Exploitation of Fallen Diaspores by Ants: Are there Ant–Plant Partner Choices?

Ants frequently interact with fleshy fallen diaspores (fruits or seeds) not adapted for ant‐dispersal. Such interactions are usually considered as opportunistic, but recent evidence has indicated that these ants may differ in their effects on diaspore survival and plant recruitment. We investigated if partner choices are recognizable among genera of ants and plants, and if ant and plant traits may influence such preferences in cerrado (savanna‐like vegetation) from southeast Brazil. During a 2‐yr period, 521 ant–diaspore interactions were recorded through various methods, yielding 71 ant species and 38 plant species. Exploitation of fallen diaspores was common among several ant genera, and included carnivorous, omnivorous, and fungivorous ants. Contrary to others areas around the world, where true myrmecochory (seed dispersal by ants) is common among shrubs, ants also exploited diaspores from several cerrado trees. Plant life form, diaspore size, and ant body size did not seem to explain the pattern of interactions observed. Two subsets of preferential interactions, however, segregated fungivorous ants from another group composed of carnivorous and omnivorous ants, probably influenced by the chemical composition of the plant diaspores. Omnivorous ants usually remove the fleshy portion of diaspores on spot and probably provide limited benefits to plants. Carnivorous and fungivorous ants usually remove the whole diaspore to the nest. As each of these ant groups may influence the fitness of diaspores in different ways, there are possible subtle pathways for the evolution of partner choices between ants and these non‐myrmecochorous diaspores.     

Dispersal of non-myrmecochorous plants by a “keystone disperser” ant in a Mediterranean habitat reveals asymmetric interdependence

In contrast to other plant–animal mutualisms, seed dispersal interactions, and particularly seed dispersal by ants, are generally considered asymmetric, non-specialized relationships in which dispersers depend less on plants than vice versa. Although myrmecochory is well understood in many terrestrial ecosystems, dispersal of non-elaiosome-bearing seeds by ants has barely been studied outside the Neotropics. Aphaenogaster senilis, a common ant in Southern Spain, collects a great variety of non-myrmecochorous diaspores along with insect prey. At our study site, fleshy fruits of Arum italicum, Phillyrea angustifolia and Pistacia lentiscus represent up to one-fourth of the items collected by A. senilis from June to November. However, they are mostly ignored by other ants. In the laboratory, the addition of A. italicum fruits to A. senilis insect-based diet increased male production and both worker and queen pupae size. Seeds were transported up to 8 m away from the mother plant and deposited in a favorable habitat allowing a relatively high proportion of germination. Given important differences in seed production between species, our data suggest that A. senilis removes virtually all seeds of A. italicum, but a negligible fraction of P. lentiscus seeds. We conclude that in contrast to the common view, dispersal of non-myrmecochorous Mediterranean plants by ants might be an important phenomenon. Keystone disperser ants like A. senilis probably obtain an important fitness advantage from non-myrmecochorous diaspore collection. However, plant benefit may vary greatly according to the amount of seeds per individual plant and the existence of alternative dispersal agents.     

Diaspore removal and potential dispersers of the rare and protected Paeonia officinalis L. (Paeoniaceae) in a changing landscape

Paeonia officinalis L., a rare and protected species, mostly occurs in open and semi‐open habitats and is often threatened by forest and shrubland spread. To explore the still undocumented dispersal features of this species, we address the following questions. What are the relative roles of ants, small rodents, and birds as diaspore removers in open habitat and woodland? Which animal groups constitute the potential disperser assemblage and how do they shape the spatial patterns of seed dispersal? Do diaspores fit the ornithochory syndrome or do they only mimic fleshy fruits? Two experiments were performed to quantify diaspore fall and diaspore removal by animal groups, above ground and on the ground. Ants did not contribute to dispersal. In open habitats, no seed removal was detected, either on follicles or once diaspores had fallen to the ground. In woodland, diaspores were weakly removed by vertebrates on follicles and were mainly removed by rodents on the ground. As a consequence, we suggest that long‐distance dispersal events are very rare, weakening the possible escape into space of populations subject to forest spread. Several traits indicate that diaspores fit the ornithochory syndrome, but other traits are strongly reminiscent of mimetic diaspores deceiving bird dispersers. © 2007 CNRS. Journal compilation © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 154 , 13–25.     

