Intertidal ecology and potential power impacts, Bay of Fundy, Canada

A large-scale tidal power project in the upper reaches of the Bay of Fundy would cause major changes in the tidal elevations of the headpond which would have a pronounced effect on intertidal ecology. The new mean low water level would be, on average, slightly above the present mean sea level whilst the new mean high water level would drop about 0.7 m. This would result in a much compressed intertidal zone. Mud and sand flats, below the present mean sea level, would be converted to sub-tidal habitat, whilst the high saltmarsh, flooded at present only on the highest spring tides, would become colonized by terrestrial vegetation. There would be major shifts in the relative importance of primary production by saltmarshes, intertidal microalgae and phytopiankton, but overall, it is likely that production would increase. Changes in both primary food sources and habitat area would affect the abundance of intertidal invertebrate organisms and the fish and migratory shorebirds that feed upon them.     

Autecology of an intertidal Alaskan harpacticoid copepod Apolethon sp.

The life history and ecology of a harpacticoid copepod, Apolethon sp., were investigated for one year in the intertidal zone of a subarctic bay in southeastern Alaska. Core samples were collected biweekly at the 0 m tidal level from March, 1992 through March, 1993; zonation and vertical distribution were determined from additional samples. Large seasonal variations in density of Apolethon sp. parallelled that of the total harpacticoid community, with highest densities (124.3±12.5 10 cm^–2; mean ± standard error) in March and lowest densities in May and October (7.6±4.5). The sex ratio of Apolethon sp. varied directly with population density, varying from <0.1 to 1.2 (male:female) during the study period. The species was restricted to a relatively narrow vertical band within the intertidal zone, having its greatest density at – 1 m and being rare above + 1 m tidal height. Adult Apolethon sp. bear conspicuous ovoid mucin structures located laterally on the genital double-somite. The mucin structures are unique to the species; several putative functions are discussed.     

Spatial and temporal variation of photosynthesis in intertidal Mazzaella laminarioides (Bory)Fredericq (Rhodophyta, Gigartinales)

The red alga Mazzaella laminarioides is an economically important species with an extended latitudinal distribution along the Chilean coast. Its populations form mid-intertidal stands, several meters wide, and therefore are differentially exposed to environmental variables that result in temporal and spatial variability in productivity. We evaluated the effect of latitude and intertidal height on productivity by in situ measurement of photosynthetic performance. Daily and seasonal variations of O₂-evolution rate and maximal quantum yield (F _v/F _m) were determined in plants from the upper and lower intertidal zone at two localities 1500,km apart. Results suggest that plant responses were mainly affected by irradiation, temperature and desiccation. At local level, upper intertidal plants showed a reduced photosynthetic rate and quantum efficiency as compared to those displayed by plants from the lower intertidal, indicating their higher level of excitation energy acclimation. Stronger acclimation differences between upper and lower intertidal plants were observed in spring and summer. Differences in photosynthetic parameters between reproductive phases were recorded in autumn and winter, regardless of the position of the individuals in the intertidal zone. The effects of tidal elevation on seasonal patterns of photosynthesis were also influenced by latitude. Seasonal variation in photosynthetic efficiency was observed in plants from the northern population at both intertidal elevations, but only at the upper intertidal level in the southern population. This study shows that production variability in M. laminarioides results from differences in the intensity of environmental factors observed seasonally at local (intertidal) and latitudinal scales.     

An examination of species dispersion patterns along the intertidal gradient on Macquarie Island (sub-Antarctica) using a restricted occupancy model

It has been widely held that intertidal zonation boundaries and the tidal emersion levels are causally related: the critical tidal level hypothesis. In order to evaluate this hypothesis, the dispersion patterns of species boundaries on the intertidal rocky shores of Macquarie Island (sub-Antarctica) were examined using a restricted occupancy model to test the null hypothesis that the observed species were randomly dispersed along the tidal emersion gradient. Subsequently this investigation demonstrated that the intertidal species boundaries on Macquarie Island shores were randomly dispersed with respect to the tidal emersion gradient. Hence no prima facie evidence was found to support the critical tidal level hypothesis. This suggests that tidal emersion is not a significant factor structuring intertidal communities on Macquarie Island.     

