Agouti reintroduction recovers seed dispersal of a large-seeded tropical tree

The aim of animal reintroductions has mainly been species recovery; only few reintroduction initiatives focus on ecosystem restoration. Therefore, reintroduction consequences on ecological interactions are seldom assessed. We used the interaction between a reintroduced population of agoutis (Dasyprocta leporina) and a vulnerable tropical endemic tree (Joannesia princeps) to examine reintroduction effects on seed dispersal and seedling establishment. To test the outcomes of this interaction, we tracked seeds of J. princeps in two adjacent forest areas with and without reintroduced agoutis. We also assessed if dispersal distances affected seedling survival. To determine seed fate and dispersal distance, we used spool-and-line tracking, together with camera traps to identify dispersers. Agoutis were the only species removing J. princeps seeds, thus dispersal only occurred where agoutis had been reintroduced; in the area without agoutis, all seeds remained intact on the soil, even one year after the experiment's beginning. At the reintroduction area, most seeds were preyed upon by agoutis but 7% remained dispersed and 2% germinated after ten months. Only seeds buried by agoutis were able to germinate. Most dispersed seeds were dispersed 15 m or farther and longer dispersal distances benefited J. princeps, since seedlings farther from a conspecific adult tree had greater survival probability. Agoutis were also seen burying seeds of two other plant species; these mammals have the potential to benefit dozens of large-seeded species in our study system. Agouti reintroduction thus exemplifies the value of trophic rewilding programs to re-establish ecological interactions and restore ecosystem functioning.     

Seed Dispersal and Demography of Pioneer Trees: The Case of Hortia arborea

Abstract: This study investigates the relationships among seed dispersal, patterns of seedling recruitment and the spatial distribution of a pioneer tree (Hortia arborea, Rutaceae) in the Brazilian Atlantic forest. The study was carried out at Dois Irmãos Reserve, a 387.4 ha reserve in northeastern Brazil. Fruit shadow concentrated beneath parental trees (74.6 % < 6 m distant) and fruit removal averaged 5.6 % (0 ‐ 25.5 %). Agoutis (Dasyprocta prymnolopha, Rodentia) ate fruits in the forest and in captivity and were the only vertebrates recorded feeding on Hortia fruits. Agoutis, however, destroyed 86.7 % of the seeds they ingested. In addition, only one seed from an expected number of 1980 germinated in a treefall gap after it passed through the digestive tract of agoutis. Both seed germination and seedling recruitment were restricted to gaps, and occurred among seeds manually released from the pulp or among those from fruits that naturally rotted in gaps. Moreover, exposed seedlings were taller than those covered by plants in a gap. H. arborea appears to self‐maintain populations in the same patches of forest, which are delimited by a pool of old and newly created gaps. More specifically, there is successful recruitment in patches occupied by parents, resulting in dense clumps of H. arborea.     

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.     

Distance and non-randomness of seed dispersal by the dwarf cassowary Casuarius bennetti

To determine the sources of dispersed seeds I inserted unique tags in fallen Aglaia aff flavida seeds before dwarf cassowaries (Casuanus bennetti) ate the fruits containing the seeds Thirty naturally-dispersed, marked seeds were re-located in cassowary droppings m a 400 ha study area The distribution of seed dispersal distances did not differ significantly from a normal distribution with a mean dispersal distance of 388 m, SD= 196 8 Mean distance of dispersed seeds to nearest mature conspecific tree was 170 m, SD= 108 4, dispersed seeds usually landed closer to other conspecifics than their parent The estimated distribution of all seeds (including many undispersed seeds) was leptokurtic, creating high densities near source trees (>0 035 seeds m^-2 within 100 m of bole) that quickly tapered off (<0 002 seeds m^-2 > 100 m from the bole), any density dependent effects are liable to be manifest only near parent trees
Cassowary movement patterns and resting behavior caused non-random dispersal of seeds Seeds were preferentially moved to level sites uphill from their source trees along routes that did not cross.steep terrain Undispersed seeds generally landed downhill from source trees This population of Aglaia would probably contract downhill into smaller, fragmented populations m the absence of cassowary-mediated dispersal     

Spatial–temporal distribution of ornithochorous seeds from an Elaeagnus umbellata community dominating a riparian habitat

