Does disturbance regime change community assembly of angiosperm plant communities in the boreal forest?

Aims To examine if and how species and phylogenetic diversity change in relation to disturbance, we conducted a review of ecological literature by testing the consistency of the relationship between phylogenetic diversity and disturbance and compared taxonomic groups, type of disturbance and ecosystem/habitat context. We provide a case study of the phylogenetic diversity–disturbance relationship in angiosperm plant communities of a boreal forest region, compared with types of natural and anthropogenic disturbances and plant growth forms.Methods Using a large-scale sampling plot network along a complete (0–100%) anthropogenic disturbance gradient in the boreal biome, we compared the changes of angiosperm plant community structure and composition across plots. We estimated natural disturbance with historical records of major fires. We then calculated phylogenetic diversity indexes and determined species richness in order to compare linear and polynomial trends along disturbance gradients. We also compared the changes of community structure for different types of anthropogenic disturbances and examined how the relationships between species and phylogenetic diversity and disturbance regimes vary among three different life forms (i.e. forbs, graminoids and woody plants).Important findings Phylogenetic diversity was inconsistently related to disturbance in previous studies, regardless of taxon, disturbance type or ecosystem context. In the understudied boreal ecosystem, angiosperm plant communities varied greatly in species richness and phylogenetic diversity along anthropogenic disturbance gradients and among different disturbance types. In general, a quadratic curve described the relationship between species richness and anthropogenic disturbance, with the highest richness at intermediate anthropogenic disturbance levels. However, phylogenetic diversity was not related to disturbance in any consistent manner and species richness was not correlated with phylogenetic diversity. Phylogenetic relatedness was also inconsistent across plant growth forms and different anthropogenic disturbance types. Unlike the inconsistent patterns observed for anthropogenic disturbance, community assembly among localities varying in time since natural disturbance exhibited a distinct signature of phylogenetic relatedness, although those trends varied among plant growth forms.     

Assessing genetic diversity in clonal organisms: low diversity or low resolution? Combining power and cost efficiency in selecting markers

The increasing use of molecular tools to study populations of clonal organisms leads us to question whether the low polymorphism found in many studies reflects limited genetic diversity in populations or the limitations of the markers used. Here we used microsatellite datasets for two sea grass species to provide a combinatory statistic, combined with a likelihood approach to estimate the probability of identical multilocus genotypes (MLGs) to be shared by distinct individuals, in order to ascertain the efficiency of the markers used and to optimize cost-efficiently the choice of markers to use for deriving unbiased estimates of genetic diversity. These results strongly indicate that conclusions from studies on clonal organisms derived using markers showing low polymorphism, including microsatellites, should be reassessed using appropriate polymorphic markers.     

Does shelter enhance early seedling survival in dry environments? A test with eight Mediterranean species

Question: Do solid‐walled polyethylene tubes and mesh fabric tubes improve the short‐term survival of eight Mediterranean tree and shrub species often used in the restoration of arid environments? Location: We conducted two experimental plantations in degraded field sites in the province of Almería (SE Spain), under arid Mediterranean conditions. Methods: One‐year‐old seedlings of Ceratonia siliqua, Juniperus phoenicea, Olea europaea, Pinus halepensis, Pinus pinaster, Quercus coccifera, Quercus ilex and Tetraclinis articulata were planted either sheltered by one of the above shelter tubes, or by being left unsheltered. Survival was recorded the first growing season after planting, which was a very dry season. Results: Overall, seedling survival ranged from as little as 0% to 24%, and tree shelters consistently enhanced survival in Quercus species only, ranging from 16% in walled shelters to 8% in mesh shelters. Shelters failed to boost survival in the six remaining species. Conclusion: The results of this study suggest that both walled and mesh shelters were mostly ineffective at increasing seedling survival for the Mediterranean species used in this experiment; these species coincide with those used in restoration programs. The use of shelters in restoration programs conducted in arid environments should be reconsidered, while walled shelters might be advisable for Mediterranean Quercus species only. Further research is necessary to develop and assess improved types of shelters for arid environments.     

