Soil and vegetation features determine the nested pattern of ant–plant networks in a tropical rainforest

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Availability of food resources and habitat structure shape the individual‐resource network of a Neotropical marsupial

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The role of epiphytism in architecture and evolutionary constraint within mycorrhizal networks of tropical orchids

Characterizing the architecture of bipartite networks is increasingly used as a framework to study biotic interactions within their ecological context and to assess the extent to which evolutionary constraint shape them. Orchid mycorrhizal symbioses are particularly interesting as they are viewed as more beneficial for plants than for fungi, a situation expected to result in an asymmetry of biological constraint. This study addressed the architecture and phylogenetic constraint in these associations in tropical context. We identified a bipartite network including 73 orchid species and 95 taxonomic units of mycorrhizal fungi across the natural habitats of Reunion Island. Unlike some recent evidence for nestedness in mycorrhizal symbioses, we found a highly modular architecture that largely reflected an ecological barrier between epiphytic and terrestrial subnetworks. By testing for phylogenetic signal, the overall signal was stronger for both partners in the epiphytic subnetwork. Moreover, in the subnetwork of epiphytic angraecoid orchids, the signal in orchid phylogeny was stronger than the signal in fungal phylogeny. Epiphytic associations are therefore more conservative and may co‐evolve more than terrestrial ones. We suggest that such tighter phylogenetic specialization may have been driven by stressful life conditions in the epiphytic niches. In addition to paralleling recent insights into mycorrhizal networks, this study furthermore provides support for epiphytism as a major factor affecting ecological assemblage and evolutionary constraint in tropical mycorrhizal symbioses.     

Ant dominance hierarchy determines the nested pattern in ant–plant networks

Extrafloral nectar (EFN) is a predictable and renewable resource for many ant colonies, and different ant species compete strongly to obtain and monopolize this highly nutritious food resource. Despite the importance of competition in structuring patterns of ant–plant interactions, this biological mechanism has been largely ignored in studies involving ant–plant networks. In this study we investigate the role of ant dominance hierarchy in structuring an ecological network involving ants and EFN‐bearing plants in a tropical coastal environment in Mexico. We show that within a nested ant–plant network, ant species found in the central core of highly interacting species were competitively superior, showing massive recruitment and resource domination, compared with peripheral species with fewer interactions. Moreover, we also observed that both central and peripheral ant species have the ability to quickly find the food resource. However, after 2 h of observation, central ant species are more frequently collected on the food resource when compared with peripheral species. We hypothesize that the existence of a central core of competitive ant species may indicate that most plant species found within ant–plant networks could be better protected against herbivory by these dominant ant species. In short, our results highlight the importance of competition and monopolization in the resource use by ants in the maintenance of the nested pattern in ant–plant mutualistic networks. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 405–414.     

The topological differences between visitation and pollen transport networks: a comparison in species rich communities of the Himalaya–Hengduan Mountains

Pollination networks are usually constructed and assessed by direct field observations which commonly assume that all flower visitors are true pollinators. However, this assumption is often invalid and the use of data based on mere visitors to flowers may lead to a misunderstanding of intrinsic pollination networks. Here, using a large dataset by both sampling floral visitors and analyzing their pollen loads, we constructed 32 networks pairs (visitation versus pollen transport) across one flowering season at four elevation sites in the Himalaya–Hengduan Mountains region. Pollen analysis was conducted to determine which flower visitors acted as potential pollinators (pollen vectors) or as cheaters (those not carrying pollen of the visited plants). We tested whether there were topological differences between visitation and pollen transport networks and whether different taxonomic groups of insect visitors differed in their ability to carry pollen of the visited plants. Our results indicated that there was a significantly higher degree of specialization at both the network and species levels in the pollen transport networks in contrast to the visitation networks. Modularity was lower but nestedness was higher in the visitation networks compared to the pollen transport networks. All the cheaters were identified as peripheral species and most of them contributed positively to the nested structure. This may explain in part the differences in modularity and nestedness between the two network types. Bees carried the highest proportion of pollen of the visited plants. This was followed by Coleoptera, other Hymenoptera and Diptera. Lepidoptera carried the lowest proportion of pollen of the visited plants. Our study shows that the construction of pollen transport networks could provide a more in‐depth understanding of plant–pollinator interactions. Moreover, it suggests that detecting and removing cheater interactions when studying the topology of other mutualistic networks might be also important.     

