The “pelagic larvaton” and its role in the biology of the World Ocean,with special reference to pelagic larvae of marine bottom invertebrates

Ecological subdivision of marine organisms is often based on two characteristics: presence in a defined environment, and types of locomotion (degree of free active movement) in such an environment. The use of these characteristics results in a simple scheme: (1) Inhabitants of the boundary surface “ocean-atmosphere” (a zone including not only the surface film but also the thin subsurface water layer below it and the air layer just above it, i.e., pleuston and neuston). (2) Inhabitants of the deeper water layers of the ocean i.e., excluding the zone mentioned under (1): (a) passively drifting forms with very limited locomotory capacity, moving practically in the vertical plane only (plankton); (b) actively moving forms which migrate both vertically and horizontally (nekton). (3) Inhabitants of the “bottom”-benthos (level-bottom of oceans and coastal waters, tidal zones up to the upper supralittoral, different types of drifting and floating substrata, e.g. ship bottoms, harbour structures, buoys, driftwood, sargassum, whales, etc.). This simple scheme is essentially based on characteristics of adults. If developmental stages are considered, pelagic larvae of bottom invertebrates, eggs and larvae of fishes and other forms, usually present only temporarily in the plankton, neuston, and pleuston, can be distinguished as “mero-plankton”, “mero-neuston” and “mero-pleuston”, from the permanent “holo”-components of these groups. Division into “mero”-subgroups opposes all these larvae to those of planktonic, neustonic and pleustonic forms developing within the “parental” groups and their environments. However, the last category of larvae in the light of world-wide distribution of the seasonal reproductive pattern of marine invertebrates and some other organisms — especially in temperate and high latitudes — can also be rated to some degree as “mero”-(not “holo”-) components. The present paper proposes to unite all larvae of marine invertebrates (and of other organisms) undergoing pelagic development into one biological group, the “pelagic larvaton”. The main characteristic for all forms of this group is the presence of one and the same life-cycle stage in one and the same environment. All forms of the “pelagic larvaton” are, to various degrees, biologically different from their respective adult forms. Even the pelagic larvae of the holoplanktonic species exhibit some differences. Within the “pelagic larvaton”, 3 subgroups can be distinguished on the basis of their ecological peculiarities;

1. Larvae undergoing their whole development in an environment different from that inhabited by their parents and belonging to a group different from that of their parental forms; e.g. the pelagic larvae of bottom invertebrates which develop in the plankton, neuston or pleuston.
2. Larvae undergoing development in the same general pelagic environment, but in “non-parental” ecological groups; e.g. larvae of nektonic species developing in the plankton, neuston or pleuston; larvae of planktonic species in the neuston or pleuston; larvae of neustonic and pleustonic species in the plankton.
3. Larvae undergoing development in the “parental” groups; e.g. larvae of planktonic species in the plankton, of neustonic species in the neuston, or of pleustonic species in the pleuston.

In contrast to the 5 ecological groups: benthos, plankton, nekton, neuston and pleuston, the “pelagic larvaton” represents rather a biological than an ecological group. The “pelagic larvaton” comprises the 5 ecological groups and maintains the permanent turnover of organic substances between water and bottom. This group short-circuits the interrelations between the 5 ecological groups in all possible combinations. The existence of the “pelagic larvaton” presents another illustration of the unity of the biological nature of the oceans. The present paper also discusses the specific distributional patterns of the pelagic larvae of bottom invertebrates and their biological role in the seas.     

Seasonal variation in territorial response,and other aspects of the ecology of the Australian temperate pomacentrid fish Parma microlepis

The Australian temperate pomacentrid fish Parma microlepis Gunther maintains permanent, interspecifically defended territories. Breeding occurs in late spring (October to December), and juveniles after a pre-settling period of 4 to 6 weeks take up territories in the adult habitat. P. microlepis feed in their territories on benthic algae and invertebrates. Density of fish is significantly correlated with the availability of suitable shelter. There is a variation in the territorial response directed at different species, and a significant correlation exists between intensity of response to a species and its similarity to P. microlepis in use of living space. No significant correlation exists between intensity of response and dietary similarity. P. microlepis defends larger areas in the breeding than in the non-breeding season against those species which represent a possible threat to its breeding success. It shows a reduced tendency to attack juvenile conspecifics at the time of year when they would be settling from the plankton. Responses to species not likely to interfere with breeding success do not vary through the year. P. microlepis is thus seen to have complex territorial responses, specific to the species of intruder and time of year. In this it is quite similar to other territorial pomacentrids (chiefly tropical) for which information exists.     

