Natural variation in Pristionchus pacificus dauer formation reveals cross-preference rather than self-preference of nematode dauer pheromones

Many free-living nematodes, including the laboratory model organisms Caenorhabditis elegans and Pristionchus pacificus, have a choice between direct and indirect development, representing an important case of phenotypic plasticity. Under harsh environmental conditions, these nematodes form dauer larvae, which arrest development, show high resistance to environmental stress and constitute a dispersal stage. Pristionchus pacificus occurs in a strong association with scarab beetles in the wild and remains in the dauer stage on the living beetle. Here, we explored the circumstances under which P. pacificus enters and exits the dauer stage by using a natural variation approach. The analysis of survival, recovery and fitness after dauer exit of eight P. pacificus strains revealed that dauer larvae can survive for up to 1 year under experimental conditions. In a second experiment, we isolated dauer pheromones from 16 P. pacificus strains, and tested for natural variation in pheromone production and sensitivity in cross-reactivity assays. Surprisingly, 13 of the 16 strains produce a pheromone that induces the highest dauer formation in individuals of other genotypes. These results argue against a simple adaptation model for natural variation in dauer formation and suggest that strains may have evolved to induce dauer formation precociously in other strains in order to reduce the fitness of these strains. We therefore discuss intraspecific competition among genotypes as a previously unconsidered aspect of dauer formation.     

Unraveling the evolutionary history of the nematode Pristionchus pacificus: from lineage diversification to island colonization

The hermaphroditic nematode Pristionchus pacificus is a model organism with a range of fully developed genetic tools. The species is globally widespread and highly diverse genetically, consisting of four major independent lineages (lineages A, B, C, and D). Despite its young age (~2.1 Ma), volcanic La Réunion Island harbors all four lineages. Ecological and population genetic research studies suggest that this diversity is due to repeated independent island colonizations by P. pacificus. Here, we use model‐based statistical methods to rigorously test hypotheses regarding the evolutionary history of P. pacificus. First, we employ divergence analyses to date diversification events among the four “world” lineages. Next, we examine demographic properties of a subset of four populations (“a”, “b”, “c”, and “d”), present on La Réunion Island. Finally, we use the results of the divergence and demographic analyses to inform a modeling‐based approximate Bayesian computation (ABC) approach, where we test hypotheses about the order and timing of establishment of the Réunion populations. Our dating estimates place the recent common ancestor of P. pacificus lineages at nearly 500,000 generations past. Our demographic analysis supports recent (<150,000 generations) spatial expansion for the island populations, and our ABC approach supports c>a>b>d as the most likely colonization order of the island populations. Collectively, our study comprehensively improves previous inferences about the evolutionary history of P. pacificus.     

Variation in Gender and Flower‐Size Dimorphism in the Dioecious Tree Dombeya ciliata,an Endemic to La Reunion Island1

The genus Dombeya is an important component of tropical forests in west Africa, Madagascar, and the Mascarene islands, but little is known about the reproductive biology of its members. In this study, we quantified gender variation and flower‐size variation in three populations of the dioecious tree Dombeya ciliata, an endemic of La Reunion Island (Indian Ocean). Variation among the three populations was observed for three characters. First, the high elevation population had strict males and females, whereas in the two low elevation populations, males were inconstant, with > 50 percent of the males set a small number of fruit. Second, females showed greater seed production at high elevation. Third, flower‐size dimorphism was greater at high elevation because male flowers were larger than female flowers.     

Natural genetic variation in Arabidopsis: tools, traits and prospects for evolutionary ecology

