Heating properties of morphs of the mangrove snail Littoraria pallescens

The rate of heating and the temperature attained after 10 and 20 min have been examined for shells of the yellow, dark and orange morphs of the mangrove leaf snail Littoraria pallescens. Under the experimental conditions used, heating from 20 to 40C took 5–10 min and thereafter the temperature remained roughly constant for up to 20 min. This represents temperature conditions which the animals are likely to experience in the wild. Large shells heat more slowly than small ones and reach lower temperatures. At around 40C yellows are on average 1.5C cooler than the dark morph, and orange individuals lie between. This difference could account for the observed difference between morphs in their choice of preferred leaf surface. Besides differing in colour, yellows are thinner and less heavy than dark shells, which probably contributes to the fact that they arc less robust.     

Sexual selection on phenotypic traits in a hybrid zone of Littorina saxatilis (Olivi)

Step clinal transitions in inherited character(s) between genetically distinct populations are usually referred to as hybrid zones. An example is found in the population of the intertidal snail Littorina saxatilis in Galicia (NW Spain). We studied the shape of the overall fitness surface for sexual selection in this hybrid zone, and the position of hybrids and pure morphs on this surface. We found that sexual divergent selection acted on a combination of phenotypic traits separating the pure morphs, and therefore that sexual selection contributed to morph differentiation. The average fitness of hybrids as a group was not significantly different from that of the pure morphs, but they did show divergent sexual selection in some traits. These results are in agreement with a model of divergent selection favouring both the pure morph as well as those hybrids most resembling each morph. The finding of divergent selection is remarkable because quadratic selection gradients are usually weak in nature.     

Colour polymorphism and thermal capacities in Theba pisana (O.F. Müller 1774)

Several land snail species are highly polymorphic regarding their shell colouration. This polymorphism has been related to predatory effects as well as climatic reasons, assuming that dark morphs benefit from being more cryptic and therefore less prone to predation, whereas pale morphs are at an advantage under solar radiation, as they are suspected to heat up less. However, the assumption of different thermal capacities of these morphs is based on experiments with little standardisation or little environmental relevance. In this study, we aimed at measuring thermal capacities of two different morphs (pale versus dark-brown banded) of the Mediterranean land snail Theba pisana, applying a standardised and environmentally relevant test set-up, in order to prove whether darker morphs indeed do heat up more than lighter coloured morphs. We did not find any differences in the thermal capacity of the different morphs and conclude that thermal capacity of the shell is predominantly defined by its material rather than its coloration. These results are discussed with regard to previous studies on thermal characteristics of different land snail morphs and correlations between climate and morph distribution.     

Sexual selection and non-random mating for shell colour in a natural population of the marine snailLittorina mariae (Gastropoda: Prosobranchia)

In order to estimate the three independent components of mating behaviour, sexual selection in females, sexual selection in males and mating pattern, we studied the distribution of shell colour morphs among mating pairs and between copulating and non-copulating snails in four subsamples of a natural population ofL. mariae. The colour of the shell, the sex and a qualitative estimate of age was recorded for every snail. We found sexual selection acting against one of the two commonest colours (yellow) among the young females. However, in males none of the eight shell colour morphs was favoured during matings. Male sexual choice or differences in female sexual activity may cause the sexual fitness disadvantage of yellow females. Moreover, individuals of different colour morphs did not mate at random, rather dissasortatively. A behavioural choice among shell colour morphs or a non-random microdistribution of the morphs may cause the departure from random mating in this population.     

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)     

Wild birds prefer the familiar morph when feeding on pastry-filled shells of the landsnail Cepaea hortensis (Müll.)

Colour polymorphisms in prey could be maintained if predators concentrate on common morphs and confer a selective advantage on rare morphs. We describe experiments to test whether wild birds feed on pastry-stuffed shells of Cepaea hortensis in a manner that might lead to such apostatic selection. The birds were first given a 'pre-training' choice test of a shell population with equal numbers of yellow unbandeds and yellow five-bandeds; they were then trained on one morph alone, given a second choice test, trained on the other morph and, finally, given a third choice test. The birds preferred five-bandeds in five of the six pre-training tests. In all six experiments the first training session increased the birds' preferences for the morph that was familiar. The results were less clear-cut when selection during pre-training was compared with selection after the second training session. However, a comparison between selection after each of the two training sessions showed that in all six experiments the results were in the direction predicted from the hypothesis that familiar morphs are preferred. This set of experiments is one of the few in which behaviour which could lead to apostatic selection has been tested with morphs that differ in pattern. The findings support the idea that polymorphism in Cepaea could be maintained by apostatic selection.     

