FLUCTUATING ASYMMETRY IN A SALIX HYBRID SYSTEM: THE IMPORTANCE OF GENETIC VERSUS ENVIRONMENTAL CAUSES

To examine the effects of hybridization and environmental stress on developmental instability, we examined fluctuating asymmetry (FA), the variance in random deviations from perfect symmetry in bilaterally symmetrical traits, for leaf symmetry in a Salix hybrid system. An abiotic environmental stress (water stress), an interspecific biotic stress (pathogen attack), and an intraspecific biotic stress (competition) were examined to determine which factors increase developmental instability. None of these three environmental stressors significantly increased FA. However, genetic stress through hybridization was detected; hybrid plants showed significantly higher levels of FA than parental species. In contrast to hybridization providing greater developmental stability through heterozygosity, these results suggest that complex, nonadditive interactions provided developmental stability and that developmental instability increased when coadapted gene complexes were disrupted through hybridization. In addition, plant biomass was significantly, negatively correlated with FA, suggesting that those individuals that were more able to buffer themselves against the disruptive effects of environmental stress may have a selective advantage over those that are less able to buffer themselves against these disruptive effects.     

Is phenotypic canalization involved in the decline of the endemic Aquilegia thalictrifolia? Rethinking relationships between fluctuating asymmetry and species conservation status

Fluctuating asymmetry (FA), the deviation from the normal symmetrical condition of a morphological trait having specific morphological symmetry, increases in response to environmental and genetic stress, is related to phenotypic plasticity and is considered a tool for monitoring a species conservation status. However, FA–stress relations are dependent on measured traits or species‐specific characteristics such as mating system and habitat. This study investigates the relationships between FA, genetic diversity, population size, density and individual fitness traits (plant height, fruit and seed set), in the endemic Aquilegia thalictrifolia, a mixed breeder that is declining, but maintaining high levels of heterozygosity. Leaf and flower FA and other traits were investigated in 10 populations of A. thalictrifolia, the whole species range. As a result, we found similar patterns of FA in leaves and flowers between populations, indicating a homogenous level of stress between populations that differed for other traits. FA and the other traits were not related, including heterozygosity. Heterozygosity was not related to individual fitness traits with the exception of plant height. In accordance with other studies, we found that the role of FA as a tool for assessing the conservation status of a species or population should be reconsidered. However, we conclude that a low level of FA should not automatically be considered an indicator of good conservation status or low level of stress, because in species that evolved in highly stable environments it may indicate a scarce ability to plastically respond to environmental changes, as a consequence of environmental and genetic canalization. Further investigation of this point is needed.     

Population density, sex ratio, body size and fluctuating asymmetry of Ceroglossus chilensis (Carabidae) in the fragmented Maulino forest and surrounding pine plantations

Habitat fragmentation results in new environmental conditions that may stress resident populations. Such stress may be reflected in demographical or morphological changes in the individuals inhabiting those landscapes. This study evaluates the effects of fragmentation of the Maulino forest on population density, sex ratio, body size, and fluctuating asymmetry (FA) of the endemic carabid Ceroglossus chilensis. Individuals of C. chilensis were collected during 2006 in five locations at Los Queules National Reserve (continuous forest), in five forest fragments and in five areas of surrounding pine plantations (matrix). In each location, once a season, 40 pitfall traps (20 in the centre, 20 in the edge), were opened for 72 h. Population density of C. chilensis was higher in the small fragments than in the pine matrix, with intermediate densities in the continuous forest; sex ratio did not differ significantly from 1:1 in the three habitats. Individuals from the centre of fragments were smaller than those from the centre of continuous forest, and FA did not vary significantly among habitats. These results suggest that small forest fragments maintain dense populations of C. chilensis and therefore they must be considered in conservation strategies. Although the decrease of the body size suggests that small remnants should be connected by managing the structure of the surrounding matrix, facilitating the dispersion of this carabid across the landscape and avoiding possible antagonistic interactions inside small fragments.     

