Tidal activity rhythms and depth distribution of rocky shore fish in an altered intertidal environment

In this note we report on changes of activity level of littoral fish and their use of different microhabitats in an altered intertidal environment, where intertidal fish are never emersed and turbulence is confined to high tides. Despite these atypical conditions, the structure of the assemblage is basically the same found in a typical rocky-shore situation. Two intertidal fish known to possess internal tidal rhythms, Lipophrys pholis and Coryphoblennius galerita, retain those rhythms in these altered conditions, and the associated social changes in L. pholis are also retained. These observations support the studies of Gibson (1971), who showed that changes in hydrostatic pressure were of prime importance in keeping the tidal rhythm entrained. The subtidal Parablennius pilicornis, on the contrary, is more active during low tide than at high tide. The hypothesis that some subtidal species are excluded from the intertidal by a turbulence-avoiding mechanism is discussed.     

The Behaviour of Juvenile Horseshoe Crabs,Tachypleus Tridentatus (Xiphosura), on a Nursery Beach at Shui Hau Wan,Hong Kong

The aim of the present study was to examine some general aspects of feeding in Tachypleus tridentatus juveniles and the possible strategies employed to survive on sand flats experiencing a fluctuating environment, in terms of temperature, salinity and tidal action. Activity of Tachypleus tridentatus juveniles in the field was studied on each visit between September 1997 and June 1998, during low tides when the animals were crawling at the sand surface in the intertidal zone at Shui Hau, Lantau Island, Hong Kong. The patterns of feeding trails produced by the juveniles in the field and their crawling rates were determined and compared. Juveniles began to crawl on the sand surface after the tide had receded and the substratum exposed for about 1 h. The numbers of animals observed increased and peaked after a further 1 h. The number of active individuals gradually declined as most re-burrowed when the tide returned. Juveniles generally crawled in pool areas on the sand surface. A seasonal pattern was also evident in the number of juveniles observed at the study site. Highest numbers were recorded in summer whilst only a few juveniles were seen in winter. Possible strategies adopted by the juveniles for successful survival on the sand flats are discussed.     

Interactive effects of food availability and aerial body temperature on the survival of two intertidal Mytilus species

Intertidal organisms must episodically contend with the rigors of both the terrestrial and the marine environments. While body temperatures during high tide are driven primarily by water temperature, aerial body temperatures are driven by multiple environmental factors such that temperature of an organism during low tide is usually quite different from air temperature. Thus, whereas decades of research have investigated the effects of water temperature on intertidal species, considerably less is known about the physiological impacts of temperature during aerial exposure at low tide, especially with regard to the interaction of aerial body temperature with other stressors. We examined the interactive effects of aerial body temperature and food supply on the survival of two intertidal blue mussels, Mytilus galloprovincialis and Mytilus trossulus. Survival was monitored for nine weeks using a simulated tidal cycle, with two levels of food and three levels of aerial body temperature (30, 25, and 20 °C). Decreased food supply significantly reduced the survival of mussels, but only under the 30 °C treatment. In the other two thermal regimes there were no significant effect of food on survival. When aerial body temperatures are high, food availability may have a greater effect on intertidal organisms. Decreases in ocean productivity have been linked to increased in ocean temperatures, thus intertidal organisms may become more susceptible to thermal stress as climates shift.     

Effects of Tidal and Diel Cycles on Dugong Habitat Use

ABSTRACT Quantifying the factors influencing behaviors of aquatic mammalian grazers may enhance the generic understanding of grazer ecology. We investigated diel and tidal patterns in movements of the dugong (Dugong dugon) by Global Positioning System—tracking 12 animals in 5 inshore—intertidal and 3 offshore—subtidal habitats along the coast of Queensland, Australia. We examined effects of tide height and time of day on the dugong's distance from 1) the nearest coast, 2) water >3 m deep, 3) actual water depth (bathymetry + tide ht) experienced, and 4) distribution of the directions of movements. Both tidal and diel cycles influenced dugong movement. Tracked dugongs tended to be closer to shore at high tide than at low tide and closer to shore at night than during the day. Onshore movement was more prevalent on incoming tides and in the afternoon and evening. Offshore movement was more prevalent on outgoing tides and from midnight through the morning until midday. Tidal and diel variation in water depths used by the inshore—intertidal dugongs was small, but probably underestimated, hidden by a sampling bias in the telemetry equipment. Onshore movement at high tide allowed dugongs to exploit intertidal seagrass beds. Dugongs are closer to shore in afternoons and evenings than in mornings. This behavior may be related to the avoidance of predators or watercraft. Our findings can be used to predict spatial patterns of dugongs within areas of conservation management significance and to assess, avoid, and mitigate adverse effects of anthropogenic disturbance.     

