Matrix models for quantifying competitive intransitivity from species abundance data

In a network of competing species, a competitive intransitivity occurs when the ranking of competitive abilities does not follow a linear hierarchy (A > B > C but C > A). A variety of mathematical models suggests that intransitive networks can prevent or slow down competitive exclusion and maintain biodiversity by enhancing species coexistence. However, it has been difficult to assess empirically the relative importance of intransitive competition because a large number of pairwise species competition experiments are needed to construct a competition matrix that is used to parameterize existing models. Here we introduce a statistical framework for evaluating the contribution of intransitivity to community structure using species abundance matrices that are commonly generated from replicated sampling of species assemblages. We provide metrics and analytical methods for using abundance matrices to estimate species competition and patch transition matrices by using reverse-engineering and a colonization-competition model. These matrices provide complementary metrics to estimate the degree of intransitivity in the competition network of the sampled communities. Benchmark tests reveal that the proposed methods could successfully detect intransitive competition networks, even in the absence of direct measures of pairwise competitive strength. To illustrate the approach, we analyzed patterns of abundance and biomass of five species of necrophagous Diptera and eight species of their hymenopteran parasitoids that co-occur in beech forests in Germany. We found evidence for a strong competitive hierarchy within communities of flies and parasitoids. However, for parasitoids, there was a tendency towards increasing intransitivity in higher weight classes, which represented larger resource patches. These tests provide novel methods for empirically estimating the degree of intransitivity in competitive networks from observational datasets. They can be applied to experimental measures of pairwise species interactions, as well as to spatio-temporal samples of assemblages in homogenous environments or environmental gradients.

Climate and soil attributes determine plant species turnover in global drylands

Aim Geographical, climatic and soil factors are major drivers of plant beta diversity, but their importance for dryland plant communities is poorly known. The aim of this study was to: (1) characterize patterns of beta diversity in global drylands; (2) detect common environmental drivers of beta diversity; and (3) test for thresholds in environmental conditions driving potential shifts in plant species composition. Location Global. Methods Beta diversity was quantified in 224 dryland plant communities from 22 geographical regions on all continents except Antarctica using four complementary measures: the percentage of singletons (species occurring at only one site); Whittaker's beta diversity, β(W); a directional beta diversity metric based on the correlation in species occurrences among spatially contiguous sites, β(R2); and a multivariate abundance-based metric, β(MV). We used linear modelling to quantify the relationships between these metrics of beta diversity and geographical, climatic and soil variables. Results Soil fertility and variability in temperature and rainfall, and to a lesser extent latitude, were the most important environmental predictors of beta diversity. Metrics related to species identity percentage of singletons and β(W) were most sensitive to soil fertility, whereas those metrics related to environmental gradients and abundance (β(R2) and β(MV) were more associated with climate variability. Interactions among soil variables, climatic factors and plant cover were not important determinants of beta diversity. Sites receiving less than 178 mm of annual rainfall differed sharply in species composition from more mesic sites (> 200 mm). Main conclusions Soil fertility and variability in temperature and rainfall are the most important environmental predictors of variation in plant beta diversity in global drylands. Our results suggest that those sites annually receiving c. 178 mm of rainfall will be especially sensitive to future climate changes. These findings may help to define appropriate conservation strategies for mitigating effects of climate change on dryland vegetation.

Moving forward on facilitation research: response to changing environments and effects on the diversity, functioning and evolution of plant communities

