Disentangling biotic and abiotic drivers of intraspecific trait variation in woody plant seedlings at forest edges

In fragmented forests, edge effects can drive intraspecific variation in seedling performance that influences forest regeneration and plant composition. However, few studies have attempted to disentangle the relative biotic and abiotic drivers of intraspecific variation in seedling performance. In this study, we carried out a seedling transplant experiment with a factorial experimental design on three land‐bridge islands in the Thousand Island Lake, China, using four common native woody plant species. At different distances from the forest edge (2, 8, 32, 128 m), we transplanted four seedlings of each species into each of three cages: full‐cage, for herbivore exclusion; half‐cage, that allowed herbivore access but controlled for caging artifacts; and no‐cage control. In the 576 cages, we recorded branch architecture, leaf traits, and seedling survival for each seedling before and after the experimental treatment. Overall, after one full growing season, edge‐induced abiotic drivers and varied herbivory pressure led to intraspecific variation in seedling performance, including trade‐offs in seedling architecture and resource‐use strategies. However, responses varied across species with different life‐history strategies and depended on the driver in question, such that the abiotic and biotic effects were additive across species, rather than interactive. Edge‐induced abiotic variation modified seedling architecture of a shade‐tolerant species, leading to more vertical rather than lateral growth at edges. Meanwhile, increased herbivory pressure resulted in a shift toward lower dry matter investment in leaves of a light‐demanding species. Our results suggest that edge effects can drive rapid directional shifts in the performance and intraspecific traits of some woody plants from early ontogenetic stages, but most species in this study showed negligible phenotypic responses to edge effects. Moreover, species‐specific responses suggest the importance of interspecific differences modulating the degree of trait plasticity, implying the need to incorporate individual‐level responses when understanding the impact of forest fragmentation on plant communities.     

Patterns and some mechanisms of downstream migration of juvenile salmonids (with reference to the Utkholok and Kalkaveyem rivers in northwestern Kamchatka)

Principal patterns of the downstream migration of juvenile salmonids (Salmonidae) were discovered on the basis of data of three years of observations. It was found that the time of the beginning, the duration, and the scope of the downstream migration in each particular year are determined by a complex of environmental factors. The crucial abiotic factors are the temperature, the water level of the river, and the illumination. The biotic factors comprise the complex trophic factor. The abiotic environmental factors play the leading role in the realization of the downstream migration at the early ontogenetic stages. The role of the biotic environmental factors increases at the late ontogenetic stages. It was demonstrated that the juveniles that perform downstream migrations are a heterogeneous group consisting of fish of different species and specimens at different ontogenetic stages, of various sizes and ages, and those ready for the transition to the marine environment to variable degrees. The analysis of the patterns of the downstream migration allowed us to describe some mechanisms that control the realization of this process.     

Ökologische Untersuchungen an Hydroiden des Felslitorals von Santa Marta (Kolumbien)

The ecology of tropic hydroids on the rocky shore of Santa Marta, Colombia, was studied from January 1971 to April 1972. Main interests concentrated on vertical distribution, temporal succession, activity, maturity, population density, shape and growth of colonies, and on type of settlement. These aspects have been related to quantitative data and to variations in biotic and abiotic factors. The results are compared with those obtained on other types of shores, viz. artificial and sandy shores. They indicate that local abiotic differences in water quality are influenced, above all, by the sequence of dry and rainy seasons. In a rapidly changing abiotic environment, life cycles and ontogenetic stages (settlement, growth, maturity) follow each other faster than in a static environment. With increasing water depth, the density of the hydroid populations decreases, while the size of single colonies increases. In addition, with increasing depth, the length of the maturity period increases. For the same species, the time of attainment of maturity may be different in different localities. Hydroid species with short life cycles settle faster than species with long life cycles. Hydroid polyps may counteract environmental perturbations by changes in activity. Distribution and zonation of hydroid populations depend to a high degree on water movement, sun orbit, and shade-spending coastal profiles.     

