Differential vulnerability to predation in two sympatric whelks is mediated by juvenile traits

In many taxa, initial differences in offspring size play an important role in mediating subsequent performance; however, the consequences of interspecific variation in size for the performance of co‐occurring taxa have been rarely examined. We used the whelks Cominella virgata and C. maculosa, which co‐occur on rocky shores throughout their life cycles, to examine the vulnerability of early life‐stages to native predators under controlled laboratory conditions. Among all the predators evaluated (the cushion sea star Patiriella spp., the olive rockfish Acanthoclinus fuscus, the oyster borer snail Haustrum scobina, the smooth shore crab Cyclograpsus lavauxi, and the pebble crab Heterozius rotundifrons), hatchlings of both species (C. virgata: ~3 mm shell length [SL] and C. maculosa: ~1.5 mm SL) were especially vulnerable to the smooth shore crab Cy. lavauxi, the only potential predator in which mortality was greater than in the control treatment. Small shore crabs (~8 mm carapace width [CW]) were unable to eat hatchlings of either whelk species, whereas medium and large shore crabs (~12 and ~18 mm CW, respectively) consumed hatchlings of both prey species. Hatchlings of C. virgata were less vulnerable to predation by medium crabs than large ones, and those of C. maculosa were equally vulnerable to both sizes of crabs. In hatchlings of both prey species, shell length and shell thickness increased over time. Two months after hatching, only individuals of C. virgata had reached a size refuge from predation. Our results show that interspecific vulnerability to predators can be mitigated by larger sizes and thicker shells at hatching; nonetheless, our results also suggest that other species‐specific factors, such as juvenile growth rate, may also play key roles in determining the vulnerability of hatchling and juvenile snails to shell‐crushing predators.     

Predation by small mobile aquatic predators regulates populations of the intertidal limpet Patella vulgata (L.)

Highly mobile aquatic predators are known to forage in the intertidal during periods of immersion. There is limited quantitative information, however, on the extent to which these predators influence the abundance of grazing molluscs which are known to have a key role in structuring intertidal assemblages. Our preliminary video observations revealed that crabs and small fish were abundant on shores in southwest England during high-tide. We then used manipulative experiments to quantify the effect of small mobile aquatic predators on the abundance of limpets (Patella vulgata L.). On the lower shore at two moderately sheltered rocky shores three treatments were established: complete cage, partial cage (cage control) and uncaged (natural condition). The complete cages excluded all predators. The partial cage treatment allowed full access to small predators and the uncaged treatment allowed access to all predators. After two months, limpet abundance in uncaged and partial cage treatments had declined by around 50% compared to the complete cage treatment. Population structure also changed with survival of larger individuals being greater than smaller individuals in the open and partial cage treatments compared to the complete cage treatment. The effects of excluding predators were consistent at small (meters) and large spatial scales (kilometres) and hence, it would appear that the outcomes of our research are generally applicable to similar shores in the region.To explore the mechanism behind the differential effects of predators according to prey size, we compared the detachment force required to remove limpets of differing sizes from the shore. This was around four times greater for larger individuals than for smaller ones indicating that smaller limpets were more vulnerable to predation. These effects were also consistent between locations. Subsequent laboratory observations showed that the crabs Carcinus maenas (L.), Necora puber (L.) and Cancer pagurus (L.) which are locally abundant predators of limpets, had differing handling behaviour but were all highly efficient at removing limpets from substratum. Hence, shell width and attachment force appeared to be critical factors influencing the vulnerability of limpets to predation by these crabs. Limpets are known to control the abundance of macroalgae on shores in the North-east Atlantic and so our conclusions about the role of mobile predators in regulating the abundance of these grazers are important to our broader understanding of the ecology of these shores.     

