An ant–plant mutualism induces shifts in the protist community structure of a tank-bromeliad

Although ants may induce community-wide effects via changes in physical habitats in terrestrial environments, their influence on aquatic communities living in plant-held waters remains largely underexplored. The neotropical tank-bromeliad Aechmea mertensii (Bromeliaceae) occurs along forest edges in ant-gardens initiated by Camponotus femoratus or by Pachycondyla goeldii. Its leaves form wells that hold rainwater and provide suitable habitats for many aquatic organisms. We postulated that these ant–plant mutualisms indirectly affect the microbial community structure via changes in the environmental conditions experienced by the plants. To test this hypothesis, we analyzed the protist communities from 63 tank-bromeliads associated with either C. femoratus or P. goeldii (hereafter Cf-Aechmea and Pg-Aechmea) along a forest edge in French Guiana. For each plant, a large number of environmental variables (including habitat structure, food resources, incident radiation and the presence of aquatic invertebrates) were quantified to determine their relative importance in driving any observed differences across ant-associated plants. Pg-Aechmea are located in sun-exposed areas and hold low volumes of water and low amounts of detritus, whereas Cf-Aechmea are located in partially shaded areas and accumulate higher amounts of water and detritus. Protists (i.e., protozoa and algae) inhabiting Cf-Aechmea exhibit greater richness and abundances than those in Pg-Aechmea. Variations in detritus content, number of leaves, incident radiation, and the epiphyte richness of the ant-garden were the main factors explaining the variation in protist richness. A shift in the functional group composition of protists between bromeliads tended by different ant species suggested that mutualistic ants indirectly mediate changes in the microbial food web.     

Long-term temporal variation in the organization of an ant–plant network

Background and Aims

Functional groups of species interact and coevolve in space and time, forming complex networks of interacting species. A long-term study of temporal variation of an ant–plant network is presented with the aims of: (1) depicting its structural changes over a 20-year period; (2) detailing temporal variation in network topology, as revealed by nestedness and modularity analysis and other parameters (i.e. connectance, niche overlap); and (3) identifying long-term turnover in taxonomic structure (i.e. switches in ant resource use or plant visitor assemblages according to taxa).

Methods

Fieldwork was carried out at La Mancha, Mexico, and ant–plant interactions were observed between 1989 and 1991, between 1998 and 2000, and between May 2010 and 2011. Occurrences of ants on extrafloral nectaries (EFNs) were recorded. The resulting ant–plant networks were constructed from qualitative presence–absence data determined by a species–species matrix defined by the frequency of occurrence of each pairwise ant–plant interaction.

Key Results

Network variation across time was stable and a persistent nested structure may have contributed to the maintenance of resilient and species-rich communities. Modularity was lower than expected, especially in the most recent networks, indicating that the community exhibited high overlap among interacting species (e.g. few species were hubs in the more recent network, being partly responsible for the nested pattern). Structurally, the connections created among modules by super-generalists gave cohesion to subsets of species that otherwise would remain unconnected. This may have allowed an increasing cascade-effect of evolutionary events among modules. Mutualistic ant–plant interactions were structured 20 years ago mainly by the subdominant nectarivorous ant species Camponotus planatus and Crematogaster brevispinosa, which monopolized the best extrafloral nectar resources and out-competed other species with broader feeding habits. Through time, these ants, which are still present, lost their position as network hubs and diminished in their importance in structuring the network; simultaneously, plants gained in importance.

Conclusions

The long-term network analysis reveals a decrease in attended plant species richness, a notable increase in plant species participation from 1990 to 2010 (sustained by less plant taxonomic similarity in the older 1990 network), an increase in the number of ant species and a diminishing dominance of super-generalist ants. The structure of the community has remained highly nested and connected with low modularity, suggesting overall a more participative, homogeneous, cohesive interaction network. Although previous studies have suggested that interactions between ants and EFN-bearing plants are susceptible to seasonality, abiotic factors and perturbation, this cohesive structure appears to be the key for biodiversity and community maintenance.     

