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Seed dispersal by vertebrates is fundamental for the persistence of plant species, forming networks of interactions that are often nested and modular. Networks involving angiosperms and frugivorous birds are relatively well-studied in the Neotropical region, but there are no previous studies of networks involving waterbirds. Here, we describe the structure of a Neotropical waterfowl seed-dispersal network and identify the species that have an important role for the network structure. We used information on 40 plant taxa found in fecal samples of five common waterfowl species to calculate the nestedness (NODF), weighted nestedness (WNODF), modularity, and weighted modularity of the network. We found that the network was nested, with yellow-billed teal showing the highest contribution both to nestedness and weighted nestedness. Twenty-four plant species contributed positively to weighted nestedness, with Salzmann's mille graines presenting the highest influence both to nestedness and weighted nestedness. The network was modular, but the weighted modularity was not significant. These results need to be considered with caution due to incomplete interaction sampling for two species. Ringed teal, Brazilian teal, and yellow-billed teal were considered hub modular species. Among plants, beak sedges and water snowflake were considered modular hub species, while Salzmann's mille graines and spikerush were network connectors. The structure of this Neotropical waterbird seed-dispersal network differed from the only previous waterfowl network study, from Europe, which found similar level of nestedness but no significant modularity. We include several possible explanations for this discrepancy and identified priorities for future research into waterbird–plant interaction networks. GGS received a postdoctoral fellowship from CNPq (Grant No. 150887/2022-1). MAP (Grant No. 304742/2019-8), LB (311409/2018-0) and LM thank the Brazilian Council for Development of Science and Technology (CNPq) for Research Productivity Grants. AJG was supported by Spanish National Plan projects CGL2016-76067-P (AEI/FEDER, EU) and PID2020-112774GB-I00/AEI/10.13039/501100011033. ESG received the grant RYC2019-027216-I funded by MCIN/AEI/10.13039/501100011033 and by ESF Investing in your future. This research was supported by funds from CNPq (Grant No. 52370695-2).

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.

Omnivorous waterbirds play an important role in aquatic ecosystems as dispersal vectors via direct ingestion, transportation, and egestion of plant and invertebrate propagules (i.e. endozoochory). Predatory birds also have the potential to disperse plants and invertebrates that were first carried internally or externally by their prey animals. However, the potential contribution of predatory waterbird species to propagule dispersal in aquatic ecosystems remains understudied. We chose the grey heron Ardea cinerea (Ardeidae) to study the potential of predatory waterbirds to disperse propagules within and among aquatic ecosystems. We hypothesised that: (1) herons disperse a wide variety of plant and invertebrate propagules, from different habitats, with different morphologies (i.e. dispersal syndromes), and including both native and alien species; (2) propagules are ingested with prey species that are primary dispersal vectors (i.e., herons are secondary dispersers); (3) heron pellets show a similar abundance and richness of propagules across their widespread range. We collected 73 regurgitated heron pellets containing undigestible remains from 12 locations across the U.K. and The Netherlands, and examined the taxonomic diversity of plant seeds, invertebrates and prey remains. Pellets were dominated by mammal hairs (99% by volume), and bones confirmed the ingestion of small mammals (prevalence of 38%, e.g. water voles Arvicola amphibius), fish (14%), and birds or amphibians (6%). A total of 266 intact plant seeds were recovered from 71% of the pellets, representing 50 taxa from 17 plant families, including the alien Cotula coronopifolia. The cumulative number of plant species dispersed was lower at higher latitudes. Eight plant species recorded had not previously been recorded as dispersed via waterbirds, and only three species have an endozoochorous dispersal syndrome. Plant taxa were dominated by Caryophyllaceae, Cyperaceae, Juncaceae, and Poaceae, with 24 species from the littoral zone (Ellenberg moisture values of 7–12) and 21 terrestrial species (Ellenberg moisture values of 4–6). Intact invertebrate propagules were found in 30% of the pellets, dominated by Cladocera (Daphniidae) and Bryozoa (including the alien Plumatella casmiana). Our results demonstrate that grey herons disperse plant seeds and aquatic invertebrates widely in north-western Europe. Herons regurgitate pellets that contain plant and invertebrate propagules from both aquatic or terrestrial habitats, for which secondary dispersal via ingestion along with prey is the likely underlying mechanism (i.e. propagules either attached to or in the digestive systems of the various prey). Our findings showcase the potential of predatory waterbirds as vectors of plants and invertebrates, and how they may facilitate connectivity between freshwater and terrestrial habitats.

Funder: Brazilian Program for Biodiversity Research, Funder: Fundação de Amparo à Pesquisa do Estado do Amazonas; Id: http://dx.doi.org/10.13039/501100004916, Funder: National Institute for Amazonian Biodiversity, Aim: To investigate the geographic patterns and ecological correlates in the geographic distribution of the most common tree dispersal modes in Amazonia (endozoochory, synzoochory, anemochory and hydrochory). We examined if the proportional abundance of these dispersal modes could be explained by the availability of dispersal agents (disperser‐availability hypothesis) and/or the availability of resources for constructing zoochorous fruits (resource‐availability hypothesis). Time period: Tree‐inventory plots established between 1934 and 2019. Major taxa studied: Trees with a diameter at breast height (DBH) ≥ 9.55 cm. Location: Amazonia, here defined as the lowland rain forests of the Amazon River basin and the Guiana Shield. Methods: We assigned dispersal modes to a total of 5433 species and morphospecies within 1877 tree‐inventory plots across terra‐firme, seasonally flooded, and permanently flooded forests. We investigated geographic patterns in the proportional abundance of dispersal modes. We performed an abundance‐weighted mean pairwise distance (MPD) test and fit generalized linear models (GLMs) to explain the geographic distribution of dispersal modes. Results: Anemochory was significantly, positively associated with mean annual wind speed, and hydrochory was significantly higher in flooded forests. Dispersal modes did not consistently show significant associations with the availability of resources for constructing zoochorous fruits. A lower dissimilarity in dispersal modes, resulting from a higher dominance of endozoochory, occurred in terra‐firme forests (excluding podzols) compared to flooded forests. Main conclusions: The disperser‐availability hypothesis was well supported for abiotic dispersal modes (anemochory and hydrochory). The availability of resources for constructing zoochorous fruits seems an unlikely explanation for the distribution of dispersal modes in Amazonia. The association between frugivores and the proportional abundance of zoochory requires further research, as tree recruitment not only depends on dispersal vectors but also on conditions that favour or limit seedling recruitment across forest types.

How flightless animals disperse to remote oceanic islands is a key unresolved question in biogeography. The flightless Pachyrhynchus weevils represent repetitive colonization history in West Pacific islands, which attracted our interests about how some weevils have successfully dispersed in the reverse direction against the sea current. Here, we propose endozoochory as a possible mechanism that the eggs of the weevils might be carried by embedded in the fruits as the food of frugivorous birds. In this study, Pachyrhynchus eggs were embedded in small pieces of persimmon fruits (Diospyros kaki) and fed to captive frugivorous birds. After digestion, 83%–100% eggs were retrieved from the feces of a bulbul (Hypsipetes leucocephalus) and two thrushes (Turdus chrysolaus). The retrieved eggs had hatching rates higher than 84%, which were not different from the control. In contrast, no egg was retrieved from the feces of the frugivorous pigeon (Treron sieboldii), which took a longer retention time in the guts. Our study identified that the eggs of Pachyrhynchus weevils are possible to be transported by internal digesting in some bird species., How flightless weevils disperse to remote oceanic islands is a fantastic issue in biogeography. Our study identified that the eggs of the weevils are possible to be transported by internal digesting in some frugivorous birds. ​

Despite being one of the most diverse families, scant attention has been paid to the seed dispersal system in Orchidaceae, owing to the widely accepted notion that wind dispersal is the dominant strategy. However, the indehiscent fruits, with seeds immersed in fleshy tissue, evoke the possibility of endozoochory in Apostasioideae, the earliest diverging lineage of orchids. In the present study, I investigated the seed dispersal system of Apostasia nipponica by direct observation, time-lapse photography, and investigation of the viability of seeds passing through the digestive tract of orthopterans. This study revealed a previously undocumented seed dispersal system in A. nipponica, in which the cricket, Eulandrevus ivani, and the camel cricket, Diestrammena yakumontana, consume the fruit and defecate viable seeds. Orthopterans are rarely considered seed dispersers, but the gross fruit morphology and pigmentation patterns of some Apostasia species parallel those seen in A. nipponica, suggesting that similar seed dispersal systems could be widespread among Apostasia species. Whether seed dispersal by orthopteran frugivores is common in Apostasioideae warrants further investigation.