ASYMMETRICAL DEVELOPMENTAL PLASTICITY IN AN INTERTIDAL SNAIL

Field and laboratory experiments were used to investigate the variation and phenotypic plasticity in the adhesive abilities of the intertidal snail Nucella lapillus between high- and low-wave-energy environments. Whelks from an exposed coast produced a larger pedal surface area and were more resistant to dislodgement than were similar-sized individuals from a protected shore. Tenacity (g/cm²) was similar between individuals from exposed and protected shores, indicating that variation in resisting dislodgement was solely a function of pedal surface area. Whelks from exposed and protected shores did not differ in pedal surface area as they emerged from egg capsules or when reared in the laboratory under uniform conditions, suggesting that variation between populations does not represent genetic differentiation. Individuals from high-and low-wave-energy environments reared intertidally produced a larger pedal surface area than did those reared in the laboratory. The extent to which pedal surface area increased corresponded to the intensity of wave action. These findings suggest that pedal surface area is a highly plastic character modulated or induced by the water turbulence accompanying breaking waves. A reciprocal-transplant experiment confirmed this notion but revealed an asymmetry in the plasticity. Snails from the protected site transplanted to an exposed shore formed a much larger pedal surface area than did controls reared on the protected shore. In contrast, whelks from a wave-swept shore transplanted to a protected shore differed little from their controls reared on the exposed shore. The asymmetric response parallels a possible asymmetry in the risks of acclimating to a temporally unpredictable environmental cue, such as wave action.     

Effect of crab effluent and scent of damaged conspecifics on feeding,growth, and shell morphology of the Atlantic dogwhelk Nucella lapillus (L.)

Juvenile Nucella lapillus of two different shell phenotypes, exposed shore and protected shore, were maintained in running seawater under each of three experimental conditions for 94 d: a) laboratory control, b) exposed to the effluent of crabs (Cancer pagurus) fed frozen fish (fish-crab), and c) exposed to the effluent of crabs fed live conspecific snails (snail-crab). Rates of barnacle consumption and rates of body weight change varied significantly between phenotypes and among experimental conditions. Individuals from the protected-shore consumed consistently fewer barnacles and grew consistently less than those from the exposed shore. Body weight increases in the fish-crab treatments were from 25 to 50% less than those in the controls and body weights in the snail-crab treatment either did not change or actually decreased. The perceived risk of predation thus appears to have a dramatic effect on the rates of feeding and growth of N. lapillus.At the end of the experiment, size-adjusted final shell weights for both phenotypes were consistently higher than controls (no crab) in both the fish-crab and snail-crab treatments. In addition, apertural tooth height, thickness of the lip, and retractability (i.e. the extent to which a snail could withdraw into its shell), with few exceptions all varied in an adaptive manner in response to the various risk treatments. Similar changes in the shell form of starved snails exposed to the same stimuli suggest very strongly that the morphological responses of both phenotypes were not just due to differences in rates of growth. These differences, at least in part, represented a direct cueing of the shell form of Nucella lapillus to differences in the perceived risk of predation. Somewhat surprisingly, the extent of phenotypic plasticity appeared to differ between the populations examined. Both field and laboratory evidence suggest that the exposed-shore population was much more labile morphologically than the protected-shore population.In many instances, particularly among starved snails, the development of antipredatory shell traits was greater in the fish-crab treatment than in the snail-crab treatment. Because the scent of crabs was present in both treatments, these results suggest a) that, at the frequency/concentration used in the experiments, the scent of damaged conspecifics may have been a supernormal stimulus and b) that the morphological response in these treatments might have been greater if the stimulus had been provided at a lower level.     

Aspects of the ecology of an exposed shore population of dogwhelks Nucella lapillus (L.)

Summary Up to the third year of life Nucella lapillus (L.) could be aged in the laboratory and in the field using shell characters. Monthly counts along transects on an exposed rocky shore have shown that the age groups behave differently, dogwhelks in their first year migrating upshore, and returning to lower levels in the second year. Adults (three years or older) lived and laid egg capsules in the low shore. The most favourable situation for subsequent hatching of these capsules was where they were permanently submerged in sea water.The first requirement of recently hatched young was probably protection from water movement, but they soon gegan feeding on small mussels. The shell height attained by the end of the first winter was related to winter temperatures, but the main growing season was June to November. Adult size was attained in two years, after which the individuals did not usually grow, but those which did grow during the third or succeeding years may have been nonbreeders or have been castrated by the trematode Parorchis acanthus.The age structure of the population suggested that mortality rate decreased with age, and this was confirmed by calculating annual mortality rates for the first three years of life. Heavy mortality in the low shore during the first winter was due mainly to predation by purple sandpipers. In succeeding winters mortality was low, and predation, the main mortality factor, was restricted to the summer months. At this time Cancer pagurus was the most voracious predator, but it was active mainly in the low shore, and the upshore distribution of second year dogwhelks, which were preferred to other age groups, age groups also led to a more economical use of environmental resources, namely food, winter aggregation sites and breeding sites.     

