Recolonization pattern of the polychaete Lanice conchilega on an intertidal sand flat following the severe winter of 1995/96

The recolonization process of Lanice conchilega was studied on an intertidal sand flat in the northern Wadden Sea after an intertidal population had been wiped out during the severe winter of 1995/96. After very low recruitment in the first two years, the population fully recovered in the third year after its destruction. By then, the distribution pattern closely resembled the pattern before the severe winter with >95% of all Lanice occurring below –0.5 m mean tidal level and dominance of the density categories 6–50 individuals (ind) m^–2 and 51–200 ind m^–2. Lanice larvae have been observed to attach preferentially on the tubes of adults. However, the high recruitment in the third year was also seen in areas devoid of adult Lanice, indicating that settlement facilitation by adults was of little importance for the recolonization process. Instead, we suggest that the observed recruitment pattern was a consequence of a large-scale decline in the metapopulation in the tidal basin and the onshore coastal subtidal which resulted in insufficient larval supply onto the tidal flats and low recruitment in the first two years following the severe winter. Received in revised form: 7 May 2001 Electronic Publication     

Recruitment of shore crabsCarcinus maenas on tidal flats: Mussel clumps as an important refuge for juveniles

During the late summer and early fall, juvenile shore crabs (Carcinus maenas L.) occurred in high abundances in mussel clumps scattered on tidal flats of the Wadden Sea. Abundances were much lower on bare tidal flats without mussel clumps and decreased substantially from July to November, whereas numbers in mussel clumps remained high. Large crabs left the tidal flats in early fall, whereas juveniles undertook tidal migrations only in the late fall. In March very few shore crabs were found in the intertidal area. The size of juvenile shore crabs living between mussels did not increase significantly during fall. On the bare tidal flats surrounding the mussels, a size increase was observed. Mussel beds and mussel clumps serve as a spatial refuge for the early benthic phases of juvenile shore crabs. Between mussels they can hide effectively from their epibenthic predators. Juvenile shore crabs do not leave the intertidal area and the mussel habitats before their major predators have left the area. Mussel clumps scattered over the tidal flats may be a critical refuge for juvenile shore crabs settling on tidal flats. Intensified efforts in mussel culturing in the European Wadden Sea during recent decades may have caused an increased abundance of mussel clumps on tidal flats, thus enhancing habitat availability for some mussel-clump inhabitants.     

Differential effects of the severe winter of 1995/96 on the intertidal bivalves Mytilus edulis, Cerastoderma edule and Mya arenaria in the Northern Wadden Sea

Intertidal mussel beds were severely damaged by scouring ice floes during the winter of 1995/96. Aerial surveys before and after the winter showed that more clusters of mussel beds vanished in a region with a higher areal share of tidal flats and a lower salinity, suggesting that the amount of ice present determined the magnitude of the disturbance on beds of Mytilus edulis. Nehls and Thiel [(1993) Neth J Sea Res 31:181–187] observed a strikingly similar spatial pattern of disturbances caused by severe storms in the Wadden Sea. Areas on mussel beds mechanically undisturbed by ice showed no reduced abundance and biomass of mussels, indicating that temperature alone was of little importance as a lethal factor. Conversely, Cerastoderma edule was strongly affected by low temperature. On average 80% died during the winter with extinctions up to 100% in the high tidal zone. At the lowest tidal level, surviving cockles were larger than those killed by the frost. A reinvestigation of sampled sites in autumn revealed that substantial further mortality had occurred during spring and summer which may constitute a time-lag effect of the preceding winter. There was no increased mortality in juvenile and adult Mya arenaria during the winter of 1995/96, confirming that this clam is a hard-winter species like Macoma balthica. Received in revised form: 7 May 2001 Electronic Publication     

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.     

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.     

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.     