BACKGROUND: The model plant Arabidopsis thaliana (Arabidopsis) shows a wide range of genetic and trait variation among wild accessions. Because of its unparalleled biological and genomic resources, the potential of Arabidopsis for molecular genetic analysis of this natural variation has increased dramatically in recent years. SCOPE: Advanced genomics has accelerated molecular phylogenetic analysis and gene identification by quantitative trait loci (QTL) mapping and/or association mapping in Arabidopsis. In particular, QTL mapping utilizing natural accessions is now becoming a major strategy of gene isolation, offering an alternative to artificial mutant lines. Furthermore, the genomic information is used by researchers to uncover the signature of natural selection acting on the genes that contribute to phenotypic variation. The evolutionary significance of such genes has been evaluated in traits such as disease resistance and flowering time. However, although molecular hallmarks of selection have been found for the genes in question, a corresponding ecological scenario of adaptive evolution has been difficult to prove. Ecological strategies, including reciprocal transplant experiments and competition experiments, and utilizing near-isogenic lines of alleles of interest will be a powerful tool to measure the relative fitness of phenotypic and/or allelic variants. CONCLUSIONS: As the plant model organism, Arabidopsis provides a wealth of molecular background information for evolutionary genetics. Because genetic diversity between and within Arabidopsis populations is much higher than anticipated, combining this background information with ecological approaches might well establish Arabidopsis as a model organism for plant evolutionary ecology.     

The role of selection and historical factors in driving population differentiation along an elevational gradient in an island bird

Adaptation to local environmental conditions and the range dynamics of populations can influence evolutionary divergence along environmental gradients. Thus, it is important to investigate patterns of both phenotypic and genetic variations among populations to reveal the respective roles of these two types of factors in driving population differentiation. Here, we test for evidence of phenotypic and genetic structure across populations of a passerine bird (Zosterops borbonicus) distributed along a steep elevational gradient on the island of Réunion. Using 11 microsatellite loci screened in 401 individuals from 18 localities distributed along the gradient, we found that genetic differentiation occurred at two spatial levels: (i) between two main population groups corresponding to highland and lowland areas, respectively, and (ii) within each of these two groups. In contrast, several morphological traits varied gradually along the gradient. Comparison of neutral genetic differentiation (F_ST) and phenotypic differentiation (P_ST) showed that P_ST largely exceeds F_ST at several morphological traits, which is consistent with a role for local adaptation in driving morphological divergence along the gradient. Overall, our results revealed an area of secondary contact midway up the gradient between two major, cryptic, population groups likely diverged in allopatry. Remarkably, local adaptation has shaped phenotypic differentiation irrespective of population history, resulting in different patterns of variation along the elevational gradient. Our findings underscore the importance of understanding both historical and selective factors when trying to explain variation along environmental gradients.     

Extremely reduced dispersal and gene flow in an island bird

The Réunion grey white-eye, Zosterops borbonicus, a passerine bird endemic to Réunion Island in the Mascarene archipelago, represents an extreme case of microgeographical plumage colour variation in birds, with four distinct colour forms occupying different parts of this small island (2512 km²). To understand whether such population differentiation may reflect low levels of dispersal and gene flow at a very small spatial scale, we examined population structure and gene flow by analysing variation at 11 microsatellite loci among four geographically close localities (<26 km apart) sampled within the distribution range of one of the colour forms, the brown-headed brown form. Our results revealed levels of genetic differentiation that are exceptionally high for birds at such a small spatial scale. This strong population structure appears to reflect low levels of historical and contemporary gene flow among populations, unless very close geographically (<10 km). Thus, we suggest that the Réunion grey white-eye shows an extremely reduced propensity to disperse, which is likely to be related to behavioural processes.     

Naïveté in novel ecological interactions: lessons from theory and experimental evidence

The invasion of alien species into areas beyond their native ranges is having profound effects on ecosystems around the world. In particular, novel alien predators are causing rapid extinctions or declines in many native prey species, and these impacts are generally attributed to ecological naïveté or the failure to recognise a novel enemy and respond appropriately due to a lack of experience. Despite a large body of research concerning the recognition of alien predation risk by native prey, the literature lacks an extensive review of naïveté theory that specifically asks how naïveté between novel pairings of alien predators and native prey disrupts our classical understanding of predator–prey ecological theory. Here we critically review both classic and current theory relating to predator–prey interactions between both predators and prey with shared evolutionary histories, and those that are ecologically ‘mismatched’ through the outcomes of biological invasions. The review is structured around the multiple levels of naïveté framework of Banks & Dickman (2007), and concepts and examples are discussed as they relate to each stage in the process from failure to recognise a novel predator (Level 1 naïveté), through to appropriate (Level 2) and effective (Level 3) antipredator responses. We discuss the relative contributions of recognition, cue types and the implied risk of cues used by novel alien and familiar native predators, to the probability that prey will recognise a novel predator. We then cover the antipredator response types available to prey and the factors that predict whether these responses will be appropriate or effective against novel alien and familiar native predators. In general, the level of naïveté of native prey can be predicted by the degree of novelty (in terms of appearance, behaviour or habitat use) of the alien predator compared to native predators with which prey are experienced. Appearance in this sense includes cue types, spatial distribution and implied risk of cues, whilst behaviour and habitat use include hunting modes and the habitat domain of the predator. Finally, we discuss whether the antipredator response can occur without recognition per se, for example in the case of morphological defences, and then consider a potential extension of the multiple levels of naïveté framework. The review concludes with recommendations for the design and execution of naïveté experiments incorporating the key concepts and issues covered here. This review aims to critique and combine classic ideas about predator–prey interactions with current naïveté theory, to further develop the multiple levels of naïveté framework, and to suggest the most fruitful avenues for future research.     

