Jeremy S. Hindell

Fish responses to experimental fragmentation of seagrass habitat.

Understanding the consequences of habitat fragmentation has come mostly from comparisons of patchy and continuous habitats. Because fragmentation is a process, it is most accurately studied by actively fragmenting large patches into multiple smaller patches. We fragmented artificial seagrass habitats and evaluated the impacts of fragmentation on fish abundance and species richness over time (1 day, 1 week, 1 month). Fish assemblages were compared among 4 treatments: control (single, continuous 9-m(2) patches); fragmented (single, continuous 9-m(2) patches fragmented to 4 discrete 1-m(2) patches); prefragmented/patchy (4 discrete 1-m(2) patches with the same arrangement as fragmented); and disturbance control (fragmented then immediately restored to continuous 9-m(2) patches). Patchy seagrass had lower species richness than actively fragmented seagrass (up to 39% fewer species after 1 week), but species richness in fragmented treatments was similar to controls. Total fish abundance did not vary among treatments and therefore was unaffected by fragmentation, patchiness, or disturbance caused during fragmentation. Patterns in species richness and abundance were consistent 1 day, 1 week, and 1 month after fragmentation. The expected decrease in fish abundance from reduced total seagrass area in fragmented and patchy seagrass appeared to be offset by greater fish density per unit area of seagrass. If fish prefer to live at edges, then the effects of seagrass habitat loss on fish abundance may have been offset by the increase (25%) in seagrass perimeter in fragmented and patchy treatments. Possibly there is some threshold of seagrass patch connectivity below which fish abundances cannot be maintained. The immediate responses of fish to experimental habitat fragmentation provided insights beyond those possible from comparisons of continuous and historically patchy habitat.

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.

Resource distribution influences positive edge effects in a seagrass fish

According to conceptual models, the distribution of resources plays a critical role in determining how organisms distribute themselves near habitat edges. These models are frequently used to achieve a mechanistic understanding of edge effects, but because they are based predominantly on correlative studies, there is need for a demonstration of causality, which is best done through experimentation. Using artificial seagrass habitat as an experimental system, we determined a likely mechanism underpinning edge effects in a seagrass fish. To test for edge effects, we measured fish abundance at edges (0-0.5 m) and interiors (0.5-1 m) of two patch configurations: continuous (single, continuous 9-m2 patches) and patchy (four discrete 1-m2 patches within a 9-m2 area). In continuous configurations, pipefish (Stigmatopora argus) were three times more abundant at edges than interiors (positive edge effect), but in patchy configurations there was no difference. The lack of edge effect in patchy configurations might be because patchy seagrass consisted entirely of edge habitat. We then used two approaches to test whether observed edge effects in continuous configurations were caused by increased availability of food at edges. First, we estimated the abundance of the major prey of pipefish, small crustaceans, across continuous seagrass configurations. Crustacean abundances were highest at seagrass edges, where they were 16% greater than in patch interiors. Second, we supplemented interiors of continuous treatment patches with live crustaceans, while control patches were supplemented with seawater. After five hours of supplementation, numbers of pipefish were similar between edges and interiors of treatment patches, while the strong edge effects were maintained in controls. This indicated that fish were moving from patch edges to interiors in response to food supplementation. These approaches strongly suggest that a numerically dominant fish species is more abundant at seagrass edges due to greater food availability, and provide experimental support for the resource distribution model as an explanation for edge effects.

Seagrass patch size affects fish responses to edges.

1. Patch area and proximity of patch edge can influence ecological processes across patchy landscapes and may interact with each other. Different patch sizes have different amounts of core habitat, potentially affecting animal abundances at the edge and middle of patches. In this study, we tested if edge effects varied with patch size. 2. Fish were sampled in 10 various-sized seagrass patches (114-5934 m(2)) using a small (0.5 m(2)) push net in three positions within each patch: the seagrass edge, 2 m into a patch and in the middle of a patch. 3. The two most common species showed an interaction between patch size and the edge-interior difference in abundance. In the smallest patches, pipefish (Stigmatopora nigra) were at similar densities at the edge and interior, but with increasing patch size, the density at the edge habitat increased. For gobies (Nesogobius maccullochi), the pattern was exactly the opposite. 4. This is the first example from a marine system of how patch size can influence the magnitude and pattern of edge effects. 5. Both patch area and edge effects need to be considered in the development of conservation and management strategies for seagrass habitats.