Intertidal exposure zones: A way to subdivide the shore

A new way of subdividing the shore is described and Doty's concept of critical tide levels (CTLs), on which the scheme is based, is revised. The various cycles of the tide (e.g., daily, monthly, and annual cycles) define several different orders of CTL at which the duration of continuous exposure or submergence increases abruptly by a discrete interval, the magnitude of which is determined by cycle period. Contrary to popular belief, most CTLs are common to all tidal types (i.e., mixed, semi-diurnal, and diurnal) and thus they can be used as reference levels for cross-comparing different intertidal regions, regardless of tidal type. CTLs naturally subdivide the intertidal region into discrete exposure zones, here called the atmozone, amphizone, and aquazone, which have upper and lower subzones. The amphizone, the intertidal core, experiences exposure and submergence extremes in terms of hours (? 1 lunar day). The overlying lower atmozone and underlying upper aquazone experience exposure and submergence extremes, respectively, of the order of days (?2 to ≈10 days), while in the upper atmozone and lower aquazone extremes are measured in months or years (> ≈20 days).In most intertidal environments (tidal flats, saltmarshes, brackish marshes, and sheltered rocky shores) CTLs (including exposure zone boundaries) probably limit some biological zones. For tidal flats experiencing large tidal ranges it is possible that biological zones in the fringes of the intertidal region shift in response to long-term fluctuations in the levels of CTLs, due to the effects of the 18.6 year soli-lunar cycle.     

Population dynamics of the ribbed mussel,Geukensia demissa: The costs and benefits of an aggregated distribution

Summary Although mussel beds are common in many intertidal habitats, the ecological significance of the aggregated distribution of mussels has not been examined. The ribbed mussel, Geukensia demissa, is found in dense aggregations on the seaward margin of many salt marshes in New England. Here, we examine the population structure of G. demissa in a New England salt marsh and investigate experimentally the costs and benefits of aggregation.Size, growth rate, and settlement rates of mussels decrease with increasing tidal height, whereas survivorship and longevity increase with increasing tidal height. Winter ice dislodges mussels from the substratum, resulting in mortality over all size classes, whereas crab predation results in the mortality of smaller mussels. The intensity of each of these mortality agents decreases with increasing tidal height. Effects of intraspecific competition on individual growth and mortality also decrease with increasing tidal height.At high densities, individual growth rates were reduced, with depression of growth rates most pronounced on smaller individuals. Mortality from sources other than intraspecific crowding, however, was reduced at high mussel densities, including mortality due to winter ice and crab predators. As a result, our data suggest that the mussel population at our study site would be reduced by 90% in only five years and no juveniles would survive through their second year without an aggregated distribution.Juveniles settle gregariously with or without adults present. The aggregated distribution of settlers and the postsettlement movement of smaller mussels to favorable microhabitats result in size and age class segregation within the population. This probably reduces intraspecific competition for food, while maintaining the survivorship advantages of an aggregated distribution.     

Contribution of unvegetated tidal flats to coastal carbon flux

Unvegetated flats occupy a large area in the intertidal zone. However, compared to vegetated areas, the carbon sequestration of unvegetated tidal flats is rarely quantified, even though these areas are highly threatened by human development and climate change. We determined benthic maximum gross primary production (GPP_m), net primary production (NPP) and total respiration (TR) during emersion on seven tidal flats along a latitudinal gradient (from 22.48°N to 40.60°N) in winter and summer from 2012 to 2016 to assess the spatial and temporal variability of carbon dioxide flux. In winter, these processes decreased by 89%–104% towards higher latitudes. In summer, however, no clear trend was detected across the latitudinal gradient. Quadratic relationships between GPP_m, NPP and TR and sediment temperature can be described along the latitudinal gradient. These curves showed maximum values of GPP_m and NPP when the sediment temperatures reached 28.7 and 26.6°C respectively. TR increased almost linearly from 0 to 45°C. The maximum daily NPP across the latitudinal gradient averaged 0.24 ± 0.28 g C m^?2 day^?1, which was only 10%–20% of the global average of NPP of vegetated coastal habitats. Multiplying with the global area of unvegetated tidal flats, our results suggest that the contribution of NPP on unvegetated tidal flats to the coastal carbon cycle is small (11.04 ± 13.32 Tg C/year). If the land cover of vegetated habitats is continuously degraded to unvegetated tidal flats, the carbon sequestration capacity in the intertidal zone is expected to reduce by at least 13.10 Tg C/year, equivalent to 1% of global carbon emissions from land‐use change.     

