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Reproduction by perennial plants varies from being relatively constant over years to the production of massive and synchronous seed crops at irregular intervals, a reproductive strategy called mast seeding. The sources of interspecific differences in the extent of interannual variation in seed production are largely unknown. We conducted a global meta‐analysis of animal‐dispersed species to quantify how the interannual variability in seed crops produced by plants can be explained by the seed mass, dispersal mode, phylogeny, and climate. Phylogenetic analysis indicated that the interannual variations in seed production and seed mass tended to be similar in related species due to their shared evolution. The interannual variation in seed production was 1.22 times higher in synzoochorous species dispersed by scatter‐hoarders compared with endozoochorous species dispersed by frugivores. Furthermore, the production of small seeds was associated with higher interannual variation in seed production, although synzoochorous species produced larger seeds than endozoochorous species. Precipitation rather than temperature had a significant positive effect on the interannual variation in seed production. The seed mass and dispersal mode contributed more to the interannual variation in seed production than phylogeny, climate, and fruit type. Our findings support a long‐standing hypothesis that interspecific variation in the masting intensity is largely shaped by interactions between plants and animals. [ABSTRACT FROM AUTHOR]

The article explores the internal seed dispersal of Rhynchotechum discolor by a freshwater crab, Geothelphusa sakamotoana. The study suggests that the crab acts as an internal seed disperser, with intact seeds being defecated and showing a slightly higher germination rate compared to non-consumed seeds. This research sheds light on the diverse mechanisms of seed dispersal and highlights the potential role of invertebrates in plant establishment. Further studies are recommended to understand the broader impact of invertebrates on seed dispersal and plant species diversity. [Extracted from the article]

Recent evidence supports that, prior to the impact of modern humans, the temperate zone of Europe was not dominated by a closed continuous forest, but consisted of heterogeneous landscapes with grasslands, light woodlands and forests. Until the late Pleistocene, this region held a diverse community of megaherbivores that maintained the open and semi‐open vegetation through grazing and browsing. This new baseline is congruent with the fact that many European woody species are light‐demanding and fail to regenerate in the shaded interiors of temperate forests. Notably, most of these species are dispersed by frugivorous and seed‐caching animals, mostly birds. Nowadays, avian seed dispersers play a key role for the connectivity and expansion of woodland patches in fragmented anthropogenic landscapes. Indeed, avian seed dispersal within and across the deforested matrix is the norm rather than the exception. Here, I reflect on the current patterns of avian seed dispersal in the fragmented anthropogenic landscapes of Europe from a Pleistocene perspective. The aim of this exercise is to discuss linkages between past and present landscapes, seeking an historical understanding of the high spatial complementarity of avian seed dispersal in and out of the forests. Synthesis: I conclude that the spatial patterns of avian seed dispersal in anthropogenic landscapes are congruent with an ecological and evolutionary history with open and semi‐open habitats created and maintained by megaherbivores (ecological memory). By shaping the environmental conditions for millions of years, megaherbivores would have created selective pressures on animals and plants for life in landscape mosaics with forests and open habitats (niche construction). Finally, I discuss differences in landscape patterns today and in the Pleistocene, as well as the generality of these ideas to other biogeographical regions. [ABSTRACT FROM AUTHOR]

The article explores the unique relationship between ants, camel crickets, and a non-photosynthetic plant called Balanophora subcupularis. The study reveals that these insects play a dual role as pollinators and seed dispersers for the plant, challenging traditional notions of plant-animal interactions. The findings suggest a rare instance of double mutualism involving invertebrates, highlighting the importance of these insects in the ecosystem. The study was conducted in Japan and sheds light on the evolutionary adaptations of plants in unique ecological niches. [Extracted from the article]

Flowering plants are essentially sessile organisms that disperse their genes through pollination, expanding their areas of occurrence through seed dispersal. In orchids, seed dispersal is commonly mediated by air currents. Conversely, members of several genera have evolved seeds adapted to endozoochory. This is the case for Vanilla, the most economically important genus in the orchid family.The role of indehiscent fruits in the attraction and rewarding of Vanilla seed dispersers was investigated based on field observations, analysis of fragrances, reward substances, and investigation of seed viability through the digestive tract.Indehiscent Vanilla fruits are consumed exclusively by herbivores, i.e. agoutis. Besides providing nutritional rewards, the fruits are rich in polyphenols that are unpalatable to omnivores. The most dominant compound in fruits is attractive only to agoutis. This is the first study showing synzoochory in Orchidaceae and specificity of seed dispersal in orchids.Indehiscent fruits may have evolved early in Neotropical Vanilla in response to selection pressures mediated by large herbivores as the genus emerged ca. 34 Mya in South America, concomitantly with megafauna diversification in the Oligocene. Extinction of the megafauna during the Pleistocene has left agoutis as inheritors of seed dispersal in species with large fleshy fruits. Apart from the effects on omnivores, this study shows that indehiscent fruits of V. chamissonis are consumed exclusively by agoutis, providing the first evidence of target mutualism in orchid seed dispersal. [ABSTRACT FROM AUTHOR]

Cyrtosia septentrionalis, an orchid species, is notable for its sausage‐shaped red fleshy fruits, which can reach up to 10 cm in length. Previous research identified frugivorous and omnivorous birds, especially the brown‐eared bulbul Hypsipetes amaurotis (Pycnonotidae, Passeriformes), as key seed dispersers of C. septentrionalis in natural habitats. This finding challenges conventional belief that orchid seeds are primarily wind‐dispersed. Moreover, given the rarity of specialized co‐evolutionary relationships between fruits and frugivores, C. septentrionalis may also rely on mammals for seed dispersal in certain regions.We aimed to explore the fruit consumption habits of animals other than birds. Motion sensor‐equipped cameras were used to observe interactions with the fruits of C. septentrionalis and assess the viability of seeds post‐consumption.The observations revealed three animal species consuming the fruits, with the masked palm civet Paguma larvata (Viverridae, Carnivora) identified as a dominant consumer in the investigated population. Microscopy analysis of seeds retrieved from P. larvata faeces indicated that the seeds remained intact and viable after passing through the digestive system of this carnivorous mammal.The above discovery suggests a potential role for carnivorous mammals in seed dispersal of C. septentrionalis, alongside birds, thus broadening our understanding of the complex seed dispersal strategies employed by orchids. [ABSTRACT FROM AUTHOR]

Browsing by alien vertebrate herbivores can have both positive and negative impacts on plant invasion. We studied the European hare (Lepus europaeus) diet and foraging behavior focusing on interactions with two invasive non-native shrub species, Cytisus scoparius and Rosa rubiginosa. Fecal pellets were used for microhistological analysis of diet and seed identification. Foraging use of both species was evaluated by browsing rankings. The diet consisted of 28 items, predominating native grasses such as Poa spp. and Festuca pallescens. Among the non-native species, the grasses Hordeum spp. leaves and stems of the shrub C. scoparius, and seed fragments of Rumex acetosella, and rosehip stand out. Fecal pellets contained 431 whole seeds from nine plant species, predominating non-native species. In diet analysis, a portion of damaged rosehip seeds was observed; however, a significant number of viable seeds were previously extracted. This suggests that consumption of R. rubiginosa fruits contributes to seed dispersal, facilitating its spread. In contrast, the browsing of L. europaeus on the vegetative parts of C. scoparius juveniles significantly reduced their cover, which could slow their growth or prevent their spread. These results are an important input for management decisions to prevent or delay the spread of these invasive species. [ABSTRACT FROM AUTHOR]