Comparative development of the rattan ocrea,a structural innovation that facilitates ant–plant mutualism

The ocrea is an extension of the leaf sheath that occurs in a wide range of angiosperms, including some rattan palms (Arecaceae/Palmae). In some rattan species, the ocrea is an inflated sac-like structure that acts as a domatium. Typically, ants occupy this domatium, tending their young and husbanding aphids, and potentially providing a defence mechanism for the rattan. We present a comparative study of early leaf development in five palm species, including both ocreate and non-ocreate taxa. Early leaf development was examined using SEM in Calamus longipinna, which has a pronounced ocrea, and compared with two other Calamus species with an inconspicuous ocrea, and two non-ocreate palms, Chamaedorea pochutlensis and Rhapis humilis. All three Calamus species examined develop an extension from the top of the leaf sheath, in contrast with the non-ocreate palms examined for comparison. The ocrea is a vascularized outgrowth from the top of the leaf sheath, initiated shortly after differentiation of the lamina and sheath. Calamus longipinna differs from the other two Calamus species in that plications develop on the surface of the ocrea, and persist as folds on the ocrea surface. The highly unusual ocreate plications of C. longipinna form axial folds that permit the sac-like ocrea to become inflated, and give it the properties of a domatium, thus facilitating the ant–plant interaction in this species. The homologies of the ocrea in rattans require further review because the interpretation of leaf-derived appendages such as ligules and stipules remains controversial based on contradictory evidence from location and vasculature.     

Some progress in the study of plant–soil interactions in China

Plants and soil are the base for sustainably surviving human beings on the globe as the role of materials, energy, resources and environment (Shao & Chu 2008; Shao et al. 2008, 2009, 2010, 2012a,b; Liu & Shao, 2010; Ruan et al. 2010; Xu et al. 2010, 2012; Shao 2012; Huang et al. 2013). This topic has been extensively investigated for 100 years with more achievements in many sectors and practical significance in conducting high-efficient agriculture and eco-environmental construction. The plant–soil interaction is the core issue of this topic, which has been given much attention for the past 30 years (Wu et al. 2007, 2010; Zhang et al. 2011, 2013; Xu et al. 2012, 2013).     

Sugar secretion and ant protection in Ficus benguetensis: Toward a general trend of fig–ant interactions

The relationship between plants and ants is often mediated by the presence of extrafloral nectaries (EFNs) that attract ants and provide rewards by protecting plants from herbivores or parasites. Ficus trees (Moraceae) and their pollinators (Hymenoptera: Agaonidae) are parasitized by many nonpollinating fig wasp species (Hymenoptera: Chalcidoidea) that decrease the reproductive output of the mutualistic partners. Previous studies have shown that ants living on and patrolling Ficus species can efficiently deter parasitic wasps. The aim of this study was to verify the presence of EFNs on figs of Ficus benguetensis and test the hypothetical protection service provided by ants. Figs in different developmental stages were collected from Fu-Yang Eco Park, Taipei, Taiwan. Sugars on the fig surface were collected and analyzed through high-performance anion-exchange chromatography. Moreover, ants were excluded from the figs to determine the effect of ants on the nonpollinating fig wasps. We identified three oligosaccharides whose relative proportions varied with the fig developmental phase. In addition, results showed that the ant-excluded figs were heavily parasitized and produced three times less pollinators than did the control figs. Finally, the specific interactions of Ficus benguetensis with ants and the relationship between figs and ants in general are discussed.     

Comparing the conservatism of ecological interactions in plant–pollinator and plant–herbivore networks

Conservatism in species interaction, meaning that related species tend to interact with similar partners, is an important feature of ecological interactions. Studies at community scale highlight variations in conservatism strength depending on the characteristics of the ecological interaction studied. However, the heterogeneity of datasets and methods used prevent to compare results between mutualistic and antagonistic networks. Here we perform such a comparison by taking plant–insect communities as a study case, with data on plant–herbivore and plant–pollinator networks. Our analysis reveals that plants acting as resources for herbivores exhibit the strongest conservatism in species interaction among the four interacting groups. Conservatism levels are similar for insect pollinators, insect herbivores and plants as interacting partners of pollinators, although insect pollinators tend to have a slightly higher conservatism than the two others. Our results thus clearly support the current view that within antagonistic networks, conservatism is stronger for species as resources than for species as consumer. Although the pattern tends to be opposite for plant–pollinator networks, our results suggest that asymmetry in conservatism is much less pronounced between the pollinators and the plant they interact with. We discuss these differences in conservatism strength in relation with the processes structuring plant–insect communities.     