EVIDENCE FOR PREDATION AS A FACTOR IN DETERMINING SHELL COLOR FREQUENCIES IN A MANGROVE SNAIL LITTORINA SP. (PROSOBRANCHIA: LITTORINIDAE)

It was hypothesized that in Littorina populations living on Avicennia marina in Moreton Bay, Queensland, yellow shelled individuals are at a selective advantage over other shell colors and that this advantage is due to differential selection by predators. Yellow shelled individuals were more likely to be recaptured than others, indicating a higher survival rate of yellows. When predation was restricted on ten mangrove trees, the apparent advantage of yellow shells was removed. After 18 months, the relative frequency of yellow shelled individuals was significantly lower on experimental trees than on control trees. A combination of selection for crypsis and for the less common morph is suggested as the mechanism maintaining the high levels of variation in this species.     

Lake Erie water snakes revisited: Morph- and age-specific variation in relative crypsis

Summary Populations of water snakes (Nerodia sipedon insularum) on islands in western Lake Erie are variable in colour pattern, consisting of unbanded, intermediate, and banded morphs. In contrast, mainland populations (N. s. sipedon) consist solely of banded morphs. Previous investigators hypothesized that natural selection favoured unbanded morphs on exposed island shorelines and banded morphs in overgrown mainland habitats and that gene flow from mainland populations was responsible for the persistence of banded morphs on islands. To clarify the potential role of natural selection, I quantified relative crypsis among morphs and age classes of water snakes by comparing the size of patches making up their colour patterns with the size of patches in island and mainland backgrounds. This analysis reveals that if unbanded morphs are more cryptic than intermediate and banded morphs on islands, it is only in the young-of-the-year age class. For older snakes on islands and for all snakes on the mainland, unbanded morphs are consistently less cryptic than intermediate and banded morphs. Given these results, the net direction of selection in island populations should depend on the intensity of predation on different age classes of snakes. Overall, selection may favour unbanded morphs (e.g. if predation occurs primarily on young-of-the-year), intermediate and banded morphs (e.g. if predation occurs primarily on older snakes), or be weak or absent (e.g. certain combinations of predation on young-of-the-year and older snakes). Using estimates of relative crypsis to guide reanalysis of morph frequency data, I find support for the hypothesis that unbanded morphs are favoured by natural selection in island populations.     

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)     

Pollinator visitation in populations of tristylous Eichhornia paniculata in northeastern Brazil

Summary The frequencies of floral morphs in populations of tristylous Eichhornia paniculata often deviate from the theoretical expectation of equality. This variation is associated with the breakdown of tristyly and the evolution of self-fertilization. Differences in morph frequencies could result from selection pressures due to variable levels of insect visitation to populations and contrasting foraging behavior among the floral morphs. We estimated pollinator densities in 16 populations and quantified visitation sequences to morphs in five populations of E. paniculata in northeastern Brazil. Foraging behavior among floral morphs was measured as the frequency of visits to morphs relative to their frequency in the population (preference) and number of flights between inflorescences of the same versus different morphs (constancy). Pollinator density (number/m²/minute) was not correlated with population size, plant density or morph diversity. Pollinator densities varied most among populations of less than 200 plants. Whether pollinators discriminated among the morphs, depended on whether they primarily collected nectar or pollen. In four populations, nectar-feeding bees (Ancyloscelis and Florilegus spp.) and butterflies showed no consistent preference or constancy among the morphs. In contrast, pollen-collecting bees (Trigona sp.) visited a lower proportion of longstyled inflorescences than expected and tended to visit more mid-and short-styled inflorescences in succession, once they were encountered. Pollinator constancy for morphs did not result from differences in inflorescence production or spatial patchiness among the morphs. Although non-random pollinator visitation to morphs in heterostylous populations could potentially affect mating and hence morph frequencies, the observed visitation patterns in this study do not provide evidence that pollinators play a major role in influencing floral morph frequencies.     

