Life history of Littorina scutulata and L. plena, sibling gastropod species with planktotrophic larvae

Abstract. The intertidal, sibling species Littorina scutulata and L. plena (Gastropoda, Proso‐branchia) are sympatric throughout most of their ranges along the Pacific coast of North America. Both species release disc‐shaped, planktonic egg capsules from which planktotrophic veliger larvae hatch. Here I review existing data and present new observations on these species' life history, including age at first reproduction, spawning season, maximum fecundity rates, capsule morphology, egg size and number, pre‐hatching development, larval growth at three food concentrations, potential settlement cues, planktonic period, and protoconch size. Previous classification of egg capsule morphologies used to distinguish the species is inaccurate; instead, capsules can be categorized into three types of which each species may produce two. Females of L. scutulata produced capsules with either two rims of unequal diameter or one rim, while females of L. plena produced capsules with one rim or two rims of nearly equal diameter. Females of each species spawned sporadically from early spring to early fall in Puget Sound. Larvae of L. plena hatched one day earlier than those of L. scutulata, and both species grew fastest in the laboratory at intermediate food concentrations. Larvae metamorphosed in the presence of a variety of materials collected from their adult habitat, including conspecific adults, algae, rocks, and barnacle tests. This is the first report of planktotrophic larvae in this genus metamorphosing in the laboratory. The total planktonic period of 8 larvae of L. scutulata raised in the laboratory was 37–70 days, and a single larva of L. plena metamorphosed after 62 days. Protoconch diameter of shells collected from the field was 256–436 μm and did not differ significantly between the species. Previous allozyme and mitochondrial DNA work has suggested high levels of genetic variability in both species and greater genetic population structure in L. plena, despite the long spawning season and long‐lived larvae in both species. The interspecific life history differences described here appear insufficient to produce consistent differences in gene flow patterns.     

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)     

Ecology of the intertidal snail Littorina acutispira Smith

Littorina acutispira Smith, a minute gastropod of < 3 mm shell height, lives at great densities in pools and on rock-surfaces at the highest levels of sea-shores in New South Wales. Populations from pools and dry areas were sampled on two shores for 18 months to investigate seasonal changes in density, size-structure, rates of growth and reproductive biology of the snails. Densities of snails increased between February and May, due to an influx of juveniles, and then declined until the following February, when they increased again. The decrease in density was due to the death of the largest snails at the end of summer, and the mortality of medium-sized snails between June and January. Longevity was estimated as 1–2 yr, but most individuals died by ≈ 16 months from settlement on the shore. Newly-settled snails grew to merge in size with those of the previous year's population by winter. During the summer months, the rate of growth of snails from a sheltered shore was greater than that of snails on a shore exposed to wave-action. Laboratory experiments revealed that this could be attributed to the presence of better quality food, or food in greater abundance on the sheltered shore, compared with the exposed shore.During winter months, but not during the summer, snails from dry areas grew more slowly than those from pools. An experiment demonstrated that some snails from dry areas might be able to compensate for reduced periods of feeding by being able to feed faster when submersed. This could not explain the differences in natural rates of growth.L. acutispira bred from October–November to March–April. Spawning in the laboratory was greatest during late summer (January to March). The percentage of mature oocytes in the gonads was small in winter and increased in early summer. Among the largest-sized snails, females outnumbered males. Two experiments, on unsexed and pre-sexed snails, demonstrated that the biased sex-ratio of the largest snails was due to faster growth by females.There was a greater density of snails on the exposed shore, which was correlated with the presence of barnacles. When barnacles were removed from experimental areas, the density of the snails declined within 24 h. This suggested that barnacles provided a refuge from wave-shock, rather than shelter from desiccation or high temperature. In laboratory experiments, snails were exposed to higher temperatures and less humidity than they would normally encounter on the shore. There was negligible mortality of small or large snails after 24 h of these conditions.This minute species grows quickly, recruits annually and has a short life-span. This type of life-history is discussed in comparison with similar small species from other habitats.     

