Climate warming increases insect-driven seed removal of two elaiosome-bearing invasive thistle species.

Ants and other insects are often a source of localized secondary dispersal for wind-dispersed plants and thus play an important ecological role in their spatial dynamics, but there is limited information on how climate change will affect such dispersal processes. Here, we use field experiments to investigate how climate warming affects seed removal, as this initiation of movement represents the first step in insect-driven secondary dispersal. Our results indicate that for the invasive thistles Carduus nutans and Carduus acanthoides, increased growing temperature influences seed attractiveness to insect dispersers, with seeds from maternal plants grown at temperatures 0.6°C above ambient removed by insect dispersers at higher rates than their unwarmed counterparts. We also observe that seed elaiosomes in these two species play an important role in dispersal, as seeds without elaiosomes were significantly less likely to be removed over the same period. Significant interactions between elaiosome presence/absence and warming treatment were also observed, though only for C. acanthoides, with the boost in seed removal from warming dampened when the elaiosome was present compared to when it was absent. These findings provide evidence that climate warming may alter aspects of dispersal such as seed removal by secondary dispersers, with potential ramifications for dispersal in future climates since seed-bearing plants around the world may be subject to increased growing temperatures, and many of these plant species bear elaiosomes and experience seed dispersal by insects.

Evaluation of seed-dispersal services by ants at a temperate pasture: Results of direct observations in an ant suppression experiment.

Ants disperse seeds of many plant species adapted to myrmecochory. While advantages of this ant-plant mutualism for myrmecochorous plants (myrmecochores) have been previously studied in temperate region mostly in forests, our study system was a pasture. Moreover, we used a unique combination of observing the effect of ant-activity suppression on ant dispersal and comparison of the contribution of ant and unassisted dispersal to the distance from mother plant. We established plots without and with ant-activity suppression (enclosures). We offered diaspores of a myrmecochorous (Knautia arvensis), and a non-myrmecochorous (Plantago lanceolata) species in a choice test and followed ants carrying diaspores during days and nights (focus of previous studies was on diurnal dispersal). We measured frequency and distances of ant dispersal and compared them with unassisted dispersal recorded using sticky trap method. The dispersal frequency was lower in enclosures (3.16 times). Ants strongly preferred diaspores of the myrmecochore to non-myrmecochore with 586 and 42 dispersal events, respectively (out of 6400 diaspores of each species offered). Ant dispersal resulted in more even and on average longer distances (maximum almost tenfold longer, 994 cm) in comparison to unassisted dispersal. Ant dispersal altered the distribution of distances of the myrmecochore from roughly symmetric for unassisted dispersal to positively skewed. Ants dispersed heavier diaspores farther. Ants dropped the majority of diaspores during the dispersal (which reduces clustering of seeds), while several (11%) were carried into anthills. Anthills are disturbed microsites presumably favorable for germination in competitive habitats. Ants provided non-negligible dispersal services to myrmecochorous K. arvensis but also, to a lesser extent, of non-myrmecochorous P. lanceolata.

EuDiS - A comprehensive database of the seed dispersal syndromes of the European flora.

Background: 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. New information: 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.

Ecology of fear: predator avoidance reduces seed dispersal in an ant.

The ecology of fear refers to the non-fatal cost that predators and parasites impose on prey populations. These non-consumptive effects (NCEs) can influence animal-plant interactions, but evidence thereof comes mainly from vertebrate systems with less focus on invertebrates. Here, I investigated whether the foraging behaviour of the ant Ectatomma ruidum was influenced by its primary predator, the forest toad Rhinella alata. In field tests, the probability of seed removal by the ants was 25% for seeds placed with the forest toad compared to 32% for control seeds, suggesting that toads reduce ant foraging rates. A further experiment revealed that ants which had previously encountered the predator and its faeces were more likely (59%) than inexperienced ants (50%) to avoid the exit with the predator faeces. This outcome suggests that ants are capable of learning cues associated with predation risk, possibly leading to NCEs. This indicates that predators can exert NCEs on invertebrate prey with potential cascading effects on seed dispersal, extending results previously seen only in vertebrate seed dispersal systems.