Regulation of algal community development in a Macrocystis pyrifera forest

The effects of small and large-scale roughness, overstory development, competition for space with sessile animals, and grazing on algal community development in a subtidal Macrocystis pyrifera forest were examined using specially prepared concrete blocks as substrata. Variation in small-scale roughness (crevices and grooves in the order of 0.1 to 3 mm width and depth) had no significant effects on community composition. However, M. pyrifera colonization, algal diversity, and sessile animal biomass were higher near the upper horizontal edges of blocks and concrete prisms. This “edge” effect may result from a combination of increased spore and larval settlement and enhanced growth of plants and animals associated with the turbulent eddies formed around these obstructions. Natural and experimentally produced variations in the algal overstory demonstrated that the presence of an overstory can reduce algal diversity and cover beneath. Caging experiments suggest that predatory fishes and sea-stars indirectly affect the algal community by removing sessile animals (primarily bryozoans) which compete with the algae for space. The exclusion of grazers resulted in increased growth of Gigartina spp. Selective grazing on this genus may account for its reduced abundance in the study area.     

Exploited species impacts on trophic linkages along reef-seagrass interfaces in the Florida Keys

The removal of fish biomass by extensive commercial and recreational fishing has been hypothesized to drastically alter the strength of trophic linkages among adjacent habitats. We evaluated the effects of removing predatory fishes on trophic transfers between coral reefs and adjacent seagrass meadows by comparing fish community structure, grazing intensity, and invertebrate predation potential in predator-rich no-take sites and nearby predator-poor fished sites in the Florida Keys (USA). Exploited fishes were more abundant at the no-take sites than at the fished sites. Most of the exploited fishes were either omnivores or invertivores. More piscivores were recorded at no-take sites, but most (approximately 95%) were moderately fished and unexploited species (barracuda and bar jacks, respectively). Impacts of these consumers on lower trophic levels were modest. Herbivorous and smaller prey fish (< 10 cm total length) densities and seagrass grazing diminished with distance from reefs and were not negatively impacted by the elevated densities of exploited fishes at no-take sites. Predation by reef fishes on most tethered invertebrates was high, but exploited species impacts varied with prey type. The results of the study show that, even though abundances of reef-associated fishes have been reduced at fished sites, there is little evidence that this has produced cascading trophic effects or interrupted cross-habitat energy exchanges between coral reefs and seagrasses.     

Screening and bioassays for biologically-active substances from forty marine sponge species from San Diego,California, USA

Biologically-active substances were investigated from 35 demosponge species and 5 calcareous sponge species collected from intertidal or shallow subtidal habitats near San Diego, California, USA, from 1978 to 1980. Crude methanolic extracts of each species were tested for suppression of growth by bacteria (7 species) and a yeast. Antimicrobial activity was found in 26 Demospongiae and 2 Calcarea. Strong activity was found in 11 demosponges and, subsequently, 38 natural products with antimicrobial activity were isolated from 8 of these species (Aplysina fistularis, Dysidea amblia, Leiosella idia, Euryspongia sp., Toxadocia zumi, Axinella sp., Haliclona?cinerea and ?Pachychalina lunisimilis). Twenty-eight of these natural products (usually as pure compounds) were assayed for: (1) suppression of growth of marine fungi (3 spp.) and a red alga; (2) behavior modifications of invertebrate adults (4spp.); (3) toxicity to a goldfish; (4) inhibition of sexual reproduction of a brown alga; (5) inhibition of settlement and/or metamorphosis of late larvae or invertebrate juveniles (4spp.). Many of the natural products were also incorporated into pelleted fish food and tested for feeding-behavior modifications of fishes (5 spp.). Three of the compounds from Dysidea amblia were inactive in all tests. All other natural products were active in at least one assay, although none was active in all assays. The discussion relates the possession of biologically-active substances to the ecology of each sponge species; for example, sponges with antimicrobial substances are rarely overgrown.     