We tested if seed germination of the tree Cecropia pachystachya is induced after being consumed by the frugivorous bat Carollia perspicillata. We carried out two essays; in the first, we compared germination of seeds from C. perspicillata feces, seeds chemically scarified in acid solutions, and seeds from fresh C. pachystachya infructescences. In the second essay we evaluated seed viability after four months storage. Percentage of seed germination was reduced after C. perspicillata ingestion, but nevertheless remained high (76%). Seeds ingested by this bat also maintain their viability after storage, and C. perspicillata can be considered an effective seed disperser of C. pachystachya seeds. Resumo Desenvolvemos dois experimentos para testar se C. perspicillata pode agir como indutor da germinação de sementes de C. pachystachya. Primeiro comparamos a germinação das sementes retiradas das fezes do morcego, de sementes tratadas em soluções ácidas e sementes retiradas de infrutescências da planta. Avaliamos também a germinação das sementes após 4 meses de armazenamento. A porcentagem de germinação das sementes foi reduzida após a ingestão do morcego, entretanto, ainda com altas taxas de germinação (76%). Sementes ingeridas pelo morcego mantiveram sua viabilidade após o período de armazenamento. C. perspicillata pode ser considerado um dispersor eficiente de sementes de C. pachystachya.

Seed dispersal is an important ecological process in wetland ecosystems and helps maintain community structure and ecosystem biodiversity. Waterbird-mediated endozoochory is an effective and feasible dispersal mechanism for wetland plants; however, the influence of vectors and seed traits on this mechanism remains unclear. To investigate the effects of vector species and seed traits (length and lignin) on retention time, retrieval and germination of gut-surviving seeds, we fed Baikal teals (Anas formosa) and green-winged teals (Anas crecca) eight common plant seeds (Polygonum aviculare, Rumex dentatus, Polygonum orientale, Vallisneria natans, Ranunculus polii, Polygonum hydropiper, Carex cinerascen and Euphrasia pectinata) in the Shengjin Lake wetland (a Ramsar site). We collected fecal samples at intervals of 2–6 h for 36 h, and found that the percentage of recovered seeds differed significantly among teal and plant species (3%~30%); 94% of viable seeds were recovered within 12 h after feeding. Moreover, the germination rate of the recovered seeds (25%~56%) was higher than that of the control. The seed retention time was affected by seed lignin and disperser species; higher lignin made digestion difficult with higher retrieval. Smaller seeds passed through the guts but had no significant effect on recovered seeds. Seed length and disperser species showed no significant correlation with germination. These findings suggested endozoochory by dabbling ducks as an effective wetland seed dispersal mechanism.

During the last 24 years, the mutualistic interaction between the dead horse arum, Helicodiceros muscivorus, and the Balearic lizard, Podarcis lilfordi, was studied on Aire Island (Balearic Islands, Spain). From a small population of a hundred plants, the dead horse arum expanded extraordinarily throughout the island, reaching the highest known densities of the species and occupying areas of the island where it was not previously present. The current abundance of plants is a direct effect of the frugivorous activity of the Balearic lizard, which is the main, if not the only, effective seed disperser of the plant on Aire Island. However, abiotic factors predominated over biotic factors in driving abundance of plants. Over the years, plant densities varied significantly depending on the aridity of the island, with higher densities recorded in drier years. Lizards’ frugivorous activity and dispersal intensity was inversely correlated with annual rainfall. We found higher dispersal intensity in years with lower rainfall. We propose that the years of lower rainfall are those in which there is a lower prey availability. In such years, lizards compensate the shortage of other trophic resources with a more intense consumption of dead horse arum fruits. The mutualistic interaction is therefore asymmetric, since there is a greater influence of the frugivorous activity of the lizards on the plants than of the plants on lizards. It is, in short, a system chronically out of balance.

A 12-month survey of cyanogenesis in the fruits and leaves of Nandina domestica (Berberidaceae) revealed that the leaves are strongly cyanogenic throughout the year, as measured by the Feigl-Anger test. Fruits vary in their cyanogenic potential depending on the degree of ripeness and clone. Green fruits are strongly and rapidly cyanogenic, but most ripe fruits are weakly and slowly cyanogenic. Some fruits tested negative for cyanide. Fruits tend to become less cyanogenic the longer they remain on the plant, so birds feeding on older fruits are at a lower risk of poisoning. More than other frugivores, cedar waxwings (Bombycilla cedrorum) are susceptible to cyanide poisoning by N. domestica fruits because of their tendency to consume large numbers of fruits in a single feeding bout. Hypotheses are examined that might explain the evolutionary paradox of why N. domestica fruits are toxic to potential dispersers.

Endozoochory refers to the ingestion of fruits by animals and their release through faeces as it plays an important role in maintaining ecosystems. This study aimed to evaluate the effect of cumbaru fruit ingestion by tapirs on cumbaru seed germination. In latrines, fruits ingested and excreted by the animals were collected among tapir faeces. The collected material was taken to the laboratory and placed in a germination chamber and greenhouse. Fruit ingestion and its subsequent defection propitiated a higher germination rate and germination vigor compared to non-ingested fruit grown in a germination chamber and greenhouse. The results in the greenhouse showed that the latrines (decomposing organic matter) have different thermal conditions (higher temperature), in relation to the external environment, which probably accelerated the germination processes. Evaluation of seedlings grown in a greenhouse and germination chamber indicated that growth in the greenhouse was better due to humic and fulvic acids, which are common stimulants of seed germination and seedling growth. Therefore, tapirs can be considered good fruit dispersers and inducers of cumbaru seeds germination.

Seed dispersal by animals is important for the ecology of plants. It is particularly important to understand which animals are involved and how they move seeds differently from one another. Some seed dispersers are understudied despite ample evidence they consume fruits and seeds. This includes animals commonly referred to as carnivores in the order Carnivora. The overall goal of my dissertation was to describe the extent and quality of seed dispersal by Carnivorans, estimate important aspects of seed dispersal for a specific Carnivoran, the coyote, and estimate how differences between a coyote and songbirds affect where plants will occur in the future and if that changes how much carbon plants store on the landscape. To achieve these goals, I first systematically reviewed existing research on frugivory and seed dispersal by Carnivorans. Then, I experimentally evaluated how long it takes for seeds to pass through a coyote, and if the consumption of seeds by coyotes negatively affects seed germination or viability. Finally, I modeled how differences in the distances that coyotes carry seeds vs. songbirds affect plant migrations. I found that Carnivoran frugivory and seed dispersal are common, involve many plant species, and occur worldwide across most ecosystems. Carnivorans also rarely damage seeds or hamper seed viability and germination when they consume and disperse seeds. Furthermore, I found that coyotes generally take between 4 and 24 hours to pass seeds from the fruits they consumed and deliver seeds to new landscapes without harming them. Given coyote travel speeds, these results suggest that coyotes regularly disperse seeds up to 5 km. This seed dispersal distance is substantially greater than songbirds and led to a 2.5 times larger expansion of where junipers grew in models extending 80 years into the future. Coyote seed dispersal would also result in 3.4 times greater conversion of grasslands and an increase total biotic carbon storage by 1.1 Pg. My findings show that understudied seed dispersers like Carnivorans can greatly impact plant ecology and ecosystem services and highlights the need for further studies on the impacts of Carnivora on seed dispersal.