Take me for a ride: Herbivores can facilitate plant reinvasions.

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

Historical forest disturbance results in variation in functional resilience of seed dispersal mutualisms.

Mutualistic interactions provide essential ecosystem functions that contribute to promoting and maintaining diversity in ecosystems. Understanding if functionally important mutualisms are "resilient" (i.e., able to resist or recover) to anthropogenic disturbance is essential for revealing the capacity for diversity to recover. Animal-mediated seed dispersal supports plant population growth and influences community structure, and disturbance affecting seed dispersal can contribute to low resiliency of plant diversity. Ant-seed dispersal mutualisms are sensitive to anthropogenic disturbance, as they rely on one to a few high-quality dispersal partners. In North American eastern deciduous forests, ants in the genus Aphaenogaster are "keystone dispersers" of understory forbs adapted to dispersal by ants (myrmecochores), which make up more than one-third of the understory herbaceous community. The majority of forests within this region have regenerated from previous disturbance in the form of clearing for agriculture. Previous studies have revealed that myrmecochore diversity is not resilient to previous clearing. Here, we ask if seed dispersal mutualisms are resilient to historical forest disturbance and if decreases in mutualistic interactions with partners, Aphaenogaster sp., or increases in antagonistic interactions cause degradation of function. In a large-scale natural experiment (20 sites), we measured seed removal, the abundance of mutualistic partners and other invertebrates interacting with seeds, myrmecochore cover, and diversity, along with ant habitat and forest structure. We found lower and more variable seed removal in secondary forests compared with remnant forests. A path analysis of all forests revealed that the abundance of mutualists was the primary determinant of the variation in seed removal, and that seed damage by antagonists (invasive slugs) negatively affected dispersal and was higher in secondary forests. In a path analysis of remnant forests, the link between mutualist abundance and seed removal was absent, but present in the secondary forest path, suggesting that seed dispersal is more variable and dependent on the mutualist abundance in secondary forests and is stable and high in remnant forests. Our results suggest that functional resilience to disturbance is variable, where seed dispersal is low in some secondary forests and not others. This work provides key insights into the effects of disturbance on mutualistic interactions and how the resilience of critical ecosystem functions impacts the capacity for diversity resiliency.

Extrafloral nectar as entrée and elaiosomes as main course for ant visitors to a fireprone, mediterranean-climate shrub.

Thousands of plants produce both extrafloral nectaries (EFNs) on their leaves and nutrient-rich appendages on their diaspores (elaiosomes). Although their individual ecology is well-known, any possible functional link between these structures has almost always been ignored. Here, we recognized their co-presence in the shrub, Adenanthos cygnorum (Proteaceae), and studied their function and interaction. We observed that the same ants frequently visit both structures, seeds are attractive to vertebrate granivores but are released into a leafy cup from where they are harvested by ants and taken to their nests, from which seeds, lacking elaiosomes, germinate after fire. We showed that juvenile plants do not produce EFNs and are not visited by ants. We conclude that EFNs are not just an indirect adaptation to minimize herbivory via aggressive ant visitors (the role of a minority) but specifically enhance reproductive success in two ways: First, by inducing ants to visit the plant as a reliable food source throughout the year. Second, by promoting discovery of the seasonally available, elaiosome-bearing seeds for transport to their nests (the majority of visitors), so avoiding the risk of granivory should seeds instead fall to the ground. Parasitoid wasps play a supporting role in controlling the main insect herbivore whose larvae devour the reproductive apices. Thus, the EFN-elaiosome relationship has three components that enhance species fitness: foliage protection, seed transport, and granivore escape. A similar system has been described only once before (in an unrelated biome) and, consistent with the objectives of ecology as an integrative science, deserves wider study.

Oak Galls Exhibit Ant Dispersal Convergent with Myrmecochorous Seeds.

