SHELL SCARRING IN CALLIOSTOMA ZIZYPHINUM (PROSOBRANCHIA: TROCHIDAE) FROM STRANGFORD LOUGH, NORTHERN IRELAND

Pink and white shells of Calliostoma zizyphinum show both undamagedand damaged scars. White shelled individuals predominate inStrangford Lough, Northern Ireland where Calliostoma densitiesand scarring levels are high. No white shells were found intertidally,outside the entrance channel to the Lough. Significant differencesoccur in the size of Calliostoma shells between sampling sitesbut not between different colour morphs at each site. Howeverin pooled samples, pink individuals are significantly largerthan white individuals. When sites and morphs are consideredseparately, pink Calliostoma density is negatively correlatedwith water movement. At sites where pink Calliostoma occur,the percentage of pink shelled individuals is negatively correlatedwith total Calliostoma density. Damaged and undamaged scarring values per unit area of shell,show highly significant differences between sites and damagedscars are significantly higher in white individuals. Undamagedscars are not correlated with any of the environmental parametersrecorded, but are positively correlated with damaged scars suggestinga common causative factor. The level of damaged scarring ispositively correlated with crab/total Calliostoma ratio at allsites and where each colour morph was considered separately.Multiple regression analyses reveal that crab/Calliostoma ratiosaccount for 42% of the between site variation in damaged scars.Significantly higher levels of damaged scars are found at siteswith high crab densities and significantly larger individualsare found at sites where crab densities are intermediate invalue. The largest and most highly scarred individuals occur at siteswith most coarse substrata where Calliostoma are present intheir lowest densities. The higher scarring levels and smallersize of white individuals reflect either higher mortality orreduced growth in white shelled Calliostoma. (Received 25 November 1991; accepted 5 November 1992)     

SHELL STRENGTH OF COLOUR MORPHS OF THE MANGROVE SNAIL LITTORARIA PALLESCENS

The load required to break shells of the snail Littor-aria pallescens(Philippi) (Gastropoda: Littorinidae) has been established.Mass increases more than proportionally with size. Size forsize, relatively heavy shells are thicker than relatively lightones, and the thicker shells are stronger. Three samples fromdifferent localities differ significantly in strength. Littorariapallescens is polymorphic for shell colour. It is shown thatin two of the three samples, both from Papua New Guinea, individualsof the dark morph are significantly heavier and stronger thanthose of the yellow morph, while there is no difference betweenmorphs in the third sample, from Thailand. In the sample withthe most robust shells the dark individuals are stronger, massfor mass, than yellows. Thus, colour morph is sometimes, butnot always, associated with differences in shell characteristicswhich are likely to affect fitness. Where there is a difference,the dark morph is estimated to be on average about 22% strongerthan the yellow morph. The difference between morphs is smallcompared with the large variation in shell characters associatedwith size. (Received 20 May 1992; accepted 22 June 1992)     

SIZE, SHAPE AND SHELL MORPHOLOGY IN THE ANTARCTIC LIMPET NACELLA CONCINNA AT SIGNY ISLAND, SOUTH ORKNEY ISLANDS

Comparison of the size, shape and shell morphology in littoraland sub-littoral morphs of the Antarctic limpet Nacella concinnareveal differences in shell morphology which are enhanced bystructural anomalies within the shells of the two types. Infestationof sub-littoral shells by the conchocelis phase of an endolithicalga significantly affects shell density and total chlorophylllevels in the two shell morphs. The surface sculpture of sub-littoralshells is characterised by a series of grooves, the configurationof which closely resembles that of the radular teeth in N. concinna.Limpets utilise the available food supply within the shell matrixof other limpets by grazing the shell material. Epibiotic growthof calcareous algae prevent erosion and preserve underlyingshell layers. In severe cases, where protection is lacking,intraspecific shell grazing may remove parts of the shell exposingthe internal tissues. The Dominican Gull, Larus dominicanus, is a major shore predatorof both shell morphs. Gull middens contain both shell typesbut are dominated by the more accessible littoral shells. Comparisonof living populations and midden assemblages indicates thatsize and shape selection of prey occurs, with pear-shaped limpetsbetween 21 mm and 29 mm in length being taken preferentially. Apparent differences in shell form are induced by physical,biological and behavioural influences. Littoral animals arerobust in nature, resist avian pre-dation and are not extensivelygrazed whereas those of the sub-littoral are not subject tothe same degree of predatory attention but suffer a gradualdepletion of their shallower shell form through a combinationof algal infection and intraspecific shell grazing. (Received 21 February 1990; accepted 5 July 1990)     