Dispersal traits and dispersal patterns in an oro-Mediterranean thorn cushion plant formation of the eastern High Atlas, Morocco

This study describes the dispersal traits and dispersal patterns of 51 perennial plant species belonging to 19 families in an oro-Mediterranean thorn cushion plant formation on the High Atlas, Morocco. Diaspore type, mass, number, antitelechoric mechanisms and dispersal time were studied with respect to growth forms, dispersal modes and spatial dispersal. Species spanned 10⁵ range of diaspore mass, which coincided with those found in other high mountain regions. Diaspore mass was significantly higher in trees and shrubs than in semi-shrubs and perennial herbs. Barochorous and zoochorous species are more likely to have heavy diaspores, whereas anemochorous and ballistic species have a medium diaspore mass and semachorous and ombro-hydrochorous species have low diaspore mass. Diaspore number was significantly higher in trees and shrubs than in semi-shrubs and perennial herbs. The barochorous, ombro-hydrochorous and zoochorous species tended to produce higher diaspore numbers than species with other dispersal modes. Bradyspory was well-developed by trees and large shrubs dispersed by biotic vectors. Synaptospermy was represented by its long dispersal component. Myxospermy was significantly associated with semi-shrubs and perennial herbs with restricted spatial dispersal. It seems that ombro-hydrochory combined with myxospermy and a high number of light seeds is an efficient mechanism that ensures successful establishment of the most typical and endemic thorn cushion plant species, such as Alyssum, Vella and Ormenis. In our study area, the highest dispersal availability was synchronized with the dry summer season (July–August) and the beginning of the rainy months (September). The dispersal peak for the wind-dispersed species, which is the most effective primary dispersal mode, occurs during the dry season, while dispersal for the biotic-dispersed species takes place throughout the year.     

Importance of elaiosome size to removal of ant-dispersed seeds

Several studies relate removal of diaspores from different ant-dispersed plant species to the size of their claiosomes. This study is the first one to relate intraspecific variation in claiosome size to removal of diaspores by ants. This approach circumvents the problem posed by interspecific variation in chemistry and morphology of elaiosomes. We used the systemHepatica nobilis Mill. (Ranunculaceae) andMyrmica ruginodis Nyl. The elaiosome was the attractive part of the diaspore and its attractiveness decreased quickly after release from the fruit. In experimental runs where ants collected diaspores, the elaiosome size of accepted diaspores was larger than of those rejected and the largest diaspores were collected first. Since size of diaspore, elaiosome and achene were correlated, the diaspores that were removed first had both the largest elaiosome and achene. However, our experiments suggested that elaiosome size was more important to removal than achene size or the elaiosome:achene size ratio. If ant dispersal improves plant fitness, elaiosome size and hence diaspore size would be expected to increase over time. However, such directional selection mediated by the ants is probably counterbalanced by the plant. Seed predators and a negative trade-off between number and size of seeds would, among other factors, select for smaller diaspore size,i.e. counteract the effect of the ants' preference for larger elaiosomes.     