Although many plants are dispersed by wind and seeds can travel long distances across unsuitable matrix areas, a large proportion relies on co‐evolved zoochorous seed dispersal to connect populations in isolated habitat islands. Particularly in agricultural landscapes, where remaining habitat patches are often very small and highly isolated, mobile linkers as zoochorous seed dispersers are critical for the population dynamics of numerous plant species. However, knowledge about the quali‐ or quantification of such mobile link processes, especially in agricultural landscapes, is still limited. In a controlled feeding experiment, we recorded the seed intake and germination success after complete digestion by the European brown hare (Lepus europaeus) and explored its mobile link potential as an endozoochoric seed disperser. Utilizing a suite of common, rare, and potentially invasive plant species, we disentangled the effects of seed morphological traits on germination success while controlling for phylogenetic relatedness. Further, we measured the landscape connectivity via hares in two contrasting agricultural landscapes (simple: few natural and semi‐natural structures, large fields; complex: high amount of natural and semi‐natural structures, small fields) using GPS‐based movement data. With 34,710 seeds of 44 plant species fed, one of 200 seeds (0.51%) with seedlings of 33 species germinated from feces. Germination after complete digestion was positively related to denser seeds with comparatively small surface area and a relatively slender and elongated shape, suggesting that, for hares, the most critical seed characteristics for successful endozoochorous seed dispersal minimize exposure of the seed to the stomach and the associated digestive system. Furthermore, we could show that a hare's retention time is long enough to interconnect different habitats, especially grasslands and fields. Thus, besides other seed dispersal mechanisms, this most likely allows hares to act as effective mobile linkers contributing to ecosystem stability in times of agricultural intensification, not only in complex but also in simple landscapes., The most critical seed characteristics for successful endozoochorous seed dispersal by hares minimize exposure of the seed to the digestive system (i.e., dense seeds with a small surface area and an elongated shape). In addition, a hare's retention time is long enough to interconnect different habitats, especially grasslands and crop fields. Thus, besides other seed dispersal mechanisms, this most likely allows hares to act as effective mobile linkers contributing to ecosystem stability during times of agricultural intensification.

We conducted a two-year study in New Mexico, USA, to determine the role of birds, lagomorphs, mesocarnivores, and porcupines in one-seed juniper (J. monosperma (Englem.) Sarg.) seed dispersal. We established random plots: (1) around cone-bearing juniper trees in the woodland; (2) around non-cone-bearing juniper trees in the woodland; (3) in woodland/grassland transition zones; and in (4) grassland habitats near juniper woodlands. We estimated seed density and tallied the number of plots with seeds deposited by each disperser group. Birds deposited the highest number of seeds/ha under the canopy of cone-bearing trees. Mesocarnivores were responsible for the highest average seed deposition in all other habitats. In juniper cone-bearing tree habitats, birds deposited seeds in 100% of plots under-canopy and 93% of plots outside-canopy. Seeds deposited by lagomorphs were observed in plots across all sampled habitats. Overall, seed deposition was greatest around cone-bearing juniper trees, followed by non-cone-bearing juniper trees, woodland-grassland transition zones, and grassland habitats. Birds deposited seeds primarily under cone-bearing tree perch sites. Lagomorphs deposited seeds widely in relatively high numbers across all habitats and are likely responsible for the greatest number of one-seed juniper seeds deposited on the soil surface at our research site.

Members of the order Carnivora are a unique and important seed disperser who consume and deposit undamaged seeds while providing regular long‐distance seed dispersal opportunities. Some members of Carnivora, such as coyotes (Canis latrans), are undergoing range expansions which may help the plant species they consume colonize new locations or replace dispersal services provided by recently extirpated species. In this study, we evaluated aspects of the seed dispersal effectiveness of coyotes and gut passage time to determine the potential dispersal distances for three commonly consumed and commonly occurring plant species (Amelanchier alnifolia, Celtis ehrenbergiana, and Juniperus osteosperma). We also investigated the potential effects of secondary dispersal of seeds away from scats by comparing seedling emergence from whole scats to those where seeds were first removed from scats. We found that seeds generally took between 4 and 24 h to pass through the digestive tract of coyotes, which could result in regular seed dispersal up to 7 km. Gut passage through coyotes had no effect on seed viability or emergence for any of the three plant species, including that gut passage for A. alnifolia and J. osteosperma does not replace cold stratification for breaking physiological dormancy. By simulating secondary dispersal, we found that 22% (±8.2%) more C. ehrenbergiana seedlings emerged when seeds were removed from scats and those seedlings emerged 7 d earlier (±5 d) compared to seeds that remained in the coyote scat. Coyotes are effective seed dispersers, with the potential for regular long‐distance dispersal services and for providing opportunities for secondary seed dispersal, which could aid in climate migration or serve to replace extirpated dispersal mutualists.

Epi- and endozoochory are well explored phenomena that contribute to the distribution patterns of plant seeds, spores or fruits by vertebrates. It is less known how soil algae may be redistributed due to analogous interactions. We describe the participation of saprophage Rossiulus kessleri (Julidae, Diplopoda) in soil algae redistribution. The research was carried out in a park area of a large industrial megacity in the Dnipro steppe region (Ukrainian North Steppe subzone), Dnipro city. Research material was collected according to zoological and algological methods. The leaf litter of tree species (ash maple Acer negundo, Italian poplar Populus deltoides, locust Robinia pseudoacacia, European white elm Ulmus laevis, Norway maple Acer platanoides) and living diplopods were collected within 1 m2 quadrats along transects. Millipedes were maintained without feeding for five days, after which they were fed with the litter collected previously. Identification of algoflora was conducted in washes from the surface of diplopods’ bodies, its gut washes, in diplopods’ excretions and leaf litter washes. In the leaf litter, as R. kessleri’s feeding base, 14 soil algae species were identified—Nostoc punctiforme, Bracteacoccus minor, Mychonastes homosphaera, Neospongiococcum sp., Chlamydomonas sp., Chlorella vulgaris, Stichococcus bacillaris, Pseudococcomyxa simplex, Desmococcus olivaceus, Trebouxia spp., Klebsormidium flaccidum, Nephrodiella phaseolus, Navicula pelliculosa and Vischeria magna. In body surface washes, five soil algae species were identified, in gut washes seven algae species were found, while in excretions, just four were observed. It was established that not all algae species from the gut washes of R. kessleri were present in excretions. The presence of some representatives of soil algae, for example, Chlorella vulgaris and Vischeria magna, in natural park litter, body surface washes, gut washes and in excretions of R. kessleri, suggests that the given species pass through the diplopod’s gut and stay undamaged. This indicates that soil saprophage R. kessleri contributes to the dispersal of some soil algae representatives through epi- and endozoochory. Our results represent novel contributions to the knowledge of zoochory. It was shown that animals can be involved in dispersal of not only plant parts but also entire organisms, although it is unknown at what scale diplopods contribute to the diversity and dispersal of algae.

Megafaunal extinctions are recurring events that cause evolutionary ripples, as cascades of secondary extinctions and shifting selective pressures reshape ecosystems. Megafaunal browsers and grazers are major ecosystem engineers, they: keep woody vegetation suppressed; are nitrogen cyclers; and serve as seed dispersers. Most angiosperms possess sets of physiological traits that allow for the fixation of mutualisms with megafauna; some of these traits appear to serve as exaptation (preadaptation) features for farming. As an easily recognized example, fleshy fruits are, an exaptation to agriculture, as they evolved to recruit a non-human disperser. We hypothesize that the traits of rapid annual growth, self-compatibility, heavy investment in reproduction, high plasticity (wide reaction norms), and rapid evolvability were part of an adaptive syndrome for megafaunal seed dispersal. We review the evolutionary importance that megafauna had for crop and weed progenitors and discuss possible ramifications of their extinction on: (1) seed dispersal; (2) population dynamics; and (3) habitat loss. Humans replaced some of the ecological services that had been lost as a result of late Quaternary extinctions and drove rapid evolutionary change resulting in domestication. Introduction Lost Seed-Dispersal Services - Small-Seeded Grains and Legumes - Large Fleshy Fruiting Plants Loss of Herbivory and Disturbance Regimes Plant Domestication - Exaptation Traits Supporting Domestication Anthropogenic Ecosystem Services

Background Seed dispersal allows plants to colonize new habitats that has an significant influence on plant distribution and population dynamics. Orchids produce numerous tiny seeds without endosperm, which are considered to be mainly wind-dispersed. Here, we report avian seed dispersal for an early diverging orchid species, Neuwiedia singapureana, which produces fleshy fruits with hard seed coats in the understory of tropical forests. Results Neuwiedia singapureana produced fleshy fruits that turned red in autumn, and birds were confirmed to be the primary seed dispersers. As compared to its sister species, N. veratrifolia with dehiscent capsular fruits, embryos of N. singapureana were larger and enclosed by thickened and lignified seed coats. After passing through the digestive tracts of birds, the seeds still stayed alive, and the walls of seed coat contained several cracks. The germination percentage increased significantly for digested seeds as compared with seeds from intact fruits. Conclusion The thickened and lignified seed coat may protect seeds as they passed through the digestive tracts of birds. Taken together with a recent report of insect-mediated seed dispersal system in the subfamily Apostasioideae, the animal-mediated seed dispersal may be an adaptive mechanism promoting the success of colonization in dark understory habitats.

