A species well travelled – the Dodonaea viscosa (Sapindaceae) complex based on phylogenetic analyses of nuclear ribosomal ITS and ETSf sequences

Aim To determine the evolutionary history of the widely studied, cosmopolitan polymorphic species Dodonaea viscosa (hop bush, varnish tree). Location All continents except Antarctica, extending from 44° S (in the South Island of New Zealand) to 33° N (in California and Arizona). Methods For 50 samples across the worldwide distribution, Bayesian analyses of nuclear ribosomal transcribed spacers (ITS 1, ITS 2 and partial ETS) were performed. The alignment was partitioned by secondary structure and analysed using separate models of sequence evolution for each spacer’s stem and loop partition. Bayesian relaxed‐clock estimations of divergence times were used to investigate the tempo of the transoceanic dispersal history of Dodonaea viscosa. Results The widely distributed Dodonaea viscosa evolved in Australia from its most recent common ancestor in the Late Pliocene to Early Pleistocene and subsequently split into two distinct, geographically based, intraspecific lineages. There are a number of regional subclades that are also supported by specific molecular elements. Main conclusions Dodonaea viscosa is not an old lineage, as has often been speculated based primarily on its vast distribution. It diverged from its most recent common ancestor and subsequently dispersed and established around the world within the last 2 Myr. Two distinct lineages within Dodonaea viscosa, which have been shaped by Quaternary climatic change, have separate dispersal histories. The molecular investigations have identified that, although there are least two evolutionary lineages within the complex, they do not correlate with any distinct morphological subunits. It is proposed that Dodonaea viscosa (including D. biloba and D. procumbens) be recognized as an ochlospecies.     

Within‐species patterns challenge our understanding of the leaf economics spectrum

The utility of plant functional traits for predictive ecology relies on our ability to interpret trait variation across multiple taxonomic and ecological scales. Using extensive data sets of trait variation within species, across species and across communities, we analysed whether and at what scales leaf economics spectrum (LES) traits show predicted trait–trait covariation. We found that most variation in LES traits is often, but not universally, at high taxonomic levels (between families or genera in a family). However, we found that trait covariation shows distinct taxonomic scale dependence, with some trait correlations showing opposite signs within vs. across species. LES traits responded independently to environmental gradients within species, with few shared environmental responses across traits or across scales. We conclude that, at small taxonomic scales, plasticity may obscure or reverse the broad evolutionary linkages between leaf traits, meaning that variation in LES traits cannot always be interpreted as differences in resource use strategy.     

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta‐analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole‐plant (e.g. plant height) vs. organ‐level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait‐based community and ecosystem studies.     

Intraspecific trait variation influences physiological performance and fitness in the South Africa shrub genus Protea (Proteaceae)

Background and AimsGlobal plant trait datasets commonly identify trait relationships that are interpreted to reflect fundamental trade-offs associated with plant strategies, but often these trait relationships are not identified when evaluating them at smaller taxonomic and spatial scales. In this study we evaluate trait relationships measured on individual plants for five widespread Protea species in South Africa to determine whether broad-scale patterns of structural trait (e.g. leaf area) and physiological trait (e.g. photosynthetic rates) relationships can be detected within natural populations, and if these traits are themselves related to plant fitness.MethodsWe evaluated the variance structure (i.e. the proportional intraspecific trait variation relative to among-species variation) for nine structural traits and six physiological traits measured in wild populations. We used a multivariate path model to evaluate the relationships between structural traits and physiological traits, and the relationship between these traits and plant size and reproductive effort.Key ResultsWhile intraspecific trait variation is relatively low for structural traits, it accounts for between 50 and 100 % of the variation in physiological traits. Furthermore, we identified few trait associations between any one structural trait and physiological trait, but multivariate regressions revealed clear associations between combinations of structural traits and physiological performance (R² = 0.37–0.64), and almost all traits had detectable associations with plant fitness.ConclusionsIntraspecific variation in structural traits leads to predictable differences in individual-level physiological performance in a multivariate framework, even though the relationship of any particular structural trait to physiological performance may be weak or undetectable. Furthermore, intraspecific variation in both structural and physiological traits leads to differences in plant size and fitness. These results demonstrate the importance of considering measurements of multivariate phenotypes on individual plants when evaluating trait relationships and how trait variation influences predictions of ecological and evolutionary outcomes.     

