Reproductive condition in two Malayan freshwater viviparid gastropods

Siamopaludina martensi and Filopaludina sumatrensis have typical viviparid reproductive systems with dimorphic spermatozoa and copulatory right tentacle in the male, and a large pallial brood-pouch in the female. Females outnumber males as in several other viviparids. Brood-pouches contained overall monthly averages of 0–14 developing young in S. martensi , 9–23 in F. sumatrensis , more than half of which were well-developed, later juveniles. Brood-pouch counts and ovary condition of both species varied in broad accordance with monthly rainfall, giving an irregularly seasonal pattern of annual change resembling those of several terrestrial Malayan snails. Juveniles emerge from the brood-pouch mostly at night. Alteration of aquarium conditions leads to temporarily accelerated release of young but does not permanently raise brood-pouch counts or rates of release.     

Oviposition Behavior and Offspring Emergence Patterns in Succinea thaanumi, an Endemic Hawaiian Land Snail

In this field study, newly laid egg clutches of Succinea thaanumi, an endemic Hawaiian land snail, were labeled and observed weekly over a 12‐mo period. Snails usually laid their clutches under leaf tips and, moreover, this placement affected snail emergence by mitigating the effects of dehydration through the leaf‐drip phenomenon. During droughts, the gel surrounding egg clutches dehydrated, pasting the embryos to the backs of leaves. The hygroscopic gel rehydrated once rains resumed. The majority of newly laid clutches contained shell‐less embryos which required more time for emergence than embryos laid with visible shells. More shelled embryos were laid subsequent to a 3‐wk drought, indicating that snails retained their eggs during adverse environmental conditions and laid them when conditions improved. The snails’ abilities to retain eggs in adverse conditions, but to lay more unshelled embryos over their lifetimes, contributed to their overall reproductive success.     

Shell shape and land-snail habitat in a Mediterranean and desert fauna

The bimodal distribution of shell shape (height: diameter), that is found in various geographically widely separate and taxonomically distinct land snail faunas of many different regions of the world, occurs also in a Mediterranean fauna and in a desert fauna that is derived from it. The desert fauna is, however, closer to the bisector than the Mediterranean one. High-spired snails are mainly rock-dwellers, and equidimensional to low-spired snails are bush-dwellers or soil-diggers, with a few rock-dwellers; litter-dwellers are small-sized species that may have either high- or low-spired shells. These results are discussed in adaptive terms. Litter is probably the more primitive of these micro-environments. Many of the small, litter-dwelling snails are ovo-viviparous rather than oviparous, perhaps so as to avoid attacks on the eggs by saprophytic fungi. The shift away from the litter environment is accompanied by a trend to abandon the ovo-viviparous strategy, in favour of oviparity, the snail using its foot to dig into the soil and lay eggs. The conchometric differences between bush-, ground- and rock-dwelling snails may perhaps reflect selective pressure to increase the size of the foot; and constraints of a habitat that consists of narrow interspaces between rocky boulders. Snails that habitually dig into the ground during periods of inactivity, and roam over the ground when active, requires a very large foot and, consequently, a very large-mouthed shell to accommodate it; the result is an equidimensional shell, globose or turbiniform in shape. Snails that climb up vertical vegetation would also require a large foot, and consequently a large-mouthed shell to contain it. A fully globose shell would however be disadvantageous, since it might cause undesired torque. Hence, bush-dwellers tend to be flatter than soil-diggers. Snails that habitually live in rock crevices, and on hard substrata, would not require a very large foot; they would need a narrow shell, both to enable easy manoeuvring through crevices and to reduce torque, the result being a small-mouthed, usually high-spired shell. The classification of land snails into bush-, soil- or rock-dwellers closely follows the taxonomic classification. In those species that depart from the habitat that is typical of their taxonomic group towards another habitat, the shell alters its shape accordingly.     