AbstractAnts disperse oak galls of some cynipid wasp species similarly to how they disperse seeds with elaiosomes. We conducted choice assays in field and laboratory settings with ant-dispersed seeds and wasp-induced galls found in ant nests and found that seed-dispersing ants retrieve these galls as they do myrmecochorous seeds. We also conducted manipulative experiments in which we removed the putative ant-attracting appendages ("kapéllos") from galls and found that ants are specifically attracted to kapéllos. Finally, we compared the chemical composition and histology of ant-attracting appendages on seeds and galls and found that they both have similar fatty acid compositions as well as morphology. We also observed seed-dispersing ants retrieving oak galls to their nests and rodents and birds consuming oak galls that were not retrieved by ants. These results suggest convergence in ant-mediated dispersal between myrmecochorous seeds and oak galls. Based on our observations, a protective advantage for galls retrieved to ant nests seems a more likely benefit than dispersal distance, as has also been suggested for myrmecochorous seeds. These results require reconsideration of established ant-plant research assumptions, as ant-mediated seed and gall dispersal appear strongly convergent and galls may be far more abundant in eastern North American deciduous forests than myrmecochorous seeds.

Seed Dispersal by Ants in Three Early-Flowering Plants.

Interactions between ants and plants vary from being occasionally beneficial to neutral and negative. Ant-mediated dispersal of obligatory myrmecochorous plants is considered mutualistic interaction, providing benefits to plants in terms of seed dispersal. Ants are rewarded by providing elaiosome, sugar, lipid and protein-rich appendages attached to seeds (diaspores). We experimentally examine rates of diaspore removal rates among three species of plants (snowdrop Galanthus nivalis, hollow root Corydalis cava and European wild ginger Asarum europaeum) under field conditions in two study sites in Central Europe. Diaspore morphology is altered by manipulating both elaiosome and seed size. The small-sized acorn ant Temnothorax crassispinus interacts with the snowdrop and hollow root and the moderately-sized red ant Myrmica ruginodis interacts with European wild ginger. Experimental manipulation with elaiosomes yields largely non-significant results. Diaspore removal rates are generally low (snowdrop 10%, hollow root 26%, European wild ginger 34%) probably due to the small size of ants relative to heavy diaspores. Many ants are observed to consume elaiosomes in situ (cheating). We conclude that ant-plant relationships in this case are not mutualistic but rather neutral/slightly negative, because the plants do not obtain any apparent benefits from their interactions with ants.

Fire-released seed dormancy - a global synthesis.

Seed dormancy varies greatly between species, clades, communities, and regions. We propose that fireprone ecosystems create ideal conditions for the selection of seed dormancy as fire provides a mechanism for dormancy release and postfire conditions are optimal for germination. Thus, fire-released seed dormancy should vary in type and abundance under different fire regimes. To test these predictions, we compiled data from a wide range of fire-related germination experiments for species in different ecosystems across the globe. We identified four dormancy syndromes: heat-released (physical) dormancy, smoke-released (physiological) dormancy, non-fire-released dormancy, and non-dormancy. In fireprone ecosystems, fire, in the form of heat and/or chemical by-products (collectively termed 'smoke'), are the predominant stimuli for dormancy release and subsequent germination, with climate (cold or warm stratification) and light sometimes playing important secondary roles. Fire (heat or smoke)-released dormancy is best expressed where woody vegetation is dense and fires are intense, i.e. in crown-fire ecosystems. In such environments, seed dormancy allows shade-intolerant species to take advantage of vegetation gaps created by fire and synchronize germination with optimal recruitment conditions. In grassy fireprone ecosystems (e.g. savannas), where fires are less intense but more frequent, seed dormancy is less common and dormancy release is often not directly related to fire (non-fire-released dormancy). Rates of germination, whether controls or postfire, are twice as fast in savannas than in mediterranean ecosystems. Fire-released dormancy is rare to absent in arid ecosystems and rainforests. The seeds of many species with fire-released dormancy also possess elaiosomes that promote ant dispersal. Burial by ants increases insulation of seeds from fires and places them in a suitable location for fire-released dormancy. The distribution of these dormancy syndromes across seed plants is not random - certain dormancy types are associated with particular lineages (phylogenetic conservatism). Heat-released dormancy can be traced back to fireprone floras in the 'fiery' mid-Cretaceous, followed by smoke-released dormancy, with loss of fire-related dormancy among recent events associated with the advent of open savannas and non-fireprone habitats. Anthropogenic influences are now modifying dormancy-release mechanisms, usually decreasing the role of fire as exaptive effects. We conclude that contrasting fire regimes are a key driver of the evolution and maintenance of diverse seed dormancy types in many of the world's natural ecosystems.