Dynamics of Spatial Pattern for the Gastrotrich Tetranchyroderma bunti in the Surface Sand of High Energy Beaches

This paper is a quantitative study of the spatial pattern of the intertidal gastrotrich. Tetranchyroderma bunti, and its migration pattern during a tidal cycle on three high energy beaches of Bogue Banks, North Carolina, USA. Included in the study were the macrodistribution and micro-distribution. T. bunti is gregarious and forms a clumped spatial pattern in the sand. The four clump sizes found were 2.2, 3.3, 4.4, and 5.5 cm with a ratio of 9: 6: 4: 1, respectively. The spatial pattern of these clumps suggests that the general spatial pattern of the population in its most dense area is aggregated clumped. Field experiments show that this is a stable pattern during at least part of the low tide period, and laboratory experiments suggest that the low water content of the sand at low tide contributes to this stability by decreasing the movement of the gastrotrichs. A significant difference (p<0.05) was found in the density of the gastrotrichs parallel to the shore line, but part of this difference was due to local shifts in the spatial pattern of the population instead of a change in the longshore abundance. T. bunti showed a tidal migration pattern opposite to that of the ground water movement.     

New ecological information onScytalina cerdale (Pisces: Scytalinidae) from a central California rocky intertidal zone

Synopsis New information regarding the ecology ofScytalina cerdale was obtained over a four year period as a consequence of a long-term marine ecological study at the Diablo Canyon Power Plant (DCPP), San Luis Obispo County, California. Twenty intertidal fish surveys were conducted at approximately quarterly intervals, between March 1979 and June 1983, at three separate rocky shore locations (stations). During each survey, a total of 108 square meters (36 m² per station) was searched for fish during periods of low tide. A total of 280S. cerdale were collected, identified, measured, and released back into the same 4 m² area, from the same intertidal station (Diablo Cove), throughout the study period. This limited intertidal occurrence most likely reflectsS. cerdale microhabitat requirements; a combination of intertidal elevation (mean = +0.3 MLLW), substratum specificity (loose gravel, 5–10mm size range, overlying a base of sand and shell fragments), and degree of wave exposure (semi-protected). Throughout the study,S. cerdale was seasonally more abundant during summer months (June through August) and less abundant during winter months (November through February). The only exception to this abundance trend followed the 1982 winter storms, which coincided with an El Niño event, whenS. cerdale abundance uncharacteristically dropped during the subsequent 1983 spring and summer surveys. Mean total lengths did not vary greatly, reflecting the absence of early juvenile fishes, and relatively high mean fish densities (3.5 fish per m² for 20 surveys) were recorded. Qualitative comparisons betweenS. cerdale abundance and seasonal changes in water temperature indicated an inverse relationship. Gravid females, demersal egg masses, and early juvenile individuals were never observed during the four years of the study. This suggests that unlike most other intertidal fishes, particularly other blennioids,S. cerdale may not utilize the intertidal zone for reproductive and recruitment purposes; Diablo Canyon is the most southerly reported distribution for this intertidal species, and therefore, may not entirely represent the species' biology as a whole.     

Larval and juvenile fishes occurring with flood tides on an intertidal mudflat in the Tama River estuary, central Japan

Intertidal movements of fish larvae and juveniles on a mudflat in the Tama River estuary, central Japan, were investigated by comparing the abundance and sizes of fishes caught in the intertidal zone during flood tides with those in the subtidal zone during low tides. A total of 28465 individuals, belonging to 9 families and 20 species, were collected by small purse seine. Among the abundant species, planktonic larvae and juveniles of gobiids and Konosirus punctatus were more abundant in the intertidal zone at flood tide than the subtidal zone at low tide. Similar occurrence patterns were found in juvenile Plecoglossus altivelis altivelis and Lateolabrax japonicus, having fully developed swimming abilities. In contrast to these species, much higher abundances of epibenthic juveniles of 2 gobiids (Acanthogobius flavimanus and Gymnogobius macrognathos) were found in the subtidal zone at low tide, although they also utilized the intertidal zone at flood tide.     

