The tidal rhythm of feeding and digestion in the pacific oyster, Crassostrea gigas (Thunberg)

A study has been made of the processes of extra- and intracellular digestion in the Pacific oyster, Crassostrea gigas (Thunberg) over two 24-h cycles in winter and summer.The results show that the digestive processes are discontinuous and related to the tidal cycle. Variations in tidal height resulting from a diurnal inequality of the tide affect both the relative dissolution of the crystalline style and the relative lengths of the constituent phases of the intracellular digestive process in the digestive diverticula. On a seasonal basis the style is present for a greater length of time in winter and, conversely, remains dissolved longer in summer. A seasonal variation in the structure of the digestive tubules has also been found.The results confirm conclusions reached earlier that the processes of extracellular digestion in the stomach and intracellular digestion in the digestive diverticula of intertidal bivalves are both discontinuous, alternate, and irrevocably interrelated since breakdown of the digestive cells of the digestive diverticula following intracellular digestion results in the dissolution of the crystalline style. The arrival of food in the stomach has a minimal effect upon the style. Moreover, the cytological structure of the digestive diverticula of C. gigas undergoes a series of synchronized changes related to the tidal cycle.     

MONOPHASIC AND DIPHASIC DIGESTIVE CYCLES IN VENERUPIS DECUSSATA AND CHLAMYS VARIA

Feeding and digestive cycles in Chlamys varia and Venerupisdecussata are identified and shown to relate to a tidal cycle.The processes of digestion within the diverticula of the twospecies are very different. The tubules of Venerupis are synchronizedand exhibit a monophasic cycle completed within a twelve-hourperiod, facilitating feeding at the next cycle of the tide.The digestive process of Chlamys requires twenty-four hoursand to accommodate feeding at each twelve-hour tidal cycle thetubules within the diverticula exhibit two different digestivephases simultaneously so that a diaphasic cycle is apparent.The pH of the mantle cavity and regions of the digestive tractand the formation and dissolution of the crystalline style areshown to be related to the tidal cycle. ^*Department of Brewing and Biological Sciences, Heriot-WattUniversity, Chambers Street, Edinburgh, EH1 1HX. (Received 22 March 1978;     

Feeding rhythms of the green-lipped mussel, Perna viridis (Linnaeus, 1758) (Bivalvia: Mytilidae) during spring and neap tidal cycles

Food availability and feeding responses of the green mussel Perna viridis were investigated for two complete tidal cycles during both spring and neap tides. Temporal changes in total particulate matter (TPM), particulate inorganic matter (PIM) and particulate organic matter (POM), were smaller during neap than spring tides. Seston characteristics at different times of a tidal cycle were compared for both spring and neap tides. Only during spring tides were TPM and PIM significantly higher at high tides while POM remained relatively constant (P>0.05). The clearance rate of the mussels underwent temporal variations with tides, and was a negative power function of TPM and a positive linear function of f (organic content), during both spring and neap tides. f was the key factor influencing filtration rate, organic ingestion rate, absorption rate and absorption efficiency. All feeding rates increased linearly with increases in organic content. Pseudofaeces were produced only during spring but not neap tides. Feeding rates and absorption efficiency were highest at low and lowest at high tides (P<0.01). There was no significant temporal change in the wet weight and protein content of the crystalline style with the tidal regime. For the digestive gland, alpha-amylase activity was higher at spring than at neap tides, and higher during high tides in a tidal cycle. The digestive gland cellulase activity did not change significantly with the tides. For the crystalline style, both the activity of cellulase and alpha-amylase were not significantly different (P>0.05) between spring and neap tides. Tidal rhythms in feeding and digestion in this species were likely controlled by temporal variations in food availability in the seawater. By adjusting feeding rates and enzymatic activities, absorption in Perna viridis remained constant, irrespective of the changes in food availability.     