Resting-egg hatching and early population development in rotifers: a review and a hypothesis for differences between shallow and deep waters

This paper reviews our very limited knowledge about resting-egg hatching and early population development in planktonic rotifers. Hatching of stem females from resting eggs may occur soon after resting eggs are produced, but perhaps usually it is delayed by a minimum obligate diapause, by a requirement for seasonal temperature changes, or by sinking to sediment environments that prevent hatching. In deep-water sediments, hatching probably is inhibited by low oxygen, darkness, or low temperature, so that eggs likely hatch following resuspension during water-column turnover. Populations should develop primarily by female parthenogenesis and have relatively low clonal diversity. By contrast, in shallow-water sediments, eggs are more likely to experience conditions conducive to hatching and to be resuspended into the water column. Populations may develop by massive emergence of stem females as well as by female parthenogenesis and thus have a very high clonal diversity. Stem females of some species are particularly fit for colonization of new habitat. First, compared to females hatched from parthenogenetic eggs, they can have a greater lipid reserve that enhances survival and reproduction. Second, amictic stem females can contain a transmissible factor that inhibits sexual reproduction and diapause for several to many successive generations, thus promoting rapid reproduction via female parthenogenesis.     

Shell-banding polymorphism of the land snail Xeropicta vestalis along the coastal plain of Israel

Along the coastal plain of Israel, shell darkness of the polymorphic land snail Xeropicta vestalis is positively related to the extent of perennial vegetation. It is not related to rain, temperature, geographic position or darkness of the ground.
White shells reflect more radiation than dark ones do, and therefore in Israel's coastal plain, where solar radiation is very strong, they are favoured. Perennial vegetation, where it occurs, shields the snails from the sun by absorbing radiation, so that snails amongst them can afford to be darker, and thus more cryptic. Hence, the more perennial vegetation in the habitat, the darker the shells.
Also during the Pleistocene, when temperatures were 5–10°C lower than today, the shells of the coastal plain were as pale as recent ones are. The distribution of a snail-predator, Gerbillus allenbyi , which is restricted to sandy biotopes where perennial bushes occur, and which was absent from Israel during the Pleistocene, could perhaps explain the distribution of X. vestalis morphs, both today and in the past.     

The effect of isolation on the life-history traits of Pseudosuccinea columella (Pulmonata: Lymnaeidae)

A population of Pseudosuccinea columella was raised under laboratory conditions and its life tables were determined in isolated and paired snails. Isolated snails were significantly larger in shell size than paired snails from five weeks of age onward. Also, statistically significant differences were found for the number of eggs per mass per individual from week 5 to 9, isolated snails exhibiting the highest values. The intrinsic and finite rates of increase were greater in isolated than in paired snails. Either an inhibition of the reproductive output between individuals or the advantage of selfing may be the cause of the differences in this species, acting as a possible mechanism that increase the fitness of isolated snails.     

The life cycle, population dynamics and productivity of Melanoides tuberculata (Muller, 1774) (Gastropoda: Prosobranchia: Thiaridae) in Hong Kong

An investigation of the life cycle, population dynamics and secondary productivity of the viviparous snail Melanoides tuberculuta inhabiting flooded furrows and irrigation ditches at Ping Long, New Territories, Hong Kong, was undertaken between March 1982 and March 1983. There was a single peak in juvenile recruitment coinciding with the warmer months; hatchlings grew quickly and were sexually mature before the next breeding season. Large animals experienced high mortality and few individuals survived until a second breeding season. Secondary productivity and the production: biomass (P:B) ratio of the population (13.43 g shell–free dry weight m⁻² yr⁻¹ and 4.81, respectively) were high and exceeded values of these parameters recorded for freshwater prosobranch populations in temperate regions. Ping Long M. tuberculata were exclusively female, as is typical of most populations of this species. Animals > 3.0 mm shell width (90–120 days old) had eggs and developing larvae in the brood pouch, and fully–developed larvae were recorded from brood pouches throughout the study although release of hatchlings was distinctly seasonal.
A comparison of life cycle parameters of Malaysian M. turberculata with those of the Ping Long population revealed a marked similarity. The significance of such similarity and the importance of parthenogenicity in the life cycle of this widely distributed snail is discussed. Melanoides tuberculata exhibits the characteristics of an opportunist or 'fugitive' species and, by virtue of its reproductive strategies and high productivity, is able to colonize rapidly and temporarily dominate vacant habitats.     