Positive indirect effects of reef fishes on kelp performance: the importance of mesograzers

It has been suggested that microcarnivorous reef fishes may play an important role in giant kelp forest communities by preventing infestations of mesograzers that could severely impact or potentially destroy recovering kelp forests after extreme disturbance events. However, these trophic linkages, specifically the direct and indirect effects of fishes on the biomass of mesograzers, grazing intensity, and the performance of giant kelp, have not been sufficiently quantified and evaluated as to their importance and in the absence of such disturbance events. We examined experimentally the effects of mesograzers on the growth and performance of giant kelp in the presence and absence of their fish predators near Santa Catalina Island, California (U.S.A.). Mesograzer biomass and grazing intensity were significantly higher when fishes were excluded from giant kelp, which in turn, lowered kelp performance. This pattern was consistent both on experimental plots of kelp as habitat isolates, and on a continuous reef. Moreover, the abundance of mesograzers was inversely related to the abundance of kelp perch among several kelp-forested reefs, suggesting that these effects can occur at larger spatial scales. Because of differences in the diet and behavior of two microcarnivorous fishes, the kelp perch and se?orita, we conducted an experiment manipulating each species and its density independently to determine their separate effects on mesograzers and kelp performance. Concurrently we examined the growth and mortality of juvenile kelp. Grazing intensity decreased, estimates of kelp performance increased, and the growth of juvenile kelp increased with increasing densities of fish but with no detectable effects between fishes. Our results demonstrate that these microcarnivorous fishes have positive indirect effects on kelp performance by reducing mesograzer biomass and grazing intensity, and the early life stages of other fishes also may be important. More specifically, these fishes have a positive effect on the density of fronds of giant kelp that can result in greater recruitment success and the abundance of kelp-associated invertebrates and fishes. Indeed, this study suggests that mesograzers have the potential to be one of the most important herbivores in kelp forest ecosystems.     

Russell's definite description analysis of production and limitation of phytoplankton species

The majority of phytoplankton species observed in the southwestern Atlantic Ocean produce new cells to equal cells lost, if any, to grazing and sinking. At the same time these species must limit new cells to numbers equalling any lost, since their abundance remains about the same in both deep and shallow areas. Of each of these species one may assert: the producer is a limiter (1). One species, coccolithus huxleyi, produces cells to offset any lost in deep water but does not limit, as it drifts into shallow water, new cells to equal the number lost in deep water, since it is more abundant in shallow than in deep water. For C. huxleyi it is true that the producer is not a limiter (2). However, once in the shallows, C. huxleyi is in a steady state, so that nutrient concentration both produces and limits new cells to equal any lost. Thus, (2) converts to (1) by having nutrient concentration the producer and the limiter. “The producer is a limiter” is a covering statement. Asserting it covers all observable species; denying it covers species not observed, those rare species which can only be found by more exhaustive sampling.     

On predation of pelagic larvae and early juveniles of marine bottom invertebrates by adult benthic invertebrates and their passing alive through their predators

The dynamic quantitative balance between prey and predator invertebrate species inhabiting the same shallow-shelf (sublittoral level bottom) benthic communities was first discussed by Thorson (1953). Thorson considered the exact timing of larval settlement of prey and predator species possessing pelagic development and temporal supression of the adult predators' feeding activities during reproduction at the time of the preys' settlement to constitute the major factors which facilitate survival of the prey species in such communities. However, information obtained demonstrates that Thorson's “mechanism of balance between predator and prey species of benthic communities” is not always effective in securing survival from predation not only of the prey's spat but even sometimes of the predator's spat also. Because of this, the “mechanism” can not be rated as universally effective in all situations. Analysis of the data so far published demonstrates that, in marine benthic communities, especially in shallow-shelf waters, it is not uncommon for gametes, larvae, or early juveniles of different prey species to pass alive through suspension (filter)-feeding and deposit-feeding adult invertebrates preying on them. Sometimes development can even continue after excretion by predators. The hypothesis of Voskresensky (1948) and Goycher (1949) of the importance of this phenomenon for the maintenance and recruitment of the mussel Mytilus edulis and other filter-feeding lamellibranchs of nearshore waters preying on their own and other lamellibranch pelagic larvae must be rejected on the basis of accumulated data on their feeding and general biology and on the adverse influence of the mucous of their faecal pellets and pseudofaeces on the larvae excreted by them alive. The data considered here demonstrate that, although the passing alive of larvae and spat of benthic invertebrates through benthic predators is not uncommon in shallow-shelf bottom-communities, it plays no important role in the processes of maintenance and recruitment of the species and communities involved nor of the marine benthos as a whole. The actual ecological significance of predation on pelagic larvae and bottom spat of benthic invertebrate prey species by all three main trophic groups of marine benthos (suspension or filter-feeders, deposit-feeders, carnivores) and its importance to predator-prey dynamics in marine benthic communities remains open to debate until more reliable quantitative data become available.     