Post-dispersal seed predation and the establishment of vertebrate dispersed plants in Mediterranean scrublands

The post-dispersal fate of seeds and fruit (diaspores) of three vertebrate-dispersed trees, Crataegus monogyna, Prunus mahaleb and Taxus baccata, was studied in the Andalusian highlands, south-eastern Spain. Exclosures were used to quantify separately the impact of vertebrates and invertebrates on seed removal in relation to diaspore density and microhabitat. The three plant species showed marked differences in the percentage of diaspores removed, ranging from only 5% for C. monogyna to 87% for T. baccata. Although chaffinches (Fringilla coelebs) fed on diaspores, rodents (Apodemus sylvaticus) were the main vertebrate removers of seed and fruit. Two species of ant (Cataglyphis velox and Aphaenogaster iberica) were the only invertebrates observed to remove diaspores. However, the impact of ants was strongly seasonal and they only removed P. mahaleb fruit to any significant extent. While removal of seed by rodents was equivalent to predation, ants were responsible for secondary dispersal. However, their role was limited to infrequent, small-scale redistribution of fruit in the vicinity of parent trees. Rodents and ants differed in their use of different microhabitats. Rodents foraged mostly beneath trees and low shrubs and avoided open areas while the reverse was true of ants. Thus, patterns of post-dispersal seed removal will be contigent on the relative abundance and distribution of ants and rodents. Studies which neglect to quantify separately the impacts of these two guilds of seed removers may fail to elucidate the mechanisms underlying patterns of post-dispersal seed removal. The coincidence of both increased seed deposition by the main avian dispersers (Turdus spp.) and increased seed predation with increasing vegetation height suggested that selection pressures other than post-dispersal seed predation shape the spatial pattern of seed dispersal. Rather than providing a means of escaping post-dispersal seed predators, dispersal appears to direct seeds to microhabitats most suitable for seedling survival. Nevertheless, the reliance of most vertebrate-dispersed trees on regeneration by seed and the absence of persistent soil seed banks imply that post-dispersal seed predators may exert a strong influence on the demography of the plants whose seeds they consume. Even where microsites are limited, the coincidence of the most suitable microhabitats for seedling establishment with those where seed predation is highest provide a means by which selective seed predators can influence community composition. Received: 19 August 1996 / Accepted: 25 January 1997     

New mutualism for old: indirect disruption and direct facilitation of seed dispersal following Argentine ant invasion

The indirect effects of biological invasions on native communities are poorly understood. Disruption of native ant communities following invasion by the Argentine ant (Linepithema humile) is widely reported to lead indirectly to the near complete collapse of seed dispersal services. In coastal scrub in southeastern Australia, we examined seed dispersal and handling of two native and two invasive alien plant species at Argentine ant-invaded or -uninvaded sites. The Argentine ant virtually eliminates the native keystone disperser Rhytidoponera victoriae, but seed dispersal did not collapse following invasion. Indeed, Argentine ants directly accounted for 92% of all ant-seed interactions and sustained overall seed dispersal rates. Nevertheless, dispersal quantity and quality among seed species differed between Argentine ant-invaded and -uninvaded sites. Argentine ants removed significantly fewer native Acacia retinodes seeds, but significantly more small seeds of invasive Polygala myrtifolia than did native ants at uninvaded sites. They also handled significantly more large seeds of A. sophorae, but rarely moved them >5 cm, instead recruiting en masse, consuming elaiosomes piecemeal and burying seeds in situ. In contrast, Argentine ants transported and interred P. myrtifolia seeds in their shallow nests. Experiments with artificial diaspores that varied in diaspore and elaiosome masses, but kept seed morphology and elaiosome quality constant, showed that removal by L. humile depended on the interaction of seed size and percentage elaiosome reward. Small diaspores were frequently taken, independent of high or low elaiosome reward, but large artificial diaspores with high reward instead elicited mass recruitment by Argentine ants and were rarely moved. Thus, Argentine ants appear to favour some diaspore types and reject others based largely on diaspore size and percentage reward. Such variability in response indirectly reduces native seed dispersal and can directly facilitate the spread of an invasive alien shrub.     