Testing the evolutionary constraints of metamorphosis: The ontogeny of head shape in Triturus newts

In vertebrates with complex, biphasic, life cycles, larvae have a distinct morphology and ecological preferences compared to metamorphosed juveniles and adults. In amphibians, abrupt and rapid metamorphic changes transform aquatic larvae to terrestrial juveniles. The main aim of this study is to test whether, relative to larval stages, metamorphosis (1) resets the pattern of variation between ontogenetic stages and species, (2) constrains intraspecific morphological variability, and (3) similar to the “hour‐glass” model reduces morphological disparity. We explore postembryonic ontogenetic trajectories of head shape (from hatching to completed metamorphosis) of two well‐defined, morphologically distinct Triturus newts species and their F1 hybrids. Variation in head shape is quantified and compared on two levels: dynamic (across ontogenetic stages) and static (at a particular stage). Our results show that the ontogenetic trajectories diverge early during development and continue to diverge throughout larval stages and metamorphosis. The high within‐group variance and the largest disparity level (between‐group variance) characterize the metamorphosed stage. Hence, our results indicate that metamorphosis does not canalize head shape variation generated during larval development and that metamorphosed phenotype is not more constrained relative to larval ones. Therefore, metamorphosis cannot be regarded as a developmental constraint, at least not for salamander head shape.     

Factors linked to interannual variation in the metazoan parasite communities of black skipjack,Euthynnus lineatus (Pisces: Scombridae)

Marine parasite communities can exhibit temporal and spatial changes in response to seasonal and local variations in several biotic and abiotic environmental factors. Limited attention has been given to the influence of abiotic factors, so their effects on parasite community structure remain unclear. A total of 496 specimens of Euthynnus lineatus were collected over a 7‐year period (2012–2018) from Acapulco Bay, Mexico. Their parasite communities were analyzed to determine if they experience interannual variations due to local biotic and abiotic factors. Thirty‐three metazoan parasite species were recovered and identified: four species of Monogenea (adults); 16 of Digenea (one larvae and 15 adults); two of Acanthocephala (adults); two of Cestoda (larvae); three of Nematoda (two larvae and one adult); and six of Crustacea (three Copepoda, and three Isopoda). Species richness was greatest among the digeneans, which represented 48% of the total species recovered, followed by the crustaceans (19% of total species). Species richness at the component community level (14–24 species) was similar to reported richness in other small tuna species. The component communities and infracommunities of E. lineatus exhibited a similar pattern: high species richness and diversity, and numerical dominance by a single species, mainly by one of the didymozoids Allopseudocolocyntotrema claviforme or Pseudocolocyntotrema yaito. Parasite community structure and species composition varied among sampling years. Variations were possibly caused by a combination of abiotic and biotic factors which generated notable changes in the infection levels of several component species during the study period. These communities may therefore be unpredictable in terms of structure and species composition, as has been suggested for other communities of marine parasites.     

Spatial variation in abiotic and biotic factors in a floodplain determine anuran body size and growth rate at metamorphosis

Body size at metamorphosis is a critical trait in the life history of amphibians. Despite the wide-spread use of amphibians as experimental model organisms, there is a limited understanding of how multiple abiotic and biotic factors affect the variation in metamorphic traits under natural conditions. The aim of our study was to quantify the effects of abiotic and biotic factors on spatial variation in the body size of tadpoles and size at metamorphosis of the European common toad (Bufo b. spinosus). Our study population was distributed over the riverbed (active tract) and the fringing riparian forest of a natural floodplain. The riverbed had warm ponds with variable hydroperiod and few predators, whereas the forest had ponds with the opposite characteristics. Spatial variation in body size at metamorphosis was governed by the interactive effects of abiotic and biotic factors. The particular form of the interaction between water temperature and intraspecific tadpole density suggests that abiotic factors laid the foundation for biotic factors: intraspecific density decreased growth only at high temperature. Predation and intraspecific density jointly reduced metamorphic size. Interspecific density had a negligible affect on body size at metamorphosis, suggesting weak inter-anuran interactions in the larval stage. Population density at metamorphosis was about one to two orders of magnitudes higher in the riverbed ponds than in the forest ponds, mainly because of lower tadpole mortality. Based on our results, we conclude that ponds in the riverbed appear to play a pivotal role for the population because tadpole growth and survival is best in this habitat.     