Temperature-related diversity of shell colour in the intertidal gastropod Batillaria

The intertidal snail Batillaria exhibits remarkable variationin the shell colour within and among populations. Field studywas conducted to determine the factors in maintaining observedshell colour polymorphism. Geographical variations in shellcolour polymorphisms in B. attramentaria were significantlycorrelated with the temperature of the locality of the population.Darker morphs were predominant in colder regions, whereas lightermorphs increase their proportion in warmer regions. A consistentassociation was also found in B. multiformis that co-existedwith B. attramentaria. Strong predatory pressure imposed bydigenean trematode parasites was observed in B. attramentaria.However, it is unlikely to affect the colour variations, becauseno correlation exists between colour morphs and trematode parasitism.Although visual selection may also contribute to colour variationin Batillaria, no evidence is found for the existence of visualpredators that affect colour patterns of these snails. The deficitof variation in cold regions is possibly due to selection againstbrighter morphs, because bright colours reflect heat. Althoughdark shells absorb sunlight and may therefore be exposed tothe risks of overheating and drying up in a hot habitat, thedarkest morph was frequently observed in the warmer regions,suggesting that physical selection on the colour morphs canbe relaxed in the warmer environment. Our results suggest thatclimatic selection is one of the significant factors maintainingshell colour polymorphism in these intertidal snails. (Received 13 December 2006; accepted 14 April 2007)     

Shell colour variation in Bullia digitalis, a sand-dwelling, intertidal whelk (Gastropoda: Prosobranchia)

Bullia digitalis is an intertidal whelk that lives on sandy beaches in South Africa. It is highly variable in shell colour, with individuals varying from white to dark brown. This paper describes shell colour variation of B. digitalis at seven sites, along a 230 km coastline east of the Cape Peninsula. Seven colour forms were found: striped, violet, banded violet, banded brown, orange, pale yellow and white. These forms are probably genetically determined morphs. The striped form is the most common at all sites, constituting 53–62% of each sample. The violet is the second most common morph. Its frequencies are remarkably stable at 15–17%. The striped form blends well into the sandy environment and may therefore be of considerable cryptic value in concealing B. digitalis from predators. The violet form is highly conspicuous. Its stable frequency throughout the study area may represent a genetic balance that is not relevant to any visual advantages of the violet colour.     

ALTITUDINAL VARIATION AND ADAPTIVE ZONES IN ARIANTA ARBUSTORUM: A NEW LOOK AT A WIDESPREAD SPECIES

Arianta arbustorum is a European land snail, occurring in bothlowland and mountainous regions. Its shell is usually globular,which makes the species aberrant among the Campylaeinae in shellshape, habitat, and wide, not exclusively montane range. Withincreasing altitudes, the shells usually decrease in size andbecome more conical. However, in some mountain stocks thereare populations with depressed shells, provided with an openumbilicus. This form (subspecies) is only known from formerPleistocene refugia. It is hypothesized that in the Pliocene, the primary montaneA. arbustorum, with a depressed shell, entered an additional,adaptive zone, while conquering damp lowland habitats. Thisresulted in an enormous range extension. In the new habitata different (globular) shell shape evolved and the species becamepolytypic. The subsequent glaciations wiped out most of theoriginal montane A. arbustorum with depressed shells. Wherethis had occurred, post-Pleistocene re-colonizing conspecificsnails with globular shells, found vacant territory and coulddevelop the dines mentioned before. The invading populationsdid not regain the original, montane, depressed shell shape.This shape was only retained in a few refugia, where the populationssurvived amidst the glaciers. (Received 9 February 1990; accepted 31 May 1990)     