Riverine wetland vegetation: importance of small-scale and large-scale environmental variation

Abstract. The purpose of this study was to evaluate the relative importance of small-scale variation in abiotic factors and large-scale spatio-temporal variation on the distribution of wetland vegetation of a section of the Upper St. Lawrence River in Québec. Vegetation data have been classified with agglomerative clustering into 11 community types, from Acer rubrum, Acer saccharinum and Fraxinus swamps, to scrubs dominated by Salix petiolaris, Alnus rugosa var. americana or Myrica gale, to Typha, Typha/Lythrum, Carex lacustris and Calamagrostis canadensis marshes. Canonical Correspondence Analysis (CANOCO) suggested that peat thickness and water level are the most important abiotic variables correlated with plant community composition. As a whole, small-scale variation accounts for 25.6 % of the species variation. Another 21.6% is explained by large-scale variation based on location data summarizing large-scale spatial distribution, and historical landscape dynamics differentiated into (a) no net loss of wetlands, (b) net loss of wetland, and (c) changes within wetlands, which are correlated with the actual variation in herbaceous and scrubby vegetation. The interaction between small-scale and large-scale variations explains another 1.7 %. In total, 48.9% of the species variation is explained by the two data sets, leaving 51.1% unexplained. Whereas the omission of some abiotic variables is possible, it is hypothesized that the abiotic conditions measured in this study play an important role, especially in the distribution of forested swamps. In addition, past history, particularly that of human interventions, becomes another important factor leading to the observed importance of large-scale spatio-temporal variables. This is particularly true for Alnus rugosa var. americana shrublands. Time lag between a relative stabilization of species distribution and the reduction of natural disturbances (water level fluctuations and fires) could be a possible cause of the importance of spatio-temporal variables and the undetermined portion of species variation.     

Extreme,continuous variation in an island snail: local diversification and association of shell form with the current environment

On Rosemary Island, a small continental island (11 km²) in the Dampier Archipelago, Western Australia, snails of the genus Rhagada have extremely diverse morphologies. Their shells vary remarkably in size and shape, with the latter ranging from globose to keeled‐flat, spanning the range of variation in the entire genus. Based primarily on variation in shell morphology, five distinct species are currently recognized. However, a study of 103 populations has revealed continuity of shell form within a very closely‐related group. A phylogenetic analysis of specimens from Rosemary Island, and other islands in the Dampier Archipelago, indicates that much of the morphological variation has evolved on the island, from within a monophyletic group. Within the island, snails with distinct shell morphologies could not be distinguished based on variation in mitochondrial DNA or their reproductive anatomy. The shell variation is geographically structured over a very fine scale, with clines linking the extreme forms over distances less than 200 m. Although there is no evidence that the different forms have evolved in isolation or as a consequence of drift, there is a strong association between keeled‐flat shells and rocky habitats, suggesting that shell shape may be of adaptive significance. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 756–769.     

