Shape variation in the rough periwinkle Littorina saxatilis on the west and south coasts of Britain

Variation in the shape of the shell in Littorina saxatilis Olivi has been shown to be due largely to the same variables on both the west and the south coasts of Britain, and it exhibits various clines. Two important aspects are the size of the aperture, which becomes relatively larger from the Isle of Man southwards to Cornwall and eastwards from Devon to the Isle of Wight, and the jugosity of the shell, which increases with distance from Cornwall both northwards as far as the Isle of Man and eastwards as far as Kent. Superimposed on the clines are domains of shape, notably one in Lewis/Harris, where the shells have a relatively large aperture, which is long and narrow, coupled with a rather globose second whorl. The local and geographical aspects of shell shape variation are discussed.     

Distribution of the species of rough periwinkle (Littorina) in Great Britain

The status of the four currently recognised species in the Littorina saxatilis species-complex, i.e. Littorina nigrolineata Gray, L. arcana Hannaford Ellis, L. saxatilis (Olivi) and L. neglecta Bean is reviewed briefly, with notes on their characteristic features and location on the shore. Since the taxonomy of these rough periwinkles has only become stable relatively recently much of the previously published information on distribution is of little or no use. In this paper their distribution around the coastline of England and Wales (with some notes on Scotland) is described and discussed. L. saxatilis is found in a wide range of habitats from exposed peninsulas to estuaries on all shores where there is a suitable rocky or stony substrate; also in salt marsh pools. L. arcana has a more restricted distribution and is notably absent along much of the south English coast, central Cardigan Bay and possibly northern Scotland; it is not found in estuaries. L. nigrolineata has an even more restricted distribution, although it occurs both on exposed coasts and in estuaries; it has only been found by us in one locality on the east coast. L. neglecta is probably fairly widely distributed but we have few details so far. The implications of the different patterns of distribution are discussed.     

Estimation of Fine-Scale Recombination Intensity Variation in the <Emphasis Type="Italic">white–echinus</Emphasis> Interval of <Emphasis Type="Italic">D. melanogaster</Emphasis>

Accurate assessment of local recombination rate variation is crucial for understanding the recombination process and for determining the impact of natural selection on linked sites. In Drosophila, local recombination intensity has been estimated primarily by statistical approaches, by estimating the local slope of the relationship between the physical and genetic maps. However, these estimates are limited in resolution and, as a result, the physical scale at which recombination intensity varies in Drosophila is largely unknown. Although there is some evidence suggesting as much as a 40-fold variation in crossover rate at a local scale in D. pseudoobscura, little is known about the fine-scale structure of recombination rate variation in D. melanogaster. Here we experimentally examine the fine-scale distribution of crossover events in a 1.2-Mb region on the D. melanogaster X chromosome using a classic genetic mapping approach. Our results show that crossover frequency is significantly heterogeneous within this region, varying approximately 3.5-fold. Simulations suggest that this degree of heterogeneity is sufficient to affect levels of standing nucleotide diversity, although the magnitude of this effect is small. We recover no statistical association between empirical estimates of nucleotide diversity and recombination intensity, which is likely due to the limited number of loci sampled in our population genetic data set. However, codon bias is significantly negatively correlated with fine-scale recombination intensity estimates, as expected. Our results shed light on the relevant physical scale to consider in evolutionary analyses relating to recombination rate and highlight the motivations to increase the resolution of the recombination map in Drosophila.     

Invasive Mahonia plants outgrow their native relatives

Invasive populations often grow more vigorously than conspecific populations in the native range. This has frequently been attributed to evolutionary changes resulting either from founder effects, or from natural selection owing to enemy release. Another mechanism contributing to evolutionary change has largely been neglected in the past: Many invasive plant species do actually descend from cultivated plants and were therefore subject to breeding, including hybridization and artificial selection. In a common garden experiment, we compared invasive Central European populations of the ornamental shrub, Mahonia, with native populations of its putative parental species, Mahonia aquifolium and M. repens, from North America. We hypothesized that plants of invasive populations show increased growth and retained high levels of heritable variation in phenotypic traits. Indeed, invasive Mahonia plants grew larger in terms of stem length, number of leaves and above-ground biomass than either of the two native species, which did not differ significantly from each other. Since there are no hints on release of invasive Mahonia populations from natural enemies, it is likely that hybridization and subsequent selection by breeders have lead to an evolutionary increase of plant vigour in the introduced range. Further on, heritable variation was not consistently reduced in invasive populations compared with populations of the two native species. We suggest that interspecific hybridization among the Mahonia species has counteracted the harmful effects of genetic bottlenecks often associated with species introductions. Based on this case study, we conclude that, more attention has to be paid on the role of plant breeding when assessing the mechanisms behind successful plant invasions in future.     

