Climate change affects colonization dynamics in a metacommunity of three Daphnia species

Climate change is expected to alter the range and abundance of many species by influencing habitat qualities. For species living in fragmented populations, not only the quality of the present patches but also access to new habitat patches may be affected. Here, we show that colonization in a metacommunity can be directly influenced by weather changes, and that these observed weather changes are consistent with global climate change models. Using a long‐term dataset from a rock pool metacommunity of the three species Daphnia magna, Daphnia longispina and Daphnia pulex with 507 monitored habitat patches, we correlated a four‐fold increase in colonization rate with warmer, drier weather for the period from 1982 to 2006. The higher colonization rate after warm and dry summers led to an increase in metacommunity dynamics over time. A mechanistic explanation for the increased colonization rate is that the resting stages have a higher exposure to animal and wind dispersal in desiccated rock pools. Although colonization rates reacted in the same direction in all three species, there were significant species‐specific effects that resulted in an overall change in the metacommunity composition. Increased local instability and colonization dynamics may even lead to higher global stability of the metacommunity. Thus, whereas climate change has been reported to cause a unidirectional change in species range for many other species, it changes the dynamics and composition of an entire community in this metacommunity, with winners and losers difficult to predict.     

Fish kairomone regulation of internal swarm structure in Daphnia pulex (Cladocera: Crustacea)

In this laboratory experiment, swarms of D. pulex were artificially created by placing thirty individuals in small chambers containing 130 ml of water. The swimming behaviour of the animals was measured using a video camera and a programme to digitize observations. The D. pulex swimming in water with fish kairomones had a more uniform swimming speed compared with animals in control water. If aggregated prey individuals benefit from a reduced predation risk, and this risk is further reduced by uniformity of swarm members, the uniformity of swimming speed can be interpreted as an behavioural adjustment to minimize the vulnerability to predation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Changes in vulnerability of Daphnia to an insecticide application depending on the population phase

1. Laboratory populations of cloned Daphnia pulex at three population phases (growing phase, density peak, declining phase) were exposed to the insecticide carbaryl at 15 μg L^?1, which killed juveniles but not adults. 2. The population decline was largest and recovery was slowest when the chemical was applied at the density peak, indicating that the population was most sensitive to the chemical at that stage. Populations in the declining phase were least affected by the treatment. 3. We propose that food condition, which varies with population density, was a major factor affecting vulnerability to carbaryl. In that case, Daphnia populations in lakes would be most vulnerable to toxic chemicals during food‐limited condition such as the spring clear‐water phase.     

Costs, benefits and the evolution of inducible defences: a case study with Daphnia pulex

Phenotypic plasticity is one major source of variation in natural populations. Inducible defences, which can be considered threshold traits, are a form of plasticity that generates ecological and evolutionary consequences. A simple cost-benefit model underpins the maintenance and evolution of these threshold, inducible traits. In this model, a rank-order switch in expected fitness, defined by costs and benefits of induction between defended and undefended morphs, predicts the risk level at which individuals should induce defences. Here, taking predator-induced morphological defences in Daphnia pulex as a threshold trait, we provide the first comprehensive investigation into the costs and benefits of a threshold trait, and how they combine to reflect fitness and predict the switchpoint at which induction should occur. We develop reaction norms that show genetic variation in switchpoints. Further experiments show that induction can confer a survival benefit and a cost in terms of lifetime reproductive success. Together, these two traits combine to estimate expected fitness and can predict the switchpoint between an undefended and a defended strategy. The predictions match the reaction norm data for clones that experience these costs and benefits, and correspond well to independent field data on induction. However, predictions do not, and cannot, match for clones that do not gain a benefit from induction. This study confirms that a simple theory, based on life history costs and benefits, is a sufficient framework for understanding the ecology and evolution of inducible, threshold traits.     

