Effect of rodents on seed fate of five hornbill-dispersed tree species in a tropical forest in north-east India

Hornbills are important dispersers of a wide range of tree species. Many of these species bear fruits with large, lipid-rich seeds that could attract terrestrial rodents. Rodents have multiple effects on seed fates, many of which remain poorly understood in the Palaeotropics. The role of terrestrial rodents was investigated by tracking seed fate of five horn bill-dispersed tree species in a tropical forest in north-cast India. Seeds were marked inside and outside of exclosures below 6-12 parent fruiting trees (undispersed seed rain) and six hornbill nest trees (a post-dispersal site). Rodent visitors and seed removal ere monitored using camera traps. Our findings suggest that several rodent species. especially two species of porcupine were major on-site seed predators. Scatter-hoarding was rare (1.4%). Seeds at hornbill nest trees had lower survival compared with parent fruiting trees, indicating that clumped dispersal by hornbills may not necessarily improve seed survival. Seed survival in the presence and absence of rodents varied with tree species. Some species (e.g. Polyalthia simiarum) showed no difference, others (e.g. Dysoxylum binectariferum) experienced up to a 64%. decrease in survival in the presence of rodents. The differing magnitude of seed predation by rodents can have significant consequences at the seed establishment stage.

Spatio-temporal variation in seed production in three Euphorbia species and the role of animals on seed fate

Comparative studies on the reproductive biology of co-occurring related plant species have provided valuable information for the interpretation of ecological and evolutionary phenomena, with direct application in conservation management of plant populations. The aims of this thesis were to identify the causes of pre-dispersal reproductive losses in three Euphorbia species (the Mediterranean E. characias and the narrow endemics E. pedroi and E. welwitschii) and evaluate the variation of their effects in time, space and between individuals and species. Furthermore, we intended to study elaiosomes’ fatty acid profiles for the three Euphorbia and assess the role played by the elaiosome in ant attraction. Finally, we aimed to identify the major seed dispersal agents for each Euphorbia species in each site and study differences in short term seed fate due to differences in ant behaviour. The results indicated that intact seed production differed significantly between the three Euphorbia, mostly due to differences in cyathia production. Losses to pre-dispersal seed predators were proportionately larger for the endemic species which also suffered higher losses resulting in flower, fruit (in E. welwitschii) and seed abortion (in E. pedroi). The elaiosomes of E. pedroi are poor in fatty acids and for this reason seeds of this species were removed in lower proportion by mutualistic dispersers than those of their congeners, being more prone to seed predation. Two larger ant species – Aphaenogaster senilis and Formica subrufa – were responsible for a larger percentage of removals with seeds being transported at larger distances and being discarded in the vicinity of their nests following elaiosome removal. Our results highlight the role of insect-plant interactions as major determinants of seed survival for the three study plants and call for the need to include more information on insect-plant interactions in plant conservation programmes.

Seed fate and seedling dynamics after masting in two African rain forest trees

How the effects of biotic factors are moderated by abiotic factors, and their consequences for species interactions, is generally understudied in ecology. A key abiotic feature of forests is regular canopy disturbances that create temporary patches, or “gaps,” of above-average light availability. Co-occurring in lowland primary forest of Korup National Park (Cameroon), Microberlinia bisulcata and Tetraberlinia bifoliolata are locally dominant, ectomycorrhizal trees whose seeds share predator guilds in masting years. Here, we experimentally tested the impact of small mammal predators upon seedling abundance, growth, and survivorship. In 2007, we added a fixed density of seeds of each species to exclosures at 48 gap–understory locations across 82.5 ha within a large Microberlinia grove, and at 15 locations outside it. For both species, small mammals removed more seeds in gaps than in understory, whereas this was reversed for seeds killed by invertebrates. Nonetheless, Microberlinia lost twice as many seeds to small mammals, and more to invertebrates in exclosures, than Tetraberlinia, which was more prone to a pathogenic white fungus. After six weeks, both species had greater seedling establishment in gaps than understory, and in exclosures outside compared to exclosures inside the grove. In the subsequent two-year period, seedling growth and survivorship peaked in exclosures in gaps, but Microberlinia had more seedlings' stems clipped by animals than Tetraberlinia, and more than twice the percentage of leaf area damaged. Whereas Microberlinia seedling performance in gaps was inferior to Tetraberlinia inside the grove, outside it Microberlinia had reduced leaf damage, grew taller, and had many more leaves than Tetraberlinia. No evidence was found for “apparent mutualism” in the understory as seedling establishment of both species increased away from (>25 m) large stems of either species, pointing to “apparent competition” instead. In gaps, Microberlinia seedling establishment was lower near Tetraberlinia than conspecific adults because of context-dependent small mammal satiation. Stage-matrix analysis suggested that protecting Microberlinia from small mammals could increase its population growth rate by 0.06. In the light of prior research we conclude that small mammals and canopy gaps play an important role in promoting species coexistence in this forest, and that their strong interaction contributes to Microberlinia's currently very poor regeneration.