Once seen as anomalous, facilitative interactions among plants and their importance for community structure and functioning are now widely recognized. The growing body of modelling, descriptive and experimental studies on facilitation covers a wide variety of terrestrial and aquatic systems throughout the globe. However, the lack of a general body of theory linking facilitation among different types of organisms and biomes and their responses to environmental changes prevents further advances in our knowledge regarding the evolutionary and ecological implications of facilitation in plant communities. Moreover, insights gathered from alternative lines of inquiry may substantially improve our understanding of facilitation, but these have been largely neglected thus far. Despite over 15 years of research and debate on this topic, there is no consensus on the degree to which plant–plant interactions change predictably along environmental gradients (i.e. the stress-gradient hypothesis), and this hinders our ability to predict how plant–plant interactions may affect the response of plant communities to ongoing global environmental change. The existing controversies regarding the response of plant–plant interactions across environmental gradients can be reconciled when clearly considering and determining the species-specificity of the response, the functional or individual stress type, and the scale of interest (pairwise interactions or community-level response). Here, we introduce a theoretical framework to do this, supported by multiple lines of empirical evidence. We also discuss current gaps in our knowledge regarding how plant–plant interactions change along environmental gradients. These include the existence of thresholds in the amount of species-specific stress that a benefactor can alleviate, the linearity or non-linearity of the response of pairwise interactions across distance from the ecological optimum of the beneficiary, and the need to explore further how frequent interactions among multiple species are and how they change across different environments. We review the latest advances in these topics and provide new approaches to fill current gaps in our knowledge. We also apply our theoretical framework to advance our knowledge on the evolutionary aspects of plant facilitation, and the relative importance of facilitation, in comparison with other ecological processes, for maintaining ecosystem structure, functioning and dynamics. We build links between these topics and related fields, such as ecological restoration, woody encroachment, invasion ecology, ecological modelling and biodiversity–ecosystem-functioning relationships. By identifying commonalities and insights from alternative lines of research, we further advance our understanding of facilitation and provide testable hypotheses regarding the role of (positive) biotic interactions in the maintenance of biodiversity and the response of ecological communities to ongoing environmental changes.

Using plant functional traits and phylogenies to understand patterns of plant community assembly in a seasonal tropical forest in Lao PDR

Plant functional traits reflect different evolutionary responses to environmental variation, and among extant species determine the outcomes of interactions between plants and their environment, including other plant species. Thus, combining phylogenetic and trait-based information can be a powerful approach for understanding community assembly processes across a range of spatial scales. We used this approach to investigate tree community composition at Phou Khao Khouay National Park (18°14’-18°32’N; 102°38’- 102°59’E), Laos, where several distinct forest types occur in close proximity. The aim of our study was to examine patterns of plant community assembly across the strong environmental gradients evident at our site. We hypothesized that differences in tree community composition were being driven by an underlying gradient in soil conditions. Thus, we predicted that environmental filtering would predominate at the site and that the filtering would be strongest on sandier soil with low pH, as these are the conditions least favorable to plant growth. We surveyed eleven 0.25 ha (50x50 m) plots for all trees above 10 cm dbh (1221 individual trees, including 47 families, 70 genera and 123 species) and sampled soils in each plot. For each species in the community, we measured 11 commonly studied plant functional traits covering both the leaf and wood economic spectrum traits and we reconstructed a phylogenetic tree for 115 of the species in the community using rbcL and matK sequences downloaded from Genebank (other species were not available). Finally we compared the distribution of trait values and species at two scales (among plots and 10x10m subplots) to examine trait and phylogenetic community structures. Although there was strong evidence that an underlying soil gradient was determining patterns of species composition at the site, our results did not support the hypothesis that the environmental filtering dominated community assembly processes. For the measured plant functional traits there was no consistent pattern of trait dispersion across the site, either when traits were considered individually or when combined in a multivariate analysis. However, there was a significant correlation between the degree of phylogenetic dispersion and the first principle component axis (PCA1) for the soil parameters.Moreover, the more phylogenetically clustered plots were on sandier soils with lower pH. Hence, we suggest that the community assembly processes across our sitemay reflect the influence ofmore conserved traits that we did not measure. Nevertheless, our results are equivocal and other interpretations are possible. Our study illustrates some difficulties in combining trait and phylogenetic approaches that may result from the complexities of integrating spatial and evolutionary processes that vary at different scales.

Worldwide evidence of a unimodal relationship between productivity and plant species richness

The search for predictions of species diversity across environmental gradients has challenged ecologists for decades. The humped-back model (HBM) suggests that plant diversity peaks at intermediate productivity; at low productivity few species can tolerate the environmental stresses, and at high productivity a few highly competitive species dominate. Over time the HBM has become increasingly controversial, and recent studies claim to have refuted it. Here, by using data from coordinated surveys conducted throughout grasslands worldwide and comprising a wide range of site productivities, we provide evidence in support of the HBM pattern at both global and regional extents. The relationships described here provide a foundation for further research into the local, landscape, and historical factors that maintain biodiversity.