Leaf trait variation on Erythroxylum tortuosum (Erythroxylaceae) and its relationship with oviposition preference and stress by a host‐specific leaf miner

It has been suggested that plant physical and chemical traits vary considerably in space and time. Hence, leaf‐mining insects may adjust their oviposition in response to leaf attributes representing high quality. Moreover, herbivorous insects can modify leaf morphology by acting as stressors, increasing, for example, fluctuating asymmetry (FA) levels. Here, we investigate oviposition preference in Agnippe sp.2, a leaf‐mining moth of Erythroxylum tortuosum, in relation to differences in leaf nutritional quality (i.e. levels of water, nitrogen and tannin content), leaf area (i.e. quantity of resource hypothesis) and FA. We also verify whether temporal variation in plant nutritional quality emerges as an alternative hypothesis to explain oviposition distribution in time, and whether this leaf miner is a stress‐causing agent, increasing FA during larval development. Mined leaves and leaves with and without eggs were periodically collected from plants located in a Cerrado fragment in Brazil. In the laboratory, leaf traits were assessed (using image analysis software) and quantified (biochemical analysis) according to the aims previously determined. Oviposition probability did not change in relation to variations in nitrogen, tannins and FA of leaves. However, leaf‐miner females preferred to oviposit on leaves having large areas and low water contents. It was also verified that new leaves of E. tortuosum, which carried most leaf‐miner eggs, presented significantly lower tannins and greater levels of nitrogen and water than old leaves. The oviposition choice exhibited by leaf miners was found to be non‐random because they appear to use resource quantity and water content as cues as where to lay their eggs. The temporal variation of plant nutritional quality is likely to influence the time of leaf‐miner oviposition; and leaf FA was not increased during larval feeding, suggesting that these herbivores do not cause variations in FA levels.     

A tropical epiphytic orchid uses a low‐light interception strategy in a spatially heterogeneous light environment

Light is considered a non‐limiting factor for vascular epiphytes. Nevertheless, an epiphyte's access to light may be limited by phorophyte shading and the spatio‐temporal environmental patchiness characteristic of epiphytic habitats. We assessed the extent to which potential light interception in Rodriguezia granadensis, an epiphytic orchid, is determined by individual factors (plant size traits and leaf traits), or environmental heterogeneity (light patchiness) within the crown of the phorophyte, or both. We studied 104 adult plants growing on Psidium guajava trees in two habitats with contrasting canopy cover: a dry tropical forest edge, and isolated trees in a pasture. We recorded the number of leaves and the leaf area, the leaf position angles, and the potential exposure of the leaf surface to direct irradiance (silhouette area of the leaf blade), and the potential irradiance incident on each plant. We found the epiphytes experience a highly heterogeneous light environment in the crowns of P. guajava. Nonetheless, R. granadensis plants displayed a common light interception strategy typical of low‐light environments, resembling terrestrial, forest understory plants. Potential exposure of the total leaf surface to direct irradiance correlated positively with plant size and within‐plant variation in leaf orientation. In many‐leaved individuals, within‐plant variation in leaf angles produced complementary leaf positions that enhanced potential light interception. This light interception strategy suggests that, in contrast to current wisdom, enhancing light capture is important for vascular epiphytes in canopies with high spatio‐temporal heterogeneity in light environments.     

Climate‐related genetic variation in drought‐resistance of Douglas‐fir (Pseudotsuga menziesii)

There is a general assumption that intraspecific populations originating from relatively arid climates will be better adapted to cope with the expected increase in drought from climate change. For ecologically and economically important species, more comprehensive, genecological studies that utilize large distributions of populations and direct measures of traits associated with drought‐resistance are needed to empirically support this assumption because of the implications for the natural or assisted regeneration of species. We conducted a space‐for‐time substitution, common garden experiment with 35 populations of coast Douglas‐fir (Pseudotsuga menziesii var. menziesii) growing at three test sites with distinct summer temperature and precipitation (referred to as ‘cool/moist’, ‘moderate’, or ‘warm/dry’) to test the hypotheses that (i) there is large genetic variation among populations and regions in traits associated with drought‐resistance, (ii) the patterns of genetic variation are related to the native source‐climate of each population, in particular with summer temperature and precipitation, (iii) the differences among populations and relationships with climate are stronger at the warm/dry test site owing to greater expression of drought‐resistance traits (i.e., a genotype × environment interaction). During midsummer 2012, we measured the rate of water loss after stomatal closure (transpiration_min), water deficit (% below turgid saturation), and specific leaf area (SLA, cm² g^?1) on new growth of sapling branches. There was significant genetic variation in all plant traits, with populations originating from warmer and drier climates having greater drought‐resistance (i.e., lower transpiration_min, water deficit and SLA), but these trends were most clearly expressed only at the warm/dry test site. Contrary to expectations, populations from cooler climates also had greater drought‐resistance across all test sites. Multiple regression analysis indicated that Douglas‐fir populations from regions with relatively cool winters and arid summers may be most adapted to cope with drought conditions that are expected in the future.     