Ducks are known to be important seed dispersers, but the role of geese in plant dispersal is less clear. Wintering populations of migratory geese are undergoing rapid changes in distribution and habitat use in response to climate change and changes in land use, and the implications for seed dispersal have not previously been studied. At the southern end of Eurasian goose flyways, numbers are declining through short‐stopping.The Doñana wetlands, or 'marismas del Guadalquivir', in south‐west Spain were formerly Europe's most important wintering grounds for the greylag goose Anser anser. There, we collected 151 faecal samples to compare seed dispersal by endozoochory in early and late winter in natural marshes and ricefields. We also tested seed germinability and simulated potential seed dispersal distances to assess the importance of this decreasing wintering population for dispersal of dry‐fruited plants previously assumed to rely on abiotic seed dispersal over short distances.We retrieved 1196 intact seeds belonging to 24 different taxa, including eight species not previously reported from waterfowl (Anatidae) vectors. Seeds were present in 47% of samples, with a peak of 90% in natural marshes in November, compared to only 27% in ricefields at the same time, or in the same marsh site in February. Seed abundance and richness per sample were significantly higher in early than late winter, and in natural compared to in artificial wetlands. Major differences in plant species composition between sampling sites were partly related to habitat differences and moisture requirements of individual species. Germinability in aquatic plants was higher when gut passage was followed by cold storage for 2 months prior to germination tests.We simulated seed dispersal events to and from our sampling sites using GPS tracking of three geese. This suggested that seeds can be dispersed up to 25 km during daily movements, with a median of 0.2–5 km, depending on the sampling site. The most frequent dispersal syndrome assigned to the plant species dispersed by geese was barochory (gravity), and geese disperse plants much farther than the mechanisms predicted by their syndromes.Geese likely have an important role in the dispersal and connectivity of plant populations within and beyond Mediterranean wetlands, providing an ecosystem service which is threatened by changes in migratory behaviour. Our results highlight the importance of studying seed dispersal interactions involving migratory waterbirds in the current context of species loss and distribution shifts, as many crucial interactions may be disappearing even before they are discovered. [ABSTRACT FROM AUTHOR]

Internal transport (endozoochory) and external transport (epizoochory) by migratory waterbirds are key mechanisms of long-distance dispersal for seeds and other diaspores of plants lacking a fleshy fruit. Beginning with Darwin in 1859, we review how opinions about the relative importance of epizoochory and endozoochory have changed repeatedly over time and how this allows us to reassess our modern understanding of plant dispersal. Darwin was mistaken in asserting that diaspores cannot survive passage through the gut of waterbirds or other granivorous birds. This "digestion myth" led him to underestimate endozoochory and overstate the importance of epizoochory, an approach which is echoed throughout the literature until the present day. Darwin also focused on aquatic plants, yet it is now clear that waterbirds are also major vectors of terrestrial plants. Based on their empirical observations and experiments, other less influential scientists (notably Hesselman in 1897, Guppy in 1906 and Proctor in the 1960s) argued that endozoochory is the more important mechanism for waterbirds. Modern field and experimental studies demonstrate the dominant role for endozoochory. Unfortunately, avian endozoochory of dry-fruited plants continues to be ignored as a dispersal mechanism by many plant ecologists, which we attribute to Darwin's continuing influence. However, this endozoochory has major implications for plant biogeography and requires wider recognition and research. [ABSTRACT FROM AUTHOR]

Seed dispersal has been an indispensable ecosystem process mediated by biotic and abiotic vectors. Animal-mediated seed dispersal, such as endozoochory, has supported plants to sustain and establish in new locations. However, increasing urbanisation make feces deposition sites as a detrimental factor for the successful seed establishment. In this regard, this study explores the deposition of feces across the eight natural urban forest fragments with varying urbanisation rates in Tokyo-Yokohama, one of the largest urban agglomerations in the world. Paved roads within the forest fragments were surveyed for feces deposition and the urbanisation rate was estimated for all the studied sites. We found 1381 feces deposited on the paved roads, with the highest in 'Sagamihara Chuo green space' (n = 673) and the second highest in 'Yokohama National University' forest area (n = 488). This study revealed a strong influence of urbanisation on feces deposition in the forest fragments. Birds were the prominent group of animals that deposited feces on paved roads. This is the first study systematically showing the influence of urbanisation on transforming the seed dispersal service by animals into potential low services due to seed wastage in feces deposited on paved roads. Urban forest management needs to consider the impact of urbanisation on not just species diversity, but the interactions and services provided by species. [ABSTRACT FROM AUTHOR]

Societal Impact Statement This study illuminates the underappreciated role of invertebrates in seed dispersal, extending beyond the well‐documented contributions of ants. Focusing on Monotropastrum humile (銀竜草 [silver dragon plant] or 水晶蘭 [crystal orchid]), a non‐photosynthetic plant known for its minuscule, dust‐like seeds, the present research uncovers their seed dispersal roles of woodlice and earwigs in Japan. Remarkably, these invertebrates include the smallest known endozoochorous seed dispersers. Summary Endozoochory, or internal seed dispersal through the digestive tracts of animals, has been less studied in invertebrates compared with vertebrates. Nonetheless, endozoochory is plausible whenever seeds are small enough for ingestion by frugivorous animals, suggesting a potential role for invertebrates in seed dispersal, especially for plants with minute seeds. Monotropastrum humile (Ericaceae), characterized by its fleshy fruits and dust‐like seeds, is known to utilize invertebrate agents such as camel crickets and cockroaches for seed dispersal. Here, we investigate this seed dispersal mechanism using time‐lapse photography, feeding experiments, and seed coat anatomy analysis, particularly focusing on interactions among undocumented invertebrate internal seed dispersers. Field observations indicated that in the studied population, M. humile fruits were primarily consumed by camel crickets, woodlice, and earwigs. Their effectiveness as seed dispersers varied, with camel crickets primarily acting as dispersers, whereas earwigs and woodlice were more inclined toward seed predation. Nonetheless, some seeds defecated by earwigs and woodlice remained intact, suggesting that they could also function as dispersal agents. The woodlouse Porcellio scaber is now recognized as the world's smallest internal seed dispersal agent. Combined with earlier discoveries, such as seed dispersal by camel crickets, cockroaches, and ants, and the fact that P. scaber is an exotic species in the study site, M. humile likely depends on a broad spectrum of local invertebrates. The engagement of multiple invertebrate dispersers may enhance seed dispersal across diverse habitats.

Societal Impact Statement: This study illuminates the underappreciated role of invertebrates in seed dispersal, extending beyond the well‐documented contributions of ants. Focusing on Monotropastrum humile (銀竜草 [silver dragon plant] or 水晶蘭 [crystal orchid]), a non‐photosynthetic plant known for its minuscule, dust‐like seeds, the present research uncovers their seed dispersal roles of woodlice and earwigs in Japan. Remarkably, these invertebrates include the smallest known endozoochorous seed dispersers. Summary: Endozoochory, or internal seed dispersal through the digestive tracts of animals, has been less studied in invertebrates compared with vertebrates. Nonetheless, endozoochory is plausible whenever seeds are small enough for ingestion by frugivorous animals, suggesting a potential role for invertebrates in seed dispersal, especially for plants with minute seeds.Monotropastrum humile (Ericaceae), characterized by its fleshy fruits and dust‐like seeds, is known to utilize invertebrate agents such as camel crickets and cockroaches for seed dispersal. Here, we investigate this seed dispersal mechanism using time‐lapse photography, feeding experiments, and seed coat anatomy analysis, particularly focusing on interactions among undocumented invertebrate internal seed dispersers.Field observations indicated that in the studied population, M. humile fruits were primarily consumed by camel crickets, woodlice, and earwigs. Their effectiveness as seed dispersers varied, with camel crickets primarily acting as dispersers, whereas earwigs and woodlice were more inclined toward seed predation. Nonetheless, some seeds defecated by earwigs and woodlice remained intact, suggesting that they could also function as dispersal agents.The woodlouse Porcellio scaber is now recognized as the world's smallest internal seed dispersal agent. Combined with earlier discoveries, such as seed dispersal by camel crickets, cockroaches, and ants, and the fact that P. scaber is an exotic species in the study site, M. humile likely depends on a broad spectrum of local invertebrates. The engagement of multiple invertebrate dispersers may enhance seed dispersal across diverse habitats. [ABSTRACT FROM AUTHOR]

Frugivory and seed dispersal are fundamental ecological interactions influencing plant population dynamics and distribution. Still, understanding how the ingestion of seeds by fruit-eating animals affects seedling emergence performance remains poorly documented. Seed dispersal through animal ingestion can either promote or hinder germination. In this study, we assessed how seed ingestion by Saguinus bicolor, a critically endangered primate from the Brazilian Amazon, affects germination rate and germinability. As primate seed ingestion often improves seed germination, we predicted that germination rate and germinability would be higher in seeds defecated by the tamarins if compared to hand-extracted seeds. Of the 23 species, five did not germinated in either the control or gut-passage treatments. Among the ones that germinated, nine had an average increase in germinability, two decreased in germinability, and seven had a neutral effect. The germination rate increased in four species, reduced in eight species and had a neutral effect in six of the species. Most of the seeds that passed through the tamarins' digestive tract successfully germinated, demonstrating that seed ingestion by pied tamarins can benefit the plant populations by dispersing viable seeds through the landscape. [ABSTRACT FROM AUTHOR]

