Interactive effects of habitat selection, food supply and predation on recruitment of an estuarine fish

 Seagrass meadows are often important habitats for newly recruited juvenile fishes. Although substantial effort has gone into documenting patterns of association of fishes with attributes of seagrass beds, experimental investigations of why fish use seagrass habitats are rare. We performed two short-term manipulative field experiments to test (1) the effects of food supply on growth and densities of fish, and (2) effects of predation on the density and size distribution of fish recruits, and how this varies among habitat types. Experiments were conducted in Galveston Bay, Texas, and we focused on the common estuarine fish, pinfish Lagodon rhomboides. In the first experiment, replicate artifical seagrass and sand plots were either supplemented with food or left as controls. Recruitment of pinfish was significantly greater to seagrass than sand habitats; however, we detected no effect of food supplementation on the abundance of recruits in either habitat. Pinfish recruits in artifical seagrass grew at a significantly faster rate than those in sand habitats, and fish supplemented with food exhibited a greater growth rate than controls in both sand and artifical grass habitats. In our second experiment, we provided artificial seagrass and sand habitats with and without predator access. Predator access was manipulated with cages, and two-sided cages served as controls. Recruitment was significantly greater to the cage versus cage-control treatment, and this effect did not vary between habitats. In addition, the standard length of pinfish recruits was significantly larger in the predator access than in the predator exclusion treatment, suggesting size-selective predation on smaller settlers or density-dependent growth. Our results indicate that the impact of predation on pinfish recruits is equivalent in both sand and vegetated habitats, and thus differential predation does not explain the higher recruitment of pinfish to vegetated than to nonvegetated habitats. Since predators may disproportionately affect smaller fish, and a limited food resource appears to be more effectively utilized by fish in vegetated than in unvegetated habitats, we hypothesize that pinfish recruits may select vegetated habitats because high growth rates allow them to achieve a size that is relatively safe from predation more quickly. Received: 10 October 1996 / Accepted: 5 April 1997     

Consistently outstanding recruitment of five species of fish to a seagrass bed in Botany Bay,NSW

Abstract Variability in abundance of fish associated with beds of the seagrass Zostera capricorni was documented at 16 sites in NSW. Samples were collected from each site by seine net every 3 months for 2.5 years. One site had significantly greater abundances of recruits of five species of economically important fish during all three recruitment seasons. This site was within a small, constructed harbour in Botany Bay. Between June and March of each year abundances of five species (Acanthopagrus australis, Rhabdosargus sarba, Girella tricuspidata, Achoerodus viridis and Meuschenia trachylepis) were up to 73 times greater than abundances at the other 15 sites. However, at other times during the year, there was no significant difference between this site and the 15 others. Understanding the processes that contribute to the success of this modified site as a recruitment sink should aid in increasing the potential of other areas as fish nursery habitat, and contribute to the management and productivity of fish populations.     

Fish Assemblages on Estuarine Artificial Reefs: Natural Rocky-Reef Mimics or Discrete Assemblages?

If the primary goal of artificial reef construction is the creation of additional reef habitat that is comparable to adjacent natural rocky-reef, then performance should be evaluated using simultaneous comparisons with adjacent natural habitats. Using baited remote underwater video (BRUV) fish assemblages on purpose-built estuarine artificial reefs and adjacent natural rocky-reef and sand-flat were assessed 18 months post-deployment in three south-east Australian estuaries. Fish abundance, species richness and diversity were found to be greater on the artificial reefs than on either naturally occurring reef or sand-flat in all estuaries. Comparisons within each estuary identified significant differences in the species composition between the artificial and natural rocky-reefs. The artificial reef assemblage was dominated by sparid species including Acanthopagrus australis and Rhabdosargus sarba. The preference for a range of habitats by theses sparid species is evident by their detection on sand-flat, natural rocky reef and artificial reef habitats. The fish assemblage identified on the artificial reefs remained distinct from the adjacent rocky-reef, comprising a range of species drawn from naturally occurring rocky-reef and sand-flat. In addition, some mid-water schooling species including Trachurus novaezelandiae and Pseudocaranx georgianus were only identified on the artificial reef community; presumably as result of the reef''s isolated location in open-water. We concluded that estuarine artificial reef assemblages are likely to differ significantly from adjacent rocky-reef, potentially as a result of physical factors such as reef isolation, coupled with species specific behavioural traits such as the ability of some species to traverse large sand flats in order to locate reef structure, and feeding preferences. Artificial reefs should not be viewed as direct surrogates for natural reef. The assemblages are likely to remain distinct from naturally occurring habitat comprised of species that reside on a range of adjacent natural habitats.     