Shell colour polymorphism in a mangrove snail Littorina sp. (Prosobranchia: Littorinidae)

Shell colour polymorphism was examined in populations of a mangrove snail Littorina sp. in Queensland, Australia. Three morphs were recognized, yellow, red and brown, and morph frequencies varied both between widely spaced populations and between islands less than 1 km apart. Morph frequencies also varied with time of year. There was a relationship between shell colour and position on the tree, with yellow snails more often occurring amongst the foliage and brown snails more often on trunks and branches. In some populations yellow snails appeared to survive better than the other morphs, while in other populations there was no difference. The evidence for the maintenance of the polymorphism by natural selection is discussed.     

Site selection in a polymorphic mangrove snail

Three species of littorinid snail live on mangroves of the genus Rhizophora on the north coast of Papua New Guinea. They have recently been placed in the genus Littoraria. The species are L. scabra and L. intermedia , which are monomorphic and cryptic and live on bark, and L. pallescens which is polymorphic and lives in foliage. Littoraria intermedia was found mostly on mangroves facing on to partially fresh water, while L. scabra occurred only at sites facing on to the sea; L. pallescens occurs at both types of site. When kept together in plastic containers, L. pallescens shows a greater propensity to climb than the other two species. The morphs of L. pallescens are principally yellow, orange or dark. Orange is present at a frequency of 0–9% and yellow at 10–50%. There is little evidence of geographical variation in frequency over 80 km of coast, although one central locality has a significantly higher frequency of yellow than sites to the north or south of it. Throughout the tropics where the species occurs, orange is the rarest and yellow usually the next rarest morph. Larger individuals are on average higher in the foliage than smaller ones, and there is a higher frequency of yellows among larger shells than smaller ones. As the air temperature increases, a progressively larger fraction of a sample was found to be on the under-surfaces of leaves rather than on the upper epidermis or on flower or leaf buds. When over 30% of a sample is situated on lower surfaces there is a significant excess of yellows and a deficiency of darks on the upper surfaces. This indicates that individuals selectively take up different positions depending on morph colour, resulting in a degree of optimal background matching. Such a choice of location may be of direct relevance to the maintenance of the polymorphism. A difference in morph frequency between the same sites examined in 1979 and 1984 is interpreted as being a result of site selectivity.     

Ontogeny of sexual dimorphism and phenotypic integration in heritable morphs

In this study we investigated the developmental basis of adult phenotypes in a non-model organism, a polymorphic damselfly (Ischnura elegans) with three female colour morphs. This polymorphic species presents an ideal opportunity to study intraspecific variation in growth trajectories, morphological variation in size and shape during the course of ontogeny, and to relate these juvenile differences to the phenotypic differences of the discrete adult phenotypes; the two sexes and the three female morphs. We raised larvae of different families in individual enclosures in the laboratory, and traced morphological changes during the course of ontogeny. We used principal components analysis to examine the effects of Sex, Maternal morph, and Own morph on body size and body shape. We also investigated the larval fitness consequences of variation in size and shape by relating these factors to emergence success. Females grew faster than males and were larger as adults, and there was sexual dimorphism in body shape in both larval and adult stages. There were also significant effects of both maternal morph and own morph on growth rate and body shape in the larval stage. There were significant differences in body shape, but not body size, between the adult female morphs, indicating phenotypic integration between colour, melanin patterning, and body shape. Individuals that emerged successfully grew faster and had different body shape in the larval stage, indicating internal (non-ecological) selection on larval morphology. Overall, morphological differences between individuals at the larval stage carried over to the adult stage. Thus, selection in the larval stage can potentially result in correlated responses in adult phenotypes and vice versa.     