Seed dispersal by frugivorous animals is a key process for plant populations and communities. When frugivores consume fruits, seeds are temporarily retained inside their guts until seed deposition. Hence, information on retention times is essential to estimate seed‐dispersal distances. Although it is well known that retention times are affected by the body size of frugivores, there is an important knowledge gap on the effects of plant species traits, such as seed size and pulp content.Here, we comprehensively address whether and which plant traits affect seed retention times in frugivorous birds. On the one hand, we conducted a first set of experiments to assess variation in retention times among 31 plant species dispersed by a single bird species, and a second set to assess variation among five plant species dispersed by a guild of five frugivorous passerines. On the other hand, we conducted a literature review retrieving retention times for 231 interactions involving 155 plant species and 55 bird species from nine different avian orders.Seed size had negative effects on retention times at the three levels addressed in this study: the larger the seeds, the shorter the times. The effects of seed size were higher within a single bird species, intermediate for the small assemblage of five passerine species, and lower for all frugivorous birds included in the compilation. Notably, the effects of seed size on retention times were comparable to that of frugivore body size in Passeriformes. We analysed the effects of the pulp content on a single bird species, but these lacked predictive power. Importantly, the effects of seed size were indirect and mediated by the type of seed ejection (digestive processing): birds generally defecate smaller seeds and regurgitate the larger ones.We provide comprehensive evidence that retention times are intrinsically linked to plant traits and to frugivores' response to such traits, particularly to seed size and the type of seed ejection, respectively. Thus, our study unveils a source of interspecific variability in the capacity of plants to disperse at long distances. Moreover, we provide methodological improvements to trait‐based models that estimate mean retention times and seed‐dispersal distances. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Zoysia japonica, a low growing grass, is tolerant to grazing and trampling. Kinkazan Island in northern Japan is inhabited by sika deer (Cervus nippon). The deer population increased in the 1970s at a shrine garden in the western part of the island, leading to expansion of Zoysia swards around the shrine garden and their colonization of remote open patches. The expansion around the garden is due to elongation of the rhizome, but expansion to remote places may be due to endozoochory by deer. This appears to be a good example of the “foliage is the fruit” hypothesis (the FF hypothesis) proposed by Janzen (1984; American Naturalist 123:338–353). To demonstrate this, we confirmed the expansion of the Zoysia swards and tested the traits of Zoysia by field surveys and indoor experiments. The Zoysia peduncles stood among the leaves, and sika deer fed on both the seeds and leaves. One deer fecal pellet contained about 20 seeds at its peak in June. In the feeding experiment, the survival rate through digestion was 38%. In the greenhouse experiment, the germination rate was 72%. An outdoor experiment showed that germination rates of the ingested seeds were 5% in a dark habitat and 58% in a bright habitat. Zoysia exhibited many of the traits presented by the FF hypothesis, and sika deer functioned as seed dispersal agents. Therefore, the Zoysia—sika deer relationship seems to support the FF hypothesis. [ABSTRACT FROM AUTHOR]

Seed dispersal distance is an important indicator of how well a plant species can cope with environmental changes. Seeds of wetland plants are primarily dispersed by hydrochory (floating on the water surface) or endozoochory (ingestion and egestion by animals). However, both mechanisms can also be combined (diplochory), which increases dispersal distances. In wetlands, seeds often float on the water surface before ingestion and transport by animals, such as waterbirds. Here, we consider what happens next after endozoochory and test whether seeds dispersed inside waterbirds can then continue to disperse by hydrochory. We experimentally assessed the buoyancy capacity of 41 wetland plant species for 14 weeks before and after simulated passage through avian digestive systems. This revealed that (1) seeds of plants previously assigned a hydrochory dispersal syndrome floated longer than those from other syndromes, but with considerable overlap; (2) fully aquatic and shoreline plant species had seeds with stronger buoyancy than terrestrial plants; and (3) digestive processes negatively affected seed buoyancy capacity for all plant species, which included wetland species with a typical hydrochory syndrome. The capacity for hydrochory is more limited after endozoochory than beforehand, with strong implications for the effectiveness of seed dispersal. [ABSTRACT FROM AUTHOR]

We analysed internal dispersal of zooplankton by waterbirds (endozoochory) in New Zealand, quantifying zooplankton eggs in faecal droppings collected at two lakes, Lake Rotoroa (Hamilton) and Lake Rotorua. Sixty-seven faecal droppings were collected from Mallard Ducks (20), Canada Geese (11), Greylag Geese (6), Black Swans (20) and Australian Coots (10). Fifty eggs were found, with a mean of 0.75 eggs per dropping, indicating that waterbirds consume zooplankton eggs, and that these pass through the digestive system. No significant difference was observed in the abundance of eggs among waterbird species, and no eggs hatched in the laboratory. Our results suggest that waterbird dispersal of zooplankton in New Zealand is occurring, but numbers being transported are low. Further, as non-native waterbirds such as mallard ducks and geese do not migrate in New Zealand to the extent they do elsewhere, they are likely not primary vectors for zooplankton dispersal. [ABSTRACT FROM AUTHOR]

Summary: Mycorrhizal symbiosis, seed dispersal, and pollination are recognized as the most prominent mutualistic interactions in terrestrial ecosystems. However, it remains unclear how these symbiotic relationships have interacted to contribute to current plant diversity.We analyzed evolutionary relationships among mycorrhizal type, seed dispersal mode, and pollination mode in two global databases of 699 (database I) and 10 475 (database II) tree species. Although database II had been estimated from phylogenetic patterns and therefore had lower certainty of the mycorrhizal type than database I, whose mycorrhizal type was determined by direct observation, database II allowed analysis of many more taxa from more regions than database I.We found evidence of joint evolution of all three features in both databases. This result is robust to the effects of both sampling bias and missing taxa. Most arbuscular mycorrhizal‐associated trees had endozoochorous (biotic) seed dispersal and biotic pollination, with long dispersal distances, whereas most ectomycorrhizal‐associated trees had anemochorous (abiotic) seed dispersal and wind (abiotic) pollination mode, with shorter dispersal distances.These results provide a novel scenario in mutualistic interactions, seed dispersal, pollination, and mycorrhizal symbiosis types, which have jointly evolved and shaped current tree diversity and forest ecosystem world‐wide. See also the Commentary on this article by Sinnott‐Armstrong, 243: 1290–1292. [ABSTRACT FROM AUTHOR]

Buttonweed (Cotula coronopifolia) is native to South Africa but invasive in wetlands in Europe, North America, and Australasia, where it excludes native plants. Despite being dry-fruited, field studies suggest migratory waterbirds can disperse its seeds via gut passage (endozoochory), aiding its expansion. To explore the potential for endozoochory in different regions and habitats, we collected seeds from six populations in Spain, Sweden, and the UK. Germination was tested under different salinity levels (0, 5, 10, 15 g/L) and simulated gut passage treatments: scarification, acidification, or both. No germination occurred at 15 g/L. Higher salinity reduced and delayed germination, but full gut passage treatment (i.e., both scarification and acidification) increased germinability and accelerated germination. Scarification or acid treatment alone resulted in intermediate germination patterns. There were significant salinity × population and gut passage × population interactions on germinability. The acceleration effect of gut passage on germination was stronger at 5–10 g/L than at 0 g/L. This study highlights how migratory birds can facilitate the spread of alien plants introduced by humans. Endozoochory by waterbirds is an understudied mechanism for the long-distance dispersal of dry-fruited alien plants. Further research on C. coronopifolia, including population genetics, is necessary to understand dispersal mechanisms and facilitate management strategies. [ABSTRACT FROM AUTHOR]