DISTRIBUTION OF DIGESTIVE TUBULES AND FINE STRUCTURE OF DIGESTIVE CELLS OF APLYSIA PUNCTATA (CUVIER, 1803)

The distribution of digestive tubules of Aplysia punctatahas been studied in animals under experimental feeding conditions.Histological analysis of the digestive gland has revealed twotypes of tubules, called tubules A and B. Tubules of type Awere composed of basiphilic cells (calcium, excretory and thincells) and tubules of type B were lined by large digestive cellsand basiphilic cells. The latter occur in small groups, usuallyin the corners of the tubules. Type A tubules are involved inion metabolism and show a diphasic cycle (absorptive and reconstitutive)according to the height and the stage of calcium cells. TypeB tubules are involved in digestive processes and display atetraphasic cycle (holding, absorption, fragmentative and reconstitutive)depending upon the height and the stage of the digestive cells.The tetraphasic cycle was compared with the four categoriesof tubules in bivalves. It is proposed that digestive processesmay be continuous in digestive cells of A. punctata. (Received 16 November 1999; accepted 1 October 2000)     

CYCLIC ACTIVITY AND EPITHELIAL RENEWAL IN THE DIGESTIVE GLAND TUBULES OF THE MARINE PROSOBRANCH MAORICRYPTA MONOXYLA (LESSON)

Three cell types are present in tubules of the digestive glandof the marine prosobranch Maoricrypta monoxyla (Lesson). Histochemistry,and feeding and starvation experiments established that themain type, the digestive cell, is involved in endocytotic uptakeof food material from the lumen. Digestion of this materialoccurs within vacuoles, and indigestible material (indicatedby the dye trypan blue) accumulates in basal residual bodiescontaining lipofuscin pigment. Another cell type, the cryptcell, appears to secrete a glycoprotein, probably enzymaticin function. The third cell type contains large vacuoles butit has not been Established whether the contents are secretoryor excretory. The tubules undergo a cycle of digestive activity not relatedto the tidal cycle as in some marine molluscs, but probablyan indirect result of the feeding regime. The cycle begins withimmature tubules in which some endocytosis occurs. These developinto absorbing tubules involved mainly in food uptake. In maturetubules intracellular digestion occurs. At a later stage thetubules fragment to produce spherules which may conserve usefulmaterial, and finally they, disintegrate completely. Eighty per cent of the dividing cells in the epithelium occurin crypts, which are therefore considered to be the main sitesof epithelial renewal. The processes by which tubules may bereformed after breakdown are discussed (Received 28 September 1978;     

Morphofunctional changes of the digestive gland in the bivalve mollusk Crenomytilus grayanus (Dunker, 1853) in normal conditions and after parasitic invasion by trematodes

Specifics of the digestive cycle were studied in normal individuals of Gray’s mussel Crenomytilus grayanus and in those infested by trematodes from an unpolluted area of the sublittoral zone of Peter the Great Bay in May (during the day), June, July, and August. Four types of tubules corresponding to four phases of the digestive cycle were identified, and three varieties of destructing tubules were distinguished. It was shown that normally phases of absorption and digestion prevailed in the digestive gland. The portion of absorptive tubules (type II) was 95% in spring and 79% in summer. The total portion of destructing and restoring tubules (types III and IV) did not exceed 10% in spring and 20% in summer, and portion of tubules at the initial condition (type I) was 4% in spring and 2% in summer. During the day, insignificant variations in the ratio of different type tubules were recorded. It was revealed that, normally, granulocytomes were formed during utilization of necrotic tubules. Parasitic invasion of trematodes entailed swelling of the gland and a change in the proportion of tubules: the number of absorptive tubules decreased to 20%, the number of destructing and restoring tubules increased up to 60% and 12%, respectively.     

Feeding and digestion in two intertidal gammarids: Marinogammarus obtusatus and M. pirloti

The diet of two intertidal species of gammarid, Marinogammarus obtusatus Dahl and M. pirloti Sexton and Spooner is described. Both species feed largely, but not exclusively, on decaying algae, but may eat some living algae and, more rarely, carrion.
Their feeding activities are influenced by the rhythmic pattern of tidal ebb and flow, feeding only when covered by water. When immersed by the incoming tide they ingest food rapidly for a while and then feed more intermittently, ingesting small amounts at a time until they are uncovered once more by the receding tide. They probably eat sufficient food to fill the gut twice in 24 hours. The optimum rate for the digestion of food is approximately five hours, although food remains in the gut for about 12 hours.
The pH of the contents of the proventriculus, digestive gland and intestine of M. obtusatus is distinctly acid. Digestive enzymes are secreted only by the digestive gland. Carbohydrases are present which will hydrolyse soluble starch, glycogen, sucrose, maltose, arbutin, salicin, raffinose and melibiose. Inulin, lactose and cellulose are not hydrolysed. Non-specific esterase and true lipase were detected by histochemical methods.
It is concluded that the food supply available to these animals is plentiful and it is unnecessary to utilize the food supply to the maximum.     