Odor of achlorophyllous plants' seeds drives seed-dispersing ants.

Seed dispersal by ants is an important means of migration for plants. Many myrmecochorous plants have specialized appendages in their seeds called elaiosome, which provides nutritional rewards for ants, and enable effective seed dispersal. However, some nonmyrmecochorous seeds without elaiosomes are also dispersed by ant species, suggesting the additional mechanisms other than elaiosomes for seed dispersal by ants. The seeds of the achlorophyllous and myco-heterotrophic herbaceous plant Monotropastrum humile are very small without elaiosomes; we investigated whether odor of the seeds could mediate seed dispersal by ants. We performed a bioassay using seeds of M. humile and the ant Nylanderia flavipes to demonstrate ant-mediated seed dispersal. We also analyzed the volatile odors emitted from M. humile seeds and conducted bioassays using dummy seeds coated with seed volatiles. Although elaiosomes were absent from the M. humile seeds, the ants carried the seeds to their nests. They also carried the dummy seeds coated with the seed volatile mixture to the nest and left some dummy seeds inside the nest and discarded the rest of the dummy seeds outside the nest with a bias toward specific locations, which might be conducive to germination. We concluded that, in M. humile seeds, volatile odor mixtures were sufficient to induce seed-carrying behavior by the ants even without elaiosomes.

Nest substrate, more than ant activity, drives fungal pathogen community dissimilarity in seed-dispersing ant nests.

Myrmecochory is a widespread mutualism in which plants benefit from seed dispersal services by ants. Ants might also be providing seeds with an additional byproduct benefit via reduced plant pathogen loads in the ant nest environment through their antimicrobial glandular secretions. We investigate this byproduct benefit by identifying fungal communities in ant nests and surrounding environments and quantifying fungal community change (1) through time, (2) between different nest substrates, and (3) as a function of average ant activity levels within nests (based on observed ant activity at nest entrances throughout the summer). We split fungal communities by functional guild to determine seed-dispersing ant-induced changes in the overall fungal community, the animal pathogen fungal community, the plant pathogen fungal community, and the myrmecochore pathogen fungal community. Nest substrate (soil or log) explained much of the variation in fungal community dissimilarity, while substrate occupation (ant nest or control sample) and time had no influence on fungal community composition. Average ant activity had no effect on the community turnover in fungal communities except for the myrmecochore pathogenic fungal community. In this community, higher ant activity throughout the summer resulted in more fluctuation in the pathogenic community in the ant nest. Our results are not consistent with a byproduct benefit framework in myrmecochory, but suggest that nest substrate drives dissimilarity in fungal communities. The influence of nest substrate on fungal communities has important implications for seeds taken into ant nests, as well as ant nest location choice by queens and during nest relocation.

Interactions between seed-dispersing ant species affect plant community composition in field mesocosms.

