Habitat harshness and morphodynamics: life history traits of the mole crab Emerita brasiliensis in Uruguayan sandy beaches

The habitat harshness hypothesis (HHH) postulates that in reflective beaches the harsh environment forces organisms to divert more energy towards maintenance and they therefore have lower abundance, fecundity, growth and survival rates than in dissipative beaches. Recent investigations have tested this hypothesis through single comparisons of only two beaches, and thus the observed trends in population level variables cannot be attributed incontestably to the beach state, but only to location. Here, abundance, reproduction, recruitment, population structure and body size of the intertidal mole crab Emerita brasiliensis were compared between populations from eight microtidal exposed sandy beaches with contrasting morphodynamics, sampled bimonthly during 22 months throughout the 180 km Uruguayan Atlantic coast. Physical variables and compound indices of the beach state were used to categorize sandy beaches. The results of this bi-annual large-scale analysis were fully consistent with the predictions of the HHH: abundance (total and population components), duration of the reproduction and recruitment seasons and the individual size of megalops and females of the mole crab E. brasiliensis decreased from dissipative to reflective beaches. This was reflected by linear or, mostly, nonlinear relationships between biological and both physical variables and compound indices of beach state. In conclusion, this multi-beach sampling provides compelling evidence of a consistent response of demographic and life history traits of an intertidal beach species to morphodynamic characteristics.     

Tisular and population level responses to habitat harshness in sandy beaches: the reproductive strategy of <Emphasis Type="Italic">Donax hanleyanus</Emphasis>

Microscopic analysis and field sampling procedures were used to compare demographic and reproductive strategies of the intertidal wedge clam Donax hanleyanus (Bivalvia: Donacidae) in two exposed sandy beaches with contrasting morphodynamics (reflective vs. dissipative) during 13 consecutive months. Histological analysis showed that: (1) the reproductive cycle of D. hanleyanus was more extended in the dissipative beach, and this was true for all the three pre-active (beginning of gonadal activity), active (maturation) and spawning stages; and (2) males and females showed significantly smaller sizes at sexual maturity at the reflective beach. Even though successive increments in proportion and mature at size were observed, the sigmoid function was significantly steeper at the reflective Arachania for both sexes, suggesting an abrupt transition to maturity. Field sampling revealed a more extended recruitment period at the dissipative beach, where recruits were also significantly more abundant than at the reflective beach. These results give support for the habitat harshness hypothesis, which predicts that in intertidal species capable of sustaining populations across a wide spectrum of physical conditions, such as D. hanleyanus, abundance, recruitment, size at maturity and extent of reproductive and recruitment seasons increase from reflective to dissipative beaches. However, a recent hypothesis suggests that reflective beaches acting as sink populations were not sustained, because mature and spawning individuals of both sexes were found in the reflective beach throughout the study period. Thus, we suggest that post-settlement processes are critical in modulating population patterns for this bivalve.     

Temporal variability in functional responses and prey selectivity of the pelagic mysid, Mysis mixta, in natural prey assemblages

Predation rates and prey selection of the pelagic mysid shrimp, Mysis mixta, were studied experimentally in the northern Baltic Sea in 1998 during their most intensive growth period, from June to October. Functional responses during 5 months were determined by providing the mysids with a natural zooplankton assemblage, diluted to several different concentrations. The results show that ingestion rate increased, along with mysid growth, from early summer to autumn and that saturation level was reached between 400 and 500 μg C l⁻¹. Ingestion rates increased with increasing prey concentration, and sigmoidal curves explained mostly the variation in ingestion rates (explanatory levels of 86–97%). Prey selection was evident in June, July and August, though weaker during the latter 2 months. Selection differed between the studied months but, generally, copepods were more positively selected than cladocerans. Rotifers were the main prey during June and July, when mysids were small, while larger mysids fed on copepods and cladocerans. Of the copepods, Eurytemora affinis was a truly selected species. This study shows that mysids feed on many zooplankton taxa and that they undergo ontogenetic diet shifts. Received: 19 July 2000 / Accepted: 19 October 2000     

