Primate communities vary in their level of redundancy, or overlap, in seed dispersal function, which could be due to body size, degree of frugivory or intraspecific variation, among other factors. In this study, we quantified redundancy in seed dispersal among three sympatric primate species: Lagothrix lagothricha, Alouatta seniculus and Sapajus apella in gallery forests in Meta, Colombia. We compared the median seed width dispersed and the number and species richness of large (≥5.9 mm wide) and very large (>7.5 mm wide) seeds per faecal sample. The medium-sized, highly folivorous A. seniculus mostly dispersed large seeds, the larger, highly frugivorous L. lagothricha dispersed very large and small seeds, and the smaller, partially frugivorous S. apella dispersed the smallest seeds. However, for L. lagothricha and S. apella, we did not find the expected results that adults disperse larger seeds than juveniles. Across species, there is complementarity in seed dispersal in relation to seed size, with L. lagothricha being unique in its contribution to the dispersal of very large seeds both in terms of quantity and richness.

Understanding the quality of seed dispersal effectiveness of frugivorous species can elucidate how endozoochory structures tropical forests. Large seeds, containing more resources for growth, and gut passage by frugivores, which remove seed pulp, both typically enhance the speed and probability of germination of tropical seeds. However, the interaction of seed size and gut passage has not been well studied. We assessed the role of two species of toucans (Ramphastos spp.) in seed germination of the tropical tree Eugenia uniflora, which produces seeds that vary considerably in size (3.7–14.3 mm), using 151 control and 137 regurgitated seeds in germination trials. We found that toucan regurgitation did not increase germination success, although 93.4% germinated compared to 76.8% of control seeds; however, larger seeds germinated more often at faster rates. Although only marginally significant, germination rates were 3.6× faster when seeds were both large and regurgitated by toucans, demonstrating that toucan regurgitation can disproportionally benefit larger E. uniflora seeds. As tropical forests are increasingly disturbed and fragmented by human activities, the ability of toucans to continue providing seed dispersal services to degraded habitats may be vital to the persistence of many tropical plants that contain larger seeds and depend on larger dispersers.

Large herbivores can disperse seeds over long distances through endozoochory. The Korean water deer (Hydropotes inermis argyropus), an internationally vulnerable species but locally considered a vermin, is a potential endozoochorous seed dispersal vector. In this study, feeding experiments were conducted to test the efficiency of seed dispersal through gut ingestion by the Korean water deer, its temporal pattern and the effect of gut passage on seed recovery and germination rate. Eight plant species, including species that formerly germinated from its faeces, were used to feed three Korean water deer. Once the deer had consumed all the provided seeds, their faeces were collected after 24, 48, 72 and 96 h. The collected faeces were air-dried, and the number of seeds retrieved from the faeces was counted every 24 h (0–24, 24–48, 48–72 and 72–96 h). Among the eight plant species, six species were retrieved with intact seeds. Panicum bisulcatum had the highest recovery rate of 33.7%, followed by Amaranthus mangostanus (24.5%) and Chenopodium album (14.4%). Most of the seeds were recovered within the 24–48 h time interval. Germination tests were conducted on the ingested and uningested seeds for the four species which had a sufficient recovery rate. The effects of gut passage on seed germination differed according to plant species. The germination rate substantially decreased after gut passage. The results suggest that the Korean water deer can disperse seeds, potentially over long distances albeit at a high cost of low seed recovery and germination rate.