THE EARLY LIFE-CYCLE STAGES OF HYDROBIA VENTROSA AND HYDROBIA NEGLECTA WITH OBSERVATIONS ON POTAMOPYRGUS JENKINSI

The egg mass of Hydrobia neglecta contains a single egg, whilethat of H. ventrosa contains up to three eggs. At hatching H.neglecta has a significantly greater shell length than H. ventrosaand the late embryos and young snails of the two species canbe separated according to the pigmentation of the head region.The young snails can also be separated on the basis of surfaceornamentation of the shell and comparisons with the closelyrelated H. ulvae and Potamopyrgus jenkinsi show how the basicpattern of shell sculpturing differs in the four species. Thesedifferences are discussed in relation to the mode of development. (Received 14 February 1980;     

Oviparous reproduction of Vestia ranojevici moravica confirms taxonomic value of life history characters in carpathian clausiliids (Gastropoda: Pulmonata)

Based on five-year observation of 36 snails collected in the ?eladná valley (Moravia, Czech Republic) and their progeny reared in laboratory, it was found that Vestia ranojevici moravica (Brabenec, 1952) is oviparous. The oviparity in V. ranojevici is in line with the previous assumption of its reproductive strategy based on anatomical studies alone. It supports the separate position of the species in the genus Vestiaa (subgenus Brabenecia). V. ranojevici was kept at room temperature (18–25°C) between March and October; in winter snails were stored at 3°C. The egg-laying period started in late March, ca. 1 month after raising the temperature, and lasted till September, with one maximum in spring. In the laboratory the snails laid eggs in batches (usually 4–6 eggs, max 11) or singly. The number of batches per pair per season was 1–8; the corresponding number of eggs — 3–40. The eggs were gelatinous, with separate calcium carbonate crystals in the external envelope. The average egg size was 1.80 × 1.51 mm, but actual size and shape were highly variable: larger and more elongated eggs were produced mainly by laboratory reared animals. Freshly deposited eggs did not contain shelled embryos nor the reproductive tract of dissected individuals contained eggs. At room temperature the eggs hatched after ca. 14–16 days, then the shell growth lasted 20–29 weeks; the snails laid eggs 30 weeks after completion of shell growth indicating one-year life cycle under laboratory conditions. Adult individuals collected in the field reproduced during 4–5 consecutive years. Comparison of V. ranojevici reproduction to the results of breeding of other Carpathian clausiliids in the same laboratory shows that similar reproductive strategies may have evolved independently in different lineages of the group: both the oviparous and egg retaining species belong to the genera Vestia and Balea.     

Comparison of heart rate in embryonic,young and adult Achatina fulica

We have studied the embryonic heart in the gastropod Achatina fulica using eggs soon after their deposition. We also followed the development of organs, with the main emphasis on the heart and its physiology. The embryos were examined at different stages by carefully removing egg shell until hatching. In this context the heart rate (HR) was analysed in embryonic snails and compared with that of young (from hatching) and adults. The HR of embryos dissected from newly deposited eggs was irregular, with a mean frequency of 0.42?±?0.14?Hz. The heartbeat during the ontogeny was more rhythmic and HR increased to 0.82?±?0.12?Hz. The intersystolic intervals (ISI) in real time showed a Gaussian distribution for all animals. The peak ISI from Gaussian fit closely matched the cumulative probability for corresponding data. Although the HR increased after hatching, comparisons yield a weak correlation (R?=?0.56?±?0.27) with the total body weight of embryonic, young and adult snails.     

Natural history of Theba pisana in Israel (Pulmonata: Helicidae)

Theba pisana in southern Israel is a semelparous species with an annual life cycle. Young snails hatch in early winter, grow rapidly in spring and by summer they reach, first adult shell-size, then adult shell-weight. Shell-size is negatively related to density, suggesting that, as in some other landsnails, there may be a density-dependent inhibition of growth. Summer forces the snails into 3–4 months of aestivation.
After the first winter rains the snails, awakened perhaps by the sudden drop in barometric pressure, emerge from aestivation. Only now are most of the spermatozoa and mature oocytes formed (maturation of gametes actually starts some two weeks before the first rains), and the snails copulate. No evidence of precocious breeding was found. The snails sometimes dig a few "sham" holes, perhaps to discourage egg-eating predators, before finally depositing their eggs (mean clutch size: 80 eggs) in the last hole.
In mid-winter, the snails die. In a habitat where the sheltering conditions of the vegetation offer the snails more protection, the dying off process of the population is stretched over a few more months. This could, perhaps, be the first step towards the evolution of a bi-annual cycle, such as is typical of T. pisana in some other areas of its distribution.     