Intertidal slope of coral sand beach as a unique habitat for fish: meiobenthic diet of the transparent sand dart, <Emphasis Type="Italic">Kraemeria cunicularia</Emphasis> (Gobiidae)

Coral sand beaches harbor gobiid sand darts (genus Kraemeria), the only fish known to live in the sand throughout their adult life. Despite the uniqueness of sand-dwelling habitat as a vertebrate, the biology of this fish remains unclear. To explore how this unique fish utilize an unusual habitat, we investigated diurnal patterns of microhabitat use and prey consumption by the transparent sand dart, Kraemeria cunicularia, at a sandy beach on Iriomote Island, the Ryukyu Archipelago, Japan. Sand darts were found in sediment in the lower intertidal zone throughout the daytime regardless of changes in tidal level, whereas at nighttime these fish were found swimming. Gut content analyses revealed that the sand dart diet was dominated by harpacticoid copepods throughout the day. Analyses of meiobenthic distribution indicated that these copepods were most abundant at lower intertidal zones where highest numbers of sand darts were found during the daytime; thus, it is possible that microhabitat use of the fish is largely determined by food availability. An extensive distributional survey throughout the Ryukyu Archipelago further indicated that sand darts prefer sandy beaches with well-sorted, coarse sand. These results provide novel insights into how sand darts respond to the tidal rhythm and highlight putative key environmental factors that determine their distribution at both regional and microhabitat scales. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Food resource use in a tropical eastern Pacific tidepool fish assemblage

An understanding of the trophic organization patterns of tropical littoral fish assemblages can contribute to the knowledge of key ecosystem processes while simultaneously assisting to validate large-scale biogeographic patterns (i.e. latitudinal patterns in fish herbivory). In the present study, the diets of eight fish species inhabiting the tide pools of a rocky shore on the western coast of Colombia (tropical eastern Pacific) are documented. A total of 17 prey items were identified, with a major representation (average percent by weight) of crabs and macroalgae items in the guts of all species. Small crustacean prey items (crabs, shrimps, copepods and amphipods) dominated the diets of most species, but consumption of macroalgae and diatoms by a significant number of species was also observed. We identified four significant trophic guilds within the assemblage using multivariate techniques (cluster analysis and nMDS): an omnivorous guild, consisting of Malacoctenus zonifer and the smallest size class of Bathygobius ramosus; a small-prey carnivorous guild, consisting of the intermediate size classes of B. ramosus and the smallest size class of Gobiesox adustus; a large-prey carnivorous guild, consisting of both largest size classes of B. ramosus and G. adustus; and an herbivorous guild consisting of Abudefduf concolor, A. troschelii and Chaenomugil proboscideus. The diet of two species slightly overlapped those of the rest of the assemblage and did not conform to any guild (Echidna nocturna and Halichoeres aestuaricola). It is hypothesised that guild formation may be a consequence of aggregation of species at abundant resources in the intertidal zone rather than a direct consequence of inter-specific competition. Ontogenetic changes in diets were observed in two resident species of the assemblage (B. ramosus and G. adustus). The latitudinal trend for herbivory inside this tropical assemblage is discussed in comparison with similar temperate studies in the eastern Pacific.     

Habitat associations and diet of young-of-the-year Pacific cod (<Emphasis Type="Italic">Gadus macrocephalus</Emphasis>) near Kodiak,Alaska