Evolution in populations of Swedish sand lizards: genetic differentiation and loss of variability revealed by multilocus DNA fingerprinting

The Swedish sand lizard (Lacerta agilis) is a relict species from the period of warmth following the last glacial episode and has a fragmented distribution in central Sweden and a more continuous distribution in the southern part of the country. We used this model system of colonization–extinction for a study of genetic variability within and among Swedish populations from different parts of the distribution range using multilocus DNA fingerprinting. The results from the Swedish populations are then contrasted with those from a large Hungarian population in the centre of the species geographical distribution range, which is likely to closely resemble the ancestral founding population of Sweden. Swedish populations have a low level of genetic variability compared with the Hungarian reference population, which showed a genetic variability within the range described for outbred populations. Within the Swedish populations, the average bandsharing was 0.61, the mean heterozygosity 0.45 and the estimated number of alleles 2.7. The figures for the Hungarian population were a bandsharing of 0.19, a heterozygosity of 0.89 and an estimated number of alleles of 9.8. A population bottleneck, common to all Swedish sand lizards, is indicated by less than 20% of the alleles in the Hungarian population being retained in the Swedish populations, and higher bandsharing similarity between different Swedish populations (0.33) as opposed to the Hungarian population (0.19). The limited variability found in Swedish sand lizards is strongly subdivided between populations, with an average F_ST of 0.32, indicating a very limited gene flow between the isolated populations, as well as between populations in the region where the sand lizard has a more or less continuous distribution (F_ST = 0.41).     

Leaf fluctuating asymmetry of common plantain as an indicator of habitat quality

Abstract

Although developmental instability (DI), measured as fluctuating asymmetry (FA), is expected to be positively related to environmental stress and negatively to habitat quality, the pattern found here was the reverse. Developmental instability of leaf traits (leaf width and vein distances within a leaf) was estimated (using two indices of FA: FA₄ and σ_i ²) and compared between three populations of Plantago major L. (Plantaginaceae) from northern Serbia. Two of the populations are from chronically polluted areas (Karaburma & Zemun), while Crni Lug is from an unpolluted, natural area. Results obtained using both FA indices were the same; higher asymmetry levels in the unpolluted area than in the polluted sites, were found for both traits. Between the two polluted sites, FA values were significantly higher in Karaburma site for vein distances within a leaf. Concerning differences in FA₄ values between samples, in two cases, results are similar to those found for σ_i ² values, for vein distances within leaf. These are the first quantitative data on P. major indicating that (i) plants living in the stressful sites are more symmetrical and (ii) leaf FA for plant species with wide ecological distribution such as P. major should be considered as an ‘index of habitat quality.’     

Characterization of fatty acid composition in healthy and bleached corals from Okinawa, Japan

Under bleaching conditions, corals lose their symbiotic zooxanthellae, and thus, the ability to synthesize fatty acids (FAs) from photosynthetically derived carbon. This study investigated the lipid content and FA composition in healthy and bleached corals from the Odo reef flat in Okinawa, southern Japan, following a bleaching event. It was hypothesized that the FA composition and abundance would change as algae are lost or die, and possibly microbial abundance would increase in corals as a consequence of bleaching. The lipid content and FA composition of three healthy coral species (Pavona frondifera, Acropora pulchra, and Goniastrea aspera) and of partially bleached and completely bleached colonies of P. frondifera were examined. The FA composition did not differ among healthy corals, but differed significantly among healthy, partially bleached, and completely bleached specimens of P. frondifera. Completely bleached corals contained significantly lower lipid and total FA content, as well as lower relative amounts of polyunsaturated FAs and higher relative amounts of saturated FAs, than healthy and partially bleached corals. Furthermore, there was a significantly higher relative concentration of monounsaturated FAs and odd-numbered branched FAs in completely bleached corals, indicating an increase in bacterial colonization in the bleached corals.     

Ethoxyresorufin O-deethylase (EROD) activity and fluctuating asymmetry (FA) in Zosterisessor ophiocephalus (Teleostei, Gobiidae) as indicators of environmental stress in the Venice lagoon

The impact of environmental stress on the grassgoby, Zosterisessor ophiocephalus, wasstudied at both biochemical and morphologicallevels, by comparing three sites in the Venicelagoon (the Northern, Central and Southernbasins) which differ in terms of the degree ofanthropogenic disturbance, in particular withregard to organic sediment contamination. At abiochemical level, the hepatic ethoxyresorufinO-deethylase (EROD) enzyme activity wasmeasured as a biomarker of the degree oforganic pollution (in particular regardingcontamination by PCBs and Dioxins). At amorphological level, the amount of fluctuatingasymmetry (FA) with regard to four bilateraltraits (postorbital length; otolith area;minimum otolith diameter; maximum and minimumotolith radii) was measured as a long termindicator of developmental stress. The ERODactivity was higher in grass gobies from theCentral basin, whereas the amount of FA washigher in the gobies collected from theNorthern basin. Both measurements indicatedthat the Southern basin was affected by lowlevels of environmental stress (low ERODinduction and a lower amount of FA). Theresults were discussed in the light of both theenvironmental differences among the threebasins and the different levels of biologicalorganisation at which the two indicators act.     