Post-larval French grunts (Haemulon flavolineatum) distinguish between seagrass, mangrove and coral reef water: Implications for recognition of potential nursery habitats

Environmental cues like sound, magnetic field, oceanic currents, water chemistry or habitat structure are believed to play an important role in the orientation of reef fish towards their settlement habitat. Some species of coral reef fish are known to use seagrass beds and mangroves as juvenile habitats. Once oceanic larvae of these fish have located a coral reef from the open ocean, they still have to find embayments or lagoons harbouring these juvenile habitats. The sensory mechanisms that are used for this are still unknown. In the present study, experiments were conducted to investigate if recruits of the French grunt (Haemulon flavolineatum) respond to habitat differences in water type, as mangrove/seagrass water may differ in biotic and abiotic compounds from coral reef water. Our results show that post-larvae of a reef fish that is highly associated with mangroves and seagrass beds during its juvenile life stage, choose significantly more often for water from mangroves and seagrass beds than for water from the coral reef. These results provide a more detailed insight in the mechanisms that play a role in the detection of these juvenile habitats.     

海草床育幼功能及其机理

海草床是近岸海域中生产力极高的生态系统,是许多海洋水生动物的重要育幼场所。从生物幼体的密度、生长率、存活率和生境迁移4个方面阐述海草床育幼功能,并从食源和捕食压力两个方面探讨海草床育幼功能机理。许多生物幼体在海草床都呈现出较高的密度、生长率和存活率,并且在个体发育到一定阶段从海草床向成体栖息环境迁移。丰富的食物来源或较低的捕食压力可能是海草床具有育幼功能的主要原因,但不同的生物幼体对海草床的利用有差异,海草床育幼功能的机理在不同环境条件下也存在差异。提出未来海草床育幼功能的重点研究方向:(1)量化海草床对成体栖息环境贡献量;(2)全球气候变化和人类活动对海草床育幼功能的影响;(3)海草床育幼功能对海草床斑块效应和边缘效应的响应,以期为促进我国海草床育幼研究和海草床生态系统保护提供依据。     

Different Surrounding Landscapes may Result in Different Fish Assemblages in East African Seagrass Beds

Few studies have considered how seagrass fish assemblages are influenced by surrounding habitats. This information is needed for a better understanding of the connectivity between tropical coastal ecosystems. To study the effects of surrounding habitats on the composition, diversity and densities of coral reef fish species on seagrass beds, underwater visual census surveys were carried out in two seagrass habitat types at various locations along the coast of Zanzibar (Tanzania) in the western Indian Ocean. Fish assemblages of seagrass beds in a marine embayment with large areas of mangroves (bay seagrasses) situated 9 km away from coral reefs were compared with those of seagrass beds situated on the continental shelf adjacent to coral reefs (reef seagrasses). No differences in total fish density, total species richness or total juvenile fish density and species richness were observed between the two seagrass habitat types. However, at species level, nine species showed significantly higher densities in bay seagrasses, while eight other species showed significantly higher densities in reef seagrasses. Another four species were exclusively observed in bay seagrasses. Since seagrass complexity could not be related to these differences, it is suggested that the arrangement of seagrass beds in the surrounding landscape (i.e. the arrangement on the continental shelf adjacent to the coral reef, or the arrangement in an embayment with mangroves situated away from reefs) has a possible effect on the occurrence of various reef-associated fish species on seagrass beds. Fish migration from or to the seagrass beds and recruitment and settlement patterns of larvae possibly explain these observations. Juvenile fish densities were similar in the two types of seagrass habitats indicating that seagrass beds adjacent to coral reefs also function as important juvenile habitats, even though they may be subject to higher levels of predation. On the contrary, the density and species richness of adult fish was significantly higher on reef seagrasses than on bay seagrasses, indicating that proximity to the coral reef increases density of adult fish on reef seagrasses, and/or that ontogenetic shifts to the reef may reduce adult density on bay seagrasses.     