Dimorphic migration,growth, and fecundity in a seasonally split population of Littorina brevicula (Mollusca: Gastropoda) on a boulder shore

Littorina brevicula (Philippi) is an abundant herbivorous gastropod species inhabiting intertidal zones of the north-western Pacific and has internal fertilization with planktonic eggs and a larval stage. A population of L. brevicula on a shore in Amakusa, Japan, splits into an upper and a mid shore sub-population during the mating season in winter, while all individuals occurred in the upper zone in summer. Transplant and recapture experiments have shown that snails from each sub-population have a distinct pattern of seasonal migration, suggesting that the snail population consists of two behavioral morphs. To explore the relationship between migration pattern and life history characteristics in the snails of the two migration morphs, monthly quantitative samplings and a mark-release and recapture survey were carried out for the population of L. brevicula over a 1-year period. In autumn, some of the adults migrated downward to the mid shore and formed the lower sub-population from October to April. Migration between the upper and lower sub-populations was minimal from December to February, when L. brevicula copulate actively. The shell growth rates of the snails in the lower zone were slightly higher than those in the upper zone, but the shell width of females was not a good predictor of the number of mature ova they contained. The number of mature ova was not significantly different between the females of the two migration morphs. In addition, the mortality of marked snails did not differ significantly between the upper and lower zones. Thus, no evidence was obtained to indicate a difference in the reproductive fitness between the two migration morphs.     

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.     

The relationship between the lethal freezing temperatures and the amounts of ice formed in the foot muscle of marine snails (Mollusca: Gastropoda).

The differences in the lethal freezing temperatures of the foot muscles of the marine snails used in this study were related to the vertical distributions of the snails on the shore. The muscles of the subtidal species Thais lapillus and Nassarius obsoletus were injured at temperatures that were significantly higher than those of the muscles of the intertidal species Littorina obtusata, Littorina littorea, and Littorina saxatilis. The lethal freezing temperatures also varied among the intertidal species. The foot muscle of the high-intertidal species, L. saxatilis, was injured at a significantly lower temperature than the foot muscles of the low-intertidal species L. obtusata.Calorimetry was used to show that the differences in the lethal freezing temperatures between the subtidal and intertidal snails were related to the amounts of tissue ice formed. The ability of the muscles of the intertidal snails to tolerate lower subfreezing temperatures was associated with an increased tolerance to greater quantities of tissue ice. In contrast, the differences in the lethal freezing temperatures among the intertidal species were independent of the amounts of tissue ice formed. The percentage of water frozen in the muscles of these snails at their respective lethal freezing temperatures were not significantly different and were equal to 82%. Thus, the physiological mechanism responsible for the differences in the lethal freezing temperatures of the muscles of the intertidal snails is associated with an increased tolerance to a factor other than the amounts of tissue ice formed.     

Vertical variation in fecundity of the intertidal gastropodMonodonta labio caused by different growth rates between tidal zones

Monodonta labio (Gastropoda: Trochidae) occurs in a wide tidal zone on a boulder-covered shore in Amakusa, Japan. To investigate sources of variation in reproductive output within a population, the fecundity ofM. labio was estimated. Regressions of gonad egg counts on shell width were calculated from samples collected bi-weekly at three tidal zones: high, mid and low intertidal. Seasonal fluctuations in the regression revealed that individual females spawn at least three times a year. Variation in fecundity between the three tidal zones was not detected at any time in standard 12 mm snails. Due to the high growth rate in the low zone during the reproductive season, annual fecundity in the low zone was larger than that in the high and mid zones. Thus, tidal zone variation in fecundity ofM. labio was a result of growth variation between tidal zones.     