For endozoochorous species, the quality component of seed dispersal effectiveness depends in part on the treatment seeds receive in the animal's gut. Covering a variety of taxa, diet, digestion system and body size of Prosopis flexuosa seed dispersers, we analysed differences among species in (1) mean retention time of ingested seeds, (2) recovery of viable seeds, (3) seed germination in comparison with seeds collected from trees and (4) germination of seeds after two different periods of retention in the gut. Feeding experiments were conducted with captive individuals of Dolichotis patagonum, Lycalopex gymnocercus, Rhea americana, Chelonoidis chilensis and Lama guanicoe. On the first day, we provided them with fruits containing controlled amounts of seed, and on the subsequent days, we collected faeces in order to recover seeds. We performed germination and viability tests on seeds coming from faeces and collected from trees. The results showed differences among species in the mean retention time of seeds. Chelonoidis chilensis had the longest mean retention time, but its effect on seed recovery and germination was similar to that of the other species, except for L. guanicoe, which showed the lowest seed recovery. When scarification and promotion of seed germination were considered, herbivorous mammals and tortoises (L. guanicoe, D. patagonum and C. chilensis) were the ones increasing germinability, whereas R. americana and L. gymnocercus did not significantly increase final seed germination percentage, which was similar to that for seeds collected from trees. P. flexuosa seeds receive a variety of treatments from endozoochorous dispersers, which might result in an overall fitness benefit for a plant living in unpredictable environments.

Large animals tend to disperse seeds over long distances via ingestion and defecation due to their large home range and capacity to move among different habitats for feeding. The aim of this study was to investigate the potential of endozoochorous seed dispersal by five herbivores: Ovis vignei, Capra aegagrus, Gazella subgutturosa, Cervus elaphus, Capreolus capreolus and two omnivores: Sus Scrofa and Ursus arctos in the Golestan National Park, northeast of Iran, by a greenhouse germination experiment. A total of 3107 seedlings belonging to 154 different plant taxa were germinated from 655 dung samples collected in three different habitats. Plant families that most frequently germinated were Poaceae and Brassicaceae. Urtica dioica was the most abundant germinating seed, accounting for 20% of all the seedlings recorded in our dung samples, whereas the most frequently observed species was Portulaca oleracea, which occurred in 24% of our samples. We showed that 54% of the seeds germinating were dispersed by only one of the mammals studied. Herbs and graminoids were the most frequently dispersed growth forms by the herbivores and the wild boar, whereas brown bears mostly dispersed shrubs. The seedling composition in the dung samples was strongly correlated with the local flora especially for non-selective feeders, like red deer. The differences observed in the number of plants and frequency of different growth forms dispersed among the studied mammals reflect their body size, digestive physiology, and dietary and habitat preferences. Our findings highlight the different and complementary roles of large herbivores and omnivores as long-distance seed dispersal vectors.

Madagascar is one of the most threatened biodiversity hotspots, and protection of its biodiversity is becoming increasingly urgent as deforestation of the island continues. For the long-term success of conservation efforts it is essential that key ecological processes, such as seed dispersal, are protected and restored. Therefore, the identification of ecological gaps is a vital task. For Madagascar, only little is known about plant–animal interactions, and traditional methods of ecological research are too time-consuming to provide crucial information about breakdowns in these interactions. To identify likely dispersal gaps we therefore used a theoretical approach to analyse plant–disperser interactions in Madagascar. We used data science tools to impute missing data on relevant plant traits to subsequently predict the most likely dispersal agents for each of Madagascar's endemic plant species. We found that 38% of the endemic species (N = 8,784) are endozoochorous, and among these 26–41% display a primate syndrome and 17–19% a bird syndrome (depending on the definition of syndromes). This lower percentage of endozoochorous species and higher percentage of species with a primate syndrome in Madagascar compared to other tropical areas reflects the unusual disperser guild on the island. Only five bird species but 20 lemur species are frugivorous, and 16 of those lemur species are currently threatened with extinction. The disappearance of frugivorous lemurs would significantly change the vegetation dynamics of Madagascar's ecosystems, and a high proportion of Madagascar's endemic plants would enter an extinction vortex.

Balanites maughamii appears to be an obligate elephant-dispersed species. Its fruits are strongly scented and produce a large range of volatiles, particularly aliphatic acids such as hexanoic and isovaleric acid. The seed coat can resist a compression force of >1.5 kN. Seeds removed from the seed coat, and those from compressed seeds or from elephant dung, have better germination than seeds enclosed by undamaged seed-coats, which largely failed to germinate over a 2-mo period. Mastication by elephants may be necessary to split the seed coat to allow germination and the germinant to emerge from the seed coat. We identify four traits of elephant-dispersed fruits; they are dull-coloured, presented on the ground, strongly scented and have exceptionally strong seed coats.