Dislodgement force and shell morphology vary according to wave exposure in a tropical gastropod (Cittarium pica)

Wave exposure has strong influences on population density, morphology and behaviour of intertidal species in temperate zones, but little is known about how intertidal organisms in tropical regions respond to gradients in wave exposure. We tested whether dislodgement force and shell shape of a tropical gastropod, Cittarium pica, differs among shores that vary in wave exposure. After adjusting for body size, we found that C. pica from exposed shores required greater dislodgement force to remove them from the shore, had slightly larger opercula (the closure to the shell aperture), and were slightly squatter in shape (reduced in shell height relative to shell width) than C. pica from sheltered shores. These morphological adjustments are consistent with those observed in temperate gastropods, which are argued to represent adaptive responses to the risk of mortality associated with dislodgement.     

Chitons’ apparent camouflage does not reduce predation by green crabs Carcinus maenas

Chitons are very common molluscs on European rocky shores. They are common prey of fish and crabs and often display several colour morphs within a given habitat. Predation is one of the potential mechanisms accounting for chiton colour polymorphism. The colour variation is considered to provide a camouflage protection through a match with the substratum surface typology. However, the effectiveness of chiton polymorphism as a predation defence requires further investigation. Previously we found a relationship between chiton colour morphs and substrate characteristics, with chitons most commonly found on substrates that were of similar colour to their shells. Here, we examined whether each morph displayed an active choice for matching the substratum. Next, we assessed if the predation success of the intertidal common crab Carcinus maenas varied significantly with the absence/presence of an apparent camouflage effect created between the chiton colour morph and the substratum type. The present study indicates that chiton colour morphs probably actively choose substratum types where they blend in. Carcinus maenas was able to prey on all Lepidochitona cinereus colour morphs, regardless of the substrate camouflage effect. Surprisingly, the predation frequency was higher on camouflaged chitons than on contrasting chitons. It was concluded that chiton camouflage is probably not a defence mechanism against predation by the crab C. maenas, and that chiton colour polymorphism is probably promoted by other, more visual predators.     

Natural selection on inflorescence color polymorphisms in wild Protea populations: The role of pollinators, seed predators, and intertrait correlations

? Premise of the study: The processes maintaining flower color polymorphisms have long been of evolutionary interest. Mechanistic explanations include selection through pollinators, antagonists, local environments, drift, and pleiotropic effects. We examined the maintenance of inflorescence color polymorphisms in the genus Protea (Proteaceae) of South Africa, in which ～40% of species contain different color morphs. ? Methods: We studied 10 populations of four bird-pollinated Protea species and compared adult performance, floral and leaf morphology, vegetative pigmentation, germination, and seedling survival between co-occurring pink and white morphs. We also tested for differences in pollination success and pre-dispersal seed predation. ? Key results: White morphs produced seeds 10% heavier and 3.5 times more likely to germinate, which all else being equal, should fuel positive selection on white. In one studied population per species, however, white morphs were more susceptible to seed predation by endophagous larvae. Pollinators had no morph-specific effects on female fecundity, as measured by amount or probability of seed set. Differences in stem color indicated that white morphs produced smaller quantities of pigment and associated compounds throughout, possibly explaining their higher seed palatability. ? Conclusions: Our findings suggest a mechanism for some white protea polymorphisms: deleterious pleiotropic effects on pink morphs are occasionally offset by reduced losses to seed-eating larvae. Because trends were repeated across species, we suggest that similar processes may also occur in other proteas, placing a new emphasis on seed predators for influencing some of South Africa's amazing floral diversity.     