Revision of the genus Acrochordiceras Hyatt, 1877 (Ammonoidea,Middle Triassic): morphology,biometry, biostratigraphy and intra‐specific variability

Abstract: The family Acrochordiceratidae Arthaber, 1911 ranges in age from latest Spathian to the middle/late Anisian boundary, and it represents a major component of ammonoid faunas during that time. The middle Anisian genus Acrochordiceras Hyatt, 1877 is the most widespread taxon of the family and occurs abundantly worldwide within the low paleolatitude belt. However, there is a profusion of species names available for Acrochordiceras. This excessive diversity at the species level essentially results from the fact that sufficiently large samples were not available, thus leading to a typological approach to its taxonomy. Based on new extensive collections obtained from the Anisian (Middle Triassic) Fossil Hill Member (Star Peak Group, north‐west Nevada) for which a high resolution biostratigraphic frame is available, the taxonomy and biostratigraphy of the genus Acrochordiceras Hyatt, 1877 is herein revised with respect to its intra‐specific variation. Morphological and biometric studies (c. 550 bedrock‐controlled specimens were measured) show that only one species occurs in each stratigraphic level. Continuous ranges of intra‐specific variation of studied specimens enable us to synonymize Haydenites Diener, 1907, Silesiacrochordiceras Diener, 1916 and Epacrochordiceras Spath, 1934 with Acrochordiceras Hyatt, 1877. Three stratigraphically successive species are herein recognized in the low paleolatitude middle Anisian faunas from Nevada: A. hatschekii (Diener, 1907), A. hyatti Meek, 1877 and A. carolinae Mojsisovics, 1882. Moreover, an assessment of intra‐specific variation of the adult size range does not support recognition of a dimorphic pair (Acrochordiceras and Epacrochordiceras) as previously suggested by other workers (Epacrochordiceras is the compressed and weakly ornamented end‐member variant of Acrochordiceras). The successive middle Anisian species of Acrochordiceras form an anagenetic lineage characterized by increasing involution, adult size and intra‐specific variation. This taxonomic revision based on new bedrock‐controlled collections is thus an important prerequisite before studying the evolution of the group.     

Floral color variation and associations with fitness‐related traits in Malva moschata (Malvaceae)

Genetic polymorphisms for floral color are interesting phenomena to study because they are likely to be maintained by opposing selective forces. Pollinator preferences may exert direct selection on floral color; however, floral color might also be the indirect target of selection through genetic associations with other traits under selection. Malva moschata (Malvaceae) is a North American species that produces either red or white flowers. In the present study, we present reflectance spectrophotometry data that characterize the nature of floral color variation in this species and show that honey bees and bumble bees should be able to distinguish between the morphs through differential sensitivity at the green (long‐wavelength) photoreceptor. Second, we use a series of phenotypic measures to investigate whether the color morphs differ with respect to other floral traits, vegetative traits or female reproductive success, and use a series of correlation analyses to infer the relative independence of color from these other traits. We found that red‐flowered morphs produced more anthers per flower and had greater leaf area, and that white‐flowered morphs had greater percentage fruit set; however, there were no reproductive success differences between the morphs. The relationship between flower size and anther number was the only correlation that differed between the morphs. Finally, a series of pollinator‐choice experiments showed that bumble bees strongly prefer red morphs in terms of visit frequency and duration, but honey bees have no preference. Taken together, our results suggest that color is rather independent of other phenotypic traits, and that honey bee abundance is likely to play a role in maintaining color variation in this system.     

Plant intraspecific functional trait variation is related to within‐habitat heterogeneity and genetic diversity in Trifolium montanum L.