Visual selection of shell colour in two littoral prosobranchs

The rough periwinkle Littorina “saxatilis” exhibits a wide range of shell colours. In current literature it is claimed that these colours are not related to the environment, that they arise randomly as genetic accidents and that they have little positive survival value. In a previous paper it has been shown that, in Wales, “saxatilis” consists of four separate, fully sympatric species. This paper reports an investigation as to whether the colour of two of these species, nigrolineata and rudis, is related to the colour of the background. Because of difficulties in quantitatively describing the colour of the background it was decided to concentrate mainly upon one aspect: whether or not the frequency of red shells is larger upon shores consisting of red sandstone than elsewhere in Wales. In both species the association between red shells and red sandstone is highly significant. All nigrolineata samples in which red was found at a frequency above 15% are from red sandstone. On red sandstone, red shells of this species were found mainly upon sheltered shores, their frequency decreasing and that of white shells increasing with exposure. This may be because barnacles, which occupy the same vertical zone as nigrolineata, are more abundant upon exposed shores. Partly covering the red rock, barnacles create a white background upon which white shells, rather than red, are cryptic. Yellow shells are found mainly on sheltered shores, where the brown alga Fucus is abundant. It was observed that when the tide is in and the algae spread out, a yellow shell situated beneath them is well concealed. Yellow is also found upon barnacles which because of fungal and lichen infections, are dirty yellow. It is suggested that a striped (“nigrolineated”) pattern breaks up the shape of the shell. It also resembles the colour of the dark rock and the dark sutures between barnacle plates. In rudis 80% of the samples containing over 25% red are from red sandstone. Contrast to nigrolineata, in this species the relative frequency of red decreases on sheltered shores. This could be because the brown alga Pelvetia, which occupies the same vertical zone as rudis, is more abundant in sheltered conditions and its colour partly covers over that of the rock beneath. The fact that in both species red shells are more frequent upon red sandstone than elsewhere in the study area, suggests that visual selection is restricting their distribution to the background that they match most. Rock pipits and shore-crabs prey upon winkles. They have colour perception and could be partly responsible for this selection.     

Variation in shell morphology of Calliostoma zizyphinum (Gastropoda: Trochidae)

Differences in shell morphology in the intertidal prosobranchmollusc Calliostoma zizyphinum were studied from a number ofsites within four geographical regions of the British Isleswith varying exposures to wave action and crab predation. Meanvalues of damage scarring were highest in shells sampled fromStrangford Lough, Northern Ireland, and lowest in individualsfrom the Atlantic Coast. Shells collected from the Isle of Manwere smaller than those from either the Atlantic coast of Irelandor Strangford Lough. Shells from Strangford Lough had tallershells (higher aspect ratio) than shells from the County Downand Atlantic Coast and shells from the County Down coast weremore squat (lower aspect ratio) than those from all other areas.Shells from Strangford Lough were significantly thinner thanshells from all other geographical areas. The relationshipsbetween shell damage scarring and shell size and tallness werenot consistent among areas. Shells from Strangford Lough arenotable in showing a steep, positive relationship between shellsize and scarring and a steep, negative relationship betweenscarring and tallness. The unusual shell phenotypes observedin shells from Strangford Lough may be explained by rapid shellgrowth, which would not only allow Calliostoma to attain a sizerefuge from crab predators but also to recover successfullyfrom repeated crab attacks on the shell lip. Such a strategywould result in larger, thinner shells with a high number ofdamage scars. (Received 6 June 2006; accepted 20 December 2006)     

