83 Comparative analysis of vertically migrating euglena viridis populations in tidal and non‐tidal benthic environments

Benthic populations of Euglena viridis exhibit vertical migration behavior on high energy intertidal beaches and along the sand banks of freshwater streams. This study examines similarities and differences in the migratory behavior and cell morphology of populations of E. viridis inhabiting Scripps Beach, La Jolla, California and Coble Brook, Burlington, North Carolina. The timing of migration was measured by counting the number of cells in samples collected from the sediment surface throughout the day. Sediment cores were extracted and sectioned to determine the vertical distribution of the population. Neutral density filters and opaque canisters were used to shade the substratum to 56%, 22%, 2%, and 0% of incident irradiance (Io) to examine the effect of light on cell morphology and migratory behavior. On intertidal beaches, E. viridis exhibited a tidal rhythm in vertical migration with cells migrating below the sediment surface at night (>15 cm) and during daytime high tides. In this habitat, the upward migration response was enhanced at irradiances lower than 100% Io but cell morphology was not altered by shading. On the banks of freshwater streams, E. viridis exhibited a diurnal migratory rhythm with both tear‐drop and spherical morphologies observed throughout the day. The population was most concentrated at the surface around solar noon and at night it was located between 1 and 2 cm below the surface. Shading did not enhance upward migration but it did affect cell morphology. These results will be interpreted in the context of the dominant selection pressures in each environment.     

84 Linking environmental forcing, kelp forest habitat dynamics, and community structure in the northeast pacific

Benthic populations of Euglena viridis exhibit vertical migration behavior on high energy intertidal beaches and along the sand banks of freshwater streams. This study examines similarities and differences in the migratory behavior and cell morphology of populations of E. viridis inhabiting Scripps Beach, La Jolla, California and Coble Brook, Burlington, North Carolina. The timing of migration was measured by counting the number of cells in samples collected from the sediment surface throughout the day. Sediment cores were extracted and sectioned to determine the vertical distribution of the population. Neutral density filters and opaque canisters were used to shade the substratum to 56%, 22%, 2%, and 0% of incident irradiance (Io) to examine the effect of light on cell morphology and migratory behavior. On intertidal beaches, E. viridis exhibited a tidal rhythm in vertical migration with cells migrating below the sediment surface at night (>15 cm) and during daytime high tides. In this habitat, the upward migration response was enhanced at irradiances lower than 100% Io but cell morphology was not altered by shading. On the banks of freshwater streams, E. viridis exhibited a diurnal migratory rhythm with both tear-drop and spherical morphologies observed throughout the day. The population was most concentrated at the surface around solar noon and at night it was located between 1 and 2 cm below the surface. Shading did not enhance upward migration but it did affect cell morphology. These results will be interpreted in the context of the dominant selection pressures in each environment.     

EFFECT OF SUBSURFACE NUTRIENT SUPPLIES ON THE VERTICAL MIGRATION OF EUGLENA PROXIMA (EUGLENOPHYTA)1

Euglena proxima Dangeard inhabits intertidal sand flats and displays a tidal rhythm in vertical migration. During daytime low tides when the sand flat is emersed, millions of cells are visible on the sediment surface, but the population remains below the surface at all other times. An earlier study demonstrated that the extent of downward migration of E. proxima is reinforced by the presence of a subsurface layer of black sediment. The present study was designed to test the hypothesis that the higher availability of inorganic nutrients or organic substrates in or above the black layer is responsible for the enhancement of downward migration in E. proxima. This hypothesis was tested experimentally by manipulating the bottom water in 24 mesocosm containers in a tidal tank. Six replicates of each of the following nutrient treatments were tested: seawater control; deep porewater collected from 70 cm below the sediment surface; seawater enriched with ammonium, nitrate, and phosphate; and seawater enriched with acetate, glucose, and the preceding inorganic nutrients. Multivariate analysis of variance revealed that the chl a biomass and chl a‐weighted mean depth of the population at high tide were significantly greater for replicates receiving inorganic nutrients. There was no difference between those receiving only inorganic nutrients and those enriched with inorganic nutrients, acetate, and glucose. These findings represent the first experimental evidence that subsurface nutrients are an important resource that reinforces the maintenance of vertical migration behavior in benthic microalgae.     

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.     