In generalized mutualisms, species vary in the quality of services they provide to their partners directly via traits that affect partner fitness and indirectly via traits that influence interactions among mutualist species that play similar functional roles. Myrmecochory, or seed dispersal by ants, is a generalized mutualism with ant species varying in the quality of dispersal services they provide to their plant partners. Variation in ant species identity can directly impact seed dispersal patterns and plant community composition; however, we know less about how interactions among seed-dispersing ant species indirectly influence plant partners. The invasive ant Myrmica rubra, is a high-quality seed-disperser in its native range that interacts with myrmecochores (ant-dispersed plants) and the high-quality seed disperser Aphaenogaster sp. in its invaded range. We use this system to examine how interactions between two functionally similar mutualist ant species influence the recruitment and community composition of ant-dispersed plants. We performed a field mesocosm experiment and a laboratory behavioural experiment to compare discovery and dominance behaviours between ant species, and seed dispersal and seedling recruitment of four myrmecochore species among intraspecific interaction treatments of each ant species and an interspecific interaction treatment. We found that M. rubra was better at discovering and dispersing seeds, but Aphaenogaster sp. was dominantly aggressive over M. rubra. Interspecific interactions dampened seed dispersal relative to dispersal by the better disperser. Despite this dampening, we found no effect of interspecific interactions on seedling recruitment. However, community composition of seedlings in the interspecific interaction treatment was more similar to composition in the aggressively dominant ant (Aphaenogaster sp.) treatment than in the better discoverer ant M. rubra treatment. We show that interspecific interactions between mutualist species in the same functional guild affect the outcome of mutualistic interactions with partner species. Despite the native ant dispersing fewer seeds, its dominance over the subordinate (invasive) ant has the potential to allow for some level of biotic resistance against the effects of M. rubra on plant communities when these species coexist.

The dynamic eggs of the Phasmatodea and their apparent convergence with plants.

The egg stages of animal life cycles are underappreciated in terms of their capacity for dispersal, protection, and biotic and abiotic interactions. Some of the most intriguing egg morphologies are seen in stick and leaf insects (Phasmatodea). Phasmids are charismatic insects, particularly due to their incredible camouflage, though a lesser-known fact is that their eggs are incredibly diverse in shape and structure, reflecting varying ecological niches. Perhaps most remarkable are those eggs which appear to resemble plant seeds in both their appearance and means of dispersal, such as via water and animal vectors. Numerous hypotheses surrounding the function of these egg morphologies and their apparent convergence with seeds have been proposed; however, empirical evidence remains lacking. Here, we present an initial synthesis of available evidence surrounding the ecology and dispersal strategies of phasmid eggs and weigh up the evidence for convergent evolution between phasmid eggs and seeds. In doing so, we highlight areas where further research is needed and discuss how the ecology of phasmid eggs may interplay with other aspects of phasmid ecology, distribution, and evolution.

Reduced dispersal at nonexpanding range margins: A matter of disperser identity.

The evolution of dispersal at range margins received much attention recently, especially in the context of dynamic range shifts, such as those following climate change. However, much less attention has been devoted to study variation in and selection on dispersal at nonexpanding range margins, where populations are often small and isolated, and empirical test is dearly missing. To fill this gap, we tested whether dispersal of an ant-dispersed perennial plant (Sternbergia clusiana) is quantitatively and/or qualitatively reduced toward a nonexpanding range margin. We evaluated plant investment in dispersal structures (elaiosome), seed removal rates, and the relative abundance, activity, and behavior of low- and high-quality seed-dispersing ants in six sites ranging from mesic Mediterranean site to arid site (>600 to <100 mm of annual rainfall, respectively), which marks the southern range margin of the species. In a set of cafeteria and baiting experiments, we found that overall seed removal rates, the contribution of high-quality dispersers, maximum dispersal distance and dispersal-conducive ant behavior decreased toward range margins. These findings agree with a lower investment in reward by range margin plant populations, as reflected by lower elaiosome/seed ratio, but not by variation in the reward chemistry. More than variation in traits controlled by the plants, the variation in ant-seed interactions could be attributed to reduced presence and activity of the more efficient seed-dispersing ants in the marginal populations. Specifically, we found a mismatch between local distribution of potentially effective seed dispersers and that of the plant, even though those dispersers were observed in the study site. Interestingly, although the observed variation in the outcome of ant-seed interactions supported the prediction of reduced dispersal at nonexpanding range margins with small and isolated populations, the underlying mechanism seems to be incidental difference in the seed-dispersing ant community rather than a plant-mediated response to selection.