Societal Impact Statement Summary Dust seeds, which are minute and contain minimal energy reserves, are often associated with heterotrophy (plants that obtain carbon without photosynthesis). Consequently, previous studies have mainly focused on the relationships between dust seeds and heterotrophy. However, dust seeds are also found in green plants. This manuscript focuses on the seed ecology of the apparently autotrophic shrub Rhynchotechum discolor that produces dust seeds. Using time‐lapse photography, feeding experiments, and germination tests, we show that camel crickets effectively disperse the seeds of this autotrophic shrub. This is the first study to document insect‐mediated internal seed dispersal of an autotrophic plant in regions inhabited by terrestrial mammals, offering new insights into the evolutionary ecology of dust seeds. Although angiosperms exhibit a wide range of seed sizes, the mechanisms driving these differences are poorly understood. The evolution of dust seeds, which contain minimal energy reserves, is traditionally linked to heterotrophy, in which external carbon sources are provided to the embryo or seedling. Dispersal by small animals may be another important but underexplored evolutionary driver of minute seed size. However, insect endozoochory has been documented only in the seeds of heterotrophic species and autotrophic species in New Zealand (i.e., in communities without native terrestrial mammals). Here, we investigate this seed dispersal mechanism in the Japanese shrub Rhynchotechum discolor (Gesneriaceae), which produces white berries with numerous tiny ellipsoid seeds, using time‐lapse photography, feeding experiments, and germination tests, particularly focusing on potential interactions with insect internal seed dispersers. Our time‐lapse photography indicated that camel crickets predominantly feed on fallen fruits of R. discolor. Field sampling revealed that camel crickets excreted numerous intact R. discolor seeds. Feeding experiments confirmed that some camel crickets excreted a high proportion of intact seeds, many of which successfully germinated into autotrophic seedlings. The present study provides the first evidence of insects acting as endozoochorous seed dispersers in fully autotrophic plants within regions inhabited by terrestrial mammals. Our findings suggest that orthopteran‐mediated seed dispersal is more widespread than previously thought, and that dust seeds can evolve from selective pressures beyond heterotrophy, specifically through insect‐mediated seed dispersal. [ABSTRACT FROM AUTHOR]

The role of large frugivores in the dispersal of tropical trees has been well‐studied, whereas the importance of the world's largest rodents for plant dispersal has previously been ignored. We studied plant dispersal by gut passage (endozoochory) in Neotropical wetlands by the largest extant rodent, the capybara Hydrochoerus hydrochaeris. We compared it with another of the world's 10 largest rodents, the nutria or coypu Myocastor coypus.We collected 96 faecal samples (50 capybara, 46 nutria) from 10 sites within the Taim Ramsar site in southern Brazil, and extracted intact propagules from 10 g subsamples. Propagules from 27 plant taxa (24 angiosperms, and three aquatic ferns) were recovered, representing 13 plant families. Whole Spirodela (duckweed) plants were dispersed. Seeds from 14 angiosperm taxa were germinated in the laboratory.Taxonomic composition of propagules was significantly different for the two mammals, but propagule size was not. Capybara dispersed more terrestrial propagules per sample, and nutrias more aquatic propagules. When total faecal production was taken into account, an estimated 1025 propagules (including 133 fern spores) were dispersed by each capybara per day, compared with 691 propagules for nutrias (including 566 fern spores). Capybaras have larger home ranges and likely dispersed plants over a greater distance.Synthesis: This is the first demonstration that rodents disperse aquatic pteridophytes as well as flowering plants. Our results suggest that capybaras and nutrias play important, complementary roles in dispersing a variety of aquatic and terrestrial plant species within Neotropical wetlands. Our study indicates that the role of herbivorous mammals as vectors of plants with small seeds and non‐fleshy fruits has been overlooked in the neotropics and that defaunation of large herbivores can have negative consequences, in a manner similar to that previously highlighted for frugivores. [ABSTRACT FROM AUTHOR]

The dispersal of fish into distant and isolated habitats remains a topic of continuous discussion in the field of fish biogeography. This is particularly relevant due to the perceived limitation of fish movement to what is known as active dispersal. Fish migration is often confined to interconnected water bodies, underscoring the significance of dispersal for fish inhabiting isolated aquatic habitats. However, empirical evidence for a natural (i.e. not human‐mediated) mechanism has been limited. Here we explore and provide evidence for waterbird‐mediated endozoochory as a possible dispersal mechanism in various fish species and families. We force‐fed mallardsAnas plathyrynchoswith fertilised eggs of nine bony fish species, covering nine taxonomic families. We recovered viable embryos of five fish taxa in the faeces of mallard, proving the ability of fish eggs to survive the passing of the digestive system of waterbirds. Moreover, the recovered eggs successfully hatched into larvae in two fish species. Taking into the flight speed and numerosity of mallards, as well as the high abundance of fish eggs, our results highlight endozoochory of fish eggs by waterbirds as a possible significant, although likely rare natural dispersal mechanism that can occur across more species than previously known in freshwater fish. [ABSTRACT FROM AUTHOR]

Current global change scenarios demand knowledge on how anthropogenic impacts affect ecosystem functioning through changes in food web structure. Frugivorous mesocarnivores are a key link in trophic cascades because, while their abundance and behaviour are usually controlled by apex predators, they can provide high‐quality seed‐dispersal services to plant communities. Thus, the recent rewilding of large carnivores worldwide can trigger cascading effects for plants.We investigated the top‐down effects of an apex predator (Iberian lynx Lynx pardinus) on seed‐dispersal services mediated by two mesocarnivore species (red fox Vulpes vulpes and stone marten Martes foina) at the plant community level by comparing areas with and without lynx in a Mediterranean mountain range in Southern Spain.We collected scats of mesocarnivores (n = 1575) to assess frugivory and seed dispersal of 15 plant species over two consecutive fruiting seasons and two habitat types (open and forest). Specifically, we assessed the effect of lynx presence on (i) seed occurrence and fleshy‐fruit biomass per scat, (ii) number of scats containing seeds and (iii) diversity of dispersed seeds.The quantity and diversity of dispersed seeds drastically decreased under predation risk for both mesocarnivore's species. Seed dispersal by stone martens was negatively affected by the presence of lynx, with a marked reduction in the number of scats with seeds (93%) and the diversity of dispersed seeds (46%). Foxes dispersed 68% fewer seeds in open habitats when coexisting with lynx, probably leading to differential contributions to seed‐dispersal effectiveness among habitats.Our study reveals a novel trophic cascade from apex predators to plant communities. The behavioural responses of frugivorous mesocarnivores to predation risk and the reduction in the intensity of their faecal deposition pattern are probably related to their lower abundance when co‐occurring with apex predators. While rewilding apex predators is a successful conservation tool, attention should be paid to cascading effects across food webs, particularly where frugivore megafauna is missing and mesocarnivores provide unique services to plants. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Frugivores and seed dispersers are important in maintaining biodiversity and ecosystems. Yet, these agents receive less attention, in particular from ubiquitous or garden-cultivated species like Amorphophallus paeoniifolius and Alocasia odora. Despite extensive ecological studies carried out across the distribution regions of both species in tropical and subtropical, the only known disperser of A. paeoniifolius is the bird-of-paradise that was reported from Papua New Guinea. Meanwhile, there are reports of Pycnonotus and Eudynamys being the dispersers for both wild and cultivated Amorphophallus in India. Despite this, the frugivores and dispersal agents for both A. paeoniifolius and A. odora, in general, are not known and may be different in other parts of the world. In this study, I set up camera traps at five sites (two for A. paeoniifolius and three for A. odora) between 2017 and 2019 at the Xishuangbanna Tropical Botanical Garden to observe the frugivores and dispersal agents for these two ubiquitous cultivated aroid species. Apart from morphological observations on the infructescence of A. paeoniifolius and A. odora, the size of berries were measured and number of seeds per berry were counted. Six frugivores (Pycnonotus aurigaster, Copsychus saularis, Alophoixus pallidus, Niltiva davidi, Orthotomus sutorius, and Pycnonotus jocosus) were identified as potential dispersal agents, attracted to the colourful, succulent, and smaller (< 2 cm) berries. Pycnonotus aurigaster visited both A. odora and A. paeniifolius. The latter four frugivores visited A. paeniifolius, whereas P. jocosus visited only A. odora. All frugivores demonstrate endozoochory, and consumed the entire berries without showing any peculiar behaviour patterns. The berries are red-yellowish on ripening, with an average size of 1.63 cm ± 0.23 cm × 0.88 cm ± 0.19 (L × W) for A. paeniifolius, and 1.04 ± 0.09 cm × 0.67 ± 0.09 cm for A. odora. This study provides insight into how frugivores may depend on and connect to their regions of distribution as well as the distributions of plant taxa. Similar studies could be comparatively expanded into different distribution localities of A. paeoniifolius and A. odora for a wider understanding of their relationship linkage to the dispersal agents. [ABSTRACT FROM AUTHOR]