Indirect effects of alternative food resources in an ant–plant interaction

The seeds of many plant species present a food body that is consumed by animal dispersers. In theory, if the animals are polyphagous, the availability of alternative food resource other than the diaspore itself may influence its dispersal and survival. We used the myrmecochore Helleborus foetidus L. (Ranunculaceae), the seeds of which are attached to a lipid-rich elaiosome that is attractive to ants, as a model system to investigate (1) whether alternative foods that are present along with the plant affect ant foraging behavior and diaspore removal and (2) whether food availability in an ant nest affects seed predation and germination. In a field experiment, artificial diaspore depots were offered together with either sugar, insect corpses, seed, or no food (control). Contrary to the prediction that ants would rather concentrate their foraging effort on the highly rewarding alternative foods only, many workers, attracted by the sugar, switched to the hellebore diaspores, which significantly enhanced removal rate. Results obtained in the laboratory further indicated that the larvae of Aphaenogaster iberica (a major seed disperser) predated more on the H. foetidus embryos when no alternative food was available. This, in turn, slightly reduced seed germination. Overall, these results shed light, for the first time, on the potential indirect effects of alternative resources on the fate of diaspores adapted for ant dispersal.     

Effect of parameters of myrmecochorous syndrome on rates of removal of Corydalis cava and Pulmonaria obscura by Formica polyctena ants]

Complex of adaptations of myrmecochorous plants to the ant dispersal composes so called myrmecochorous syndrome. To study the effect of some morphological and anatomical adaptations on diaspore attractiveness for ants, the field experiments with diaspores and their extracts were carried out. It was shown that chemical cues attracting ants are located in different parts of diaspores: in elaiosome only (C. cava) or in elaiosome, coverage of external layer of the fruit and in the fruit collar (P. obscura). Anatomy of adaptive structure of diaspores is also discussed together with the results of field experiments.     

Size and lipid content of nonmyrmecochorous diaspores: effects on the interaction with litter-foraging ants in the Atlantic rain forest of Brazil

Ants are often attracted to diaspores not adapted for dispersal by ants. These diaspores may occasionally benefit from this interaction. We selected six nonmyrmecochorous plant species (Virola oleifera, Eugenia stictosepala, Cabralea canjerana, Citharexylum myrianthum, Alchornea glandulosa and Hyeronima alchorneoides) whose diaspores differ in size and lipid content, and investigated how these features affect the outcome of ant-diaspore interactions on the floor of a lowland Atlantic forest of Southeast Brazil. A total of 23 ant species were seen interacting with diaspores on the forest floor. Ants were generally rapid at discovering and cleaning the diaspore pulp or aril. Recruitment rate and ant attendance were higher for lipid-rich diaspores than for lipid-poor ones. Removal rate and displacement distance were higher for small diaspores. The large ponerine ant Pachycondyla striata, one of the most frequent attendants to lipid-rich arillate diaspores, transported the latter into their nests and discarded clean intact seeds on refuse piles outside the nest. Germination tests with cleaned and uncleaned diaspores revealed that the removal of pulp or aril may increase germination success in Virola oleifera, Cabralea canjerana, Citharexylum myrianthum and Alchornea glandulosa. Gas chromatography analyses revealed a close similarity in the fatty acid composition of the arils of the lipid-rich diaspores and the elaiosome of a typical myrmecochorous seed (Ricinus communis), corroborating the suggestion that some arils and elaiosomes are chemically similar. Although ant-derived benefits to diaspores – secondary dispersal and/or increased germination – varied among the six plant species studied, the results enhanced the role of ant-diaspore interactions in the post-dispersal fates of nonmyrmecochorous seeds in tropical forests. The size and the lipid-content of the diaspores were shown to be major determinants of the outcome of such interactions.     