The role of seed mass on the caching decision by agoutis, Dasyprocta leporina (Rodentia: Agoutidae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Differential seed germination of a keystone palm (Euterpe edulis) dispersed by avian frugivores

The effectiveness of seed dispersal by vertebrates has been analysed by examining both quantitative and qualitative components (Jordano & Schupp 2000, Schupp et al. 2010). While the quantitative component is relatively easily assessed in the field (e.g. visitation rate, number of fruits eaten per visit), the qualitative component (e.g. fate of dispersed seeds, seed treatment in the digestive system of the disperser) is rarely studied under natural conditions, because it is difficult to measure the effects on seeds once ingested by the dispersers (Cortes et al. 2009). © Cambridge University Press 2012.

Differential seed germination of a keystone palm (Euterpe edulis) dispersed by avian frugivores

FAPESP (BIOTA Program) [2007/03392-6, 2010/04927-3]

Flying vs. climbing: Factors controlling arboreal seed removal in oak-beech forests

Nuts are heavy and nutritious seeds that need animals to be successfully dispersed. Most studies address nut removal by a single animal species once seeds fall onto the ground. However, nuts are also accessible before the seed drop and usually to a wide guild of seed foragers. This study examines the factorscontrollingarborealseedremoval in oak–beechforests within the whole guild of nut foragers. We found that seed-dispersing rodents (Apodemus sylvaticus) were the main acorn removers in the oaks (up to 3.75 m height), with a rapid seed encounter and a high removal rate. However, rodents did not climb the beech trees, probably due to their smoother bark in comparison to oak bark and/or the lower nutritional value of beechnuts with regard to acorns. Jays (Garrulus glandarius) were more abundant in oak stands (both dense and scattered) and clearly preferred acorns to beechnuts whereas nuthatches (Sitta europaea) were more abundant in beech stands and preferred beechnuts to acorns. Non-storing birds such as great tits (Parus major) also removed acorns and beechnuts, especially in the stands where oaks are dominant. Jays and rodents preferred sound seeds over insect-infested seeds but such a preference was not found for nuthatches. This study highlights that pure beech stands showed a reduced guild of arboreal nut foragers in comparison to oak stands. This different guild could probably affect the spatial patterns of seed dispersal, with a proportionally higher number of long dispersal events for acorns (mostly jay-dispersed) than for beechnuts (mostly nuthatch-dispersed). Long-distance dispersal of beechnuts (by jays) is determined by the presence of other preferred species (oaks) and their frequency of non-mast years. Seed location in different habitats strongly determines the contribution of different arboreal removers (including climbing rodents) and their removal speed, leading to a differential seed fate that will eventually affect tree regeneration. As nuthatches are sedentary birds, it is important to maintain old and dead trees where they can breed (crevices), forage (arthropods) and store seeds in order to favor beechnut dispersal and gene flow. By maintaining or favoring oak trees within beech stands we will ensure a wider guild of arboreal nut dispersers.

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.

Studies on oxamyl : analytical method development and investigation of fate in peach seedlings and corn seeds