European grassland ecosystems: threatened hotspots of biodiversity

Biodiversity is not homogenously distributed over the globe, and ecosystems differ strongly in the number of species they provide. With this special issue we highlight the ecology and endangerment of one of the most diverse ecosystem of Europe: the European grassland ecosystems. The selected 16 contributions describe interactions from below-ground to the atmosphere and focus on (1) effects of abiotic and biotic on species diversity, (2) the impact of various factors along spatial and temporal gradients, (3) the relevance of falling abandoned and eutrophication—including countervailing management strategies like encroachment; and (4) intraspecific effects based on physiology, genetics and intraspecific plasticity. The contributions cover fungi, plants, and invertebrates and highlight effects taking place at the level of ecosystem, species community, species, populations, and also within individuals (physiology and genetics).     

Characterizing Tropical Tree Species Growth Strategies: Learning from Inter-Individual Variability and Scale Invariance

Understanding how tropical tree species differ in their growth strategies is critical to predict forest dynamics and assess species coexistence. Although tree growth is highly variable in tropical forests, species maximum growth is often considered as a major axis synthesizing species strategies, with fast-growing pioneer and slow-growing shade tolerant species as emblematic representatives. We used a hierarchical linear mixed model and 21-years long tree diameter increment series in a monsoon forest of the Western Ghats, India, to characterize species growth strategies and question whether maximum growth summarizes these strategies. We quantified both species responses to biotic and abiotic factors and individual tree effects unexplained by these factors. Growth responses to competition and tree size appeared highly variable among species which led to reversals in performance ranking along those two gradients. However, species-specific responses largely overlapped due to large unexplained variability resulting mostly from inter-individual growth differences consistent over time. On average one-third of the variability captured by our model was explained by covariates. This emphasizes the high dimensionality of the tree growth process, i.e. the fact that trees differ in many dimensions (genetics, life history) influencing their growth response to environmental gradients, some being unmeasured or unmeasurable. In addition, intraspecific variability increased as a power function of species maximum growth partly as a result of higher absolute responses of fast-growing species to competition and tree size. However, covariates explained on average the same proportion of intraspecific variability for slow- and fast-growing species, which showed the same range of relative responses to competition and tree size. These results reflect a scale invariance of the growth process, underlining that slow- and fast-growing species exhibit the same range of growth strategies.     

Biogeography of parasitism in freshwater fish: spatial patterns in hot spots of infection

Contrary to species occurrence, little is known about the determinants of spatial patterns of intraspecific variation in abundance, particularly for parasitic organisms. In this study, we provide a multi‐faceted overview of spatial patterns in parasite abundance and examine several potential underlying processes. We first tested for a latitudinal gradient in local abundance of the regionally most common parasite species and whether these species achieve higher abundances at the same localities (shared hot spots of infection). Secondly, we tested whether intraspecific similarity in local abundance between sites follows a spatial distance decay pattern or is better explained by variation in extrinsic biotic and abiotic factors between localities related to local parasite transmission success. We examined the infection landscape of a model fish host system (common and upland bullies, genus Gobiomorphus: Eleotridae) across its entire distributional range. We applied general linear models to test the effect of latitude on each species local abundance independently, including the abundance of each co‐infecting species as another predictor. We computed multiple regressions on distance matrices among localities based on abundance of each of the four most common trematode species, as well as for geographic distance, biotic and abiotic distinctness of the localities. Our results showed that the most widely distributed parasites of bullies also achieve the highest mean local abundances, following the abundance – occupancy relationship. Variation in local abundance of any focal parasite species was independent of latitude, the abundance of co‐occurring species and spatial distance or disparity in biotic attributes between localities. For only one parasite species, similarity of abundance between sites covaried with the extent of abiotic differences between sites. The lack of association between hot spots of infection for co‐occurring species reinforces the geographic mosaic scenario in which hosts and parasites coevolve by suggesting non‐deterministic, species‐specific variation in parasite abundance across space.     