Relationship between soil nutrient availability and plant species richness in a tropical semi‐arid environment

Question: What is the relationship between soil fertility and plant species richness in the ‘fertile islands’ occurring beneath two species of legume (Cercidium praecox and Prosopis laevigata)? Location: Tehuacán‐Cuicatlán region, central Mexico. Methods: Plant richness was measured in three micro‐environments (below canopies of C. praecox, below canopies of P. laevigata and in areas without canopies). The concentration of soil nutrients (C, N and P), C and N in the microbiota, and processes of ecosystem functioning (net C mineralization rate and N mineralization) were measured. The relationship between soil variables and plant richness were assessed with ANCOVAs. Results: Soil nutrients and species richness increases markedly under fertility islands. There were higher concentrations of C and N in the soil, faster rates of C mineralization, and higher species richness under P. laevigata canopies. The relationship between soil fertility and species richness was always positive except for total N, ammonium and net C mineralization rate under C. praecox, and for available P under P. laevigata. Conclusions: The sign of the relationship between soil fertility and species richness varies according to the nutrient and the micro‐environment. Positive relationships could result from between species complementarity and facilitation. Negative relationships could be explained by a specific limitation threshold for some soil resources (P and N for plants and C for the soil microbiota) which eliminate the possibilities of between species complementarity and facilitation above that threshold. As in all observational studies, these relationships should be considered only correlational.     

The impact of life form on the architecture of orchid mycorrhizal networks in tropical forest

Understanding the processes that determine the architecture of interaction networks represents a major challenge in ecology and evolutionary biology. One of the most important interactions involving plants is the interaction between plants and mycorrhizal fungi. While there is a mounting body of research that has studied the architecture of plant–fungus interaction networks, less is known about the potential factors that drive network architecture. In this study, we described the architecture of the network of interactions between mycorrhizal fungi and 44 orchid species that represented different life forms and co‐occurred in tropical forest and assessed the relative importance of ecological, evolutionary and co‐evolutionary mechanisms determining network architecture. We found 87 different fungal operational taxonomic units (OTUs), most of which were members of the Tulasnellaceae. Most orchid species associated with multiple fungi simultaneously, indicating that extreme host selectivity was rare. However, an increasing specificity towards Tulasnellaceae fungal associates from terrestrial to epiphytic and lithophytic orchids was observed. The network of interactions showed an association pattern that was significantly modular (M = 0.7389, M_random = 0.6998) and nested (NODF = 5.53, p < 0.05). Terrestrial orchids had almost no links to modules containing epiphytic or lithophytic orchids, while modules containing epiphytic orchids also contained lithophytic orchids. Within each life form several modules were observed, suggesting that the processes that organize orchid–fungus interactions are independent of life form. The overall phylogenetic signal for both partners in the interaction network was very weak. Overall, these results indicate that tropical orchids associate with a wide number of mycorrhizal fungi and that ecological rather than phylogenetic constraints determine network architecture.     

Ecological,historical and evolutionary determinants of modularity in weighted seed‐dispersal networks

Modularity is a recurrent and important property of bipartite ecological networks. Although well‐resolved ecological networks describe interaction frequencies between species pairs, modularity of bipartite networks has been analysed only on the basis of binary presence–absence data. We employ a new algorithm to detect modularity in weighted bipartite networks in a global analysis of avian seed‐dispersal networks. We define roles of species, such as connector values, for weighted and binary networks and associate them with avian species traits and phylogeny. The weighted, but not binary, analysis identified a positive relationship between climatic seasonality and modularity, whereas past climate stability and phylogenetic signal were only weakly related to modularity. Connector values were associated with foraging behaviour and were phylogenetically conserved. The weighted modularity analysis demonstrates the dominating impact of ecological factors on the structure of seed‐dispersal networks, but also underscores the relevance of evolutionary history in shaping species roles in ecological communities.     