Savanna fires increase rates and distances of seed dispersal by ants

Myrmecochory (seed dispersal by ants) is a prominent dispersal mechanism in many environments, and can play a key role in local vegetation dynamics. Here we investigate its interaction with another key process in vegetation dynamics—fire. We examine ant dispersal of seeds immediately before and after experimental burning in an Australian tropical savanna, one of the world’s most fire-prone ecosystems. Specifically, our study addressed the effects of burning on: (1) the composition of ants removing seeds, (2) number of seed removals, and (3) distance of seed dispersal. Fire led to higher rates of seed removal post-fire when compared with unburnt habitat, and markedly altered dispersal distance, with mean dispersal distance increasing more than twofold (from 1.6 to 3.8 m), and many distance dispersal events greater than the pre-fire maximum (7.55 m) being recorded. These changes were due primarily to longer foraging ranges of species of Iridomyrmex, most likely in response to the simplification of their foraging landscape. The significance of enhanced seed-removal rates and distance dispersal for seedling establishment is unclear because the benefits to plants in having their seeds dispersed by ants in northern Australia are poorly known. However, an enhanced removal rate would enhance any benefit of reduced predation by rodents. Similarly, the broader range of dispersal distances would appear to benefit plants in terms of reduced parent–offspring conflict and sibling competition, and the location of favourable seedling microsites. Given the high frequency of fire in Australian tropical savannas, enhanced benefits of seed dispersal by ants would apply for much of the year.     

黄土丘陵沟壑区80种植物繁殖体形态特征及其物种分布

植物繁殖体的形态特征及其物种分布是植物长期适应环境的结果,对退化生态系统的更新与恢复具有重要的理论与实践指导意义。研究了80种植物的繁殖体形态特征及与其物种分布的关系,结果表明：1）繁殖体重量差别较大（0. 020—357.428 mg）,主要集中在1—9.999 mg（占52.5%）。2）繁殖体的形状指数FI变化范围较大（1.06—12.93）,61.3%的植物繁殖体的形状近圆球形。3）33种植物繁殖体具有明显的附属物,包括毛、翅、芒和刺等。4）87.5%的植物繁殖体表面具有各种纹饰（棱、被毛、纹和颗粒状等）;繁殖体颜色以褐色和黑色为主,少数繁殖体颜色鲜艳;6种植物繁殖体具有吸水分泌粘液的特性。5）黄土丘陵沟壑区繁殖体重量小或形状近圆球形的植物分布数量多、范围广。6）繁殖体具有毛、翅和芒附属物或吸湿分泌粘液的植物在黄土丘陵沟壑区较繁殖体无附属物的植物能够分布更多、更广。繁殖体这些有利于物种分布的形态特征对于植物适应该区干旱与土壤侵蚀干扰具有重要的生态学意义,而且可用于指导该区人工补播促进植被恢复的物种选择。     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

D3: The Dispersal and Diaspore Database – Baseline data and statistics on seed dispersal

Seed dispersal is hard to measure, and there is still a lack of knowledge about dispersal-related traits of plant species. Therefore, we developed D³, the Dispersal and Diaspore Database (available at www.seed-dispersal.info), which aims at simplifying ecological and evolutionary analyses by providing and integrating various items related to seed dispersal: empirical studies, functional traits, image analyses and ranking indices (quantifying the adaptation to dispersal modes).Currently, the database includes data for more than 5000 taxa and 33 items as well as digital images of diaspores (i.e. the dispersal units), seeds, fruits and infructescences. The included items cover common traits like diaspore mass, size, shape, terminal velocity and seed number per diaspore. Furthermore, we present newly or further developed items like ecomorphological categorizations of the diaspore and fruit as well as information from literature on prevailing dispersal modes. Finally, we introduce several items which are not covered in other databases yet: surface structure and form of the diaspore, the exposure of the diaspores in the infructescence and dispersal rankings. Dispersal rankings allow estimations of how well certain species are adapted to a specific dispersal mode in comparison to a larger species set. They are calculated as the percentile rank of an indicator of species’ dispersal potential in relation to a larger species set.Especially for the new and further developed items we outline the basic concepts in detail, describe the measurement and categorization methods and show how to interpret and integrate these data for single species as well as for larger species sets. Thereby, we calculate baseline statistics of seed dispersal of the Central European flora. We found that diaspores of 72% of the taxa show specializations related to long-distance dispersal, i.e. most often elongated appendages or nutrient-rich tissues. Diaspore masses, sizes and terminal velocities vary over several orders of magnitude and can be approximated by lognormal distributions.     