Results of testing the comparatory method: The curvature of the shell valve frontal section is inappropriate as a systematic character for the freshwater pearl mussel of the genus Margaritifera

This paper continues a discussion on the number of pearl mussel species of the genus Margaritifera in northern Europe. A biometric study of 1711 pearl mussel Margaritifera margaritifera shells from 15 rivers in Russia and Latvia (basins of the White and Baltic seas) has been conducted. All the examined samples fall into two groups: the northern group (with the shells more flattened on average, f. margaritifera) and the southern one (with more convex shells, f. elongata); the boundary between these groups is at 63° N. Analysis of intrapopulation variation has shown that the samples contain individuals that correspond to f. margaritifera, f. elongata, and f. borealis. However, any hiatus between these forms is absent in all the samples, and individuals belonging to two intermediate forms are rather frequent. The hypothesis on the species specificity of the shell valve frontal section has not been confirmed based on examination of large shell samples. The pearl mussels inhabiting rivers of Northern Europe belong to a single species, M. margaritifera.     

Enigmatic septa in shells of some Middle Jurassic Pholadomya (Bivalvia) from Poland

The interior of 36 specimens of Pholadomya Sowerby (Bivalvia) from the Middle Jurassic of Poland reveals the presence of unusual septa that separate sediment‐filled chambers from the shell interior. The septa occupy one or several recurrent loci in shells of various individuals, that is within umbones, in pallial sinuses and along the shell margins. Based on the location and shape of the septa, eight forms grouped into types and varieties are identified. A possible cause for the formation of septa is sediment toxicity, but intrusion of sediment to the shell interior must have been linked to shell breakage or rupture of the free periostracum. One form of septa occurs in perforated umbones, common in Pholadomya; other forms occur in intact shells, which suggest damage to free periostracum. The most likely cause for the latter is the presence of parasites, especially digenean trematodes, for which clams were intermediate hosts. The morphological aberrations presented here were hitherto unknown in both fossil and extant bivalves. This study is also the first report of pathologies in Anomalodesmata.     

SPATIAL PATTERNS OF SHELL SHAPE OF THREE SPECIES OF CO-EXISTING LITTORINID SNAILS IN NEW SOUTH WALES, AUSTRALIA

The shape and relative weight of the shell have been shown tovary intraspecifically and interspecifically in a number ofspecies of gastropods, including many different littorinids.These differences give rise to different shell forms in differenthabitats. In those species which have non-planktotrophic development,differences in shell form among shores have been usually explainedin terms of natural selection because exposure to waves supposedlyfavours light shells with large apertures, while predation bycrabs on sheltered shores favours elongated, thick shells withsmaller apertures. Differences in shell shape among speciesfound at different heights on the shore have been explainedin terms of resistance to desiccation and temperature. Suchvariables would tend to act on a relatively broad-scale, i.e.causing differences among heights on a shore or among shores.Rates of growth, which might vary at much smaller scales withina shore, have also been shown to affect the shapes of many shells. In this study, the shape and relative weight of shells of threespecies of co-existing littorinids (Littorina unifasciata, Bembiciumnanum and Nodilittorina pyramidalis) were measured. These speciesall haveplanktotrophic development and they are found on manyshores where there is no evidence that they are preyed uponby crabs. Before explanations of shell shape are proposed, itis necessary that patterns of variation, within different partsof ashore and among different shores are clearly documented.These patterns were measured at a number of different spatialscales within and among replicate shores with different amountsof wave exposure. Large and small specimens were included toallow intraspecific comparisons among snails of different sizesfound at different heights on the shore. The results showedsignificant differences among shores in shape and relative weightof shells, but these differences could not be explained by exposureto waves. In addition, snails of different sizes and differentspecies did not show the same patterns although they were collectedfrom the same sites. Importantly, the shell shape of Liuorinaunifasciata varied significantly among sites at approximatelythe same height within a shore. These differences could notbe clearly correlated with density, mean size nor exposure towaves. The only consistent pattern was a decrease in relativeaperture size in specimens living higher on the shore. Modelsthat have commonly been proposed to explain shape and relativeweight of shells in other species of gastropods are not adequateto explain the small- and large-scale variation of the measurementsdescribed here. It is proposed that any selective advantageof shell morphology and the effects of any variables on thedevelopment of shell morphology in these species can only beidentified after appropriately designed and replicated fieldexperiments. (Received 4 March 1994; accepted 13 September 1994)     