Differences in predatory pressure on terrestrial snails by birds and mammals

The evolution of shell polymorphism in terrestrial snails is a classic textbook example of the effect of natural selection in which avian and mammalian predation represents an important selective force on gene frequency. However, many questions about predation remain unclear, especially in the case of mammals. We collected 2000 specimens from eight terrestrial gastropod species to investigate the predation pressure exerted by birds and mice on snails. We found evidence of avian and mammalian predation in 26.5% and 36.8% of the shells. Both birds and mammals were selective with respect to snail species, size and morphs. Birds preferred the brown-lipped banded snail Cepaea nemoralis (L.) and mice preferred the burgundy snail Helix pomatia L. Mice avoided pink mid-banded C. nemoralis and preferred brown mid-banded morphs, which were neglected by birds. In contrast to mice, birds chose larger individuals. Significant differences in their predatory pressure can influence the evolution and maintenance of shell size and polymorphism of shell colouration in snails.     

Perch color preference in juvenile green tree pythons,Chondropython viridis

Green tree pythons, Chondropython viridis, are polymorphic for color as juveniles, commonly being primarily yellow or brown until becoming mostly green at about 1 year of age. We tested the hypothesis that the different morphs arose as a result of selection for differential background matching, yellow morphs selecting light-colored backgrounds, and brown morphs selecting dark-colored backgrounds. Twelve yellow and eight brown morphs were placed repeatedly in individual testing enclosures and allowed to choose between black and white or yellow and brown halves of a t-perch. Trials showed that both color morphs preferred dark over light perches. We tentatively suggest that individuals chose dark-colored perches for purposes of concealment. © 1994 Wiley-Liss, Inc.     

The ecology of Mytilus edulis L. (Lamellibranchiata) on exposed rocky shores

Summary Growth studies in M. edulis L. have shown that rates vary considerably according to age, size and environmental conditions. This may in part be attributed to its sessile habit, being unable to move away from the variable external conditions.The use of modal length frequency distributions is somewhat limited, since with three or more year groups represented, the growth of the majority of the population is so slow that individual year classes lose their identity. Growth boxes, containing marked animals of different ages, and set out in a variety of habitats gave information regarding local and seasonal growth rates. Disturbance rings were shown to be annual, and from them growth curves have been constructed.Growth is particularly seasonal, little or none occurring during the winter. Growth rates varied considerably with a variety of environmental factors (both biotic and physical) and some of these are discussed. Variable individual growth rates, together with slow growth of the majority of animals in mixed populations, are perhaps the major factors in producing population structures typical of this species on open shores.Survivorship curves for mussels in a variety of habitats have been constructed by following the survival of groups of marked animals. Whilst high mortalities occurred in the mid and low shore, survival in the upper shore in the absence of major predators, was greatly enhanced, resulting in established populations of considerable age. Periods of maximum mortality during spring and summer could be correlated with the abundance of major predators.The almost cosmopolitan distribution of M. edulis in the N. hemisphere has been made possible by virtue of its high reproductive capacity, successful larval dispersion and wide tolerance of environmental conditions, Its patchy and apparently erratic distribution both from one shore to annother and even on the same shore, is greatly influenced by the local and seasonal abundance of major predators. Whilst the upward extension of mussels is prevented, ultimately, by physical factors (e.g. temperature, dessication), its lower limits (and absence from the shallow sub littoral on many shores) are governed essentially by predators. The upward extension of many predators such as crabs or starfish, resulting in locally intense predation, may in turn be partly influenced by the actual topography of the shore itself.     