INVERTEBRATE‐MEDIATED NUTRIENT LOADING INCREASES GROWTH OF AN INTERTIDAL MACROALGA1

Even in nitrogen‐replete ecosystems, microhabitats exist where local‐scale nutrient limitation occurs. For example, coastal waters of the northeastern Pacific Ocean are characterized by high nitrate concentrations associated with upwelling. However, macroalgae living in high‐zone tide pools on adjacent rocky shores are isolated from this upwelled nitrate for extended periods of time, leading to nutrient limitation. When high‐intertidal pools are isolated during low tide, invertebrate‐excreted ammonium accumulates, providing a potential nitrogen source for macroalgae. I quantified the influence of mussels (Mytilus californianus Conrad) on ammonium accumulation rates in tide pools. I then evaluated the effects of ammonium loading by mussels on nitrogen assimilation and growth rates of Odonthalia floccosa (Esp.) Falkenb., a common red algal inhabitant of pools on northeastern Pacific rocky shores. Odonthalia was grown in artificial tide pool mesocosms in the presence and absence of mussels. Mesocosms were subjected to a simulated tidal cycle mimicking emersion and immersion patterns of high‐intertidal pools on the central Oregon coast. In the presence of mussels, ammonium accumulated more quickly in the mesocosms, resulting in increased rates of nitrogen assimilation into algal tissues. These increased nitrogen assimilation rates were primarily associated with higher growth rates. In mesocosms containing mussels, Odonthalia individuals added 41% more biomass than in mesocosms without mussels. This direct positive effect of mussels on macroalgal biomass represents an often overlooked interaction between macroalgae and invertebrates. In nutrient‐limited microhabitats, such as high‐intertidal pools, invertebrate‐excreted ammonium is likely an important local‐scale contributor to macroalgal productivity.     

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.     

HABITAT DIFFERENCES IN THE TIMING OF REPRODUCTION OF THE INVASIVE ALGA SARGASSUM MUTICUM (PHAEOPHYTA,SARGASSACEAE) OVER TIDAL AND LUNAR CYCLES1

Sargassum muticum (Yendo) Fensholt is an invasive species that is firmly established on intertidal and subtidal rocky shores of Europe and the Pacific coast of North America. Local success and spread of S. muticum is thought to rely on its reproductive potential that seems dependent on exogenous factors like tidal and lunar cycles. This study is the first to compare the reproductive patterns (periodicity of egg expulsion and embryo settlement) of this invader in two different habitats: the middle and low intertidal. The combination of monthly, daily, and tidal samples at triplicate sites within each habitat showed a semilunar periodicity of egg expulsion and embryo settlement coincident with increasing tidal amplitude just before full and new moons. In both habitats, duration of each egg expulsion event was ～1 week, and embryo settlement occurred during the first daily low tide and with the incoming high tide during spring tides. However, both expulsion and settlement started 1–2 d earlier, expulsion saturation was faster, and settlement was higher in the mid‐ compared to the low intertidal. Our results suggest that the exact timing of gamete expulsion and embryo release of S. muticum responds to local factors, including tidal cues, which result in differences between mid‐ and low‐intertidal habitats.     

EFFECT OF LIGHT ON VERTICAL MIGRATION AND PHOTOSYNTHESIS OF EUGLENA PROXIMA (EUGLENOPHYTA)

During daytime low tides, sediment-inhabiting populations of Euglena proxima Dangeard migrate to the surface of intertidal sand flats to form large monospecific green patches near the Duke University Marine Laboratory in Beaufort, North Carolina. The effect of incident irradiance on the upward migration of E. proxima was studied by shading the sediment surface in the field with neutral density filters to 2, 20, 55, and 100% incident irradiance. The results of these in situ experiments revealed that E. proxima 's upward migration, as measured by population density or chl a concentration, was stimulated by decreasing incident irradiance: population densities and chl a concentrations on the sediment surface at 2% incident irradiance were greater statistically than at 100% full sunlight. In situ measurements of photosynthesis were performed by incubating 3-mL aliquots of a cell suspension in scintillation vials containing ¹⁴C inorganic carbon beneath neutral density filters. These experiments revealed that E. proxima exhibited photoinhibition at 100% full sunlight, but this depression in photosynthetic performance at high light was absent when the cells were incubated in vials containing acid-cleaned sediment. This amelioration of photoinhibition in vials containing sediment is most likely due to the downward migration of the population away from photoinhibitory light levels. These results cannot be interpreted as strictly photic effects because the temperature of the vials varied with the light treatment. The results of this study suggest that the vertical migration behavior of E. proxima and its photosynthetic physiology are intimately linked. Vertical migration provides a mechanism to deal with short-term changes in irradiance and avoid photoinhibitory light levels.     