BackgroundLong-distance seed dispersal (LDD) has strong impacts on the spatiotemporal dynamics of plants. Large animals are important LDD vectors because they regularly transport seeds of many plant species over long distances. While there is now ample evidence that behaviour varies considerably between individual animals, it is not clear to what extent inter-individual variation in behaviour alters seed dispersal by animals.MethodsWe study how inter-individual variation in the movement and feeding behaviour of one of Europe’s largest herbivores (the red deer,Cervus elaphus) affects internal seed dispersal (endozoochory) of multiple plant species. We combine movement data of 21 individual deer with measurements of seed loads in the dung of the same individuals and with data on gut passage time. These data serve to parameterize a model of passive dispersal that predicts LDD in three orientations (horizontal as well as upward and downward in elevation).With this model we investigate to what extent per-seed probabilities of LDD and seed load vary between individuals and throughout the vegetation period (May–December). Subsequently, we test whether per-seed LDD probability and seed load are positively (or negatively) correlated so that more mobile animals disperse more (or less) seeds. Finally, we examine whether non-random associations between per-seed LDD probability and seed load affect the LDD of individual plant species.ResultsThe studied deer dispersed viable seeds of at least 62 plant species. Deer individuals varied significantly in per-seed LDD probability and seed loads. However, more mobile animals did not disperse more or less seeds than less mobile ones. Plant species also did not differ significantly in the relationship between per-seed LDD probability and seed load. Yet plant species differed in how their seed load was distributed across deer individuals and in time, and this caused their LDD potential to differ more than twofold. For several plant species, we detected non-random associations between per-seed LDD probability and seed load that generally increased LDD potential.ConclusionsInter-individual variation in movement and feeding behaviour means that certain deer are substantially more effective LDD vectors than others. This inter-individual variation reduces the reliability of LDD and increases the sensitivity of LDD to the decline of deer populations. Variation in the dispersal services of individual animals should thus be taken into account in models in order to improve LDD projections.

Many plants and invertebrates rely on internal transport by animals for long-distance dispersal. Their dispersal capacity is greatly influenced by interactions with the animal's digestive tract. Omnivorous birds adjust their digestive tract morphology to seasonally variable diets. We performed feeding trials in waterfowl to unravel how changing organ size, in combination with seed size, affects dispersal potential. We subjected captive mallards to mimics of summer (animal-based), winter (plant-based), and intermediate diets, and analyzed gut passage of seeds before and after the treatment (trials 1 and 2). To test the effect of gut morphology on seed digestion, we measured digestive organ size after euthanasia. Three hours before euthanasia, differently sized seeds were fed to test how seed size affects gut passage by determining their relative position in the digestive tract (trial 3). Trials 1 and 2 showed that intact seed passage was lower in the plant-based than in the animal-based diet group. Retention time changed only within groups, decreasing in the animal-based, and increasing in the plant-based diet group. No post-diet differences in organ size were detected, probably due to large between-individual variation within groups. Digestive tract measures did not explain variation in seed survival or retention time. Trial 3 revealed that small seeds pass the digestive tract rapidly, while large seeds are retained longer, particularly in the gizzard. Differential retention in the gizzard, the section where seeds can be destroyed, is likely why larger seeds have a lower probability to pass the digestive tract intact. Our results confirm that rapid, flexible adaptation to diet shifts affects seed digestion in waterfowl, although we could not conclusively relate this to organ size. Large interindividual variation in digestive efficiency between mallards feeding on the same diet may provide opportunities for seed dispersal in the field throughout the annual cycle.

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.

Premise of the Study: In animal‐dispersed plants, seed dispersal patterns of the same species in the same habitat can greatly differ among individuals and temporally. Many studies have revealed the pervasive effects of spatial variation in fruit abundance on seed dispersal of individual plants. By contrast, very few studies have investigated the temporal variation in seed dispersal. Methods: We investigated the effects of conspecific and community‐level fruit abundance on fruit removal rate and seed dispersal distance of six bird‐dispersed woody species in a Japanese temperate forest for 3 yr. The six species share similar fruit characteristics but have different fruiting seasons. In addition, we also considered the effects of bird seasonality in diet and species composition on seed dispersal. Key Results: Inter‐annual and inter‐seasonal variation in community‐level fruit abundance determined both fruit removal and the seed dispersal distance across species, but the effect differed with seasonal changes in bird behavior. Abundant fruit satiated fruit removal by birds only during fruit‐feeding periods, not during insect‐feeding periods. A scarcity of fruit increased the dispersal distance, but only during the migratory period. This difference was probably due to the different foraging behaviors of passing migrants and territorial birds. Conclusions: Our results illustrated that temporal variation in community‐level fruit abundance could be a determining factor for seed dispersal in temperate forests. The effects of temporal variations in fruit abundance on seed dispersal patterns should be investigated in other ecosystems in order to understand their importance and associations with frugivore phenology., 動物による種子散布メカニズムの一端を解明 --森の果実の豊凶が鳥のタネまきを左右する--. 京都大学プレスリリース. 2018-10-15.

Background and aims – Exploring the role of mammalian carnivores as seed dispersers in Mediterranean environments is crucial for understanding biotic interactions and preserving mutualistic networks in areas with high biodiversity. We examine the potential role of the Sardinian fox ( Vulpes vulpes subsp. ichnusae ) as a seed-disperser of two juniper species ( Juniperus phoenicea subsp. turbinata and J. oxycedrus subsp. macrocarpa ) in Mediterranean coastal environments.Methods – Observational and manipulative experiments were conducted in five coastal sites in north-western Sardinia (Italy) between 2010 and 2013.Key results – We found that Sardinian fox actively disperses seeds of J. phoenicea subsp. turbinata, whereas no evidence was obtained for the fox dispersing seeds of J. oxycedrus subsp. macrocarpa. Fox scat contained, on average, 73–86 J. phoenicea subsp. turbinata seeds, accounting 16.3–17.8 % of the average dung weight. The role of Sardinian fox as a primary disperser of J. phoenicea subsp. turbinata is by directly dispersing juniper seeds (via defecation) to a specific microhabitat (i.e. 80–90 % of dung was released on dwarf plants, mainly Helichrysum italicum subsp. microphyllum ), which positively affected the survival of emerged seedlings). We quantified that fox dispersed 30 to 100 seeds per day per hectare (3 500–10 500 seeds per hectare in one winter season).Conclusions – We reported that Sardinian fox is a direct disperser of J. phoenicea subsp. turbinata, thus playing a major role in secondary successional dynamics in Mediterranean coastal environments. Evolutionary implications are discussed, in that the positive interaction between Sardinian fox and J. phoenicea subsp. turbinata could be recent, following the introduction of fox to the Tyrrhenian islands during the 7 th –6 th millennium BC.

Fish have somehow colonized isolated water bodies all over the world without human assistance. It has long been speculated that these colonization events are assisted by waterbirds, transporting fish eggs attached to their feet and feathers, yet empirical support for this is lacking. Recently, it was suggested that endozoochory (i.e., internal transport within the gut) might play a more important role, but only highly resistant diapause eggs of killifish have been found to survive passage through waterbird guts. Here, we performed a controlled feeding experiment, where developing eggs of two cosmopolitan, invasive cyprinids (common carp, Prussian carp) were fed to captive mallards. Live embryos of both species were retrieved from fresh feces and survived beyond hatching. Our study identifies an overlooked dispersal mechanism in fish, providing evidence for bird-mediated dispersal ability of soft-membraned eggs undergoing active development. Only 0.2% of ingested eggs survived gut passage, yet, given the abundance, diet, and movements of ducks in nature, our results have major implications for biodiversity conservation and invasion dynamics in freshwater ecosystems.