Microhabitat use by a post-settlement stage estuarine fish: evidence from relative abundance and predation among habitats

Seagrass beds provide food and shelter for many fish species. However, the manner in which fishes use seagrass bed habitats often varies with life stage. Juvenile fishes can be especially dependent on seagrass beds because seagrass and associated habitats (drift macroalgae) may provide an effective tradeoff between shelter from predation and availability of prey. This study addressed aspects of habitat use by post-settlement pinfish, Lagodon rhomboides (Linneaus), an abundant and trophically important species in seagrass beds in the western North Atlantic and Gulf of Mexico. Abundance of post-settlement fish in seagrass beds was positively related to volume of drift macroalgae, but not to percent cover of seagrass, indicating a possible shelter advantage of the spatially complex algae. Tethering experiments indicated higher rates of predation in seagrass without drift macroalgae than in seagrass with drift macroalgae. Aquarium experiments showed lower predation with higher habitat complexity, but differences were only significant for the most extreme cases (unvegetated bottom, highest macrophyte cover). Levels of dissolved oxygen did not differ between vegetated and unvegetated habitats, indicating no physiological advantage for any habitat. Seagrass beds with drift macroalgae provide the most advantageous tradeoff between foraging and protection from predation for post-settlement L. rhomboides. The complex three-dimensional shelter of drift macroalgae provides an effective shelter that is embedded in the foraging habitat provided by seagrass. Drift macroalgae in seagrass beds is a beneficial habitat for post-settlement L. rhomboides by reducing the risk of predation, and by providing post-settlement habitat within the mosaic (seagrass beds) of adult habitat, thus reducing risks associated with ontogenetic habitat shifts.     

Variation in fish density,assemblage composition and relative rates of predation among mangrove,seagrass and coral reef habitats

We tested the hypothesis for several Caribbean reef fish species that there is no difference in nursery function among mangrove, seagrass and shallow reef habitat as measured by: (a) patterns of juvenile and adult density, (b) assemblage composition, and (c) relative predation rates. Results indicated that although some mangrove and seagrass sites showed characteristics of nursery habitats, this pattern was weak. While almost half of our mangrove and seagrass sites appeared to hold higher proportions of juvenile fish (all species pooled) than did reef sites, this pattern was significant in only two cases. In addition, only four of the six most abundant and commercially important species (Haemulon flavolineatum, Haemulon sciurus, Lutjanus apodus, Lutjanus mahogoni, Scarus iserti, and Sparisoma aurofrenatum) showed patterns of higher proportions of juvenile fish in mangrove and/or seagrass habitat(s) relative to coral reefs, and were limited to four of nine sites. Faunal similarity between reef and either mangrove or seagrass habitats was low, suggesting little, if any exchange between them. Finally, although relative risk of predation was lower in mangrove/seagrass than in reef habitats, variance in rates was substantial suggesting that not all mangrove/seagrass habitats function equivalently. Specifically, relative risk varied between morning and afternoon, and between sites of similar habitat, yet varied little, in some cases, between habitats (mangrove/seagrass vs. coral reefs). Consequently, our results caution against generalizations that all mangrove and seagrass habitats have nursery function.     

Evaluating the impact of predation by fish on the assemblage structure of fishes associated with seagrass (Heterozostera tasmanica) (Martens ex Ascherson) den Hartog, and unvegetated sand habitats