The dispersal efficiency and potential distribution of ornithochorous seeds of Elaeagnus umbellata in a riparian habitat were evaluated to clarify this species' establishment site in relation to the disturbance regime of the floodplain. Fruit removal by avian frugivores was monitored using fruit bags, and the spatial distribution of excreted seeds was quantified by seed traps set randomly on a gravel bar as an isolated seed source in the Yoshino River throughout an autumn fruiting season. Although more than 45% of the fruits remained on the twigs in the fruit bags, almost all fruits on the control twigs without fruit bags were exploited by the beginning of January. The fruit removal rate and seed dispersal distance were positively correlated with an increase in wintering bird species and their abundance. Intact bird‐dispersed seeds of E. umbellata were trapped within a 400‐m range and damaged seeds were limited to traps set within 50 m from the seed source. Frugivore behavior, such as feces excretion on rocks near water drinking sites and perching on surrounding woodland, greatly influenced the spatial and temporal dispersal pattern of the seed rain. In the present study, the avian frugivores showed upstream seed dispersal; thus, in years with stochastic autumnal floods, secondary dispersal via hydrochory downstream may be facilitated. The intensive seed dispersal in E. umbellata indicates that the present distribution of parent trees in the restricted elevation range of the gravel bars is the result of survival through disturbance, rather than seed dispersal limitation.     

Factors Limiting Post‐logging Seedling Regeneration by Big‐leaf Mahogany (Swietenia macrophylla) in Southeastern Amazonia,Brazil, and Implications for Sustainable Management1

Post‐logging seedling regeneration density by big‐leaf mahogany (Swietenia macrophylla), a nonpioneer light‐demanding timber species, is generally reported to be low to nonexistent. To investigate factors limiting seedling density following logging within the study region, we quantified seed production rates, germinability, dispersal patterns, and seed fates on the forest floor through germination and the first seedling growing season in southeastern Amazonia, Brazil. Fruit production rates were low by three logged and one unlogged populations compared to reports from other regions. Commercial‐sized trees (>60 cm diameter) were more fecund than noncommercial trees (30–60 cm diameter) at two sites, averaging 14.5 vs. 3.9 fruits/tree/year, respectively, at Marajoara, a logged site, over 8 yr. Fruit capsules contained an average of 60.3 seeds/fruit, 70 percent of which appeared viable by visual inspection. Sixty‐seven to 72 percent of apparently viable seeds germinated in nursery beds 2.5 mo after the dispersal period, when wet season rains began. Dry season winds blew most seeds west‐northwest of parent trees, with median dispersal distances of 28 and 9 m on west and east sides of parent trees, respectively. Nearly 100 percent of seeds fell within an area of 0.91 ha. On the forest floor beneath closed canopies, mammals, invertebrates, and fungal pathogens killed 40 percent of apparently viable seeds, while 36 percent germinated. Nine months after seedling establishment—midway through the first logging season following seed dispersal—14 percent of outplanted seeds survived as seedlings, representing 5.8 seeds/fruit. We conclude that seedlings are likely to survive in logging gaps at appreciable densities only in rare cases where previous year fruit production rates by logged trees were high (4–12.5% of commercial‐sized trees/year at Marajoara) and where tree crowns were felled in west or northwest directions.     

Standard yet unusual mechanisms of long-distance dispersal: seed dispersal of Corymbia torelliana by bees

Mellitochory, seed dispersal by bees, has been implicated in long-distance dispersal of the tropical rain forest tree, Corymbia torelliana (Myrtaceae). We examined natural and introduced populations of C. torelliana for 4 years to determine the species of bees that disperse seeds, and the extent and distance of seed dispersal. The mechanism of seed dispersal by bees was also investigated, including fruit traits that promote dispersal, foraging behaviour of bees at fruits, and the fate of seeds. The fruit structure of C. torelliana , with seed presented in a resin reward, is a unique trait that promotes seed dispersal by bees and often results in long-distance dispersal. We discovered that a guild of four species of stingless bees, Trigona carbonaria, T. clypearis, T. sapiens , and T. hockingsi, dispersed seeds of C. torelliana in its natural range. More than half of the nests found within 250 m of fruiting trees had evidence of seed transport. Seeds were transported minimum distances of 20–220 m by bees. Approximately 88% of seeds were dispersed by gravity but almost all fruits retained one or two seeds embedded in resin for bee dispersal. Bee foraging for resin peaked immediately after fruit opening and corresponded to a peak of seed dispersal at the hive. There were strong correlations between numbers of seeds brought in and taken out of each hive by bees ( r =  0.753–0.992, P  < 0.05), and germination rates were 95 ± 5%. These results showed that bee-transported seeds were effectively dispersed outside of the hive soon after release from fruits. Seed dispersal by bees is a non-standard dispersal mechanism for C. torelliana, as most seeds are dispersed by gravity before bees can enter fruits. However, many C. torelliana seeds are dispersed by bees, since seeds are retained in almost all fruits, and all of these are dispersed by bees.     