The influence of environmental variables and habitat on growth and survival of juvenile gadoid species in the Atlantic has been clearly demonstrated; conversely, in the North Pacific little is known about the habitat and ecology of juvenile Pacific cod (Gadus macrocephalus Tilesius). The hypothesis that density of young-of-the-year (YOY) Pacific cod in nearshore habitats is predicted by shallow depth and the presence of eelgrass and macroalgae was tested in a variety of nearshore habitats adjacent to commercial fishing grounds near Kodiak Island, AK. From 10 to 22 August 2002, a beach seine and small-meshed beam trawl were used to capture YOY Pacific cod (n = 254) ranging from 42 to 110 mm fork length. Depth, water temperature, salinity, sediment grain size, and percent cover by emergent structure (i.e. tube-dwelling polychaetes, sea cucumber mounds, macroalgae) were measured prior to fishing. Density of YOY Pacific cod was highest in areas of moderate depth (15–20 m) and positively and linearly related to percent cover by sea cucumber mounds and to salinity. No previous studies have documented fish utilizing sea cucumber mounds as habitat. Furthermore, eelgrass and macroalgae were inconsequential to cod distribution. Diets consisted mainly of small calanoid copepods, mysids, and gammarid amphipods and were significantly related to cod length and depth. This work provides important information on previously undocumented factors that affect distribution and feeding of YOY Pacific cod, which ultimately influence growth and survival in this species.     

Diet and prey selectivity of scyphomedusae from Port Phillip Bay,Australia

Scyphomedusae collected from Port Phillip Bay, Victoria, Australia, between 1984 and 1986, consumed a variety of zooplankton. The percentage composition of gut contents of Cyanea capillata (Linné) in order of decreasing importance was larvaceans 31%, cladocerans 29%, fish eggs 14%, copepods 11%, hydromedusae 9%, and ascidian tadpoles 3%. The percentage composition of gut contents of Pseudorhiza haeckeli Haacke was fish eggs 41%, copepods 33%, larvaceans 8%, cladocerans 4%, crab zoea 4%, and decapod larvae 1%. Both species of scyphomedusae showed strong positive selection for fish eggs and yolk-sac larvae, and negative selection for other prey items. When fish eggs were omitted from the selectivity analyses, C. capillata showed positive selection for amphipods, decapods, crab zoea, Podon spp., larvaceans and ascidian tadpoles, and negative selection for Evadne spp. and all copepod taxa. Pseudorhiza haeckeli showed positive selection for amphipods, decapod larvae, crab zoea and cladocerans, and negative selection for cirripede larvae, larvaceans and hydromedusae. Amongst copepods, P. haeckeli showed positive selection for calanoid and harpacticoid copepods and negative selection for cyclopoid copepods.     

Diel trophic interactions between vertically-migrating zooplankton and their fish predators in an eelgrass community

Diel changes in the composition of crustacean zooplankton and the diets of fish predators from an intertidal eelgrass flat were monitored concurrently. The zooplankton is characterized by two major components. The obligate zooplankters (holoplanktonic calanoid copepods and meroplanktonic decapod larvae) appear to exhibit vertical migration, being present in higher densities near the surface of the water column at night. The facultative zooplankton (amphipods and ostracods) are benthic during the day, but move up into the water column at night. Planktivorous midwaterdwelling fish consume calanoid copepods and decapod larvae during the day and cease feeding or switch their diet to amphipods at night. Benthic-dwelling fish consume some amphipods during both day and night. The factors important in prey selection by fish and the functional significance of vertical migration in both components of the zooplankton are discussed in the light of the changing patterns of fish predation.This paper is Publication No. 183 in the Ministry for Conservation of Victoria, Environmental Studies Series.     

Diet of orange roughy,Hoplostethus atlanticus (Pisces: Trachichthyidae) on the Challenger Plateau,New Zealand

Stomach contents of 687 orange roughy [Hoplostethus atlanticus (Collett, 1889)] from the Challenger Plateau, sampled in March, July and November 1984, were examined. The relative importance of different components of the diet was assessed using three feeding indices which combine, in different ways, percentage frequency of occurrence, percentage number, and percentage weight of prey categories. For both sexes, for all size classes and at any time of the year, the most frequent and abundant components of the diet were prawns, followed by fish, squid, amphipods and mysids. The main natant decapod families were Oplophoridae, Pasiphaeidae, and Sergestidae. Fish, mainly Chauliodontidae and Myctophidae, were the most important prey by weight. Most prey species were benthopelagic and mesopelagic organisms which move towards the surface at night. However, H. atlanticus can be caught by bottom trawl (between 750 and 1 200 m) during any 24 h period, and there was no evidence of vertical migration in search of their prey. When the fish grow in length, there is a transition in the diet from prawns, mysids, and fish, to prawns, fish and squid. Squid were not found in the stomach contents of fish smaller than 20 cm. These dietary changes may be linked to modifications in morphology with growth. The relative proportions of the main dietary items, and in particular the natant decapod families, varied with time of the year. Also, the stomach data seem to indicate an increasing importance of fish and squid in the diet in deeper water. H. atlanticus appears to be an opportunistic predator, consuming a wide variety of invertebrates and fishes. Our results provide evidence to support the notion that benthopelagic predators which consume vertically migrating mesopelagic fish, have an important role in the transfer of energy to the benthos.     

