的其它因子存在较大关联。同时,滇榄仁土壤种子库数量与植冠种子库的脱落过程存在紧密联系,而植冠种子脱落可能与该区域气候因子存在着较大依存关系,其中,与风力可能存在着正相关,与降雨量和空气相对湿度存在着负相关,这可能是滇榄仁长期以来对干热河谷环境的一种适应特征。     

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’.     

荒漠孑遗植物裸果木种子时空扩散特性

裸果木(Gymnocarpos przewalskii)是亚洲中部荒漠区少有的第三纪孑遗物种,由于气候变化及人为干扰,其自然种群分布范围不断缩小。种子扩散作为植物生活史过程中的重要阶段,不仅对物种生存及其多样性至关重要,还影响物种分布范围和局部丰度。2015年和2016年分别在新疆哈密地区,采用布设种子收集器的方法,对其自然种群种子扩散的时空动态进行了定点连续观测。结果表明:该物种于当年6月上旬开始扩散,2015年略早于2016年。每年种子扩散持续时间约两个月,扩散趋势为单峰曲线,且呈集中大量扩散的模式,扩散高峰期与当年初次月降水高峰期吻合;在顺风的正南和东南方向上,种子扩散密度大且距离远;种子扩散主要集中在母株冠幅下,随着距母株距离的增加,种子扩散密度减少,二者间存在极显著的负相关性(P0.01),由于裸果木枝条繁多,对风力强度起到了一定的阻碍作用,可能是造成种子集中扩散在母株下的原因。裸果木种子扩散受外界环境(降水、风向)和自身因素等方面的影响,当种子在大量降水前完成扩散,将有利于种子在适宜的微生境萌发,是对多风、干旱的恶劣生境的一种长期适应。     

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.