Does population size affect genetic diversity? A test with sympatric lizard species

Genetic diversity is a fundamental requirement for evolution and adaptation. Nonetheless, the forces that maintain patterns of genetic variation in wild populations are not completely understood. Neutral theory posits that genetic diversity will increase with a larger effective population size and the decreasing effects of drift. However, the lack of compelling evidence for a relationship between genetic diversity and population size in comparative studies has generated some skepticism over the degree that neutral sequence evolution drives overall patterns of diversity. The goal of this study was to measure genetic diversity among sympatric populations of related lizard species that differ in population size and other ecological factors. By sampling related species from a single geographic location, we aimed to reduce nuisance variance in genetic diversity owing to species differences, for example, in mutation rates or historical biogeography. We compared populations of zebra-tailed lizards and western banded geckos, which are abundant and short-lived, to chuckwallas and desert iguanas, which are less common and long-lived. We assessed population genetic diversity at three protein-coding loci for each species. Our results were consistent with the predictions of neutral theory, as the abundant species almost always had higher levels of haplotype diversity than the less common species. Higher population genetic diversity in the abundant species is likely due to a combination of demographic factors, including larger local population sizes (and presumably effective population sizes), faster generation times and high rates of gene flow with other populations.     

Polyploidy and microsatellite variation in the relict tree Prunus lusitanica L.: how effective are refugia in preserving genotypic diversity of clonal taxa?

Refugia are expected to preserve genetic variation of relict taxa, especially in polyploids, because high gene dosages could prevent genetic erosion in small isolated populations. However, other attributes linked to polyploidy, such as asexual reproduction, may strongly limit the levels of genetic variability in relict populations. Here, ploidy levels and patterns of genetic variation at nuclear microsatellite loci were analysed in Prunus lusitanica, a polyploid species with clonal reproduction that is considered a paradigmatic example of a Tertiary relict. Sampling in this study considered a total of 20 populations of three subspecies: mainland lusitanica (Iberian Peninsula and Morocco), and island azorica (Azores) and hixa (Canary Islands and Madeira). Flow cytometry results supported an octoploid genome for lusitanica and hixa, whereas a 16‐ploid level was inferred for azorica. Fixed heterozygosity of a few allele variants at most microsatellite loci resulted in levels of allelic diversity much lower than those expected for a high‐order polyploid. Islands as a whole did not contain higher levels of genetic variation (allelic or genotypic) than mainland refuges, but island populations displayed more private alleles and higher genotypic diversity in old volcanic areas. Patterns of microsatellite variation were compatible with the occurrence of clonal individuals in all but two island populations, and the incidence of clonality within populations negatively correlated with the estimated timing of colonization. Our results also suggest that gene flow has been very rare among populations, and thus population growth following founder events was apparently mediated by clonality rather than seed recruitment, especially in mainland areas. This study extends to clonal taxa the idea of oceanic islands as important refugia for biodiversity, since the conditions for generation and maintenance of clonal diversity (i.e. occasional events of sexual reproduction, mutation and/or seed immigration) appear to have been more frequent in these enclaves than in mainland areas.     

Issues affecting the measurement of disturbance response patterns in herbaceous vegetation – A test of the intermediate disturbance hypothesis

The 'intermediate disturbance hypothesis' (IDH) predicts maximum species diversity at intermediate levels of disturbance. Disturbance levels are measured by intensity or frequency of disturbance or by the time since disturbance. The IDH has been tested over a wide range of studies and communities with results either accepting or rejecting the hypothesis. The objectives of this study were to determine how observed disturbance response patterns for the same herbaceous plant community are influenced by modelling techniques, particularly in relation to the expression of disturbance gradients, choice of species diversity indices, and time of assessment since disturbance. Response patterns were examined using a multi-factorial disturbance experiment involving grazing (absent and present); fire (absent and present); soil cultivation (none, 5 and 20 cm); and amendment (none, fertiliser, and fertiliser plus clover seeds). Generalized linear models with a b-spline function were used to define response patterns for five different disturbance gradients over a 24-month period using three indices of diversity. Seven basic disturbance response models were recognized ranging from the classical IDH bell-shaped pattern through increasing or decreasing trends, to a no-change model with increasing levels of disturbance. Only 6.7% of all models were consistent with the IDH. The no-change model was found in nearly half the cases investigated and increased in occurrence with time since disturbance. The choice of the disturbance intensity gradient, the species diversity index used and the time of assessment after disturbance significantly influenced the frequency of occurrence of the disturbance response models observed. Consequently, the responses of vegetation to disturbance gradients show many patterns depending on how they are defined and modelled rather than the simple bell-shaped curve as predicted by the IDH, but the ecological mechanisms supporting some of these patterns need to be further investigated.     

Does environmental disturbance also influence within-stream beta diversity of macroinvertebrate assemblages in tropical streams?