Questions: Free‐ranging herbivores are often seen as important endozoochorous dispersal vectors for plant propagules. Dispersal by free‐ranging animals, in contrast to domestic animals, and its effects on vegetation have rarely been studied. We ask what are the effects of deposition of dung on open landscape vegetation, how does it influence vegetation development after disturbance by soil removal, and does it change species richness or composition? Location: The Doupov Mountains, Czech Republic. Methods: We established a six‐year experiment in 2015 to study the effects of dung deposition on vegetation of an open grassland landscape. The experiment comprised 300 plots distributed across five fenced site locations and treatments were implemented in partially crossed design, with soil removed (disturbance) and dung pellets deposited; treatments were applied once to five replicates, from June to October 2015. We recorded plant species cover at the beginning of the vegetation seasons of 2016, 2017 and 2021. Results: Dung deposition increased species richness and Ellenberg Indicator Values for nutrients, and these effects were stronger in disturbed plots; however, these effects diminished with time. There were no effects of dung deposition on species composition. In contrast, disturbance affected species diversity, composition, Ellenberg Indicator Values for nutrients, proportion of dung‐dispersed species, but not species richness. Surprisingly, there were no effects of timing of dung deposition. Conclusions: Our results indicate that endozoochorous seed dispersal by red deer impacts vegetation dynamics of this open grassland landscape, but effects are weak and diminish with time. Considering the large seed load in dung, it is possible that endozoochorous seeds from wild herbivores are incorporated into the soil seed bank, where they persist until appropriate ecological conditions for germination arise. [ABSTRACT FROM AUTHOR]

Plants inhabiting open landscapes are often dispersed by ungulates and are expected to be adapted to this type of dispersal through their seed traits. To find which traits help seeds survive the passage through digestion of wild ungulates, we conducted a comprehensive feeding experiment with almost forty species of plants and three species of ungulates. We fed specified numbers of seeds to the animals, collected the dung, and germinated the dung content. We explored whether seed morphological traits and seed nutrient contents are good predictors of seed survival after passage through the ungulate digestive system. We also tested how the seed survival differed after the passage through different ungulate species. To find answers, we used GLMM with beta-binomial distribution and animal and plant species as random factor, respectively. We found that species survival and germination success were negatively correlated to seed elongation and the thickness of the seed coat. Even though phylogenetically correct GLMM did not yield significant results, when we tested species from commonly represented families, separately (legumes and grasses compared to all other species) different traits had statistically significant effects. In the case of seed elongation, the effect changed direction from negative to positive when legumes and grasses were left out. Our results suggest that seed traits enabling species survival after passage through the digestive tract are strongly phylogenetically conserved and different groups of plants evolved different ways of adapting to grazing pressure and utilize it for dispersal. [ABSTRACT FROM AUTHOR]

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Copyright of Biotropica is the property of Wiley-Blackwell and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Carnivorous mammals have been reported to feed on fleshy fruits and disperse seeds, but these interactions are seldom observed and are poorly understood in hyper‐arid regions. Scent may play a key role in these interactions, given that most carnivores have sensitive olfactory systems and dichromatic vision. The !nara (Acanthosicyos horridus: Cucurbitaceae) in the Namib Desert produces large (10–20 cm diameter) melons that remain green while undergoing increased volatile emissions with a changing chemical profile as they ripen. Using extensive camera trapping, we found that the fruits are consumed mainly by black‐backed jackals (Canis mesomelas) and rarely also by other carnivores such as cape fox (Vulpes chama) and brown hyaena (Hyaena brunnea). We found that scent cues play a crucial role as jackals mainly sought fruits at night and easily located ripe fruits that were experimentally buried beneath the sand to remove visual cues. Seeds retrieved from jackal scat showed improved germination relative to un‐ingested seeds. This study highlights the importance of fruit scent in mediating seed dispersal mutualisms involving carnivores. [ABSTRACT FROM AUTHOR]

Questions: How do seed rain biomass and richness change from old‐growth tropical forests to pastures at different elevations? How do seed mass and seed dispersal mode change from forests to pastures across these elevations? What implications do these changes have for the recovery of deforested areas in tropical mountains? Location: Old‐growth montane forests and livestock pastures along an elevational gradient (1,000–3,000 m a.s.l.), located at Podocarpus National Park, San Francisco Reserve and surrounding agricultural lands, Andes of southern Ecuador. Methods: We collected seed rain for a 3‐month period using 324 traps installed at eighteen 1‐ha plots across elevations. Half of the traps were installed in nine 1‐ha forest plots, and the other half in nine 1‐ha pasture plots. For each trap, we identified the seeds and measured seed rain biomass and richness. We also recorded seed traits and calculated community‐weighted means of seed mass and seed dispersal mode (proportion of endozoochory). Results: Forests received a higher seed rain biomass than pastures, but only at the lowest elevation. Seed rain richness did not differ between habitat types at all elevations. Community‐weighted means of seed mass and the proportion of endozoochorous species declined from forests to pastures, especially at the lower elevations. Conclusions: Although seed rain biomass and richness were overall similar between forests and pastures, large‐seeded and endozoochorous species were generally poorly represented in the seed rain of pastures compared with that of forests. These findings show that biomass and richness of seed rain may be insufficient to quantify the restoration potential of natural seed rain in deforested areas. Information on seed traits, such as seed mass and seed dispersal mode, is important to optimize restoration efforts towards the regeneration of diverse old‐growth forests along elevational gradients in tropical mountains. [ABSTRACT FROM AUTHOR]

The process of seed dispersal that underpins ecosystem maintenance is performed by diverse arrays of fruit‐eating animals. However, seed dispersal studies are primarily focused on a subset of these animal communities that disperse seeds by endozoochory. Stomatochory (seed dispersal in which seeds are carried externally and are not swallowed) is rarely considered to be effective, despite an increasing number of taxa‐focused studies that indicate otherwise.We collated the available information on stomatochory to provide a quantitative overview of the dispersal mechanism, including plant types and fruit traits dispersed, dispersal distances and germination potential for all available taxa. We compared seed sizes dispersed, dispersal distances and germination potential with corresponding data on endozoochory for bats and primates. We also identified the main taxa dispersing seeds by stomatochory and assessed what factors influenced the distances that they carried seeds to.Stomatochoric dispersers can displace large quantities of seeds, including large seeds and those of large fruits, and over short to long distances (>1 km). Compared with similar‐sized endozoochoric dispersers, they can disperse larger seeds, but over shorter distances, on average. Similar to endozoochory, seed handling by stomatochory also improves the germination potential of dispersed seeds. Dispersal distances achieved by stomatochory were influenced by body mass, daily path length, seed width, fruit type and seed handling techniques.Five main taxonomic groups of stomatochoric dispersers were identified: bats, parrots, squirrels, corvids and Old World monkeys (cercopithecines). Parrots perform dispersal services for the largest fruits and over the longest distances. However, given the lack of research on stomatochory, it is likely other taxa are also important stomatochoric dispersers but have not been identified yet.More research attention must be directed towards seed dispersal services that are not provided by endozoochory. Many stomatochoric dispersers are common animals within communities and could be playing dominant seed dispersal roles, even without swallowing seeds. Community‐wide studies should incorporate all seed dispersal interactions, rather than focusing solely on endozoochory. This will ensure a more robust understanding of community‐wide patterns. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Megafauna are important seed dispersers because they can disperse large quantities of seeds over long distances. In Hokkaido, Japan, the largest terrestrial animal is the brown bear (Ursus arctos) and other megafauna seed dispersers are lacking. Thus, brown bears are expected to have an important function as seed dispersers in Hokkaido. In this study, we, for the first time, evaluated the seed dispersal function of brown bears in Hokkaido using three fleshy-fruited trees and studied: (1) gut passage time (GPT) in feeding experiments, (2) seed dispersal distance using tracking data of wild bears, and (3) the effect of gut passage and pulp removal on germination rate. Most seeds were defecated intact, and less than 6% were broken. The average GPT without pulp was 3 h and 56 min to 6 h and 13 min, depending on the plant and trial. Each plant's average simulated seed dispersal distance was 202–512 m. The dispersal distance of Actinidia arguta seeds with pulp was significantly longer than those without pulp because of their longer GPT. The germination rate of defecated seeds without pulp was 19–51%, depending on the plant, and was significantly higher or not different comparing with that of seeds with pulp. We concluded that brown bears in Hokkaido are effective seed dispersers. In managing brown bears in Hokkaido, such ecological functions should be considered along with conserving the bear population and reducing human–bear conflicts. [ABSTRACT FROM AUTHOR]