Characterization of tidal pool algae in the Mexican Tropical Pacific coast

Tidal pools in the Mexican Tropical Pacific coast have received relatively little attention in spite of their considerable richness in species and wide distribution in the region.This paper presents the first characterization of the algal flora of this region. It analyzes the number and composition of species of the tidal pools of six localities with regard to geographical distribution and its seasonal variations as well as tidal level. 97 species are reported, 25 Chlorophyta, 23 Phaeophyta, 34 Rhodophyta and 15 Cyanophyta.Of that total of species, 63% were found in one locality, 23.7% in two, 11.3% in three and 1 % in 4 or 5 localities. Not one species was common to all of the pools.The highest number of species was found on pools of the middle and low intertidal where the Chlorophyta, Rhodophyta and Phaeophyta were the most abundant algae. Cyanophyta was more common in the supralittoral and high intertidal pools.     

Zonation of intertidal macrobenthos in the estuaries of Schelde and Ems

Based on data, collected in 1980–1990, the intertidal benthic macrofauna of the Schelde and Ems estuaries was compared. The spatial occurrence of the benthic macrofauna along the salinity gradient, including the freshwater tidal area was emphasized. Both estuaries appeared to have a very similar species composition, especially at genus level. The higher number of species observed in the Schelde estuary was probably due to a greater habitat diversity. In both estuaries species diversity decreased with distance upstream. The total density did not vary along the estuarine gradient, whereas biomass is highest in the polyhaline zone.In both estuaries distinct intertidal benthic communities were observed along the salinity gradient: a marine community in the polyhaline zone, a brackish community in the mesohaline zone, and a third community in the oligohaline and freshwater tidal zones of the estuary. These three communities were very similar between both estuaries. Their main characteristics were discussed together with the occurrence and distribution of the dominant species.For the Schelde estuary and to a lesser extent also for the Ems estuary, there was evidence that anthropogenic stress had a negative effect on the intertidal macrobenthic communities of the oligohaline/freshwater tidal zone. Only Oligochaeta were dominating, whereas the very euryhaline and/or true limnetic species were missing. In the mesohaline zone, the Schelde estuary was dominated by large numbers of short-living, opportunistic species, whereas in the Ems estuary relatively more stable macrobenthic communities were observed. A comparison with some other European estuaries showed in general similar trends as those observed for the Schelde and Ems estuaries.     

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.     

Thermal constraints on daily patterns of aggregation and density along an intertidal gradient in the periwinkle Echinolittorina peruviana

Variations in environmental temperature have both direct and indirect effects that affect organisms at levels ranging from intra-cellular physiological processes to ecological patterns. These variations are especially important for intertidal marine ectotherms such as littorinids since they alternate between periods of immersion in seawater, and must also experience long periods of emersion. In central Chile, Echinolittorina peruviana is one of the most conspicuous species on rocky intertidal shores, occurring at high tidal levels and in the splash zone. The species is known to resist direct exposure to the sun for long periods, although juveniles tend to be restricted to protected microhabitats. Adults show seasonal variations in abundance and vertical distribution and may form aggregations that have been shown to help reduce water loss and body temperature. In this study, we evaluate the relationship between daily thermal variations throughout the vertical distribution of this species and how these affect the patterns of density and aggregation. Our results suggest that one of the leading determinants of the spatio-temporal variation of density in E. peruviana may be operative temperature (T_O: the amount of stored heat resulting from the balance between heat fluxes into and out of the body, measured with taxidermic mounts mimicking heat transfer properties of the snail). T_O showed a strong negative relationship with density and a strong positive relationship with aggregation in the highest intertidal level monitored. The strength of these relationships decreased in importance at lower levels. While T_O alone cannot explain the abundance of E. peruviana throughout its range of distribution, our results show that it does have a strong influence that should be considered in addition to other ecological factors affecting the density of intertidal littorinids.     

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.     

Empty habitat in coastal sediments for populations of macrozoobenthos

Species with wide dispersal are expected to have little empty habitat. This was tested by analysing habitat use in the Wadden Sea near the island of Sylt (North Sea). Sampling covered 222 intertidal sites spread across the K?nigshafen tidal flats (4.5 km², mapping approach conducted once) and 270 subtidal sites along 12 km of a tidal channel system (stratified random sampling in spring and autumn). For any single species, 'suitable habitat' was extrapolated from the ranges of water depth and sediment composition present at the ten sites with the highest frequency or abundance of the species. On average, macrobenthic species actually used less than half of the suitable sites within the scale of local populations; this was far less than expected from a lognormal distribution. In the subtidal, abundance of most species changed between the two sampling seasons. A 50% increase in overall abundance was accompanied by a decrease in empty habitat of only about 25%. Thus, a doubling of abundances would not fill the empty space but just of half of it. The polychaete Scoloplos armiger was an exception in occupying almost all of its suitable habitat in the intertidal and subtidal sediments. A distinct patch of high species richness occurred where flood waters persistently form a large gyre which may enhance larval settlement. We suggest that limitations to larval settlement and/or juvenile survival primarily cause the extent of the observed habitat emptiness. Electronic Publication     