Diversity and evolution of a trait mediating ant–plant interactions: insights from extrafloral nectaries in Senna (Leguminosae)

Background and Aims

Plants display a wide range of traits that allow them to use animals for vital tasks. To attract and reward aggressive ants that protect developing leaves and flowers from consumers, many plants bear extrafloral nectaries (EFNs). EFNs are exceptionally diverse in morphology and locations on a plant. In this study the evolution of EFN diversity is explored by focusing on the legume genus Senna, in which EFNs underwent remarkable morphological diversification and occur in over 80 % of the approx. 350 species.

Methods

EFN diversity in location, morphology and plant ontogeny was characterized in wild and cultivated plants, using scanning electron microscopy and microtome sectioning. From these data EFN evolution was reconstructed in a phylogenetic framework comprising 83 Senna species.

Key Results

Two distinct kinds of EFNs exist in two unrelated clades within Senna. ‘Individualized’ EFNs (iEFNs), located on the compound leaves and sometimes at the base of pedicels, display a conspicuous, gland-like nectary structure, are highly diverse in shape and characterize the species-rich EFN clade. Previously overlooked ‘non-individualized’ EFNs (non-iEFNs) embedded within stipules, bracts, and sepals are cryptic and may represent a new synapomorphy for clade II. Leaves bear EFNs consistently throughout plant ontogeny. In one species, however, early seedlings develop iEFNs between the first pair of leaflets, but later leaves produce them at the leaf base. This ontogenetic shift reflects our inferred diversification history of iEFN location: ancestral leaves bore EFNs between the first pair of leaflets, while leaves derived from them bore EFNs either between multiple pairs of leaflets or at the leaf base.

Conclusions

EFNs are more diverse than previously thought. EFN-bearing plant parts provide different opportunities for EFN presentation (i.e. location) and individualization (i.e. morphology), with implications for EFN morphological evolution, EFN–ant protective mutualisms and the evolutionary role of EFNs in plant diversification.     

Arabidopsis thaliana as a model for the study of plant–virus co-evolution

Understanding plant–virus coevolution requires wild systems in which there is no human manipulation of either host or virus. To develop such a system, we analysed virus infection in six wild populations of Arabidopsis thaliana in Central Spain. The incidence of five virus species with different life-styles was monitored during four years, and this was analysed in relation to the demography of the host populations. Total virus incidence reached 70 per cent, which suggests a role of virus infection in the population structure and dynamics of the host, under the assumption of a host fitness cost caused by the infection. Maximum incidence occurred at early growth stages, and co-infection with different viruses was frequent, two factors often resulting in increased virulence. Experimental infections under controlled conditions with two isolates of the most prevalent viruses, cauliflower mosaic virus and cucumber mosaic virus, showed that there is genetic variation for virus accumulation, although this depended on the interaction between host and virus genotypes. Comparison of Q_ST-based genetic differentiations between both host populations with F_ST genetic differentiation based on putatively neutral markers suggests different selection dynamics for resistance against different virus species or genotypes. Together, these results are compatible with a hypothesis of plant–virus coevolution.     

Intra- and interspecific tree diversity promotes multitrophic plant–Hemiptera–ant interactions in a forest diversity experiment

Interactions between species of different trophic levels have long been recognized as fundamental processes in ecology. Although mounting evidence indicates that plant species diversity (PSD) or plant genetic diversity (PGD) can influence the plant-associated arthropod community, these two fundamental levels of biodiversity are not often manipulated simultaneously to assess their effects on species interactions. We used a large tree diversity experiment (BEF-China), which manipulates PSD and PGD in a crossed design to test individual and combined effects of PSD and PGD on multitrophic interaction networks and interaction partner species richness and occurrence. We focused on two tree species, on which sap-sucking Hemiptera and interacting ant species commonly occur. This tri-trophic interaction can be divided into the antagonistic plant–Hemiptera interaction and the mutualistic Hemiptera–ant interaction, known as trophobioses. Qualitative evaluation of tri-trophic interaction networks at different PSD and PGD combinations showed increased interaction partner redundancy at high PSD and PGD. This was supported by increased Hemiptera species richness at high PSD and PGD. Furthermore, the data indicate higher occurrence of Hemiptera and trophobioses and higher trophobiotic ant species richness with increasing PSD and PGD. As no plant diversity component alone caused an effect we conclude that the combined effect of high PGD and high PSD might be additive. In summary, as plant genetic diversity, especially at low species richness, seems to increase the interaction partner redundancy in interaction networks and the diversity of interacting communities, we suggest that genetic diversity should be considered in forest conservation and restoration programs.     