CALCIUM PROVISION TO EGGS IN TWO POPULATIONS OF HELIX ASPERSA BY PARENTS FED A DIET HIGH IN LEAD

The reproductive performance of two populations of Helix aspersawith different histories of exposure to lead was assessed bytheir egg production, hatching success and the metal contentof the eggs. By using laboratory-bred parents raised in a Pb-freeenvironment, the possibility of an inherited response to a highPb exposure was assessed. Following a period of dormancy, six individuals in each population werepaired for copulation and allowed to oviposit. During this time theywere fed a diet with 500 µg.g^-1 Pb, for a total of 14days. After oviposition, the egg mass from each pairing was weighed,counted and samples analysed. Metal concentrations in four tissuefractions of the parents (shell, hepatopancreas, reproductive systemand rest9) were also analysed. There was no significant transfer of Pb to the eggs in either population.There were differences in their egg massesthe population froman uncontaminated site, Rottingdean (RD), laid, on average,twice the number of eggs of the snails from Minera (MI), a contaminatedsite in North Wales. However the latter had a higher averagepercentage emergence (90% compared to 55% for RD). There wasalso considerable hatching asynchrony in the RD offspring, whilehatching within each MI egg mass was completed within two days.This is discussed in relation to the Ca status of the parentand possible differences in reproductive strategy between thetwo populations. (Received 22 February 2000; accepted 8 May 2000)     

Schistosoma mansoni infections in neonatal Biomphalaria glabrata snails.

In areas endemic for schistosomiasis, the population dynamics of the snail intermediate hosts have a direct effect on parasite transmission. The present study focused on the potential for neonatal Biomphalaria glabrata snails to become infected with Schistosoma mansoni and to produce cercariae under various conditions. It was found that snails as small as 0.74 mm in shell diameter could survive miracidial penetration and could release cercariae when as small as 1.6 mm in diameter. Cercariae produced by small snails were equally infectious for mice when compared with those shed by larger snails. Likewise, histological examination of neonatally exposed snails revealed normally developing parasites at all stages of infection. It was found that in 2 snail populations expressing either high or low susceptibility to the parasite, peak susceptibility occurred at 25 days of age in both groups. Daily cercarial production for neonatally exposed snails was initially low but increased dramatically as the snails grew, eventually reaching values as high as 2,100 cercariae/snail/day. A moderate to high percentage of snails infected as neonates was eventually capable of simultaneously producing both eggs and cercariae. These studies emphasize the potential importance of neonatal and preadult snails in helping to maintain foci of S. mansoni infection in endemic areas.     

ANNUAL AND LUNAR REPRODUCTIVE CYCLES IN LITTORINA SAXATILIS (OLIVI) AND DIFFERENCES BETWEEN BREEDING IN THE MARINE FIRTH OF FORTH AND THE FORTH ESTUARY

Gonads were most commonly reduced in October and fullest inMar-Jun with more pronounced seasonal differences in the estuarythan in the firth. Brood-pouches accordingly were fullest inspring-early summer and emptiest in autumn-early winter. Eggsand early embryos predominated in brood-pouches of early springbut were few in Aug-Oct whereas well-developed juveniles werefew in early spring, increased in spring-summer and came topredominate in Jun-Nov. Brood-pouch loads varied with size offemale but, for a standard 10 mm female, were significantlylower in the 2 estuarine sites than in the firth. Changes inmean brood-pouch count varied only 2.5-fold during the yearin the firth but 4.10-fold in the estuary. In vitro development of eggs to shelled juveniles with the velumresorbed took some 68 d at 10°C and 32%. Eggs and late brood-pouchjuveniles were significantly heavier in the estuary than inthe firth (dry weight, ash-free dry weight (AFDW), ash weight).AFDW changed little during development while ash weight (mostlyshell) increased 3.5-4.4-fold. In a laboratory tide tank, femalesreleased 60 d means of <1.3 juveniles d^–1; more inmid- and late year than in Jan-Mar, more from upshore firthfemales than from low shore or estuary females, more when heldat high tide-tank levels than at lower levels, and generallyat rates comparable to release in containers held at the siteof origin. Egg production in laboratory females did not keeppace with release of young and, at the end of trials, brood-pouchcounts were mostly lower than in contemporary free winkles.Females that had released most (e.g. at high tank levels) containedleast brood-pouch embryos and young subsequently. Release of young in the tide tank was markedly faster at timesnear new moon (often reaching or exceeding 2 female^–1d^–1) than near full moon (often <1 female^–1 d^–1).This lunar fluctuation was clearest in fast-releasing sets heldat high- or mid-tide tank levels. It was not observed in Jan-Mar(when release from most females was slow) nor in females atlow tank levels. (Received 3 September 1990; accepted 8 November 1990)     