A high performance liquid chromatographic method employing two columns connected in series and separated~y·a.switching valve has been developed for the analysis of the insecticide/ nematicide oxamyl (methyl-N' ,N'-dimethyl-N-[(methylcarbamoyl) oxy]-l-thiooxarnimidate) and two of its metabolites. A variation of this method involving two reverse phase columns was employed to monitor the persistence and translocation of oxamyl in treated peach seedlings. It was possible to simultaneously analyse for oxamyl and its corresponding oxime (methyl-N',N'-dimethyl-N-hydroxy-l-thiooxamimidate}, a major metabolite of oxamyl in plants, without prior cleanup of the samples. The method allowed detection of 0.058 pg oxamyl and 0.035 p.g oxime. On treated peach leaves oxamyl was found to dissipate rapidly during the first two-week period, followed by a period of slow decomposition. Movement of oxamyl or its oxime did not occur in detectable quantities to untreated leaves or to the root or soil. A second variation of the method which employed a size exclusion column as·the first column and a reverse phase column as the second was used to monitor the degradation of oxamyl in treated, planted corn seeds and was suitable for simultaneous analysis of oxamyl, its oxime and dimethylcyanoformamide (DMCF), a metabolite of oxamyl. The method allowed detection of 0.02 pg oxamyl, 0.02 p.g oxime and 0.005 pg DMCF. Oxamyl was found to persist for a period of 5 - 6 weeks, which is long enough to permit oxamyl seedtreatment to be considered as a potential means of protecting young corn plants from nematode attack. Decomposition was found to be more rapid in unsterilized soil than in sterililized soil. DMCF was found to have a nematostatic effect at high concentrations ( 2,OOOpprn), but at lower concentrations no effect on nematode mobility was observed. Oxamyl, on the other hand, was found to reduce the mobility of nematodes at concentrations down to 4 ppm.

Seed dispersal and spatial distribution of Attalea geraensis (Arecaceae) in two remnants of Cerrado in Southeastern Brazil

The seed dispersal system of Attalea geraensis (Arecaceae), an acaulescent palm, was investigated during one year in two Cerrado fragments in the state of São Paulo, southeastern Brazil. A. geraensis had inflorescences and infrutescences throughout the year. Two scatter-hoarding rodents (the spiny rat, Clyomys bishopi and agoutis, Dasyptocta azarae) were identified as seed predators/dispersers, able to move seeds up to 30 in from the palms, although most of the fruits (57.5%) were dispersed less than 2 in. The removal rates were high and after 20 days, 97.2% of the fruits were removed. Fruit fate was not related to fruit mass, length and diameter. The application of Morisita's index showed a more clumped distribution of adults in the smaller fragment, probably because of the absence of agoutis. Higher seed removal by rodents in the large Cerrado remnant may decrease seed predation by beetles. (C) 2007 Elsevier Masson SAS. All rights reserved.

Seasonal variation in the fate of seeds under contrasting logging regimes

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Seed dispersal and reproduction patterns among Everglades plants

In this study three aspects of sexual reproduction in Everglades plants were examined to more clearly understand seed dispersal and the allocation of resources to sexual reproduction—spatial dispersal process, temporal dispersal of seeds (seedbank), and germination patterns in the dominant species, sawgrass (Cladium jamaicense). Community assembly rules for fruit dispersal were deduced by analysis of functional traits associated with this process. Seedbank ecology was investigated by monitoring emergence of germinants from sawgrass soil samples held under varying water depths to determine the fate of dispersed seeds. Fine-scale study of sawgrass fruits yielded information on contributions to variation in sexually produced propagules in this species, which primarily reproduces vegetatively. It was hypothesized that Everglades plants possess a set of functional traits that enhance diaspore dispersal. To test this, 14 traits were evaluated among 51 species by factor analysis. The factorial plot of this analysis generated groups of related traits, with four suites of traits forming dispersal syndromes. Hydrochory traits were categorized by buoyancy and appendages enhancing buoyancy. Anemochory traits were categorized by diaspore size and appendages enhancing air movement. Epizoochory traits were categorized by diaspore size, buoyancy, and appendages allowing for attachment. Endozoochory traits were categorized by diaspore size, buoyancy, and appendages aiding diaspore presentation. These patterns/trends of functional trait organization also represent dispersal community assembly rules. Seeds dispersed by hydrochory were hypothesized to be caught most often in the edge of the north side of sawgrass patches. Patterns of germination and dispersal mode of all hydrochorous macrophytes with propagules in the seedbank were elucidated by germination analysis from 90 soil samples collected from 10 sawgrass patches. Mean site seed density was 486 seeds/m2 from 13 species. Most seeds collected at the north side of patches and significantly in the outer one meter of the patch edge (p = 0.013). Sawgrass seed germination was hypothesized to vary by site, among individual plants, and within different locations of a plant’s infructescence. An analysis of sawgrass fruits with nested ANOVAs found that collection site and interaction of site x individual plant significantly affect germination ability, seed viability, and fruit size (p ≤ 0.050). Fruit location within a plant’s infructescence did not significantly affect germination. As for allocation of resources to sexual reproduction, only 17.9% of sawgrass seeds germinated and only 4.8% of ungerminated seeds with fleshy endosperm were presumed viable, but dormant. Collectively, only 22% of all sawgrass seeds produced were viable.