Seed dispersal and predation in the endemic Atlantic rainforest palm <Emphasis Type="Italic">Astrocaryum aculeatissimum</Emphasis> across a gradient of seed disperser abundance

Tropical forests have been subject to intense hunting of medium and large frugivores that are important in dispersing large-seeded species. It has been hypothesized that in areas with extinction or low abundance of medium and large-bodied animals the density of small rodents may increase. Therefore, this increment in the density of small rodents may compensate for the absence or low abundance of medium and large frugivores on seed removal and seed dispersal. Here, we fill up this gap in the literature by determining if seed removal, seed dispersal, and seed predation by small rodents (spiny rats, Trinomys inheringi and squirrels, Sciurus ingrami) are maintained in defaunated areas. We accessed seed removal, seed dispersal, seed predation, and seedling recruitment of an endemic Atlantic rainforest palm, Astrocaryum aculeatissimum, in a gradient of abundance of agoutis. We found that seed removal, scatter hoarding, and seed predation increase with the abundance of agoutis. In contrast, the proportion of dispersed but non-cached seeds decreased with the abundance of agoutis. We did not find any effect of the abundance of agoutis on seed dispersal distance, but we did find a positive trend on the density of seedlings. We concluded that small rodents do not compensate the low abundance of agoutis on seed removal, scatter hoarding, and seed predation of this palm tree. Moreover, areas in which agoutis are already extinct did not present any seed removal or scatter hoarding, not even by small rodents. This study emphasizes both the importance of agoutis in dispersing seeds of A. aculeatissimum and the collapse in seed dispersal of this palm in areas where agoutis are already extinct.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

Rapid Assessment of Seed Removal and Frugivore Activity as a Tool for Monitoring the Health Status of Tropical Forests

Ecological processes in tropical forests are being affected at unprecedented rates by human activities. Yet, the continuity of ecological functions like seed dispersal is crucial for forest regeneration. It thus becomes increasingly urgent to be able to rapidly assess the health status of these processes in order to take appropriate management measures. We tested a method to rapidly evaluate seed removal rates on two animal‐dispersed tree species, Virola kwatae and V. michelii (Myristicaceae), at three sites in French Guiana with increasing levels of anthropogenic disturbance. We counted fallen fruits, fruit valves, and seeds of each focal fruiting tree in a single 1 m² quadrat, and calculated two indices: the proportion of seeds removed and the proportion of fruits opened by mammals. They both provide an indirect and rapid assessment of frugivore activity. Our results showed a significant decrease in the proportion of removed seeds (16%) and fruits opened (19%) at the most impacted site in comparison with the other two sites (79% for seeds, 60% and 35% for fruits). This testifies to an increased impoverishment of the primate and toucan communities at the disturbed sites. This standardized protocol provides fast information about the health status of the community of seed dispersers and predators and of their seed removal services. It is time‐ and cost‐effective and is not species‐specific, allowing comparisons among sites or over time. We suggest using it with the pantropical Myristicaceae and any other capsule‐producing family to rapidly assess the health status of seed removal processes across the tropics.     

Seed dispersal by animals: exact identification of source trees with endocarp DNA microsatellites

A long-standing challenge in studies of seed dispersal by animal frugivores has been the characterization of the spatial relationships between dispersed seeds and the maternal plants, i.e. the seed shadow. The difficulties to track unambiguously the origin of frugivore-dispersed seeds in natural communities has been considered an unavoidable limitation of the research field and precluded a robust analysis of the direct consequences of zoochory. Here we report that the multilocus genotype at simple sequence repeat (SSR; microsatellite) loci of the woody endocarp, a tissue of maternal origin, provides an unequivocal genetic fingerprint of the source tree. By comparing the endocarp genotype against the complete set of genotypes of reproductive trees in the population, we could unambiguously identify the source tree for 82.1% of the seeds collected in seed traps and hypothesize that the remaining 17.9% of sampled seeds come from other populations. Identification of the source tree for Prunus mahaleb seeds dispersed by frugivores revealed a marked heterogeneity in the genetic composition of the seed rain in different microhabitats, with a range of 1-5 distinct maternal trees contributing seeds to a particular landscape patch. Within-population dispersal distances ranged between 0 and 316 m, with up to 62% of the seeds delivered within 15 m of the source trees. Long distance dispersal events, detected by the exclusion of all reproductive trees in the population, accounted for up to 17.9% of the seeds sampled. Our results indicate strong distance limitation of seed delivery combined with infrequent long-distance dispersal events, extreme heterogeneity in the landscape pattern of genetic makeup, and a marked mosaic of multiple parentage for the seeds delivered to a particular patch.     