Influence of territoriality on adult density in two rockfishes of the genus Sebastes

Nonterritorial Sebastes carnatus and S. chrysomelas existed, along with territorial individuals, at 3 tagging sites off southern California, USA, which were monitored for nearly 1 yr. To test the hypothesis that territoriality affected adult density in these species, territorial fish were removed and the subsequent utilization of vacated territories by other fish was monitored. Intrusion of other fish into vacated territories increased significantly in 90% of the removals. Other fish colonized both the feeding and sheltering parts of the vacated territories, indicating that the previous owners had successfully defended both parts of their territories. Many of the colonizers had already possessed territories; they expanded their territories or moved into presumably better havitat. Several previously-nonterritorial fish also moved into vacated areas, and at least some of them appeared to establish territories. These fish, then, had previously been capable of establishing territories, but were prevented from doing so by resident territory holders. Thus territoriality, rather than such other factors as predation or low recruitment, limited the number of territorial fish at each site. However, territorial fish did not inhibit the settlement of larval recruits, and the relative mortality rates of older territorial vs nonterritorial fish were not determined. Thus the question of whether territoriality was a major factor controlling total density remains unresolved.     

Feeding ecology of the green turtle (Chelonia mydas) at rocky reefs in western South Atlantic

Feeding ecology of green turtles was investigated between January 2005 and April 2008 at Arvoredo Reserve, Brazil (27°17′S, 48°18′W). Data were obtained through the performance of observational sessions, geo-referenced counts, benthic surveys, capture and recapture of individuals, and oesophageal lavages. This protected area was identified as an important green turtle feeding ground, used year-round by juveniles (curved carapace length = 32–83 cm). Turtles fed close to the rocky shores of the area and selected grazing sites commonly at hard-to-reach, near-vertical portions of the rocks. They were less active in cold months, and more abundant at shallow areas of the reef (0–5 m), where their preferred food items occurred. Their diet was dominated by macroalgae species but invertebrates were also present. Their main food item was the red algae Pterocladiella capillacea, which seems to be eaten through periodical cropping of its tips. Observational methods such as the ones applied here could be incorporated to other research programs aiming to understand the relationships between Chelonia mydas feeding populations and their environment.     

Tropical coastal habitats as surrogates of fish community structure, grazing, and fisheries value

Habitat maps are frequently invoked as surrogates of biodiversity to aid the design of networks of marine reserves. Maps are used to maximize habitat heterogeneity in reserves because this is likely to maximize the number of species protected. However, the technique's efficacy is limited by intra-habitat variability in the species present and their abundances. Although communities are expected to vary among patches of the same habitat, this variability is poorly documented and rarely incorporated into reserve planning. To examine intra-habitat variability in coral-reef fishes, we generated a data set from eight tropical coastal habitats and six islands in the Bahamian archipelago using underwater visual censuses. Firstly, we provide further support for habitat heterogeneity as a surrogate of biodiversity as each predefined habitat type supported a distinct assemblage of fishes. Intra-habitat variability in fish community structure at scales of hundreds of kilometers (among islands) was significant in at least 75% of the habitats studied, depending on whether presence/absence, density, or biomass data were used. Intra-habitat variability was positively correlated with the mean number of species in that habitat when density and biomass data were used. Such relationships provide a proxy for the assessment of intra-habitat variability when detailed quantitative data are scarce. Intra-habitat variability was examined in more detail for one habitat (forereefs visually dominated by Montastraea corals). Variability in community structure among islands was driven by small, demersal families (e.g., territorial pomacentrid and labrid fishes). Finally, we examined the ecological and economic significance of intra-habitat variability in fish assemblages on Montastraea reefs by identifying how this variability affects the composition and abundances of fishes in different functional groups, the key ecosystem process of parrotfish grazing, and the ecosystem service of value of commercially important finfish. There were significant differences in a range of functional groups and grazing, but not fisheries value. Variability at the scale of tens of kilometers (among reefs around an island) was less than that among islands. Caribbean marine reserves should be replicated at scales of hundreds of kilometers, particularly for species-rich habitats, to capture important intra-habitat variability in community structure, function, and an ecosystem process.     