In a world of white,flower colour matters: A white–purple transition signals lack of reward in an alpine Euphrasia

The New Zealand alpine flora displays a range of unusual characteristics compared with other alpine floras, in particular the high frequency of species with small white flowers. The presence of both white and bright purple flowers on the same plant in the New Zealand alpine annual creeping eyebright (Euphrasia dyeri Wettst.) provides an ideal opportunity to investigate the significance of flower colour in an environment where coloured flowers are rare. The relationships among flower age, gender phase, reward availability and petal colour were assessed in natural populations of E. dyeri. The effect of pollination on flower colour was tested using hand pollination of bagged flowers. Direct observations and videos of flowers were used to assess patterns of flower visitation by native and introduced pollinators. Unpollinated white E. dyeri flowers changed from white to purple within 6 days. However, pollination of white flowers triggered a significantly faster colour change, typically within 1–2 days. White flowers had receptive stigmas, large amounts of lipid‐rich pollen and small amounts of nectar, whereas stigmas of purple flowers are not receptive and flowers did not provide pollen or nectar rewards. Flowers were mainly visited by native syrphid flies. Both native syrphids and introduced Bombus bees showed a marked avoidance of purple flowers, tending to preferentially visit white flowers. Our study suggests that flower colour change from white to bright purple in E. dyeri functions to direct pollinators to rewarding, receptive flowers. As many Euphrasia L. species are described as having variably coloured flowers, this mechanism may be more widespread in the genus. Furthermore, our results add to the growing evidence that the dominance of white flowers in the New Zealand alpine is not simply due to a lack of colour discrimination among pollinators.     

HABITAT CONSTRAINTS ON SHELL-COLOUR VARIATION OF A DESERT LANDSNAIL, TROCHOIDEA SIMULATA

Trochoidea simulata, a landsnail widespread in desert regionsof Israel, varies greatly in shell colour. We have previouslyshown that predator and thermoegulatory mediated selection cannotexplain the maintenance of this colour variation. In this paper,we test the hypothesis that observed morph frequencies are aneffect of habitat variation, and that the colour variation hasno functional significance. Because white snails have thickershells than striped or brown snails, we propose that the colourof an individual snail depends on its access to CaCO₃. We testedthis hypothesis by comparing the frequency of white, striped,and brown snails at 9 locations. Each location consisted ofa paired hillside and loess wadi-bed site with high and lowCaCO₃ content respectively. There were significantly more whitesnails at hillside than loess sites. In addition, these habitatsvaried mostly in abiotic characteristics, and not in terms ofsnail population parameters or vegetation characteristics. Hillsidesites had significantly more CaCO₃ and limestone rocks thanloess sites. Soil CaCO₃ content was the strongest predictorof morph frequency among paired sites. Shell-colour variationin this species may be an effect of individual access to resources,mainly CaCO₃ that is ingested with food. (Received 5 August 1996; accepted 25 October 1996)     

AN ANALYSIS OF SHELL SHAPE VARIATION IN THE PAINTED TOPSHELL CALLIOSTOMA ZIZYPHINUM (L.) (PROSOBRANCHIA: TROCHIDAE

Littoral and sublittoral Calliostoma zizyphinum (L.) from fourteensites around Ireland are compared by a new method of shell shapeanalysis. The apparatus used is described and results show clearvariation between populations. It is also used to show similaritybetween the variegated and white morphs found in StrangfordLough. Patterns of growth are shown to be largely in responseto the habitat conditions encountered at each site. The meanapical angle of each population is shown to be of great importancein distinguishing populations of topshells, as it varies withthe relative exposure of each site. (Received 30 July 1986;     