Apes are important long-distance dispersers of large seeds in African tropical forests. Seed size and shape are likely to affect the ease of swallowing for an animal species. If an endozoochorous seed is larger than the digestive tract of an animal, the seed cannot be swallowed, and a round seed is more difficult to swallow than an elongated seed of the same length. In order to test if such a correlation exists between the seed size and its shape, we investigated the length and width of chimpanzee-dispersed seeds at the Mahale Mountains National Park, Tanzania. Among the 14 species of seeds, longer seeds had significantly narrower relative widths, and thus, they were more ovoid. Since the chimpanzee is the largest arboreal frugivore at Mahale, their food selection might have influenced the shape of larger seeds. The chimpanzee's selective consumption of such fruits with longer, elongated seeds may have facilitated the selective dispersal of such plant species in that area.

Mimosa luisana is functionally important in the Tehuacán-Cuicatlán Valley, Mexico, since it is able to create favourable microsites for the establishment of other plant species. The endozoochory of M. luisana seeds by goats was evaluated in terms of excrement deposition pattern, seed survival and germination. The excrement deposition pattern was evaluated by collecting pellets in four plots of 25 × 2 m randomly placed in a grazing area and recording the microhabitat where pellets were found. Seed survival and germination were evaluated by feeding the goats with seeds and collecting dung pellets at 8-h intervals for 80 h. Seeds from goat pellets (treatment) and seeds collected from pods (control) were placed in a germination chamber for 24 d. Goats mainly deposited M. luisana seeds in viable sites (open areas) for growth. Mimosa luisana seeds survived the goat digestive treatment (5.91% ± 2.86%) and most of them (67% ± 25.9%) were recovered 8–32 h after ingestion. Goat gut treatment increased M. luisana final germination (47.5% ingested, 5.83% control) and shortened initial and mean time of germination. Our results indicate that goats may be an efficient disperser of M. luisana seeds.

Fossilized hackberry (Celtis) seeds were found within the shells of two Stylemys individuals excavated from Oligocene sediments from South Dakota. The presence of in situ skeletal elements indicates that the tortoises were buried without extensive disarticulation. Abiotic transport of the seeds into the carcasses is unlikely given the anatomically correct placement of both skeletal elements and seeds and the comparative settling velocities of the encasing sediment versus modern Celtis seeds. Ecological evidence from modern Celtis and Stylemys analogues suggests that tortoises are commonly seed dispersal agents. The fossils are therefore interpreted as enterolites, providing the oldest reliable evidence of tortoise frugivory.

The dispersal of plants within botanically rich grassland is an important area of study if such swards are to be maintained. The application of farmyard manure provides a possible mechanism by which seeds contained within hay can be returned/introduced to grasslands. In this study, hay, dung and manure samples taken from farms with botanically rich meadows contained very limited quantities of seeds of desirable species. Digestion by cattle, and storage within a manure heap for 6 months or longer, further reduced seed germination. Samples were characterized by an abundance of the grass, Poa trivialis, with few forbs present. Confirmation of the negative effect of the digestive processes of cattle on seed viability was achieved using an in vitro laboratory experiment. However, this experiment did show that the perennial herbs, Filipendula ulmaria and Sanguisorba officinalis, were able to survive digestion at least as well as P. trivialis. The burial of known quantities of seeds in a manure heap also showed these perennial herbs to be at least as resistant to damage as P. trivialis. The results demonstrate that, given appropriate timing of the hay cut, seeds of species with high conservation value could become incorporated into manure for subsequent dispersal. However, manure dispersal would appear to be of limited value for many species desirable from a conservation viewpoint.