Edge disturbance shapes liana diversity and abundance but not liana‐tree interaction network patterns in moist semi‐deciduous forests,Ghana

Edge disturbance can drive liana community changes and alter liana‐tree interaction networks, with ramifications for forest functioning. Understanding edge effects on liana community structure and liana‐tree interactions is therefore essential for forest management and conservation. We evaluated the response patterns of liana community structure and liana‐tree interaction structure to forest edge in two moist semi‐deciduous forests in Ghana (Asenanyo and Suhuma Forest Reserves: AFR and SFR, respectively). Liana community structure and liana‐tree interactions were assessed in 24 50 × 50 m randomly located plots in three forest sites (edge, interior and deep‐interior) established at 0–50 m, 200 m and 400 m from edge. Edge effects positively and negatively influenced liana diversity in forest edges of AFR and SFR, respectively. There was a positive influence of edge disturbance on liana abundance in both forests. We observed anti‐nested structure in all the liana‐tree networks in AFR, while no nestedness was observed in the networks in SFR. The networks in both forests were less connected, and thus more modular and specialised than their null models. Many liana and tree species were specialised, with specialisation tending to be symmetrical. The plant species played different roles in relation to modularity. Most of the species acted as peripherals (specialists), with only a few species having structural importance to the networks. The latter species group consisted of connectors (generalists) and hubs (highly connected generalists). Some of the species showed consistency in their roles across the sites, while the roles of other species changed. Generally, liana species co‐occurred randomly on tree species in all the forest sites, except edge site in AFR where lianas showed positive co‐occurrence. Our findings deepen our understanding of the response of liana communities and liana‐tree interactions to forest edge disturbance, which are useful for managing forest edge.     

Floodplain ecosystem dynamics under extreme dry and wet phases in semi‐arid Australia

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Vegetation connectivity increases ant activity and potential for ant‐provided biocontrol services in a tropical agroforest

In natural and managed systems, connections between trees are important structural resources for arboreal ant communities with ecosystem‐level effects. However, ongoing agricultural intensification in agroforestry systems, which reduces shade trees and connectivity between trees and crop plants, may hinder ant recruitment rates to resources and pest control services provided by ants. We examined whether increasing connectivity between coffee plants and shade trees in coffee plantations increases ant activity and enhances biological control of the coffee berry borer, the most devastating insect pest of coffee. Further, we examined whether artificial connections buffer against the loss of vegetation connectivity in coffee plants located at larger distances from the nesting tree. We used string to connect Inga micheliana shade trees containing Azteca sericeasur ant nests to coffee plants to compare ant activity before and after placement of the strings, and measured borer removal by ants on coffee plants with and without strings. Ant activity significantly increased after the addition of strings on connected plants, but not on control plants. Borer removal by ants was also three times higher on connected plants after string placement. Greater distance from the nesting tree negatively influenced ant activity on control coffee plants, but not on connected plants, suggesting that connections between coffee plants and nest trees could potentially compensate for the negative effects that larger distances pose on ant activity. Our study shows that favoring connectivity at the local scale, by artificially adding connections, promotes ant activity and may increase pest removal in agroecosystems. Abstract in Spanish is available with online material.     

Movement patterns of catfishes (Ariidae) in a tropical semi‐arid estuary

From December 2005 to November 2006, 216 samples were taken from the main channel of the Goiana Estuary, representing a total sampled area of 23 ha. Ariidae species were the most abundant in density (1600 individuals ha^?1, 53%) and biomass (18 813 g ha^?1, 63%). Cathorops spixii was the most abundant in density (1340 individuals ha^?1) and biomass (14 203 g ha^?1). The variables: number of species, total density and biomass, showed significant interactions between the factors of areas and seasons (P < 0·01). The highest total density (7394 individuals ha^?1) and biomass (70 292 g ha^?1) occurred in the middle and upper estuaries, respectively, during the early‐rainy season. The density of C. spixii differed significantly between areas and seasons (P < 0·01), while Cathorops agassizii differed significantly only between seasons (P < 0·01). The biomass differed significantly for the species C. spixii, C. agassizii and Sciades herzbergii between seasons, and C. spixii and C. agassizii between areas. Also, the density and biomass of C. spixii and C. agassizii showed interaction between areas and seasons. Only the biomass of S. herzbergii showed interaction between areas and seasons (P < 0·01). This indicates that seasonal variations influenced the distribution of Ariidae species in the different areas of the Goiana Estuary. Moreover, canonical correspondence analysis highlighted a strongly significant correlation (P < 0·01) between the seasonal variations of the environmental gradients (salinity and water temperature) and distribution of catfishes. The management and conservation of estuaries should take into account the life cycle of these species in different estuarine areas and hydrological seasons.     