The role of fish predation in structuring assemblages of fish over unvegetated sand and seagrass was examined using enclosure and exclusion cages to manipulate the abundance of predatory fish from November 1998 to January 1999. In our exclusion experiment, piscivorous fish were excluded from patches of unvegetated sand and seagrass to measure how they altered abundances of small fishes, i.e., fish <10 cm in length. Habitats from which piscivorous fish were excluded contained more small fish than those with partial cages, which in turn contained more fish than uncaged areas. These patterns were consistent between unvegetated sand and seagrass areas, although the relative differences between predator treatments varied with habitat. Overall, small fish were more abundant in unvegetated sand than seagrass. Atherinids and syngnathids were the numerically dominant families of small fish and varied in complex ways amongst habitats and cage treatments. The abundance of atherinids varied inconsistently between cage treatments through time. Only during the final two sampling times did the abundance of atherinids vary significantly across cage treatments. Syngnathids were strongly associated with seagrass and were significantly more abundant in caged than uncaged habitats. In our enclosure experiment, five individuals of a single species of transient piscivorous fish, Western Australian salmon (Arripidae: Arripis truttacea Cuvier), were enclosed in cages to provide an estimate of the potential for this species to impact on small fish. The abundance of small fish varied significantly between cage treatments. Small fish were more abundant in enclosure cages and exclusion cages than uncaged areas; however, there was no difference in the abundance of small fish in enclosure cages and partial cages, and no difference between exclusion cages and partial cages. These patterns were consistent amongst habitats. Atherinids and syngnathids were again the numerically dominant families of small fish; atherinids varied more with cage structure while syngnathids did not vary statistically between cages, blocks (locations within which a single replicate of each cage treatment was applied) or habitats. Dietary analysis of caged A. truttacea demonstrated the potential for this species to influence the assemblage structure of small fish through predation - atherinids were consumed more frequently in unvegetated sand than seagrass, and syngnathids were consumed only in seagrass, where they are most abundant. Observations of significant cage or predation effects depended strongly on the time at which sampling was undertaken. In the case of the atherinids, no predation or cage effects were observed during the first two sampling times, but cage effects and predation effects strongly influenced abundances of fish during the third and fourth sampling times, respectively. Our study suggests that transient piscivorous fish may be important in structuring assemblages of small fish in seagrass and unvegetated sand, and seagrass beds may provide a refuge to fishes. But the importance of habitat complexity and predation, in relation to the potentially confounding effects of cage structure, depends strongly on the time at which treatments are sampled, and the periodicity and multiplicity of sampling should be considered in future predation studies.     

Coastal Fish Assemblages Reflect Geological and Oceanographic Gradients Within An Australian Zootone

Distributions of mobile animals have been shown to be heavily influenced by habitat and climate. We address the historical and contemporary context of fish habitats within a major zootone: the Recherche Archipelago, southern western Australia. Baited remote underwater video systems were set in nine habitat types within three regions to determine the species diversity and relative abundance of bony fishes, sharks and rays. Constrained ordinations and multivariate prediction and regression trees were used to examine the effects of gradients in longitude, depth, distance from islands and coast, and epibenthic habitat on fish assemblage composition. A total of 90 species from 43 families were recorded from a wide range of functional groups. Ordination accounted for 19% of the variation in the assemblage composition when constrained by spatial and epibenthic covariates, and identified redundancy in the use of distance from the nearest emergent island as a predictor. A spatial hierarchy of fourteen fish assemblages was identified using multivariate prediction and regression trees, with the primary split between assemblages on macroalgal reefs, and those on bare or sandy habitats supporting seagrass beds. The characterisation of indicator species for assemblages within the hierarchy revealed important faunal break in fish assemblages at 122.30 East at Cape Le Grand and subtle niche partitioning amongst species within the labrids and monacanthids. For example, some species of monacanthids were habitat specialists and predominantly found on seagrass (Acanthaluteres vittiger, Scobinichthys granulatus), reef (Meuschenia galii, Meuschenia hippocrepis) or sand habitats (Nelusetta ayraudi). Predatory fish that consume molluscs, crustaceans and cephalopods were dominant with evidence of habitat generalisation in reef species to cope with local disturbances by wave action. Niche separation within major genera, and a sub-regional faunal break, indicate future zootone mapping should recognise both cross-shelf and longshore environmental gradients.     

The effects of predation by juvenile fish on the meiobenthic community structure in a natural pond

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Effect of simulated macroalgae on the fish assemblage associated with a temperate reef system

Increased habitat complexity is supposed to promote increased diversity, abundance and biomass. This study tested the effect of the macroalgal cover on temperate reef fishes by mimicking macroalgae on artificial reefs in NW Sicily (Mediterranean Sea). Macroalgal cover affected reef fishes in different ways and independently of intrinsic temporal trends. The fish assemblages of manipulated and control artificial reef units differed in the relative abundances of the associated species, but little in species composition. In line with studies in seagrass habitats, fishes were most abundant in reefs covered by artificial macroalgae. Three species (Boops boops, Serranus scriba and Symphodus ocellatus) exhibited consistently greater abundance on vegetated reef units than on control reef units. The total number of species and the abundance of three particular species (S. scriba, S. ocellatus and Thalassoma pavo) displayed temporal trends which were independent on short and large temporal scales. Only fish total biomass and one species (Spicara flexuosa) displayed strong effects of interaction among the experimental factors. Mechanisms to explain these findings are discussed from observational evidence on habitat use and interactions among multiple species. This study highlights that manipulative experiments involving repeated sampling of fish in artificial habitats appear to be a valid approach to study fish-habitat relationships in fluctuating environments. It is also concluded that macroalgae mimics may serve as a tool for restoring lost marine vegetated habitats when current human-induced conditions prevent the recovery of pristine macroalgal stands.     