FLUCTUATING ASYMMETRY IN CENTRAL AND MARGINAL POPULATIONS OF LYCHNIS VISCARIA IN RELATION TO GENETIC AND ENVIRONMENTAL FACTORS

Developmental instability in the form of increased fluctuating asymmetry can be caused by either genetic or environmental stress. Because extinctions can be attributed broadly to these factors, fluctuating asymmetry may provide a sensitive tool for detecting such stresses. We studied the level of fluctuating asymmetry of flowers of a perennial outcrossing plant species, Lychnis viscaria, both in natural and common-garden populations. The degree of flower asymmetry was higher in small, isolated, and marginal populations of the species range. These marginal populations also were the most homozygous. In the core area of the species' range, flowers were more symmetrical The level of asymmetry was correlated with both population size and heterozygosity. However, a partial correlation analysis revealed that when the impact of population size was controlled for, there was a negative relationship between fluctuating asymmetry and heterozygosity, whereas when controlling for heterozygosity, no relationship between population size and fluctuating asymmetry was found. This indicates that genetic consequences of small population size probably underlie the relationship between the level of asymmetry and population size. Results from a transplantation experiment showed that individuals subjected to a higher environmental stress had an increased level of asymmetry compared to control plants. In the common-garden conditions the level of fluctuating asymmetry did not differ between the central and marginal populations. This suggests that presumably both genetic and environmental factors affected to the higher level of asymmetry among marginal populations compared to central ones. In all we conclude that even though fluctuating asymmetry seems to be a sensitive tool for detecting stresses, results from studies focusing on only one factor should be interpreted with caution.     

Lizards from Urban Areas Are More Asymmetric: Using Fluctuating Asymmetry to Evaluate Environmental Disturbance

The increase in human activities that leads to wildlife decline and species extinction poses an urgent need for simple indicators of environmental stress in animal populations. Several studies have suggested that fluctuating asymmetry (FA) can be an easy, direct measure of developmental instability because it is associated to environmental stress and, as such, it can be a useful indicator of population disturbance. We examined three different morphological traits in urban and rural populations of the common wall lizard (Podarcis muralis) to test whether anthropogenic disturbance causes an increase in FA. Compared to rural populations, urban ones showed higher levels of FA in all analyzed traits, thus providing evidence that FA can respond to anthropogenic disturbance. However, we also found significant differences in FA among traits, where femoral pores and subdigital lamellae, traits with a functional relevance, were more stable developmentally compared to supracilliar granules which have no evident function. Unsigned FA [abs(right-left)] exhibited significant, but weak, positive correlations among traits, indicating that developmental noise does not have a uniform effect across characters and thus questioning the view of developmental stability as an organism-wide property. The degree of signed FA (right-left) was more similar between structurally associated traits, possibly as an outcome of morphological integration. In conclusion, our results demonstrate that FA can be a reliable indicator of disturbance provided that it is analyzed on multiple traits simultaneously and examined at the population level.     

Effect of directional selection for body size on fluctuating asymmetry in certain morphological traits in <Emphasis Type="Italic">Drosophila ananassae</Emphasis>

Variation in the subtle differences between the right and left sides of bilateral characters or fluctuating asymmetry (FA) has been considered as an indicator of an organism’s ability to cope with genetic and environmental stresses during development. However, due to inconsistency in the results of empirical studies, the relationship between FA and stress has been the subject of intense debate. In this study, we investigated whether stress caused by artificial bidirectional selection for body size has any effect on the levels of FA of different morphological traits in Drosophila ananassae. The realised heritability (h²) was higher in low-line females and high-line males, which suggests an asymmetrical response to selection for body size. Further, the levels of FA were compared across 10 generations of selection in different selection lines in both sexes for sternopleural bristle number, wing length, wing-to-thorax ratio, sex comb-tooth number and ovariole number. The levels of FA differed significantly among generations and selection lines but did not change markedly with directional selection. However, the levels of FA were higher in the G10 generation (at the end of selection) than G0 (at the start of selection) but lower than the G5 generation in different selection lines, suggesting that the levels of FA are not affected by the inbreeding generated during the course of selection. Also, the levels of FA in the hybrids of high and low lines were significantly lower than the parental selection lines, suggesting that FA is influenced by hybridisation. These results are discussed in the framework of the literature available on FA and its relationship with stress.     