An ethological study on chimpanzees at the artificial feeding place in the Mahale mountains,Tanzania—with special reference to the booming situation

A provisioned chimpanzee group in the Mahale Mountains was observed. The characteristic feature of chimpanzee behavior is that many types of behavior occur at one time as a bout of interactions. This can be seen in the booming situation in the wild. Similar situations were observed at the artificial feeding place. The behavior of the initiators of the interactions differed according to age, sex and social status. The chimpanzees' behavior was studied as a system. The relationships between each behavior pattern were investigated by analysis of the sequential behaviors of the same individual, and that of chains of signal-response behaviors, as well as that of behaviors which appeared when a third individual joined the initial interaction. Appeasement behaviors were more flexible, while others were more stereotyped. Part of the appeasement behavior resembled begging behavior. Appeasement behavior is thought to have a rather new phylogenetic origin. It is suggested that the various chimpanzee behaviors have evolved as a harmonious system which can be observed in the booming situation. The origin of chimpanzee society based on the evolution of their behavior system is discussed. Analysis of triplet interactions indicated that the types of triplet interaction in chimpanzees are quite different from those of multiple male group species such as macaques and baboons.     

Effect of livestock grazing in the partitions of a semiarid plant–plant spatial signed network

In recent times, network theory has become a useful tool to study the structure of the interactions in ecological communities. However, typically, these approaches focus on a particular kind of interaction while neglecting other possible interactions present in the ecosystem. Here, we present an ecological network for plant communities that consider simultaneously positive and negative interactions, which were derived from the spatial association and segregation between plant species. We employed this network to study the structure and the association strategies in a semiarid plant community of Cabo de Gata-Níjar Natural Park, SE Spain, and how they changed in 4 sites that differed in stocking rate. Association strategies were obtained from the partitions of the network, built based on a relaxed structural balance criterion. We found that grazing simplified the structure of the plant community. With increasing stocking rate species with no significant associations became dominant and the number of partitions decreased in the plant community. Independently of stocking rate, many species presented an associative strategy in the plant community because they benefit from the association to certain ‘nurse’ plants. These ‘nurses’ together with species that developed a segregating strategy, intervened in most of the interactions in the community. Ecological networks that combine links with different signs provide a new insight to analyze the structure of natural communities and identify the species which play a central role in them.     

Dynamics of distribution and performance of ramets constructing genets: a demographic�Cgenetic study in a clonal plant, Convallaria keiskei

Background and Aims

In clonal plants producing vegetative offspring, performance at the genet level as well as at the ramet level should be investigated in order to understand the entire picture of the population dynamics and the life history characteristics. In this study, demography, including reproduction and survival, the growth patterns and the spatial distributions of ramets within genets of the clonal herb Convallaria keiskei were explored.

Methods

Vegetative growth, flowering and survival of shoots whose genets were identified using microsatellite markers were monitored in four study plots for 3 years (2003–2005). The size structures of ramets in genets and their temporal shifts were then analysed. Their spatial distributions were also examined.

Key Results

During the census, 274 and 149 ramets were mapped in two 1 × 2 m plots, and 83 and 94 ramets in two 2 × 2 m quadrats. Thirty-eight genotypes were identified from 580 samples. Each plot included 5–18 genets, and most ramets belonged to the predominant genet(s) in each plot. Shoots foliated yearly for several years, but flowering ramets did not have an inflorescence the next year. A considerable number of new clonal offspring persistently appeared, forming a bell-shaped curve of the size structure of ramets in each genet. Comparing the structures modelled by the normal distributions suggested variation among ramets belonging to a single genet and variation among genets. Furthermore, spatial analyses revealed clumped and distant distributions of ramet pairs in a genet, in which the distant patterns corresponded to the linearly elongating clonal growth pattern of this species.