Effects of dietary calcium on growth and oviposition of the African land snail Limicolaria flammea (Pulmonata: Achatinidae)

In an attempt to elucidate the role of calcium in the life of the edible Achatinid snail, Limicolaria flammea (Miller) I investigated short and long term effects of calcium added to the food. The short term experiments lasted for 18, 30 and 32 weeks respectively, while the long term experiment to determine life time utilization of calcium carbonate lasted for 15 months. In the short term experiments, hatchlings were divided into densities of one, ten and 50 snails. In the 10 snail group, there was a positive correlation between calcium provision, body weight (t test, p < 0.01; r = 0.96, p < 0.0001) and shell length (t test, p < 0.01; r = 0.96, p < 0.00001). There was also a positive correlation between increase in shell length and availability of calcium in the 1 snail group (t test, p< 0.01; r = 0.99, p < 0.00001). In the 50-snail group, the correlation was positive for shell length of the snails (t test, p < 0.05; r = 0.99, p < 0.0001) and body weight (t-test, p < 0.05; r = 99, p < 0.00001). Mortality was very high in the snails deprived of calcium and they did not produce eggs. In the long term experiment, there were three feeding peaks in L. flammea. In the first feeding peak, amount of food and calcium ingested by the snails increased in the first three months of life. The second feeding peak occurred at six months of age, while the last occurred at 10 months of age. The amount of calcium ingested during the second peak decreased gradually in the 4th and 5th month. The amount of calcium ingested was lowest during the 3rd feeding peak. The period of highest weight gained by the snails was between the 1st and 6th month and then dropped at between six and 12 months of age which corresponds to the period of egg production. There were also three peaks of egg production; the first was between six and eight months (535 eggs), the second at between 10 and 11 months (350 eggs) and the third at 13 to 14 months (310 eggs) respectively.     

A comparative study of the ultrastructure and mineralogy of calcified land snail eggs (Pulmonata: Stylommatophora)

This study indicates that eggs containing calcium carbonate crystals occur in at least 36 of the 65 known families of the land snails (class Gastropoda: order Stylommatophora). Eggs from 22 of these families were available for examination. The x-ray diffraction data, available for the first time for 21 of these families, shows that these egg shells are all made of calcite only, or of a combination of calcite with smaller amounts of aragonite. All of the snail (body) shells examined were made of aragonite only. This is the first ultrastructural investigation of these egg shells, and it indicates that the eggs exhibit enough structural diversity to allow identification of parental animals to genus, and often to species level solely on the basis of egg shell ultrastructure. All of the calcified eggs may be divided into two groups: (1) partly calcified, with discrete crystals of CaCo₃ dispersed in the jelly layer, and (2) heavily calcified, with a hard, brittle egg shell made of fused crystals of CaCO₃ much like an avian egg. Both types of calcified eggs occur in oviparous as well as in ovoviviparous snails. Because of the wide distribution of calcified eggs in the Stylommatophora, and because of the occurrence of heavily calcified eggs in ancient families such as Partulidae, Endodontidae, and Zonitidae, the calcified egg is viewed as a primitive land snail trait associated with terrestrial adaptation. The function of the calcified egg shell, in addition to mechanical support of egg contents, is to supply the developing embryo with enough calcium to form the embryonic shell by the time of hatching.     

Cultivating Lithoglyphopsis aperta Temcharoen, a new snail host for Schistosoma japonicum, Mekong strain.

Lithoglyphopsis aperta Temcharoen, a new intermediate host for Schistosoma japonicum, Mekong strain, was cultivated successfully through the F2 generation with the use of a petri dish aquarium provided with mud and alga. The incubation period of eggs was about 4 weeks; the young grew to maturity in 16 to 20 weeks; egg-laying followed about 6 to 8 weeks later, the cycle taking roughly 6 months. The husked eggs were deposited exclusively at the edge and wall of the aquarium. Eggs were hemispherical and with the husks were between 0.37 and 0.43 mm in diameter. The newly hatched young were only 0.24 mm in shell length and the full-grown snails were 3.11 to 3.36 mm long. Logarithmic growth began between the 12th and 14th weeks.     