Does fluctuating asymmetry constitute a sensitive biomarker of nutritional stress in house sparrows (Passer domesticus)?

Small random deviations from left–right symmetry in bilateral traits, termed fluctuating asymmetry (FA), are theoretically predicted to increase with environmental stress and believed to constitute a potential biomarker in conservation. However, reported relationships between FA and stress are generally weak and variable among organisms, traits and stresses. Here we test if, and to what extent, FA increases with nutritional stress, estimated from independent feather growth measurements, in free-ranging house sparrows (Passer domesticus). Ptilochronological feather marks showed significant heterogeneity among study plots, indicating that house sparrow populations were exposed to variable levels of nutritional stress during development. However, individuals from more stressed populations did not show increased levels of fluctuating asymmetry in tarsus or rectrix length, nor was there evidence for significant between-trait concordance in FA at the individual or the population level. Lack of support for FA in tarsus and rectrix length as estimator of nutritional stress in house sparrows may indicate that developmental instability is insensitive to nutritional stress in this species, poorly reflected in patterns of fluctuating asymmetry due to ecological or statistical reasons, or highly context-specific. Such uncertainty continues to hamper the use of FA as a biomarker tool in conservation planning.     

Water stress and light intensity effects on growth and nocturnal acid accumulation in a terrestrial CAM bromeliad (Bromelia humilis Jacq.) under natural conditions

Summary Seasonal variations in CAM performance of sunexposed and partially shaded populations of Bromelia humilis were measured under natural conditions in a semi-arid region in northern Venezuela. The sun population consisted of smaller plants, with lower chlorophyll and total nitrogen contents per unit leaf area compared with plants from the partial-shade population. During the dry season CAM activity, assessed as nocturnal acid accumulation, was higher in the partial-shade population. Acid accumulation was stimulated by irrigation in both populations within 24 h after treatment. Daily changes in concentration of soluble sugars were opposite to leaf acidity indicating their role as carbon source for acid synthesis during the night. The change in nocturnal sugar concentration was always more than the amount required for acid accumulation, suggesting other carbohydrate-consuming processes such as transportation of sugars out of the leaf. CAM activity was higher during the rainy season, and differences between populations were smaller. At the end of the rainy season reduction of CAM activity caused by drought was first detected in the sun population. Measured ratios of glucan/soluble sugar show a higher proportion of readily utilizable sugars during periods of active CAM and growth. Under conditions of continuous high light intensity and air temperature leading to all year round high potential evaporation in semiarid tropical regions, fully exposed populations of B. humilis show a pronounced reduction of metabolic activity. Partial shade favours growth and CAM activity in this constitutive CAM species. It is concluded that water stress, and not light intensity, is the predominant limiting factor for growth of this species under natural conditions.     

Environmental stress increases skeletal fluctuating asymmetry in the moor frog Rana arvalis

Whether fluctuating asymmetry (FA) provides a useful metric indicator of the degree of environmental stress experienced by populations is still a contentious issue. We investigated whether the degree of FA in skeletal elements is useful in elucidating the degree of environmental stress experienced by frog populations, and further, tested the proposition that a trait’s sensitivity to stress—as reflected in the degree of FA—is related to the degree of directional selection experienced by the given trait. We compared the degree of FA in four bilateral skeletal elements of male and female moor frogs (Rana arvalis) originating from low (acidified) and neutral pH populations. While the degree of uncorrected FA was unrelated to the degree of acidity, the growth rate and age of the individuals, the size-corrected FA was significantly higher in low than in neutral pH populations and decreased with individual ages and growth rates. In addition, both measures of FA were significantly higher in males and in particular in traits presumably under high sexual selection as indicated by the degree of sexual size dimorphism. All in all, the results indicate that individuals from acidified localities are smaller, younger and exhibit a significantly higher degree of FA than individuals from neutral pH populations. These results constitute the first assessment of FA in amphibians and suggest that the degree of FA in skeletal traits can be a useful indicator of the degree of environmental stress experienced by amphibian populations.     