Hunting of Mammals Reduces Seed Removal and Dispersal of the Afrotropical Tree Antrocaryon klaineanum (Anacardiaceae)

Throughout the tropics, mammalian seed dispersers are being driven to local extinction by intense hunting pressure, generating concern not only about the loss of these species, but also about the consequences for the plants they disperse. We compared two rain forest sites in Cameroon—one with heavy hunting pressure and one protected from hunting—to appraise the loss of mammalian seed dispersers and to assess the impact of this loss on seed removal and seed dispersal of Antrocaryon klaineanum (Anacardiaceae), a mammal-dispersed tree. Surveys of arboreal frugivores indicate that three of the five monkey species, as well as chimpanzee and gorilla, have been extirpated from the hunted forest. Diaspore counts underneath A. klaineanum adults (six trees per site) indicate that seed removal is severely reduced in the hunted forest. Finally, genetic maternity exclusion analysis (using 3–7 nuclear microsatellite loci) of maternally inherited endocarp tissue from diaspores collected under the canopies of 12 fruiting "mother" trees (six trees per site) revealed that seed dispersal in the hunted forest is also greatly reduced. In the hunted forest with reduced mammal dispersal agents, only 1 of the 53 assayed endocarps (2%) did not match the mother and was determined to be from a dispersed diaspore. By contrast, in the protected forest, 20 of the 48 assayed endocarps (42%) were from dispersed diaspores. This study provides strong evidence that loss of dispersal agents can lead to reduced seed removal and loss of seed dispersal, disrupting the seed dispersal cycle.     

Signals Can Trump Rewards in Attracting Seed-Dispersing Ants

Both rewards and signals are important in mutualisms. In myrmecochory, or seed dispersal by ants, the benefits to plants are relatively well studied, but less is known about why ants pick up and move seeds. We examined seed dispersal by the ant Aphaenogaster rudis of four co-occurring species of plants, and tested whether morphology, chemical signaling, or the nutritional quality of fatty seed appendages called elaiosomes influenced dispersal rates. In removal trials, ants quickly collected diaspores (seeds plus elaiosomes) of Asarum canadense, Trillium grandiflorum, and Sanguinaria canadensis, but largely neglected those of T. erectum. This discrepancy was not explained by differences in the bulk cost-benefit ratio, as assessed by the ratio of seed to elaiosome mass. We also provisioned colonies with diaspores from one of these four plant species or no diaspores as a control. Colonies performed best when fed S. canadensis diaspores, worst when fed T. grandiflorum, and intermediately when fed A. canadense, T. erectum, or no diaspores. Thus, the nutritional rewards in elaiosomes affected colony performance, but did not completely predict seed removal. Instead, high levels of oleic acid in T. grandiflorum elaiosomes may explain why ants disperse these diaspores even though they reduce ant colony performance. We show for the first time that different elaiosome-bearing plants provide rewards of different quality to ant colonies, but also that ants appear unable to accurately assess reward quality when encountering seeds. Instead, we suggest that signals can trump rewards as attractants of ants to seeds.     