Incorporating tides into the acoustic ecology of humpback whales

North Pacific humpback whales (Megaptera novaeangliae) migrate annually to foraging grounds in Southeast Alaska that are characterized by semidiurnal tidal cycles. Tidal activity is an important driver of marine mammal behavior on foraging grounds, but is often omitted in studies of acoustic ecology. To better understand the role of sound in this vocal species we investigated the influence of tidal height and direction on humpback whale nonsong calling behavior in Frederick Sound and described new call types for this population. The likelihood of detecting a call from the low‐frequency‐harmonic, pulsed, or noisy‐complex call classes was independent of tidal activity. The likelihood of detecting a call from the tonal call class, and a feeding call in particular, was 2.1 times higher during flood tides than during ebb tides (95% CI 1.1–4.4). This likely reflects an indirect relationship between humpback whale foraging and tides.     

An entrainment model for semilunar rhythmic swimming behaviour in the marine isopod Eurydice pulchra Leach

Entrainment experiments have been carried out with geographically widely separated populations of the sand beach isopod Eurydice pulchra Leach subjected to periods of simulated tidal agitation imposed concurrently with a 24-h light: dark (L: D) cycle. Circatidal swimming rhythms of greatest amplitude were induced when agitation was applied with the subjective timing, within the L: D cycle, of local spring high tides. This occurred in a normal L: D regime and also when the L: D regime was phase shifted through 90°. Animals previously maintained in constant darkness (D: D) and subsequently exposed to simulated tidal disturbance at various times in constant darkness were unable to modulate the amplitude of circatidal swimming activity. Isopods previously maintained in a normal L: D cycle and subsequently subjected to artificial tidal agitation in constant darkness were, however, able to modulate circatidal activity. This indicates that E. pulchra is capable of detecting tidal agitation and daily light cues and using them in conjunction with its circadian “clock” to modulate its endogenous circatidal rhythmicity. The free-running semilunar rhythm of swimming activity entrained only when the timing of agitation within the day/night cycle mimicked the pattern of local spring high tides. Agitation with the timing of neap high tides entrained no free-running circa-semilunar activity pattern.     

Progress of migrating Atlantic salmon (Salmo salar) along an estuary, observed by ultrasonic tracking

Adult Atlantic salmon were captured in the Miramichi esturary, tagged with ultrasonic transmitters, and released. Nine fish were tracked for a total of 425 h, spanning 71 flood and ebb tides. There was drifting with the tidal currents, and holding of position relative to land. Fish that achieved overall upstream progress did so by drifting with flood tidal currents and by stemming the ebb currents. Fish which did not achieve upstream progress also drifted with flood tidal currents, but these fish did not stem the ebb currents and dropped back downstream during ebb tides. No differences in movement patterns were apparent for different times of day or wind conditions.     

Flux of decapod larvae and juveniles at a station in the lower Canal de Mira (Ria de Aveiro,Portugal) during one lunar month