Riparian deforestation is a major threat to the ecological integrity of streams and aquatic biodiversity, influencing microhabitat availability and susceptibility to disturbances. Here we tested if riparian deforestation of tropical streams influenced beta diversity of macroinvertebrate assemblages, by comparing indices that weighted differentially rare and dominant taxa, and testing if nestedness in community composition increased in deforested streams. Within-stream beta diversity was higher in deforested than forested streams, mainly due to taxon loss and higher dominance. In disturbed streams, higher sedimentation in pool mesohabitats resulted in larger differences in community composition, whereas mesohabitats in forested streams were more stable.     

Does disturbance affect bud bank size and belowground structures diversity in Brazilian subtropical grasslands?

Brazilian Campos grasslands are ecosystems under high frequency of disturbance by grazing and fires. Absence of such disturbances may lead to shrub encroachment and loss of plant diversity. Vegetation regeneration after disturbance in these grasslands occurs mostly by resprouting from belowground structures. We analyzed the importance of bud bank and belowground bud bearing organs in Campos grasslands. We hypothesize that the longer the intervals between disturbances are, the smaller the size of the bud bank is. Additionally, diversity and frequency of belowground organs should also decrease in areas without disturbance for many years. We sampled 20 soil cores from areas under different types of disturbance: grazed, exclusion from disturbance for two, six, 15 and 30 years. Belowground biomass was sorted for different growth forms and types of bud bearing organs. We found a decrease in bud bank size with longer disturbance intervals. Forbs showed the most drastic decrease in bud bank size in the absence of disturbance, which indicates that they are very sensitive to changes in disturbance regimes. Xylopodia (woody gemmiferous belowground organs with hypocotyl-root origin) were typical for areas under influence of recurrent fires. The diversity of belowground bud bearing structures decreased in the absence of disturbance. Longer intervals between disturbance events, resulting in decrease of bud bank size and heterogeneity of belowground organs may lead to the decline and even disappearance of species that relay on resprouting from the bud bank upon disturbance.     

Does disturbance drive the collapse of biotic interactions at the severe end of a diversity–biomass gradient?

It has been recently proposed that the decrease in diversity towards the severe end of the humped-back diversity–biomass model of Grime was driven by a collapse of facilitation due to extreme conditions of either stress or physical disturbance. In order to test the hypothesis that disturbance is the primary direct factor driving the collapse of interactions occurring along environmental severity gradients, we conducted a removal experiment in the highly stressed French coastal dunes along a gradient of disturbance due to sand burial. Four dune species were used as targets and transplanted with and without neighbours in four communities along the gradient. The experiment was conducted twice, a dry and an average year. Results of the experiment showed that during the dry year the effect of the environment was prominent and only one species was facilitated for survival in the least disturbed community. During the average year, interactions for growth were important only in the same community, with positive or negative responses depending on the natural position of the target species within the coastal dune gradient. In accordance with our hypothesis, most interactions for both survival and growth were observed in the least disturbed community exhibiting the highest diversity. There were no interactions in the most disturbed community with the lowest diversity.     

Does pollution influence small mammal diet in the field? A metabarcoding approach in a generalist consumer

Mammals are mainly exposed to trace metals (TMs) via consuming contaminated food. Several studies have demonstrated relationships between metal concentrations in food and in animal tissues. However, potential effects of TMs on feeding behaviour of wildlife have been poorly documented under field conditions, despite experimental evidence showing that food selection is impacted by resource contamination. Here, we test the hypothesis that the diet of a generalist rodent, the wood mouse (Apodemus sylvaticus), is altered by soil TM contamination in the field. Wood mice were sampled in spring and in autumn along a gradient of soil contamination in the surroundings of a former smelter located in northern France. Available resources in the field were inventoried, and the diet of the animals was analysed using DNA “metabarcoding.” We demonstrated that (a) relationship between the resource richness in the diet and their richness in the field was altered by soil metal contamination. Wood mice specialized their diet along the gradient of soil metal contamination for both plant and invertebrate resources in spring. We also showed that (b) preference for Salicaceae, a plant family accumulating metals, decreased when soil contamination increased. These results suggest that environmental TM pollution could act as a force modulating trophic interactions in terrestrial food webs, thereby affecting wildlife exposure to contaminants by trophic route.     