Species-specific detection of Lobaria pulmonaria (lichenized ascomycete) diaspores in litter samples trapped in snow cover

The foliose lichen Lobaria pulmonaria has suffered a substantial decline in central and northern Europe during the twentieth century and is now considered to be critically endangered in many European lowland regions. Based on demographic studies, it has been proposed that under the present environmental conditions and forest management regimes, dispersal of diaspores and subsequent establishment of new thalli are insufficient to maintain the remnant small lowland populations. Chances of long-term survival may therefore be reduced. The data and analytical power of these demographic studies are limited. Since lichen diaspores show very few species-specific morphological characteristics, and are therefore almost indistinguishable, the accurate assessment of diaspore flux would be a fundamental first step in better understanding the life cycle of L. pulmonaria. Here we present a new molecular approach to investigate the dispersal of L. pulmonaria diaspores in its natural environment by specifically identifying small amounts of DNA in snow litter samples at varying distances from known sources. We used a species-specific polymerase chain reaction (PCR) primer pair to amplify the ribosomal internal transcribed spacer region (ITS rDNA) and a sensitive automated PCR product detection system using fluorescent labelled primers. We detected considerable amounts of naturally dispersed diaspores, deposited as far as 50 m away from the closest potential source. Diaspores were only found in the direction of the prevailing wind. Diaspore deposition varied from 1.2 diaspores per m(2) per day at 50 m distance from the source to 15 diaspores per m(2) per day at 1 m distance. The method described in this paper opens up perspectives for studies of population dynamics and dispersal ecology mainly in lichenized ascomycetes but also in other organisms with small, wind-dispersed diaspores.     

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.     

Role of nurse plants in Araucaria Forest expansion over grassland in south Brazil

Abstract Araucaria Forest expansion over grassland takes place under wet climate conditions and low disturbance and it is hypothesized that isolated trees established on grassland facilitate the establishment of forest woody species beneath their canopies. Forest with Araucaria angustifolia is a particular type of Brazilian Atlantic Forest and the main forest type on the highland plateau in south Brazil, often forming mosaics with natural Campos grassland. The objectives of this paper were to evaluate the role of isolated shrubs and trees as colonization sites for seedlings of Araucaria Forest woody species on grassland, to determine which species function as preferential nurse plants in the process and the importance of vertebrate diaspore dispersal on the structure of seedling communities beneath nurse plants. The study was conducted in São Francisco de Paula, Rio Grande do Sul State, where we sampled isolated shrubs and trees in natural grassland near Araucaria Forest edges. Seedlings were counted and identified, and seedling diaspore dispersal syndromes, size and colour were registered. We detected 11 woody species with a potential role in nucleating grassland colonization by forest species. Beneath the canopies of nurse plants more forest species seedlings were found compared with open field grassland and the seedlings had diaspores mostly dispersed by vertebrates. Also, more seedlings were found under the canopy of A. angustifolia than beneath other nurse plant species. We conclude that A. angustifolia trees established on grassland act as nurse plants, by attracting disperser birds that promote colonization of the site by other forest species seedlings, and that under low level of grassland disturbance, conservation of frugivorous vertebrate assemblages may increase forest expansion over natural grassland and also facilitate the regeneration of degraded forest areas.     

DISPERSAL ECOLOGY OF CAREX PEDUNCULATA (CYPERACEAE), A NEW NORTH AMERICAN MYRMECOCHORE

Carex pedunculata is the first North American species of the Cyperaceae that is identified as a myrmecochore. Many morphological and phenological features of this species and its breeding system are interpreted as adaptive for seed dispersal by ants. In laboratory tests, workers of the ant species Aphaenogaster rudis carry the diaspores to the nest, eat the elaiosomes, carry larvae to the elaiosomes to feed, and deposit diaspores whose elaiosomes have been eaten with other nest debris. The achenes then germinate. Achenes will also germinate without any handling by ants. Workers will also transport diaspores with uneaten elaiosomes when the nest is disturbed. Greenhouse tests show that seedling growth is greatly inhibited if a diaspore remains near the parent plant and cohort seedlings. Field studies of natural populations identify rotting logs (the location of ant nests) as forest floor microsites for colonization of C. pedunculata and other myrmecochores. Ant nesting behavior may pattern much of the herb stratum. This species is self-compatible, and single seeds may start successful new populations. Three processes contribute to population growth: vegetative growth, germination of untransported diaspores, and germination of ant-transported diaspores.