1. The interplay between dispersal and adaptation to local environments ultimately determines the distribution of plant species, but their relative contribution remains little understood. Tropical ...mistletoes provide the opportunity to dissect these contributions of dispersal and adaptation, because as hemiparasitic plants, they are typically adapted to grow on a handful of species within diverse tropical communities and are non-randomly dispersed by mutualistic frugivorous birds. 2. Here we hypothesized that the primary determinant of the abundance of a tropical mistletoe (Dendropemon caribaeus, Loranthaceae) in Puerto Rico will be the compatibility between the mistletoe and plant species in a community. Alternatively, the mistletoe's abundance could be primarily shaped by other factors such as the availability of potential host plants, or factors that determine how mistletoe seeds are dispersed by avian frugivores. We conducted surveys and experiments to assess the capacity of this mistletoe to grow on trees available in the local community, and measured the monthly phenology and seed dispersal patterns of the mistletoe and other bird-dispersed plants in the community over a period of 4 years. A path model was used to evaluate how the abundance of the mistletoe was shaped by host abundance, fruiting phenology, bird dispersal and compatibilities with host plants. 3. Our analyses show that the compatibility between mistletoe and host tree species, measured by mistletoe survival and growth rate, was the most important factor for mistletoe abundance. The next most important factor was the phenological characteristics of the hosts; this outcome likely arose because frugivory and seed dispersal services for mistletoes and hosts are performed by the same birds. 4. Synthesis. Mistletoes often parasitize only a subset of the suitable plant species that are available in a given community. Our results indicate that such patterns are not only determined by host quality and abundance but also by the phenological patterns of trees that influence the probabilities of mistletoe seeds being deposited on them by shared avian seed dispersers.
Network models of frugivory and seed dispersal are usually static. To date, most studies on mutualistic networks assert that interaction properties such as species' degree (k) and strength (s) are ...strongly influenced by species abundances. We evaluated how species' degree and strength change as a function of temporal variation not only in species abundance, but also in species persistence (i.e., phenology length). In a two-year study, we collected community-wide data on seed dispersal by birds and examined the seasonal dynamics of the above-mentioned interaction properties. Our analyses revealed that species abundance is an important predictor for plant strength within a given sub-network. However, our analyses also reveal that species' degree can often be best explained by the length of fruiting phenology (for plants degree) or by the number of fruiting species (for dispersers degree), which are factors that can be decoupled from the relative abundance of the species participating in the network. Moreover, our results suggest that generalist dispersers (when total study period is considered) act as temporal generalists, with degree constrained by the number of plant species displaying fruits in each span. Along with species identity, our findings underscore the need for a temporal perspective, given that seasonality is an inherent property of many mutualistic networks.
Oceanic islands are places where biological assemblages are relatively simple, as compared to the mainland. On islands, however, pollinator assemblages may to be composed of a taxonomically disparate ...group of organisms (e.g. insects, lizards, and birds), some of them with opportunistic nectar-feeding behaviour. Here we investigated some components of pollination effectiveness of Lotus maculatus (Fabaceae), an endangered Canary Islands endemic. In a flower exclusion experiment, we bagged flowers and compared their subsequent fruit and seed set to that of control flowers. Number of interactions with vertebrate and invertebrate flower visitors was counted and it was recorded whether interactions were legitimate (potentially pollinating) or non-legitimate (nectar robbing). Additionally, we estimated pollen loads on lizards and looked for any relationship between reproductive success of individual plants and number of visits made by the top three flower-visiting species (in terms of both frequency of occurrence at censuses and number of floral visits). Bagged flowers fruited less and with fewer seeds than control flowers. The only observed flower-visiting vertebrate was the Tenerife lizard Gallotia galloti, whose interactions were always legitimate and with around a half of captured individuals carrying pollen grains. The most frequent flower-visiting insect was the honeybee Apis mellifera followed by the solitary bee Lasioglossum arctifrons. The honeybee, however, was only a nectar robber, and the solitary bee was not an effective pollinator, but rather a pollen gatherer. Fruit set by individual plants was positively related only to frequency of visits by the lizard. Thus, the lizard seems to play a key role in the conservation management of L. maculatus.