Intraspecific variation in the population dynamics and growth of the limpet,Cellana tramoserica

Summary Populations of the limpet Cellana tramoserica (Sowerby) from high, mid, and low intertidal regions, and from a subtidal zone, were studied at Cape Banks (N.S.W.), Australia. Individuals from the subtidal population had the largest mean and maximum shell size, the low and high shore populations were intermediate, while the midshore population had both the smallest mean and maximum size. The density of adults showed the reverse trend: the midshore region had the greatest adult population density while the subtidal population had the smallest density. The density of juveniles and recruits was negatively correlated with tidal height in the intertidal areas, but the density of both age-classes was smallest in the subtidal region. The rate of growth of individuals was negatively correlated to the adult density of a site; individuals from the subtidal population grew the fastest, while those from the midshore region grew the slowest. The level of adult mortality of the four populations was similar, with an annual rate of between 50–60%. Juvenile mortality did, however, differ among populations; reduced percentages of juveniles reached adult size in the mid and low shore populations than in the highshore and subtidal populations.The expected lifetime fecundity of individual females differed among the four populations. Subtidal individuals were expected to spawn 40 times the amount of gonad material during their life than were individuals from the midshore population. Individuals from the high and low populations probably spawn 10 times more than those from the midshore region. Differences in the fecundities of individuals were not likely to be the result of genetic differences because experimental transplants and manipulations of density showed that individuals from all the intertidal populations could increase their growth rate to match that of subtidal individuals. The implications of the difference in lifetime fecundity among populations are discussed.     

Holocene reef evolution in a macrotidal setting: Buccaneer Archipelago,Kimberley Bioregion,Northwest Australia

This study uses information derived from cores to describe the Holocene accretion history of coral reefs in the macrotidal (up to 11 m tidal range) Buccaneer Archipelago of the southern Kimberley coast, Western Australia. The internal architecture of all cored reefs is broadly similar, constituting well-preserved detrital coral fragments, predominantly branching Acropora, in a poorly sorted sandy mud matrix. However, once the reefs reach sea level, they diverge into two types: low intertidal reefs that maintain their detrital character and develop relatively narrow, horizontal or gently sloping reef flats at approximately mean low water spring, and high intertidal reefs that develop broad coralline algal-dominated reef flats at elevations between mean low water neap and mean high water neap. The high intertidal reefs develop where strong, ebb-dominated, tidal asymmetry retains seawater over the low tide and allows continued accretion. Both reef types are ultimately constrained by sea level but differ in elevation by 3–4 m.     

Patterns in the utilization of the intertidal salt marsh by larvae and juveniles of Fundulus heteroclitus (Linnaeus) and Fundulus luciae (Baird)

Larval and juvenile fishes were collected at low tide from the surface of an intertidal salt marsh on Sapelo Island, Georgia every 6 days from 25 May through 20 December 1982. Larval fishes were present in shallow puddles of tidal water on the marsh from the beginning of the sampling period until the end of October. Juveniles were present throughout the year. Fundulus heteroclitus (Linnaeus) and F. luciae (Baird) accounted for 96.3% (67.0% and 29.3%, respectively) of the 4355 fishes collected. These fishes exhibited synchronous, temporal pulses in larval abundance, suggesting that reproduction was discontinuous and controlled by the same factor(s) in both species. Larval F. heteroclitus inhabited aquatic micro-habitats near the upland edge of the intertidal marsh, but as the larvae grew to juvenile size (≈ 10 mm standard length) they moved to lower elevations near a tidal creek. Large juveniles and adults of F. heteroclitus infrequently occurred in the samples, presumably because they leave the intertidal marsh as the tide ebbs. F. luciae were rarely found in low marsh areas, but all age classes (including adults) occurred at higher elevations, supporting previous suggestions that this species prefers high marsh habitats. The vegetated, intertidal salt marsh appears to be the principal nursery habitat for both of these cyprinodontid species.     