Activity and shelter use of an intertidal snail: effects of sex, reproductive condition and tidal cycle

Although activity patterns are an important component of the ecology of animals as they search for food, mates and appropriate habitats, substantial variation can be observed even within a single population. The source of this variation is the combination of intrinsic and extrinsic factors acting at an individual level. We documented the activity patterns of the ovoviviparous intertidal gastropod Littorina saxatilis (Olivi) over a series of tidal cycles to assess how differences in movements of individuals with different reproductive attributes (non-gravid females, gravid females and males) might explain intra-population variability in activity. Four behaviors were measured for four consecutive tidal cycles for each of 3 months: the magnitude, frequency, and orientation of movements, and the use of shelters (i.e., crevices). In addition, we correlated movements with variations in several environmental factors (temperature, wind, humidity) during low tide to investigate the possible risk sensitivity of the different reproductive groups. All groups were more active and moved greater distances during high tide. Females, especially gravid females, appeared to be more risk-sensitive as they moved shorter distances and used shelters more often. Gravid females were also the only group with a negative relationship between air temperature and movement during low tide. We interpret these behaviors as a means to increase the probability of survival of the brooded offspring carried by the female. Males were the only group to consistently have directionality in their movements and showed evidence of an “up-down” behavior for remaining in a restricted zone. Reproductive attributes appear to be able to explain part of the intra-population variability in foraging activity in natural populations and should be taken into account in the design and interpretation of behavioral and ecological studies.     

Entrainment of the activity rhythm of the mole crab Emerita talpoida

The mole crab Emerita talpoida migrates with the tide in the swash zone of sand beaches. A circatidal rhythm in vertical swimming underlies movement, in which mature male crabs show peak swimming activity 1-2 h after the time of high tides at the collection site. In addition, there is a secondary rhythm in activity amplitude, in which crabs are maximally active following low amplitude high tides and minimally active following high amplitude high tides. The present study determined the phase response relationship for entrainment of the circatidal rhythm with mechanical agitation and whether the cycle in activity related to tidal amplitude could be entrained by a cycle in the duration of mechanical agitation at the times of consecutive high tides. After entrainment with mechanical agitation on an orbital shaker, activity of individual crabs was monitored in constant conditions with a video system and quantified as the number of ascents from the sand each 0.5 h. Mechanical agitation at the times of high tide, mid-ebb and low tide reset the timing of the circatidal rhythm according to the timing relationship to high tide. However, mechanical agitation during flood tide had no entrainment effect. In addition, a cycle in duration of mechanical agitation entrained the rhythm in activity amplitude associated with tidal amplitude. Both rhythms and entrainment effectiveness over the tidal cycle may function to reduce the likelihood of stranding above the swash zone.     

Effect of immersion at low tide on distribution and movement of the mud snail, Ilyanassa obsoleta (Say), in the upper Bay of Fundy, eastern Canada

Habitat heterogeneity often affects movement behaviours of animals, and consequently their spatial distribution. We evaluated the effect of immersion at low tide on the distribution, fine-scale movement patterns and daily movement patterns of the mud snail Ilyanassa obsoleta on a mudflat in the upper Bay of Fundy, Canada. Mud snails migrate onto intertidal mudflats in the summer, and our field survey showed that their density was higher inside tide pools relative to adjacent areas that are exposed at low tide. Using time-lapse videography, we evaluated the effect of snail size, snail density, and immersion at low tide on fine-scale movement patterns of I. obsoleta. Time until snails stopped moving and burrowed was unaffected by snail size, but snails at low and high densities burrowed somewhat faster than those at intermediate densities. Snail size and snail density had no detectable effect on displacement speed or linearity of displacement. Immersion affected snail movement: snails within tide pools delayed burrowing and traveled in more convoluted paths compared to those on exposed mud. Snails increased their turning angles within tide pools, which is probably the mechanism by which aggregations are formed. We also performed a mark-recapture experiment to compare daily movement patterns of snails released inside and outside tide pools. Snails released in tide pools moved shorter distances, but did not orient themselves differently than snails released outside tide pools. Both groups exhibited significant directionality, moving against the mean water current direction over 24 h. In sum, immersion at low tide affected the behaviour and spatial distribution of snails, resulting in snail aggregations within tide pools. These snail aggregations, in turn, may be a major factor influencing spatial dynamics on mudflats, including causing changes in distribution patterns of the burrowing amphipod Corophium volutator, a dominant inhabitant and key species in the food web of mudflats.     