Social facilitation affects longevity and lifetime reproductive success in a self-fertilizing land snail

Factors that reduce the reproductive output of self-fertilizing hermaphrodites are receiving increasing attention. The combined effects of reduced fecundity of selfing parents and inbreeding depression of the progeny have been referred to as self-fertilization depression. In isolated freshwater snails the reproductive output of selfing individuals also decreases due to the lack of social facilitation (absence of a conspecific). We examined the effect of social facilitation on lifetime reproductive success (number of young produced and longevity) over two generations in the simultaneously hermaphroditic land snail Balea perversa . In a parallel study we showed that B. perversa kept singly and in pairs reproduced exclusively by self-fertilization. In the parent generation, snails kept singly produced less offspring than snails kept in pairs. The difference in lifetime number of young was mainly due to differences in adult life span. Snails of the two groups did not differ in reproductive rate (number of young produced per 100 d of reproductive life) and hatchling size. In the offspring generation, snails kept singly did not differ from individuals kept in pairs in the lifetime number of young and hatchling survival. As in the parent generation, snails kept singly reproduced during a shorter period than snails kept in pairs. However, the shorter reproductive life span of snails kept singly was compensated for by a slightly (but not significantly) higher reproductive rate which resulted in a similar number of offspring produced for both groups. In both generations, snails of the two groups did not differ in size at first reproduction, adult growth rate and size at death. These findings suggest that social facilitation may affect longevity in selfing B. perversa .     

Seasonality and life histories of two endemic New Zealand carabid beetles (Coleoptera: Carabidae): Mecodema oconnori Broun and Megadromus capito (White)

Abstract

Seasonal activity and life histories of two large endemic carabid beetles, Mecodema oconnori Broun and Megadromus capito (White), were investigated in two lowland forest remnants in the Horowhenua region, North Island, New Zealand. Seasonal activity was estimated from manual searches of surface refuges, plus pitfall trapping from October to March. Adults of both species were present throughout the year, although M. oconnori was much more abundant in spring. Megadromus capito larvae were caught from October to March, and the three instars overlapped temporally, implying a long period of larval emergence and development. Adult females were dissected to investigate their reproductive phenology. Both species carried eggs for at least 6 months of the year. M. oconnori females contained very few eggs, with a maximum of three eggs each and a mean of 1.6 eggs per female. Mg. capito females had up to 28 eggs each and a mean of 10.5. Both species appear to have low egg loads, a relatively long period of reproductive activity and long life spans, but longer‐term studies on the pre‐adult stages and reproductive activity of adults are required to complete understanding of their life cycles.     

Gene regulation of shell banding in a land snail from Israel

The Israeli land snail, Xeropicta vestalis, offers a particularly clear example of gene regulation in relation to natural selection, in that within each population the appropriate phenotype is generated only at the correct part of the animals life cycle, and a contrasting phenotype develops when the forces of natural selection change. In the mountains of Jerusalem, where the winter is cold, the shells are dark. Westwards, towards the coastal plain where the winter is warmer, the shells gradually become paler. As dark shells absorb more radiation than pale ones, this clinal variation in morph frequencies can be explained in thermal terms. (Banded shells are also more cryptic than non-banded shells, so that in the mountains visual selection by predators may be an additional force which favours dark shells.) Xeropicta vestalis is an annual, semelparous species: the snails hatch in winter, become mature within one year, reproduce, and then die. In the coastal plain the snails are active throughout most of the year, and they have a long period in which to grow to reach adult size. In the mountains and hills, on the other hand, the snails are active for only a very short period. They spend most of their lives as small snails, in a state of aestivation. Xeropicta vestalis must be dark in mountains because when it finally awakens, it must very rapidly and hastily reach reproduction size. A dark shell, by speeding up temperature-dependent processes in this critical stage, assists the snail to mature rapidly. Shell darkness varies with age: in the mountains and hills the shells are moderately dark when they hatch, but become darker whilst growing in early winter. In the coastal plain also, the snails are moderately dark when they hatch; but here they become paler, whilst growing in winter and spring. In both cases, each snail is darker in the colder months and paler in the hot ones. A strategy of gene regulation of shell colour is thus favoured when the subsequent forces of the environment are very contrasting in their direction, very severe—yet also very predictable.