Frequency- and density-dependent sexual selection in natural populations of Galician Littorina saxatilis Olivi

Galician exposed shore populations of the direct developing periwinkle Littorina saxatilis are strikingly polymorphic, with an ornamented and banded upper shore form and a smooth and unbanded lower shore form. Intermediates between the two pure forms occur in a narrow mid shore zone together with the parental forms. We have previously shown that the two pure forms share the same gene pool but that mating between them is non-random. This is due to a non-random microdistribution in the zone of overlap, and also to assortative mating. In this study we present data which show that intermediate (hybrid) females mate less often than pure females in micropatches dominated by either of the pure forms, but not in micropatches in which the two pure forms are equally common. Thus, sexual fitness in intermediate females depends on the frequency of both pure morphs. Furthermore, sexual selection against intermediate females also varies with the densities of snails within each micro patch. The biological mechanisms which may explain this particular reduction of female hybrid fitness are discussed. Assortative mating between the pure morphs is sometimes almost complete, while both morphs do not mate the intermediates assortatively. In the light of this, sexual selection against intermediate females may contribute considerably to restrict gene flow between the pure forms.     

Trade‐offs in female signal apparency to males offer alternative anti‐harassment strategies for colour polymorphic females

Colour polymorphisms are known to influence receiver behaviour, but how they affect a receiver's ability to detect and recognize individuals in nature is usually unknown. I hypothesized that polymorphic female damselflies represent an evolutionary stable strategy, maintained by trade‐offs between the relative apparency of morphs to male receivers. Using field experiments on Enallagma hageni and focal studies of E. hageni and Enallagma boreale, I tested for the first time the predictions that (i) green heteromorphs and blue andromorphs gain differential protection from sexual harassment via background crypsis and sexual mimicry, respectively, and (ii) female morphs behaviourally optimize their signal apparency to mate‐searching males. First, based on male reactions elicited by females, against a high‐contrast background, the two morphs did not differ in being detected by males, and once detected, they did not differ in being recognized (eliciting sexual reactions). However, on green ferns, heteromorphs were less likely to be detected (elicited only fly‐bys) than andromorphs, but once detected, the morphs did not differ in being recognized. In contrast, when perched on a dowel with two male signal distractors, andromorphs were detected less often, and once detected, they were recognized less often than heteromorphs. Second, in fields where females foraged, andromorphs perched higher on vegetation than heteromorphs and were more often in the vicinity of males. Neither harassment rates nor evasive behaviours differed between morphs. Males aggregated in high density near shore where solitary females were rare. Equilibrium frequencies of these and other colour morphs should reflect the relative ease with which receivers detect and recognize them in the context where they are encountered.     

Adaptation to a steep environmental gradient and an associated barrier to gene exchange in Littorina saxatilis

Steep environmental gradients offer important opportunities to study the interaction between natural selection and gene flow. Allele frequency clines are expected to form at loci under selection, but unlinked neutral alleles may pass easily across these clines unless a generalized barrier evolves. Here we consider the distribution of forms of the intertidal gastropod Littorina saxatilis, analyzing shell shape and amplified fragment length polymorphism (AFLP) loci on two rocky shores in Britain. On the basis of previous work, the AFLP loci were divided into differentiated and undifferentiated groups. On both shores, we have shown a sharp cline in allele frequencies between the two morphs for differentiated AFLP loci. This is coincident with a habitat transition on the shore where the two habitats (cliff and boulder field) are immediately contiguous. The allele frequency clines coincide with a cline in shell morphology. In the middle of the cline, linkage disequilibrium for the differentiated loci rises in accordance with expectation. The clines are extremely narrow relative to dispersal, probably as a result of both strong selection and habitat choice. An increase in F(ST) for undifferentiated AFLPs between morphs, relative to within-morph comparisons, is consistent with there being a general barrier to gene flow across the contact zone. These features are consistent either with an episode of allopatric divergence followed by secondary contact or with primary, nonallopatric divergence. Further data will be needed to distinguish between these alternatives.     