Dung beetles form an insect group that fulfils important functions in terrestrial ecosystems throughout the world. These include nutrient cycling through dung removal, soil bioturbation, plant growth, secondary seed dispersal and parasite control. We conducted field experiments at two sites in the northern hemisphere temperate region in which dung removal and secondary seed dispersal were assessed. Dung beetles were classified in three functional groups, depending on their size and dung manipulation method: dwellers, large and small tunnelers. Other soil inhabiting fauna were included as a fourth functional group. Dung removal and seed dispersal by each individual functional group and combinations thereof were estimated in exclusion experiments using different dung types. Dwellers were the most diverse and abundant group, but tunnelers were dominant in terms of biomass. All dung beetle functional groups had a clear preference for fresh dung. The ecosystem services in dung removal and secondary seed dispersal provided by dung beetles were significant and differed between functional groups. Although in absolute numbers more dwellers were found, large tunnelers were disproportionally important for dung burial and seed removal. In the absence of dung beetles, other soil inhabiting fauna, such as earthworms, partly took over the dung decomposing role of dung beetles while most dung was processed when all native functional groups were present. Our results, therefore, emphasize the need to conserve functionally complete dung ecosystems to maintain full ecosystem functioning.

Summary Dispersal of plant seeds by ducks and other waterbirds is of great importance to the ecology of freshwater habitats. To unravel the mechanisms of waterbird-mediated seed dispersal, numerous laboratory experiments have been conducted, but effects of seed and waterbird traits on dispersal potential have rarely been investigated under field conditions. Through analysis of the digestive tracts of 100 wild mallards (Anas platyrhynchos) across a winter season in the Netherlands, we assessed (i) the inter-individual and seasonal variability of seeds in the digestive tract, (ii) the variability of digestive tract organ size and gizzard grit mass, and (iii) the potential effects of seed species traits and gut traits on the survival potential of ingested seeds. We found 4548 ingested seeds of at least 66 plant species from a wide range of habitats, most of which were retained in the gizzard. Nineteen species had not previously been reported from mallard diets. Individual tracts contained anywhere between 0 and 1048 seeds, of up to 14 species (median of three species). Diet composition and digestive tract size varied substantially between individuals and over the course of the winter season. As predicted from controlled feeding studies, we found that also in wild mallards, size-dependent gut passage survival favours the dispersal of small-seeded species. Despite the large variation in gizzard and small intestine size in this study, their effect on the dispersal potential of ingested seeds in the field remains unclear. We found no difference in dispersal potential between plants species growing in wet or dry habitats. This study demonstrates that wild mallards are opportunistic seed consumers with a very diverse diet as reflected by seed species composition in both the foregut and hindgut. However, we also show that serious limitations of field-based analyses compared to controlled experiments can impede drawing conclusions about gut passage survival of seeds. The large variability in diet composition among individuals and over time indicates high endozoochorous dispersal potential for a wide range of plant species by wild mallard in aquatic and wetland, as well as surrounding terrestrial habitats.

International audience; Herbivorous ungulates are key species in the functioning of terrestrial ecosystems, and their recent demographic and geographic expansion in some temperate regions is likely to influence ecological processes, particularly if we consider plants and the frequency of mobile links among plant populations. In forests, long-distance seed dispersal essentially relies on animal movements. Owing to their ability to cover daily long distances, large herbivorous ungulates are assumed to be important vectors for long-distance seed dispersal (hundreds to thousands of metres). To quantify patterns of seed dispersal, we analysed the paths of roe deer (Capreolus capreolus), red deer (Cervus elaphus) and wild boar (Sus scrofa) which had been equipped with GPS collars. We used seed gut passage times from a recent companion paper (Picard et al., Ecology and Evolution, 2015;5:2621–2632). We combined the animal movements and seed gut passage times to estimate seed dispersal curves for herbivorous ungulates. On average, the forest ungulates might disperse consumed seeds over distances greater than 2 km, during a 48-h path in the forest. Maximal dispersal distance was longer for red deer (3.5 km) and wild boar (3.2 km) than for roe deer (2.0 km). By combining high excretion rates and long dispersal distance, wild boar appears to be potentially an efficient seed disperser. Seed dispersal by common herbivorous ungulates is likely to play a crucial role in the today's context of rapid environmental changes such as global warming and landscape fragmentation. Differences in the distribution of seed gut passage times and seasonal distances covered underline the relevance of cross comparative approaches and the complementarity of herbivorous ungulates in long-distance seed dispersal.

Seeds dispersed by endozoochory by large herbivores are disseminated over long distances, which has potential impacts on vegetation dynamics within a region. The Korean water deer (Hydropotes inermis argyropus) is a dominant deer species in South Korea, but is considered a vulnerable species internationally. In the present study, we aimed to elucidate the role of Korean water deer in seed dispersal in lowland areas where land-use change is occurring. We assessed seasonal differences in seed dispersal in Korean water deer, and the traits of seeds dispersed in this manner. We identified species dispersed by controlled germination using Korean water deer fecal samples from the Civilian Control Zone adjacent to the Demilitarized Zone between South and North Korea, a mixed lowland habitat of forest and open area. A total of 208 fecal pellet groups were collected throughout the year from April 2017 to March 2018. We found a total of 35 plant species from the deer feces. More seeds and species were dispersed in fall, the main seeding period in South Korea. Dispersed plant species were mainly forbs that were specific to lowland areas and had medium (1–2 mm) seeds with no specific dispersal adaptations. Our results suggest that Korean water deer preferentially disperse particular plant species. They disperse disproportionately more graminoids, species from open areas, shorter than 2 mm, with no special adaptation, and specifically without fleshy/edible fruits, which may result in ecological filtering. We conclude that Korean water deer, as an endozoochorous seed dispersal vector in the lowland of South Korea, can affect plant populations and communities in the region.

Made available in DSpace on 2020-12-12T01:27:33Z (GMT). No. of bitstreams: 0 Previous issue date: 2020-07-01 Japan Society for the Promotion of Science Numerous studies have evaluated the qualitative and quantitative effectiveness of several animal species in each phase of the endozoochory. For the “seed germination and growth” phase, the ideal method of evaluation is to track the germination and seedling growth of seeds where they were dispersed by animals, but few studies have done so. Furthermore, because these studies were limited to measurements such as the seedling density and seedling growth over time or comparison with surrounding areas, survival rates, and factors specific to seedling mortality are unclear. This study aimed to qualitatively evaluate the effectiveness of one of the main seed dispersers in temperate forest ecosystems in Asia, raccoon dogs (Nyctereutes procyonoides), in the seed germination and growth phase. Seedlings of four tree species germinated in their latrines: Ginkgo biloba, Diospyros kaki, Aphananthe aspera, and Celtis sinensis. It was previously thought that most seeds in latrines germinated but died off quickly through competition within seedlings. However, this study has verified that in specific environmental conditions, seedlings of these four species live for at least one year after germination. In particular, G. biloba has a large seed size, and its survival rate of 85% could indicate that the raccoon dog latrines are suitable for its dispersal. These results demonstrate the effectiveness of seed dispersal raccoon dogs in the seed germination and growth phase. United Graduate School of Agricultural Science Tokyo University of Agriculture and Technology, 3-5-8 Saiwai Department of Ecology UNESP São Paulo State University Institute of Agriculture Tokyo University of Agriculture and Technology, 3-5-8 Saiwai Institute of Global Innovation Research Tokyo University of Agriculture and Technology, 3-5-8 Saiwai Department of Ecology UNESP São Paulo State University Japan Society for the Promotion of Science: JP17H00797 Japan Society for the Promotion of Science: JP25241026

Long-distance seed dispersal is an important ecosystem service provided by migratory animals. Plants inhabiting discrete habitats, like lakes and wetlands, experience dispersal limitation, and rely heavily on zoochory for their spatial population dynamics. Granivorous waterbirds may disperse viable seeds of wetland plants over long distances during migration. The limited knowledge of waterbird migration has long hampered the evaluation of the importance of waterbirds in seed dispersal, requiring key metrics such as realistic dispersal distances. Using recent GPS tracking of mallards during spring migration, we built a mechanistic seed dispersal model to estimate realistic dispersal distances. Mallards are abundant, partially migratory ducks known to consume seeds of >300 European plant species. Based on the tracking data, we informed a mallard migration simulator to obtain a probabilistic spring migration model for the mallard population wintering at Lake Constance in Southern Germany. We combined the spring migration model with seed retention curves to develop seed dispersal kernels. We also assessed the effects of pre-migratory fasting and the availability of suitable deposition habitats for aquatic and wetland plants. Our results show that mallards at Lake Constance can disperse seeds in the northeastern direction over median distances of 293 and 413 km for seeds with short and long retention times, respectively, assuming a departure immediately after foraging. Pre-migratory fasting strongly affected the dispersal potential, with only 1-7% of ingested seeds left for dispersal after fasting for 12 h. Availability of a suitable deposition habitat was generally

