Thermal tolerance,evaporative water loss,air-water oxygen consumption and zonation of intertidal prosobranchs: a new synthesis

Duration of emergence increases with tidal height on rocky shores therefore, emergence adaptations in intertidal species such as littorine and other prosobranch gastropods have been considered correlated with zonation patterns; temperature tolerance, desiccation resistance and aerial respiration rate all commonly assumed to increase progressively with increasing zonation level. Such direct correlations are rarely observed in nature. Maximal aerial gas exchange occurs in mid-shore, not high shore species. Temperature tolerance and desiccation resistance do not increase directly with shore height. Thus, hypotheses regarding physiological correlates of zonation require revaluation. A new hypothesis is presented that the high tide mark presents a single major physiological barrier on rocky shores. Above it, snails experience prolonged emergence and extensive desiccation; below it, predictable submergence and rehydration with each tidal cycle. Thus, desiccation stress is minimal below the high tide mark and maximal above it. Therefore, species restricted below high tide (the eulittoral zone) should display markedly different adaptive strategies to emergence than those above it (the eulittoral fringe). A review of the literature indicated that adaptations in eulittoral species are dominated by those allowing maintenance of activity and foraging in air including: evaporative cooling; low thermal tolerance; elevated aerial O₂ uptake rates; and high capacity for radiant heat absorption. Such adaptations exacerbate evaporative water loss. In contrast, species restricted to the eulittoral fringe display adaptive strategies that minimize desiccation and prolong survival of emergence including: foot withdrawal, preventing heat conduction from the substratum; aestivation in air; elevated thermal tolerance reducing necessity for evaporative cooling; position maintenance by cementation to the substratum and increased capacity for heat dissipation. In order to test of this hypothesis the upper thermal limits, tissue and substratum temperatures on emergence in direct sunlight and evaporative water loss and tissue temperatures on emergence in 40 °C were evaluated for specimens of six species of eulittoral and eulittoral fringe gastropods from a granite shore on Princess Royal Harbour near Albany, Western Australia. The results were consistant with adaptation to the proposed desiccation barrier at high tide. The eulittoral species, Austrocochlea constricta, Austrocochlea concamerata, Nerita atramentosa and Lepsiella vinosa, displayed adaptations dominated by maintenance of activity and foraging during emergence while the eulittoral fringe littorine species, Bembicium vittatum and Nodilittorina unifasciata displayed adaptations dominated by minization of activity and evaporative water loss during emergence. The evolution of adaptations allowing tolerance of prolonged desiccation have allowed littorine species to dominate high intertidal rocky shore gastropod faunas throughout the world's oceans.     

Horizontal distribution of the plankton rotifers Keratella cochlearis (Bory de St Vincent) and Polyarthra vulgaris (Carlin) in a small eutrophic lake

The plankton rotifers Keratella cochlearis and Polyarthra vulgaris were sampled at 10 cm below the surface at different distances from two dissimilar shores and in the centre of a small eutrophic lake. Light and depth were measured at each sampling point. In each case the numbers of rotifers per liter increased with distance from the shore. There was a significant correlation between the numbers for the two species for the two shores, but none in the centre. In the case of one shore there was a strong correlation between rotifer numbers and supra-surface ambient light.     

Establishment of the Baikalian endemic amphipod Gmelinoides fasciatus Stebb. in Lake Ladoga

Gmelinoides fasciatus Stebb., a small amphipod from the Lake Baikal basin, was discovered in July 1988 in Lake Ladoga, the largest European lake. G. fasciatus likely invaded Lake Ladoga as a consequence of its intentional introduction, aimed at enhancing fish production, in some Karelian Isthmus lakes close to Lake Ladoga's western shore in the early 1970's. Benthos studies conducted in 1989 and 1990 revealed that G. fasciatus was well established in littoral communities along the western and northern shores of Lake Ladoga. G. fasciatus was the dominant species in these littoral communities and contributed over 70% of the macroinvertebrate biomass. The species was abundant in different macrophyte beds and stony littoral areas, both in heavily polluted and undisturbed sites. The maximum abundance and biomass of G. fasciatus was about 54,000 ind m^–2 and 160 g (wet wt.) m^–2. Negative impacts of G. fasciatus on native species have been observed. Studies are needed to identify the effects of G. fasciatus on the functioning of littoral communities and to predict and control the spread of this amphipod.     