Seed dispersal by lizards on a continental‐shelf island: predicting interspecific variation in seed rain based on plant distribution and lizard movement patterns

Aim We estimated the patterns of seed deposition provided by the eyed lizard, Timon lepidus, and evaluated whether these patterns can be generalized across plant species with different traits (fruit and seed size) and spatial distributions. Location Monteagudo Island, Atlantic Islands National Park (north‐western Spain). Methods We radio‐tracked seven lizards for 14 days and estimated their home ranges using fixed kernels. We also geo‐referenced all fruit‐bearing individuals of four plant species dispersed by eyed lizards in the study area (Corema album, Osyris alba, Rubus ulmifolius and Tamus communis), measured the passage time of their seeds through the lizard gut, and estimated seed predation in four habitats (bare sand, grassland, shrub and gorse). Seed dispersal kernels were estimated using a combination of these data and were combined with seed predation probability maps to incorporate post‐dispersal seed fate (‘seed survival kernels’). Results Median seed gut‐passage times were around 52–98 h, with maximum values up to 250 h. Lizards achieved maximum displacement in their home ranges within 24–48 h. Seed predation was high (80–100% of seeds in 2 months), particularly under Corema shrub and gorse. Seed dispersal kernels showed a common pattern, with two areas of preferential seed deposition, but the importance of these varied among plant species. Interspecific differences among dispersal kernels were strongly reduced by post‐dispersal seed predation; hence, seed survival kernels of the different plant species showed high auto‐ and pairwise‐correlations at small distances (< 50 m). As a result, survival to post‐dispersal seed predation increased with dispersal distance for O. alba and T. communis, but not for C. album. Main conclusions Seed dispersal by lizards was determined primarily by the interaction between the dispersers’ home ranges and the position of the fruit‐bearing plants. As a result, seed rain shared a common template, but showed considerable variation among species, determined by their specific spatial context. Seed predation increased the spatial coherence of the seed rain of the different species, but also resulted in contrasting relationships between seed survival and dispersal distance, which may be of importance for the demographic and evolutionary processes of the plants.     

Leaf-shape variation of Paederia foetida in Japan: reexamination of the small, narrow leaf form from Miyajima Island

Variations in Paederia foetida L. leaf shape were examined to evaluate the taxonomic validity of the small, narrow leaf form of P. foetida f. microphylla Honda from Miyajima Island, Honshu, Japan. There is considerable variation in P. foetida individuals in terms of leaf size and leaf index (leaf length:leaf width ratio). On Miyajima Island, some individuals have narrow leaves with a high leaf index value, a phenotype represented by the type specimen of P. foetida f. microphylla, and some do not. Given that the leaf size of individuals from Miyajima Island is smaller than that of individuals from other localities in Japan, and that the small leaf phenotype is stable even under cultivation, P. foetida f. microphylla is classified as the form having the smallest leaf size. Anatomical examination of leaf blades revealed that the large variation in leaf size was attributable to variation in the number of leaf cells but not to differences in cell size or cell shape. Based on these results, we discuss the endemism of P. foetida f. microphylla.     

Methylenedioxy- and methoxyflavones from Melicope coodeana syn. Euodia simplex

Three new natural products, 3,8-dimethoxy-5,7-dihydroxy-3′,4′-methylenedioxyflavone, 3,6,8-trimethoxy-5,7-dihydroxy-3′,4′-methylenedioxyflavone and 3,6,8,3′,4′-pentamethoxy-5,7-dihydroxyflavone were isolated from Melicope coodeana syn. Euodia simplex (Rutaceae) along with 3,6,3′-trimethoxy-5,7,4′-trihydroxyflavone and 3,3′-dimethoxy-5,7,4′-trihydroxyflavone. The structural assignments are based on ¹H and ¹³C NMR data, including discussion of the chemical shifts of C-2 in 3,5-dihydroxy- and 3-methoxy-5-hydroxyflavones. The presence of highly methoxylated and methylenedioxyflavones is characteristic of the genus Melicope, and the present findings support the recent transfer of Euodia simplex to Melicope.     

The Ibis: Transformations in a Twentieth Century British Natural History Journal

The contents of the British Ornithologists' Union's journal, The Ibis, during the first half of the 20th century illustrates some of the transformations that have taken place in the naturalist tradition. Although later generations of ornithologists described these changes as logical and progressive, their historical narratives had more to do with legitimizing the infiltration of the priorities of evolutionary theory, ecology, and ethology than analyzing the legacy of the naturalist tradition on its own terms. Despite ornithologists' claim that the journal's increasing focus on "biology" represented a natural development after the preliminary phase of systematics and geographical ornithology, in fact a small group campaigned to bring the priorities of population ecology, behavior, and selection theory into the journal and British ornithology more generally. The problems involved in this transition highlight the importance of methodological and institutional context in determining and reinforcing appropriate research programs for ornithologists. Comparing the discipline-building rhetoric of moderns with the contents of the past illustrates how modern evaluations of 19th century research programs have been enmeshed in ornithologists' endeavors to forge new identities for traditional disciplines.     