A study of the endogenous activity rhythms of the marine isopod Exosphaeroma truncatitelson

The endogenous activity rhythms of the South African endemic isopod Exosphaeroma truncatitelson were explored. Isopods were collected on days coinciding with either a neap or spring tide. High- and low-energy beaches, which vary in the intensity of wave action, were selected as study sites. Isopods, in groups of 10–30, were placed in glass tanks with different experimental conditions: darkness + no sand, darkness with sand and natural ambient light (no sand). Activity of the isopods was measured for 50 h by recording the number of isopods swimming actively during a 30-s period. These isopods exhibited an endogenous activity rhythm which coincided with the tidal cycle even though removed from their natural habitat. Cosine curves were fitted to the observed activity data and statistical differences across variables were investigated. Peak activity was shown to occur shortly after high tide. Similar activity cycles were observed for both neap and spring tides and for different times of the year. Results also indicated that the activity cycles at both low- and high-energy beaches were similar. Two survival advantages of this rhythm are proposed: prevention of stranding on high shores and competitive avoidance with other local isopod species.     

Meiobenthos of hypersaline tropical mangrove sediment in relation to spring tide inundation

Tropical intertidal sediments often contain porewater of relatively high salinity, especially in areas exposed to longer periods without seawater inundation and high evaporation. Such an area exists on the west coast of Zanzibar: a high intertidal mangrove plateau, flooded only during spring high tides, with sediment porewater salinities commonly exceeding 100 ppt. A field survey was conducted in this area to examine variations in population density of major meiofaunal taxa and the assemblage structure of free-living marine nematodes during spring-neap tidal cycles. Samples were taken on seven occasions for two months, starting from the end of the rainy season. Porewater salinity remained high throughout the sampling period, ranging from 89 to 160 ppt. Neither spring tide inundation nor heavy rains lowered the salinity markedly. The meiofauna consisted only of four taxa, present on all sampling occasions: nematodes, harpacticoid copepods, plathyhelminthes and chironomids. Densities in surface sediments (0–5 cm) were low compared to other mangrove areas, ranging from 271 to 656 animals 10 cm^-2 with nematodes dominant on all sampling occasions (58–87%). Density fluctuations could not be explained by the effects of spring tide inundation, but the meiofauna showed significant correlations with grain size and organic material. Despite the wide range of salinity, only the numbers of chironomids were negatively correlated with increased salinity. Nematode species diversity was low in all samples, although altogether 28 species were recorded in the samples. Four species occurred in more than 50% of the samples (Microlaimus sp. (100%), Metalinhomoeus sp. (76%), Daptonema sp.l (56%), Chromadorina sp. (56%)) while 12 species were found only in one or two samples. Multidimensional scaling ordination (MDS) of the nematode species abundance data indicated little effects of spring tide inundation on the assemblage structure, but rather a successive change from wet to dry season with a reduction in species diversity and increased numbers of the dominant nematode species Microlaimus sp.     

Diel and tidal changes in the distribution and feeding habits of Japanese temperate bass Lateolabrax japonicus juveniles in the surf zone of Ariake Bay

Japanese temperate bass Lateolabrax japonicus juveniles recruit to the surf zone and grow by feeding on commonly occurring coastal copepods. However, little is known about diel and tidal patterns in their migration and feeding habits. We sampled wild juveniles during the neap and spring tides, over periods of 24 h, with small seine nets in the sand flat of the eastern part of Ariake Bay, Kyushu, western Japan. In both the neap and spring tides, abundance of juveniles significantly increased during the daytime, being highest around the time of high tide. The relative gut fullness indices of juveniles drastically increased in the morning (0700–0900) and during the flood tide in the daytime, while major prey copepod (Paracalanus spp.) density in the ambient water was relatively constant. We summarized that L. japonicus juveniles would migrate to the surf zone after sunrise to feed on copepods, and then emigrate from the surf zone after sunset. The migratory behavior of L. japonicus juveniles would be influenced by light (daytime) and feeding activity influenced by both light (morning) and tidal condition (flood tide). The intertidal region of the tidal flat was recognized to be one of the important habitats for L. japonicus during their early life history.     