The complexity of phenotypic plasticity in the intertidal snail Nodilittorina australis

The intertidal snail Nodilittorina australis has distinct nodulose and striate morphs, which are plastic, developmental responses to sloped and vertical rocky shores, respectively. The nodulose morph cools more quickly, which should be advantageous on the sunnier, sloped shores. In contrast, the striate morph responds more quickly to submersion, which should be advantageous on the more wave-intense vertical shores. Transplant experiments revealed that even large snails can alter development of new shell. The developmental transitions indicate partial dissociation of colour and sculpture. However, the incomplete association of component traits is seen in very few natural populations, indicating that canalization of nodulose and striate pathways has evolved to the extent that is adaptively relevant. There is a distinct asymmetry of the lability of development, whereby nodulose snails are much more likely to switch to development of striate shell than the reverse; this directionality occurs naturally as well as in transplant experiments. Rather than representing an underlying constraint, the asymmetry probably reflects a greater advantage of nodules in smaller snails. The only clear developmental constraint affecting this dimorphism is the association of the production of nodules with slower growth, an expected consequence of the rate-limiting process of shell deposition.     

Effects of sewage pollution on the structure of rocky shore macroinvertebrate assemblages

The urgency to find efficient indices and indicators to prevent further deterioration of coastal areas is one of the hot topics in today’s scientific publication. However, a detailed knowledge of community responses to anthropogenic impacts is essential to sustain those indices. The studies on the response of benthic community to sewage pollution on intertidal rocky shores are generally based on visual census and do not take into account the tidal levels. In order to fulfil this gap in this study: (i) the sampling was performed by destructive sampling, with all individuals identified to the species level; (ii) the sampling was done at all levels of the intertidal (sublittoral fringe, eulittoral, and littoral fringe). Sewage pollution changed the environmental variables and the abundance of macroinvertebrates, being Mytilus galloprovincialis, Melarhaphe neritoides, and Chthamalus montagui the species most responsible for the dissimilarities observed. Effects were different on the three intertidal zones: community structure changed in the sublittoral fringe; suspension-feeders abundances and species richness increased in the eulittoral; no differences were detected in the littoral fringe. Moreover, the results confirm that the presence of sewage discharges tended to benefit suspension feeders, and that the sensitive species were replaced by opportunistic ones.     

Orientation of the sandhopperTalitrus saltator (Amphipoda,Talitridae) living on dynamic sandy shores

Summary The littoral sandhopperTalitrus saltator Montagu maintains its position along the beach mainly by means of an innate solar orientation. Different populations show different directional tendencies depending on the direction of the sealand axis. Experiments were designed which aimed to disentangle the innate and acquired components of the escape orientation. Therefore, the solar orientation of experienced adults living on shores subject to fluctuations of the shoreline and of their laboratory-born (inexperienced) offspring were compared. The innate sun compass orientation shown by the young is modified by learning during life under natural conditions, depending on the variability of the shore. Results showed: (1) an improvement in the solar orientation in individuals living on relatively constant shores; (2) a loss of precision in individuals living on highly dynamic shores; (3) a change to a new ecologically efficient escape direction in individuals living on a relatively stable lagoon shore.Abbreviation TED theoretical escape direction     

The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

A field investigation into the effects of interspecific competition on the distribution and cover of Ulothrix flacca