Fish Assemblages in Different Shallow Water Habitats of the Venice Lagoon

The small-sized fish assemblages of the Venice Lagoon were investigated and compared among five shallow subtidal habitats (seagrass beds, sparsely vegetated habitats, unvegetated sand bottoms, mudflats and saltmarsh creeks) in the Northern lagoon basin. Sampling was carried out seasonally (Spring, Summer and Autumn of 2002) in 4–7 stations for each habitat type, by means of a fine-mesh, small beach seine. Two-way analysis of variance was applied to assess the differences in species richness, fish diversity, density and standing stock amongst habitats, whereas fish assemblage composition was investigated by using multivariate analyses (MDS, ANOSIM, SIMPER). The analyses indicated that seagrass beds and saltmarsh creeks are relevant shallow habitats in structuring the small-sized fish assemblages of the Venice Lagoon, supporting specialized and recognizable fish assemblages. Those in seagrass beds, in particular, were characterized by higher species richness and standing stock with respect to all the others. The structuring role of these habitats was discussed in terms of both habitat complexity and degree of confinement. In contrast, sandy bottoms, mudflats and sparsely vegetated habitats were identified as “transition” habitats, with highly variable fish assemblages, influenced by the contribution of the adjacent habitats, and acting probably as both ‘buffer zones’ between the other habitats and migration routes for many fish species in the lagoon.     

Seagrass beds and intertidal invertebrates: an experimental test of the role of habitat structure

The majority of field experiments have been carried out on relatively small spatial and short temporal scales, but some of the most interesting ecological processes operate at much larger scales. However, large-scale experiments appropriate to the landscape, often have to be carried out with minimal plot replication and hence reduced statistical power. Here, we report the results of such a large scale, un-replicated field experiment on the Mondego estuary, Portugal, which nevertheless provides compelling evidence of the importance of habitat structure for invertebrate community composition and dynamics. In this estuary, seagrass beds have suffered a dramatic decline over the last 20 years, associated with changes in invertebrate assemblages. In addition, the most abundant species in the system, Hydrobia ulvae, displays distinctly different population structures in those sites. The aim of the field experiment was to test the hypothesis that these differences are related to enhanced survival of snails due to protection from avian or fish predators so that they can grow to larger body sizes in the more complex habitat provided by seagrass. We tested this hypothesis through a large-scale experiment using artificial seagrass beds over a 12-month period. Adult snail densities were higher in the artificial bed plots compared to controls. However, these differences emerged only slowly, related to snail growth rate. This suggests that protection from epibenthic predators can have a significant effect on population structure and hence biomass and productivity of key species in this system. However, the invertebrate assemblage in artificial seagrass plots and the natural seagrass bed, remained statistically separate by the end of the experiment. Handling editor: K. Martens     

Juvenile sparids (Rhabdosargus holubi) consistently select structurally dense vegetated habitat in nursery seascapes

Seagrass habitats provide structural complexity in coastal estuarine and marine environments, which offer fish optimal foraging grounds and refuge from predation. However, seagrasses are some of the most threatened ecosystems globally, with anthropogenic activities such as population growth and environmental degradation leading to the fragmentation, thinning, and loss of these habitats. Rhabdosargus holubi is one of only a few vegetation-associated marine fish species in South African estuaries. Although field studies have shown a strong association with seagrass over other aquatic vegetation for the juveniles of this species, habitat choice has never been empirically tested. Here, we used artificial vegetation units to test habitat choice (different structural complexities) for this species. We also tested whether habitat choice is influenced by a predatory threat, with fish preferentially selecting dense habitat in the presence of a predator and whether this effect may be more apparent in smaller individuals. We found that R. holubi significantly prefer greater structural complexity over less complex habitats, in both the absence and presence of a predator and for both small and large juveniles, showing that R. holubi actively choose more complex structures and are attracted to the structure per se irrespective of the threat of predation. This study highlights the importance of dense seagrass as nursery areas for this species and demonstrates how the loss of these habitats could impact the nursery function of estuaries.     