Biogenesis, evolution, and functions of plant microRNAs

This review focuses on the biological role of one class of plant small RNAs, ～22-nt microRNAs (miRNAs). The majority of plant miRNA targets are genes encoding the effector factors of cell signaling pathways. The regulation of their expression is necessary for both ontogenesis and rapid response of plants to biotic and abiotic stress factors. We also summarized current views on the biogenesis and evolution of plant miRNAs as well as the techniques used for their investigation.     

Disparate relatives: Life histories vary more in genera occupying intermediate environments

Species within clades are commonly assumed to share similar life history traits, but within a given region some clades show much greater variability in traits than others. Are variable clades older, allowing more time for trait diversification? Or do they occupy particular environments, providing a wider range of abiotic or biotic opportunities for the establishment and maintenance of diverse trait attributes? Does environmental opportunity increase trait variability across all species, or is it specific to species belonging to the same clade, increasing only within-clade trait variability? We studied the variability of six life-history traits (initiation of flowering, duration of flowering, plant life span, seed mass, stress tolerance, type of reproduction) within 383 angiosperm genera from Central Europe distributed along six abiotic gradients. We compared patterns of within-genus variability to those present in the entire dataset, independent of genus membership. We found that trait variability differed strongly between genera, but did not depend on their age. Trait variability was higher within genera occupying intermediate positions along regional abiotic environmental gradients, compared with patterns across the entire dataset (and unbiased by geographical sampling, family membership or species richness). Increasing trait variability within genera reflected increasing independence of traits from the abiotic environment. We conclude that intermediate abiotic environments play an important role in maintaining and possibly generating the striking diversity of life history traits present within certain clades. They may do so by relaxing the abiotic constraints on the evolution and maintenance of species traits within clades.     

Accounting for dispersal and biotic interactions to disentangle the drivers of species distributions and their abundances

Although abiotic factors, together with dispersal and biotic interactions, are often suggested to explain the distribution of species and their abundances, species distribution models usually focus on abiotic factors only. We propose an integrative framework linking ecological theory, empirical data and statistical models to understand the distribution of species and their abundances together with the underlying community assembly dynamics. We illustrate our approach with 21 plant species in the French Alps. We show that a spatially nested modelling framework significantly improves the model's performance and that the spatial variations of species presence-absence and abundances are predominantly explained by different factors. We also show that incorporating abiotic, dispersal and biotic factors into the same model bring new insights to our understanding of community assembly. This approach, at the crossroads between community ecology and biogeography, is a promising avenue for a better understanding of species co-existence and biodiversity distribution.     

Biogeography of parasite abundance: latitudinal gradient and distance decay of similarity in the abundance of fleas and mites,parasitic on small mammals in the Palearctic,at three spatial scales

We tested whether biogeographic patterns characteristic for biological communities can also apply to populations and investigated geographic patterns of variation in abundance of ectoparasites (fleas and mites) collected from bodies of their small mammalian hosts (rodents and shrews) in the Palearctic at continental, regional and local scales. We asked whether (i) there is a relationship between latitude and abundance and (ii) similarity in abundance follows a distance decay pattern or it is better explained by variation in extrinsic biotic and abiotic factors. We analysed the effect of latitude on mean intraspecific abundance using general linear models including proportional abundance of its principal host as an additional predictor variable. Then, we examined the relative effect of geographic distance, biotic and abiotic dissimilarities among regions, subregions or localities on the intraspecific dissimilarity in abundance among regions, subregions or localities using Generalized Dissimilarity Modelling. We found no relationship between latitude and intraspecific flea or mite abundance. In both taxa, environmental dissimilarity explained the largest part of the deviance of spatial variation in abundance, whereas the effect of the dissimilarity in the principal host abundance was of secondary importance and the effect of geographic distance was minor. These patterns were generally consistent across the three spatial scales, although environmental variation and dissimilarity in principal host abundance were equally important at the local scale in fleas but not in mites. We conclude that biogeographic patterns related to latitude and geographic distance do not apply to spatial variation of ectoparasite abundance. Instead, the geographic distribution of abundance in arthropod ectoparasites depends on their responses, mainly to the off-host environment and to a lesser extent the abundance of their principal hosts.     

