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Light availability and habitat complexity are two key drivers of community assembly. Urbanisation has been shown to affect both, with important consequences to ecological communities. On the intertidal, for instance, studies have shown that light intensity is greater on natural rocky shores than on less complex artificial habitats (seawalls), though different habitats can also experience similar light intensities, for example when shaded by urban structures. Understanding therefore how these factors individually, and combined, affect communities is important to understand the mechanisms driving changes in community structure, and consequently provide solutions to tackle the increasing homogenisation of habitats and lightscapes in urbanised spaces through smart infrastructure designs. Here, we assessed how different light levels affect the recruitment of communities in rock pools and on emergent rock on an intertidal rocky shore. We cleared 30 patches of emergent rock and 30 rock pools and manipulated light using shades with different light transmissions (full light, procedural control, 75%, 35%, and 15% light transmission, full shade) and assessed mobile and sessile communities monthly for 6 months. Effects of reducing light levels were generally stronger on rock than in pools. Fully shaded plots supported double the amount of mobile organisms than plots in full sunlight, in both habitats. Algal cover was higher in pools compared to rock, and at intermediate light levels, but effects varied with site. This study highlights the importance of variable light conditions and different habitats for rocky shore communities, which should be considered in future coastal developments to retain natural biodiversity. [ABSTRACT FROM AUTHOR]

The co-occurrence of several species, all of which share similar resource requirements, remains a paradox in ecology. Here, I evaluated resource use along multiple environmental axes to understand the potential for niche partitioning and complementarity in a guild of suspension-feeding rocky shore invertebrates. I focused on the mussels Perna canaliculus, Mytilus galloprovincialis, Aulacomya maoriana, and Xenostrobus pulex, all of which coexist along the coastline of the South Island of New Zealand. I quantified the mussels' distributions, both vertically on the shore and within the three-dimensional mussel bed matrix, and used carbon (δ13C) and nitrogen (δ15N) stable isotope ratios to compare species' diets. Mussels exhibited niche partitioning along all resource axes. Given the mussels' use of different spatial and food resources, I evaluated the potential for complementarity with respect to mussels' roles as mediators of an important spatial subsidy, carbon inputs from the nearshore ocean into rocky-shore ecosystems. In these systems, mussels are basal species, capturing and consuming particulate organic matter in the ocean and making it available for local consumption within the benthic community. I found that mussel diversity matters; even the most productive species—Perna canaliculus—only contributed about half of the mussel-mediated carbon that accumulated over a year. Multiple co-occurring species are, therefore, likely to provide more carbon at the base of the intertidal food web than any one species living independently, and complementarity in resource use along multiple environmental axes is likely an important mechanism underlying this relationship. [ABSTRACT FROM AUTHOR]

The effective use of ecosystem engineers to conserve biodiversity requires an understanding of the types of resources an engineer modifies, and how these modifications vary with biotic and abiotic context. In the intertidal zone, oysters engineer ecological communities by reducing temperature and desiccation stress, enhancing the availability of hard substrate for attachment, and by ameliorating biological interactions such as competition and predation. Using a field experiment manipulating shading, predator access and availability of shell substrate at four sites distributed over 900 km of east Australian coastline, we investigated how the relative importance of these mechanisms of facilitation vary spatially. At all sites, and irrespective of environmental conditions, the provision of hard substrate by oysters enhanced the abundance and richness of invertebrates, in particular epibionts (barnacles and oyster spat) and grazing gastropods. Mobile arthropods utilised the habitat provided by disarticulated dead oysters more than live oyster habitat, whereas the abundance of polychaetes and bivalves were much greater in live oysters, suggesting the oyster filter-feeding activity is important for these groups. In warmer estuaries, shading by oysters had a larger effect on biodiversity, whereas in cooler estuaries, the provision of a predation refuge by oysters played a more important role. Such knowledge of how ecosystem engineering effects vary across environmental gradients can help inform management strategies targeting ecosystem resilience via the amelioration of specific environmental stressors, or conservation of specific community assemblages. [ABSTRACT FROM AUTHOR]

Thermal performance curves (TPCs) represent an increasingly popular tool in ecology for anticipating species responses to climate change. TPC theory has been developed using species that experience similar temperatures during activity and at rest and consequently exhibit thermal ranges for activity that closely coincide with their physiological thermal tolerances. Many species, however, experience other stressors, such as desiccation, that limit active behaviour at temperatures below the maximum values experienced. As a result, activity is constrained to a narrow thermal window that is a subset of the range of temperatures that can be tolerated physiologically. This results in a decoupling of behavioural and physiological TPCs that does not conform to the present paradigm. To test the generality of TPC theory, we measured thermal responses for behaviour (crawling speed) and physiological tolerance (heart rate) for six rocky shore gastropods spanning a thermal/desiccation stress gradient. We hypothesized a positive relationship between shore level and the degree of decoupling of behavioural and physiological TPCs. This prediction was confirmed, and was explained by the extension of the physiological TPC beyond the range of the behavioural TPC. Decoupling of behavioural and physiological TPCs is central to predicting accurately the fitness dynamics of ectothermic species subject to multiple stressors. We believe that this decoupling should be explicitly considered as an adaptive trait defining an organism's thermal niche. [ABSTRACT FROM AUTHOR]