Endogenous rhythms and entrainment cues of larval activity in the horseshoe crab Limulus polyphemus

The American horseshoe crab, Limulus polyphemus (Linnaeus), typically inhabits estuaries and coastal areas with pronounced semi-diurnal and diurnal tides that are used to synchronize the timing of spawning, larval hatching, and emergence. Horseshoe crabs spawn in the intertidal zone of sandy beaches and larval emergence occurs when the larvae exit the sediments and enter the plankton. However, L. polyphemus populations also occur in areas that lack significant tidal changes and associated synchronization cues. Endogenous activity rhythms that match predictable environmental cycles may enable larval horseshoe crabs to time swimming activity to prevent stranding on the beach. To determine if L. polyphemus larvae possess a circatidal rhythm in vertical swimming, larvae collected from beach nests and the plankton were placed under constant conditions and their activity monitored for 72 h. Time-series analyses of the activity records revealed a circatidal rhythm with a free-running period of ≈ 12.5 h. Maximum swimming activity consistently occurred during the time of expected falling tides, which may serve to reduce the chance of larvae being stranded on the beach and aid in seaward transport by ebb currents (i.e., ebb-tide transport). To determine if agitation serves as the entrainment cue, larvae were shaken on a 12.4 h cycle to simulate conditions during high tide in areas with semi-diurnal tides. When placed under constant conditions, larval swimming increased near the expected times of agitation. Thus, endogenous rhythms of swimming activity of L. polyphemus larvae in both tidal and nontidal systems may help synchronize swimming activity with periods of high water and inundation.     

Evidence for tidally-induced vertical migration of some gelatinous zooplankton in the Wadden Sea area near Sylt

Gelatinous zooplankton is more abundant in the Wadden Sea area of North Sylt than in the adjacent North Sea. The hypothesis is tested that medusae maintain their position in the North Sylt Wadden Sea by ascending to the surface at flood tides and descending to the bottom during ebb tides, thus avoiding the strong ebb currents which could carry them out of this food-rich area. Surface sampling at a main tidal channel revealed great differences between high tide and low tide abundances of five species of medusae in the surface layer.Rathkea octopunctata, Sarsia tubulosa, Eucheilota macultata andPleurobrachia pileus were all more abundant around high tide than during low tide. Bongo net sampling in different depth layers revealed thatPleurobrachia pileus, Bougainvillia ramosa andEucheilota maculata showed a preference for the surface layers around high tide, whereas most of the individuals were found in the deepest layer around low tide. The results suggest tidally-induced vertical migration of medusae in tidal channels. This may assist maintenance of the populations in the Wadden Sea area near Sylt.     

Heat-shock response and antioxidant defense during air exposure in Patagonian shallow-water limpets from different climatic habitats

Climate warming involves not only a rise of air temperature means, but also more frequent heat waves in many regions on earth, and is predicted to intensify physiological stress especially in extremely changeable habitats like the intertidal. We investigated the heat-shock response (HSR) and enzymatic antioxidant defense levels of Patagonian shallow-water limpets, adapted to distinct tidal exposure conditions in the sub- and intertidal. Limpets were sampled in the temperate Northern Patagonia and the subpolar Magellan region. Expression levels of two Hsp70 genes and activities of the antioxidants superoxide dismutase (SOD) and catalase (CAT) were measured in submerged and 2- and 12-h air-exposed specimens. Air-exposed Patagonian limpets showed a tiered HSR increasing from South to North on the latitudinal gradient and from high to low shore levels on a tidal gradient. SOD activities in the Magellan region correlated with the tidal rhythm and were higher after 2 and 12 h when the tide was low at the experimental site compared to the 6 h value taken at high tide. This pattern was observed in intertidal and subtidal specimens, although subtidal individuals are little affected by tides. Our study shows that long-term thermal adaptation shapes the HSR in limpets, while the oxidative stress response is linked to the tidal rhythm. Close to the warm border of their distribution range, energy expenses to cope with stress might become overwhelming and represent one cause why the limpets are unable to colonize the shallow intertidal zone.     

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.     

Physiological aspects of sound communication in crickets (Gryllus campestris L.)

