Interaction rewiring and the rapid turnover of plant–pollinator networks

Whether species interactions are static or change over time has wide‐reaching ecological and evolutionary consequences. However, species interaction networks are typically constructed from temporally aggregated interaction data, thereby implicitly assuming that interactions are fixed. This approach has advanced our understanding of communities, but it obscures the timescale at which interactions form (or dissolve) and the drivers and consequences of such dynamics. We address this knowledge gap by quantifying the within‐season turnover of plant–pollinator interactions from weekly censuses across 3 years in a subalpine ecosystem. Week‐to‐week turnover of interactions (1) was high, (2) followed a consistent seasonal progression in all years of study and (3) was dominated by interaction rewiring (the reassembly of interactions among species). Simulation models revealed that species’ phenologies and relative abundances constrained both total interaction turnover and rewiring. Our findings reveal the diversity of species interactions that may be missed when the temporal dynamics of networks are ignored.     

A year‐long plant‐pollinator network

Abstract In this work we analyse the pollination community in a South American forest known as ‘talar’. This is a vegetal woody community that inhabits fossil coastal banks characterized by seasonal temperate weather and calcareous soil, at the coast of the Río de la Plata, in the province of Buenos Aires, Argentina. We obtained data of the interactions between anthophylous insects and entomophylous flowering plants over an extensive period of time. We showed that pollination system parameters, such as partners’ identity, system size, and connectance, fluctuated among months, when sampled year‐long. Maximal network size occurred in early spring and early autumn, when both the number of mutualistic species and the number of interactions peaked, and this was also when network asymmetry was higher than average. Monthly connectance of the plant‐flower visitor matrix decreased to its lowest values at these peaks. Available data suggest that cumulative traditional connectance (i.e. the connectance calculated as the whole number of interactions registered in the community divided by the full size system) underestimates actual connectance values by a factor of c. 3 ×. Monthly values of connectance decreased exponentially as system size increased, and the distribution of interactions per species followed power‐law regimes for animals, and truncated power‐law regimes for plants, in accordance with patterns previously deduced from among‐network cumulative communities studies. We think that either within or and among pollination networks, systems that are organized as power‐law regimes may be a basic property of these webs, and provide examples of the fact. Both seasonal changes and interactions between mutualists like competition, and some degree of facilitation, may be very important to understand the performance of the system as a whole, and the role and importance of different species in the community. We suggest that communities of plant – pollinators that exhibit extended activity, such as temperate or tropical seasonal ones, should be studied through consecutive plant‐pollinator webs rather than cumulative ones. The partition of the system into smaller serial parts allows us to obtain outstanding information of every short period. This information is flattened by the average effect when we considered the combined analysis of the whole data.     

Inferring coevolution in a plant–pollinator network

Mutualistic interactions are at the core of community dynamics, determining dispersal, colonization and differential survival and reproduction among individuals and species. Mutualistic interactions therefore affect the fitness of interaction partners, hence modifying their respective evolutionary trajectories reciprocally, potentially leading to coevolution. Although mathematical models predict coevolution in mutualistic interaction networks, no empirical data are available. By taking into account the patterns of interactions and reconstructing evolutionary change in plant and pollinator traits, we tested the hypothesis that coevolution occurs between plants and insects that interact more frequently, or more symmetrically. To test this hypothesis, we built an interaction network with data from five flowering seasons, measured plant and insect morphology, mapped morphology on the plant and insect phylogenies, and reconstructed ancestral character changes based on maximum parsimony. We calculated an index, called the coevolutionary ratio, which represents the amount of correlated change in traits that mediate the interaction between plants and pollinators (i.e. proboscis versus corolla length, and body width and corolla aperture). Our results suggest that high frequency of interaction, i.e. the number of times two species interact, does not lead to coevolution. Instead, symmetry of interaction strength, i.e. the reciprocal similarity in the mutual effect of interaction partners, may lead to coevolution, in spite of a pervasive lack of reciprocal specialization and high interaction frequency. Although the statistical signal is quite weak, our results hold for three statistical tests of very different nature. The most specialized species, expected to be under directional selection, do not show more evolutionary change than do generalist species, expected to be under different, perhaps opposing, selective pressures. By dissecting the complexity of an interaction network we show that coevolution may partially shape functional morphology of interaction partners, thus providing the closest evidence to date of mutualistic adaptation of organisms within a community.     

Vegetation changes along a precipitation gradient in Central Argentina

Changes in vegetation along a precipitation gradient in Central Argentina were studied. Floristic samples were taken along an east-west transect of about 300 km. Correlation analysis between precipitation and ordination axes was used to provide an environmental interpretation of vegetation variability.Floristic analysis produced an ordination of plant communities from evergreen forests (precipitation >500 mm) to desert shrublands and therophyte communities (precipitation <200 mm). Results showed a trend of floristic and structural impoverishment towards the west. There is a replacement of species along the transect and a shift in dominant growth forms. The first ordination axis is significantly, negatively correlated with annual precipitation.     