基于胁迫梯度假说和互惠理论的海三棱藨草种群恢复技术

滨海湿地生态修复已成为阻止海岸带生态系统退化、保护生物多样性以及提供生态服务的关键措施。以长江口原生盐沼植物海三棱藨草(Scirpus mariqueter)为研究对象,选取崇明东滩新生滩涂湿地为研究区域,通过沿潮滩高程梯度的海三棱藨草植株斑块的移植实验,探究胁迫梯度假说和互惠理论(即种内的正相互作用)对长江口海三棱藨草种群恢复的指导意义。研究结果显示:(1)在一定的胁迫梯度范围内(潮滩高程2.0 m以上),增大种植斑块可以促进海三棱藨草的种内正相互作用,显著提高种植斑块的存活率和植株密度(P0.05);(2)潮滩水文动力沉积条件与潮滩高程梯度密切相关(P0.05),水文动力沉积作用对海三棱藨草定居和生长的胁迫随高程梯度下降而增强。潮滩高程2.0 m以下处强烈的水文动力条件干扰限制了生物-物理因素的正反馈作用。滨海湿地盐沼植被修复工作的成功率可以通过改进种植方式,增强种内的正相互作用得到极大的提高。研究可为开展大规模滨海湿地盐沼植被修复工程和提高生态修复效率提供科学依据和技术支持。     

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.     

Predation on 0-group and older year classes of the bivalve Macoma balthica: interaction of size selection and intertidal distribution of epibenthic predators

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea.Shrimps Crangon crangon, adult crabs Carcinus maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinus stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+group Macoma.Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation.     

Tower construction by the manicure crab <Emphasis Type="Italic">Cleistostoma dilatatum</Emphasis> during dry periods on an intertidal mudflat

Animals living on upper intertidal mudflats experience habitat desiccation during neap tides when water does not flood the habitat. Individuals of the manicure crab Cleistostoma dilatatum construct cone-shaped towers at the entrance of their burrows, in which they remain during neap tides. These towers are the tallest known structures compared to body size built by crabs living on intertidal flats. The frequency of tower construction followed semilunar tidal cycles with most building done prior to neap tides when few crabs were active on the mudflat surface. Bigger crabs tended to make taller and wider towers with a wider pinhole on the top. These towers may regulate the microclimate in burrows.     

Middle Triassic horseshoe crab reproduction areas on intertidal flats of Europe with evidence of predation by archosaurs

A systematically excavated track site in a 243.5 Myr old Middle Triassic (Karlstadt Formation, Pelsonian, middle Anisian) intertidal carbonate mud‐flat palaeoenvironment at Bernburg (Saxony‐Anhalt, central Germany) has revealed extensive horseshoe crab trackways attributable to the Kouphichnium Nopsca, 1923 ichnogenus. The exposed track bed of a Germanic Basin‐wide spanned intertidal megatrack site is a mud‐cracked biolaminate surface on which detailed tracks have been preserved because of rapid drying and cementation as a result of high temperatures, followed by rapid covering with a protective layer of arenitic storm or tsunami sediments. The different trackway types and their orientations have allowed a tidal sequence to be reconstructed, with the initial appearance of swimming horseshoe crabs followed by half‐swimming/half‐hopping limulids under the shallowest water conditions. The Bernburg trackways, which have mapped lengths of up to 40 m, were all produced by adult animals and exhibit a variety of shapes and patterns that reflect a range of subaquatic locomotion behaviour more typical of mating than of feeding activities. The closest match to the proportions and dimensions of the horseshoe crab tracks at Bernburg is provided by the largest known Middle Triassic limulid Tachypleus gadeai, which is known from the north‐western Tethys in Spain. The horseshoe crab body fossils recognized in the German Mesozoic intertidal zones, instead, are from juveniles. The uniformly adult size indicated by the trackways therefore suggests that they may record the oldest intertidal reproductive zones of horseshoe crabs known from anywhere in the world, with the track‐makers having possibly migrated thousands of kilometres from shallow marine areas of the north‐western Tethys to reproduce in the intertidal palaeoenvironments of the Germanic Basin. Chirotherium trackways of large thecodont archosaurs also appeared on these flats where they appear to have fed on the limulids. With the tidal ebb, smaller reptiles such as Macrocnemus (Rhynchosauroides trackways) appeared on the dry intertidal flats, probably feeding on marine organisms and possibly also on horseshoe crab eggs. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 76–105.     