Ulothrix flacca is a filamentous green alga found in its greatest abundance in the littoral fringe of the rocky intertidal during the winter months. It is much less abundant below this level. This experiment was undertaken tot determine if the lower abundance below the littoral fringe is due to the effects of interspecific competition or physiological limitations. Clearing experiments were performed on the Coast of Maine, USA to eliminate competition. Belts 10 cm wide that ran from the littoral fringe downward into the eulittoral zone were cleared of all attached organisms down to bare rock. A second condition was set up in which only the brown algal canopy was removed from a 60 cm wide belt. Unaltered belts were designated as controls. Squares 0.25 m² in area were cleared in the littoral fringe and eulittoral zone. U. flacca was able to colonize the cleared areas of the eulittoral zone and reach abundances equal to those found in the unaltered control areas of the littoral fringe. This indicates that physiological limitations do not restrict the abundance of U. flacca in the eulittoral zone. However, evidence of grazing in cleared areas adjacent to high concentrations of the barnacle Semibalanus balanoides suggest that competition alone is not responsible for the limited abundance of U. flacca in the eulittoral zone. Herbivores associated with the presence of barnacles may also have an impact on the abundance of U. flacca, even during the winter months.     

Maintenance of zonation patterns in two species of flat periwinkle,Littorina obtusata and L. mariae

The zonation patterns of Littorina obtusata (L.) and Littorina mariae Sacchi et Rastelli were shown to be quite distinct on a sheltered rocky shore. L. obtusata was found at all the heights sampled; it reached peak numbers at mid shore on the alga Ascophyllum nodosum L. (Le Jol). There was no difference in the tidal height occupied by adults or juveniles; or in the mean size of L. obtusata along the vertical gradient of the shore. In contrast L. mariae occurred exclusively low on the shore, on Fucus serratus L. Translocation of the two species within their respective levels resulted in random movement after 4 days, although initial movements after 1 and 2 days were sometimes directional. Animals transplanted to the normal level of the other species showed directional movement towards their home zone; this was most pronounced after 4 days. There was no difference in the distance moved by the two species, although the distance moved did vary with tidal height, both species moving further at mid shore than low shore. Distances moved by littorinids at replicate areas in the low shore were similar but those at mid shore did vary. There was an interaction between the species and the different tidal heights which revealed that transplanted species moved further than translocated species at the same tidal level. However, this was only significant in the case of L. mariae. It is suggested that the close relationship between the winkles and their host algae may direct the homing behaviour of displaced individuals.     

Oxidative processes as indicators of chemical stress in marine bivalves

Deleterious effects of environmental contaminants could be due to enhanced prooxidant forces overcoming antioxidant defences. Before practical biomarkers based on free radical biology will be generally accepted and validated in situ, additional research is required concerning normal physiological and environmental influences on the relevant systems. The aims of this study were to evaluate in situ the importance of oxyradical production in the presence and absence of pollutants and to characterize some antioxidant systems in Mytilus edulis L. Specimens of M. edulis L. were transplanted from a reference site (Franquelin) to Baie Comeau (Baie des Anglais), on the North shore of the St. Lawrence maritime estuary, where are found aluminium and pulp and paper plants. An oxidative stress was observed in mussels submitted to a chronic exposure in the polluted environment. Variations of pro-and anti-oxidant molecules involved in oxidative processes were related in part to seasonal and physico-chemical influences. Catalase activity, malondialdehyde and glutathione concentrations will be useful as biomarkers of stress in situ since they react to anthropogenic influence and to abiotic factors such as emersion period and temperature.     

Distribution of the species of rough periwinkle (Littorina) in Great Britain

The status of the four currently recognised species in the Littorina saxatilis species-complex, i.e. Littorina nigrolineata Gray, L. arcana Hannaford Ellis, L. saxatilis (Olivi) and L. neglecta Bean is reviewed briefly, with notes on their characteristic features and location on the shore. Since the taxonomy of these rough periwinkles has only become stable relatively recently much of the previously published information on distribution is of little or no use. In this paper their distribution around the coastline of England and Wales (with some notes on Scotland) is described and discussed. L. saxatilis is found in a wide range of habitats from exposed peninsulas to estuaries on all shores where there is a suitable rocky or stony substrate; also in salt marsh pools. L. arcana has a more restricted distribution and is notably absent along much of the south English coast, central Cardigan Bay and possibly northern Scotland; it is not found in estuaries. L. nigrolineata has an even more restricted distribution, although it occurs both on exposed coasts and in estuaries; it has only been found by us in one locality on the east coast. L. neglecta is probably fairly widely distributed but we have few details so far. The implications of the different patterns of distribution are discussed.     