Side effects of water liming on body size and eggs development of Daphnia species in Lake Orta (N. Italy)

Life-tables of the Daphnia species (D. longispina and D. obtusa), currently inhabiting the previously acidified Lake Orta, were followed to study the side effects of liming on the demographic parameters (e.g. longevity, body size increment, eggs and new-borns production), of both species. Different media were prepared to simulate the lake water pH's (5, 6 and 7), and copper concentrations (40 μg l^-1), before and after liming, using GF/C filtered lake water in such simulation and as a reference medium. The obtained results have shown that, the individuals of D. longispina were only able to survive after the liming and the consequent reduction in copper concentrations. However, due to shift occurred in the different media pH's after liming toward slightly alkaline conditions, no significant variations in body size increment or in eggs and newborns production were detected of this species in the different media (p = 0.38, p = 0.29, p = 0.09 respectively). On the other hand, the effect of the variations in pH on the body size increment of the B. obtusa individuals was marginally significant (p = 0.07). A highly significant (p = 0.002), increase in eggs production by D. obtusa individuals was detected in the different media after the CaCO₃ addition. Conversely, a corresponding highly significant (p = 0.0007), decrease in the production of new-borns, accompanied by the degeneration of about 78% of the produced eggs, was detected in the different media after liming. Our results suggest that in Lake Orta the actual irregular abundance of the studied Daphnia species and the eggs degeneration phenomenon, observed in our study, could be largely attributed to the interspecific competition resulting from species-specific sensitivity to the continuous variations in the lake water Chemistry after the liming operation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Population divergence of genetic (co)variance matrices in a subdivided plant species,Brassica cretica

Abstract The present study of Brassica cretica had two objectives. First, we compared estimates of population structure (Q_st) for seven phenotypic characters with the corresponding measures for allozyme markers (F_st) to evaluate the supposition that genetic drift is a major determinant of the evolutionary history of this species. Secondly, we compared the genetic (co)variance ( G ) matrices of five populations to examine whether a long history of population isolation is associated with large, consistent differences in the genetic (co)variance structure. Differences between estimates of F_st and Q_st were too small to be declared significant, indicating that stochastic processes have played a major role in the structuring of quantitative variation in this species. Comparison of populations using the common principal component (CPC) method rejected the hypothesis that the G matrices differed by a simple constant of proportionality: most of the variation involved principal component structure rather than the eigenvalues. However, there was strong evidence for proportionality in comparisons using the method of percentage reduction in mean‐square error (MSE), at least when characters with unusually high (co)variance estimates were included in the analyses. Although the CPC and MSE methods provide different, but complementary, views of G matrix variation, we urge caution in the use of proportionality as an indicator of whether genetic drift is responsible for divergence in the G matrix.     

Sperm number assessed by flow cytometry in species of Daphnia (Crustacea,Cladocera)

Daphnia magna and Daphnia pulex are two important model species in ecotoxicology. In daphniids, studies of the effects of contaminants have mostly focused on female life history traits, yet it would also be important to examine male reproductive traits, particularly in relation to endocrine disruptors. In this study, we developed a protocol that uses flow cytometry to measure sperm number in individual males of different species of Daphnia. We tested our protocol on 114 males from several clones of three common species of Daphnia. Sperm count varied widely among individuals and reached high numbers (up to 1.45 × 10⁵). Positive relationships between male length and sperm number were observed in D. pulex and Daphnia pulicaria, but not in D. magna. Important inter‐clonal differences in sperm production were observed in all species, with some clones producing very little sperm. Duplicated sperm samples showed on average only 6% difference in sperm counts. Sperm counts were stable at least over a 2‐hr period and up to 5 hr for most samples. This sperm isolation protocol and flow cytometric enumeration approach will be of major interest to ecotoxicologists.     