Effect of salinity changes on the bacterial diversity, photosynthesis and oxygen consumption of cyanobacterial mats from an intertidal flat of the Arabian Gulf

The effects of salinity fluctuation on bacterial diversity, rates of gross photosynthesis (GP) and oxygen consumption in the light (OCL) and in the dark (OCD) were investigated in three submerged cyanobacterial mats from a transect on an intertidal flat. The transect ran 1 km inland from the low water mark along an increasingly extreme habitat with respect to salinity. The response of GP, OCL and OCD in each sample to various salinities (65 per thousand, 100 per thousand, 150 per thousand and 200 per thousand) were compared. The obtained sequences and the number of unique operational taxonomic units showed clear differences in the mats' bacterial composition. While cyanobacteria decreased from the lower to the upper tidal mat, other bacterial groups such as Chloroflexus and Cytophaga/Flavobacteria/Bacteriodetes showed an opposite pattern with the highest dominance in the middle and upper tidal mats respectively. Gross photosynthesis and OCL at the ambient salinities of the mats decreased from the lower to the upper tidal zone. All mats, regardless of their tidal location, exhibited a decrease in areal GP, OCL and OCD rates at salinities > 100 per thousand. The extent of inhibition of these processes at higher salinities suggests an increase in salt adaptation of the mats microorganisms with distance from the low water line. We conclude that the resilience of microbial mats towards different salinity regimes on intertidal flats is accompanied by adjustment of the diversity and function of their microbial communities.     

Root morphology is related to the phenotypic variation in waterlogging tolerance of two populations of Suaeda salsa under salinity

The effects of waterlogging and salinity on seedling emergence, seedling growth and ion accumulation in a euhalophyte Suaeda salsa in an intertidal zone and on saline inland soil were investigated. Seedlings of S. salsa from the intertidal zone emerged more rapidly than those of the inland population under both waterlogged and drained conditions. Waterlogging and salinity had no adverse effects on seedling emergence of S. salsa from the intertidal zone, but markedly inhibited this parameter in the inland population. Waterlogging did not affect the seedling survival, shoot dry mass, and shoot height in high salinity in S. salsa from the intertidal zone, while the opposite trend was shown in the inland population. The root dry mass was higher in S. salsa from the intertidal zone as compared to the inland population, in waterlogged treatments by 1.9, 1.3, and 1.5 times in 1, 200, and 600 mM NaCl, respectively, and in drained treatments by 1.8, 2.3, and 3.0 times in 1, 200, and 600 mM NaCl, respectively. Waterlogging increased Na⁺ and K⁺ concentrations in high salinity, but waterlogging had no effect on Cl^- concentration in shoots of S. salsa from the intertidal zone. In all NaCl treatments, waterlogging had no effect on concentrations of these ions in shoots of S. salsa from the saline inland site. In a field investigation, the fresh mass of shoots and roots were lower, whereas the root/shoot ratio was 1.5 times higher in S. salsa from the intertidal zone, compared with the inland population. These findings indicate that S. salsa population from the intertidal zone is more waterlogging tolerant than the inland population. S. salsa from the intertidal zone produced relatively more root biomass and this might help anchor plants against tidal action in the intertidal zone. The physiological and morphological characteristics may determine the natural distributions of the two S. salsa populations in their different saline environments.     

Physical habitat attribute mediates biotic resistance to non-indigenous species invasion

A soft-shelled non-indigenous clam, Nuttallia obscurata, has invaded coastal soft-sediment habitats of the northeastern Pacific. In a survey of 35 sites within the San Juan Islands, Washington, USA, Nuttallia was found almost exclusively in sandy substrates, higher in the intertidal than most native clams (>1 m above mean lower low water). The distinctive distribution of Nuttallia suggested that tidal height and sediment composition may be important physical factors that control its refuge availability, regulating its exposure to predation and ultimately the success of its invasion. I tethered Nuttallia for 24 h in the high intertidal where it is typically found and in the low intertidal at an elevation where it was never found. Clams restrained to the surface suffered high mortality from crab predation at both tidal heights, whereas control clams with unrestricted burrowing movement exhibited high mortality rates only in the low intertidal. In a second experiment, I transplanted sediment within and between the two intertidal heights to measure effects of tidal height and sediment type on survival and burial depth of Nuttallia. At both tidal heights all clams placed on mud-cobble substrate, naturally common in the low intertidal, suffered high mortality rates (>60% in 24 h). Nuttallia on loosely packed sand substrate, naturally found in the upper intertidal, survived much better, however, because they buried deeper than in the tightly packed mud. Caged control clams at both tidal heights suffered no mortality. Apparently native predators are mitigating community level impacts of an invader by excluding Nuttallia completely from some beaches with improper sediment characteristics or relegating it in others to a zone not often inhabited by native species, thereby reducing potential competitive interactions. These findings show that a physical habitat characteristic can mediate biotic resistance to an invader and thus control invasion success and community-level impacts. Generally, such physical-biological interactions may explain some of the reported site-to-site variability in invasion success, as well as the patchy distribution of many soft-sediment infaunal species.     