INTRASPECIFIC AND INTERSPECIFIC VARIATION IN SHELL MORPHOLOGY OF BIOMPHALARIA PFEIFFERI (KRAUSS, 1848) AND HELISOMA DURYI (WETHERBY, 1879) FROM MOSHI, TANZANIA, ANALYSED BY MORPHOMETRIC STUDIES

Helisoma duryi, a planorbid snail species, which is a potentialbiological control agent of the intermediate hosts of schistosomiasis,was introduced in an irrigation scheme (T.P.C.) in northernTanzania in 1972. The species has been present at T.P.C. since,and during a snail survey in January 1981, a great variationin the shell morphology of H. duryi, ranging from typical H.duryi forms to forms resembling Biomphalaria pfeifferi, theintermediate host of Schistosoma mansoni in the area, was observed.The Biomphalaria- Wkc forms of H. duryi could constitute anentire population but also occasionally occurred in populationsof typical H. duryi. This study was undertaken to determine whether simple shellmeasurements, or ratios between some of these could be usefulin discriminating between H. duryi and B. pfeifferi. Followingparameters were considered: shell height, shell diameter, diameterof the umbilicus, ratio between shell height and diameter ofthe umbilicus, and the number of whorls. (Received 29 July 1983;     

Quantitative measures and 3D shell models reveal interactions between bands and their position on growing snail shells

The nature of shell growth in gastropods is useful because it preserves the ontogeny of shape, colour, and banding patterns, making them an ideal system for understanding how inherited variation develops, is established and maintained within a population. However, qualitative scoring of inherited shell characters means there is a lack of knowledge regarding the mechanisms that control fine variation. Here, we combine empirical measures of quantitative variation and 3D modeling of shells to understand how bands are placed and interact. By comparing five‐banded Cepaea individuals to shells lacking individual bands, we show that individual band absence has minor but significant impacts upon the position of remaining bands, implying that the locus controlling band presence/absence mainly acts after position is established. Then, we show that the shell grows at a similar rate, except for the region below the lowermost band. This demonstrates that wider bands of Cepaea are not an artifact of greater shell growth on the lower shell; they begin wider and grow at the same rate as other bands. Finally, we show that 3D models of shell shape and banding pattern, inferred from 2D photos using ShellShaper software, are congruent with empirical measures. This work therefore establishes a method that may be used for comparative studies of quantitative banding variation in snail shells, extraction of growth parameters, and morphometrics. In the future, studies that link the banding phenotype to the network of shell matrix proteins involved in biomineralization and patterning may ultimately aid in understanding the diversity of shell forms found in molluscs.     

Maternal inheritance of plant variegation in cowpea, Vigna unguiculata (L.) Walp.

A chimeric plant was observed in the F₂ generation of a cross between a mutant cultivar, Ife BPC, and a germplasm line, TVu 2, in cowpea, Vigna unguiculata (L.) Walp. The chimeric plant had four lateral branches, one of which was intensely variegated, while the others were mostly green with few white sectors. F₃ progeny from the intensely variegated branch of this plant were all variegated, while seed derived from the mostly green branches produced only green progeny. In subsequent generations, the descendants of the variegated branch bred true for the variegated trait, while those of the mostly green branches were also true-breeding for green colour. No pure-green or pure-white shoots were observed in any of the variegated plants examined in this study. Consequently, no pure-green or pure-white seedlings were produced from seeds harvested from the variegated plants. The results of reciprocal crosses between variegated and normal green plants indicate that variegation is inherited in a strictly uniparental maternal fashion. This is the first report of a cytoplasmically inherited mutation affecting foliage colour in cowpea. Received: 10 March 2000 / Accepted: 16 May 2000     