Effects of seed morphology and elaiosome chemical composition on attractiveness of five Trillium species to seed-dispersing ants.

Morphological and chemical attributes of diaspores in myrmecochorous plants have been shown to affect seed dispersal by ants, but the relative importance of these attributes in determining seed attractiveness and dispersal success is poorly understood. We explored whether differences in diaspore morphology, elaiosome fatty acids, or elaiosome phytochemical profiles explain the differential attractiveness of five species in the genus Trillium to eastern North American forest ants. Species were ranked from least to most attractive based on empirically-derived seed dispersal probabilities in our study system, and we compared diaspore traits to test our hypotheses that more attractive species will have larger diaspores, greater concentrations of elaiosome fatty acids, and distinct elaiosome phytochemistry compared to the less attractive species. Diaspore length, width, mass, and elaiosome length were significantly greater in the more attractive species. Using gas chromatography-mass spectrometry, we found significantly higher concentrations of oleic, linoleic, hexadecenoic, stearic, palmitoleic, and total fatty acids in elaiosomes of the more attractive species. Multivariate assessments revealed that elaiosome phytochemical profiles, identified through liquid chromatography-mass spectrometry, were more homogeneous for the more attractive species. Random forest classification models (RFCM) identified several elaiosome phytochemicals that differed significantly among species. Random forest regression models revealed that some of the compounds identified by RFCM, including methylhistidine (α-amino acid) and d-glucarate (carbohydrate), were positively related to seed dispersal probabilities, while others, including salicylate (salicylic acid) and citrulline (L-α-amino acid), were negatively related. These results supported our hypotheses that the more attractive species of Trillium-which are geographically widespread compared to their less attractive, endemic congeners-are characterized by larger diaspores, greater concentrations of fatty acids, and distinct elaiosome phytochemistry. Further advances in our understanding of seed dispersal effectiveness in myrmecochorous systems will benefit from a portrayal of dispersal unit chemical and physical traits, and their combined responses to selection pressures.

Invasive ants disperse seeds farther than native ants, affecting the spatial pattern of seedling recruitment and survival.

Mutualists can vary in the quantity and quality of service which they provide to their partners. Variation in seed disperser quality depends on seed-processing traits, dispersal distance, and deposition location, all of which ultimately affect plant fitness. Here, we compared these aspects of seed dispersal quality between a native and an invasive ant species, and examined how they affect competition and plant performance. Using experimental mesocosm communities, we examined how these two ant species affect the spatial pattern of recruitment and establishment for four myrmecochorous plant species, including one invasive species. We measured the locations of dispersed seedlings relative to ant nests, adult plants, and other dispersed seedlings, as well as measured the effects of location on plant performance. The invasive ant, Myrmica rubra, secondarily dispersed seeds farther from its nests, creating a less clumped pattern of seedling recruitment compared to the native ant, Aphaenogaster rudis. Plant species responded differently to dispersal. Invasive seedlings recruited farther from adult plants than native seedlings, and had higher survival the farther they were from conspecifics. In contrast, native plants had higher survival and grew taller when dispersed farther from invasive plants. We show that seed-dispersing ant partners differ in mutualist quality creating differences in dispersal distance and deposition location that affects a plant's competitive environment. Our results reveal the potential for long-term consequences on plant community structure with changing ant partner identity. We emphasize the need to examine dispersal quality in addition to quantity to uncover the importance of partner identity in structuring communities.

Origin and divergence of Afro-Indian Picrodendraceae: linking pollen morphology, dispersal modes, fossil records, molecular dating and paleogeography.