The thermal and energetic significance of clustering in the speckled mousebird,Colius striatus

Summary The effect of clustering behaviour on metabolism, body temperature, thermal conductance and evaporative water loss was investigated in speckled mousebirds at temperatures between 5 and 36°C. Within the thermal neutral zone (approximately 30–35 °C) basal metabolic rate of clusters of two birds (32.5 J·g^-1·h^-1) and four birds (28.5 J·g^-1·h^-1) was significantly lower by about 11% and 22%, respectively, than that of individuals (36.4 J·g^-1·h^-1). Similarly, below the lower critical temperature, the metabolism of clusters of two and four birds was about 14% and 31% lower, respectively, than for individual birds as a result of significantly lower total thermal conductance in clustered birds. Body temperature ranged from about 36 to 41°C and was positively correlated with ambient temperature in both individuals and clusters, but was less variable in clusters. Total evaporative water loss was similar in individuals and clusters and averaged 5–6% of body weight per day below 30°C in individuals and below 25°C in clusters. Above these temperatures total evaporative water loss increased and mousebirds could dissipate between 80 and 90% of their metabolic heat production at ambient temperatures between 36 and 39°C. Mousebirds not only clustered to sleep between sunset and sunrise but were also observed to cluster during the day, even at high ambient temperature. Whereas clustering at night and during cold, wet weather serves a thermoregulatory function, in that it allows the brrds to maintain body temperature at a reduced metabolic cost, clustering during the day is probably related to maintenance of social bonds within the flock.Abbreviations BMR basal metabolic rate - bw body weight - C _totab total thermal conductance - EWI evaporative water loss - M metabolism - RH relative humidity - T _a ambient temperature - T _b body temperature - T _ch chamber temperature - T _cl cluster temperature - TEWL total evaporative water loss - LCT lower critical temperature - TNZ thermal neutral zone     

Factors limiting the distribution of the annual legumeKummerowia striata in a river floodplain

The annual legumeKummerowia striata often predominates in river floodplains forming a large population, but it is rare in gravelly areas adjacent to it. In order to clarify the factors limiting the distribution ofK. striata, its distribution pattern, germination and survival of seeds sown artificially were studied in the floodplain of the Ohta River. There was a positive correlation between the cover value ofK. striata and the percentage of fine particles (<0.2 mm) in the soil. In the winter season, there was a number of seeds (up to 47 seeds 20 cm⁻²) on the ground where aK. striata population had existed in the previous year, but they were very sparse in areas apart from the population. Only a small proportion of seeds sown in the gravelly area germinated. Many of the germinated individuals died before the first leaf unfolded, but the rest survived to maturity despite repeated floodings. Plants germinated in the gravelly area and those inoculated with soil from the gravelly area bore nodules, showing the presence ofRhizobium in the gravelly area. It was concluded that the colonization ofK. striata in the gravelly area was limited by slow dispersal, and by a low percentage of germination and high mortality in the early stage of establishment.     

Biotic zonation on rocky shores of Heard Island

Summary The zonation of rocky shore biota on Heard Island is described for the first time, related to a universal zonation scheme and compared with that recorded for other sub-Antarctic localities. The Kelp Zone of holdfasts of the giant kelp, Durvillea antarctica (Chamisso) Hariot (one of the characteristic features of these regions) is confirmed as a sublittoral fringe. The occurrence of a Bare Zone within the littoral zone on sub-Antarctic shores is discussed. The possible roles of submersion, spray, freezing and predation by gulls in influencing the extent and composition of zones are discussed.     