Abstract
We provide an overview of the current state of knowledge of island frugivory and seed dispersal and identify knowledge gaps that are important for fundamental research on—and applied ...conservation of—island ecosystems. We conducted a systematic literature search of frugivory and seed dispersal on islands, omitting large, continental islands. This revealed a total of 448 studies, most (75%) published during the last two decades, especially after 2010. Nearly 65% of them were focused on eight archipelagos. There is a paucity of studies in Pacific archipelagos near Asia and Australia, and in the Indian Ocean. Data on island frugivory and seed dispersal are diverse but highly uneven in geographic and conceptual coverage. Despite their limited biodiversity, islands are essential reservoirs of endemic plants and animals and their interactions. Due to the simplicity of insular ecosystems, we can assess the importance of seed dispersal theory and mechanisms at species and community levels. These include the ecological and biogeographical meaning and prevalence of non-standard mechanisms of seed dispersal on islands; the seed dispersal effectiveness and the relative roles of different frugivore guilds (birds and reptiles being the most important); and patterns of community organization and their drivers as revealed by interaction networks. Island systems are characterized by the extinction of many natives and endemics, and high rates of species introductions. Therefore, understanding how these losses and additions alter seed dispersal processes has been a prevailing goal of island studies and an essential foundation for the effective restoration and conservation of islands.
Frugivores shape plant communities via seed dispersal of fleshy‐fruited plant species. However, the structural characteristics that frugivores impart to plant communities are little understood. ...Evaluating how frugivores structure plant communities via the nonproportional use of available fruit resources is critical to understand the functioning of ecosystems where fleshy‐fruited plant species are dominant, such as tropical forests.
We performed a seed‐addition field experiment to investigate how frugivorous birds shape the composition and richness of forests during early stages of secondary succession in cleared areas in Puerto Rico. The experiment tested whether the bird‐generated seed rain and the subsequent early successional plant communities were proportional representations of the fleshy‐fruited species that dominated the surrounding community. Experimental treatments consisted of patches with (a) seed‐additions by wild birds attracted to experimental patches with pole perches, (b) manual seed‐additions proportional to fruit abundance at the local scale (≤50 m from experimental plots) and (c) manual seed‐additions proportional to fruit abundance at the landscape scale (entire study site).
Birds' seed‐additions differed in composition and abundance to expectations based on fruit availability at local and landscape scales. Treatments with seeds added by birds had the highest species richness in both the seed rain and the emergence stages despite how, on average, the monthly richness in the landscape‐scale treatment was double that of birds and the local‐scale treatment. This phenomenon was explained by the highest heterogeneity from the bird seed‐addition treatment across months, and the lowest seed per capita emergence rates in landscape treatments. Rather than reflecting relative fruit abundance, birds biased seed rain and per capita emergence towards a non‐random mixture of both small‐ and large‐seeded species, resulting in richer and distinct plant communities.
Because frugivory and seed dispersal patterns depart from random encounters between frugivores and plants in communities, successional forests are characterized by an overrepresentation of proportionally rare plant species, and decreases in the dominance of many common species. Thus, for regenerating tropical forests, frugivory can function as mechanism that promotes persistence of rare plant species and their coexistence with more abundant plants.
A plain language summary is available for this article.
Plain Language Summary
The behavioural complementarity of fruit‐eating animals is thought to exert a key role in plant community assembly. However, a mechanistic understanding of the causal links between the two processes ...is still lacking.
This study assesses whether complementarity between dispersers in feeding and microhabitat‐use behaviour enhances community‐scale dispersal services, resulting in a more diverse community of seedlings.
We used a Bayesian approach to connect a comprehensive database of seed dispersal effectiveness at a community scale with a transition probability model that accounts for behavioural complementarity. Our model system was the thermosclerophyllous shrubland of the Canary Islands. There, fleshy‐fruited plants rely on two types of frugivores: lizards and birds.
Lizards consumed all plant species and preferentially used open areas, whereas birds foraged for small single‐seeded fruits and dispersed their seeds beneath plants. Through feeding on different sets of plants, they generated a rich seed‐rain community. By diversifying the microhabitat of deposition, more species could find suitable recruitment sites.
Distinct foraging and microhabitat‐use choices led to complementary dispersal services. Lizards ensured that all plant species were present in the seedling community, while birds promoted a more even distribution of them. As a result, diversity in the community of seedlings was enhanced.
Overall, our work underscores that behavioural complementarity promotes diversity in the early‐regenerating plant communities. These enhanced dispersal services rely on the presence of all functional groups. Thus, in communities where frugivores display unique behaviours, preserving a diverse community of dispersers should be a conservation target.
A free Plain Language Summary can be found within the Supporting Information of this article.
A free Plain Language Summary can be found within the Supporting Information of this article.