Whether fish mediate plant dispersal in temperate freshwaters is largely unknown. A prerequisite for successful dispersal is ingestion and surviving the journey in the intestinal tract. This study asks whether plant propagules are being ingested under field conditions and what factors shape dispersal potential, focusing on differences across plant species and propagule form (seed or fragment), seasonal differences and plant and fish traits that facilitate dispersal. We focused on three common fish species reported to differ in foraging strategy. Fish were caught monthly over a 1-year period in a Dutch lowland stream. Before they were returned to the stream, fish were kept in water for 26 h and their feces were collected, resulting in 150 fecal samples. Excreted animal remains and plant propagules were identified and enumerated. Plant propagules were tested for viability. In total, 88,579 vegetative fragments of vascular plants, 316 of mosses and 14 of charophytes were identified. Viability was low (<

Birds commonly regurgitate indigested particles after foraging. Many frugivorous birds regurgitate seeds ingested with the flesh of fruits, which importantly contributes to plant dispersal in terrestrial ecosystems. In freshwater ecosystems, waterbirds are known to defecate viable seeds in their faeces, but little is known about regurgitation as a potential additional seed dispersal mechanism. We experimentally fed eight mallards (Anas platyrhynchos) with a high and low volume of seeds of ten wetland plant species, and monitored regurgitation and defecation of intact seeds over 24 h. Regurgitation occurred at least once in all individual mallards and was induced by two different mechanisms: (i) feeding of high food volumes was significantly associated with regurgitation of all seed species from the crop after retention times of 1–3 h, and (ii) large indigestible seeds were expelled from the gizzard 11 or more hours after feeding. Seed regurgitation was much less plant species-specific than survival of seeds passing digestion, which suggests it is a particularly suitable dispersal mechanism for plant species unable to disperse by endozoochory (such as plant species with large, soft-bodied seeds). Observations of regurgitation by wild waterbirds are needed to improve our knowledge on this additional role of waterbirds in ecosystem functioning.

Fish can have an important role in the passive dispersal of freshwater zooplankton. In the Paraná River system the migratory fish Prochilodus lineatus (Valenciennes) constitutes about 60% of the fish biomass, the adult individuals have an ilyophagous feeding mode adapted to feed on soft bottom sediments. We hypothesize that P. lineatus ingests resting stages of zooplankton along with bottom sediments and that these resting stages are able to hatch after passing through the digestive tract. Forty adult P. lineatus individuals were caught in a lake located in the floodplain of Middle Parana River. Content of the last part of intestine was removed and divided into two equal portions and stored (3 months) at 4°C and at room temperature. Later, both portions were incubated for 27 weeks at 25°C. Hatching was controlled at 4-day intervals during the first 9 weeks of the experiment and later less frequently. At the end of the experiment, 8016 individuals were recorded, belonging to 18, mostly littoral species (15 rotifers, 2 cladocerans, and one copepod). Incubation preceded by a cooling period resulted in hatching of more species and individuals. Our results show that migratory fish may be an important vector for zooplankton dispersal. Fil: Battauz, Yamila Soledad. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Santa Fe. Instituto Nacional de Limnología. Universidad Nacional del Litoral. Instituto Nacional de Limnología; Argentina Fil: Jose, Susana Beatriz. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Santa Fe. Instituto Nacional de Limnología. Universidad Nacional del Litoral. Instituto Nacional de Limnología; Argentina. Universidad Nacional del Litoral. Facultad de Bioquímica y Ciencias Biológicas; Argentina Fil: Paggi, Juan Cesar. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Santa Fe. Instituto Nacional de Limnología. Universidad Nacional del Litoral. Instituto Nacional de Limnología; Argentina

The name Elatine campylosperma Seub. is generally treated as one of the synonyms of E. macropoda Guss. However, recent morphological, phylogenetic and karyological studies indicate that this judgement should be revised. In the present paper we typify the name E. campylosperma, review its taxonomic history and provide a thorough description, with compilation of previously published data and our new measurements from in vitro cultures. Based on our herbarium survey, we outline its Atlantic-Mediterranean distribution area (Spain, France, Italy, Greece, Turkey and Algeria). Habitat preferences are summarized from our field observations, water quality measurements and the label information of the herbarium specimens examined. Intact E. campylosperma seeds were found in faecal samples of the Eurasian Coot (Fulica atra L.) in southern Spain and two of them were germinated, suggesting that E. campylosperma has a capacity for long distance dispersal via endozoochory.

Fruit–frugivore interactions are critical for the dynamics and evolution of many plant communities. The strength of the interactions between a given plant species and different frugivore guilds (e.g., seed dispersers, seed predators) often vary in space due to changes in plant extrinsic factors (e.g., frugivore abundances) and plant traits (e.g., fruit size and reward). By reciprocally translocating Pyrus bourgaeana ripe fruits representative of five Mediterranean localities during 2 consecutive years, we experimentally quantified guild-specific variations among populations in frugivory strength, while accounted for plant-intrinsic and- extrinsic factors. Though overall fruit removal did not differ among localities, there were strong guild-specific differences in fruit removal strength. Fruit removal by pulp feeders, seed dispersers, and fruit predators varied among populations up to 8.5-, 5.6-, and 4.0-folds, respectively. These strong variations seemed mediated by changes in frugivore relative abundances rather than on availability of alternative fruits. As expected, all fruit traits considered (e.g., fruit size, pulp amount) markedly varied among tree populations. However, no frugivore guild showed preference for fruits from any locality, suggesting that fruit traits did not contribute much to differences in frugivory strength among populations. Since the functional diverse frugivore guilds played contrasting roles in P. bourgaeana dynamics (e.g., seed dispersal vs. seed predation), our study highlights the importance of accounting for functional diversity in frugivore guilds when estimating spatial variations in the strenght of seed dispersal. This investigation also illustrates a neglected but widely applicable experimental approach to identify the relative importance of extrinsic factors and fruit traits in mediating fruit–frugivore interactions.

Increasing land abandonment in many areas of the world presents an opportunity for ecosystem recovery, which is often driven by seed dispersal by vertebrate frugivores. However, we are far from understanding the most effective way of using common management actions (i.e. planting fruiting trees) to stimulate animal seed dispersal and thus the restoration of human-altered abandoned habitats. To investigate how to stimulate animal seed dispersal, we combined long-term field data with individual-based, spatially explicit simulation models. We used our approach to assess the effectiveness of contrasting Iberian pear Pyrus bourgaeana planting strategies in enhancing restoration of abandoned lands through seed dispersal by red foxes Vulpes vulpes and Eurasian badgers Meles meles in the Doñana World Biosphere Reserve (South West Spain). Our simulation results indicate that planting trees in an aggregated fashion is less efficient in terms of seed arrival than planting them regularly or randomly. For aggregated planted trees, the increase in the area of the oldfield that received seeds was only 7%–9% compared to the baseline scenario of no intervention, whereas for regularly distributed planted trees the increment was up to 40%. Doubling the number of planted P. bourgaeana trees appeared cost-effective for regular and random tree distributions, but not for the aggregated one. For example, while doubling the number of trees planted regularly leads up to 12% increase in the number of seeds arriving into the oldfield, no increment on the number of arrived seeds was detected when trees were planted aggregately. Synthesis and applications. Choosing the spatial distribution and density of planted trees in abandoned lands depends on a number of ecological and socio-economical factors. Given our results, the strong seed dispersal limitation of the target tree population and that our study site was fully protected for conservation, planting Pyrus bourgaeana trees regularly appeared to be the most efficient strategy to enhance seed arrival into the target oldfield. Combining long-term field data with individual-based, spatially explicit simulation models have the potential to guide local restoration efforts in diverse human-altered habitats and thus bridge the existing gap between basic and applied research on animal seed dispersal.

Most studies and reviews on endozoochorous seed dispersal by ungulates overlook that ruminants spit viable seeds from the cud, i.e. after they were ingested and carried inside the rumen. To assess the importance of this modality of endozoochory, we performed feeding trials with three captive red deer (Cervus elaphus) and four common plant species (Chamaerops humilis, Crataegus monogyna, Celtis australis and Ceratonia siliqua) differing in fruit and seed traits. Experimental deer regurgitated seeds from the rumen and ejected them, and also defecated some seeds of the four species, at a ratio of about 6:10. This high ratio (about 37%) of spitted seeds would be overlooked by studies considering only dispersal of seeds contained in faeces.