hyDRaCAT Spectral Reflectance Library: Hyperspectral Field Spectroscopy and Field Spectro-Goniometry of Siberian and Alaskan Tundra

hyDRaCAT Spectral Reflectance Library for tundra provides the surface reflectance data and the bidirectional reflectance distribution function (BRDF) of important Arctic tundra vegetation communities at representative Siberian and Alaskan tundra sites. The aim of this dataset is the hyperspectral and spectro-directional reflectance characterization as basis for the extraction of vegetation parameters, and the normalization of BRDF effects in off-nadir and multi-temporal remote sensing data. The spectroscopic and field spectro-goniometric measurements were undertaken on the YAMAL2011 expedition of representative Siberian vegetation fields and on the North American Arctic Transect NAAT2012 expedition of Alaskan vegetation fields both belonging to the Greening-of-the-Arctic (GOA) program. For the field spectroscopy each 100 m2 vegetation study grid was divided into quadrats of 1 × 1 m. The averaged reflectance of all quadrats represents the spectral reflectance at the scale of the whole grid at the 10 × 10 m scale. For the surface radiometric measurements two GER1500 portable field spectroradiometers (Spectra Vista Corporation, Poughkeepsie, NY, USA) were used. The GER1500 measures radiance across the wavelength range of 350-1,050 nm, with sampling intervals of 1.5 nm and a radiance accuracy of 1.2 × 10**-1 W/cm**2/nm/sr. In order to increase the signal-to-noise ratio, 32 individual measurements were averaged per one target scan. To minimize variations in the target reflectance due to sun zenith angle changes, all measurements at one study location have been performed under similar sun zenith angles and during clear-sky conditions. The field spectrometer measurements were carried out with a GER1500 UV-VIS spectrometer The spectrogoniometer measurements were carried out with a self-designed spectro-goniometer: the Manual Transportable Instrument platform for ground-based Spectro-directional observations (ManTIS, patent publication number: DE 10 2011 117 713.A1). The ManTIS was equipped with the GER1500 spectrometer allowing spectro-directional measurements with up to 30° viewing zenith angle by full 360° viewing azimuth angles. Measurements in central Yamal (Siberia) at the research site 'Vaskiny Dachi' were carried out in the late summer phenological state from August 12 2011 to August 28 2011. All measurements in Alaska along the North South transect on the North Slope were taken between 29 June and 11 July 2012, ensuring that the vegetation was in the same phenological state near peak growing season.

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea

The distribution of calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea was quantitatively analysed. The samples contain 11 cyst species and the vegetative coccoid Thoracosphaera heimii. Cyst abundance increases towards the deeper parts of the basins and is generally higher in the eastern Mediterranean Sea. Three major distribution characteristics exist: (1) different assemblages in oceanic and neritic regions, (2) little agreement with the associations of areas studied so far like the Atlantic Ocean, and (3) a unique oceanic assemblage in the eastern Mediterranean Sea. A gradual change in cyst assemblages from the western to the eastern Mediterranean Sea was observed and statistically compared with the main environmental gradients in the upper water column. Temperature, nitrate concentration and possibly salinity appear to be the most important factors controlling cyst production. Three groups containing cysts with similar environmental preferences can be distinguished: (1) an eastern Mediterranean group related to relatively high temperature and salinity but low nitrate concentration, (2) a group of more or less consistently abundant cosmopolitan species tolerating or even preferring relatively low temperature and salinity but high nitrate concentration, and (3) a group containing species that are possibly adapted to neritic environments and have probably been transported from coastal areas into the studied regions. In contrast to other calcareous plankton, calcareous dinoflagellate cysts correlate strongly with the main environmental gradients in the Mediterranean Sea, bearing a high potential for palaeoenvironmental reconstructions.