Mangrove Habitat Use by Juvenile Reef Fish: Meta-Analysis Reveals that Tidal Regime Matters More than Biogeographic Region

Identification of critical life-stage habitats is key to successful conservation efforts. Juveniles of some species show great flexibility in habitat use while other species rely heavily on a restricted number of juvenile habitats for protection and food. Considering the rapid degradation of coastal marine habitats worldwide, it is important to evaluate which species are more susceptible to loss of juvenile nursery habitats and how this differs across large biogeographic regions. Here we used a meta-analysis approach to investigate habitat use by juvenile reef fish species in tropical coastal ecosystems across the globe. Densities of juvenile fish species were compared among mangrove, seagrass and coral reef habitats. In the Caribbean, the majority of species showed significantly higher juvenile densities in mangroves as compared to seagrass beds and coral reefs, while for the Indo-Pacific region seagrass beds harbored the highest overall densities. Further analysis indicated that differences in tidal amplitude, irrespective of biogeographic region, appeared to be the major driver for this phenomenon. In addition, juvenile reef fish use of mangroves increased with increasing water salinity. In the Caribbean, species of specific families (e.g. Lutjanidae, Haemulidae) showed a higher reliance on mangroves or seagrass beds as juvenile habitats than other species, whereas in the Indo-Pacific family-specific trends of juvenile habitat utilization were less apparent. The findings of this study highlight the importance of incorporating region-specific tidal inundation regimes into marine spatial conservation planning and ecosystem based management. Furthermore, the significant role of water salinity and tidal access as drivers of mangrove fish habitat use implies that changes in seawater level and rainfall due to climate change may have important effects on how juvenile reef fish use nearshore seascapes in the future.     

The Mangrove Nursery Paradigm Revisited: Otolith Stable Isotopes Support Nursery-to-Reef Movements by Indo-Pacific Fishes

Mangroves and seagrass beds have long been perceived as important nurseries for many fish species. While there is growing evidence from the Western Atlantic that mangrove habitats are intricately connected to coral reefs through ontogenetic fish migrations, there is an ongoing debate of the value of these coastal ecosystems in the Indo-Pacific. The present study used natural tags, viz. otolith stable carbon and oxygen isotopes, to investigate for the first time the degree to which multiple tropical juvenile habitats subsidize coral reef fish populations in the Indo Pacific (Tanzania). Otoliths of three reef fish species (Lethrinus harak, L. lentjan and Lutjanus fulviflamma) were collected in mangrove, seagrass and coral reef habitats and analyzed for stable isotope ratios in the juvenile and adult otolith zones. δ¹³C signatures were significantly depleted in the juvenile compared to the adult zones, indicative of different habitat use through ontogeny. Maximum likelihood analysis identified that 82% of adult reef L. harak had resided in either mangrove (29%) or seagrass (53%) or reef (18%) habitats as juveniles. Of adult L. fulviflamma caught from offshore reefs, 99% had passed through mangroves habitats as juveniles. In contrast, L. lentjan adults originated predominantly from coral reefs (65–72%) as opposed to inshore vegetated habitats (28–35%). This study presents conclusive evidence for a nursery role of Indo-Pacific mangrove habitats for reef fish populations. It shows that intertidal habitats that are only temporarily available can form an important juvenile habitat for some species, and that reef fish populations are often replenished by multiple coastal habitats. Maintaining connectivity between inshore vegetated habitats and coral reefs, and conserving habitat mosaics rather than single nursery habitats, is a major priority for the sustainability of various Indo Pacific fish populations.     

Contrasting effects of habitat structure on the recruitment and mortality of an epibiotic macroalga