二色仙人掌，中国仙人掌科一新归化种

2019年4-6月,在江苏省西南部句容市郭庄镇虬山,首次发现二色仙人掌(Opuntia cespitosa Rafinesque)归化种群。该种群主要生长在海拔46～90 m的废弃采石场上,分布范围东西长510 m,南北宽260 m,面积约13.26 hm2。二色仙人掌开花结实的成年植株共计910棵,其中最大灌木平卧茎长轴上叶状茎10个,树龄可达10 a,大概在2010年就生长于虬山。在句容虬山采石场关闭时,采石场工作人员可能将盆栽的二色仙人掌就地丢弃,从而导致该归化种群的发展。比较了二色仙人掌与中国仙人掌属已知其余4种归化植物的形态特征,并编制了分种检索表。此外,讨论了二色仙人掌的传播机制及其入侵风险。     

Range-edge effects promote clonal growth in peripheral populations of the one-sided wintergreen Orthilia secunda

Existing in suboptimal conditions is a frequent occurrence for species inhabiting the cusp of their ecological range. In range‐edge populations of plants, the scarcity of suitable habitat may be reflected in small population sizes which may result in increased self‐pollination and/or inbreeding and an increase in the incidence of clonal reproduction. These factors may result in a decrease in levels of genetic diversity and a loss of potential adaptive variation that may compromise species’ ability to cope with changes in their environment, an issue that is particularly relevant today with the current concern surrounding global climate change and its effect on species’ distributional ranges. In the present study, we have compared the levels of clonal reproduction in the one‐sided wintergreen Orthilia secunda (L.) House in (1) populations from its main continuous distribution range, (2) populations occurring on the limits of the continuous range, and (3) peripheral populations outwith the species’ continuous distribution range. Range‐edge populations in Scotland and Sweden displayed significantly lower genotypic richness and diversity than those from the main area of the species’ distribution in these countries. Populations from Ireland, which occur in the temperate zone rather than the boreal conditions that are the preferred habitat for the species, and which represent relict populations left over from cooler periods in the Earth's history, displayed no within‐population genetic diversity, suggesting a complete lack of sexual reproduction. Furthermore, the genetic distinctiveness of the Irish populations, which contained alleles not found in either the Scottish or the Swedish populations, highlights the value of ‘trailing edge’ populations and supports the concept of ‘parochial conservation’, namely the conservation of species that are locally rare but globally common.     

水菜花种群潜在生境选择与空间格局预测

国家二级保护野生植物水菜花(Ottelia cordata)，喜生于清洁的水环境中，对环境变化极为敏感，是检验湿地环境及气候变化的关键指示物种之一，在我国仅零星分布于海南北部的火山熔岩湿地区，生存状况不容乐观。研究水菜花种群潜在生境选择及其空间格局演变，有利于加强濒危物种保护保育及湿地生态系统修复、管理。该研究基于GIS平台和MaxEnt模型，结合气候、地形和土壤因子，探究水菜花种群环境限制因子及其在气候变化背景下潜在适宜生境的演变格局。结果表明，水菜花种群对温差与降水量变化敏感，等温性、最冷季度降水量、土壤类型和年均降水量对水菜花种群分布影响显著；全新世中期-当前-2070年气候变化背景下，水菜花适宜生境面积先减小后增大，分布重心呈西南-东北-西南转移格局；未来气候情景下，水菜花种群高度和中度适宜生境缩减，低适宜生境增加，南部地区将出现新增适宜生境，东北、西北及西南部适宜生境将发生消减。该研究从气候环境角度论证了水菜花种群的潜在生境选择及空间变化特征，可为濒危物种保护保育、湿地管理及其生物多样性维护工作提供参考和指导。     

Fluctuating asymmetry in leaves and flowers of sympatric species in a tropical montane environment