Microhabitat selection in the common lizard: implications of biotic interactions,age, sex,local processes,and model transferability among populations

Modeling species' habitat requirements are crucial to assess impacts of global change, for conservation efforts and to test mechanisms driving species presence. While the influence of abiotic factors has been widely examined, the importance of biotic factors and biotic interactions, and the potential implications of local processes are not well understood. Testing their importance requires additional knowledge and analyses at local habitat scale. Here, we recorded the locations of species presence at the microhabitat scale and measured abiotic and biotic parameters in three different common lizard (Zootoca vivipara) populations using a standardized sampling protocol. Thereafter, space use models and cross‐evaluations among populations were run to infer local processes and estimate the importance of biotic parameters, biotic interactions, sex, and age. Biotic parameters explained more variation than abiotic parameters, and intraspecific interactions significantly predicted the spatial distribution. Significant differences among populations in the relationship between abiotic parameters and lizard distribution, and the greater model transferability within populations than between populations are in line with effects predicted by local adaptation and/or phenotypic plasticity. These results underline the importance of including biotic parameters and biotic interactions in space use models at the population level. There were significant differences in space use between sexes, and between adults and yearlings, the latter showing no association with the measured parameters. Consequently, predictive habitat models at the population level taking into account different sexes and age classes are required to understand a specie's ecological requirements and to allow for precise conservation strategies. Our study therefore stresses that future predictive habitat models at the population level and their transferability should take these parameters into account.     

Abiotic factors shape temporal variation in the structure of an ant–plant network

Despite recognition of key biotic processes in shaping the structure of biological communities, few empirical studies have explored the influences of abiotic factors on the structural properties of mutualistic networks. We tested whether temperature and precipitation contribute to temporal variation in the nestedness of mutualistic ant–plant networks. While maintaining their nested structure, nestedness increased with mean monthly precipitation and, particularly, with monthly temperature. Moreover, some species changed their role in network structure, shifting from peripheral to core species within the nested network. We could summarize that abiotic factors affect plant species in the vegetation (e.g., phenology), meaning presence/absence of food sources, consequently an increase/decrease of associations with ants, and finally, these variations to fluctuations in nestedness. While biotic factors are certainly important, greater attention needs to be given to abiotic factors as underlying determinants of the structures of ecological networks.     

Cultural and climatic changes shape the evolutionary history of the Uralic languages

Quantitative phylogenetic methods have been used to study the evolutionary relationships and divergence times of biological species, and recently, these have also been applied to linguistic data to elucidate the evolutionary history of language families. In biology, the factors driving macroevolutionary processes are assumed to be either mainly biotic (the Red Queen model) or mainly abiotic (the Court Jester model) or a combination of both. The applicability of these models is assumed to depend on the temporal and spatial scale observed as biotic factors act on species divergence faster and in smaller spatial scale than the abiotic factors. Here, we used the Uralic language family to investigate whether both ‘biotic’ interactions (i.e. cultural interactions) and abiotic changes (i.e. climatic fluctuations) are also connected to language diversification. We estimated the times of divergence using Bayesian phylogenetics with a relaxed‐clock method and related our results to climatic, historical and archaeological information. Our timing results paralleled the previous linguistic studies but suggested a later divergence of Finno‐Ugric, Finnic and Saami languages. Some of the divergences co‐occurred with climatic fluctuation and some with cultural interaction and migrations of populations. Thus, we suggest that both ‘biotic’ and abiotic factors contribute either directly or indirectly to the diversification of languages and that both models can be applied when studying language evolution.     

Temporal variation in plant-soil feedback controls succession

Soil abiotic and biotic factors play key roles in plant community dynamics. However, little is known about how soil biota influence vegetation changes over time. Here, we show that the effects of soil organisms may depend on both the successional development of ecosystems and on the successional position of the plants involved. In model systems of plants and soils from different successional stages, we observed negative plant–soil feedback for early-successional plant species, neutral feedback for mid-successional species, and positive feedback for late-successional species. The negative feedback of early-successional plants was independent of soil origin, while late-successional plants performed best in late- and worst in early-successional soil. Increased performance of the subordinate, late-successional plants resulted in enhanced plant community diversity. Observed feedback effects were more related to soil biota than to abiotic conditions. Our results show that temporal variations in plant–soil interactions profoundly contribute to plant community assemblage and ecosystem development.     

