Lack of metabolic temperature compensation in the intertidal gastropods,Littorina saxatilis (Olivi) and L. obtusata (L.)

Two intertidal snails, Littorina saxatilis (Olivi, 1972) (upper eulittoral fringe/maritime zone) and Littorina obtusata (Linnaeus, 1758) (lower eulittoral) were collected from a boulder shore on Nobska Point, Cape Cod, Massachusetts, in July and acclimated for 15–20 days at 4 ° or 21 °C. Oxygen consumption rate (Vo₂) was determined for 11–15 subsamples of individuals at 4 °, 11 ° and 21 °C with silver/platinum oxygen electrodes. Multiple factor analysis of variance (MFANOVA) of lo₁₀ transformed values of whole animal Vo₂ with log₁₀ dry tissue weight (DTW) as a covariant revealed that increased test temperature induced a significant increase in Vo₂ in both species (P<0.00001). In contrast, MFANOVA revealed that temperature acclimation did not affect Vo₂ in either L. saxatilis (P= 0.35) or L. obtusata (P= 0.095). Thus, neither species displayed a capacity for the typical metabolic temperature compensation marked by an increase in Vo₂ at any one test temperature in individuals acclimated to a lower temperature that is characteristic of most ectothermic animals. Lack of capacity for metabolic temperature acclimation has also been reported in other littorinid snail species, and may be characteristic of the group as a whole. Lack of capacity for respiratory temperature acclimation in these two species and other littorinids may reflect the extensive semi-diurnal temperature variation that they are exposed to in their eulittoral and eulittoral fringe/maritime zone habitats. In these habitats, any metabolic benefits derived from longer-term temperature compensation of metabolic rates are negated by extreme daily temperature fluctuations. Instead, littorinid species appear to have evolved mechanisms for immediate metabolic regulation which, in L. saxatilis and L. obtusata and other littorinids, appear to centre on a unique ability for near instantaneous suppression of metabolic rate and entrance into short-term metabolic diapause at temperatures above 20–35 °C, making typical seasonal respiratory compensation mechanisms characteristic of most ectotherms of little adaptive value to littorinid species.     

Effects of ambient temperature and of temperature acclimation on nitrogen excretion and differential catabolism of protein and nonprotein resources in the intertidal snails,Littorina saxatilis (Olivi) and L. obtusata (L.)

Nitrogenous excretion in two snails, Littorina saxatilis (high intertidal) and L. obtusata (low intertidal) was studied in relation to temperature acclimation (at 4° and 21°C), including total N excretion rates, the fraction of urea in N excretion, corresponding O:N ratios and the partitioning of deaminated protein between catabolic and anabolic processes at 4°, 11° and 21°C. Aggregate N excretion rates in both species showed no significant compensatory adjustments following acclimation. Total weight specific N excretion rates at 21°C were higher in standard 3 mg L. saxatilis (739 ng N mg⁻¹ h⁻¹) than standard 5 mg L. obtusata (257 ng N mg⁻¹ h⁻¹) for snails acclimated to 21°C. Comparisons of Q₁₀ values of total weight specific N excretion to Q₁₀ values for weight specific oxygen consumption ({xxV}O₂) between 4° to 11 °C and 11° to 21°C indicated that, while total rates of catabolic metabolism ({xxV}O₂) and protein deamination in L. obtusata were essentially parallel, the relationship between N excretion and {xxV}O₂ in L. saxatilis revealed the partitioning of a larger share of deaminated protein carbon into anabolism at 4° and 21°C than at 11°C. Urea N accounted for a larger share of aggregate N excreted in L. saxatilis than in L. obtusata, but in both species urea N is a greater proportion of total N excreted when acclimated at 4°C (urea N: ammonia N ratio range: 1 to 2.15) than in snails acclimated to 21°C (urea N: ammonia N ratio range: 0.46 to 1.39). Molar O:N ratios indicate that the proportion of metabolism supported by protein catabolism is greater in L. saxatilis (O:N range: 2.5–8.4) than in L. obtusata (O:N range: 7.3–13.0). In both species, regardless of acclimation temperature, the O:N ratios are generally lowest (high protein catabolism) at 4°C and highest at 21°C.     

Seasonal migration promoting assortative mating in Littorina brevicula on a boulder shore in Japan

Littorina brevicula Philippi is one of the most common snails found in the upper intertidal zone of Japan. In Amakusa, some of the population of L. brevicula migrate to the lower zone in the winter, while the rest stay in the upper zone. Thus, during the winter, which is its reproductive season, the population of L. brevicula divides into two sub-populations. This leads to a hypothesis that the migration pattern in winter is genetically controlled and this behavioural dimorphism is maintained by reproductive isolation between the two sub-populations. In order to test this hypothesis, the following three points were investigated: (1) whether the same snails migrate in a similar way every winter, (2) whether there is a significant tidal level preference in snails, and (3) whether reproductive isolation occurs between the two sub-populations. The results showed (1) the migration behaviour of each snail was consistent over two successive winters, i.e. the same group of snails migrated downward every winter and the same group of snails stayed in the upper zone every winter, (2) transplanted snails moved toward the original zones where they were caught, suggesting that the snails actively selected their tidal zone in winter, and (3) most of the snails copulated within each sub-population. Therefore, reproductive isolation between the two sub-populations was considered to be established to some extent by the dimorphic migration behaviour. In conclusion, the migratory behaviour of L. brevicula is determined separately for each individual and might be genetically controlled, and the behavioural dimorphism may be maintained by partial reproductive isolation between the two sub-populations.     