Frugivory and Seed Dispersal by Crocodilians: An Overlooked Form of Saurochory?

Saurochory (seed dispersal by reptiles) among crocodilians has largely been ignored, probably because these reptiles are generally assumed to be obligate carnivores incapable of digesting vegetable proteins and polysaccharides. Herein we review the literature on crocodilian diet, foraging ecology, digestive physiology and movement patterns, and provide additional empirical data from recent dietary studies of Alligator mississippiensis. We found evidence of frugivory in 13 of 18 (72.2%) species for which dietary information was available, indicating this behavior is widespread among the Crocodylia. Thirty-four families and 46 genera of plants were consumed by crocodilians. Fruit types consumed by crocodilians varied widely; over half (52.1%) were fleshy fruits. Some fruits are consumed as gastroliths or ingested incidental to prey capture; however, there is little doubt that on occasion, fruit is deliberately consumed, often in large quantities. Sensory cues involved in crocodilian frugivory are poorly understood, although airborne and waterborne cues as well as surface disturbances seem important. Crocodilians likely accrue nutritional benefits from frugivory and there are no a priori reasons to assume otherwise. Ingested seeds are regurgitated, retained in the stomach for indefinite and often lengthy periods, or passed through the digestive tract and excreted in feces. Chemical and mechanical scarification of seeds probably occurs in the stomach, but what effects these processes have on seed viability remain unknown. Because crocodilians have large territories and undertake lengthy movements, seeds are likely transported well beyond the parent plant before being voided. Little is known about the ultimate fate of seeds ingested by crocodilians; however, deposition sites could prove suitable for seed germination. Although there is no evidence for a crocodilian-specific dispersal syndrome similar to that described for other reptiles, our review strongly suggests that crocodilians function as effective agents of seed dispersal. Crocodilian saurochory offers a fertile ground for future research.

Seed Dispersal and Reproduction Patterns Among Everglades Plants

In this study three aspects of sexual reproduction in Everglades plants were examined to more clearly understand seed dispersal and the allocation of resources to sexual reproduction— spatial dispersal process, temporal dispersal of seeds (seedbank), and germination patterns in the dominant species, sawgrass (Cladium jamaicense). Community assembly rules for fruit dispersal were deduced by analysis of functional traits associated with this process. Seedbank ecology was investigated by monitoring emergence of germinants from sawgrass soil samples held under varying water depths to determine the fate of dispersed seeds. Fine-scale study of sawgrass fruits yielded information on contributions to variation in sexually produced propagules in this species, which primarily reproduces vegetatively. It was hypothesized that Everglades plants possess a set of functional traits that enhance diaspore dispersal. To test this, 14 traits were evaluated among 51 species by factor analysis. The factorial plot of this analysis generated groups of related traits, with four suites of traits forming dispersal syndromes. Hydrochory traits were categorized by buoyancy and appendages enhancing buoyancy. Anemochory traits were categorized by diaspore size and appendages enhancing air movement. Epizoochory traits were categorized by diaspore size, buoyancy, and appendages allowing for attachment. Endozoochory traits were categorized by diaspore size, buoyancy, and appendages aiding diaspore presentation. These patterns/trends of functional trait organization also represent dispersal community assembly rules. Seeds dispersed by hydrochory were hypothesized to be caught most often in the edge of the north side of sawgrass patches. Patterns of germination and dispersal mode of all hydrochorous macrophytes with propagules in the seedbank were elucidated by germination analysis from 90 soil samples collected from 10 sawgrass patches. Mean site seed density was 486 seeds/m2 from 13 species. Most seeds collected at the north side of patches and significantly in the outer one meter of the patch edge (p = 0.013). Sawgrass seed germination was hypothesized to vary by site, among individual plants, and within different locations of a plant’s infructescence. An analysis of sawgrass fruits with nested ANOVAs found that collection site and interaction of site x individual plant significantly affect germination ability, seed viability, and fruit size (p < 0.050). Fruit location within a plant’s infructescence did not significantly affect germination. As for allocation of resources to sexual reproduction, only 17.9% of sawgrass seeds germinated and only 4.8% of ungerminated seeds with fleshy endosperm were presumed viable, but dormant. Collectively, only 22% of all sawgrass seeds produced were viable.