Patterns of selective predation by juvenile,benthivorous fish on estuarine macrofauna

Benthic feeding on macrofauna was studied in juveniles of the sparids Lithognathus lithognathus and Rhabdosargus holubi in the upper reaches of the Gamtoos Estuary, South Africa. Fish and benthic macrofauna were sampled simultaneously, and the selection of invertebrate prey assessed. Both fish species strongly selected for corophioid amphipods and consumed other benthic taxa in low numbers. R. holubi also exploited aquatic autotrophs, while L. lithognathus had a narrow prey-spectrum, feeding almost exclusively on the tube-dwelling amphipod Grandidierella lignorum. G. lignorum was the most abundant prey species, both in the benthos and the fish's diet. This species also showed prominent behavioural differences between the sexes; males were markedly more active on the sediment surface than females, who rarely left their tubes during the day. Males switched from an infaunal to epifaunal microhabitat in search of receptive females, concurrently increasing their exposure to fish predators. Consequently, L. lithognathus selected significantly more males than female amphipods, causing a marked bias towards females in the sex ratio and age-structure of the amphipod population. Juvenile amphipods were less preyed upon, presumably as a result of lower prey-detection or capture efficiency by the predators. Accepting current notions about predation as an important structuring element for benthic communities, our data also stress the prominence of size-and sex-selective predation in structuring individual prey populations.     

Predatory feeding of two marine mysids

Predatory feeding of the marine mysids Mysidopsis bigelowi and Neomysis americana on several species of co-occurring copepods was examined in laboratory experiments. M. bigelowi exhibited a curvilinear functional response; there was a negative logarithmic relationship between prey density and clearance rates. N. americana also exhibited higher clearance rates at lower prey densities. Increased clearance rates at lower prey densities were probably due to increased swimming speed or reaction distance as hunger increased. This response occurred only when mysids could visually locate prey; in complete darkness clearance rates were significantly lower and independent of prey density. Feeding rates on different prey species were only partially dependent on prey size; prey movement patterns and escape behavior also strongly affected feeding rates. M. bigelowi showed active prey selection when offered a choice of different prey species. Estimates of predation rates of estuarine mysid populations indicate that they could have a significant effect on co-occurring copepod populations.     

Fatty acid signatures of stomach contents reflect inter- and intra-annual changes in diet of a small pelagic seabird,the Thin-billed prion <Emphasis Type="Italic">Pachyptila belcheri</Emphasis>

In diet analyses of seabirds, fatty acid signatures (FAS) can be used to overcome biases due to differential digestion of prey and enable the analysis of very digested diet samples. We applied FAS analysis to stomach contents of a small sub-Antarctic seabird, the Thin-billed Prion Pachyptila belcheri, which feeds mainly on squid during incubation and on crustacea during chick rearing. This seasonal dietary switch of Thin-billed prions was reflected in differences in FAS in regurgitates, as were inter-annual differences in diet composition. A discriminant function analysis correctly classified 93.4% of cases with respect to year (2006–2008) and stage of the breeding cycle (incubation versus chick rearing). The dominant types of crustacea in the diet of Thin-billed prions (amphipods Themisto gaudichaudii, euphausiids, decapods Munida gregaria, and calanoid copepods) were distinguished well by characteristic FAS patterns. However, the FAS of the two main types of prey by volume, amphipods T. gaudichaudii and squid Gonatus antarcticus, were similar to each other. Although FAS were successfully applied in the analysis of prey in stomach contents of prions, FAS of some prey species were similar and may not be distinguishable from each other if used in quantitative diet analyses.     