Historical habitat connectivity affects current genetic structure in a grassland species

Many recent studies have explored the effects of present and past landscape structure on species distribution and diversity. However, we know little about the effects of past landscape structure on distribution of genetic diversity within and between populations of a single species. Here we describe the relationship between present and past landscape structure (landscape connectivity and habitat size estimated from historical maps) and current genetic structure in a perennial herb, Succisa pratensis. We used allozymes as co‐dominant markers to estimate genetic diversity and deviation from Hardy–Weinberg equilibrium in 31 populations distributed within a 5 km² agricultural landscape. The results showed that current genetic diversity of populations was related to habitat suitability, habitat age, habitat size and habitat connectivity in the past. The effects of habitat age and past connectivity on genetic diversity were in most cases also significant after taking the current landscape structure into account. Moreover, current genetic similarity between populations was affected by past connectivity after accounting for current landscape structure. In both cases, the oldest time layer (1850) was the most informative. Most populations showed heterozygote excess, indicating disequilibrium due to recent gene flow or selection against homozygotes. These results suggest that habitat age and past connectivity are important determinants of distribution of genetic diversity between populations at a scale of a few kilometres. Landscape history may significantly contribute to our understanding of distribution of current genetic structure within species and the genetic structure may be used to better understand landscape history, even at a small scale.     

Landscape scale genetic effects of habitat fragmentation on a high gene flow species: Speyeria idalia (Nymphalidae)

Detection of the genetic effects of recent habitat fragmentation in natural populations can be a difficult task, especially for high gene flow species. Previous analyses of mitochondrial DNA data from across the current range of Speyeria idalia indicated that the species exhibited high levels of gene flow among populations, with the exception of an isolated population in the eastern portion of its range. However, some populations are found on isolated habitat patches, which were recently separated from one another by large expanses of uninhabitable terrain, in the form of row crop agriculture. The goal of this study was to compare levels of genetic differentiation and diversity among populations found in relatively continuous habitat to populations in both recently and historically isolated habitat. Four microsatellite loci were used to genotype over 300 individuals from five populations in continuous habitat, five populations in recently fragmented habitat, and one historically isolated population. Results from the historically isolated population were concordant with previous analyses and suggest significant differentiation. Also, microsatellite data were consistent with the genetic effects of habitat fragmentation for the recently isolated populations, in the form of increased differentiation and decreased genetic diversity when compared to nonfragmented populations. These results suggest that given the appropriate control populations, microsatellite markers can be used to detect the effects of recent habitat fragmentation in natural populations, even at a large geographical scale in high gene flow species.     

Population genetic structure of three tree species in the mangrove genus Ceriops (Rhizophoraceae) from the Indo West Pacific

Ceriops is a viviparous mangrove with widespread species Ceriops decandra and C. tagal, and an endemic species C. australis. Genetic diversity of the three species was screened in 30 populations collected from 23 locations in the Indo West Pacific (IWP) using Inter-simple sequence repeats (ISSR) and sequences of partial nuclear gene (G3pdh) and chloroplast DNA (trnV-trnM). At the species level, the total gene diversity (Ht) revealed by ISSRs was 0.270, 0.118, and 0.089 in C. decandra, C. tagal, and C. australis, respectively. A total of six haplotypes of G3pdh and five haplotypes of trnV-trnM were recognized among the three species. Only C. decandra was detected containing more than one haplotype from each sequence data set (four G3pdh haplotypes and three trnV-trnM haplotypes). At the population level, genetic diversity of Ceriops was relatively low inferred from ISSRs (He = 0.028, 0.023, and 0.053 in C. decandra, C. tagal, and C. australis, respectively). No haplotype diversity within population was detected from any of the three species. Cluster analysis based on ISSRs identified three major geographical groups in correspond to the East Indian Ocean (EIO), South China Sea (SCS), and North Australia (NA) in both C. decandra and C. tagal. The cladogram from DNA sequences also detected the same three geographical groups in C. decandra. Analysis of molecular variance (AMOVA) revealed that most of the total variation was accounted for by differentiation between the three major geographical regions of both C. decandra and C. tagal. The significant genetic structure may result from the geological events in these regions during the recent Pleistocene glaciations. This study also provided insights into the phylogenetics of Ceriops. Yelin Huang and Fengxiao Tan contributed equally to this work.     