LIFE CYCLE AND HABITAT SHIFT OF THE TROCHID SNAIL DILOMA SUAVIS (PHILIPPI) WITHIN INTERTIDAL MUSSEL ZONES

The life cycle of the trochid snail Diloma suavis (Philippi, 1849),was studied on an intertidal rocky shore at Shirahama, Wakayama Prefecture,where two mytilid bivalves, Septifer virgatus (Wiegmann) andHormomya mutabilis (Gould), formed vertically contiguous musselbeds in the upper-middle and lower zones, respectively. At lowtide in April, the snail density increased with decreasing shoreheight and was greatest at the middle level of the H. mutabilisbed. Then, the density decreased towards the lower littoralfringe. Newly settled juveniles smaller than 2 mm in shell heightappeared abundantly in late summer and autumn within algal turfon the lower shore. As snails grew larger than 2 mm, they appearedwithin the gaps of the H. mutabilis bed and the S. virgatusbed. They increased in size monotonically towards the next summer,but rate of growth in shell height tended to be great in autumnand small in winter. Seasonal change in the density of snailsfound within the gaps of the mussel beds was remarkable during athree year period, increasing from autumn to winter and then decreasingtowards next summer. Reproduction occurred in summer, and adultsnails disappeared by September. It is thus suggested that this specieshas a one year lifespan and shows a habitat shift from algal turfto the gaps of the mussel beds with growth. (Received 12 October 1998; accepted 2 March 1999)     

Mortality, behavior and the effects of predators on the intertidal distribution of littorinid gastropods

Predators can affect the vertical distribution of mobile intertidal invertebrates in two ways: they can (1) cause greater mortality of prey at certain intertidal levels, and (2) induce prey to seek safer intertidal areas. In this study, we investigate whether low-intertidal and subtidal predators affect the intertidal distribution of two congeneric species of small herbivorous gastropods of northeastern Pacific shores, Littorina sitkana Philippi 1846, and L. scutulata Gould 1849. In particular, we tested the hypothesis that predators affect the distribution of these snails by inducing them to seek higher and safer intertidal areas. On a wave-sheltered shore in Barkley Sound, British Columbia, L. sitkana and L. scutulata were both killed by predatory crabs (e.g., Cancer productus) more frequently when tethered near the lower limit of their intertidal distribution ( approximately 1 m) than when tethered where they were most common ( approximately 2.5 m), suggesting that high mortality rates are partly responsible for the lower-limit of these snails' intertidal distribution. However, two field mark-recapture experiments indicated that the snails' behavioral response to predation risk also influences their distribution. In the first experiment, snails from the 2.5-m level (low risk) transplanted to the 1.0-m level (high risk) displayed a strong and consistent tendency to move shoreward, especially L. sitkana, some traveling 10-15 m in 2-3 days to regain their original level. These shoreward movements were especially precise in the northern part of the study area, where predation rates on tethered snails were greatest. Furthermore, larger more vulnerable snails were more strongly oriented shoreward than smaller individuals, indicating that antipredator behavior might also contribute to intertidal size gradients in these species. In the second mark-recapture experiment, we manipulated predation risk using small cages and found that snails exposed to the odors of C. productus crabs foraging on conspecific and heterospecific snails displayed more precise (L. sitkana and L. scutulata) and longer (L. sitkana) shoreward movements than snails held in control conditions. These results provide the first experimental evidence that antipredator behavior may contribute to the intertidal distribution patterns of littorinids.     

Population dynamics,growth and reproductive rates of Littorina nigrolineata Gray from a moderately sheltered locality in North Wales