The role of invertebrates in seedling establishment in a heterocarpic plant,Atriplex sagittata

Seed heteromorphism is a marked character of many Chenopodioideae (Amaranthaceae). Seed morphs differ in dormancy, germination and seedling biology, but differences in their predation have not yet been studied. Atriplex sagittata produces small black dormant and large brown non‐dormant seeds. In this study, the timing of seed release and seedling establishment were ascertained, and their consumption by invertebrates (carabids, isopods and slugs) was studied. Seeds dispersed in the autumn passed the winter on the ground surface, protected from invertebrate predation by low temperatures. In the following vegetative season, ungerminated black seeds exposed to predation on ground surface were preferred by a large carabid species, Pseudoophonus rufipes. Some black and all brown seeds escaped predation by germinating in early spring. The seedlings were little endangered by carabids and isopods but were preferred by an invasive slug, Arion vulgaris, the feeding of which can exterminate seedlings at places in which slugs are abundant. Invertebrate predation is important factor of seed and seedling mortality of A. sagittata and seed heteromorphism modifies its intensity and timing.     

More Ornamented Males Exhibit Increased Predation Risk and Antipredatory Escapes,but not Greater Mortality

Secondary sexual traits not only confer benefits to their bearer through increased mate acquisition, but may also have inherent costs, including the attraction of predators. Here, we examined the relationship between conspicuous secondary sexual traits and predation costs using two male morphs of Schizocosa wolf spiders: brush‐legged and non‐ornamented. In the field, we ran two predation experiments using artificial enclosures to directly test mortality costs of predation on the two male morphs. Using a natural predator, a larger wolf spider in the genus Hogna, we found no difference in predation on brush‐legged vs. non‐ornamented males. However, predation was depends on environmental conditions. More individuals were preyed upon at night (vs. during the day) and on rock litter (vs. leaf litter), but the two male morphs were preyed upon equally to each other across environmental treatments. A laboratory experiment incorporated staged interactions between a single predator (Hogna) and each male morph to examine finer details of predation events. Again, we found no differential mortality between brush‐legged and non‐ornamented males. However, brush‐legged males were attacked sooner and were more likely to escape the attack. Our results show an association between sexual ornamentation and predation risk as well as escape behavior.     

The adaptive significance of aggregation behaviour in the dogwhelk Nucella lapillus (L.)

Summary Three kinds of aggregation behaviour were observed in an exposed shore population of Nucella lapillus. Aggregations on the open rock surface during the summer protected dogwhelks from water movement, and were not found on shores where the topography conferred protection. Feeding occurred mainly within these aggregations, probably because individuals experienced less disturbance there than when isolated. Physical contact was important in holding animals together.Winter and pre-breeding aggregations were usually found in clefts or pools. All age groups formed winter aggregations, but those of immature dogwhelks were not as permanent as those of adults, and the latter merged with the prebreeding aggregations. Winter aggregations protected dogwhelks from dislodgment when their ability to regain a foothold was reduced by low temperatures, while pre-breeding aggregations brought the sexes together for fertilization, but the permanence of adult winter aggregations suggested that reproductive activity may have been occurring within them.     

Aposematic coloration does not deter corallivory by fish on the coral <Emphasis Type="Italic">Montastraea cavernosa</Emphasis>

Predation on corals by visual predators is a significant source of partial or total mortality on coral reefs, and corals have evolved strategies, including chemical defenses, to deter predation. One mechanism that organisms use to communicate the presence of chemical defenses is aposematic coloration, or the display of bright coloration as a warning to visual predators such as fish. Corals exhibit multiple colors, and it has been hypothesized that one role for this variability in coloration is as an aposematic warning of adverse palatability. Here, we test green and orange color morphs of the Caribbean coral Montastraea cavernosa for the presence of chemical defenses and whether their differences in coloration elicited different feeding responses. While M. cavernosa is chemically defended, there is no difference in feeding deterrence between color morphs; thus, the different color morphs of this coral species do not appear to represent an example of aposematic coloration.     