Population dynamics of hoolock gibbons (Hylobates hoolock) in Assam,India

Data are presented on group size, composition, and reproduction of several hoolock gibbon groups studied in different areas of Assam, India, between 1986 and 1988. Presence of fewer young than reported in earlier studies signals a grim future for these gibbons unless strict conservation measures are undertaken.     

Ingestion Rates of Some Daphnia Species in a Shallow Lake (Lake Balaton,Hungary)

Filtering rates of three species of Daphnia were studied in situ by the ¹⁴C technique during the summer in a lake with a water temperature of 20-25°C (Lake Balaton, Hungary). The renewal time of gut contents proved to be 3-5 min in each of the three species whether fed on natural or artificial food. From May to September, filtering rates of D. cucullata, D. hyalina and D. galeata averaged 7.8, 10.3 and 9.2 ml ind.⁻¹ day⁻¹, respectively. Identically long individuals of all three species had almost the same filtering rates in spring, whereas in summer animals of identical weights had similar filtering rates regardless of species because of the length variability of the helmets. The grazing rate of adult filter-feeding crustaceans is suggested to be approximately 7 % per day.     

黄土高原不同气象因子对中国沙棘亚居群遗传多样性的影响

顶生金花茶(Camellia terminalis)是一种濒危植物,主要分布在广西天等县石灰岩山钙质土上的常绿阔叶林内.采用样地调查、统计分析和回归模型预测等方法,对顶生金花茶种群的分布环境、资源量现状、大小和高度结构等进行了研究.结果表明:该种群主要依存于天等县小山乡的蚬木-石山樟林(P1)和任豆-蚬木林(P2)、福新乡的枝花李榄-石山楠林(P3)和齿叶大风子-闭花木林(P4)4种群落,构成灌木层的优势种和共优种;P1、P3和P4种群处于群落较稳定的典型常绿季雨林中,结构较完整,属于稳定或增长型;P2种群由于恶劣环境和频繁人为干扰的共同胁迫,有衰退的趋势;与小山乡生境相比,福新乡相对冷凉湿润的气候条件、丰厚肥沃的土层以及较轻度的人为干扰有利于顶生金花茶种群的发展.目前顶生金花茶种群的资源量有限,建议列为"严重濒临灭绝"(CR)等级.     

Population genetic structure,migration, and polyploidy origin of a medicinal species Gynostemma pentaphyllum (Cucurbitaceae)

Gynostemma pentaphyllum, a member of family Cucurbitaceae, is a perennial creeping herb used as a traditional medicinal plant in China. In this study, six polymorphic nSSR and four EST‐SSR primers were used to genotype 1,020 individuals in 72 wild populations of G. pentaphyllum. The genetic diversity and population structure were investigated, and ecological niche modeling was performed to reveal the evolution and demographic history of its natural populations. The results show that G. pentaphyllum has a low level of genetic diversity and high level of variation among populations because of pervasive asexual propagation, genetic drift, and long‐term habitat isolation. Results of the Mantel test demonstrate that the genetic distance and geographic distance are significantly correlated among G. pentaphyllum natural populations. The populations can be divided into two clusters on the basis of genetic structure. Asymmetrical patterns of historical gene flow were observed among the clusters. For the contemporary, almost all the bidirectional gene flow of the related pairs was symmetrical with slight differences. Recent bottlenecks were experienced by 34.72% of the studied populations. The geographic range of G. pentaphyllum continues to expand northward and eastward from Hengduan Mountains. The present distribution of G. pentaphyllum is a consequence of its complex evolution. Polyploidy in G. pentaphyllum is inferred to be polygenetic. Finally, G. pentaphyllum is a species in need of protection, so in situ and ex situ measures should be considered in the future.     