Natural grasslands, one of the main ecosystems worldwide, are under threat and native species are desired when addressing restoration issues. The main processes for grasslands restoration success are plant establishment and seed addition in the soil. Thus, it requires overcoming barriers that prevent germination and seed dispersal strategies. This study aimed to assess the germination potential, treatments to overcome seed dormancy and endozoochory dispersal of forage native species by cattle. Seeds from 21 native species from the four most common botanical families (Poaceae, Asteraceae, Fabaceae and Cyperaceae) were collected in a natural grassland area (Pampa Biome) and used in the experiments. Experiment I: germination under controlled conditions of all species was carried out and compared to commercial forage (Paspalum notatum cv. "Pensacola"). Experiment II: three treatments to overcome seed dormancy (mechanical and chemical scarification; priming agent) were tested in six species. Experiment III: the recovery, viability and germination of native P. notatum Flüggé and Desmodium incanum DC seeds from cattle faeces were evaluated. The results show that Paspalum umbrosum Trin. and Saccharum angustifolium Nees have high germination potential when compared to commercial forage. The treatments to overcome seed dormancy increased the germination of D. incanum, Paspalum plicatulum Michx. and Paspalum dilatatum Poir. Total seed recovery of 15% (P. notatum) and 26% (D. incanum), without affecting seed viability, show that cattle are able to supply the soil seed bank with native species. Germination potential, treatments to overcome seed dormancy and endozoochory dispersal of forage native species may be exploited in natural grassland restoration. [ABSTRACT FROM AUTHOR]

The plant species of the Caatinga biome are food sources of great importance for herds. Goat farming is one of the most explored agricultural activities in the Northeast region of Brazil, where goats are important seed dispersers in semiarid areas. In this study, we evaluate the passage of seeds of four Caatinga forest species through goats' digestive system, their recovery, and the effect on seed germination (Ger). A completely randomized experimental design was used. The seeds of four species Piptadenia stipulacea, Mimosa tenuiflora, Desmanthus virgatus, and Mimosa caesalpiniifolia were provided to eight goats and evaluated after nine excretion periods (12, 24, 36, 48, 60, 72, 84, 96, and 108 h). Seed recovery was low, with values of 10.87% for M. tenuiflora, 3.41% for P. stipulacea, 2.04% for M. caesalpiniifolia, and 1.54% for D. virgatus. The seeds of D. virgatus and M. tenuiflora are small and pass faster through goats' digestive system, with the largest amount recovered 48 h after ingestion. In contrast, P. stipulacea and M. caesalpiniifolia seeds are retained for a longer period in the intestine of the animals, with the highest number obtained 72 and 96 h after, respectively. The passage of seeds through the digestive system of D. virgatus for 24 h and P. stipulacea for 72 h, promoted the overcoming of dormancy, reaching Ger values of 66.7 and 35.7%, respectively. The Ger of M. tenuiflora seeds as well as the Ger and survival of M. caesalpiniifolia seeds decreased after passing through the digestive system. The recovery of seeds from the four species indicates that goats are efficient seed dispersers in Caatinga areas. [ABSTRACT FROM AUTHOR]

Macrocyclic lactone anthelmintics are widely used to control invertebrate pests in livestock, such as sheep. While anthelmintic effects on non‐target animals, such as dung‐dwelling insects, are well studied, effects on seed germination are largely unknown. Seeds can come into contact with anthelmintics either during passage through the gastro‐intestinal tract of grazing animals or when anthelmintics are excreted with their dung into the environment, which may result in changed germination patterns.We used four commonly applied macrocyclic lactones to assess their effects on germination: moxidectin, ivermectin, abamectin and doramectin as pure substances; moxidectin and ivermectin also in formulated form. We tested these pharmaceuticals on 17 different temperate grassland species from five plant families. Seeds were exposed to three concentrations of macrocyclic lactones (0.1, 1.0 and 10.0 mg·l−1) under controlled conditions, and germination was assessed over a 6‐week period. From these data, we calculated germination percentage, mean germination time and germination synchrony.Most of the tested species were significantly affected in germination percentage and/or mean germination time by at least one of the tested pharmaceuticals, with formulated moxidectin having the largest impact. In general, the effects found were species‐ and pharmaceutical‐specific. While formulated substances generally reduced germination percentage and increased mean germination time, pure substances increased germination percentage. Synchrony showed less clear patterns in all pharmaceuticals.Although effect size and sign varied between species, our study shows that non‐target effects of macrocyclic lactones commonly occur in terrestrial plants. This may impede successful seed exchange between habitats via sheep, and even translate into profound changes to grazed ecosystems. [ABSTRACT FROM AUTHOR]

The success of plant reproduction is highly dependent on effective seed dispersal. This study aimed to evaluate the potential seed dispersal effectiveness of cattle for Malus sieversii. The impact of cattle on the dispersal quantity and dispersal quality of M. sieversii seeds was explored based on camera trapping, GPS tracking, and germination trials. The results showed that, on average, cattle visited M. sieversii trees 477.33 times during a two-month observation period. Out of these visits, 315 were specifically for fruit removal. The fruit removal rate per cattle visit was as high as 96.67%. Additionally, cattle were able to disperse M. sieversii seeds up to a maximum distance of 533.67 m, with an average dispersal distance of 134.62 m. The average distance of cattle movement was recorded as 176.95 m/h, with peak activity observed during 11:00–13:00 and 19:00–21:00. The germination rate of M. sieversii seeds that passed through the digestive tract of cattle was significantly higher than that of control seeds. Finally, the emergence rate and survival rate of seeds dispersed by cattle to forest edges and gaps were significantly higher than those dispersed to understory. These findings suggest that cattle can serve as effective long-distance dispersers of M. sieversii seeds and may play a crucial role in the regeneration and expansion of M. sieversii populations in the Ili Botanical Garden. [ABSTRACT FROM AUTHOR]

Seed dispersal is a critical process in plant colonisation and demography. Fruits and seeds can be transported by several vectors (typically animals, wind and water), which may have exerted strong selective pressures on plant’s morphological traits. The set of traits that favour dispersal by a specific vector have been historically considered as seed dispersal syndromes. As seed dispersal syndromes have a great potential to predict how seeds move (i.e. the relative importance of the standard mechanisms of seed dispersal), they have attracted the attention of naturalists and researchers for centuries. However, given that observations of actual dispersal events and colonisation are seldom reported, there is still much confusion in current studies failing to properly discriminate between seed dispersal syndromes (i.e. sets of traits that favour a particular mechanism) and actual seed dispersal (i.e. the vector that moves a given seed in one dispersal event). This distinction is important because the presence of any seed dispersal syndrome does not preclude the seed being occasionally dispersed by other non-standard mechanisms (i.e. different from the one predicted). Similarly, the absence of seed dispersal syndromes does not prevent seeds from being dispersed. The correct coding of seed dispersal syndromes thus requires a systematic and evolutive, rather than a phenomenological approach. Unfortunately, such approach has rarely been implemented at a community-level and no comprehensive datasets of seed dispersal syndromes are yet available for any entire flora.This database contains categorisation of the native European flora into eight seed dispersal syndromes. Information for a total of 9,874 species retrieved from the volumes of Flora Europaea were analysed. Earlier versions of this database, which only coded for the presence of four long-distance dispersal syndromes (endozoochorous, epizoochorous, thalassochorous and anemochorous diaspores), were used in four previous studies. Here, we present a fully revised and expanded database, including the presence of four additional short-distance dispersal syndromes (myrmecochorous, vertebrate hoarding, freshwater hydrochorous and ballochorous diaspores), a nomenclatural update for all species and the codification of 416 additional species.Roughly half (51.3%) of the native European flora produce diaspores without traits clearly associated with facilitating seed dispersal. The other half (48.7%) of the European plant species produces diaspores with some specialised traits associated with seed dispersal, most of which (79.9%) with a potential to facilitate long-distance dispersal events. The most common diaspores are those with anemochorous (23.5%), epizoochorous (8.0%), endozoochorous (7.8%), myrmecochorous (7.2%), thalassochorous (2.3%), freshwater dispersal (2.1%), ballochorous (4.6%) and vertebrate hoarding associated traits (0.2%). Two-thirds (66.3%) of the European shrub and tree species have diaspores with some specialisation for biotic seed dispersal.