<Emphasis Type="Italic">Lyngbya majuscula</Emphasis> blooms and the diet of small subtropical benthivorous fishes

Increasing concerns about the ecological impacts of ongoing and possibly worsening blooms of the toxic, carcinogenic cyanobacteria Lyngbya majuscula in Moreton Bay, Australia, led us to assess differences in meiofaunal prey assemblages between bloom and non-bloom substrates and the potential dietary impacts of dense L. majuscula blooms on the omnivorous benthivore, the Eastern Long-finned Goby, Favonigobius lentiginosus and the obligate meiobenthivorous juveniles of Trumpeter Whiting, Sillago maculata. Marked differences in invertebrate communities were found between sandy and L. majuscula bloom foraging substrates, with copepods significantly more abundant (18.49% vs. 70.44% numerical abundance) and nematodes significantly less abundant (55.91% vs. 1.21% numerical abundance) within bloom material. Gut analyses showed that bentho-planktivorous fishes exposed to L. majuscula in captivity had consumed a significantly greater quantity of prey by both total number (P < 0.0019) and volume (P < 0.0006) than fish exposed to sand treatments. Thus, it is likely for such fishes that L. majuscula blooms increase rates of prey encounter and consumption, with consequent changes in trophic relationships through shifts in predator–prey interactions between small benthivorous fishes and their meiofaunal prey.     

The role of the generally unrecognised microprey source as food for larval fish in the Irish Sea

Few studies have studied the food supply to larval fish in the Irish Sea; thus, we have assessed the full prey-field available to larval fish, ranging from protozoa to copepods. Specifically we assessed if fish larvae feed on protozoa, as suggested by others, and if densities of the protozoa and the appropriate size of metazoan prey were previously underestimated. By examining the gutcontents of fish larvae, the prey available to them, and the potential accessibility of prey to fish, we develop a simple food web, presented as a box-model. By doing so, we indicate that the lack of focus on small metazoa and protozoa has underestimated the food available to fish larvae; without these, we might have concluded that prey levels were too low to support the growth of the larval fish assemblage. Our methods were as follows. Sampling was at two sites, off the Isle of Man, with distinct physical and biological structures, soon after fish spawning: the southwest coast, where many species occur in spring-summer (23 April; 6, 19 May; 1, 22 June; 12 July) and the east coast, where only herring larvae occur in September–November (12, 28 October). Microplankton (15–200 μm), mesozooplankton, and larval fish were collected at 1, 15, and 25 m: microplankton with 1.5 L bottles and a 64 μm-mesh net; mesozooplankton and larval fish with a Gulf VII high-speed sampler (280 μm mesh). The 64 μm mesh net, mounted on the Gulf VII, provided simultaneous hauls. Fixed samples were evaluated to determine species composition, abundance, and biomass. Larval fish diet was determined from fish collected by short net hauls: fixed guts were examined and prey, including protozoa, analysed. Using physical data as a guide, plankton data were integrated through the water column to determine standing stocks. Size-based food availability to larval fish was estimated from the gut contents. The role of protozoa was examined, assuming that they are digested at the same rate as metazoan and if they are digested 2.5–10 times faster; increased digestion rates indicated that they contributed substantially to the larval fish diet.     

Sediment characteristics and colonization of soft-bottom benthos: a field manipulation experiment

A field manipulation experiment was carried out to test the effects of sediment characteristics (particle size and organic content) on colonization of soft-bottom benthos. Defaunated mud and sand in experimental trays were exposed at sub-tidal levels of two sites with sandy and muddy sea bottom, and retrieved monthly for examination of benthic composition. Seventy-eight out of the 107 species recorded, and 50% of the dominant species identified in the present study, were common in both the sand and mud trays. Classification analysis revealed higher faunal similarity of benthic composition in both sediment types within each of the two study sites. Results of three-way ANOVA (Sediment, Site and Time) showed that sediment effect was only significant for colonization of bivalves and gastropods, but not for polychaetes, amphipods, total species and individual numbers, or overall benthic diversity and evenness. This suggested that many of the benthic species can colonize both sediment types, and that factors other than sediment particle size and organic content may play a significant role in determining colonization of these species. The faunal composition of the same sediment type between sites was found to be different, indicating that the availability and abundance of benthic larvae/adults may be more important than sediment characteristics in determining benthic settlement. An increase in faunal similarity between mud and sand was observed over the study period, implying that the importance of sediment characteristics to benthic colonization decreased with exposure time. Received: 29 November 1996 / Accepted: 16 January 1997     