The pantropical Picrodendraceae produce mostly spheroidal to slightly oblate, echinate pollen grains equipped with narrow circular to elliptic pori that can be hard to identify to family level in both extant and fossil material using light microscopy only. Fossil pollen of the family have been described from the Paleogene of America, Antarctica, Australia, New Zealand, and Europe, but until now none have been reported from Afro-India. Extant pollen described here include representatives from all recent Picrodendraceae genera naturally occurring in Africa and/or Madagascar and south India and selected closely related tropical American taxa. Our analyses, using combined light microscopy and scanning electron microscopy, show that pollen of the Afro-Indian genera encompass three morphological types: Type 1, comprising only Hyaenanche; Type 2, including Aristogeitonia, Mischodon, Oldfieldia and Voatamalo; Type 3, comprising the remaining two genera, Androstachys and Stachyandra. Based on the pollen morphology presented here it is evident that some previous light microscopic accounts of spherical and echinate fossil pollen affiliated with Arecaceae, Asteraceae, Malvaceae, and Myristicaceae from the African continent could belong to Picrodendraceae. The pollen morphology of Picrodendraceae, fossil pollen records, a dated intra-familial phylogeny, seed dispersal modes, and the regional Late Cretaceous to early Cenozoic paleogeography, together suggest the family originated in the Americas and dispersed from southern America across Antarctica and into Australasia. A second dispersal route is believed to have occurred from the Americas into continental Africa via the North Atlantic Land Bridge and Europe.

Investment in reward by ant-dispersed plants consistently selects for better partners along a geographic gradient.

Seed dispersal by ants (myrmecochory) is an asymmetric, presumably mutualistic interaction, where a few ant species benefit many plants. Myrmecochorous plants express specialized adaptations, most notably a large elaiosome, which promote interactions with efficient seed dispersers while decreasing interactions with poor dispersers, resulting in de facto partner choice. However, because variation in plants' investment in reward and ant response to them may vary spatially and temporally, it is unclear whether such specialization is consistent along geographic gradients; especially towards myrmecochory's range margin. To answer this question on context-dependent partner choice, we first estimated variation in reward investment by co-occurring myrmecochores along a steep environmental gradient in a Mediterranean region. Second, we tested whether variation in plant investment in reward was positively and consistently correlated with the quality of dispersal plant received along the same gradient. Using in situ cafeteria experiments, we simultaneously presented diaspores of locally co-occurring myrmecochorous species to ants of two guilds representing high- and low-quality dispersers. We then recorded ant-seed behaviour, seed preference and seed removal rates for each ant guild. We found both overall and within-site high variation among plant species in the total and relative investment in elaiosomes. Both ant guilds removed substantial proportion of the seeds. However, scavenging ants (high-quality dispersers) clearly preferred diaspores with larger elaiosomes, whereas granivorous ants (low-quality dispersers) exhibited no preference. Furthermore, both the variation in plant traits and the corresponding response of different ant guilds were consistent along the studied geographic gradient. This consistency holds even when granivores, which removed seeds in a non-selective fashion and provided apparently low-quality seed dispersal services, were, at least numerically, the dominant ant guild. This dominance and the consistency of the partner choice shed light on the functionality of elaiosomes at the margins of myrmecochory's distribution.

Variation in ant-mediated seed dispersal along elevation gradients.

Ant-mediated seed dispersal, also known as myrmecochory, is a widespread and important mutualism that structures both plant and ant communities. However, the extent to which ant functional types (e.g., granivorous generalists vs. myrmecochorous ants) across environmental gradients affect seed removal rates is not fully understood. We used a replicated, standardized seed removal experiment along elevation gradients in four mountain ranges in the southwestern United States to test predictions that: (1) seed removal rates would be greater at lower elevations, and (2) seed species identity influences seed removal rates (i.e., seeds from their native elevation range would be removed at higher rates than seeds outside of their native elevation range). Both predictions were supported. Seed removal rates were ∼25% higher at lower elevation sites than at higher elevation sites. The low elevation Datura and high elevation Iris were removed at higher rates in their respective native ranges. We attribute observed differences in dispersal rates to changes in ant community composition, functional diversity, and abundance. We also suggest that temperature variation along the elevation gradient may explain these differences in seed removal rates.