The population dynamics of Littorina nigrolineata Gray on a moderately sheltered boulder shore was characterized by an almost constant mortality rate of post-juvenile snails both within and between years. Cohorts of post-juvenile snails had a “half life” of about 5 months. Juvenile snails had higher mortality rates than post-juveniles. Mortality rate was independent of size in post-juveniles. The population of post-juvenile snails fluctuated only about two-fold during the 3 yr of study. Maxima of population size occurred each September as a result of an autumnal pulse of recruitment into the post-juvenile size classes. These recruits originated from eggs laid in the summer of the previous year. The size-frequency structure of the population lacked well defined modes except during the spring and summer when the pulse of recruits formed a distinct mode, which later disappeared due to growth and mortality of the recruits. Oviposition continued throughout the year with a pulse in summer. Newly laid eggs took 4–7 wk to hatch. The hatchlings had a mean shell height of 0.5 mm and were estimated to take 6–9 months to grow to a shell height of about 4 mm. Growth increments over periods of 3 months. estimated from tagged cohorts of snails ranging from 3 to 16 mm, released on a stone jetty, revealed seasonal and size-specific changes in growth rate. Growth rate was significantly lower between December and March than at other times of the year. Seasonal differences in growth rate did not, however, significantly affect the yearly size increments of snails hatching in summer or winter. Growth increments did not decline linearly with increasing body size, as is assumed by the von Bertalanffy growth model, but increased to a maximum at about 6–7 mm shell height. Beyond the maximum, the growth increments decreased linearly with increasing shell height. This non-linearity caused an inflexion in the growth curve, so that the shell height. This non-linearity caused an inflexion in the and 12.4 mm by the von Bertalanffy growth curve. The deviation from the von Bertalanffy growth model was possibly due to departures from the assumptions that the assimilation rate is proportional to the square of a linear size dimension and that the metabolic rate is proportional to the cube of the same dimension. A von Bertalanffy growth curve, fitted to growth increments measured from “growth checks” on tagged snails on the boulder shore, predicted slightly faster growth rates than the von Bertalanffy growth curve fitted to increments of mean cohort size on the stone jetty. The closeness of the two curves, however, suggested that both methods of measuring growth increments were tolerably accurate.     

Why are intertidal snails rare in the subtidal? Predation, growth and the vertical distribution of Littorina littorea (L.) in the Gulf of Maine

The North Atlantic gastropod Littorina littorea exhibits a characteristic “intertidal” distribution: the snail is abundant in the littoral zone but scarce in the shallow subtidal and the relatively few subtidal individuals are larger (in shell size) on average than those in the intertidal zone. For highly mobile species like L. littorea, this vertical distribution is primarily determined by directional movement. Biotic and abiotic factors vary across tidal heights, and natural selection for movement to shore levels where fitness is maximized provides the ultimate (evolutionary) explanation for vertical distribution patterns. In this study, we asked whether variation in growth rate and/or predation pressure among tidal heights provide an ultimate explanation for vertical gradients in L. littorea size and abundance. We used a cage experiment to compare juvenile growth rate among tidal heights and a series of field and laboratory experiments to examine variation in predation pressure among tidal heights and snail size classes. Juvenile growth rates were highest in the low intertidal zone, declining at both higher and lower levels. Predation risk for tethered L. littorea increased with both decreasing tidal height and decreasing body size (shell height). Almost all tethered prey were consumed by shell- breaking predators and a census revealed that the two most abundant such predators were the crabs Carcinus maenas and Cancer borealis. Laboratory feeding experiments were used to compare size-dependent prey vulnerability and prey-size preferences for these two key predators. We found that L. littorea vulnerability decreased with increasing snail size and increased with increasing size of both predator species. However, whereas C. borealis were capable of consuming even the largest L. littorea, most Carcinus were unable to feed on individuals larger than 10 mm in shell height. Additionally, C. borealis preferred larger sizes of L. littorea than did Carcinus. Thus, Carcinus, which co-occurs with L. littorea in the intertidal, is a much less effective predator than C. borealis, which is found primarily in the subtidal. We conclude that predation on L. littorea by C. borealis and other subtidal consumers has resulted in the scarcity of this ecologically important grazer in the subtidal. This effect has been produced both through direct predation and by imposing strong selection for movement of L. littorea to higher tidal zones.     

The role of growth,predation, and habitat selection in the population distribution of the crown conch,Melongena corona Gmelin

The intertidal gastropod Melongena corona Gmelin exhibits a size gradient along the shore as a function of habitat. Small, juvenile snails were found on the sand beach and larger adults were found on hard substratum habitats (shell rubble beach and oyster bar). A transplant experiment was performed to test three proximate explanations for this pattern:

• 1.(1) differential growth rate of conchs in different habitats,
• 2.(2) differences in predation intensity between habitats and
• 3.(3) active habitat selection by snails.

Shell scars on the last shell whorl were used as an index of predation pressure. Growth and predation were not significantly different for snails of similar size in different habitats, but snails were found to return to their original habitat when displaced. The ability to home did not differ between sexes or juveniles and adults. Active habitat selection appears to be a significant proximate factor maintaining the population distribution. A number of potential ultimate causes of the size class segregation are suggested.     