Microsatellite analysis of seed dispersal and parentage of saplings in bur oak, Quercus macrocarpa

Microsatellite analysis was used to examine parentage and spatial distributions of 62 adult bur oaks Quercus macrocarpa, and 100 saplings in a single stand. Using genotypes scored by PCR products at four microsatellite loci, we determined that 94 saplings matched at least one parent in the stand. Saplings often occur as dense clusters of half-sibs around the presumed maternal parent, and only four adults were seed parents to a large proportion of the saplings sampled. A stump apparently was the seed parent of the largest cluster of half-sibs, which occupied a sizeable light gap opened up by the death of their maternal tree. Approximately half of the saplings appeared to have grown from seeds that had not been removed after falling from the tree, and half from seeds that were dispersed beyond the crown of their maternal parent. Long-distance seed dispersal may be more common than has been previously reported. Extremely high levels of long-distance pollination were indicated, and pollen donors within the stand were generally distributed randomly around maternal trees. More than half of the saplings had paternal parents outside of the stand. This study demonstrates the utility of microsatellite analysis for studying mating systems, seed dispersal and seedling establishment in natural plant populations.     

Evidence for secondary seed dispersal by rodents in Panama

Summary The data presented show thatVirola nobilis (Myristicaceae), a bird/mammal-dispersed tree species in Panama, may also be dispersed by a terrestrial rodent, the agouti (Dasyprocta punctata). Using a thread-marking method, we observed that agoutis scatterhoardedV. nobilis seeds that they found both singly or in clumps. Seed removal and seed burial rates were strongly affected by features of forest habitats, such asV. nobilis tree richness (rich vs poor) and/or forest age (old vs young), but not by seed dispersal treatment (scattered vs clumped). Predation (mostly post-dispersal) of unburied seeds by weevils was independent of habitat and dispersal treatment. Seeds artificially buried in aVirola-rich area were more likely to escape predation and become established than unburied seeds under natural conditions. The food reward for agoutis is in the germinating seedlings. The seed dispersal syndrome ofV. nobilis involves long- and short-distance dispersers which both appear important for tree recruitment.     

Combining distances of ballistic and myrmecochorous seed dispersal in Adriana quadripartita (Euphorbiaceae)

The separate contributions of different vectors to net seed dispersal curves of diplochorous systems have rarely been characterised. In Australia, myrmecochory is a common seed dispersal syndrome and in the majority of such systems, seeds are initially dispersed ballistically. We measured ballistic and myrmecochorous seed dispersal distances in relation to canopies of Adriana quadripartita (Euphorbiaceae) and used a simulation model to estimate the net dispersal curve. We also compared seed removal rates and ant abundances under, and outside, plant canopies to examine how foraging patterns by ants may affect net dispersal.Overall ant abundance did not show a significant numerical response to seedfall; however, the abundance of the main seed dispersing ant, Rhytidoponera ‘metallica’ did. Despite this, seed removal rates did not differ significantly between canopy and open locations. Rhytidoponera ‘metallica’ account for 93% of observed seed dispersal events. On average, the ants dispersed seeds 1.54 m and in doing so, moved seed a mean radial distance of 0.76 m away from canopy edges. This contribution to net dispersal distance by ants is considerable since ballistic dispersal moved seeds a median distance of 7.5 cm. Our simulation model indicated that the combination of ballistic and ant seed dispersal is expected to result in seeds being transported a median net radial dispersal distance of 1.05 m from the canopy edge.Thus in this system, an important function of diplochory may simply be to move a higher proportion of seeds from under the canopy of parent plants than is possible by ballistic dispersal alone. This ‘dispersal-for-distance’ may result in reduced parent–offspring competition or may increase the probability that seeds reach rare safe sites for germination and recruitment.     