Seasonal dynamic variation of pollination network is associated with the number of species in flower in an oceanic island community

植物与传粉者相互作用的传粉网络是一个动态的实体,植物开花物候的季节性变化可以在短时间和长时间内重塑其结构。然而,很少有研究考虑到这种季节性动态变化,特别是海洋岛屿群落的传粉网络。本研究探讨了海洋岛屿群落的植物与传粉者间传粉网络的结构是如何随群落内花资源丰富度的季节性变化而动态变化的。利用春夏秋冬四个季节的植物与传粉者间相互作用的数据,分析了四个季节定性的传粉网络结构的动态变化,研究了中国南海西沙群岛的永兴岛群落传粉网络的季节性动态变化。在这四个季节中,分别收集了连续两个月的植物与传粉者相互作用的数据,并计算了四个网络水平的指标来表征传粉网络的总体结构。采用群落差异性统计分析方法,对群落四个季节的网络结构参数进行比较分析,探讨影响这种动态变化格局的潜在因素。同时计算并比较了植物和传粉功能群在物种水平的网络指标的季节动态变化。研究结果表明,永兴岛群落网络水平的特化性和模块化在四个季节的变化均与植物物种丰富度的变化呈明显相反的变化趋势。开花植物种类的增加可能促进了传粉者之间更激烈的竞争,从而导致生态位重叠的增加,引起传粉网络特化性和模块化的下降,反之亦然。进一步分析表明,传粉网络的季节动态变化的内在驱动力是植物与传粉者间连接的重新组合。因此,传粉者之间生态位重叠的季节性变化导致了植物与传粉者间相互作用的重组,从而推动了该群落内植物与传粉者间相互作用的更替变化。在物种水平上,与其它传粉功能群相比,天蛾类传粉者最特化,而蜜蜂科传粉者的物种作用强度最大。因此,在探索孤立的海洋岛屿生态系统以及其它生态系统中的植物与传粉者间相互作用时,应适当考虑到这些新的发现。     

Habitat selection by feral cats and dingoes in a semi‐arid woodland environment in central Australia

Habitat use by feral cats and dingoes was examined within a heterogeneous semi‐arid woodland site in central Australia over 2 years. Density estimates of feral cats based on tracks were higher in mulga habitat than in open habitat. Isodar analysis implied that this pattern of habitat use by feral cats was consistent with the consumer‐resource model of density‐dependent habitat selection, which is an ideal free solution. The reason why mulga supported higher densities of feral cats was unclear. Foraging success of feral cats may be higher in the mulga because the stalk and ambush hunting tactics typically employed by felids are well suited to dense cover. Mulga may also have offered feral cats more protection from dingo predation. Dingo activity was distributed uniformly across habitats. The dingo isodar was statistically non‐significant, suggesting that habitat selection by dingoes was independent of density.     

Ant community structure under Retama sphaerocarpa shrubs in a semi‐arid environment

The Tabernas Desert (Spain) is an emblematic area as it constitutes, strictly speaking, the only European desert. Under the canopy of one of the dominant shrub species, Retama sphaerocarpa, there are different microhabitat conditions that encourage the growing of a dense understory of herbs (fertile islands). However, these changes may not be enough to be also reflected in the fauna. Studying ants as bioindicators, we found higher densities of individuals and species richness below the canopy than outside it. However, while the density of individuals increased from outside the canopy to the inner position under the shrubs, species richness increased from the inner to the border position under the canopy and abruptly decreased outside it. This is because the inner positions were usually monopolized by dominant ants with populous nests that feed on aphid honeydew, such as Monomorium subopacum or Tapinoma nigerrimum. Granivorous species (genus Messor) were usually found in intermediate positions under the canopy, taking advantage of the better microclimatic conditions but close to the grass for feeding. Among the most abundant species, the only one found mainly outside the influence of the retama shrubs was the thermophilic insectivorous Cataglyphis ibericus.     