Genetic data from Staurosira (Fragilariophyceae) sedimentary DNA

Genetic investigations on eukaryotic plankton confirmed the existence of modern biogeographic patterns, but analyses of palaeoecological data exploring the temporal variability of these patterns have rarely been presented. Ancient sedimentary DNA proved suitable for investigations of past assemblage turnover in the course of environmental change, but genetic relatedness of the identified lineages has not yet been undertaken. Here, we investigate the relatedness of diatom lineages in Siberian lakes along environmental gradients (i.e. across treeline transects), over geographic distance and through time (i.e. the last 7000 years) using modern and ancient sedimentary DNA. Our results indicate that closely-related Staurosira lineages occur in similar environments and less-related lineages in dissimilar environments, in our case different vegetation and co-varying climatic and limnic variables across treeline transects. Thus our study reveals that environmental conditions rather than geographic distance is reflected by diatom-relatedness patterns in space and time. We tentatively speculate that the detected relatedness pattern in Staurosira across the treeline could be a result of adaptation to diverse environmental conditions across the arctic boreal treeline, however, a geographically-driven divergence and subsequent repopulation of ecologically different habitats might also be a potential explanation for the observed pattern.

Phytoplankton, dissolved inorganic nutrients and hydrography during the ACEx/SIMTECO campaign in the southern Brazilian shelf in June 2012 (SW Atlantic - Winter 2012)

The response of phytoplankton assemblages to hydrographical forcing across the southern Brazilian shelf was studied based on data collected during wintertime (June/2012), complemented with MODIS-Aqua satellite imagery. The in situ data set was comprised by water column structure properties (derived from CTD casts), dissolved inorganic nutrients (ammonium, nitrite, nitrate, phosphate and silicate) and phytoplankton biomass [chlorophyll a (Chl a) concentration] and composition. Phytoplankton assemblages were assessed by both microscopy and HPLC-CHEMTAX approaches. A canonical correspondence analysis associating physical, chemical and phytoplankton composition data at surface evinced a tight coupling between the phytoplankton community and hydrographic conditions, with remarkable environmental gradients across three different domains: the pelagic, outer shelf Tropical Water (TW); the mid shelf domain under influence of Subtropical Shelf Water (STSW); and the inner shelf domain mainly under influence of riverine outflow of the Plata River Plume Water (PPW). Results showed that intrusion of low salinity and nutrient-rich PPW stimulated the phytoplankton growth and diversity within the inner shelf region, with enhanced Chl a levels (>1.3 mg/m**3) and a great abundance of diatoms, ciliates, dinoflagellates, raphidophyceans and cryptophytes. Conversely, other diatoms (e.g. Rhizosolenia clevei), tiny species of prochlorophytes and cyanobacteria and a noticeable contribution of dinoflagellates and other flagellates associated with lower Chl a levels (<0.93 mg/m**3), characterized the TW domain, where low nutrient concentrations and deep upper mixed layer were found. The transitional mid shelf domain showed intermediate levels of both nutrients and Chl a (ranging 1.06-1.59 mg/m**3), and phytoplankton was mainly composed by dinoflagellates, such as Dinophysis spp., and gymnodinioids. Results have shown considerable phytoplankton diversity in winter at that section of the southwestern Atlantic Ocean.

Geographical variation in shell morphology of juvenile snails (Concholepas concholepas) along the physical-chemical gradient of the Chilean coast

Changes in phenotypic traits, such as mollusc shells, are indicative of variations in selective pressure along environmental gradients. Recently, increased sea surface temperature (SST) and ocean acidification (OA) due to increased levels of carbon dioxide in the seawater have been described as selective agents that may affect the biological processes underlying shell formation in calcifying marine organisms. The benthic snail Concholepas concholepas (Muricidae) is widely distributed along the Chilean coast, and so is naturally exposed to a strong physical-chemical latitudinal gradient. In this study, based on elliptical Fourier analysis, we assess changes in shell morphology (outlines analysis) in juvenile C. concholepas collected at northern (23°S), central (33°S) and southern (39°S) locations off the Chilean coast. Shell morphology of individuals collected in northern and central regions correspond to extreme morphotypes, which is in agreement with both the observed regional differences in the shell apex outlines, the high reclassification success of individuals (discriminant function analysis) collected in these regions, and the scaling relationship in shell weight variability among regions. However, these extreme morphotypes showed similar patterns of mineralization of calcium carbonate forms (calcite and aragonite). Geographical variability in shell shape of C. concholepas described by discriminant functions was partially explained by environmental variables (pCO2, SST). This suggests the influence of corrosive waters, such as upwelling and freshwaters penetrating into the coastal ocean, upon spatial variation in shell morphology. Changes in the proportion of calcium carbonate forms precipitated by C. concholepas across their shells and its susceptibility to corrosive coastal waters are discussed.