Changing ecological communities along an elevation gradient in seasonally dry tropical forest on Hispaniola (Sierra Martín García,Dominican Republic)

We report the results of systematic vascular plant and bird surveys in Seasonally Dry Tropical Forest (SDTF) across leeward and windward elevation gradients (31–884 masl) in the Sierra Martín García (SMG), Dominican Republic. We expected to see gradual, systematic changes in plant distributions with elevation owing to the strong effect of topoclimate. In contrast, we predicted bird community composition to be related only weakly to elevation, because we expected bird distributions to be more strongly related to vegetation structure than composition. Based on 48 vegetation transects, we identified seven groups that differed in their species composition, which was correlated with elevation and precipitation. The most distinctive vegetation community occurs in dry, warm, and low elevations on the leeward slope, featuring large numbers of non‐woody indicator species (those species found frequently within one group but not in other groups) even though most of its trees and shrubs represent species that are widespread. Low rainfall and shale bedrock (rather than limestone) may be the primary drivers of distinctiveness in the low‐elevation leeward plots. On the leeward slope, where we also surveyed the birds, the vegetation community changes gradually with elevation at mid‐ to high elevations. The most distinctive bird community also was associated with the low‐elevation forest on shale bedrock and was dominated by widespread species. At higher elevations, but still within leeward SDTF, the bird communities had a stronger component of species endemic either to Hispaniola or to Caribbean islands, and species turnover did not correspond to the elevation gradient. Abstract in Spanish is available with online material.     

Effects of climate on the radial growth of mixed stands of Nothofagus nervosa and Nothofagus obliqua along a precipitation gradient in Patagonia,Argentina

Climate models for North Patagonia in Argentina project dryer conditions, due to a decrease in mean precipitation combined with an increase in mean temperatures. The temperate mixed Nothofagus forest, which exists along a steep precipitation gradient, could be directly impacted. For this study, we evaluated the influence of mean climate on the growth of the significant deciduous species: N. nervosa and N. obliqua. For the first time in Argentina, dendroclimatological analyses were done on both species using a network of 14 chronologies, covering their longitudinal distribution along a gradient of declining precipitation from west to east. Seasonal correlation analysis revealed that temperature has a negative effect on the growth of both species across all sites, particularly during summer of the previous and current growth season. Precipitation has a positive effect on the growth of trees for both species, which is more significant for N. nervosa. The relationship between early-summer climate and growth remained relatively stable over time for N. nervosa; however, for N. obliqua the detrimental effects of temperature increased towards the end of the 20th century, and the positive effects of precipitation decreased, particularly at the driest end of the gradient. These results suggest that a continued decrease in rainfall with a rise in temperature could impact growth for both of these species.     

Land degradation affects shrub growth responses to precipitation in a semiarid rangeland of north‐eastern Patagonia (Argentina)

Arid land degradation diminishes the proportion of precipitation conducted to infiltration and increases the proportion lost to run‐off and evaporation. Consequently, we expect that the effects of annual precipitation on shrub growth vary with land degradation as a result of changes in soil available water. Chuquiraga avellanedae is the dominant shrub and the main indicator of land degradation in semiarid rangelands of north‐eastern Patagonia. We chose two communities with a different degree of land degradation: an herbaceous steppe with shrubs (HSS) and a degraded shrub steppe (SS). Vegetative growth of C. avellanedae was determined nondestructively using a double‐sampling approach. Soil water content was estimated for the two communities using a soil water balance model. Linear regressions were used to evaluate the relationships between shrub growth and (i) annual precipitation and (ii) mean available water during the period of high vegetative growth in the soil layer that each plant community concentrates their roots. In SS, with elevated clay content, there were more roots of C. avellanedae in the upper layers of soil while in HSS, with coarse‐textured soil, C. avellanedae had more roots in deeper layers. Vegetative growth of C. avellanedae, both in HSS and SS communities, was positively related to annual precipitation but, for a given precipitation, C. avellanedae presented higher vegetative growth in HSS than in SS. We also found a positive relationship between vegetative growth and soil available water, and this relationship did not differ between communities. SS presented lower water availability because of lower infiltration rates. Our results showed that, irrespective of the degree of land degradation, plants respond directly to water content of the soil layers where most roots are present at a specific window of time.     