Summary

Emigration and immigration of decapod larvae from estuaries depend on timing of larvae occurrence in the water column relative to the tidal, tidal amplitude and day cycles. The phase relation of these natural cycles varies with tidal regime and geographically, resulting in different time-patterns of hatching of first stage larvae and of presence of late stage larvae in the water column. Vertical migration behaviour according to phase of tide also controls transport inside estuaries. These mechanisms were investigated in a field study conducted on the northwest coast of Portugal where neap ebb tides occur during the night around the quarters of the moon. Flux of decapod larvae through one sampling station was measured during one lunar month at the Canal de Mira (Ria de Aveiro) in the spring of 1990. The sampling programme was comprised of a set of 25-h fixed station studies, separated by 25-h intervals during which no sampling took place. Plankton samples were collected with a pump every hour at three depths. Current velocity and direction at the standard depths, as well as height of the water column, were also measured every hour. Hourly instantaneous flux of larvae through a 1-m-wide vertical section of the Canal de Mira was calculated for the most abundant forms. A total number of 13 combinations of species and larval stages were analyzed, belonging to the families Atelecyclidae, Pirimelidae, Portunidae, Pilumnidae, Grapsidae, Palaemonidae, Crangonidae and Thalassinidae. Patterns of net larval flux along the lunar month could be grouped into three types. Type 1 includes first zoeas that were consistently exported to the sea. Type 2 comprises late zoeas, megalops and juveniles that were consistently imported into the estuary. First zoeas that were imported during some of the 25-h studies but were exported during the others were included in Type 3; in species of this type import periods appeared to alternate with export periods according to lunar phase. Flux of Type 1 larvae followed a semi-lunar pattern. Release activity of Type 1 zoeas took place during the night and started during neap tides around the quarters of the moon, but maximum releases occurred 3–4 h after high tide of average amplitude tides, 3–4 days after the quadratures. These observations agree with the hypothesis that hatching is timed to occur on ebb tides of the largest possible amplitude so that larvae are easily dispersed from areas with a high density of predator fishes. However, based on other observations on the Portuguese coast, it cannot be ruled out that hatching might depend on a minimum number of hours of darkness experienced by the females. Larvae included in Type 2 comprise forms that may have been retained inside the estuary for the entire larval phase, as well as one form that was imported from shelf waters. No semi-lunar pattern of import was detected in this last form. Fluctuations of net flux observed in Type 3 larvae, as well in other forms that were not included in any of the types, were more difficult to explain. These larvae were first zoeas of species belonging to different taxonomic, morphological and ecological groups and may show a diversity of adaptations to the way of life of the adults. Imports and exports of larvae depended not only on time-patterns of abundance, but also on time-patterns of larval vertical distribution. As a general rule, larval stages showed patterns of depth distribution that were consistent with vertical migration rhythmic behaviours synchronized with the tidal cycle. Though the effect was not always statistically significant, first-stage larvae were closer to the surface during ebb, especially during the night, enhancing seaward transport. On the contrary, later zoeal stages, megalops and juveniles were usually closer to the surface during flood, suggesting migration to the water column during this phase of the tide and landward transport.     

Entrainment of a semilunar rhythm by a simulated moonlight cycle in the terrestrial crab,Sesarma haematocheir

Summary Sesarma haematocheir is a species of terrestrial crabs inhabiting hillsides and paddy fields near the sea. Females show a semilunar rhythm of zoea-release coinciding with days of spring tides and in addition with the time of high water occurring at the nearby seacoast about dusk. Nature of environmental stimuli or zeitgebers that induce the semilunar rhythm of zoea-release was examined experimentally. In case that a population of males and females was kept under the condition of a 24-h light-dark cycle (LD 14:10) only, females showed a free-running semilunar rhythm of egg-laying and zoea-release synchronized from field conditions. On the other hand, a semilunar rhythm of egg-laying and zoea-release was entrained by the combination of the 24-h LD and a simulated 24.8-hr moonlight cycle of which the phase was shifted in relation to the natural lunar cycle. This result suggests that the 24.8-h moonlight cycle acts as a zeitgeber of a semilunar rhythm. The 12.4-h tidal cycle parallels with the 24.8-h moonlight cycle in the field. On the basis of the perception of a distinct phase relationship between the 24-h LD and the 24.8-h moonlight cycle, it is considered that crabs substitute the 24.8-h moonlight cycle for the 12.4-h cycle of tides as a zeitgeber to synchronize the phase of the semilunar rhythm with a tidal situation.     

Patterns of temporal occurrence of brachyuran crab larvae at Saco mangrove creek, Inhaca Island (South Mozambique): implications for flux and recruitment

Temporal patterns of larval occurrence of brachyuran taxa weredescribed from Saco mangrove creek, Mozambique, based on planktonsamples. Brachyuran larvae were collected hourly in four 24h cycles during neap and spring tidal periods at a fixed station,in November 1997 and February 1998. Results indicate a semilunarcycle of larval release activity for most species. Newly-hatchedstages of sesarmids showed a peak occurrence in post-crepuscularebbing tides, Macrophthalmus spp. and Uca spp. showed highestdensities during ebbing tides at night. Dotilla fenestrata andPinnotheridae, in spite of showing a semilunar pattern, didnot display significant differences between day and night. Mostmangrove taxa showed thus a clear larval exportation behaviourduring ebb tides, with little return during the following floods,with varying degrees of dependence on the diel period. Megalopaewere restricted to the night period, and highest values wereobtained during spring tidal periods. However, maximum valuesof megalopae were obtained both during flood and ebb periods,contrary to most previous studies. This is interpreted as abi-directional transport of non-competent megalopae into andfrom the mangrove area.     