Assessing dispersal events in plants faces important challenges and limitations. A methodological issue that limits advances in our understanding of seed dissemination by frugivorous animals is identifying 'which species dispersed the seeds'. This is essential for assessing how multiple frugivore species contribute distinctly to critical dispersal events such as seed delivery to safe sites, long-distance dispersal and the colonization of non-occupied habitats. Here, we describe DNA-barcoding protocols successfully applied to bird-dispersed seeds sampled in the field. Avian DNA was extracted from the surface of defecated or regurgitated seeds, allowing the identification of the frugivore species that contributed each dispersal event. Disperser species identification was based on a 464-bp mitochondrial DNA region (COI: cytochrome c oxidase subunit I). We illustrate the possible applications of this method with bird-dispersed seeds sampled in the field. DNA-barcoding provides a non-invasive technique that allows quantifying frugivory and seed dispersal interaction networks, assessing the contribution of each frugivore species to seed rain in different microhabitats, and testing whether different frugivore species select different fruit/seed sizes. DNA barcoding of animal-dispersed seeds can resolve the distribution of dispersal services provided by diverse frugivore assemblages, allowing a robust and precise estimation of the different components of seed dispersal effectiveness, previously unattainable to traditional field studies at individual seed level. Given that seeds are sampled at the end of the dispersal process, this technique enables us to link the identity of the disperser species responsible for each dispersal event to plant traits and environmental features, thereby building a bridge between frugivory and seed deposition patterns. © 2014 British Ecological Society.

Intra- and interspecific spatially contagious seed dispersal has far-reaching implications for plant recruitment, distribution, and community assemblage. However, logistical and analytical limitations have curtailed our understanding concerning the mechanisms and resulting spatial patterns of contagious seed dispersal in most systems and, especially, in complex seed-disperser networks. We investigated mechanisms of seed aggregation using techniques of spatial point pattern analysis and extensive data sets on mutispecific endozoochorous seed rain generated by five frugivorous mammals in three Mediterranean shrublands over two seasons. Our novel analytical approach revealed three hierarchical and complementary mechanisms of seed aggregation acting at different levels (fecal samples, seeds, pairs of seed species) and spatial scales. First, the three local guilds of frugivores tended to deliver their feces highly aggregated at small and intermediate spatial scales, and the overall pattern of fecal delivery could be described well by a nested doublecluster Thomas process. Second, once the strong observed fecal aggregation was accounted for, the distribution of mammal feces containing seeds was clustered within the pattern of all feces (i.e., with and without seeds), and the density of fecal samples containing seeds was higher than expected around other feces containing seeds in two out of the three studied seeddisperser networks. Finally, at a finer level, mark correlation analyses revealed that for some plant species pairs, the number of dispersed seeds was positively associated either at small or large spatial scales. Despite the relatively invariant patterning of nested double-clustering, some attributes of endozoochorous seed rain (e.g., intensity, scales of aggregation) were variable among study sites due to changes in the ecological context in which seeds and their dispersers interact. Our investigation disentangles for the first time the hierarchy of synergic mechanisms of spatially contagious seed dispersal at a range of spatial scales in complex seeddisperser networks, thus providing a robust and widely applicable framework for future studies. © 2014 by the Ecological Society of America.

In order to clarify the interactive mechanism between grazing yak and alpine meadow on the Qinghai-Tibetan Plateau, our study assessed seed density (by species) in the topsoil of alpine meadow with different grazing intensities in the Tianzhu area, north-eastern margins of the Qinghai-Tibetan Plateau and their rates of occurrence in yak dung. Seed density in the topsoil of the lightly grazed, moderately grazed, heavily grazed and extremely grazed alpine meadows in November, 2010 were 1 551, 1 692, 2 660 and 1 830 grains m^(-2), while in the same meadows in April, 2011 densities were 1 530, 2 404, 2 530 and 2 692 grains m^(-2), respectively. In the cold season pasture, mean seed density in yak dung from November to April in the lightly grazed, moderately grazed, heavily grazed and extremely grazed sites were 121, 127, 187, and 120 grains kg^(-1) of dry yak dung. The proportion of total seed numbers in yak dung to soil seed bank in lightly grazed, moderately grazed, heavily grazed and extremely grazed alpine meadow was 1.40, 2.62, 0.69, and 0.90%. 12 species out of the 47 were not found in topsoil but were found in yak dung, 10 species out of 45 were not found in yak dung but were found in the topsoil. Endozoochorous dispersal by yaks is therefore very important for soil seed bank and plant biodiversity and population dynamics in alpine meadows.

Seed ingestion by frugivorous vertebrates commonly benefits plants by moving seeds to locations with fewer predators and pathogens than under the parent. For plants with high local population densities, however, movement from the parent plant is unlikely to result in ‘escape’ from predators and pathogens. Changes to seed condition caused by gut passage may also provide benefits, yet are rarely evaluated as an alternative. Here, we use a common bird-dispersed chilli pepper (Capsicum chacoense) to conduct the first experimental comparison of escape-related benefits to condition-related benefits of animal-mediated seed dispersal. Within chilli populations, seeds dispersed far from parent plants gained no advantage from escape alone, but seed consumption by birds increased seed survival by 370% – regardless of dispersal distance – due to removal during gut passage of fungal pathogens and chemical attractants to granivores. These results call into question the pre-eminence of escape as the primary advantage of dispersal within populations and document two overlooked mechanisms by which frugivores can benefit fruiting plants.

Endozoochory has been suggested as an important mechanism for long-distance plant dispersal that facilitates seed arrival to new and more favorable habitats. The hardness (mechanical strength) and thickness of the seed coat are important characteristics for seed survival and germination after passage through an herbivore gut. We studied the effects of seed passage through cattle gut on seed coat structure in three hard-seeded Acacia species that have physical dormancy, A. aroma, A. atramentaria and A. caven, occurring in the semiarid woodlands and shrublands of central Argentina. Histology of the seed coat was examined and a simulated cattle consumption experiment was conducted. Only A. aroma, which has the thinnest seed coat (in terms of epidermis and sclerified parenchyma) of the three species, showed a high germination percentage after seed passage through the digestive tract of cattle, whereas seeds of A. atramentaria and A. caven remained hard and viable. We conclude that the structure of the seed coat in hard-seeded species is crucial in determining the success of endozoochory.

In Europe and North America waterfowl are major dispersers of aquatic and terrestrial plants, but in New Zealand their role has yet to be investigated. Mallards were introduced to New Zealand in the late 1800s, and today they are the most abundant and widespread waterfowl in the country. To assess seed dispersal, we radiomarked 284 female mallards from two study sites during the pre-breeding (June August) and breeding (August December) periods in 2014 2015, and examined movements that occurred within 24, 48 or 72 h when seed dispersal by endozoochory is considered likely. During June and July 2015, we collected 29 faecal samples from individual female mallards during radiomarking and 24 samples from mallard flocks. We recovered 69 intact seeds from the faecal samples and identified 12 plant taxa. Of the plant seeds identified and dispersed by mallards in this study, 40% were members of the Asteraceae family, nine plant species were alien to New Zealand, and the indigenous-status of three unidentified taxa could not be determined. Two taxa (and9%of seeds) were germinated following gut passage: an unidentified Asteraceae and Solanum nigrum. During the prebreeding and breeding periods, movement of females within 24 h averaged 394m(SDD 706 m) and 222m(SD D 605 m) respectively, with maximum distances of 3,970mand 8,028 m. Maxima extended to 19,230 m within 48 h. Most plant species recorded are generally assumed to be self-dispersed or dispersed by water; mechanisms that provide a much lower maximum dispersal distance than mallards. The ability of mallards to disperse viable seeds up to 19 km within 48 h suggests they have an important and previously overlooked role as vectors for a variety of wetland or grassland plant species in New Zealand.