Do metal-rich plants deter herbivores? A field test of the defence hypothesis

Some plant species growing on metalliferous soils are able to accumulate heavy metals in their shoots up to very high concentrations, but the selective advantage of this behaviour is still unknown. The most popular hypothesis, that metals protect plants against herbivores, has been tested several times in laboratory conditions, with contradictory results. We carried out the first large-scale test of the defence hypothesis in eight natural populations of the model Zn hyperaccumulator Thlaspi caerulescens J. and C. Presl (Brassicaceae). In two climatic regions (temperate, Belgium–Luxembourg, and Mediterranean, southern France), we worked in metalliferous and in normal, uncontaminated environments, with plants spanning a wide range of Zn concentrations. We also examined the importance of glucosinolates (main secondary metabolites of Brassicaceae) as antiherbivore defences. When exposed to natural herbivore populations, T. caerulescens suffered lower herbivory pressures in metal-enriched soils than in normal soils, both in Belgium–Luxembourg and in southern France. The trapping of gastropods shows an overall lower population density in metalliferous compared to normal environments, which suggests that herbivory pressure from gastropods is lower on metalliferous soils. In addition, foliar concentration of glucosinolates was constitutively lower in all populations from metal-enriched soils, suggesting that these have evolved towards lower investment in organic defences in response to lower herbivory pressure. The Zn concentration of plants had a protective role only for Belgian metallicolous plants when transplanted in normal soils of Luxembourg. These results do not support the hypothesis that Zn plays a key role in the protection of T. caerulescens against enemies. In contrast, glucosinolates appear to be directly involved in the defence of this hyperaccumulator against herbivores.     

Does habitat heterogeneity affect the diversity of epigaeic arthropods in vineyards?

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Does competition or facilitation regulate migrant ungulate populations in the Serengeti? A test of hypotheses

Summary Interspecific competition and facilitation have both been proposed as processes promoting species separation and co-existence in African ungulates. In one group of grazers on the Serengeti plains, comprising wildebeest (Connochaetes taurinus), zebra (Equus burchelli), and Thomson's gazelle (Gazella thomsoni), these processes have also been suggested to regulate the populations. Censuses of these populations over 20 years have shown changes that allow a test of which, if either, process regulates population numbers. Wildebeest numbers have levelled off as a result of intraspecific competition for food following a five-fold increase due to release from disease and an increase in food supply. Zebra numbers have remained stationary throughout the same period. Gazelle numbers have declined in the last 10 years. These results are contrary to the facilitation hypothesis, which predicts that wildebeest numbers should not have increased if those of zebra did not, and that gazelle numbers should have increased since the wildebeest population increased. The gazelle results are consistent with the interspecific competition hypothesis, but the zebra results are contrary to it. We propose an alternative hypothesis that predation regulates the zebra population, and we suggest ways of testing this.     

Does understory vegetation reflect the history of fluvial disturbance in a riparian forest?

We have examined the relationship between the history of fluvial disturbance and understory vegetation in a riparian forest. The study site was divided into three sites, by use of aerial photographs and topographical maps, with different histories of fluvial disturbance: (1) Fagus-type on land that has not been flooded for the last 39 years, at least; (2) Populus-type on land that has not been flooded since debris flow occurred 34 years ago; and (3) Salix-type on land that has been flooded periodically from an abandoned channel since debris flow occurred 34 years ago. Species richness in the Salix-type was significantly higher than in the other types. Detrended correspondence analysis revealed obvious floristic differences among the three canopy types. Canonical correspondence analysis showed that herbaceous species were mainly found on lower plots with high moss cover, implying that moss layers may capture seeds transported by the stream. Tall herbs occurred in less shaded plots and on higher plots, suggesting that their rapid growth prevented the occurrence of other species. Fagus-type was dominated by species with ingested fruits which depended on animals for their dispersal. Populus and Salix-types were dominated by species with wind dispersal or no dispersal mechanism, which depended on physical phenomena for dispersal. Attributes of current understory vegetation were connected with historical events, suggesting that riparian vegetation reflects the history of fluvial disturbance.     