The 18S rRNA gene sequences of four commercially important seaweeds

18S rRNA gene sequences are presented forAhnfeltia plicata, Chondrus crispus, Furcellaria lumbricalis andPalmaria palmata, commercially important marine algae of the North Atlantic. The sequences range from 1765 to 1777 nucleotides in length, with guanine + cytosine content of 50.1% to 52.4%. Sequence divergence between species in different orders was 11.3–12.3%, whereas the variation betweenC. crispus andF. lumbricalis, both from the Gigartinales, was only 3.6%. Based on limited experience with other groups of Rhodophyta, these sequences obtained from single populations are likely to be representative of the species as a whole, with little variation expected among conspecifics regardless of morphological aberration or apparent genetic isolation.NRCC 34824.     

Intraspecific variation in resistance of frog eggs to fungal infection

Documenting sources of variation in host viability at pathogen exposure within and among populations is an important task in order to predict host-pathogen evolutionary dynamics. In the present study, we investigated family and population variation in the degree of embryonic infection of the pathogenic fungus Saprolegnia spp., by infecting moor frog (Rana arvalis) eggs from six populations and exposing them to two different temperatures. We found a significant family effect on the degree of Saprolegnia-infection of eggs and embryos, suggesting that there is genetic variation in resistance among embryos, or variation among females in some aspect of maternally induced resistance. Furthermore, infection level differed significantly between temperatures, with most families having more infected eggs in the relatively colder temperature. However, eggs and embryos from the different populations showed different degrees of Saprolegnia-infection in the two temperatures, i.e., there was a significant population × temperature interaction on the proportion of infected eggs. Thus, the degree of Saprolegnia-infection is sensitive to variation at the level of the family, population and environmental conditions, suggesting that responses to fungal outbreaks will vary geographically and will be difficult to predict.     

Lack of metabolic temperature compensation in the intertidal gastropods,Littorina saxatilis (Olivi) and L. obtusata (L.)

Two intertidal snails, Littorina saxatilis (Olivi, 1972) (upper eulittoral fringe/maritime zone) and Littorina obtusata (Linnaeus, 1758) (lower eulittoral) were collected from a boulder shore on Nobska Point, Cape Cod, Massachusetts, in July and acclimated for 15–20 days at 4 ° or 21 °C. Oxygen consumption rate (Vo₂) was determined for 11–15 subsamples of individuals at 4 °, 11 ° and 21 °C with silver/platinum oxygen electrodes. Multiple factor analysis of variance (MFANOVA) of lo₁₀ transformed values of whole animal Vo₂ with log₁₀ dry tissue weight (DTW) as a covariant revealed that increased test temperature induced a significant increase in Vo₂ in both species (P<0.00001). In contrast, MFANOVA revealed that temperature acclimation did not affect Vo₂ in either L. saxatilis (P= 0.35) or L. obtusata (P= 0.095). Thus, neither species displayed a capacity for the typical metabolic temperature compensation marked by an increase in Vo₂ at any one test temperature in individuals acclimated to a lower temperature that is characteristic of most ectothermic animals. Lack of capacity for metabolic temperature acclimation has also been reported in other littorinid snail species, and may be characteristic of the group as a whole. Lack of capacity for respiratory temperature acclimation in these two species and other littorinids may reflect the extensive semi-diurnal temperature variation that they are exposed to in their eulittoral and eulittoral fringe/maritime zone habitats. In these habitats, any metabolic benefits derived from longer-term temperature compensation of metabolic rates are negated by extreme daily temperature fluctuations. Instead, littorinid species appear to have evolved mechanisms for immediate metabolic regulation which, in L. saxatilis and L. obtusata and other littorinids, appear to centre on a unique ability for near instantaneous suppression of metabolic rate and entrance into short-term metabolic diapause at temperatures above 20–35 °C, making typical seasonal respiratory compensation mechanisms characteristic of most ectotherms of little adaptive value to littorinid species.     

Developing a breeding strategy to exploit quantitative variation in symbiotic nitrogen fixation

Summary This paper examines evidence which quantifies the relative importance of legume and Rhizobium genotypes as determinants of phenotypic variation in symbiotic nitrogen fixation. It demonstrates potentially large and unpredictable effects of the Rhizobium genotype. The likely importance of such effects on crop yield is considered. The information is then used to assess ways in which legume breeding programmes may be altered to encompass the effects of genetic variation in Rhizobium.     