Biogeographic variation in behavioral and morphological responses to predation risk

The expression of prey antipredator defenses is often related to ambient consumer pressure, and prey express greater defenses under intense consumer pressure. Predation is generally greater at lower latitudes, and antipredator defenses often display a biogeographic pattern. Predation pressure may also vary significantly between habitats within latitudes, making biogeographic patterns difficult to distinguish. Furthermore, invasive predators may also influence the expression of prey defenses in ecological time. The purpose of this study was to determine how these factors influence the strength of antipredator responses. To assess patterns in prey antipredator defenses based upon geographic range (north vs. south), habitat type (wave-protected vs. wave-exposed shores), and invasive predators, we examined how native rock (Cancer irroratus) and invasive green (Carcinus maenas) crab predators influence the behavioral and morphological defenses of dogwhelk (Nucella lapillus) prey from habitats that differ in wave exposure across an ~230 km range within the Gulf of Maine. The expression of behavioral and morphological antipredatory responses varied according to wave exposure, geographic location, and predator species. Dogwhelks from areas with an established history with green crabs exhibited the largest behavioral and morphological antipredator responses to green crabs. Dogwhelk behavioral responses to rock crabs did not vary between habitats or geographic regions, although morphological responses were greater further south where predation pressure was greatest. These findings suggest that dogwhelk responses to invasive and native predators vary according to geographic location and habitat, and are strongly affected by ambient predation pressure due to the invasion history of an exotic predator.     

RELATIONSHIPS BETWEEN SHELL SHAPE, WATER RESERVES, SURVIVAL AND GROWTH OF HIGHSHORE LITTORINIDS UNDER EXPERIMENTAL CONDITIONS IN NEW SOUTH WALES, AUSTRALIA

The shell morphologies of the highshore littorinids, Littorinaunifasciata Gray and Nodilittorina pyramidalis (Quoy & Gaimard)have previously been shown to vary at a variety of spatial scales,including among replicate sites at the same height, from heightto height and from shore to shore. In this study, the relationshipsbetween morphology of the shell, the reserves of water heldwithin the shell, the size of the foot and survival on differentshores and rates of growth in different habitats were examinedfor L. unifasciata and, to a lesser extent, N. pyramidalis.Reserves of water were not consistently related to size or shapeof the shell, but did increase as relative weight of shell increased.This may be due to the relatively smaller body providing moreinternal volume for extra-corporeal water. Water reserves andthe amount of free water held in the shell were also not relatedto loss of water or survival during extended periods of emersion.Although the shape of the shell on sheltered and exposed shoreswas correlated with size of the foot, with the snails on anexposed shore having larger apertures and feet than those ona sheltered shore, transplant experiments did not show differentialmortality between morphs from the different shores. All translocatedand transplanted snails disappeared from the exposed shore ata greater rate than from the sheltered shore, but this was probablydue to the snails dispersing out of the experimental areas ratherthan due to mortality. Therefore, many of the large-scale modelsthat have previously been used to describe patterns of shellshape in intertidal gastropods do not appear to be importantin these highshore littorinids. Finally, field experiments ongrowth of juvenile L. unifasciata indicated that rate of growth,largely governed by opportunity to feed rather than type andquantity of food, is the most likely explanation for the small-and large-scale patterns of shell shape that have been previouslydescribed in this species. (Received 22 October 1996; accepted 19 February 1997)     

The white colour of the Ostrich ( Struthio camelus ) egg is a trade-off between predation and overheating

Most ground nesters lay pigmented eggs, and egg pigmentation generally matches the environment. Pigmentation of eggs has evolved as a protective device against predation, but dark-pigmented eggs can be susceptible to overheating when exposed to solar radiation. The Ostrich (Struthio camelus) lays white eggs that are unattended for the first few weeks before incubation, and are quite visible to predators. To evaluate the effect of colour on the surface and core temperatures, we painted some Ostrich eggs dark brown or white, and left some unpainted (control), and exposed all of them directly to the sun during the day. The surface and core temperatures of brown eggs were significantly higher than those of the white-painted and control eggs. In addition, the core temperature of brown eggs exceeded 37.5°C, which is the temperature at which embryo mortality starts to increase. In a second experiment, we placed eggs (brown-painted and control) in various types of vegetation to study their visibility to an observer walking towards them. The white eggs were discovered from a significantly longer distance than the brown eggs, indicating that the predation risk may be much higher for white eggs. The results thus suggest that white eggs minimise overheating and allow the Ostrich to leave its eggs unattended before incubation starts, but they are more susceptible to predation.