Field surveys over 2 years in contiguous beds of the seagrassesZostera capricorni andPosidonia australis showed that the green algaCodium duthieae was consistently more abundant inZ. capricorni than inP. australis. In 1 year, mature plants were also more abundant at the boundary between the seagrass beds than in either bed. Field experiments and programmes of sampling were used to investigate three potential explanations for the unusual distribution of this alga: (1) that the availability of substrata suitable for attachment of the alga differed between the two seagrass beds; (2) that mortality of matureC. duthieae differed between the seagrass beds; and/or (3) that the intensity of recruitment was different in the two seagrass beds.C. duthieae plants were exclusively epibionts of the bivalveAnadara trapezia. Detailed sampling showed that the abundance ofA. trapezia was similar in both seagrass beds and that the distribution of bivalves suitable as substrata forC. duthieae plants was not obviously related to proximity to the boundary between the beds. Two experiments investigated the survival ofC. duthieae plants in each bed. In the first, matureC. duthieae plants transplanted into theP. australis bed suffered similar rates of mortality to plants which were disturbed and moved within theZ. capricorni bed or which were left undisturbed in theZ. capricorni bed. Fewer of the host bivalves were recovered from theZ. capricorni bed, however, indicating that the mechanism of mortality differed between the beds, hosts being more frequently dislodged in theZ. capricorni bed. Removal of the leaves of the seagrasses had consistently greater effects on near-bottom current velocities in theZ. capricorni bed than in theP. australis bed and significantly increased mortality ofC. duthieae in theZ. capricorni bed. Survival of plants was greater in plots of artificial leaves ofP. australis placed in theZ. capricorni bed than in plots of the naturalZ. capricorni leaves or plots where the natural leaves were removed. Most mortality in theZ. capricorni bed was due to dislodgement of the alga and its bivalve substratum. Corresponding manipulations of leaves in theP. australis bed had consistently smaller effects on survivorship of both the alga and its host. Patterns in the recruitment of the alga most clearly reflected the distribution of adults.C. duthieae recruits were 5 times more abundant in theZ. capricorni bed and at the boundary between the two beds than in theP. australis bed. The results demonstrate how habitat structure, provided by the canopy of leaves of the two species of seagrass, can have contrasting effects on the recruitment and mortality of a macroalga. In the case ofC. duthieae, it appears that the differential pattern of recruitment is the primary determinant of the distribution of adult plants.     

Habitat linkages: the effect of intertidal saltmarshes and adjacent subtidal habitats on abundance, movement, and growth of an estuarine fish

In this study we used pinfish (Lagodon rhomboides) in field experiments to examine linkages between intertidal saltmarsh and adjacent subtidal habitats. Pinfish are more than twice as abundant in intertidal marshes adjacent to seagrass beds than in those adjacent to the unvegetated subtidal bottom. Movement of pinfish between the marsh edge and the adjacent subtidal habitat was greater for fish captured in areas with both intertidal and subtidal vegetation than in those with intertidal vegetation and adjacent unvegetated mudflats. This movement provides an important link between habitats, allowing transfer of marsh-derived secondary production to subtidal seagrass beds and vice versa. Pinfish held in enclosures with both intertidal and subtidal vegetation were, on average, approximately 90% heavier than fish held in enclosures with intertidal vegetation and unvegetated subtidal bottom. Because saltmarshes and seagrass beds contribute to the production of living marine resources, active measures are being taken to preserve and restore these habitats. The results from this study have direct application to decisions concerning site selection and optimal spatial proximity of saltmarsh and seagrass habitats in the planning of restoration and mitigation projects. To maximize secondary production and utilization of intertidal marshes, managers may opt to restore and/or preserve marshes adjacent to subtidal seagrass beds. Received: 31 May 1996 / Accepted: 23 October 1996     

Seascape structure and complexity influence temperate seagrass fish assemblage composition

Understanding how spatial patterning relates to ecological processes is fundamental to define important species–environment associations at broader scales. Analyses targeting habitat structure (i.e. composition and configuration) in terrestrial landscapes are increasing, but similar studies in marine landscapes are still relatively uncommon. In this study, we explored how seascape structure and complexity (determined from significant spatial pattern metrics) influenced summer and autumn fish assemblage composition in 30 seagrass (Zostera marina) meadows along the west coast of Sweden. Species density was not influenced by seascape structure in any season. In contrast, the majority of significant fish assemblage variables were influenced by seascape structure during the summer (i.e. abundance and proportion of juveniles, abundance of Labridae and abundance of occasional shallow‐water visitors) whilst fewer in the autumn (i.e. abundance of occasional shallow‐water visitors and Synganthidae). For instance, less complex seascapes were more suitable for juvenile assemblages in summer, as these seascapes exhibit larger patch sizes of appropriate habitat (e.g. Z. marina) and less edge boundaries providing refuges from predators and food resources. Abundances of migrating fish, such as the sea trout Salmo trutta, also responded positively to a less complex seascape in the summer though perhaps ecological processes, such as prey availability, were additional contributing factors driving this relationship. High complexity seascapes only had a positive influence on the abundance of taxa using multiple habitats (Labridae during the summer). Our study shows that fish assemblages in temperate marine environments are significantly linked to spatial habitat patterning and seascape complexity. This offers valuable insights into species–habitat–seascape linkages, information important for coastal conservation and marine spatial planning.