Repeated, spatially explicit sampling is widely used to characterize the dynamics of sessile communities in both terrestrial and aquatic systems, yet our understanding of the consequences of errors made in such sampling is limited. In particular, when Markov transition probabilities are calculated by tracking individual points over time, misidentification of the same spatial locations will result in biased estimates of transition probabilities, successional rates, and community trajectories. Nonetheless, to date, all published studies that use such data have implicitly assumed that resampling occurs without error when making estimates of transition rates. Here, we develop and test a straightforward maximum likelihood approach, based on simple field estimates of resampling errors, to arrive at corrected estimates of transition rates between species in a rocky intertidal community. We compare community Markov models based on raw and corrected transition estimates using data from Endocladia muricata-dominated plots in a California intertidal assemblage, finding that uncorrected predictions of succession consistently overestimate recovery time. We tested the precision and accuracy of the approach using simulated datasets and found good performance of our estimation method over a range of realistic sample sizes and error rates. [ABSTRACT FROM AUTHOR]

Habitat-forming species increase spatial complexity and alter local environmental conditions, often facilitating a diversified assemblage of plants and animals. Removal of dominant species, therefore, can potentially lead to pronounced changes in diversity and community structure through a series of negative and positive interactions involving several components of the community. Here we test community responses to the deletion of the dominant, canopy-forming alga Hormosira banksii from the mid-intertidal zone of wave-protected rocky shores in southern New Zealand. This species was removed in winter (July) from three 3×3-m areas at each of two platforms (Kaikoura and Moeraki) on the east coast of the South Island. Initially, 59 taxa occurred in stands, but there were only four algal species with greater than 5% cover and three mobile invertebrate species with more than five individuals per 0.25 m2. By 6 months after Hormosira removal, most fucoid and coralline algae had burned off, and there were blooms of ephemeral algae in the removal plots, but almost no change within controls. After 2 years, diversity declined by 44% relative to controls at Kaikoura and 36% at Moeraki, and the amount of bare space had increased by tenfold at Kaikoura and twofold at Moeraki. Few sessile or mobile invertebrates were present. Recruitment of Hormosira occurred after 14 months in the removal plots. At this time, a “press” disturbance was initiated into one half of each removal plot to test the effects of continued removal of Hormosira on diversity. Similar “end-points” of the control and “press” removal plots were not reached after 2 years, and even after Hormosira recruitment into the original “pulse” experiment there was little recovery of the community. In this mid-intertidal system with considerable thermal stress, and perhaps in others with few perennial species, diversity and community structure can critically depend on positive associations with a single dominant species. [ABSTRACT FROM AUTHOR]

Experimental studies of biogeographic processes are important, but rarely attempted because of the logistical challenges of research at large spatial scales. I used a series of large-scale transplant experiments to investigate the mechanisms controlling species abundance near a poleward range boundary. The intertidal limpet Collisella scabra experiences a 100-fold decline in abundance over the northernmost 300 km of its range. Temperature and food supply both strongly influenced individual survival, growth, and maturation. Regression analysis also revealed significant interactions among these conditions: the effect of one could not be predicted without knowing the level of the other. But these relationships could not explain geographic abundance patterns. Instead, individual limpets were highly successful at sites with relatively low abundance. These results suggest that, even though temperature is important to the success of individual C. scabra populations, the primary effect of warming temperatures under climate change may not be a shift in geographic distribution. [ABSTRACT FROM AUTHOR]

The paradigmatic gradient for intertidal marine organisms of increasing physical stress from low to high elevation has long served as the basis for using direct effects of duration of water coverage to predict many biological patterns. Accordingly, changes in potential feeding time may predict the direction and magnitude of differences between elevations in individual growth rates of sessile marine invertebrates. Oysters (triploid Crassostrea ariakensis) experimentally introduced at intertidal (MLW+0.05 m) and subtidal (MLW–0.25 m) elevations in racks provided a test of the ability to use duration of water coverage to predict changes in growth. During early-to-mid winter, a depression of 38–47% in shell growth of intertidal oysters matched the 36% reduction in available feeding time relative to subtidal oysters. In late winter as solar heating of exposed oysters increased, growth differences of 52–55% departed only slightly from the predicted 39%. In spring, however, duration of water coverage failed to predict even the correct direction of growth change with elevation as intertidal oysters grew 34% faster despite 39% less feeding time. Intense seasonal development of shell fouling by other suspension feeders like ascidians, mussels, and barnacles on subtidal (94% incidence) but not on aerially exposed intertidal (21–38% incidence) oysters may explain why duration of water cover failed to predict spring growth differences. Less intense fouling develops on intertidal oysters due to the physiological stress of aerial exposure on settlers, especially during higher temperatures and longer solar exposures of spring. Fouling by suspension feeders is known to reduce growth of the host through localized competition for food and added energetic costs. Thus, in springtime, indirect effects of aerial exposure providing a partial refuge from biological enemies overwhelmed direct effects of reduced duration of water coverage to reverse the expected pattern of slower intertidal growth of a marine invertebrate. [ABSTRACT FROM AUTHOR]