Summary An extremely elaborate performance, involving endogenous timing with both tidal and lunar frequencies, has been recorded in the locomotor activity of an adult specimen of the intertidal isopod,Excirolana chiltoni. During two months of observation, under constant, non-tidal conditions, this animal showed a persistent tidal rhythm in its swimming activity. Bursts of activity were initially well synchronized with times of tide crest on the shore. The average free-running period of the tidal rhythm was about 24 h 55 min, i.e. about 5 minutes longer than the average period of the tides; thus, the loss of synchrony with the concurrent tides was very gradual. The amount of activity per burst showed a conspicuous pattern of variation, a periodic amplitude modulation which paralleled, in detail, the complex lunar cycle of changes in height of high tide. The free-running period of the bimodal, circa-lunar rhythm of amplitude modulation was one or two days longer than the natural 29-day lunar cycle of tide heights.Each feature of this recording has been qualitatively replicated in activity records from other individuals of this species. Freshly-collectedExcirolana generally show spontaneous bursts of activity at times of tide crest, bursts which are repeated as a persistent tidal rhythm, the period of which commonly departs by only a few minutes from that of the natural tidal cycle. Superimposed on the tidal rhythm is an endogenous monthly pattern of amplitude modulation, which alters the amount of activity per burst. This circa-lunar rhythm has a free-running period between about 26 and 33 days, and generally leads to maximum activity on days of highest of high tides. The net result of the tidal and lunar rhythms is an activity pattern which permits the isopods to recapitulate, in great detail, certain significant ecological aspects of the mixed, semi-diurnal tidal regime of California.The experimental data are not compatible with the hypothesis that uncontrolled environmental factors, such as vibrations from waves, were responsible for the rhythmic behavior of the animals. Neither do the data support the hypothesis that beats between the observed tidal rhythm and a hidden daily or circadian rhythm were responsible for the observed circa-lunar rhythm. Furthermore, the pattern of the circa-lunar rhythm cannot be accounted for by a single monthly oscillation in the excitability of the animals, such as might be mediated by changes in the level of an excitatory or inhibitory hormone in the circulatory system.     

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.     