Although viable seeds of Mediterranean dry-fruited shrubs are found in herbivore dung, the ecological importance of this observation is still not well understood. We analysed seed retrieval percentages, defecation time and germinability after sheep gut passage for the five most common shrub species of an area in central Spain (Retama sphaerocarpa, Cytisus scoparius, Halimium umbellatum subsp. viscosum, Cistus ladanifer and Lavandula stoechas subsp. pedunculata). Five ewes were fed seeds, and their dung was collected regularly during the following week. Seeds were hand-sorted from dung subsamples and tested for germinability. The defecated seeds were clustered in time, with a majority retrieved in the 24–40 h period, although over 1% of the seeds were retained in the gut for more than 72 h. Data suggested a possible link between seed size and retrieval, with medium-sized seeds less damaged (16–23%) than larger and smaller seeds (10–12%), although only a small number of species were studied. Germination results showed an increased percentage of germination after gut passage for H. umbellatum (x2 test, P<0.05) and a marginally significant difference for C. scoparius (P<0.1). Soft-seeded L. stoechas did not germinate after gut passage. The results indicate a potential role of herbivore endozoochory for the long-distance dispersal of dry-fruited shrubs and their potential colonization of distant sites.

It is normally assumed that Lavandula stoechas subsp. pedunculata (Miller) Samp. ex Rozeira (Labiatae) is dispersed by autochory in spite of the clear pioneer nature of the species. This paper examines the efficiency of autochorous dispersal (seed rain) and the possibility that the species is also dispersed endozoochorally by sheep. Seed rain was measured using pitfall traps and adhesive strips in summer 1999. The viable seed content in sheep dung was measured by greenhouse germination of dung collected monthly in the summers of 1998 and 1999. Two experimental tests were also conducted to ascertain whether dung-borne seeds could be established under field conditions and to evaluate the effect of added dung in the establishment of seeds taken from plants. The recorded seed rain was 2544 seeds m-2 inside the Lavandula patches, with an aggregated distribution. Autochory around the mother plant fitted a negative exponential distribution, with 90.5% concentrated in a 0–30 cm radius and a maximum distance of 1 m. Viable Lavandula seeds were found in 73% of the examined dung samples, with an average of 5.5 seeds per sample (6 g) and a high inter- and intra-annual variation. This high seed content, together with the daily sheep movements over several kilometres, make the species highly dispersible, possibly explaining its clear pioneer nature. Moreover, available data suggest that seeds in sheep dung can germinate and establish under natural conditions, and that dung addition has a positive effect on species establishment.

Effects of the elephant gut and elephant dung on seed germination and early seedling establishment/growth were investigated in Banyang-Mbo Wildlife Sanctuary using undestroyed seeds of 14 plant species sorted from fresh elephant dung between 1 June 1994 and 31 May 1995 and similar seeds extracted from fresh ripe fruits fallen on the forest floor within the same period both sown in fresh elephant dung and forest soil. Parameters measured were final germination success, germination time and seedling growth rate. Results indicated that two species, Panda oleosa and Poga oleosa, did not germinate at all after 365 d irrespective of their sources and media of planting. Germination success observed in ingested seeds was significantly different from that observed in seeds from fresh fruits. Germination success observed in elephant dung was not significantly different from that in forest soil. Mean germination time varied widely between species and treatments but was generally shorter in seeds that passed through the elephant gut than those collected from fresh ripe fruits and these differences in mean germination time were significant in 92% of the species that germinated. Growth rates of seedlings from ingested seeds were higher than those from fresh ripe fruits especially in elephant dung. Growth medium was highly significant to growth in 10 of 12 species (83%) and source was important only to two species (16%) while their combined interacting effects were significant to three of the 12 species (25%). It is concluded that ingestion of seeds by the elephant is important in the germination of some rain-forest species and the elephant dung that contains the seeds dispersed is very important in the rapid growth of the seedlings. On the basis of germination success plant species that are absolutely or exclusively dependent on elephants for dispersal/germination are absent in Banyang-Mbo Wildlife Sanctuary, while Omphalocarpum elatum and Strychnos aculeata would be on the basis of fruit morphology.