Behavioural studies on the lesser sandeel Ammodytes marinus (Raitt) III. The effect of temperature on activity and the environmental control of the annual cycle of activity

The behaviour of the lesser sandeel, Ammodytes marinus (Raitt), has been investigated at 5, 10 and 15° C, using a photographic method of recording activity. The activity patterns at 10 and 15° C were very similar, there being a high level of swimming activity during the light period, which fell to a low level at 5° C. It was also lower at 10° C at the end of the experiment than at the beginning and it is suggested that this might have been due to an increase in the fat contents of the fish. The feeding rate of the fish was measured and showed a Q ₁₀ of 2.08 for the temperature range 5–15° C. The annual cycle of activity of A. marinus is discussed in relation to seasonal changes in food availability, light and temperature, and in the fat content of the fish. It is concluded that after spawning in the December–January period the fish remain buried in the sand until April, because of the limiting effect on swimming and feeding activity of the environmental factors in the intervening period. The proportion of fish available for capture at the start of the fishery in April is related mainly to temperature, but food (as measured by numbers of copepods) light intensity and photoperiod are by then increasing rapidly. After July the fishery ceases and it is thought that this is because the fish have entered an overwintering stage, during which they remain buried in the sand. This phase is also thought to be associated with the maturation of the gonads in readiness for the winter spawning. The factors causing the fish to enter this stage are as yet undetermined but may be related to the attainment of a certain level of fat content.     

A quantitative description of the behaviour of wild juvenile plaice (Pleuronectes platessa L.)

The behaviour of young plaice (Pleuronectes platessa L.) as they migrate up and down sandy beaches with the tide is described. Their behaviour during this migration consists mainly of swimming and feeding interspersed with rarer behavioural acts. Over short periods swimming behaviour can be described by a random model in which the probability of a swim occurring remains constant. This probability varies markedly, however, from hour to hour. Two types of swimming movement are recognized: one of very short duration represents searching for food and the other, longer, category serves to transport the fish up and down the shore. Variations in the feeding rate and in the frequency, duration, and direction of swimming movements over the tidal cycle are described and related to the changing physical and biological conditions that the fish experience during their intertidal movements.     

The influence of habitat, season and tidal regime in the activity of the intertidal crab Neohelice (=Chasmagnathus) granulata

The activity pattern of intertidal crabs is influenced by factors that usually change rhythmically following tidal and/or diel cycles, and is often associated with the use of refuges. The movement activity of the burrowing crab Neohelice granulata was compared among three populations from SW Atlantic coastal areas where they face different tidal regimes, water salinities, substrata and biological factors. At each site, we examined the seasonal activity of the crabs (individuals collected in pitfall traps) in two types of habitat: mudflat and salt marsh. The working hypothesis is that the activity would vary according to the diverse environmental conditions encountered at geographical and local scales. Crab activity varied between sites and seasons showing to be more intense when habitats were covered by water. The most active groups were large males, followed by large non-ovigerous females. Ovigerous females were almost inactive. Most crabs were near or inside burrows at low tides in Mar Chiquita and Bahía Blanca, but they were active at both low and high tides in San Antonio during spring and summer. N. granulata were active in a wide range of temperatures: from 10 to 37 °C at low tides and at temperatures as low as 2 °C when covered by water. Differences of activity between mudflat and salt marsh varied among sites depending on flooding frequencies. Movement activity of N. granulata varied both in space and in time; crabs move under very different abiotic conditions (e.g., low or high tide, daylight or night, low and high temperature) and their movement may also be prevented or elicited by biotic conditions like burrow complexity, food quality and predation pressure. The wide set of conditions under which N. granulata can be active may explain why this is the only semiterrestrial crab inhabiting latitudes higher than 40°S in South America.     

The ecology,behaviour and physiology of fishes on coral reef flats,and the potential impacts of climate change

Reef flats, typically a low‐relief carbonate and sand habitat in shallow water leeward of the reef crest, are one of the most extensive zones on Pacific coral reefs. This shallow zone often supports an abundant and diverse fish assemblage that is exposed to more significant variations in physical factors, such as water depth and movement, temperature and ultraviolet (UV) radiation levels, than most other reef fishes. This review examines the characteristics of reef flat fish assemblages, and then investigates what is known about how they respond to their biophysical environment. Because of the challenges of living in shallow, wave‐exposed water, reef flats typically support a distinct fish assemblage compared to other reef habitats. This assemblage clearly changes across tidal cycles as some larger species migrate to deeper water at low tide and other species modify their behaviour, but quantitative data are generally lacking. At least some reef flat fish species are well‐adapted to high temperatures, low oxygen concentrations and high levels of UV radiation. These behavioural and physiological adaptations suggest that there may be differences in the demographic processes between reef flat assemblages and those in deeper water. Indeed, there is some evidence that reef flats may act as nurseries for some species, but more research is required. Further studies are also required to predict the effects of climate change, which is likely to have multifaceted impacts on reef flats by increasing temperature, water motion and sediment load. Sea‐level rise may also affect reef flat fish assemblages and food webs by increasing the amount of time that larger species are able to forage in this zone. The lack of data on reef flats is surprising given their size and relative ease of access, and a better understanding of their functional role within tropical marine seascapes is urgently required.     