Buttonweed (Cotula coronopifolia) is native to South Africa but invasive in wetlands in Europe, North America, and Australasia, where it excludes native plants. Despite being dry-fruited, field studies suggest migratory waterbirds can disperse its seeds via gut passage (endozoochory), aiding its expansion. To explore the potential for endozoochory in different regions and habitats, we collected seeds from six populations in Spain, Sweden, and the UK. Germination was tested under different salinity levels (0, 5, 10, 15 g/L) and simulated gut passage treatments: scarification, acidification, or both. No germination occurred at 15 g/L. Higher salinity reduced and delayed germination, but full gut passage treatment (i.e., both scarification and acidification) increased germinability and accelerated germination. Scarification or acid treatment alone resulted in intermediate germination patterns. There were significant salinity × population and gut passage × population interactions on germinability. The acceleration effect of gut passage on germination was stronger at 5–10 g/L than at 0 g/L. This study highlights how migratory birds can facilitate the spread of alien plants introduced by humans. Endozoochory by waterbirds is an understudied mechanism for the long-distance dispersal of dry-fruited alien plants. Further research on C. coronopifolia, including population genetics, is necessary to understand dispersal mechanisms and facilitate management strategies.

Migratory waterbirds disperse a broad range of angiosperms by endozoochory (seed dispersal via gut passage), especially plants in coastal wetlands. However, there is no previous information about the capacity of seeds to remain in the seed bank after waterbird endozoochory, and very little about how wetland salinity can influence the effect of gut passage on germination. We collected seeds of Juncus subulatus (Juncaceae), Bolboschoenus maritimus, and Schoenoplectus litoralis (Cyperaceae) from Doñana marshes in Spain. All three species are considered to have physiological dormancy. After gut passage following ingestion by ducks, seeds were stored in darkness in solutions with six different conductivities (1, 2, 4, 8, 16, and 32 dSm-1), for periods of 1, 6, or 12 months to simulate presence in a seed bank. After storage, 1800 seeds of each plant species assigned to these treatments were subjected to germination tests in demineralized water, together with 1800 control seeds that had not been ingested before storage. All species germinated readily after storage, with or without gut passage beforehand. Storage time and salinity both had important effects on germinability and time to germination, which differed between control and ingested seeds, and between plant species. After ≥6 months, germinability of Cyperaceae was enhanced by gut passage (≤25% higher than control seeds) at some salinities. Only J. subulatus showed consistently lower germinability after passage (≤30%). Only B. maritimus showed consistently slower germination after passage (≤33%). Salinity effects were more complex after passage, but increasing salinity did not generally have a negative impact on germination of ingested seeds. When compared to additional seeds that had not been stored before germination tests, storage reduced germinability in J. subulatus (≤39% reduction), but increased it in B. maritimus (≤17%) and S. litoralis (≤46%). Seeds dispersed by waterbird endozoochory may be easily incorporated into wetland seed banks, where they can remain halotolerant and delay germination until conditions become suitable. This can benefit wetland plants by increasing rates of long-distance dispersal, gene flow, and establishment of new populations. Avian gut passage can have positive and species-specific effects on germination in plants with persistent seed banks and/or physiological dormancy [ABSTRACT FROM AUTHOR]

Understanding the influence of non‐native herbivores on ecosystems by means of dietary foraging and seed dispersal is important for understanding how non‐native species can alter an invaded landscape, yet requires multiple methodologies. In south‐eastern Australia, introduced sambar deer (Rusa unicolor) are rapidly expanding in range and placing native ecosystems at risk through browsing and as vectors for seed dispersal. We simultaneously investigated sambar deer dietary composition and seed dispersal using DNA sequencing and germination trials, from faecal pellets collected in alpine and wet forest ecosystems. This allowed us to contrast the dietary impacts of introduced sambar deer in different environments, and to explore the potential for habitat‐specific variation in diet. DNA sequencing of the trnL, ITS2 and rbcL gene regions revealed a diverse plant species dietary composition comprising 1003 operational taxonomic units (OTUs). Sambar deer exhibited intermediate feeder behaviours dominated by forbs in alpine and shrubs in wet forest ecosystems. A large proportion of plant OTUs were considered likely to be native, however, the proportion of exotic species in the diet in both ecosystems was greater than would be expected based on the proportion of exotic species in each of the two landscapes. Seed germination trials indicated that sambar deer can disperse a substantial number of native and exotic species in both alpine and wet forest ecosystems. In alpine ecosystems, an individual sambar deer was estimated to disperse on average 816 (±193) seeds per day during the study period, of which 652 (±176) were exotic. Synthesis and applications. Our results suggest that native plant species comprise the majority of sambar deer diets in Australian ecosystems and that the introduced species is dispersing both native and exotic plant species via endozoochory. However, exotic species seedling germination numbers were significantly higher in alpine ecosystems, and given the large daily movements of sambar deer, represents a significant vector for the spread of exotic plant species. Management of native plant species and vegetation communities of conservation significance, or at risk to sambar deer browsing is of high priority, through either the removal of sambar deer or implementation of exclusion‐based methods. [ABSTRACT FROM AUTHOR]

Ducks and geese are little studied dispersal vectors for plants lacking a fleshy fruit, and our understanding of the traits associated with these plants is limited. We analyzed 507 faecal samples of mallard (Anas platyrhynchos) and Canada goose (Branta canadensis) from 18 natural and urban wetlands in England, where they are the dominant resident waterfowl. We recovered 930 plant diaspores from 39 taxa representing 18 families, including 28 terrestrial and five aquatic species and four aliens. Mallards had more seeds and seed species per sample than geese, more seeds from barochory and hydrochory syndromes, and seeds that on average were larger and from plants with greater moisture requirements (i.e., more aquatic). Mallards dispersed more plant species than geese in natural habitats. Plant communities and traits dispersed were different between urban (e.g., more achenes) and natural (e.g., more capsules) habitats. Waterfowl can readily spread alien species from urban into natural environments but also allow native terrestrial and aquatic plants to disperse in response to climate heating or other global change. Throughout the temperate regions of the Northern Hemisphere, the mallard is accompanied by a goose (either the Canada goose or the greylag goose) as the most abundant waterfowl in urbanized areas. This combination provides a previously overlooked seed dispersal service for plants with diverse traits. [ABSTRACT FROM AUTHOR]

The expansion of alien plant species is of global concern, yet our understanding of their dispersal mechanisms is limited. Here we address the potential of alien plant seeds to disperse via ingestion, transport and egestion in waterfowl (endozoochory). Based on their general rapid expansions, we expected alien plant species to have several advantages for endozoochory compared to native plant species. We hypothesised that seeds of alien species would have higher passage rate, longer gut retention times, higher germinability after gut passage and shorter time‐to‐germination after egestion by waterfowl.In order to test our hypotheses, we compared the endozoochorous dispersal ability of six pairs of congeneric alien and native wetland plant species in a feeding experiment with mallards (Anas platyrhynchos). We focused on differences in seed survival, gut retention time, germinability and time‐to‐germination. In the analyses we corrected for seed shape and volume as these seed traits are known to have important effects.With gut passage, alien species had higher passage rates and germinated slower, whereas native species had shorter retention times and greater germinability. Controlling for seed traits did not alter these conclusions, but seed traits affected all aspects of the endozoochory process. This suggests that alien species may have particular traits correlated with a higher endozoochory potential. Among control seeds, alien seeds germinated faster and their germinability was higher than natives. Seed traits explained differences in germinability and time‐to‐germination in control seeds.Seeds of alien plant species have traits that correlate with successful endozoochory. This may provide alien species with a competitive advantage over native plant species by ensuring higher endozoochory rates in new environments, potentially enabling their rapid expansions. Our study underlines the important role of seed traits in the endozoochory potential of alien and native plant species, notably through their influence on retention time and germination. [ABSTRACT FROM AUTHOR]