Long distance dispersal may have important consequences for gene flow and community structure. The dispersal of many plants depends on transport by vertebrate seed dispersers. The shapes of seed shadows produced by vertebrates depend both on movement patterns of the dispersers and on the dynamics and effects of passage through the disperser’s gut (i.e. the retention time, survival and germination of ingested seeds). A combination of experiments with captive waterbirds and aquatic plant seeds was used to analyse the following: (a) the effects of inter- and intra-specific variation in seed size and duck species on seed retention time in the gut and (b) the relationship between retention time and the percent germination and germination rates of seeds. Among the three Scirpus species used, those with smaller seeds showed higher survival after ingestion by birds and longer retention times inside their guts than those with larger seeds. For Potamogeton pectinatus, only seeds from the smaller size class (

Background: Propagule retention time is a key factor in determining propagule dispersal distance and the shape of “seed shadows”. Propagules dispersed by animal vectors are either ingested and retained in the gut until defecation or attached externally to the body until detachment. Retention time is a continuous variable, but it is commonly measured at discrete time points, according to pre-established sampling time-intervals. Although parametric continuous distributions have been widely fitted to these interval-censored data, the performance of different fitting methods has not been evaluated. To investigate the performance of five different fitting methods, we fitted parametric probability distributions to typical discretized retention-time data with known distribution using as data-points either the lower, mid or upper bounds of sampling intervals, as well as the cumulative distribution of observed values (using either maximum likelihood or non-linear least squares for parameter estimation); then compared the estimated and original distributions to assess the accuracy of each method. We also assessed the robustness of these methods to variations in the sampling procedure (sample size and length of sampling time-intervals). Results: Fittings to the cumulative distribution performed better for all types of parametric distributions (lognormal, gamma and Weibull distributions) and were more robust to variations in sample size and sampling time-intervals. These estimated distributions had negligible deviations of up to 0.045 in cumulative probability of retention times (according to the Kolmogorov–Smirnov statistic) in relation to original distributions from which propagule retention time was simulated, supporting the overall accuracy of this fitting method. In contrast, fitting the sampling-interval bounds resulted in greater deviations that ranged from 0.058 to 0.273 in cumulative probability of retention times, which may introduce considerable biases in parameter estimates. Conclusions: We recommend the use of cumulative probability to fit parametric probability distributions to propagule retention time, specifically using maximum likelihood for parameter estimation. Furthermore, the experimental design for an optimal characterization of unimodal propagule retention time should contemplate at least 500 recovered propagules and sampling time-intervals not larger than the time peak of propagule retrieval, except in the tail of the distribution where broader sampling time-intervals may also produce accurate fits

1. Dabbling ducks (Anatinae) are omnivorous birds that are widespread, numerous, highly mobile and often migratory, and therefore have great potential for (long distance) dispersal of other organisms, including plants. However, their ability to act as plant dispersal vectors has received little attention compared to frugivores and is often assumed to be relevant only for wetland species. 2. To evaluate the potential for plant dispersal by dabbling ducks, we collated and analysed existing data. We identified all plant species whose seeds have been recorded in the diets of the seven dabbling duck (Anas) species in the Western Palaearctic, as reported from gut content analyses. We then analysed the habitats and traits of these plant species to identify general patterns, and related these to data on gut passage survival and duck movements. 3. A large number of plant species (> 445 species of 189 genera and 57 families) have been recorded in the diet of dabbling ducks. These plant species represent a very wide range of habitats, including almost the full range of site fertility, moisture and light conditions, excluding only very dry and deeply shaded habitats. The ducks prefer seeds of intermediate sizes (1–10 mm3), which have good chances to survive gut passage, but also ingest smaller and larger seeds. Ingested seeds represent a wide range of dispersal syndromes, including fleshy fruits. Many species (62%) were not previously considered animal- dispersed in plant data bases, and 66% were not identified as bird-dispersed. Rarefaction analyses suggest that our analysis still greatly underestimates the total number of plant species ingested. 4. Synthesis. Dabbling ducks do not exclusively ingest seeds of wetland plants, which make up only 40% of the ingested species. Rather, they feed opportunistically on a wide cross-section of plant species available across the landscapes they inhabit. Given the millions of ducks, the hundreds to thousands of seeds ingested per individual on a daily basis, and known gut passage survival rates, this results in vast numbers of seeds dispersed by ducks per day. Internal seed dispersal by dabbling ducks appears to be a major dispersal pathway for a far broader spectrum of plant species than previously considered

1. The potential for seed dispersal by fish (ichthyochory) is likely to vary within aquatic plant species, depending on intraspecific variation in phenotypic seed traits. 2. We studied the effect of seed size variation within the unbranched burreed (Sparganium emersum) on the potential for internal dispersal by the common carp (Cyprinus carpio), by feeding them light (< 10 mg), medium (10-20 mg) and heavy (> 20 mg) seeds, seed mass being positively related to seed size. 3. We hypothesized: (i) that ingestion, retention time, survival during gut passage and viability after gut passage of S. emersum seeds would be affected by seed size; and (ii) that this would translate into intraspecific variation in dispersal probability and dispersal distance among seed size categories. 4. Ingestion was negatively related to seed size, while survival during gut passage was positively related to seed size. Seed viability after gut passage was not affected by seed size. Since the negative effect of ingestion was counterbalanced by an equally strong but positive effect on seed survival, the probability of dispersal did not differ between the tested seed-size categories. 5. The time that seeds remained in the digestive tract of carp did not differ between seed sizes, suggesting equal potential dispersal distances for all seeds. Based on optimum swimming speeds of carp, ranging from 0.9 to 1.8 km h-1, maximum dispersal distances will most likely range from 13.5 to 27 km. 6. This study highlights the importance of studying all stages of the endozoochorous dispersal process in order to estimate the effect of a phenotypic seed trait on plant dispersal.

Seed dispersal via ingestion and defecation by large herbivores (endozoochory) plays a potentially important role in structuring plant communities. In the present study we tested whether cattle disperse seeds between different plant communities in a heterogeneous coastal habitat. We surveyed the seed contents of cattle dung collected from two habitat types within a grazing system, one in dunes and the other in salt-marsh. The dunes are characterized by sandy soil, and infrequent inundation by the sea, whereas the salt-marsh features clay deposited during inundations. Seeds of both salt-marsh and dune species were dispersed into salt-marsh and dune habitats. The seed content of cattle dung collected in salt-marsh and dunes was similar with respect to species composition and seed density. However, dispersal of dune species into salt-marsh and vice-versa did not result in establishment in those communities. Seed traits per se did not differ between dune species and salt-marsh species. Species abundance in the established vegetation and seed density in dung were positively correlated. Seed abundance for most species found in dung was consistent with the availability of viable seeds during the growing season. Seed mass, seed 'roundness' and seed release height were all negatively correlated with seed numbers in dung. An indirect factor, namely, forage quality of the plant, and hence attractiveness for herbivores, could explain the higher likelihood of seed dispersal of salt-marsh species than of dune species. The salt-marsh harbours species with higher forage quality than the dune species, as derived from Ellenberg nitrogen indicator values.

1. The survival and retention of seeds was studied by feeding known quantities of seeds of 25 species to four captive Fallow Deer (Dama dama L.). To test for ecological correlates, plant species were selected to represent large variation in seed size, seed shape, seed longevity and habitat fertility.2. Seeds of 24 out of 25 fed plant species survived ingestion and defecation by Fallow Deer. Seed survival ranged between 0.5 and 42% of germinable seeds fed. Time to recover 50% of all seeds defecated by Fallow Deer in faeces averaged 25 h, and ranged from 13 to 38 h.3. Seed survival was negatively related to seed mass (R = 0.65) and variance of unit seed dimensions (R = -0.56), and positively related to seed longevity (R = 0.40), but not related to habitat fertility. Log(10) of (seed mass x variance of seed dimensions) was the best predictor of seed survival (R = -0.68).4. The ecological correlates of seed survival presented here can help us to estimate the ability of plant species to disperse seeds over long distances.

© 2015 John Wiley & Sons Ltd. Summary: Alien plant species are rapidly spreading in aquatic ecosystems around the world, causing major ecological effects. They are typically introduced by humans, after which natural vectors facilitate their further spread. Migratory waterbirds have long been recognised as important dispersal vectors for native and aquatic plants, yet little is known about their role in the spread of alien species. We determined experimentally the potential for long-distance dispersal of native and alien wetland plants in Europe by two abundant waterfowl: mallards Anas platyrhynchos and greylag geese Anser anser. We fed seeds from two plants alien to Europe and two native plants to 10 individuals of each bird species, testing for the effects of bird and plant species on the potential for dispersal. Intact seeds were retrieved from faeces for up to 4 days after ingestion. The proportion of seeds retrieved intact varied significantly between plant, but not bird, species. Retrieval was highest for the invasive water primrose Ludwigia grandiflora (>35% of ingested seeds), lowest for the invasive cordgrass Spartina densiflora (