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 Collembolan,oudemansia Esakii,an Intertidal Marine Insect in Hong Kong,with Emphasis on the Floatation Method of Collection

Summary Oudemansia esakii is shown to be a truly marine insect as all stages of the life cycle are found in intertidal sand in Hong Kong. The motile stages migrate vertically and forage during low tide, but burrow into the sand with incoming tidal conditions. An air bubble often forms around them while in the soil. Total forced immersion for 75 hours showed no harmful effects. The greatest burrowing depth is 36 cm and the optimal littoral position is three feet above low water. A floatation method using graded sieves is superior to MacFadyen's high gradient technique. Pertinent hydrographic data are included, as are biological relationships.Marine Science Laboratory, the Chinese University of Hong Kong     

Tidal movements of the macrofauna on an exposed sandy beach in South Africa

All epifaunal and shallow-burrowing, intertidal macrofauna on an exposed sandy beach appear to undergo tidal migrations which vary in extent but are all upshore on the incoming tide and downshore on the outgoing tide during day and night. In the crustaceans this is probably endogenous rhythmic behaviour and they show increased activity at night. In the molluscs this movement is not synchronised by an intrinsic mechanism but results from behavioural responses to changing physical conditions. The large wedge clam, Donax serra, shows no tidal migrations but does show a semilunar rhythm of movement from just above mean tide level (MTL) during springs to the low tide level during neap tides.     

The feeding strategy of the dunlin (Calidris alpina L.) in artificial and non-artificial habitats at Ria de Aveiro,Portugal

Usually waders (Aves, Charadrii) feed in the inter-tidal zone over the low tide period and roost over high tide. But occasionally when their energetic needs increase, due to adverse weather conditions or to a need to store fat to fuel migration, the birds also feed during high tide, sometimes in agricultural land. At the Ria de Aveiro, about 44% (corresponding to 4500 birds) of the wintering dunlin (Calidris alpina) use artificial salt pan habitats throughout the winter whilst the others utilize natural intertidal habitats, mainly mudflats. The dunlin that feed intertidally in natural areas, such as the Canal de Mira, do not feed over high tide. In contrast, the birds that use the salt pan area feed both during low tide, in abandoned salt pans whose walls are breached and thus allow access to the tide, and high tide, in active salt pans, and keep on doing so throughout the winter. The energetic needs of both groups of birds are thought to be similar. However, the benthos of the abandoned salt pans seem to be different, and poorer in energetic terms, from those of the natural mudflats. It is argued that dunlin wintering in the salt pan area must compensate for this by feeding over high tide in active salt pans. Rather than just being a means of obtaining more energy at certain times of high energy demand, it would seem that feeding in active salt pans over high tide should be considered as a normal component of the feeding strategy of these birds, a strategy that is different from those wintering in the natural mudflats. These findings highlight the importance of the salt exploitation industry to waders and also show that artificial habitats can support large numbers of wintering waders.     

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.     

Local distribution and thermal ecology of two intertidal fishes

Geographic variability in the physiological attributes of widely distributed species can be a result of phenotypic plasticity or can reflect evolutionary responses to a particular habitat. In the field, we assessed thermal variability in low and high intertidal pools and the distribution of resident fish species Scartichthys viridis and transitory Girella laevifrons along this vertical intertidal gradient at three localities along the Chilean coast: Antofagasta (the northernmost and warmest habitat), Carrizal Bajo (central coast) and Las Cruces (the southernmost and coldest habitat). In the laboratory, we evaluated the thermal sensitivity of fish captured from each locality. The response to temperature was estimated as the frequency of opercular movements and as thermal selectivity in a gradient; the former being a indirect indicator of energy costs in a particular environment and the latter revealing differential occupation of habitat. Seawater temperature in intertidal pools was greatest at Antofagasta, and within each site was greatest in high intertidal pools. The two intertidal fish species showed opposite patterns of local distribution, with S. viridis primarily inhabiting the lower sectors of the intertidal zone, and G. laevifrons occupying the higher sectors of the intertidal zone. This pattern was consistent for all three localities. Locality was found to be a very important factor determining the frequency of opercular movement and thermal selectivity of both S. viridis and G. laevifrons. Our results suggest that S. viridis and G. laevifrons respond according to: (1) the thermal history of the habitat from which they came, and (2) the immediate physical conditions of their habitat. These results suggest local adaptation rather than plasticity in thermoregulatory and energetic mechanisms.