Herbivores shape plant invasions through impacts on demography and dispersal, yet only demographic mechanisms are well understood. Although herbivores negatively impact demography by definition, they can affect dispersal either negatively (e.g., seed consumption), or positively (e.g., caching). Exploring the nuances of how herbivores influence spatial spread will improve the forecasting of plant movement on the landscape. Here, we aim to understand how herbivores impact how fast plant populations spread through varying impacts on plant demography and dispersal. We strive to determine whether, and under what conditions, we see net positive effects of herbivores, in order to find scenarios where herbivores can help to promote spread. We draw on classic invasion theory to develop a stage-structured integrodifference equation model that incorporates herbivore impacts on plant demography and dispersal. We simulate seven herbivore "syndromes" (combinations of demographic and/or dispersal effects) drawn from the literature to understand how increasing herbivore pressure alters plant spreading speed. We find that herbivores with solely negative effects on plant demography or dispersal always slow plant spreading speed, and that the speed slows monotonically as herbivore pressure increases. However, we also find that plant spreading speed can be hump shaped with respect to herbivore pressure: plants spread faster in the presence of herbivores (for low herbivore pressure) and then slower (for high herbivore pressure). This result is robust, occurring across all syndromes in which herbivores have a positive effect on plant dispersal, and is a sign that the positive effects of herbivores on dispersal can outweigh their negative effects on demography. For all syndromes we find that sufficiently high herbivore pressure results in population collapse. Thus, our findings show that herbivores can speed up or slow down plant spread. These insights allow for a greater understanding of how to slow invasions, facilitate native species recolonization, and shape range shifts with global change. [ABSTRACT FROM AUTHOR]

The dung seed bank (i.e. vegetation renewal) and greenhouse gas (GHG, e.g. carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O), which contributing to global warming) emissions (i.e. materials circulation) are two of the most important ecological functions of livestock dung in grassland ecosystems. With decomposition of faeces, dung seedling density (e.g. increase) and GHG emission (e.g. decrease) changes proceeded simultaneously. Although these two processes have so far been studied in isolation, almost no studies have explored the effects of dung seedlings on GHG emissions.We collected the dung of three livestock species—yak, sheep and horse—during late April (i.e. early warm season) in an alpine meadow in the northeastern Qinghai‐Tibetan Plateau (QTP). The livestock dung seedling composition and GHG fluxes were determined on Days 1, 6, 11, 16, 21, 26, 31, 37, 53 and 60 of germination, and we used frozen and ground faeces as the control group.The average seedling density of horse dung was significantly greater than that of yak and sheep. The GHG fluxes in the collected native dung samples were significantly greater than those of the dung controls, and the fluctuations in CO2, CH4 and N2O all showed a pattern of yak dung > horse dung > sheep dung. Over the course of the experiment, the GHG fluxes of both the experimental and control groups gradually decreased; however, the GHG emissions of the experimental group declined faster than those of the control group.There was a significant negative correlation between dung seedling density and GHG fluxes. Dung seedlings provide a reference for estimation of GHG emissions from livestock faeces.Synthesis and applications. Our research has revealed the features of and interactions between two important ecological functions of livestock dung in the grazing system. Furthermore, dung seedlings could substantially affect GHG fluxes and emission rates of livestock dung patches. The contributions of dung seeds to GHG emissions should be considered when estimating global warming potential from grazing livestock on the QTP for the purposes of determining national and regional land use policies and compiling global GHG inventories. [ABSTRACT FROM AUTHOR]

The European rabbit plays a major role in seed dispersal in its native environment. We evaluated its ecological interactions with plants through the dispersal of seeds by endozoochory in two invaded arid ecotones of Argentina. We found 855 whole seeds in 1283 fecal pellets, belonging to one non-native (Sesuvium portulacastrum) and five native plants (Arjona sp., Fabiana denudata, Frankenia juniperoides, Lycium chilensis, Poa sp.). Our results indicate that the European rabbit is a legitimate disperser of F. juniperoides, L. chilensis, and S. portulacastrum by the consumption and dissemination of viable seeds. Contrastingly, the rabbit is an illegitimate disperser of Arjona sp., F. denudata, and Poa sp. Our study identifies new interactions between an invasive herbivore and sympatric plants in the arid ecosystems of Argentina. We put forth that the mutualistic interaction between the European rabbit and both native and non-native plants highlights the complexity of trophic networks in invaded environments. [ABSTRACT FROM AUTHOR]

Aim: How plants can disperse in response to global change is a critical question, yet major knowledge gaps persist about long‐distance dispersal (LDD) mechanisms. We studied the potential a migratory waterfowl has for LDD of flowering plants via gut passage of seeds (endozoochory), comparing spring and autumn migration. Location: United Kingdom and Iceland. Taxon: Pink‐footed Goose (Anser brachyrhynchus, Baillon) and Angiosperma. Methods: We studied endozoochory by Pink‐footed geese migrating within and between the UK and Iceland by faecal sampling and GPS tracking. We collected 614 faecal samples from 14 areas in the UK and one in Iceland. Using GPS tracks to and from these areas, we estimated how far seeds can be dispersed by Pink‐footed geese, and where to or from. Results: We recorded 5507 intact seeds of 35 species (27 terrestrial) from 15 plant families, with lower seed abundance per dropping when birds were migrating northwards in the UK during spring than upon their arrival in autumn. Species richness of plant seeds was highest in Iceland and in autumn. Only four plant species dispersed had an "endozoochory syndrome". GPS movements showed that seeds retained in guts for up to 24 h can be readily dispersed in both directions between the UK and Iceland, with maximum distances exceeding 2000 km, as well as between UK localities separated by 100 s of km. Movements northwards of ≤400 km were even recorded in autumn. While at stopover sites, daily movements between roost and feeding sites often exceed 20 km. Main Conclusions: Pink‐footed geese are LDD vectors for plants previously assumed to lack an LDD mechanism. Spring migration is not the only period when geese move plants to cooler latitudes. The pink‐footed goose can allow terrestrial and aquatic plants to cross the ocean and to keep pace with climate change. [ABSTRACT FROM AUTHOR]

Migratory waterbirds disperse a broad range of angiosperms by endozoochory (seed dispersal via gut passage), especially plants in coastal wetlands. However, there is no previous information about the capacity of seeds to remain in the seed bank after waterbird endozoochory, and very little about how wetland salinity can influence the effect of gut passage on germination. We collected seeds of Juncus subulatus (Juncaceae), Bolboschoenus maritimus, and Schoenoplectus litoralis (Cyperaceae) from Doñana marshes in Spain. All three species are considered to have physiological dormancy. After gut passage following ingestion by ducks, seeds were stored in darkness in solutions with six different conductivities (1, 2, 4, 8, 16, and 32 dSm-1), for periods of 1, 6, or 12 months to simulate presence in a seed bank. After storage, 1800 seeds of each plant species assigned to these treatments were subjected to germination tests in demineralized water, together with 1800 control seeds that had not been ingested before storage. All species germinated readily after storage, with or without gut passage beforehand. Storage time and salinity both had important effects on germinability and time to germination, which differed between control and ingested seeds, and between plant species. After ≥6 months, germinability of Cyperaceae was enhanced by gut passage (≤25% higher than control seeds) at some salinities. Only J. subulatus showed consistently lower germinability after passage (≤30%). Only B. maritimus showed consistently slower germination after passage (≤33%). Salinity effects were more complex after passage, but increasing salinity did not generally have a negative impact on germination of ingested seeds. When compared to additional seeds that had not been stored before germination tests, storage reduced germinability in J. subulatus (≤39% reduction), but increased it in B. maritimus (≤17%) and S. litoralis (≤46%). Seeds dispersed by waterbird endozoochory may be easily incorporated into wetland seed banks, where they can remain halotolerant and delay germination until conditions become suitable. This can benefit wetland plants by increasing rates of long-distance dispersal, gene flow, and establishment of new populations. Avian gut passage can have positive and species-specific effects on germination in plants with persistent seed banks and/or physiological dormancy.