Population genetic diversity and structure of a naturally isolated plant species, Rhodiola dumulosa (Crassulaceae)

AIMS: Rhodiola dumulosa (Crassulaceae) is a perennial diploid species found in high-montane areas. It is distributed in fragmented populations across northern, central and northwestern China. In this study, we aimed to (i) measure the genetic diversity of this species and that of its populations; (ii) describe the genetic structure of these populations across the entire distribution range in China; and (iii) evaluate the extent of gene flow among the naturally fragmented populations. METHODS: Samples from 1089 individuals within 35 populations of R. dumulosa were collected, covering as much of the entire distribution range of this species within China as possible. Population genetic diversity and structure were analyzed using AFLP molecular markers. Gene flow among populations was estimated according to the level of population differentiation. IMPORTANT FINDINGS: The total genetic diversity of R. dumulosa was high but decreased with increasing altitude. Population-structure analysis indicated that the most closely related populations were geographically restricted and occurred in close proximity to each other. A significant isolation-by-distance pattern, caused by the naturally fragmented population distribution, was observed. At least two distinct gene pools were found in the 35 sampled populations, one composed of populations in northern China and the other composed of populations in central and northwestern China. The calculation of Nei's gene diversity index revealed that the genetic diversity in the northern China pool (0.1972) was lower than that in the central and northwestern China pool (0.2216). The populations were significantly isolated, and gene flow was restricted throughout the entire distribution. However, gene flow among populations on the same mountain appears to be unrestricted, as indicated by the weak genetic isolation among these populations.     

Effects of population size and isolation on the genetic structure of the East African mountain white‐eye Zosterops poliogaster (Aves)

Habitat size, quality and isolation determine the genetic structure and diversity of populations and may influence their evolutionary potential and vulnerability to stochastic events. Small and isolated populations are subject to strong genetic drift and can lose much of their genetic diversity due to stochastic fixation and loss of alleles. The mountain white‐eye Zosterops poliogaster, a cloud forest bird species, is exclusively found in the high mountains of East Africa. We analysed 13 polymorphic microsatellites for 213 individuals of this species that were sampled at different points in time in three mountain massifs differing in habitat size, isolation and habitat degradation. We analysed the genetic differentiation among mountain populations and estimated the effective population sizes. Our results indicate three mountain‐specific genetic clusters. Time cohorts did not show genetic divergences, suggesting that populations are large enough to prevent strong drift effects. Effective population sizes were higher in larger and geographically interconnected habitat patches. Our findings underline the relevance of ecological barriers even for mobile species and show the importance of investigating different estimators of population size, including both approaches based on single and multiple time‐points of sampling, for the inference of the demographic status of a population. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 828–836.     

Molecular genetic analysis of Daphnia in the English Lake District: species identity, hybridisation and resting egg banks

• 1 Taxonomic issues in many Daphnia species complexes are often confused by a high degree of phenotypic plasticity and by interspecific hybridisation. Here, we employ molecular genetic tools to confirm the species composition and incidence of hybridisation in extant and resting egg populations of Daphnia from Windermere and Esthwaite Water in the English Lake District.
• 2 A combination of species‐diagnostic allozymes and mtDNA, confirms that contemporary populations are dominated by a single species, D. galeata.
• 3 We present the first account of genetic characterisation of dated ‘resting’ eggs using microsatellites and mtDNA, employing PCR‐based DNA recovery, thus providing a temporal dimension to taxonomic patterns. Thirty years ago, two species were present in Esthwaite, D. galeata and D. hyalina, but Windermere populations were dominated by D. galeata only.
• 4 The use of PCR‐based mtDNA RFLP analysis as a species‐diagnostic tool, and microsatellites to monitor clonal diversity, provide a valuable approach for long‐term studies, especially in populations free from the complicating effect of frequent hybridisation. The detailed limnological records available for many large lakes, and associated changes in land‐use, pollutants and climate, combined with long‐term ephippial molecular genetic data, provide opportunities for exploring natural and anthropogenic impacts on genetic and community structure.