模拟海平面上升对红树植物秋茄的影响

研究了壤质沙土 (粗质土 )和粘土 (细质土 )条件下红树植物秋茄 (K andelia candel)对水位上升和淹水时间延长的反应。模拟海平面上升 30 cm导致红树林土壤的酸化 ,且细质土的酸化比粗质土严重 ;秋茄繁殖体的萌苗速度明显加快 ;促进秋茄的早期生长 ,尤其是导致最初 2个月茎高生长的增加 ,然而 ,后 2个月秋茄的相对生长率并不因水位的升高而增加 ;地下部 /地上部生物量比减小 ,在粗质土中尤为如此 ;幼苗粗根比例明显增加 ;叶片叶绿素 a/ b比值下降。在微型盆栽试验条件下 ,无论是高水位还是低水位 ,所有的秋茄繁殖体均成功萌发且幼苗在整个试验期间均成活。在野外条件下 ,秋茄幼苗成活率在高水位和低水位条件下均高达 90 %以上。野外条件下 ,无论是经胚轴萌发还是幼苗移栽的幼苗 ,最初 4个月的茎高生长均为低潮区高于高潮区 ,与微型试验结果相同。微型盆栽试验和野外种植试验均表明 ,海平面上升 30 cm对秋茄的萌发和早期生长具有促进作用     

Estuaries as Filters: The Role of Tidal Marshes in Trace Metal Removal

Flux calculations demonstrate that many estuaries are natural filters for trace metals. Yet, the underlying processes are poorly investigated. In the present study, it was hypothesized that intertidal marshes contribute significantly to the contaminant filter function of estuaries. Trace metal concentrations and sediment characteristics were measured along a transect from the subtidal, over an intertidal flat and marsh to a restored marsh with controlled reduced tide. Metal concentrations in the intertidal and restored marsh were found to be a factor two to five higher than values in the subtidal and intertidal flat sediments. High metal concentrations and high accretion rates indicate a high metal accumulation capacity of the intertidal marshes. Overbank sedimentation in the tidal marshes of the entire estuary was calculated to remove 25% to 50% of the riverine metal influx, even though marshes comprise less than 8% of the total surface of the estuary. In addition, the large-scale implementation of planned tidal marsh restoration projects was estimated to almost double the trace metal storage capacity of the present natural tidal marshes in the estuary.     

Low tide distribution of wintering waders and shelduck on the Severn Estuary in relation to the proposed tidal barrage

Waders and shelduck were counted at low tide on 162 sectors comprising 85% of the intertidal area (21 467 ha) of the Severn Estuary on 12 occasions during winter 1987/88. On average, 50% of birds present at low tide utilized just 13 sectors (12% of the area); 90% of birds occurred on only 56 sectors, leaving large expanses of intertidal sand virtually devoid of birdlife. Dunlin, the numerically dominant species, occurred widely on the middle and outer estuary, whereas shelduck predominantly occurred on the outer estuary and redshank around many tributary river mouths. Curlew, the most ubiquitous species, was the only one concentrated on the inner estuary. Severe gales in both late December and mid-January concentrated all main species within fewer sectors, probably by the short-term removal of surface sediment from substantial areas. It is estimated that the proposed tidal barrage would eliminate intertidal areas accounting for between c. 40% (for shelduck and curlew) and 80% (for redshank) of current total low tide usage by the internationally important populations present.     

The biology of the twaite shad, Alosa fallax fallax (Lacépède), in the Severn Estuary

The estuarine biology of the twaite shad was studied in the Severn Estuary. Adults enter the estuary at the start of the freshwater phase of their spawning migration between April and June. Peak immigration generally occurs in May and is associated with temperatures in the range 10.6–12.3°C. The mean (± s.d. ) instantaneous mortality rate for the mature population was 0.53±0.18. The effect of additional mortality on the spawning population was modelled assuming constant recruitment and no density-dependent effects.
Juvenile twaite shad are present in the estuary from July until they emigrate seaward during the autumn. A portion of these fish re-enter the estuary the following April–May and remain until late summer/early autumn before once more migrating seaward.
The 0 + age group feed mainly on harpacticoid and calanoid copepods and mysids, the relative preponderance of these in the diet being apparently related to tidal conditions. The possible implications of the proposed tidal power barrage in the Severn Estuary on the twaite shad population are discussed in relation to movement, diet and additional mortality of the mature population.