Assessing the trophic link between seagrass habitats and piscivorous fishes

Links between piscivorous fishes and seagrass habitat were investigated in Port Phillip Bay, Australia. Abundances of piscivorous fish were estimated, the contribution of fish to their diets was measured and the trophic link between these fish and seagrass beds was assessed with stable isotopes. Piscivorous fishes were common in seagrass and included eight species from seven families (33% by abundance). They consumed at least eight families of teleost prey, including juveniles of seagrass-associated taxa, such as monacanthids (e.g. Meuschenia freycineti) and syngnathids (Stigmatopora argus). No fish were exclusively piscivorous and contribution of fish to the diets of the most common species (Arripis truttacea and Kestratherina esox) varied strongly through time. Putative contributions of each source (primary producer – plant) to the nutrition of piscivorous fishes were: (1) fishes whose base for nutritional support was driven mostly (>50%) by seagrass (e.g. Platycephalus speculator and Platycephalus laevigatus); and (2) fishes whose base for nutritional support was not driven by any particular primary producer (e.g. Arripis truttacea and Pseudocaranx dentex). The propensity for piscivorous fish to include seagrass-associated fish in their diets, their perennial presence in seagrass and the strong putative contribution by seagrass to their nutrition, suggest that seagrass habitats can be valuable habitat for piscivorous fishes.

Variability in the numbers of post-settlement King George whiting (Sillaginidae: Sillaginodes punctata, Cuvier) in relation to predation, habitat complexity and artificial cage structure

The importance of predation by fish in altering abundances of juvenile King George whiting (Sillaginodes punctata) was examined at multiple locations in Port Phillip Bay, Australia, by manipulating the numbers of piscivorous fish in unvegetated sand and seagrass habitats using cages. Additional information regarding the local abundances of, and habitat use by, the most common piscivorous fish, Western Australian salmon (Arripidae: Arripis truttacea, Cuvier), was gathered using netting surveys and underwater video. Regardless of habitat, abundances of S. punctata were similar in partial cages and uncaged areas. In unvegetated sand, S. punctata were more abundant inside cages than partial cages or uncaged areas. In seagrass, there was no difference in the numbers of S. punctata between caging treatments. Patterns in abundances of S. punctata between cage treatments in each habitat were consistent between sites, but the relative difference in the abundances of S. punctata between habitats was site specific. Abundances of A. truttacea varied significantly between sites, and they consumed a variety of epibenthic fishes including atherinids, clupeids, gobiids, syngnathids and pleuronectids. At one site in Port Phillip Bay (Blairgowrie), A. truttacea occurred more commonly in patches of unvegetated sand than seagrass. Over unvegetated sand, abundances of A. truttacea varied little between partial cages and uncaged areas. The numbers of S. punctata varied between caging treatments and habitats in a manner that was consistent with a model whereby seagrass interferes with foraging by predatory fish and provides juvenile fish with a refuge from predation. The almost total absence of A. truttacea in seagrass habitats and the lack of S. punctata in their diets implies, however, that patterns in S. punctata in seagrass/unvegetated sand mosaics are driven by processes other than direct predation.