Juvenile Chinese mitten crabs (<Emphasis Type="Italic">Eriocheir sinensis</Emphasis>) in the Thames estuary: distribution,movement and possible interactions with the native crab <Emphasis Type="Italic">Carcinus maenas</Emphasis>

The Chinese mitten crab, Eriocheir sinensis is a successful invader. Whilst non-breeding adult E. sinensis have been associated with the destruction of riverbanks, little is known about the ecology of this species in its invaded areas. This is especially true of the juveniles which are a key migratory stage. Intertidal surveys along the Thames estuary indicated an increase in the population since the 1990s. Juvenile E. sinensis were abundant in the sampled upper tidal region of the Thames, refuging under boulders in the intertidal at low tide. Seasonal differences in sampled populations were observed, with a significantly lower abundance of crabs found during winter compared to summer. Mark-recaptures indicated movement in the intertidal occurring during high tide, with an influx of new crabs evident after a single tidal cycle. Endogenous rhythms were also apparent, with peaks in activity occurring corresponding with night-time high tide conditions. E. sinensis juveniles can successfully exclude similar sized native Carcinus maenas from shelters in the laboratory, regardless of which species originally inhabited the shelter. This may have implications for native estuarine Carcinus populations, which can rely on such intertidal shelters for refuges.     

Spatial and temporal patterns of platyhelminth assemblages in intertidal sediments of northeast Australia

The distribution of Platyhelminthes was surveyed along two transects in tidal flats of the northeast coast of Queensland, Australia, in May 1990 and September 1991. Species numbers and densities as well as individual abundances and diversity increased from the high towards the low intertidal in relation to sediment composition. Along this gradient, platyhelminth assemblages of low intertidal sandflats, muddy sandflats and mudflats were distinguished with multivariate analyses. The distribution pattern was affected by the sediment parameter, and also by interactions with macrofauna. Sampling and analysis of the platyhelminth assemblage in tidal flats of Hinchinbrook Channel over time (November 1988 to October 1991) revealed annual and interannual variation, but no seasonality. This revised version was published online in July 2006 with corrections to the Cover Date.     

Changes in the waterbird populations of the Oosterschelde (SW Netherlands) as a result of large-scale coastal engineering works

Between 1982 and 1987, the construction of a storm-surge barrier and two secondary dams in the eastern and northern parts of the Oosterschelde/Krammer-Volkerak area resulted in the loss of 33% of the 170 km² of intertidal area in the estuary. Consequences for non-breeding waterbirds were evaluated on the basis of monthly high-tide counts during five seasons before and three seasons after the construction period.In the entire Oosterschelde/Krammer-Volkerak area, numbers of wintering waders decreased but those of ducks increased. Peak numbers and total number of bird-days changed little, but the seasonal pattern shifted from a midwinter maximum to a peak in autumn.In the Oosterschelde (excluding the Krammer-Volkerak), where 17% of the tidal flats disappeared, species feeding mainly on open water remained stable or increased. Species dependent on intertidal areas for foraging (mainly waders and dabbling ducks) generally decreased. Total density of intertidal foragers decreased slightly. In most intertidal species, the Oosterschelde wintering population showed a stronger decrease, or smaller increase, than was shown during the same period by numbers in Britain and Ireland which were taken as an index of the total W-European winter populations. Changes varied considerably between species, and were correlated with their distribution within the estuary. Species concentrated in the eastern sector, where most habitat loss occurred, declined more than species with a more westerly distribution.Results indicate that intertidal foragers forced to move from the enclosed parts of the estuary were not generally able to settle into the remaining intertidal areas. Both dispersal to adjacent areas (mainly by dabbling ducks) and mortality during severe winter weather (in some wader species) may have contributed to the declines. Populations of intertidal foragers apparently were (and consequently still are) close to carrying capacity, and further changes in capacity, as foreseen from geomorphological changes still under way in the estuary, are likely to be reflected in bird populations.Numbers of waders moulting in the Oosterschelde in late summer declined strongly compared to numbers in other seasons. Increased disturbance due to recreational activities may have played a role during this time of the year.     