Ontogeny of diet shifts by a microcarnivorous fish, Cheilodactylus spectabilis : relationship between feeding mechanics, microhabitat selection and growth

The mechanisms leading to ontogenetic shifts in prey selectivity are examined for the temperate microcarnivore Cheilodactylus spectabilis (Cheilodactylidae) in north-eastern New Zealand. These fish prey on invertebrates associated with benthic turf and foliose algae, using a suctorial feeding mode combined with oral sorting. All sizes of fish feed in the same shallow-water habitat using the same feeding mode. Dietary analysis revealed that while all sizes of fish consumed similar taxa, the relative proportion of taxa consumed reflected fish size. Juveniles consumed mainly gammarid amphipods while large adults targeted ophiuroids, with an abrupt shift from feeding predominantly on amphipods at 250 mm standard length. This dietary shift loosely coincided with the onset of sexual maturity and a change in growth trajectory, although dietary trends did not differ between sexes. Both juveniles and adults were found to select particular taxa from the available turf micro-fauna, with juveniles consuming smaller sizes of amphipods than adults. Microhabitat use was also found to change ontogenetically. Detailed observations on feeding mechanics suggested that size-related changes in suctorial force allowed the exploitation of a broader range of microhabitats with increasing fish size. Received: 20 June 1997 / Accepted: 1 April 1998     

Locomotor activity patterns and feeding habits in the prawn Palaemon xiphias (Crustacea: Decapoda: Palaemonidae) in Alfacs Bay,Ebro Delta (northwest Mediterranean)

Diel activity rhythms of the prawn Palaemon xiphias Risso, 1818 from Alfacs Bay, Ebro Delta, were studied by time-lapse videorecordings. Activity displayed an endogenous circadian rhythm, with maximum activity at night. Feeding habits were studied by frequency of occurrence and by the points method. The food of P. xiphias consisted mainly of crustaceans; the remains of amphipods, isopods, mysids, copepods, decapods and ostracods were identified. The remaining items consisted of molluscs, polychaetes, ophiuroids, plant material, sand, and unidentified organic debris. The results indicate that P. xiphias is a predator of benthic invertebrates rather than a scavenger or detritus feeder. Diet composition changes with increasing size of the prawn.     

Reproductive plasticity in mole crabs, Emerita brasiliensis, in sandy beaches with contrasting morphodynamics

Ovarian macroscopical analysis, histological validation and field sampling procedures were used to evaluate the variability in reproductive traits of the mole crab Emerita brasiliensis Schmitt, 1935 (Decapoda: Hippidae) in two exposed sandy beaches of Uruguay with contrasting morphodynamics. All developmental stages involved in the complex life cycle exhibited lower abundance, individual size and temporal occurrence in a harsh reflective beach, compared with a more benign dissipative environment. In addition, this population showed more compressed events of the reproductive cycle (e.g oogenesis, encounter of potential mate and female parental care) and recruitment period. However, the beginning of the vitellogenesis, ovary maturity, male sexual differentiation and ending of spermatogenesis occurred at smaller sizes under dissipative conditions. These results disagree with the recent findings of delayed sexual maturity in dissipative beaches. We postulate that, in dissipative conditions, high food availability might allow an overlapping of reproductive and moulting processes, and thus females may reach optimum size and sexual maturity with fewer moults than in reflective beaches. Hence, reproductive responses must be considered not only in relation to environmental harshness, but also in the context of life history traits and their phylogenetic and allometric constraints. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.