Aggressive Mimicry in the Intra-populational Color Variation of the Tanganyikan Scale-eater Perissodus microlepis (Cichlidae)

A color morph of the Tanganyikan scale-eater, Perissodus microlepis (Cichlidae), considered to be an example of aggressive mimicry, is reported from the rocky littoral area at the southern end of Lake Tanganyika, Zambia. The anal fin of the morph is yellow, instead of the usual dark or transparent, being closely similar to that of the gregarious cyprichrominine cichlids, Cyprichromis leptosoma species complex and Paracyprichromis nigripinnis, abundant in the area. The color morph of Perissodus microlepis was observed to mingle with and attack shoals of the prey. About 24% of P. microlepis observed in less than 15m depth at Kasenga Point were the yellow-fin type. The color variation, which is likely associated with a variation in feeding mode within this population, probably evolved and has been maintained by the local community structure, which included abundant cyprichrominine fishes.     

Selection for Antimicrobial Peptide Diversity in Frogs Leads to Gene Duplication and Low Allelic Variation

Antimicrobial peptides are highly diverse pathogen-killing molecules. In many taxa, their evolution is characterized by positive selection and frequent gene duplication. It has been proposed that genes encoding antimicrobial peptides might be subject to balancing selection and/or an enhanced mutation rate, but these hypotheses have not been well evaluated because allelic variation has rarely been studied at antimicrobial peptide loci. We present an evolutionary analysis of novel antimicrobial peptide genes from leopard frogs, Rana. Our results demonstrate that a single genome contains multiple homologous copies, among which there is an excess of nonsynonymous nucleotide site divergence relative to that expected from synonymous site divergence. Thus, we confirm the trends of recurrent duplication and positive selection. Allelic variation is quite low relative to interspecies divergence, indicating a recent positive selective sweep with no evidence of balancing selection. Repeated gene duplication, rather than a balanced maintenance of divergent allelic variants at individual loci, appears to be how frogs have responded to selection for a diverse suite of antimicrobial peptides. Our data also support a pattern of enhanced synonymous site substitution in the mature peptide region of the gene, but we cannot conclude that this is due to an elevated mutation rate.     

Temperature during egg formation and the effect of climate warming on egg size in a small songbird

The predicted effects of recent climate warming on egg size in birds are controversial, as only two long-term studies have been reported, with contrasting results. Long-term data on egg size variation are analyzed in relation to ambient temperatures in a southern European population of pied flycatchers where breeding phenology has not matched the spring advancement in the last decades. Cross-sectional, population analyses indicated that egg breadth, but not egg length, has decreased significantly along the 16-year period, leading to marginally non-significant decreases in egg volume. Longitudinal, individual analyses revealed that despite females consistently laying larger eggs when they experienced warmer temperatures during the prelaying and laying periods, there was an overall negative response – i.e. decreasing egg volume and breadth with increasing spring (May) average temperatures – across individuals. This trend is hypothesised to be caused by the mismatched breeding phenology, in relation to climate warming, of this population. Except in the unlikely cases of populations capable of perfectly synchronising their phenology to changes in their environment, maladjustments are likely for traits such as egg size, which depend strongly on female condition. Slight changes or absence thereof in breeding dates may be followed by mismatched dates, in terms of food abundance, for optimal egg formation, which would be reflected in smaller average egg size, contrary to early predictions on the effects of climate warming on bird egg size.     

The taxonomy of the Littorina saxatilis speicies-complex, with particular reference to the systemaitc status of Littorina patula Jeffrys

Variation in shell shape and penis morphology of Littorina rudis Maton is examined using data from all parts of Britain. The shell shape variation within populations of L. rudis is shown to account for Liltorina patula Jeffrys at the only site where the latter species was recorded. In addition, the shell shape of I,. rudis varies with exposure, individuals on exposed shores having a relatively larger aperture than those on sheltered shores. Wave action and desiccation are considered the most likely factors maintaining this variation. The penis morphology of L. rudis varies within and between shores to an extent that renders the use of this character invalid for distinguishing L. patula from L. rudis. The radulae of adults of L. rudis, L. patula and Littorina nigrolineata (Gray) are similar in structure having blunt cusps, whilst adult Littorina neglecta Bean and juvenile L. rudis have pointed cusps. The possibility of a neotenous origin of L. neglecta from L. rudis is discussed. On the evidence presented here it is suggested that L. patula must be regarded as a synonym of L. rudis.     

Validation of a planimetric procedure to quantify stress in Littorina littorea (Gastropoda: Mollusca): is it independent of the reproductive cycle?