Stomatal Responses of Variegated Leaves to CO2 Enrichment

The responses of stomatal density and stomatal index of fivespecies of ornamental plants with variegated leaves grown attwo mole fractions of atmospheric CO₂ (350 and 700 µmolmol^-1) were measured. The use of variegated leaves allowed anypotential effects of mesophyll photosynthetic capacity to beuncoupled from the responses of stomatal density to changesin atmospheric CO₂ concentration. There was a decrease in stomataldensity and stomatal index with CO₂ enrichment on both white(unpigmented) and green (pigmented) leaf areas. A similar responseof stomatal density and index was also observed on areas ofleaves with pigmentation other than green indicating that anydifferences in metabolic processes associated with colouredleaves are not influencing the responses of stomatal densityto CO₂ concentrations. Therefore the carboxylation capacityof mesophyll tissue has no direct influence on stomatal densityand index responses as suggested previously (Friend and Woodward1990 Advances in Ecological Research 20: 59-124), instead theresponses were related to leaf structure. The stomatal characteristics(density and index) of homobaric variegated leaves showed agreater sensitivity to CO₂ on green portions, whereas heterobaricleaves showed a greater sensitivity on white areas. These resultsprovide evidence that leaf structure may play an important rolein determining the magnitude of stomatal density and index responsesto CO₂ concentrations.Copyright 1995, 1999 Academic Press Leaf structure, photosynthesis, stomatal conductance, CO₂, stomatal density, stomatal index     

Endogenous and environmental factors influence 35S promoter methylation of a maize A1 gene construct in transgenic petunia and its colour phenotype

Summary 30000 transgenic petunia plants carrying a single copy of the maize A1 gene, encoding a dihydroflavonol reductase, which confers a salmon red flower colour phenotype on the petunia plant, were grown in a field test. During the growing season plants with flowers deviating from this salmon red colour, such as those showing white or variegated phenotypes and plants with flowers exhibiting only weak pigmentation were observed with varying frequencies. While four white flowering plants were shown at the molecular level to be mutants in which part of the A1 gene had been deleted, other white flowering plants, as well as 13 representative plants tested out of a total of 57 variegated individuals were not mutants but rather showed hypermethylation of the 35S promoter directing A1 gene expression. This was in contrast to the homogeneous fully red flowering plants in which no methylation of the 35S promoter was observed. While blossoms on plants flowering early in the season were predominantly red, later flowers on the same plants showed weaker coloration. Once again the reduction of the A1-specific phenotype correlated with the methylation of the 35S promoter. This variation in coloration seems to be dependent not only on exogenous but also on endogenous factors such as the age of the parental plant from which the seed was derived or the time at which crosses were made.     

Water-borne cues from a shell-crushing predator induce a more massive shell in experimental populations of an intertidal snail

Inducible defenses are important in the life strategies of many taxa. In some species of marine gastropods, water-borne chemical cues from potential predators induce defensive changes in shell form and differences in growth rate. We examined such phenotypic plasticity in the direct-developing snail, Littorina subrotundata (Carpenter, 1864). Among experimental field populations of L. subrotundata exposed to differing intensities of predation by the purple shore crab, Hemigrapsus nudus (Dana, 1851), snails collected from predation-intense environments often had more massive shells than closely related snails from adjacent environments where predation was negligible. Snails collected from both environments were raised in tanks containing cages of H. nudus that were feeding on conspecific snails and compared to a control group raised in the absence of this stimulus. Most snails developed significantly more massive shells in the presence of the crabs suggesting that adaptive phenotypic plasticity may account for some of the variation we observed in the field. In one case, snails from a predation-intense environment did not exhibit a statistically significant amount of plasticity, but instead grew a more massive shell irrespective of the laboratory stimulus. We interpret this as evidence for a genetic difference in the plasticity of shell form among experimental populations, caused by intense selection by H. nudus. There was no statistical difference in the growth rates of snails among treatments.