Fluctuating asymmetry (FA) represents small, random variations in traits, presumably with bilateral symmetry, and is widely used as a tool to measure developmental instability in plants and animals. Because FA is a quick, simple and reliable measure, it has been frequently used for monitoring levels of environmental stress. This study investigated whether FA can be used as a predictor of individual developmental instability for four sympatric Melastomataceae species. To achieve that aim, 20 individuals of Trembleya laniflora, T. parviflora, Lavoisiera campos‐portoana and Tibouchina heteromalla were marked in southeastern Brazil and monitored before and during the flowering season. The FA index was calculated as the mean of the difference in the width or length between the left and right sides measured for each leaf or flower. All studied species exhibited asymmetry in the leaves and flowers, but the leaves of L. campos‐portoana and the petal width of T. heteromalla exhibited directional asymmetry, also an indicator of developmental instability. The highest level of leaf asymmetry was found in T. heteromalla and on flowers of L. campos‐portoana. None of the studied species exhibited a significant relationship between the FA level of the leaves and flowers on an individual basis, indicating that environmental and/or genetic sources of stress might act differently on different plant traits. For the studied species, measurements of FA can be suggested as useful tools to biomonitor levels of stress experienced by both leaves and flowers within the Melastomataceae family.     

Developmental instability in hybrids between Lychnis viscaria and Lychnis alpina (Caryophyllaceae)

Developmental instability shown by increased fluctuating asymmetry can be caused by either genetic or environmental stress. Genomic coadaptation and heterozygosity are the genetic factors that are commonly assumed to increase the level of developmental stability. Therefore, in hybrid populations the level of fluctuating asymmetry (FA) can be lower due to higher heterozygosity or higher due to disruption of coadapted gene complexes, depending on the degree of divergence between hybridizing taxa. Here I present data on FA in petals from hybrids between Lychnis viscaria (Caryophyllaceae) and Lychnis alpina and from parental species grown in a common garden environment. Petal asymmetry of hybrids was clearly higher than that of either parental species grown in common environment. Between the two parental species petal asymmetry did not differ. The mean size of the petals in hybrids was about the same as in L. viscaria, thus indicating no heterotic effect. Therefore, it seems that hybrids between L. viscaria and L. alpina do suffer from the disruption of coadapted gene complexes as indicated by higher developmental instability.     

Evolution of faster development does not lead to greater fluctuating asymmetry of sternopleural bristle number inDrosophila

Both strong directional selection and faster development are thought to destabilize development, giving rise to greater fluctuating asymmetry (FA), although there is no strong empirical evidence supporting this assertion. We compared FA in sternopleural bristle number in four populations ofDrosophila melanogaster successfully selected for faster development from egg to adult, and in four control populations. The fraction of perfectly symmetric individuals was higher in the selected populations, whereas the FA levels did not differ significantly between selected and control populations, clearly indicating that directional selection for faster development has not led to increased FA in sternopleural bristle number in these populations. This may be because: (i) development time and FA are uncorrelated, (ii) faster development does result in FA, but selection has favoured developmentally stable individuals that can develop fast and still be symmetrical, or (iii) the increased fraction of symmetric individuals in the selected populations is an artifact of reduced body size. Although we cannot discriminate among these explanations, our results suggest that the relationship between development time, FA and fitness may be far more subtle than often thought.     

Bilateral asymmetry of otoliths from Collichthys lucidus of different sizes in Haizhou Bay and Xiangshan Bay

In recent years, the frequent occurrence of most human activities has seriously affected the structure and functioning of coastal ecosystems. The asymmetric relationship between the left and right otoliths of fish is often used to test the difference in fluctuating asymmetry (FA) reflected by the square of the coefficient of asymmetric variation (CV²_a), which can be regarded as an important step in the study of marine environmental pressure and implementation of offshore ecological restoration. In this study, the authors tested the bilateral FA of Collichthys lucidus in the coastal waters of Haizhou Bay, Jiangsu, and Xiangshan Bay, Zhejiang, China, using four sagittal otolith characters (length, width, perimeter and area) as biological characters. The results showed that the value of CV²_a in otolith width (more sensitive to environmental pressure) of C. lucidus in Xiangshan Bay was higher than that in Haizhou Bay, indicating that the environmental pressure on Xiangshan Bay was relatively high. The authors did not find any significant differences in the FA of otoliths between different body sizes of C. lucidus, which may be related to the short-distance migration in different regions and the dietary shifts in the life history of this species. The results have conservation and management implications for this population.