Seed dispersal effectiveness (SDE) is the contribution of dispersers to plant recruitment and is estimated as the product of the number of seeds dispersed (quantity) and the probability of ...recruitment of each dispersed seed (quality). Although SDE is a key concept in seed dispersal ecology, few studies estimate SDE and none has a community approach. Oceanic islands, with simple communities, are ideal for this purpose. In this study, we compared the SDE of the main types of dispersers (lizards and passerine birds) at the community level in a given habitat. We estimated SDE using a stochastic simulation model parameterized with empirical data on quantity and quality components measured throughout the recruitment process. Although lizards are highly frugivorous and their density was ~20 times higher than that of birds, lizards and birds dispersed a similar quantity of seeds. This may be due to lower intake of seeds by lizards due to their slower metabolism (~20 times lower than birds). This low metabolic rate limits the importance of lizards as seed dispersers, but it is compensated by extraordinarily high lizard densities in the study area (~9600 individuals/km
2
). High densities of lizards are typical of islands, and this helps to explain why dispersal by lizards seems mainly an island phenomenon. Birds and lizards showed functional complementarity, especially regarding seed dispersal distribution patterns. In fact, lizards dispersed more seeds in shrublands and open sites, and birds in woodlands and beneath canopies, with their joint contribution helping to maximize recruitment. Lizards provided higher SDE than birds for 7 out of 11 plant species. The disperser with a higher quantity for a given plant generally had the higher quality, and plants could be classified as bird- or lizard-dependent for dispersal. This dependence increased when considering SDE instead of dispersal quantity only. Moreover, quality was a better predictor of SDE than quantity, which should be considered when parameterizing interaction networks, as this might affect inferences about their architecture.
Aim: We studied, for the first time, the effect of insularity on plant-seed disperser networks at both the community and species level. We focused on the Mediterranean shrubland, comparing different ...biogeographical scenarios (insular and continental) that share the same basic species composition. Location: The study was conducted within one mainland (southern Spain) and four island localities: two in the Canary Islands (oceanic origin) and two in the Balearic Islands (continental origin). Methods: We built qualitative (presence/absence of interaction) and quantitative seed dispersal networks (number of consumed fruits) and used different descriptor parameters, such as connectance, nestedness, interaction asymmetry and various interaction diversity indices that describe their topology. To assemble the interaction networks, we used data on the presence of different species of fruits in systematically collected droppings of the different seed dispersers; seeds in droppings were identified by means of a binocular lens whereas a microscope was used to identify pericarp tissue remains. Results: Island networks were smaller and less complex (a lower number of observed links than expected for their size) than the mainland network. As expected, connectance was higher within islands than on the mainland. By contrast, nestedness was consistently high at all sites, although relative nestedness (which accounts for network size) was lower within islands, whether continental or oceanic. At both community and species level (especially for animals) interactions tended to be more specialized and symmetric within the islands. Main conclusions: The lower species number and greater specialization in insular seed dispersal systems appears to lead to the prevalence of more symmetric interactions than those found on the mainland. This indicates that insular mutualistic interactions and interacting species are more vulnerable than mainland ones to disturbances, as previous work has suggested that asymmetrical interactions facilitate the maintenance of biodiversity and act as a resilience mechanism against species extinction.
1. Secondary seed dispersal is a multistep process with two or more phases, which involve different dispersers that usually extend the distance from the seed’s parent plant. This ecological process ...has been recorded in some subtropical oceanic islands, where predatory birds commonly consume frugivorous lizards and disperse seeds already consumed by the lizards. 2. We evaluated the overall importance of this type of secondary dispersal in the Canary Islands, the only place worldwide where it has been studied in depth. From an examination of all the islands and their suitable habitats, we found seeds from 78 plant species inside 2098 shrike pellets and 5304 kestrel pellets. A greater number of species were secondarily dispersed by kestrels (76; 97%) than by shrikes (26; 34%). 3. Forty‐four (56%) of the total species detected in pellets were identified at the species level, comprising 73% native and 27% introduced species. Seventy per cent of these identified species were fleshy fruit‐bearing plants and 84% of the interactions took place in open habitats, close to coastal areas. 4. Germination experiments showed that seeds of at least 32 plant species were viable after being removed from the bird pellets. A similar pattern of seed germination was detected for seeds from the droppings of lizards and pellets of shrikes, showing both to be effective dispersers. However, the seeds dispersed by kestrels had different levels of success depending on the number of gut passages experienced. Seeds that had undergone double gut treatment (lizard and secondary ingestion by kestrel) had reduced germination rates of many small‐ and medium‐sized seeds compared with seeds ingested by lizards and discarded inside the lizard guts by kestrels. 5. We also studied the relationship between body length and gape width of lizards in order to assess limitations on the sizes and quantities of seeds available for secondary dispersal. Kestrels can disperse a greater number and variety of seeds because they predate larger lizards that potentially carry greater seed loads. 6. Synthesis. The current results show how these non‐standard long‐distance dispersal events produced by predatory birds can be considered as a regular and generalized process on all islands of the Canary archipelago.
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