Geographical patterns of species turnover in aquatic plant communities

1. A classic theory in biogeography predicts that high latitude communities are unstable. This may be because of decreased species richness or decreased environmental predictability and productivity towards the poles.
2. We studied latitudinal patterns in long-term community persistence of aquatic vascular plants in 112 Finnish lakes, situated within a 1000-km range from the northernmost to the southernmost lake.
3. Contrary to theoretical predictions, we found that the turnover rate of plant species in 45 years was inversely related to latitude. That is, plant communities in northern lakes were more persistent than communities in southern lakes. When we used multiple regression to find the best predictors of species turnover rate (TR), latitude was the only variable that was highly significantly related to species turnover rate. Area, species number, water transparency, pH and change in transparency did not notably explain the gradient observed.
4. The latitudinal trend was mainly because of lower species immigration rates at higher latitudes, whereas extinction rate did not so strongly decrease with increasing latitude. Immigrations and extinctions in the lakes were not in balance: the species numbers between the 1930s and 1980s increased more strongly in the southern than northern lakes.
5. We suggest that the inverse relationship between latitude and plant species TR in Finland is most probably caused by human influence on lakes, especially eutrophication and immigration of new species in southern latitudes. In addition, although species richness per lake did not decrease towards the north, the total species pool probably does, which means that in the north there are fewer species that can actually immigrate.     

Pine afforestation alters rhizosphere effects and soil nutrient turnover across a precipitation gradient in Patagonia,Argentina

Plant and Soil - Pine species, when planted outside their native range, can have profound impacts on soil carbon (C) and nitrogen (N) pools, which may be related in part to their association with...     

Euglenoid morphospecies replacement along a hydraulic gradient of the Lower Paraná Basin (Argentina)

1. This study deals with the distribution of euglenoid morphospecies along the length of the Lower Paraná River floodplain. Six sites located in the Paraná River, Estevez stream and Montiel shallow lake were surveyed monthly from May 1995 to April 1996. 2. Eighty‐eight infrageneric taxa were registered though the total density percentage of euglenoids seldom exceeded 15% of total phytoplankton. The increasing species richness towards the lake is mostly the result of a high contribution of species of Trachelomonas and naked infrageneric taxa of Euglena, Lepocinclis and Phacus, while the distribution of Strombomonas species along the transect is fairly constant. 3. The richness of euglenoid morphospecies follows a pattern opposite to that found for the entire phytoplankton community. The distribution of euglenoids differs among the three habitats defined in this study: river, stream, shallow lake. These differences depend on the hydraulic condition of each habitat, as revealed by canonical correspondence analysis. 4. The degree of association with each type of aquatic habitat varied among the species and differed between water phases. Strombomonas fluviatilis was equally constant along the length of the ecotone, while S. girardiana and S. scabra were particularly associated with the Paraná River and P. longicauda, L. truncata, T. hispida and T. similis with the lake. 5. The flood pulse establishes a clear pattern both in species richness and abundance of euglenoids. The number of species recorded during high water was more than twice that recorded during low water.     

Fish communities along an oxygen-poor salinity gradient (Zeeschelde Estuary, Belgium)

Fish living in the tidal fresh and brackish water reaches of the Zeeschelde Estuary were studied in samples collected from the cooling-water inlets of three power stations. Between July 1994 and June 1995, 42 different species were recorded including 26 marine migrants, 14 freshwater species and two diadromous species. Species number as well as fish abundance were correlated significantly with salinity and oxygen concentration. The community structure was analysed with a correlation biplot based on principal component analysis of the root-root transformed fish abundance. Four separated communities could be distinguished since the first principal component expressed a salinity and the second a temperature gradient. During summer and fall Pomatoschistus microps, P. minutus, P. lozanoi and Syngnathus rostellatus were most abundant in the brackish-water reach. At this time, freshwater species seemed to avoid the freshwater area. During winter, Sprattus sprattus, Clupea harengus and Dicentrarchus labrax were the dominant species of the brackish-water zone while the freshwater reaches were dominated by Gasterosteus aculeatus . Migrating fish such as Pleuronectes flesus, Lampetra fluviatilis, Anguilla anguilla and Osmerus eperlanus were restricted to the brackish environment.     

Effects of anthropogenic habitat disturbance on local pollinator diversity and species turnover across a precipitation gradient

Anthropogenic habitat disturbance can have profound effects on multiple components of forest biotas including pollinator assemblages. We assessed the effect of small-scale disturbance on local richness, abundance, diversity and evenness of insect pollinator fauna; and how habitat disturbance affected species turnover across the landscape and overall diversity along a precipitation gradient in NW Patagonia (Argentina). We evaluated the effect of disturbance on overall pollinator fauna and then separately for bees (i.e. Apoidea) and non-bee pollinators. Locally, disturbed habitats had significantly higher pollinator species richness and abundances than undisturbed habitats for the whole pollinator assemblage, but not for bees or non-bees separately. However, significant differences in species richness between habitats vanished after accounting for differences in abundance between habitat types. At a local scale Shannon–Weaver diversity and evenness did not vary with disturbance. A β diversity index indicated that, across forest types, species turnover was lower between disturbed habitats than between undisturbed habitats. In addition, rarefaction curves showed that disturbed habitats as a whole accumulated fewer species than undisturbed habitats at equivalent sample sizes. We concluded that small patches of disturbed habitat have a negligible effect on local pollinator diversity; however, habitat disturbance reduced β diversity through a homogenization of the pollinator fauna (in particular of bees) across the landscape.     