We describe differences in life history of the intertidal arboreal snail, Littoraria filosa, among patches of mangroves ranging in size from isolated trees to large stands several square kilometres in area. Recruitment of L. filosa occurred from mid spring (October) to early winter (June), recruits grew rapidly and copulating adults were found during the following September–April. Populations within large patches of forest were annuals; all or most individuals died between October–January (spring–midsummer). In contrast, those in smaller peripheral patches were more likely to survive over the summer but survival differed among patches and years. These differences in life history were caused by a parasitoid fly (genus Sarcophaga) that attacked L. filosa 10 mm and longer and was present in all large patches, but absent from, or rare, in smaller peripheral patches. Experimental introductions to isolated trees confirmed that the fly could kill L. filosa. Another sarcophagid parasitoid that attacked L. filosa from 4 to less than 10 mm long was also found in every patch. The combined effects of these parasitoids appear to determine the metapopulation structure of L. filosa. Most adults in large patches were killed by the larger fly during early summer. Summer recruits were often killed by the smaller fly within a month of settlement and when this happened effective recruitment of L. filosa was reduced to autumn. The planktotrophic larval stage of L. filosa lasts less than 1 month, so the source of autumn recruits to all patches must have been adults that survived the early summer, most of which were in small patches or on isolated trees. Consequently these ”peripheral sources” are likely to be important for persistence of the metapopulation of L. filosa. The results of this study demonstrate that metapopulation structure may be determined by complex interactions and that common models cannot be assumed to apply in all habitats. [ABSTRACT FROM AUTHOR]

Holling’s (1992) proposition that discontinuities in biotic and abiotic processes generate structure in ecological systems is examined experimentally by imposing size-specific perturbations on marine sediment assemblages. Two kinds of perturbations were applied: organic enrichment and predation, each at two levels. Perturbations significantly affected the densities and relative abundance of the main invertebrate taxa and these effects were consistent with the known effects of enrichment and predation. However, there was little evidence of significant treatment effects on the overall benthic biomass or abundance size spectrum, supporting the contention that the spectrum is conservative and is probably constrained by habitat architecture. [ABSTRACT FROM AUTHOR]

Few comparative studies of spatial patterns at different scales have examined several species in the same habitat or the same species over a range of habitats. Therefore, variability in patterns among species or among habitats has seldom been documented. This study quantifies spatial patterns of a suite of intertidal snails and a species of barnacle using a range of statistical techniques. Variability in densities was quantified from the scale of adjacent quadrats (over a distance of centimeters) to tens of kilometers. Significant differences in abundances occurred primarily at two spatial scales. Small-scale differences were found at the scales of centimeters or 1-2 m and, for many species on many shores, these accounted for most of the variability in abundances from place to place. These are likely to be determined by behavioural responses to small-scale patches of microhabitat. Large-scale differences in abundance were also found in most species at the scale of hundreds of meters alongshore. These are likely to be due to variation in recruitment (and/or mortality) because of limited dispersal by adults of these species. There was little or no additional variation among shores, separated by tens of kilometers, than was shown among patches of shore separated by hundreds of meters. Identification of the scale(s) at which significant differences in abundance are found focus attention on the processes (and the scales at which these processes operate) that influence patterns of distribution and abundance. Some of the advantages and disadvantages of various procedures are discussed. [ABSTRACT FROM AUTHOR]