EFFECTS OF WOUNDING BY THE HERBIVOROUS SNAILS LITTORINA SITKANA AND L. SCUTULATA (MOLLUSCA) ON GROWTH AND REPRODUCTION OF THE INTERTIDAL ALGA FUCUS DISTICHUS (PHAEOPHYTA)1

Small (3–7 cm long) Fucus distichus ssp. edentatus (de la Pyl.) Pow. Plants were tagged at three sites in which densities of the herbivorous snails Littorina sitkana and L. scutulata ranged from 367–4690 animals · M^-2. From April–August 1986, the growth rate, degree of wounding, and reproductive status of individual thalli were monitored at 2–4 week intervals. Grazer-inflicted damage to the thalli varied within and among sites. Mean growth rates at the site with low densities of littorines were about twice those at the site with intermediate densities and about four times those at the high density site. At the site with high densities of littorines, F. distichus growth rates were negatively correlated with the degree to which the plants were wounded. There appeared to be no correlation of grazer density with F. distichus survivorship. Thalli at the site with few herbivores tended to reproduce earlier and at a larger size than did those at the other two sites. In all three areas, only thalli that had received little damage from herbivores became reproductive. By lowering growth rates and delaying reproduction in F. distichus, grazing by littorine snails can potentially cause variation in reproductive output among individual thalli.     

Feeding adaptations of Melagraphia aethiops (Gmelin), an intertidal trochid mollusc

The feeding behaviour of the intertidal trochid Melagraphia aethiops (Gmelin) on a sheltered stony shore was investigated. The effects of snail size, season, and shore level on feeding were examined. Feeding was examined in the laboratory in aquaria fitted with glass plates coated with algal and detrital films. Rates of radula rasping and the frequency of feeding bouts (feeding activity) were measured.

Radula rate varied little with snail size and large snails were 35% more active feeders than small snails. Small snails, however, consumed three to four times as much food per radula rasp, on a unit weight basis, as large snails. Winter acclimatization involved an increased tolerance of low water temperatures and an overall 60% decrease in radula rasping rate compared with summer acclimatized animals. Concurrently, field crawling rates decreased 71% during the summer-winter transition suggesting that lower food intake was balanced by reduced locomotory activity. High and low shore inhabiting snails showed little difference in radula rasping rates. High shore snails, however, compensated for their shorter feeding periods by being 37% more active in feeding and crawling in the field 40% faster than low shore snails.     

Growth rate differences between upper and lower shore ecotypes of the marine snail Littorina saxatilis (Olivi) (Gastropoda)

Shell polymorphisms are widespread among those intertidal gastropods that lack a pelagic spreading stage. These polymorphisms may indicate diversifying selection in a heterogeneous habitat, but to do this the variation must be at least pardy inherited. Galician populations of Littorina saxatilis (Olivi) living in exposed rocky shores are highly polymorphic in several shell traits, e.g. ornamentation, banding and size. Mature snails of the upper-shore ridged and banded (RB) morph is, for example, often twice as large as mature individuals of the lower-shore smooth and unbanded (SU) morph of the same shore.
We investigated the hypothesis that lower-shore snails grow more slowly and that differences in growth rate were at least partly inherited and could be explained by diversifying selection. We released snails of different origin (upper, mid- and lower shore) and morph (RB, SU and hybrids) at different shore levels and compared their shell increment after one month of growth. We found that despite considerable variation among individuals and among replicate samples (together about 53% of the total variation), average rates of growth differed between morphs. RB snails both from the upper and mid-shores grew at a high rate at all shore levels, SU snails grew considerably less, and hybrids grew at intermediate rates, at all levels. Inherited difference among morphs explained about 34% of the total variation while effects of shore levels and the interaction morph x shore level explained only 5 and 7%, respectively. Thus a large part of the difference in growth rate leading to different adult sizes of the two morphs has probably evolved due to spatially varying selection favouring large sizes in upper-shore and small sizes in lower-shore environments.     

Predatory behaviour of juvenile shore crab Carcinus maenas (L.)