Histological and ultrastructural characterization of the alimentary system of solifuges (Arachnida,Solifugae)

Solifuges are voracious and fast predators. Once having captured a prey item, mostly small arthropods or even small vertebrates, they start feeding on their prey by constant chewing movements with their huge chelicerae. At the same time, they squeeze out the soft tissue that passes the anterior lattice‐like part of the mouthparts. The digestion of the food takes place in the midgut, which is anatomically highly complex. It consists of the midgut tube from which numerous prosomal and opisthosomal diverticula and tubular lateral branches arise. The dimorphic epithelium of the midgut tube and the diverticula is constituted of digestive and secretory cells. The digestive cells are characterized by an apical tubulus system and contain nutritional vacuoles, lipids, spherites, and glycogen. Secretory cells contain a huge amount of rough endoplasmic reticulum and secretory vacuoles. The lateral branches are ultrastructurally similar to Malpighian tubules and are likely involved in excretion. In contrast to the midgut, the epithelium of the hindgut consists of only one type of cell overlain by a thin cuticle. Digested residuals are stored in the hindgut until defecation. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Microstructural and histochemical advances on the digestive gland of the common cuttlefish,Sepia officinalis L.

The cephalopod digestive gland is a complex organ that, although analogous to the vertebrate liver, has additional functions, with special (albeit not exclusive) note on its active role in digestion. Although the structure of the digestive cell and its main constituents are well known (among which “boules” and brown bodies are distinctive features), histological details of other cell types and the general structure of the digestive gland need still further research. By a thorough combination of histological and histochemical techniques, it is shown that the digestive gland diverticula of the common cuttlefish (Sepia officinalis L.) are comprised of three essential cell types: digestive, basal and excretory. Basal (“pyramidal”) cells are multi-functional, being responsible for cell replacement and detoxification, mainly through the production of calcic spherulae containing metals like copper and lead in a complex organic matrix of proteins and ribonucleins. Since copper- and lead-positive spherulae were almost absent from other cell types and lumen of the tubules, it appears that controlled bioaccumulation of these metals, rather than excretion, is the main detoxification mechanism. The results show that the organ is crossed by an intricate network of blood vessels, especially arteries and arterioles, whose contents share histochemical properties with a particular set of “boules” that are shed into the lumen of diverticula for elimination, suggesting that the organ actively removes unwanted metabolites from the haemolymph. Conversely, the rarer excretory cells appear to be specialized in the elimination of salts. Although the exact nature of many excretory and secretory products, as the metabolic pathways that originate them, remain elusive, the findings suggest an intricate interaction between the different cell types and between these and the surrounding media: haemolymph and digestive tract.     

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.     

Foraging strategy of the mudskipper Periophthalmus sobrinus Eggert in a Kenyan mangrove

The feeding behaviour of Periophthalmus sobrinus was studied in a population living along the channels of the Tana river delta. In November 1993 mudskippers were sampled every 2 tidal h for two consecutive spring and neap tides both during the day and night. The fishes were sexed and their stomach contents analysed. The results indicated that the mudskipper has a carnivorous diet and forages during both day and night although feeding greatly decreases after dark. Quantitative and qualitative differences in diet were found during the day and night of the two synodic and tidal phases. These differences were also demonstrable in the two sexes. The study showed that females of P. sobrinus are more selective than males and that differences between the two sexes may be related to reproductive condition. Furthermore, the mudskippers showed a tidal rhythm in feeding behaviour with peaks of activity around low tide (LT) on spring tides and around high tide (HT) on neap tides. Foraging activity seems to be influenced by the different spatial and temporal distributions of the mudskipper's prey caused by synodic, tidal and diel changes in the environment.     