The importance of intraspecific competition in a Littorina littorea population in the Wadden Sea

Two field experiments were carried out totest whether effects of intraspecific competition ina Littorina littorea population can be detectedin a short-term investigation. Different size classesof L. littorea showed no significant differencein preferences when offered four kinds of eitherpossible food or substrata (Fucus vesiculosus,Ulva lactuca, Carcinus maenas, brick).Large and medium winkles preferred Fucusvesiculosus, followed by Ulva lactuca. Deadshore crabs (Carcinus maenas) were the leastpreferred objects for all size classes. On the firstday of the experiment bricks were more attractive tosmall littorines than to larger ones. Considering allfour days, the same ranking occurred for all sizeclasses: Fucus vesiculosus > Ulvalactuca > brick > Carcinus maenas. The reactionofjuveniles to increased densities was examined using anin situ caging experiment on a mussel bed. Meshsize of the cages allowed adult densities to beincreased while juveniles could escape by passingthrough the meshes. However, there was no significantemigration of small winkles even from cages with 10 to20 times natural density of large individuals. Ofgreater importance was the original number of winklesat the site. The available resources on the musselbeds appear to be sufficient to maintain a highpopulation density. Intraspecific competition does notseem to play a major role in this L. littorea-population.     

On the taxonomy of Littorina africana (Mollusca: Gastropoda)

L. africana and L. knysnaensis are regarded as two morphs of a single species which exhibits a genetic cline along the south-eastern coast of southern Africa. The dark brown morph knysnaensis dominates the western, cooler end of the cline and is replaced by the pale blue morph africana at the warmer end of the cline. These conclusions are based on evidence from the latitudinal distributions, the complete range of intermediate forms regarding shell colour and shell morphology and the lack of differences in redular morphology, penial morphology or habitat.     

Phenotypic plasticity in gastropod shell remodeling

A laboratory experiment was conducted with the marine gastropod Littorina littorea from three different sites in Long Island Sound to test for inducible responses to the non‐native predatory crab Hemigrapsus sanguineus. Individuals of L. littorea did not exhibit decreased shell growth in response to increased predation risk, which differs from the response seen in other species of Littorina. However, snails did exhibit slower growth in response to reduced food availability, as seen in other gastropods. Surprisingly, snails from two sites exhibited differences between the crab‐exposed and reduced food‐availability treatments in how they remodeled the thickness of different microstructural layers of shell material in apical parts of the shell. Although individuals of L. littorea did not show the typical shell thickening response seen in other gastropods, they did show differences among populations in remodeling their shell. This underlies the importance of examining how the shell is constructed and remodeled in studies of inducible defenses in gastropods, as well as the importance of examining animals from multiple populations.     

The close relationship between estimated divergent selection and observed differentiation supports the selective origin of a marine snail hybrid zone

To study the role of divergent selection in the differentiation of the two morphs in a hybrid zone of the intertidal snail Littorina saxatilis, we compared the strength of the divergent selection acting on a series of shell characters (as estimated by the viability of snails in a reciprocal transplant experiment) with the contribution of these characters to the phenotypic differences between the morphs. We found a close correlation between selection and differentiation, which suggests a cause-effect relationship, i.e. that all present differentiation is the result of past divergent selection. In addition, divergent selection was a very important component of the total natural selection acting on shell measures. These novel results support previous evidence, based on allozyme analysis, of a parapatric origin for this hybrid zone. We discuss possible limitations of this interpretation and the circumstances under which allopatric differentiation would produce the same results. Phenotypic analysis of divergent selection may be a useful method of investigating the evolutionary mechanisms involved in differentiation processes.     

A preliminary study on genetic differentiation in Littorina saxatilis from Galway Bay,Ireland: Littorina tenebrosa Montagu – a valid species or ecotype?

Littorina tenebrosa is a small fragile-shelled periwinkle which lives on permanently submerged algae in coastal lagoons and non-tidal brackish pools. This periwinkle is a member of the rough periwinkle group which also comprises Littorina saxatilis, L. arcana, L. compressa and L. neglecta and is most closely related to L. saxatilis although its exact systematic status is in some doubt. Based on its unique ecology many believe L. tenebrosa to be a valid species. However, shell morphometric and allozyme analyses on Scottish and Swedish populations of L. tenebrosa and L. saxatilis have indicated that the two periwinkles are virtually identical. Preliminary results on five allozyme loci (AAT-1, GPI, PGM-1, PGM-2 and PNP) in samples of L. tenebrosa and L. saxatilis from Golam Head, Lettermullen, and other locations on the west coast of Ireland show L. tenebrosa to be genetically differentiated from L. saxatilis At Golam Head, where opportunities for gene flow occur between the two taxa, L. tenebrosa is as genetically differentiated from local L. saxatilis as it is from L. saxatilis from more geographically distant locations.     

Intersex and Littorina littorea in Cork Harbour: results of a medium-term monitoring programme

Cork Harbour, located on the south coast of Ireland, is a complex, multi-user environment. In the present study Littorina littorea (L.) is used as a bio-indicator of TBT pollution, as it has a wide range within the harbour, unlike the more common bio-indicator, Nucella lapillus (L.), which is limited to rocky substrata near the mouth of the harbour. Intersex in L. littorea is used for the first time in Ireland as a medium-term indicator of TBT pollution and results show that there are no deviations in levels of intersex over the three year period from 1994–1996. These results are discussed.