Spatial patchiness of epilithic biofilm caused by refuge-inhabiting high shore gastropods

Aggregations of Melarhaphe neritoides and Littorina saxatilis are common on the high shore in north-east England. These aggregations are frequently found in crevices which act as a refuge from desiccation. The aggregations are surrounded by patches of epilithic biofilm which are visibly lighter in colour than the biofilm present on the remainder of the high shore. Chlorophyll a levels were lower in regions of close proximity to aggregated littorinids (mean = 4.9 μg cm⁻²) as compared to areas >‰5 cm away from the edge of the visible colour change (mean = 15.9 μg cm⁻²). Manipulations of littorinid density showed that areas of high density had significantly lower levels of chlorophyll a than those where littorinids were excluded. This difference was mainly due to the presence of higher numbers of small grazed areas or an increase in size of grazed areas rather than a homogenous change in chlorophyll a levels across the entire manipulated area. These results are supported by observations showing that only 7% of the total foraging time of littorinids was spent outside visibly lighter patches of biofilm suggesting littorinids only control local biofilm levels through grazing.This paper has not been submitted elsewhere in identical or similar form, nor will it be during the first three months after its submission to Hydrobiologia     

Thoughts on the ecology and evolution of the intertidal biota of the Azores and other Atlantic islands

The special features of the intertidal ecosystems of remote islands are reviewed briefly before focusing on the littoral zone of Macaronesia (Azores, Madeira and Canaries). Distribution patterns are briefly compared with other European shores. Species missing from the Azores compared with Madeira, the Canaries and continental European shores are listed. The degree of genetic differentiation within and amongst selected important species of gastropods (Patella spp.) and barnacles (Chthamalus spp.) in Macaronesia is reviewed. The role of isolated islands in allopatric speciation of these organisms is discussed in relation to dispersal. Possible threats to the populations and communities of rocky shores in Macaronesia are considered in the context of the endemic nature of certain species. The importance of studies of basic ecology and genetics to inform resource management and conservation is highlighted. Future research directions are indicated, emphasizing the usefulness of Macaronesia as a model system to explore speciation in intertidal organisms.     

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.     

白及种质资源及其近缘种的SSR指纹图谱研究

白及(Bletilla striata Rchb.)为中国珍稀濒危药用植物,野生资源枯竭,混伪种繁多。该研究基于SSR标记技术对16个地区48份白及(Bletilla striata Rchb.)样品和4份黄花白及(Bletilla ochracea Schltr.)、4份小白及(Bletilla formosana(Hayata)Schltr.)、4份华白及(Bletilla sinensis(Rolfe)Schltr.)样品进行遗传多样性分析,构建白及种质资源的SSR指纹图谱,并对60份白及样品进行种质资源鉴定,分析白及属种内及其种间的遗传分化特征。结果表明:(1)20对白及SSR引物均能在4个白及属植物中成功扩增,条带清晰且多态性丰富,每对引物的复等位基因数(Na)在4~9之间,等位基因数(Na)总和为127,平均为6.35。(2)筛选出基因型丰富、多态性信息含量(PIC)高的7对白及SSR引物(BJSSR01、BJSSR14、BJSSR15、BJSSR16、BJSSR18、BJSSR19、BJSSR22)构建的白及SSR指纹图谱能将白及属各种质资源清楚分开。(3)白及属在种间水平均有较高的遗传多样性(Na=6.35,I=1.429 1,h=0.706 8),物种间遗传分化强烈(Gst=0.44),物种间的基因流较弱(Nm=0.475 3)。(4)UPGMA聚类分析表明,60份供试样品明显聚为4大支,同一物种的个体首先聚在一起,这与形态学分类基本一致;不同来源地的样品种内和种间的亲缘关系有明显差异,地理距离较近的白及样品具有较近的遗传关系。