The structure of rocky intertidal communities may be influenced by large-scale patterns of productivity. In this study we examine the in situ rates of production by intertidal epilithic microalgae (chlorophyll a production per unit area per month), intertidal nutrient concentrations (nitrates, nitrites, phosphates and silicates), and standing stocks of different functional-form groups of macroalgae around the South African coast, and their relationships to consumer biomass. Clear gradients of in situ intertidal primary production and nutrient concentrations were recorded around the South African coast, values being highest on the west coast, intermediate on the south and lowest on the east coast. Primary production by intertidal epilithic microalgae was correlated with nutrient availability and could also be related to nearshore phytoplankton production. The dominance patterns of different functional forms of macroalgae changed around the coast, with foliose algae prevalent on the west coast and coralline algae on the east coast. However, overall macroalgal standing stocks did not reflect the productivity gradient, being equally high on the east and west coasts, and low in the south. Positive relationships existed between the average biomass of intertidal intertebrate consumers (grazers and filter-feeders) and intertidal productivity, although only the grazers were directly 'connected' to in situ production by epilithic intertidal microalgae. The maximum body size of a widely distributed limpet, Patella granularis, was also positively correlated with level of in situ primary production. The maximal values of biomass attained by intertidal filter-feeders were not related to intertidal primary production, and were relatively constant around the coast. At a local scale, filter-feeder biomass is known to be strongly influenced by wave action. This implies that the local-scale water movements over-ride any effects that large-scale gradients of primary production may have on filter-feeders. The large-scale gradient in intertidal productivity around the coast is thus strongly linked with grazer biomass and individual body size, but any effect it has on filter-feeder biomass seems subsidiary to the local effects of wave action. [ABSTRACT FROM AUTHOR]

Recent theoretical models have considered how nutritional status and predation risk affect the decision by an individual to forage. We consider the feeding behaviour of a scavenger under risk of predation. The intertidal gastropod Nassarius dorsatus Röding was used to test the following hypotheses: (1) a damaged conspecific should be a less preferred meal than a damaged predator or another species; (2) a scavenger should be able to discriminate between a damaged conspecific in the presence and one in the absence of its predator, preferring the latter; and (3) the decision by a scavenger to feed should reflect a trade-off between nutritional status and the above preferences. Results from the laboratory and field were consistent with predictions. Two hypotheses are suggested from the work: scavengers may generally be less willing to feed on carcases of conspecifics than on those of other species, and intertidal scavengers may commonly face a relative or absolute shortage of food. [ABSTRACT FROM AUTHOR]

Mutualistic relations between plants and animals are well documented on land but have received less attention in marine systems. This study examined the relationship between the territorial intertidal limpet Patella longicosta and the crustose brown alga Ralfsia verrucosa. Adult Patella are found exclusively in association with Ralfsia, on which they feed, while Ralfsia occurs primarily, but not exclusively, in Patella territories. Ralfsia benefits directly from both the presence and the territorial behaviour of Patella. Algal productivity was assessed by measuring oxygen evolution and utilization in situ and deriving photosynthesis/irradiance curves. Productivity was increased by about 30% by the presence of Patella in both summer ( P of grazed algae 0.0098; ungrazed algae 0.0063 mg C · cm · h) and winter ( P grazed algae 0.0081; ungrazed algae 0.0053 mg · C · h). Algal growth rates were not significantly increased by the application of limpet mucus in the laboratory. We did not examine nutrient regeneration by the limpet, but the increase in photosynthetic rate may depend on the limpet's grazing pattern which creates secondary sites for growth. Ralfsia also benefited from the territorial behaviour of Patella. The effects of different grazing regimes were investigated in different seasons by removing territorial limpets and either excluding all limpets using copper-based antifouling paint, or allowing access to non-territorial limpets (mostly P. oculus) using partial paint barriers. Exclusion of all limpets resulted in rapid overgrowth of Ralfsia plants by the foliose green alga Ulva sp.. Where non-territorial limpets had access to the plants overgrowth was reduced but Ralfsia plants were entirely removed by destructive grazing. Non-territorial grazers removed 90% of Ralfsia plants within 4 weeks in summer and 60% in winter. In control treatments P. longicosta prevented overgrowth by Ulva and actively excluded vagrant grazers, preventing overgrazing. Based on these findings, the association between the limpet and alga can be regarded as a nonobligate mutualism. [ABSTRACT FROM AUTHOR]

Terrestrial and freshwater pulmonate snails exhibit a marked depression of aerobic metabolism during estivation. This is an adaptation for existence in periodically harsh environments and, though marine gastropods may undergo anaerobic metabolism, they have not been shown to adaptively depress aerobic metabolic rate. We compared the metabolic response to progressive aerial exposure of two intertidal gastropod limpets, a prosobranch and a pulmonate. The prosobranch Patella granularis maintained a constant heart rate until shortly before death. In contrast, the pulmonate Siphonaria oculus underwent facultative depression of heart rate, accompanied by a decline in oxygen consumption. Both heart rate and oxygen consumption returned to normal levels on reimmersion in water. Metabolic rate depression is energy conserving, and may account for the ability of S. oculus to extend higher up the shore than P. granularis, into areas where food availability is low. S. oculus is a primitive, marine pulmonate, periodically subject to harsh conditions, and its capacity for metabolic rate depression may represent a pre-adaptation for life on land. [ABSTRACT FROM AUTHOR]