Correlated morphological and colour differences among females of the damselfly Ischnura elegans

Abstract 1. The female‐limited colour polymorphic damselfly Ischnura elegans has proven to be an interesting study organism both as an example of female sexual polymorphism, and in the context of the evolution of colour polymorphism, as a model of speciation processes. 2. Previous research suggests the existence of correlations between colour morph and other phenotypic traits, and the different female morphs in I. elegans may be pursuing alternative phenotypically integrated strategies. However, previous research on morphological differences in southern Swedish individuals of this species was only carried out on laboratory‐raised offspring from a single population, leaving open the question of how widespread such differences are. 3. The present study therefore analysed multi‐generational data from 12 populations, investigating morphological differences between the female morphs in the field, differences in the pattern of phenotypic integration between morphs, and quantified selection on morphological traits. 4. It was found that consistent morphological differences indeed existed between the morphs across populations, confirming that the previously observed differences were not simply a laboratory artefact. It was also found, somewhat surprisingly, that despite the existence of sexual dimorphism in body size and shape, patterns of phenotypic integration differed most between the morphs and not between the sexes. Finally, linear selection gradients showed that female morphology affected fecundity differently between the morphs. 5. We discuss the relevance of these results to the male mimicry hypothesis and to the existence of potential ecological differences between the morphs.     

Polymorphic shell banding in the dog-whelk, Nucella lapillus (Mollusca)

The shell of the Common dog-whelk (Nucella lapillus (L.)) is white and unbanded at most places around the British Isles. However, high frequencies of banding occur on the Buchan coast, around Anglesey and the Menai Straits, on the Cower Peninsula, around the Devon–Somerset border in the Bristol Channel, and especially on the north Cornish coast (reaching a peak between Newquay and Padstow). The frequency of banding is significantly less in older than younger whelks in the same locality, and this change is uncorrelated with the selection against shell shape variation that takes place on exposed shores. It is concluded that banding is a pleiotropic manifestation of physiological variation, and that a study of such variation in different morphs could indicate the importance of different physiological stresses at different stages of the life history of N. lapillus.     

Natural selection for apostasy and crypsis acting on the shell colour polymorphism of a mangrove snail, Littoraria filosa (Sowerby) (Gastropoda: Littorinidae)

Littoraria filosa (Sowerby) is a member of the L. scabra group, found amongst the foliage of mangrove trees in northern Australia. The colour of the shell is polymorphic, showing two discrete ground colours, either yellow or orange-pink, with a variable degree of superimposed brown patterning. At a site on Magnetic Island, northern Queensland, colour frequencies of small snails were similar on different backgrounds. Amongst larger shells yellows were more frequent on Avicennia trees with abundant foliage, and browns on relatively bare trees, suggesting that visual selection for crypsis occurred. There was no evidence of substrate selection by the morphs. Yellow shells were cooler than brown shells, but differences in colour frequencies on sunny and shaded trees, and at different seasons, did not suggest climatic selection. By manipulating the colour frequencies of subpopulations of small snails isolated on individual trees, it was shown that the disappearance of yellow and brown shells was frequency-dependent. This result is consistent with hypotheses of mimicry of background elements by the morphs and of apostatic selection by unknown predators. Only the latter can account for the persistence of the highly conspicuous pink morph at a low frequency.     

PHYSIOLOGICAL STRESS AND COLOR POLYMORPHISM IN THE INTERTIDAL SNAIL NUCELLA LAPILLUS

The intertidal snail Nucella lapillus exhibits considerable variation in shell color both within and between populations differentially exposed to wave action. Populations from high-wave-energy shores tended to be highly polymorphic and were dominated by pigmented morphs (especially brown), while those at more sheltered locations exhibited less polymorphism and were predominantly white. Field and laboratory experiments were conducted to determine the role of physiological stress and selective predation in maintaining the observed distribution of color morphs. The results demonstrated that 1) physiological stress from high temperature and desiccation during periods of tidal emersion was greater on protected shores, 2) under similar natural conditions, brown morphs heated up faster, attained higher temperatures, desiccated more rapidly, and suffered greater mortality than did white morphs, and 3) when pairs of brown and white morphs were tethered intertidally there was virtually no mortality of either morph on the exposed shore or in shaded microhabitats on the protected shore, but brown morphs suffered much greater mortality in sunny microhabitats on the protected shore. These findings demonstrate that the interpopulation variation in shell color of N. lapillus is in part a response to a selective gradient in physiological stress. Selection for crypsis by visually hunting predators did not appear to play a prominent role; however, only adults were considered, and the predation experiments were conducted in the fall before shorebirds that prey on whelks had arrived from their summer feeding grounds. Further experimentation to quantify the effects of visual predators such as birds and fish, particularly on juvenile snails, is necessary to assess adequately the importance of predation.