Stochastic simulation of size variation in turbot: possible causes analysed with an individual-based model

Causes of size variation in a population of juvenile turbot were studied using an individual based model (IBM). Each simulation started with 800 (divided into eight groups of 100 each) 120-day-old (posthatch) juveniles and was run for 140 days, and the data gained from model simulations compared directly with the result of a laboratory study with size-graded turbot. Stochastic growth with memory, which was included in the models as an individual genetical growth rate variation, is important in explaining size variation, and the combination between individual genetic growth rate and social interactions related to size-dependent hierarchies also contributes to size variation. The use of size-dependent growth rate alone fails to explain size variation, and is of little value in predicting size variation in turbot culture. Further, the results indicate formation of different types of size hierarchies for different sizes of juvenile turbot.     

Utility of closely related taxa for genetic studies of adaptive variation and speciation: Current state and perspectives in plants with focus on forest tree species

Studies of adaptation and speciation have greatly benefited from rapid progress of DNA sequencing and genotyping technologies. Comparative genomics of closely related taxa has great potential to advance evolutionary research on genetic architecture of adaptive traits and reproductive isolation. Such studies that utilized closely related plant species and ecotypes have already provided some important insights into genomic regions and/or genes that are potentially involved in local adaptation and speciation. The choice of an appropriate species model for such research is crucial. The paper discusses current approaches used to reveal the patterns of intra‐ and interspecific divergence due to natural selection. Its outcomes in herbaceous plants and forest trees are briefly summarized and compared to reveal general regularities concerning evolutionary processes. We then highlight the importance of multispecies studies and discuss the utility of several related pine taxa as fine candidates for evolutionary inferences. Genetically similar but ecologically and phenotypically diverged taxa seem a promising study system to search for genomic patterns of speciation and adaptive variation.     

Contrasting patterns of morphological variation with dietary preferences in Micropogonias furnieri: insights from stable‐isotope and digestive‐trait analyses

The dietary preferences of populations of whitemouth croaker Micropogonias furnieri, which commonly inhabit estuarine and oceanic environments of the south‐western Atlantic Ocean, were investigated using stable‐isotope analysis and digestive traits, and compared with previous genetic and morphometric surveys of this species. Isotopic and C:N‐derived data suggested that individuals from coastal lagoons are the most differentiated from the remaining localities surveyed. In contrast, the analysis of the digestive traits did not show the same differentiation pattern. The overall correlation between isotopic, molecular and morphological variations suggests that genetic and phenotypic differences among populations are accompanied by differential resource use, supporting the idea that selective forces could be playing an important role in population differentiation.     

Spine development in Brachionus quadridentatus from an Australian billabong: genetic variation and induction by Asplanchna

Fertilized resting eggs of Australian Brachionus quadridentatus hatched 2–3 days after hydration into females with or, more frequently, without posterior lateral spines. These females then produced clones with short-spined or long-spined phenotypes. Asplanchna girodi induced females from two short-spined clones and one long-spined clone to produce daughters with significantly longer posterior lateral spines. In all clones, there were significant differences in spine development among offspring of mothers within Asplanchna and control treatments. The range of phenotypes reported in one short-spined clone is observed in the billabong and includes much of the variation described for the species, with mehleni (long-spined) phenotypes occurring with Asplanchna. In B. quadridentatus, the ecological significance of long-spined, basic phenotypes, and of the spine-development response to Asplanchna, is unclear. In laboratory cultures, females of all clones were attached to the substratum or water surface, and were safe from Asplanchna; in nature, females are epiphytic and probably rarely susceptible to Asplanchna. Most (96%) resting eggs produced in cultures and kept under culture conditions hatched after a 7-day latent period. This raises questions regarding natural conditions which might prevent hatching and allow accumulation of resting eggs in a sediment egg bank. Hatching of resting eggs in nature may be enhanced in sediments which dry and then become flooded after rains.