This study forms part of a larger project in which five planimetric parameters have been used to study changes in the digestive epithelium of Littorina littorea under different environmental and physiological conditions. Our aim was to examine the effect of the reproductive cycle on these parameters in order to assess their usefulness as indicators of stress. A one way anova shows that the absolute parameters of mean epithelial thickness (MET), mean diverticulum radius (MDR) and mean luminal radius (MLR) vary significantly depending on the time of the year. This variation is highly correlated to the amount of digestive tissue present and hence negatively correlated to the reproductive state because gonad and digestive tissue volume are inversely related. Consequently, these parameters are not good indicators of stress. However, whereas the absolute size of the digestive acinus varies with the reproductive state of the animal, the MET, MDR and MLR retain the same proportions and therefore the ratios MET/MDR and MLR/MET remain constant. This makes them useful indicators of stress because they are independent of intrinsic variables such as the reproductive cycle.     

Spatial patchiness of epilithic biofilm caused by refuge-inhabiting high shore gastropods

Aggregations of Melarhaphe neritoides and Littorina saxatilis are common on the high shore in north-east England. These aggregations are frequently found in crevices which act as a refuge from desiccation. The aggregations are surrounded by patches of epilithic biofilm which are visibly lighter in colour than the biofilm present on the remainder of the high shore. Chlorophyll a levels were lower in regions of close proximity to aggregated littorinids (mean = 4.9 μg cm⁻²) as compared to areas >‰5 cm away from the edge of the visible colour change (mean = 15.9 μg cm⁻²). Manipulations of littorinid density showed that areas of high density had significantly lower levels of chlorophyll a than those where littorinids were excluded. This difference was mainly due to the presence of higher numbers of small grazed areas or an increase in size of grazed areas rather than a homogenous change in chlorophyll a levels across the entire manipulated area. These results are supported by observations showing that only 7% of the total foraging time of littorinids was spent outside visibly lighter patches of biofilm suggesting littorinids only control local biofilm levels through grazing.This paper has not been submitted elsewhere in identical or similar form, nor will it be during the first three months after its submission to Hydrobiologia     

Assessment of interspecific interactions among parasitoids on the outcome of inoculative biological control of leafminers attacking chrysanthemum

Indigenous natural enemies occur within field grown crops at varying densities dependent upon a variety of other biotic and abiotic parameters. This natural control often does not provide adequate suppression, which results in the application of other pest management solutions including augmentative biological control. When releasing mass-reared natural enemies into a backdrop of existing natural enemy populations, competitive interactions are likely to occur. To assess the influence of these interspecific interactions on the outcome of such biological control practices studies were conducted in a simulated, field cage grown, cut chrysanthemum production system. Competitive interactions of two commercially available parasitoids were studied both in terms of parasitoid-host population dynamics and the impact of interspecific interactions on crop quality at harvest. The parasitoids Diglyphus isaea and Dacnusa sibirica attacking the leafminer Liriomyza langei were used as the model insect system. Both parasitoids are cosmopolitan and are known to occur in many ornamental production areas. Treatment comparisons included single species releases with complimentary releases of both species either simultaneously or with 2-week time lags, as well as a no release control to measure the background effects of natural mortality. Conclusions drawn from results of population-level studies replicated within and among years were that levels of interspecific competition among parasitoid species were undetectable at leafminer densities typical of field-grown ornamental crops (low densities), and thus, the efficacy of one species released into a backdrop of potentially competing parasitoids did not negatively or positively affect the outcome of the augmentative biological control, nor was there a positive outcome; however, crop quality at harvest was influenced.     

Phylogenetics of New World Astragalus: Screening of novel nuclear loci for the reconstruction of phylogenies at low taxonomic levels

This study explores methods to use information gathered from genomics technology to understand evolutionary relationships in the hyperdiverse legume group Neo-Astragalus. These species inhabit deserts and mountains of North and South America, and even though the monophyly of the group is well established, relationships within it are still poorly understood. Plastid genes, commonly used to infer phylogenies in plants, are usually not useful for closely related taxa because of low levels of genetic variation. TheMedicago truncatula genome project provided a suite of candidate nuclear loci with high levels of variation that might prove suitable for low-level phylogenetics. This paper reports the development of methods for screening a large number of these nuclear loci, and detailed analysis of four of them. Four different patterns of phylogenetic diversification occur in the loci sampled from these genomes ofAstragalus species. One locus (CNGC4) was single copy and could be directly used in phylogenetic analyses. Two loci (ARG10 and FENR) showed patterns strongly suggestive of duplication events in some taxa, and one locus (tRALS) has apparently undergone a cryptic duplication, making it very difficult to diagnose. Potential methods for using the information provided by these loci are discussed.     