Influence of freshwater flows on the distribution of eggs and larvae of black bream Acanthopagrus butcheri within a drought-affected estuary

This study tested the hypothesis that variable freshwater flow in the Gippsland Lakes, Australia, influences the location and extent of environmental conditions suitable for spawning and larval development of black bream, Acanthopagrus butcheri. Freshwater flow had a large influence on the salinity and level of stratification (difference between bottom and surface salinity) in the Gippsland Lakes. Freshwater flows throughout this study varied from no or low flows through to major flooding in 2007. Eggs of A. butcheri were found in similar concentrations throughout the lakes and rivers. More than 99% of larvae, however, were collected from within rivers, with very few larvae being collected from the lakes. A comparison of two spawning seasons revealed that the year with higher freshwater flows also had greater spawning activity and higher concentrations of larvae. Interestingly, there was a significant relationship between the distribution of eggs and larvae with the level of stratification. The highest concentrations of larvae occurred at sites with a difference in bottom and surface salinities of 15-20. This study demonstrates that despite A. butcheri spawning in the lakes and rivers, it is only locations with a halocline that function as larval nursery habitat.

Edge patterns in aquatic invertebrates explained by predictive models

Predictive frameworks for understanding and describing how animals respond to habitat fragmentation, particularly across edges, have been largely restricted to terrestrial systems. Abundances of zooplankton and meiofauna were measured across seagrass-sand edges and the patterns compared with predictive models of edge effects. Artificial seagrass patches were placed on bare sand, and zooplankton and meiofauna were sampled with tube traps at five positions (from patch edges: 12, 60 and 130 cm into seagrass; and 12 and 60 cm onto sand). Position effects consisted of the following three general patterns: (1) increases in abundance around the seagrass-sand edge (total abundance and cumaceans); (2) declining abundance from seagrass onto sand (calanoid copepods, harpacticoid copepods and amphipods); and (3) increasing abundance from seagrass onto sand (crustacean nauplii and bivalve larvae). The first two patterns are consistent with resource-distribution models, either as higher resources at the confluence of adjacent habitats or supplementation of resources from high-quality to low-quality habitat. The third pattern is consistent with reductions in zooplankton abundance as a consequence of predation or attenuation of currents by seagrass. The results show that predictive models of edge effects can apply to aquatic animals and that edges are important in structuring zooplankton and meiofauna assemblages in seagrass.

Genetic stock structure of blue-eye trevalla (Hyperoglyphe antarctica) and warehous (Seriolella brama and Seriolella punctata) in south-eastern Australian waters.

Blue-eye trevalla (Hyperoglyphe antarctica), blue warehou (Seriolella brama) and silver warehou (Seriolella punctata) from the family Centrolophidae are three commercially important species in the Australian fishery. These species are currently managed as single stocks. We tested the hypothesis that patterns of phenotypic structuring in these species reflect underlying genetic stock structure using an analysis of mitochondrial DNA control region sequences. The analysis revealed high levels of haplotype diversity within populations. The most common haplotypes for the species occurred in all geographical locations sampled. For S. brama, although structuring was not significant after Bonferroni correction, differences between two sites were sufficient to warrant caution in the management of fishery zones for this species. There were also some indications of structuring when sites were grouped into common regions. Demographic analysis suggested that S. brama might have had a history of population bottlenecks followed by sudden population expansion, potentially contributing to genetic structuring in the fishery. No structuring was detected for H. antarctica and S. punctata. The present study highlights the need for, and the utility of, multiple sources of information, that is, genetic, phenotypic, behavioural and ecological, when managing marine fisheries and the need to take a cautionary approach to the interpretation of genetic data for fisheries management.

Fine-scale spatial and temporal variations in diets of the pipefish Stigmatopora nigra within seagrass patches

Diets of the pipefish Stigmatopora nigra were analysed to determine if food availability was causing S. nigra to distribute according to habitat edge effects. Gut analysis found little difference in the diets of S. nigra at the edge and interior of seagrass patches, regardless of time of day or season. Fish diets did, however, vary with seagrass density: S. nigra in denser seagrass consumed more harpacticoid copepods and fewer planktonic copepods. The lack of difference in prey eaten by S. nigra at the edge and interior of patches suggests either that food was not determining S. nigra distribution patterns within patches or that differences in fish densities across patches meant that relative fish-prey densities were similar at edge and interior positions. Alternatively, any edge effects in diet might be masked by gradients in seagrass structure.