Biotic and abiotic variables show little redundancy in explaining tree species distributions

Abiotic factors such as climate and soil determine the species fundamental niche, which is further constrained by biotic interactions such as interspecific competition. To parameterize this realized niche, species distribution models (SDMs) most often relate species occurrence data to abiotic variables, but few SDM studies include biotic predictors to help explain species distributions. Therefore, most predictions of species distributions under future climates assume implicitly that biotic interactions remain constant or exert only minor influence on large‐scale spatial distributions, which is also largely expected for species with high competitive ability. We examined the extent to which variance explained by SDMs can be attributed to abiotic or biotic predictors and how this depends on species traits. We fit generalized linear models for 11 common tree species in Switzerland using three different sets of predictor variables: biotic, abiotic, and the combination of both sets. We used variance partitioning to estimate the proportion of the variance explained by biotic and abiotic predictors, jointly and independently. Inclusion of biotic predictors improved the SDMs substantially. The joint contribution of biotic and abiotic predictors to explained deviance was relatively small (~9%) compared to the contribution of each predictor set individually (~20% each), indicating that the additional information on the realized niche brought by adding other species as predictors was largely independent of the abiotic (topo‐climatic) predictors. The influence of biotic predictors was relatively high for species preferably growing under low disturbance and low abiotic stress, species with long seed dispersal distances, species with high shade tolerance as juveniles and adults, and species that occur frequently and are dominant across the landscape. The influence of biotic variables on SDM performance indicates that community composition and other local biotic factors or abiotic processes not included in the abiotic predictors strongly influence prediction of species distributions. Improved prediction of species' potential distributions in future climates and communities may assist strategies for sustainable forest management.     

Intraspecific taxonomic differentiation in ticks (Acari: Ixodidae) in the view of the morphological conception of species

The criterions for two most used intraspecific taxonomic ranks of ticks--subspecies and morphotype, have been formulated on the basis of the study of morphological variation in the distribution range of all active ontogenetic stages of 11 polymorphic species. All these species are vectors of transmissible diseases. They have vast distribution ranges and different types of host-parasite relationships. Subspecies have the complexes of visual morphological differences expressed in one or both sexes of mature ticks more limited, than those of related species. At immature stages differences of subspecies consist more often in morphometric characters and can be established by the methods of mathematical statistics only. Morphotypes, as a rule, differ at each corresponding stage by "their own" complexes of morphometric characters. All differential parameters of studied morphotypes are overlapped, but have statistically significant differences (by the Student's test). The concrete variations of differentiation by subspecies and morphotypes have been considered. The historical factors of intraspecific differentiation have been reconstructed for each species.     

The life form concept and the use in the analysis of life cycle evolutionary strategies

The life form is a generalized morphoecological characteristic of an animal giving an idea of the organism as a whole, its position and function and functional role in the ecosystem. This characteristic is inherent to a species or to a group of congeneric species (for applied goal it is better to use a genus, not species) considered in the framework of higher taxon (from family to type). The principal contradiction of life form concept is determined by the existence of ontogenetic stages and changes of life forms during the whole life of individual. It is usually assumed that the concept of life form should be applied only to the adult stage, thus ignoring the integral character of the life cycle as indivisible unit of selection, evolution and functioning in ecosystem. We propose that a morphologically specific ontogenetic of a given species should be used as an elementary lowest unit in the classification of life forms. Thus it can be considered as integrated internally structured morphoecological unit in time and multidimensional space of abiotic and biotic environmental factors. As an example we describe the types of reproductive strategies and classification of elementary (ontogenetic) life forms in cephalopods. We present characteristics of the life cycle of some typical cephalopod species inhabiting different biotopes and having different models of locomotion, feeding, reproduction and development.