Does the spatial variation in selective pressures explain among-site differences in seed mass? A test with Buxus balearica

Positive effects of seed size on germination and survival can be offset by a greater probability of predation or a poorer dispersal of larger seeds. We hypothesized that spatial variation in local selective pressures acting on seed mass may lead to differences in both optimal and observed seed mass among discrete populations. We first examined the variation in seed mass across a total of 14 populations of Buxus balearica; in six of such populations, we further examined the temporal variation by measuring seed mass during three consecutive years. Second, we evaluated seed mass effects on different recruitment phases (seed rain to seedling establishment) in three populations. Lastly, for these three populations, we estimated the resulting overall phenotypic selection on seed mass during recruitment and compared the observed and the predicted optimal seed masses. Most variation (c. 70%) in seed mass occurred among populations, and although we found inter-annual variation in seed mass, the differences in seed mass among populations were consistent over time. Conflicting selective pressures on seed mass appeared during recruitment, and their direction and strength varied among populations, depending on the relative local importance of seed predation vs. germination and establishment. Observed seed mass matched predicted optimal seed mass in two of the three examined populations, suggesting local adaptative responses to the spatial mosaic of selective pressures.     

Does morphological convergence imply functional similarity? A test using the evolution of quadrupedalism in ornithischian dinosaurs

Convergent morphologies are thought to indicate functional similarity, arising because of a limited number of evolutionary or developmental pathways. Extant taxa displaying convergent morphologies are used as analogues to assess function in extinct taxa with similar characteristics. However, functional studies of extant taxa have shown that functional similarity can arise from differing morphologies, calling into question the paradigm that form and function are closely related. We test the hypothesis that convergent skeletal morphology indicates functional similarity in the fossil record using ornithischian dinosaurs. The rare transition from bipedality to quadrupedality occurred at least three times independently in this clade, resulting in a suite of convergent osteological characteristics. We use homology rather than analogy to provide an independent line of evidence about function, reconstructing soft tissues using the extant phylogenetic bracket and applying biomechanical concepts to produce qualitative assessments of muscle leverage. We also optimize character changes to investigate the sequence of character acquisition. Different lineages of quadrupedal ornithischian dinosaur stood and walked differently from each other, falsifying the hypothesis that osteological convergence indicates functional similarity. The acquisition of features correlated with quadrupedalism generally occurs in the same order in each clade, suggesting underlying developmental mechanisms that act as evolutionary constraints.     

Does plasticity enhance or dampen phenotypic parallelism? A test with three lake–stream stickleback pairs

Parallel (and convergent) phenotypic variation is most often studied in the wild, where it is difficult to disentangle genetic vs. environmentally induced effects. As a result, the potential contributions of phenotypic plasticity to parallelism (and nonparallelism) are rarely evaluated in a formal sense. Phenotypic parallelism could be enhanced by plasticity that causes stronger parallelism across populations in the wild than would be expected from genetic differences alone. Phenotypic parallelism could be dampened if site‐specific plasticity induced differences between otherwise genetically parallel populations. We used a common‐garden study of three independent lake–stream stickleback population pairs to evaluate the extent to which adaptive divergence has a genetic or plastic basis, and to investigate the enhancing vs. dampening effects of plasticity on phenotypic parallelism. We found that lake–stream differences in most traits had a genetic basis, but that several traits also showed contributions from plasticity. Moreover, plasticity was much more prevalent in one watershed than in the other two. In most cases, plasticity enhanced phenotypic parallelism, whereas in a few cases, plasticity had a dampening effect. Genetic and plastic contributions to divergence seem to play a complimentary, likely adaptive, role in phenotypic parallelism of lake–stream stickleback. These findings highlight the value of formally comparing wild‐caught and laboratory‐reared individuals in the study of phenotypic parallelism.     

Does clonal integration improve competitive ability? A test using aspen (Populus tremuloides [Salicaceae]) invasion into prairie

Many clonal plants consist of many connected individual ramets, allowing them to share water and nutrients via physiological integration. Integration among ramets may also improve the ability of clonal plants to tolerate abiotic stress or improve the competitive ability of individual ramets. Here I use a field experiment to determine whether clonal integration improves ramet performance for a widespread clonal tree species invading into native prairie. Aspen (Populus tremuloides) dominates the southern treeline in western Canada, has long-lived belowground connections between mother and daughter ramets, and reproduces vegetatively via resprouting rhizomes after disturbance. I applied two competition treatments (neighbors present or absent) and two clonal integration treatments (belowground rhizomes between mother and daughter ramets either severed or left intact) to 12 replicate Populus daughter ramets at each of three sites. Neighbors improved the survivorship of Populus ramets by 25-35% after 2 yr, but decreased growth by ～20%. Clonal integration tended to improve ramet survival and growth, but these trends were often not significant. Clonal integration did not alter the effects of competition from neighboring vegetation, suggesting that connections between ramets do not necessarily improve the competitive ability of Populus invading into native prairie.