     

High genetic diversity and low structure in an endemic long-lived tree,Yucca capensis (Asparagaceae)

• Endemic species distributed in fragmented habitats are highly vulnerable to extinction because they may have low genetic diversity. However, some life-history traits can mitigate the effect of genetic drift on populations. We analysed the level and distribution of genetic variation and ancestral population size of Yucca capensis, a long-lived endemic plant of the Baja California Peninsula, Mexico. Its populations are scattered across a habitat that is suffering accelerated transformation.
• We used six nuclear microsatellites to genotype 224 individuals from 17 locations across the entire species' geographic range. We estimated polymorphisms, heterozygosity and genetic structure. We also evaluated the ancestral and recent effective size and time since the population started to change.
• We found high heterozygosity, high polymorphism and low differentiation among locations, suggesting a panmictic population across the range. We also detected a large ancestral effective population size, which suffered a strong reduction in the Mid-Holocene.
• Despite changes in environmental conditions caused by habitat modification, the high diversity and low differentiation in Y. capensis may result from its large ancestral effective size and life-history traits, such as plant longevity, clonal growth and mating system, which reduce the rate of loss of genetic variation. However, the dependence on a specialist pollinator that displays short flight range can reduce gene flow among the plant populations and could, shortly, lead them into an extinction vortex.

     

Population genetic structure of the South American species Hypochaeris lutea (Asteraceae)

The genus Hypochaeris has a recent evolutionary history caused by long‐distance dispersal in conjunction with adaptive radiation in the South American continent. Hypochaeris lutea is a perennial herb that grows mostly at altitudes of around 1000 m in cold swamps of the southern regions of Brazil. We investigated the amplified fragment length polymorphism (AFLP) in 270 individuals representing 11 Brazilian populations of H. lutea to elucidate the population genetic structure of this species. The frequencies of polymorphic loci and gene diversity ranged from 83.42% to 91.66% and from 0.26 to 0.34, respectively. Analysis of molecular variance revealed that most of the genetic variability was found within (76.67%) rather than among (23.3%) populations, agreeing with the pattern of genetic distribution within and among populations observed in other allogamous species of Hypochaeris. A Mantel test showed no correlation between genetic and geographic distances when all populations were considered. Simulations performed using a Bayesian approach consistently identified two clusters with different admixture proportions of individuals, as also revealed by a UPGMA dendrogram of populations. The pattern of genetic structure observed in H. lutea is consistent with a process of successive colonization events by long‐distance dispersal resembling the rapid and recent radiation that has been proposed to explain the origin of the South American species of Hypochaeris.     

Finding the relevant scale: clonality and genetic structure in a marine invertebrate (Crambe crambe, Porifera)

Important changes in genetic relatedness may occur at extremely small scales in benthic invertebrates, providing key information about structuring processes in populations of these organisms. We performed a small-scale study of the population structure of the sponge Crambe crambe, in which 177 individuals from the same rocky wall (interindividual distances from 0 to 7 m) were genotyped using six microsatellite markers. 101 sponges had unique genotypes and the remaining 76 individuals formed 24 groups of sponges sharing genotypes (clones). Mean intraclone distances were found to be c. 20 cm. Spatial autocorrelation analyses showed a drastic decrease in genetic relatedness over the first 100 cm of distance. If the contribution of clonality to this pattern was eliminated, the trend was attenuated, but remained a marked one and was still significant within the first distance classes (30-40 cm). Estimated mean dispersal distances per generation were c. 35 cm, and neighbourhood sizes were estimated at c. 33 sponges. Genetic similarities with sponges of the same locality, or from other Mediterranean localities, were within the same range as those found in sponges 2-7 m apart. It is concluded that asexual reproduction plays an important role in structuring populations in this species. However, over and above the effects of clonality, a strong fine-scale genetic structure was present at distances in the range of tens of centimetres, probably as a result of short dispersal of larvae. This fine-scale genetic structure may be common in invertebrates with lecitotrophic larvae.