TIDAL RHYTHMS: THE CLOCK CONTROL OF THE RHYTHMIC PHYSIOLOGY OF MARINE ORGANISMS

1. A great number of vital processes are rhythmic and the rhythms quite often persist in constant conditions. The best-known rhythms are circadian; much less is known about circalunadian rhythms, and this review was prepared in an attempt to rectify this deficiency. All through the article comparisons are drawn between circalunadian and circacian rhythms. 2. Activity rhythms. (a) The activity patterns of 28 intertidal animals are discussed. All describe a periodicity with a basic component of 24.8 hours, and this approximate period persists in the laboratory in constant light and temperature and in the absence of the tides. The duration of persistence ranges from a few cycles to months, and is a function of the species studied, the conditions imposed, and individual tenacity. (b) In those few cases where relatively long-term observations have been made, there is a trend for the period of the rhythm to become circatidal, or better, circalunadian. (c) The ‘desired’ phase relationship between rhythm and tidal cycle is species-specific. Geographical translocation experiments have shown that the phase is set by the local tides. (d) In some cases the amplitude of the persistent rhythm mimics the semidiurnal inequality of the tides. (e) In about a third of the species discussed, a circadian component has been found combined with the tidal component. Many of the other studies were of such short duration that a low-amplitude circadian component would have gone unnoticed. (f) The tidal rhythm is innate. However, the rhythm is (i) sometimes lacking in organisms living in non-tidal habitats, or (ii) fades after a spell of incarceration in constant conditions. Various treatments — some aperiodic — can induce the expression of the missing tidal rhythm. (g) In the green crab, removal of the eyestalks destroys the activity rhythm. 3. Vertical migration rhythms. (a) A rather surprisingly large number of intertidal animals have been found to undergo migration rhythms between the upper layers of the substratum and its surface. The movements are synchronized with the tides in nature, but most species have either been shown to be diurnal in constant conditions, or in cases where adequate testing has not been done, suspected of being so. (b) In only one species has confirming work shown that the fundamental frequency is truly tidal. This finding is especially important as it shows that tidal rhythms need only the single-cell level of organization for expression. Even at this level there appears to be a dictatorial override by a circadian clock. 4. Colour change. Low-amplitude tidal rhythms in colour change — superimposed on a more dominant circadian change — have been reported to be intrinsic in four species and inducible in a fifth. 5. Oxygen consumption. Tidal rhythms in oxygen consumption have been described for seven invertebrates and one alga; six of the species have superimposed solar-day rhythmic components also. 6. Translocation. A total of five geographical translocation experiments, in which the organisms were maintained in constant conditions throughout, have been tried. Unequivocally in one case, and possibly in a second, the test organisms rephased spontaneously to the times commensurate with local tidal conditions. In two other cases, the pretranslocation phase was retained. The fifth experiment has not been reproducible. 7. Determination of phase. (a) The tidal cycle on the home shoreline sets the phase of the inhabitant's rhythms. Even the location of a crab's burrow on the beach incline can play a determining role. (b) Paradoxically, the periodic wetting by inundation is not an important entraining factor for most intertidal organisms. Instead, the effective portions of the tidal cycle include one or more of the following. (i) Mechanical agitation, especially for animals living in an uprush zone where they are periodically subjected to the pounding surf, (ii) Temperature cycles, though they have not yet been systematically investigated, have very pronounced entraining roles in crabs. (iii) Pressure is probably not a generally important entraining agent for most intertidal organisms, but it is so for the green crab. (c) Light-dark cycles in general, whether daily or tidal in length, have no effect on the entrainment or phase setting of many tidal rhythms. There are two exceptions: (i) a 24-hour light-dark cycle is known to keep a tidal locomotor rhythm (one that becomes circalunadian in constant conditions) at a strict tidal frequency. (ii) In rhythms with both daily and tidal components, when the former is shifted by light stimuli, the latter is affected in a nearly identical manner. 8. Temperature. (a) The role of temperature on tidal rhythms is compared with its role on circadian rhythms. (b) The effects of different constant temperatures have so far been studied on only four tidal rhythms. All studies indicate a lack of any permanent change in period, which is not so with most circadian rhythms; the latter having temperature coefficients around 1.1. In two of the studies the rhythms under test temperatures were followed for less than a day, and a third study cannot be repeated. (c) Short exposure to very cold temperature pulses produced a response that may be interpreted as a temporary stoppage of the clock. Exposure to relatively less-cold pulses appear simply to reset the hands of the clock. The same responses have been demonstrated with circadian rhythms. (d) In the case of green crabs, which had become arrhythmic during prolongued captivity in the laboratory, a tidal rhythm could be reinitiated by a single short cold treatment. The cold pulse also set the phase of the rhythm. (e) A few superficial studies employing temperature steps or pulses have produced results which suggest that a phase-change sensitivity rhythm — just like that found associated with circadian rhythms — may underlie tidal rhythms. Certainly a determined search for this rhythm should be made in the near future. 9. Clock control of rhythms. (a) An argument is constructed claiming that tidal rhythms have a basic period of about 24–8 hours rather than the more expected tidal interval of 12.4 hours. In constant conditions, a circalunadian period is usually displayed. (b) After speculating that a frequency-transforming coupler may function between the clock and the overt rhythm, reasons are given that lead to the further speculation that both circadian and circalunadian rhythms could be generated by a single clock, via specific coupling mechanisms. (c) Two current hypotheses concerning the nature of the clockworks are reviewed and discussed. (d) Suggestions are made for future investigations.     

Short-term changes in activity rhythms in an intertidal arthropod (Acarina: Bdella interrupta Evans)

Summary On a marine saltmarsh, the intertidal mite, Bdella interrupta Evans, is exposed to periods of regular tidal submergence which alternate with periods of tidal emergence. The mite shows well defined day and night peaks of locomotory activity on the soil surface. During periods of tidal submergence the activity peaks showed an apparent periodicity, of around 12.5 h, characteristic of a tidal rhythm. In the presence of non-submerging tides the periodicity of the activity peaks changed to around 11.5 h. It is suggested that this shorter periodicity results either from free-running of the circatidal clock in the absence of entraining tides or from a sequence of transients that are phaseshifting towards a stable relationship with a second zeitgeber. It is further suggested that the alternation between a ca. 12.5 h and a ca. 11.5 h rhythm ensures that an activity peak is maintained in daylight and, also, reduces the possibility of inundation during the first critical floodings during sequences of rising spring tides.     

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.