Intraspecific trait variation (ITV), based on available genetic diversity, is one of the major means plant populations can respond to environmental variability. The study of functional trait variation and diversity has become popular in ecological research, for example, as a proxy for plant performance influencing fitness. Up to now, it is unclear which aspects of intraspecific functional trait variation (iFD_CV) can be attributed to the environment or genetics under natural conditions. Here, we examined 260 individuals from 13 locations of the rare (semi‐)dry calcareous grassland species Trifolium montanum L. in terms of iFD_CV, within‐habitat heterogeneity, and genetic diversity. The iFD_CV was assessed by measuring functional traits (releasing height, biomass, leaf area, specific leaf area, leaf dry matter content, F_v/F_m, performance index, stomatal pore surface, and stomatal pore area index). Abiotic within‐habitat heterogeneity was derived from altitude, slope exposure, slope, leaf area index, soil depth, and further soil factors. Based on microsatellites, we calculated expected heterozygosity (H_e) because it best‐explained, among other indices, iFD_CV. We performed multiple linear regression models quantifying relationships among iFD_CV, abiotic within‐habitat heterogeneity and genetic diversity, and also between separate functional traits and abiotic within‐habitat heterogeneity or genetic diversity. We found that abiotic within‐habitat heterogeneity influenced iFD_CV twice as strong compared to genetic diversity. Both aspects together explained 77% of variation in iFD_CV ( = .77, F_{2, 10} = 21.66, p < .001). The majority of functional traits (releasing height, biomass, specific leaf area, leaf dry matter content, F_v/F_m, and performance index) were related to abiotic habitat conditions indicating responses to environmental heterogeneity. In contrast, only morphology‐related functional traits (releasing height, biomass, and leaf area) were related to genetics. Our results suggest that both within‐habitat heterogeneity and genetic diversity affect iFD_CV and are thus crucial to consider when aiming to understand or predict changes of plant species performance under changing environmental conditions.     

The metabolic enzyme phosphoglucose isomerase (Pgi) affects the outcome of intra‐specific competition in a polyembryonic wasp

1. Polyembryonic parasitoid wasps in the family Encyrtidae (Hymenoptera) have evolved a caste system consisting of morphologically and functionally distinct larvae called soldiers and reproductives. 2. Two selective pressures are thought to underlie the evolution of the soldier caste: defence against competitors and resolution of the sex ratio conflict. Previous studies also indicate that soldier development time strongly affects the outcome of intra‐specific competition in the polyembryonic encyrtid Copidosoma floridanum Ashmead. This study builds on prior findings by showing that alleles of the metabolic enzyme phosphoglucose isomerase (Pgi) differentially affect soldier development time and the outcome of competition. 3. Soldier larvae with the Pgi alleles 100 or 120 emerged on average 65 h post‐parasitism, whereas soldier larvae with a third allele, 54, emerged at 67 h. In turn, C. floridanum broods homozygous for the 100 and 120 alleles outcompete broods homozygous for the 54 allele. 4. Pgi allelic diversity may be maintained through a life‐history trade‐off affecting female brood sizes with homozygous broods bearing the developmentally disadvantageous 54 allele producing more adult females than broods bearing alternate common alleles.     

Countergradient vs. cogradient variation in growth and diapause in a lichen‐feeding moth,Eilema depressum (Lepidoptera: Arctiidae)

Separating genetic and environmental causes of the latitudinal differences among populations is crucial when evaluating the potential for microevolutionary responses to the changing environment. We studied among‐population and environmental components of variation in several life‐history traits of a lichen‐feeding moth Eilema depressum when offspring of replicate Swiss and Finnish females were reared in a common‐garden factorial experiment. A partial second generation was produced only among Swiss larvae, more likely so at higher temperature regime and higher host quality, and more frequently among the offspring of particular females. Growth rates of larvae that chose the diapause development were higher in northern individuals. Our results thus reveal adaptive differences between latitudinal populations in studied life‐history traits, allowing to expect rapid adaptation of the species to further environmental changes. In contrast, invariable responses of the growth rates of the larvae to temperature and host quality support the idea that some basic parameters of insect growth show a high degree of evolutionary conservatism.     

Morphometric variation of Seminavis pusilla (Bacillariophyceae) and its relationship to salinity in inter‐dune lakes of the Badain Jaran Desert,Inner Mongolia,China

We used light and scanning electron microscope analyses to quantify morphometric features (valve length, width, stria density, lineola density and valve curvature) from the observation of valves representing Seminavis pusilla. Cluster analysis based on Gaussian mixture models and the expectation‐maximization algorithm was used for delineating two species, Seminavis pusilla sensu stricto and Seminavis lata (Krammer) Rioual comb. et stat. nov. By comparison with S. pusilla, S. lata is characterized by wider valves and lower stria density. The two species have also markedly different ecology. S. pusilla is most abundant in the most saline lakes of the dataset, while S. lata is most abundant in the less saline lakes. Our results indicate that combining the two species into S. pusilla sensu lato would lead to a loss of ecological information and a decrease of the performance of transfer functions developed for quantitative reconstruction of past salinity from fossil diatom assemblages in sediment cores.