Influence of extrafloral nectary phenology on ant–plant mutualistic networks in a neotropical savanna

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The global biogeography of semi‐arid periodic vegetation patterns

Aim Vegetation exhibiting landscape‐scale regular spatial patterns has been reported for arid and semi‐arid areas world‐wide. Recent theories state that such structures are bound to low‐productivity environments and result from a self‐organization process. Our objective was to test this relationship between periodic pattern occurrence and environmental factors at a global scale and to parametrize a predictive distribution model. Location Arid and semi‐arid areas world‐wide. Methods We trained an empirical predictive model (Maxent) for the occurrence of periodic vegetation patterns, based on environmental predictors and known occurrences verified on Landsat satellite images. Results This model allowed us to discover previously unreported pattern locations, and to report the first ever examples of spotted patterns in natural systems. Relationships to the main environmental drivers are discussed. Main conclusions These results confirm that periodic patterned vegetations are ubiquitous at the interface between arid and semi‐arid regions. Self‐organized patterning appears therefore to be a biome‐scale response to environmental conditions, including soil and topography. The set of correlations between vegetation patterns and their environmental conditions presented in this study will need to be reproduced in future modelling attempts.     

Scale‐dependent determinants of plant species richness in a semi‐arid fragmented agro‐ecosystem

Aims: (1) Understanding how the relationship between species richness and its determinants depends on the interaction between scales at which the response and explanatory variables are measured. (2) Quantifying the relative contributions of local, intermediate and large‐scale determinants of species richness in a fragmented agro‐ecosystem. (3) Testing the hypothesis that the relative contribution of these determinants varies with the grain size at which species richness is measured. Location: A fragmented agro‐ecosystem in the Southern Judea Lowland, Israel, within a desert–Mediterranean transition zone. Methods: Plant species richness was estimated using hierarchical nested sampling in 81 plots, positioned in 38 natural vegetation patches within an agricultural matrix (mainly wheat fields) among three land units along a sharp precipitation gradient. Explanatory variables included position along that gradient, patch area, patch isolation, habitat heterogeneity and overall plant density. We used general linear models and hierarchical partitioning of variance to test and quantify the effect of each explanatory variable on species richness at four grain sizes (0.0625, 1, 25 and 225 m²). Results: Species richness was mainly affected by position along a precipitation gradient and overall plant density, and to a lesser extent by habitat heterogeneity. It was also significantly affected by patch area and patch isolation, but only for small grain sizes. The contribution of each explanatory variable to explained variance in species richness varied with grain size, i.e. scale‐dependent. The influence of geographic position and habitat heterogeneity on species richness increased with grain size, while the influence of plant density decreased with grain size. Main conclusions: Species richness is determined by the combined effect of several scale‐dependent determinants. Ability to detect an effect and effect size of each determinant varies with the scale (grain size) at which it is measured. The combination of a multi‐factorial approach and multi‐scale sampling reveals that conclusions drawn from studies that ignore these dimensions are restricted and potentially misleading.     

Plant community resilience in the face of fire: experimental evidence from a semi‐arid shrubland

The ability of communities or ecosystems to recover their structure and function after a disturbance is known as resilience. According to different views, resilience can be influenced by the resource‐use strategies of the plant functional types that dominate the community or by the existence of functional redundancy within plant functional types. We investigated how the dominance of different plant functional types and species affected the resilience of a mountain shrubland after an intense fire. We took advantage from a pre‐existing long‐term removal experiment in which either whole plant functional types (deciduous shrubs, graminoids, perennial forbs and annual forbs) or the dominant species within each plant functional type were removed for 10 years. We sampled species and plant functional types cover during the first growing season after the fire. First, to test whether functional redundancy increased resilience, we analyzed the existence of functional compensation inside plant functional types. Second, to test whether the dominance of plant functional types with different resource‐use strategies affected recovery, we compared resilience at the levels of species, plant functional types and total cover, estimated on the basis of a change index and multivariate Euclidean distances. No compensation was observed in any of the plant functional types. At the level of species, we found that the assemblages dominated by conservative resource‐use strategies were the ones showing higher resilience. This was due to the high recovery of the dominant species of shrubs plant functional type. The opposite (lowest recovery of conservative resource‐use strategies) was found at the plant functional type and total cover‐levels. Our study did not support the hypothesis of resilience by functional redundancy. Instead, regeneration by buried meristems from the pre‐fire stage appeared to be the factor that most influenced recovery. Resource‐use strategies explained resilience of vegetation cover, but not of floristic composition. Regeneration traits, rather than vegetative traits or mechanism of functional compensation, appeared as the most relevant to explain the response of this system after fire.