Juvenile shore crabs Carcinus maenas (L.) were observed feeding on rock barnacles Semibalanus balanoides (L.) on a Bay of Fundy rocky shore. This previously unreported predatory behaviour was further investigated in the laboratory. When given a choice of three common and abundant gastropods, Nucella lapillus (L.), Littorina littorea (L.), and Littorina obtusata (L.), and the rock barnacle Semibalanus balanoides, juvenile shore crabs of both sexes ate mainly barnacles and consumed proportionately more barnacles than gastropods compared with adults, which ate mainly gastropods. The rock barnacle is an abundant and readily available food source which may be important in sustaining the juvenile crab through periods of moults and rapid growth. As the shore crab attains a certain age (size), it must forage lower on the shore as gastropods become more important in its diet.     

Field and laboratory studies on the life-cycle, growth and feeding preference in the hairy snail Trochulus hispidus (L., 1758) (Gastropoda: Pulmonata: Hygromiidae)

For the first time the life cycle of the common land snail Trochulus hispidus was completely described in Central Europe (Poland). This is a semelparous species predominantly with an annual life cycle and the reproductive period lasting from April till October. The first young snails hatch in spring, grow rapidly in summer and reach ca. 4 whorls until winter. In spring of the next year they mature and reproduce. After that they die. There is hardly any growth from late autumn till early spring. The average proportional growth rate is ca. 0.3 whorl/month in the wild. The fastest growth is present in the youngest snails and then gradually decreases over the course of their age. Laboratory and field observations allowed for establishing the following life cycle parameters: eggs calcified, almost spherical, ca. 1.5 mm, laid in spring and summer in batches of between 1 and 47. Time to hatching is 6–24 days, hatching is asynchronous; newly-hatched snails have approximately 1.5 whorls. Analysis of food preferences revealed, that T. hispidus tends to restrict its diet during the life. Generally the youngest snails equally consumed leaves of all four tree species offered (Fraxinus excelsior, Acer pseudoplatanus, Tilia cordata and A. platanoides) whereas adults preferred F. excelsior over A. pseudoplatanus and A. platanoides.     

Du microcosme évolutif à la théorie générale : les escargots,Steve Gould et l’évolution

From an Evolutionary Microcosm to general theory: Land Snails, Steve Gould and Evolution. From 1962 to the end of his career, Stephen Jay Gould has studied the land snails of the genera Poecilozonites and then Cerion. Snails have provided him material to develop his morphometric methods, to analyze shell growth, and may have been at the origin of his work on ontogeny and phylogeny. Their variation, which he interpreted in terms of adaptation until the beginning of the 1980s, has been later reanalyzed by him in terms of non-adaptive evolution and contingency. Growth and variation of the shell allowed Gould to define constraint and canalisation in evolution. In their taxonomic work, Gould and Woodruff have practiced a definition of biological species as the group of populations allowing maximal congruence of their morphological characters, genetic characters and geographic distribution, of which potential interfecundity is necessary, but not sufficient. To cite this article: S. Tillier, C. R. Palevol 2 (2003).     

Opposing selective pressures decouple pattern and process of parasitic infection over small spatial scale

Species face multiple selective pressures that may require opposing responses to mitigate. On rocky shorelines, fitness of the intertidal snail Littorina littorea is determined by both parasitism and predation. We experimentally demonstrated that L. littorea was at greatest risk of infection from trematode parasites high in the intertidal zone where it was in closest proximity to abundant gull feces (the vector for the snail's parasites). However, because of extreme, size‐selective predation pressure at low tidal elevations, small snails often live high in the intertidal until they have grown sufficiently large. By prolonging their exposure to infection higher on the shore, ontogenetic responses to predation risk accentuate parasite risk. Counterintuitively, snails exhibited the highest trematode prevalence at the lowest tidal elevations where they had almost no risk of contracting infection. By carrying contracted infections into the lowest tidal zones, the larger, predation‐resistant snails invert hotspots of infection risk and prevalence, underscoring that size‐dependent selection pressures can decouple infection process and pattern even over small scales.