Historic mussel beds (Mytilus edulis) in the sedimentary record of a back-barrier tidal flat near Spiekeroog Island, southern North Sea

Mussel beds show irregular cycles of appearance, disappearance and reappearance at preferred distinct locations on intertidal flats. These cycles are documented in the depth profile by the presence of shell-rich sediments intercalated with sandy layers. Such mussel bed layers were regularly found over the past 12 years on the Swinnplate, a back-barrier tidal flat south of Spiekeroog Island, southern North Sea. They are characterised by the occurrence of sub-articulated pairs of Mytilus shells. Based on life span considerations, a period of at least 35–40 years over which mussel beds may have repeatedly established themselves is suggested. A survey in spring 2000 revealed a reduced occurrence of old, embedded beds on the Swinnplate. The present results, based on core profiles from 1992 to 2000 and amino acid age determination, show that alternations of shell layers indicative of former Mytilus beds and sediment layers provide insight into the historic development of tidal flat environments. The frequent occurrence of sub-articulated valve pairs in the shell layers documents that the embedding of the mussel beds took place soon after the death of the organisms without prior physical disturbance. Successions of historic Mytilus beds provide evidence that mussel beds are a regular, but not necessarily permanent, faunal feature of back-barrier tidal flats of the North Sea. Electronic Publication     

Reproduction ofAmphiporus lactifloreus (Hoplonemertini) on tidal flats: implications for studies on the population biology of nemertines

The reproductive period and life-history parameters were investigated for the hoplonemertineAmphiporus lactifloreus found on the tidal flats of the island of Sylt in the northern Wadden Sea. Every six weeks 20 individuals were collected and then histologically examined to determine the development stage of their reproductive organs.A. lactifloreus reproduces in the late autumn, its peak reproduction being in the second half of November. Individuals from all size classes >20 mm body length produced gametes. Individuals of the cohorts that reproduced in the late autumn of 1992 persisted and grew until July 1993, indicating thatA. lactifloreus is an iteroparous species. The length of relaxed individuals was significantly correlated with their length under anaesthetized conditions, but the regression changed significantly after the reproductive period. Length under ‘relaxed’ conditions was significantly correlated with weight (wet weight, dry weight, and ash-free dry weight); these relationships did not vary significantly before or after the reproductive period. Our results show that important life-history data of intertidal nemertines can be obtained without time-consuming histological studies. On the basis of these findings, recommendations for future studies on the population biology of intertidal nemertines are given. Regular length measurements of nemertines under ‘relaxed’ conditions are proposed as a useful tool for tracking the growth and survival of annual cohorts of intertidal nemertines.     

Seasonal migrations of the common Goby,Pomatoschistusmicrops (Kroyer), in Morecambe Bay and elsewhere

Summary Seasonal changes in abundance, and also reproduction, of the gobiid Pomatoschistus microps in intertidal pools on mussel beds at Morecambe, Lancashire, were studied from 1956 to 1961. During winter the shore was largely vacated by this species which returned for breeding purposes in spring. Spawning took place in upper shore pools from April to August. After a planktonic life of between 6–9 weeks, young fishes were recruited to the demersal population usually from June to September. A correlation between seasonal migrations and sea temperature is postulated. Geographical variation in occurrence and extent of migration in P. microps and the related sublittoral P. minutus is considered in relation to minimum winter sea temperatures over the southern Boreal and Baltic sea areas. Offshore winter migration has been found to take place in areas where such temperatures normally fall below 5° C, but not where minimum sea temperature is above 7° C. The effect of the severe winter of 1962–63 on certain populations of P. microps is described.