Role of the tidal cycle in the gonadal development and spawning of the tropical wrasse Halichoeres trimaculatus

Many wrasses on coral reefs exhibit daily spawning that peaks around daytime high tides. In this study, we examined tidal-related ovarian development in the threespot wrasse, Halichoeres trimaculatus, a species common on coral reefs in the Indo-Pacific Ocean. When the fish were collected in the morning at different tidal phases, the gonadosomatic index (GSI) and ovarian histology changed; concomitant with increases in GSI towards high tide, a clutch of the most advanced oocytes developed from vitellogenic to maturation stages. Ovulated eggs and post-ovulated follicles (POF) existed in most ovaries around high tide, but only POF remained around ebb tide, suggesting that spawning occurred during or after high tide. We noticed that tidal-related spawning was considerable in the morning and that most ovaries collected on the afternoon high tide exhibited post-spawning features. This suggests that certain labrid species possess plasticity with regard to their spawning time and utilize potent environmental cues to ensure their reproductive success. When pieces of ovary were incubated with precursor steroids, high conversion of testosterone to 17beta-estradiol occurred during high and ebb tides, while that of 17alpha-hydroxyprogesterone to 17alpha,20beta-dihydroxy-4-pregnen-3-one and 17alpha,20beta,21-trihydroxy-4-pregnen-3-one was observed during low and flood tides. Incubation of pieces of ovary with human chorionic gonadotropin resulted in similar fluctuations in the steroid hormones with tidal phase. Production of these steroid hormones correlated with oocyte development in the ovaries and was probably regulated by gonadotropin. These results demonstrate that the daily cycle is fundamental for oocyte development, and that the tidal cycle is superimposed on this process.     

Changes in the digestive gland of the loliginid squid Sepioteuthis lessoniana (Lesson 1830) associated with feeding

Changes associated with feeding in the histological and cytological structure of the digestive gland of the loliginid squid Sepioteuthis lessoniana were examined, along with the nature of both the intracellular and extracellular enzymes produced by the gland. The timing of the release of the extracellular enzymes during the digestive cycle was also determined using a quantitative experimental program. Like that of all coleoid cephalopods, the digestive gland was characterised by one type of cell with several functional stages. As is the case for other loliginid squids, however, the digestive cells did not contain the large enzyme-carrying boules that characterise the digestive glands of most cephalopods. Instead, smaller secretory granules were found in the digestive cells and these may be the enzyme carriers. The prominent rough endoplasmic reticulum, large mitochondria and active Golgi complexes present in the digestive cells are characteristic of cephalopods and indicate a high metabolic activity. Like that of other cephalopods, endocytotic absorption of nutrients and intracellular digestion occurs in the digestive gland of this squid. From quantitative and qualitative examinations of structural changes in the digestive gland of S. lessoniana after feeding, a schedule of its function during the course of digestion was proposed. This indicated that digestion was very rapid, being completed in as little as 4 h in S. lessoniana. Extracellular digestive enzymes were only released after the first hour following feeding, which implies that they are stored in the stomach between meals to increase digestive efficiency.     

Behavior of Four Species of Fiddler Crabs,Genus Uca,in Southeast Sulawesi,Indonesia

We studied the behaviors of four species of sympatric fiddler crabs on Kaledupa Island, Indonesia. Species differences in activity level, grooming, burrowing and feeding were related to their habitat and food. Uca chlorophthalmus, living in muddy mangrove areas, were inactive and spent most of the time feeding in place. Females fed 50% faster than males and spent more time feeding. U. vocans was the dominant species at the beach in silty sand and was very active. Its feeding rate was about twice that of the former species, females fed more rapidly than males, and many crabs of both sexes fed in droves at the water’s edge during ebb tides. During ebb tides, they spent most of their time feeding, while at flood tide they engaged in a greater variety of activities, including burrow maintenance. They frequently walked while feeding and interacted aggressively. U. tetragonon lived in a pebbly band along one edge of the beach, by a quay. Their feeding rate was comparable in both sexes and slower than that of U. vocans; they fed largely on filamentous algae growing on the quay, which provides better food, and fed faster during flood tide than ebb tide. They spent more time in waving and other sex-related activities, and were seldom aggressive, except during the week of the full moon. Burrowing activities included placing excavated mud balls some distance away from their burrows and re-arranging them. U. dussumieri inhabited the other end of the beach in muddier substrate. They did not have sex differences in feeding rates and their rate of scooping food into their mouths was slow, but feeding claws made multiple pinches of the substrate, thus accumulating more material in each clawful of food.