Microhabitat-associated variability in survival and growth of subtidal solitary ascidians during the first 21 days after settlement

Newly settled juvenile ascidians were transplanted into shaded and unshaded sites at two subtidal depths to test the hypothesis that photonegative settlement behavior enhances juvenile survival. Silt, filamentous algae and grazing gastropods (Margarites pupillus) were identified as sources of mortality associated primarily with non-cryptic microhabitats. Silt inflicted heavy mortality on all six species tested. Algal overgrowth was important in shallow water, where it increased mortality and depressed growth rates over a 14-d period. Laboratory experiments demonstrated that gastropods can remove juvenile ascidians from the substratum. The sources of post-settlement mortality investigated may help determine distributional patterns of ascidians in the San Juan Islands, Washington, USA, and may also represent selective pressures that maintain negative phototaxis in the behavioral repertoires of ascidian tadpoles.     

Field survey of the occurrence and significance of regeneration in Amphiura filiformis (Echinodermata: Ophiuroidea) from Galway Bay,west coast of Ireland

The Boreo-Mediterranean amphiurid Amphiura filiformis (O. F. Müller) occurs in high densities within Galway Bay. This burrowing species consistently shows signs of recent regeneration (of both arms and disc “cap”) at the above locality. Specimens were collected by SCUBA in March, April and May 1980, May 1981, and April 1982. Biomass assays have revealed that individuals with up to 50% (mean ca. 25%) of their entire body weight consisting of regenerated tissue are not uncommon. The occurrence of arm breakage and regrowth was found to be more frequent in the proximal (basal) and mid-arm regions than in the exposed arm tip portions. Roughly 3% of the population were found to be in the process of regenerating the disc “cap” or covering. Preliminary results from gut analyses of potential predators at the sampling station indicate that young flatfish (plaice and dab) actively “crop” the arms of A. filiformis. Records from north western Europe of this species occurring in the diets of fish and invertebrates are considered. In this long-lived species, the continual necessity to regenerate is deemed to be a major drain on bodily resources.     

Wax ester biosynthesis by midwater marine animals

Lipid is a major energy reserve in many aquatic animals, and wax esters are the principal type of lipid present in most pelagic marine invertebrates and teleost fishes from deep water or near-surface cold waters. It has been suggested that these wax esters are biosynthesized by only a few organisms, and are then transferred along the food web to the fishes and marine mammals. We found that accessible mesopelagic myctophid (Lampanyctus ritteri, Stenobrachius leucopsarus and Triphoturus mexicanus) and gonostomatid (Cyclothone atraria and Gonodtoma gracile) fishes and crustaceans (Gaussia princeps, Calanus helgolandicus, Acanthephyra quadrispinosa and Sergestes prehensilis) biosynthesize wax esters from acetate, longchain alcohol or fatty acid precursors, in vivo or in vitro. In the latter experiments, organ tissues (hepatopancreas and gut) are more active than muscle, although, overall, fish muscle is probably a major site of wax ester biosynthesis in the species studied. Therefore, wax esters are not persistent dietary survivors in the food web of the oceanic midwaters; rather, most invertebrates and fishes in this environment make wax esters, modifying ingested fats to their own characteristic patterns.     

Herbivory by the Caribbean king crab on coral patch reefs

Caribbean coral reefs are increasingly dominated by macroalgae instead of corals due to several factors, including the decline of herbivores. Yet, virtually unknown is the role of crustacean macrograzers on coral reef macroalgae. We examined the effect of grazing by the Caribbean king crab (Mithrax spinosissimus) on coral patch reef algal communities in the Florida Keys, Florida (USA), by: (1) measuring crab selectivity and consumption of macroalgae, (2) estimating crab density, and (3) comparing the effect of crab herbivory to that of fishes. Mithrax prefers fleshy macroalgae, but it also consumes relatively unpalatable calcareous algae. Per capita grazing rates by Mithrax exceed those of most herbivorous fish, but Mithrax often occurs at low densities on reefs and its foraging activities are reduced in predator-rich environments. Therefore, the effects of grazing by Mithrax tend to be localized and when at low density contribute primarily to spatial heterogeneity in coral reef macroalgal communities.     