Mammal Abundances and Seed Traits Control the Seed Dispersal and Predation Roles of Terrestrial Mammals in a Costa Rican Forest

In Neotropical forests, mammals act as seed dispersers and predators. To prevent seed predation and promote dispersal, seeds exhibit physical or chemical defenses. Collared peccaries (Pecari tajacu) cannot eat some hard seeds, but can digest chemically defended seeds. Central American agoutis (Dasyprocta punctata) gnaw through hard‐walled seeds, but cannot consume chemically defended seeds. The objectives of this study were to determine relative peccary and agouti abundances within a lowland forest in Costa Rica and to assess how these two mammals affect the survival of large seeds that have no defenses (Iriartea deltoidea, Socratea exorrhiza), physical defenses (Astrocaryum alatum, Dipteryx panamensis), or chemical defenses (Mucuna holtonii) against seed predators. Mammal abundances were determined over 3 yrs from open‐access motion‐detecting camera trap photos. Using semi‐permeable mammal exclosures and thread‐marked seeds, predation and dispersal by mammals for each seed species were quantified. Abundances of peccaries were up to six times higher than those of agoutis over 3 yrs, but neither peccary nor agouti abundances differed across years. Seeds of A. alatum were predominantly dispersed by peccaries, which did not eat A. alatum seeds, whereas non‐defended and chemically defended seeds suffered high levels of predation, mostly by peccaries. Agoutis did not eat M. holtonii seeds. Peccaries and agoutis did not differ in the distances they dispersed seeds. This study shows that seed fates are contingent upon many factors such as seed defenses, frugivore–granivore abundances, and seed‐handling capabilities. Mammal–seed interactions are complex; the outcomes of these interactions depend on the inherent characteristics of seeds and their potential dispersers.     

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.     

Forest Fragmentation Reduces Seed Dispersal of Duckeodendron cestroides, a Central Amazon Endemic

Fragmentation that alters mutualistic relationships between plants and frugivorous animals may reduce the seed dispersal of trees. We examined the effects of forest fragmentation on the distributions of seeds and seedlings of a Central Amazon endemic tree, Duckeodendron cestroides . In the dry seasons of 2002–2004, seeds and first-year seedlings were counted within wedge-shaped transects centered around Duckeodendron adults in fragments and nearby continuous forests at the Biological Dynamics of Forest Fragmentation Project. Analyses showed that fragmentation reduced seed dispersal quantity and quality. The percent and distance of dispersed seeds were both twice as great in continuous forest as in fragments. The distances of each tree's five furthest dispersed seeds were three times greater in continuous forest than fragments. Over the 3-yr study, 20 times more seeds were dispersed more than 10 m from parent crowns in continuous forest than fragments. A regression analysis showed more dispersed seeds at all distances in continuous forest than fragments. Dispersal differences were strong in 2002 and 2004, years of large fruit crops, but weak or absent in 2003, when fruit production was low. As distance from parent crowns increased, the number of seedlings declined more rapidly in fragments than continuous forest. Distance-dependent mortality between the seed and seedling stages appeared to be more important in continuous forest than fragments. This research provides ample, indirect evidence demonstrating that forest fragmentation can result in the breakdown of a seed dispersal mutualism, potentially jeopardizing the reproduction of a rare, tropical tree.     