Conclusion

Characteristics of ramet performances such as flowering and recruitment of clonal offspring, in addition to growth, played a large part in the regulation of genet dynamics and distribution, which were different among the studied genets. These might be characteristics particularly relevant to clonal life histories.Key words: Clonal plant, Convallaria keiskei, demography, genet, genetic identification, growth pattern, life history, ramet, spatial distribution     

Distribution and pollution evaluation of fluoride in a soil–water–plant system in Shihezi,Xinjiang, China

Fluoride (F) pollution is a serious environmental problem in some areas of China, but it has yet to be reported in a soil–water–plant system in Shihezi, Xinjiang. This study was undertaken to investigate the distribution and migration rule of F in soil, water, and plants, and to evaluate F pollution of soil. Results showed that the average concentration of total F (T-F) in the topsoil in the northwest, north, and southeast of Shihezi was higher than the national average T-F (478 mg/kg), while it was lower in southwest. The highest T-F contents of the soil profile were detected in the depth of 20 cm. The F content in groundwater in the northwest region was higher than the GB/T 14848–93 (1.0 mg/L), whereas the F contents in other water samples were within the standard. The F contents (1.75?2.81 mg/kg) in plant leaves were higher than the food limits (1.0 mg/kg). The obtained comprehensive pollution index of the soil was 1.86, which means a mild concentration of F in Shihezi. This research has reference value for the study of F pollution and comprehensive control in the northwest oasis with the typical arid and saline conditions.     

Ant–plant interaction in the Neotropical savanna: direct beneficial effects of extrafloral nectar on ant colony fitness

Current evidence suggests that ant–plant relationships may influence species composition, abundance, and interactions at the community scale. The main resource that plants offer to ants is extrafloral nectar (EFN) and the major part of published studies shown benefits from ants to plants possessing EFNs. However, the complementary question of whether and how ants benefit from EFNs is rarely addressed. Here, we present the results of a long-term study to demonstrate whether EFN has a positive effect on ant colony fitness. We quantified colony growth rate, survival and the final weight of individuals as measures of benefit derived from EFN. Our results provide clear evidence that EFN can have a significant positive impact on the survivorship, growth and reproduction of the Myrmicinae Cephalotes pusillus. In fact, a diet rich in EFN (providing at least 30 cal per day) resulted in five times more individuals per colony, greater body weights, and more eggs. These results have shed new light on the relationships between ants and EFN-bearing plants such as in tropical and temperate systems. The ant C. pusillus is the first case in which we have firm evidence that EFN improves colony growth and development, corroborating more than 100 years of experimental evidence of benefits to plants in these widespread relationships.     

From species to individuals: does the variation in ant–plant networks scale result in structural and functional changes?

Predicting the outcomes of any mutualistic interaction between ants and plants can be a very difficult task, since these outcomes are often determined by the ecological context in which the interacting species are embedded. Network theory has been an important tool to improve our understanding about the organizational patterns of animal–plant interactions. Nevertheless, traditionally, network studies have focused mainly on species-based differences and ignoring the importance of individual differences within populations. In this study, we evaluated if downscaling an ant–plant network from species to the individual level results in structural and functional changes in a network involving different-sized plant individuals. For this, we studied the extrafloral-nectar producing-tree Caryocar brasiliense (Caryocaraceae) and their associated ants in a Neotropical savanna. We observed 254 interactions involving 43 individuals of C. brasiliense and 47 ant species. The individual-based ant–plant network exhibited a nested pattern of interactions, with all developmental stages contributing equally to structuring this non-random pattern. We also found that plants with greater centrality within the network were better protected by their ant partners. However, plants with higher levels of individual specialization were not necessarily better protected by ants. Overall, we presented empirical evidence that intra-population variations are important for shaping ant–plant networks, since they can change the level of protection against herbivores conferred by the ants. These results highlight the importance of individual-based analyses of ecological networks, opening new research venues in the eco-evolutionary dynamics of ant–plant interactions.     

Monocropping decouples plant–bacteria interaction and strengthens phytopathogenic fungi colonization in the rhizosphere of a perennial plant species

Plant and Soil - Whether Arbuscular mycorrhizal fungi (AMF) influence community composition by changing plant adaptation to resource limitation remains unclear. This study examined how AMF affect...