Importance of life cycle events in the population dynamics of Gonyaulax tamarensis

Life cycle changes that allow populations of the toxic dinoflagellate Gonyaulax tamarensis Lebour to inhabit the benthos and the plankton alternately are important factors regulationg the initiation and decline of blooms in restricted embavments. When the dynamics of these estuarine populations were monitored during “bloom” and “non bloom” years, it was shown that: (1) each year, germination of benthie cysts inoculated the overlying waters during the vernal warming period, but a large residual population remained in the sediments throughout the blooms; (2) the resulting planktonic population began growth under suboptimal temperature conditions; (3) the populations developed from this inoculum through asexual reproduction until sexuality (and cyst formation) were induced; (4) encystment was not linked to any obvious environmental cue and occurred under apparently optimal conditions; and (5) an increase in the number of non-mitotic swimming cells (planozygotes, the precursors to dormant cysts) accompanied the rapid decline of the planktonic population. Thus encystment, in combination with hypothesized losses due to advection and grazing, contributed substantiatly to the decline of the vegetative cell population. We conclude that the encystment/excystment cycle temporally restricts the occurrence of the vegetative population and may not be optimized for rapid or sustained vegetative growth and bloom formation in shallow embayments. The factors that distinguish “bloom” from “non-bloom” years thus appear to be operating on the growth of the planktonic population.     

Variability in grazer-mediated defensive responses of green and red macroalgae on the south coast of South Africa

Variabilities in the responses of several South African red and green macroalgae to direct grazing and the responses of one green alga to cues from grazers were tested. We used two feeding experiments: (1) testing the induced responses of three red and one green algae to direct grazing by mesograzers and (2) a multi-treatment experiment, in which the direct and indirect effects of one macrograzer species on the green alga Codium platylobium were assessed. Consumption rates were assessed in feeding assays with intact algal pieces and with agar pellets containing non-polar extracts of the test algae. Defensive responses were induced for intact pieces of Galaxaura diessingiana, but were not induced in pellets, suggesting either morphological defence or chemical defence using polar compounds other than polyphenols. In contrast, exposure to grazing stimulated consumption of Gracilaria capensis and Hypnea spicifera by another grazing species. In the multi-treatment experiment, waterborne cues from both grazing and non-grazing snails induced defensive algal traits in C. platylobium. We suggest that inducible defences among macroalgae are not restricted to brown algae, but that both the responses of algae to grazers and of grazers to the defences of macroalgae are intrinsically variable and complex.     

Larvae of a colonial ascidian use a non-contact mode of substratum selection on a coral reef

The rate at which larvae successfully recruit into communities of marine benthic invertebrates is partially dependent upon how well larvae avoid benthic predators and settle on appropriate substrata. Therefore, to be able to predict recruitment success, information is needed on how larvae search for settlement sites, whether larvae preferentially settle on certain substrata, and the extent to which there are adequate cues for larvae to find these substrata. This article describes how larvae of the colonial ascidian Diplosoma similis find settlement sites on a coral reef. Direct field observations of larval settlement were made on a fringing reef in Kaneohe Bay, Oahu, Hawaii, between September 1985 and April 1986. A comparison of the substrata that larvae contacted prior to settlement relative to the percentage cover of these substrata on the study reef suggests that larvae are using a non-contact mode of substratum identification to locate suitable settlement sites. This mode of substratum identification allowed 74% of larvae to evade predation by benthic organisms who would otherwise have eaten larvae if they had been contacted. Of those larvae that evaded predation, 88% subsequently settled on the same two substrata upon which most adults are found (dead coral or the green alga Dictyosphaeria cavernosa). This pattern of settlement was probably a result of active selection, since the two substrata cover only 14.4% of the reef's surface and currents had little effect on the direction in which larvae swam. An important contributing factor to the high success rate of larval settlement on suitable substrata was the lack of any temporal decay in substratum preference. It is concluded that for Diplosoma similis larval supply is a sufficient predictor of larval settlement rate. However, for marine invertebrates whose larvae are passively dispersed and exhibit a greater temporal decay in substratum preference, larval settlement should generally have a greater dependency on spatial variation in the abundance of benthic predators and suitable substrata.