Compensation for floral herbivory in <Emphasis Type="Italic">Solanum carolinense</Emphasis>: identifying mechanisms of tolerance

While a plant’s capacity to tolerate damage by herbivores can be studied as a single trait, it is important to recognize that tolerance is generally a result of the combined action of several different traits. Here, we report on a pair of experiments to identify mechanisms for tolerating floral herbivory in Solanum carolinense, an andromonoecious perennial herb that regularly suffers from high levels of florivory. We measured the effect of actual and simulated florivory on host-plant fitness and assessed which plant traits exhibited plasticity in response to florivory. In addition, for each of nine plant genets, we calculated tolerance indices and determined which traits were genetically correlated with tolerance. Traits that served to help S. carolinense tolerate florivory in terms of sexual reproduction included initiating more inflorescences, aborting fewer buds prior to anthesis and fewer ovaries after fertilization, and increasing the ratio of perfect:male flowers. In addition, the greater the levels of florivory, the more the plants allocated to root growth, which may promote tolerance through greater potential future reproduction. The plant population contained significant genetic variation for tolerance itself and for nearly all of the putative tolerance mechanisms, which suggests that S. carolinense has the potential to evolve greater tolerance through a variety of different routes in response to natural selection.     

Individual variation in two host plants of the ladybird beetle,Epilachna pustulosa (Coleoptera: Coccinellidae)

Individual variation in two species of host plants (thistle,Cirsium kamtschaticum, and blue cohosh,Caulophyllum robustum) of the herbivorous ladybird beetleEpilachna pustulosa was examined under laboratory conditions for their acceptability to adult beetles as a food resource, for adult preference and for larval performance. When clones of these plants were subjected to non-choice tests using posthibernating female beetles, there was found to be significant intraspecific variation among clones in terms of their acceptability, but interspecific variation was not detected. Significant intraspecific as well as interspecific variation were frequently detected in the two host plants when clones of these plants were subjected to choice tests using posthibernating female beetles; the magnitude of interspecific plant variation for beetle preference is not necessarily larger than that of intraspecific plant variation. Individual variation across plant species with respect to beetle larval performance was also significant. A positive correlation between adult preference and larval performance is suggested across the two taxonomically remote host plant species, thistle and blue cohosh, although this needs further investigation.     

Maintenance of zonation patterns in two species of flat periwinkle,Littorina obtusata and L. mariae

The zonation patterns of Littorina obtusata (L.) and Littorina mariae Sacchi et Rastelli were shown to be quite distinct on a sheltered rocky shore. L. obtusata was found at all the heights sampled; it reached peak numbers at mid shore on the alga Ascophyllum nodosum L. (Le Jol). There was no difference in the tidal height occupied by adults or juveniles; or in the mean size of L. obtusata along the vertical gradient of the shore. In contrast L. mariae occurred exclusively low on the shore, on Fucus serratus L. Translocation of the two species within their respective levels resulted in random movement after 4 days, although initial movements after 1 and 2 days were sometimes directional. Animals transplanted to the normal level of the other species showed directional movement towards their home zone; this was most pronounced after 4 days. There was no difference in the distance moved by the two species, although the distance moved did vary with tidal height, both species moving further at mid shore than low shore. Distances moved by littorinids at replicate areas in the low shore were similar but those at mid shore did vary. There was an interaction between the species and the different tidal heights which revealed that transplanted species moved further than translocated species at the same tidal level. However, this was only significant in the case of L. mariae. It is suggested that the close relationship between the winkles and their host algae may direct the homing behaviour of displaced individuals.     

Sequence differentiation in regions identified by a genome scan for local adaptation

Genome scans using large numbers of randomly selected markers have revealed a small proportion of loci that deviate from neutral expectations and so may mark genomic regions that contribute to local adaptation. Measurements of sequence differentiation and identification of genes in these regions is important but difficult, especially in organisms with limited genetic information available. We have followed up a genome scan in the marine gastropod, Littorina saxatilis, by searching a bacterial artificial chromosome library with differentiated and undifferentiated markers, sequencing four bacterial artificial chromosomes and then analysing sequence variation in population samples for fragments at, and close to the original marker polymorphisms. We show that sequence differentiation follows the patterns expected from the original marker frequencies, that differentiated markers identify independent and highly localized sites and that these sites fall outside coding regions. Two differentiated loci are characterized by insertions of putative transposable elements that appear to have increased in frequency recently and which might influence expression of downstream genes. These results provide strong candidate loci for the study of local adaptation in Littorina. They demonstrate an approach that can be applied to follow up genome scans in other taxa and they show that the genome scan approach can lead rapidly to candidate genes in nonmodel organisms.