Carbon and nitrogen dynamics across a natural precipitation gradient in Patagonia,Argentina

Abstract. Both ecosystem carbon gain and nutrient availability are largely constrained by the magnitude and seasonality of precipitation in arid and semi‐arid ecosystems. We investigated the role of precipitation on ecosystem processes along an International Geosphere Biosphere Programme (IGBP) transect in temperate South America. The transect consists of a contiguous precipitation gradient in the southern region of Argentinean Patagonia (44–45° S), from 100 mm to 800 mm mean annual precipitation (MAP) and vegetation ranging from desert scrub to closed canopy forest. Gravimetric soil water content tracked changes in seasonal and annual precipitation, with a linear increase in soil water content with increasing MAP. Above‐ground net primary production (ANPP) increased linearly along the gradient of precipitation (ANPP =– 31.2 + 0.52 MAP, r²= 0.84, p= 0.028), supporting the relationship that carbon assimilation is largely controlled by available water in these sites, and was in general agreement with regional models of ANPP and rainfall. However, inorganic soil nitrogen was also highly linearly correlated with both MAP ([N] = 0.19 MAP – 32, r²= 0.96, p= 0.003) and ANPP (ANPP = 2.6 [N_inorganic]+59.4, r²= 0.79, p= 0.042), suggesting a direct control of precipitation on nitrogen turnover and an interaction with nitrogen availability in controlling carbon gain. The asynchrony of precipitation and changes in dominant vegetation may play important roles in determining the carbon‐nitrogen interactions along this rainfall gradient.     

Rain,predators and vegetation lushness may structure web‐building spider communities along precipitation gradients

1. Web building spider communities represent a vehicle for addressing how abiotic and biotic factors interact to structure natural communities. 2. The present study investigates how intense rainfall and potential predation by ants affect the proportion of three‐dimensional (3D) versus two‐dimensional (2D) spider webs along precipitation gradients. 3. Besides capturing prey, 3D webs may provide protection against predators, but they require a much greater material investment to be built than 2D webs. If costs take precedence over predator protection benefits, the proportion of 3D webs would decrease as precipitation increases (the ‘rain intensity’ hypothesis). Alternatively, if predator protection benefits take precedence, and the abundance of ants and other predators increases with precipitation, the proportion of 3D webs would increase with precipitation (the ‘predation risk’ hypothesis). 4. Seven sites were selected along a rain gradient of relatively uniform elevation and latitude in western Ecuador. Rain intensity, ant abundance and vegetation lushness (leaf area, canopy cover, tree diameter) were all observed to increase along the gradient, as did vegetation cover immediately above each web. 5. Relative to 2D webs, 3D webs increased in frequency with annual rainfall, consistent with the predation risk hypothesis but counter to the rain intensity hypothesis. 6. In areas of greater precipitation, however, lusher vegetation provided greater immediate vegetation cover to webs. Microhabitat factors may thus mitigate the destructive power of intense rainfall, thus allowing the predator protection benefits of 3D webs to be realised despite the occurrence of strong rains.     

Mixed hummingbird: Long‐proboscid‐fly pollination in ‘ornithophilous’Embothrium coccineum (Proteaceae) along a rainfall gradient in Patagonia,Argentina

Abstract The pollination ecology of eight populations of the tree Embothrium coccineum was studied along a steep rainfall gradient in NW Patagonia, Argentina. The showy red flowers suggest an ornithophilous pollination syndrome and they have been reported to attract hummingbirds in Argentina and hummingbirds and passerines in Chile. At each population, flower visitors were recorded and floral rewards were analysed. We found a highly significant increase in nectar concentration towards the drier end of the gradient, but this change was not related to the turnover of species in the flower‐visitor assemblage of E. coccineum. In addition to the hummingbird Sephanoides sephaniodes (Green‐Backed Firecrown, Trochilidae) which is widespread throughout the temperate forest at this latitude, other species seem to be locally important as pollinators of E. coccineum in some sites in Argentina (e.g. two long‐tongued tanglewing flies (Nemestrinidae) of the genus Trichophthalma). The long‐dated occurrence of tanglewing flies in South America, relative to the more modern hummingbirds, suggests that ornithophily may be a derived character in E. coccineum, the ancestral condition being pollination by Nemestrinidae.