Seed dispersal by ants in the semi-arid Caatinga of North-East Brazil

BACKGROUND AND AIMS: Myrmecochory is a conspicuous feature of several sclerophyll ecosystems around the world but it has received little attention in the semi-arid areas of South America. This study addresses the importance of seed dispersal by ants in a 2500-km(2) area of the Caatinga ecosystem (north-east Brazil) and investigates ant-derived benefits to the plant through myrmecochory. METHODS: Seed manipulation and dispersal by ants was investigated during a 3-year period in the Xingó region. Both plant and ant assemblages involved in seed dispersal were described and ant behaviour was characterized. True myrmecochorous seeds of seven Euphorbiaceae species (i.e. elaiosome-bearing seeds) were used in experiments designed to: (1) quantify the rates of seed cleaning/removal and the influence of both seed size and elaiosome presence on seed removal; (2) identify the fate of seeds dispersed by ants; and (3) document the benefits of seed dispersal by ants in terms of seed germination and seedling growth. KEY RESULTS: Seed dispersal by ants involved one-quarter of the woody flora inhabiting the Xingó region, but true myrmecochory was restricted to 12.8 % of the woody plant species. Myrmecochorous seeds manipulated by ants faced high levels of seed removal (38-84 %) and 83 % of removed seeds were discarded on ant nests. Moreover, seed removal positively correlated with the presence of elaiosome, and elaiosome removal increased germination success by at least 30 %. Finally, some Euphorbiaceae species presented both increased germination and seedling growth on ant-nest soils. CONCLUSIONS: Myrmecochory is a relevant seed dispersal mode in the Caatinga ecosystem, and is particularly frequent among Euphorbiaceae trees and shrubs. The fact that seeds reach micro-sites suitable for establishment (ant nests) supports the directed dispersal hypothesis as a possible force favouring myrmecochory in this ecosystem. Ecosystems with a high frequency of myrmecochorous plants appear not to be restricted to regions of nutrient-impoverished soil or to fire-prone regions.     

The intermediate dispersal hypothesis: seed dispersal is maximized in areas with intermediate usage by hoarders

Seed dispersal and predation are paramount for tropical plant diversity. When encountered by scatter-hoarding frugivores, seeds can be either eaten, dispersed or ignored. But even after dispersal, seed caches are still subjected to predation. Many factors are known to influence these dynamics; however, how frequently hoarders use certain patches has seldom been related to cache predation rates. We used the interaction between agoutis (Dasyprocta leporina), a scatter-hoarding rodent, and Joannesia princeps, a tropical tree, as a model to investigate how the number of visits by hoarders in certain areas influences cache predation and seed fate. Camera-traps were used for 30 days in twenty different locations in Tijuca National Park to assess number of visits by agoutis. Thereafter, we placed seed piles on the same areas and determined their fate using the spool-and-line method to track seeds for over one hundred days. We found a non-linear relationship between how often an area is used by hoarders and the final proportion of dispersed seeds. At areas with a low number of visits, proportion of dispersed seeds was low due to low removal. As frequency of visits by hoarders increased, seed removal and the number of dispersal events increased but so did cache predation. Thus, in areas intensively used by hoarders, high cache predation resulted in a low number of dispersed seeds that remained alive in caches. As a result, dispersal was maximized in areas with intermediate use by scatter-hoarders, where there was a balance between primary seed dispersal and cache predation.

     

Seed dispersal by Galápagos tortoises

Aim Large‐bodied vertebrates often have a dramatic role in ecosystem function through herbivory, trampling, seed dispersal and nutrient cycling. The iconic Galápagos tortoises (Chelonoidis nigra) are the largest extant terrestrial ectotherms, yet their ecology is poorly known. Large body size should confer a generalist diet, benign digestive processes and long‐distance ranging ability, rendering giant tortoises adept seed dispersers. We sought to determine the extent of seed dispersal by Galápagos tortoises and their impact on seed germination for selected species, and to assess potential impacts of tortoise dispersal on the vegetation dynamics of the Galápagos. Location Galápagos, Ecuador. Methods To determine the number of seeds dispersed we identified and counted intact seeds from 120 fresh dung piles in both agricultural and national park land. To estimate the distance over which tortoises move seeds we used estimated digesta retention times from captive tortoises as a proxy for retention times of wild tortoises and tortoise movement data obtained from GPS telemetry. We conducted germination trials for five plant species to determine whether tortoise processing influenced germination success. Results In our dung sample, we found intact seeds from > 45 plant species, of which 11 were from introduced species. Tortoises defecated, on average, 464 (SE 95) seeds and 2.8 (SE 0.2) species per dung pile. Seed numbers were dominated by introduced species, particularly in agricultural land. Tortoises frequently moved seeds over long distances; during mean digesta retention times (12 days) tortoises moved an average of 394 m (SE 34) and a maximum of 4355 m over the longest recorded retention time (28 days). We did not find evidence that tortoise ingestion or the presence of dung influenced seed germination success. Main conclusions Galápagos tortoises are prodigious seed dispersers, regularly moving large quantities of seeds over long distances. This may confer important advantages to tortoise‐dispersed species, including transport of seeds away from the parent plants into sites favourable for germination. More extensive research is needed to quantify germination success, recruitment to adulthood and demography of plants under natural conditions, with and without tortoise dispersal, to determine the seed dispersal effectiveness of Galápagos tortoises.