Selective consumption and facilitation by mesograzers in adult and colonizing macroalgal assemblages

Grazing pressure on macroalgae in littoral communities may vary with algal species, the age of an algal individual and grazer identity. Previous studies on alga–grazer interactions have shown that grazer preference for an algal species may release another one from interspecific competition. We measured the impacts of four common grazer taxa and the natural grazer guild on macroalgal communities at both their colonization and adult stages, and compared the impacts to grazer exclosures. The grazer effects were stronger on colonizing than on adult macroalgae; grazers did not reduce the total density of adult algae. Grazers both feed on propagules and indirectly facilitate other algae, depending on the grazer or algal species. Hydrobia species increased the settlement of spores of the red alga Ceramium tenuicorne. Similarly, the gastropod Theodoxus fluviatilis tended to facilitate one crustose algal species, but decreased the propagule density of annual filamentous algae, suggesting a preference for one species to the advantage of another. Effects of crustacean mesograzers on the studied macroalgae were weak. These results indicate that northern Baltic macroalgae are limited to grazing mainly during their colonization stage.     

Effects of macroalgal structural complexity on nearshore larval and post-larval crab composition

Larval and post-larval crab distribution was surveyed in three different habitats in Kachemak Bay, Alaska to determine temporal and spatial variability. Distribution varied temporally and spatially from June 2005 to September 2006. Nine sites of varying habitat complexity were surveyed monthly using scuba surveys and light traps to measure habitat variables and quantify crab zoeae and megalopae. A total of 10,016 crabs belonging to seven families were identified. Four species comprised the majority (97%) of the total crab assemblages and included Cancer oregonensis, Fabia subquadrata, Telmessus cheiragonus, and Pugettia gracilis. Peak abundances occurred in summer but varied on small temporal scales with species. No single bay-wide variable determined the appearance of all species. Depending on species, appearance may be influenced by seasonality of environmental variables. Spatially, highest abundances occurred in habitats with less structural complexity. Spatial differences in crab abundance may have resulted from variability on large scale physical transport mechanisms and not kelp-mediated flow alterations.     

Interhemispheric comparison of recruitment to intertidal communities: pattern persistence and scales of variation

Recruitment variation can be a major source of fluctuation in populations and communities, making it difficult to generalize results. Determining the scales of variation and whether spatial patterns in the supply of individuals are persistent over time can provide insight into spatial generality and the application of conservation and metacommunity models. We examined these issues using eight-year-long data sets of monthly recruitment of intertidal mussels (Mytilus spp., Perumytilus purpuratus, Semimytilus algosus, Brachidontes granulata) and barnacles (Balanus glandula, Chthamalus dalli, Jehlius cirratus, Notochthamalus scabrosus) at sites spanning > 900 km along the coasts of Oregon-northern California (OR-NCA, 45.47-39.43 degrees N) and central Chile (CC, 29.5-34.65 degrees S). We evaluated four general "null" hypotheses: that despite different phylogenies and great spatial separation of these taxa, their similar life history strategies and environmental settings lead to similar patterns of recruitment (1) between hemispheres, (2) in time, (3) in space, and (4) at larger and smaller spatial scales. Hypothesis 1 was rejected: along the OR-NCA coast, rates of recruitment were between two and three orders of magnitude higher, and patterns of seasonality were generally stronger and more coherent across space and time than along CC. Surprisingly, however, further analysis revealed regularities in both time and space for all species, supporting hypotheses 2 and 3. Temporal decorrelation scales were 1-3 months, and characteristic spatial scales of recruitment were approximately 250 km. Contrary to hypothesis 4, for the ecologically dominant species in both hemispheres, recruitment was remarkably persistent at larger mesoscales (kilometers) but was highly stochastic at smaller microscales (meters). Across species, increased recruitment variation at large scales was positively associated with increased persistence. Our results have several implications. Although the two regions span distinct latitudinal ranges, potential forcing processes behind these patterns include similar large-scale climates and topographically locked hydrographic features, such as upwelling. Further, spatial persistence of the recruitment patterns of most species at the mesoscale supports the view that marine protected areas can be powerful conservation and management tools. Finally, persistent and yet contrasting spatial patterns of recruitment among competing species suggest that recent metacommunity models might provide useful representations of the mechanisms involved in species coexistence.     

绿潮藻类暴发对天鹅湖水体和沉积物磷含量的影响

在荣成天鹅湖藻类暴发区域采集新鲜沉积物和丝状硬毛藻（Chaetomorpha spp.）,进行室内模拟试验,监测了生长过程中硬毛藻的生物量、磷富集量以及不同处理水体总磷（TP）和可溶性磷（SRP）质量浓度的变化,并分析了藻类生长对沉积物中各形态磷含量的影响。结果表明,当水体磷含量较高时,硬毛藻生长较快,相对生长速率高达14.88%,之后随着水体磷浓度的下降,生长速率逐渐减小。不同处理间硬毛藻的生物量相差很大,高磷含量处理显著高于低磷处理,最大差值可达26.50 g。随着藻类的生长,水体TP和SRP含量明显降低,其中高磷含量处理的TP质量浓度由0.93 mg·L-1降至0.01 mg·L-1,低磷含量处理水体SRP质量浓度均降至0.006 mg·L-1以下。当水体磷含量降至一定水平,沉积物中磷可向水体释放,其中可还原态磷和铁铝结合态磷的降幅分别为23.98%和12.61%。在高磷含量处理组,藻体中磷的富集量显著升高,且当水体磷含量相同的条件下,有沉积物处理的富集量显著高于无沉积物处理。相关分析表明,藻体生物量与水体TP和SRP的相关性较好,其中高磷含量处理组生物量与水体TP、SRP呈高度负相关,而相对生长速率与之呈显著正相关。结果说明,水体及沉积物中磷均可作为硬毛藻生长的营养来源;另一方面,藻类生长可明显降低水体磷含量,并促进沉积物中磷的释放。     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Effects of salinity on multiplication and transmission of an intertidal trematode parasite

Salinity levels vary spatially in coastal areas, depending on proximity to freshwater sources, and may also be slowly decreasing as a result of anthropogenic climatic changes. The impact of salinity on host–parasite interactions is potentially a key regulator of transmission processes in intertidal areas, where trematodes are extremely common parasites of invertebrates and vertebrates. We investigated experimentally the effects of long-term exposure to decreased salinity levels on output of infective stages (cercariae) and their transmission success in the trematode Philophthalmus sp. This parasite uses the snail Zeacumantus subcarinatus as intermediate host, in which it asexually produces cercariae. After leaving the snail, cercariae encyst externally on hard substrates to await accidental ingestion by shorebirds, which serve as definitive hosts. We found that at reduced salinities (25 or 30 psu), the cercarial output of the parasite was lower, the time taken by cercariae to encyst was longer, fewer cercariae successfully encysted and encysted parasites had lower long-term survival than at normal seawater salinity (35 psu). The strong effect of salinity on the replication and transmission of this parasite suggests that there may be sources and sinks of transmission to birds along coastal areas, depending on local salinity conditions. Also, unless it evolves to adapt to changing conditions, the predicted reduction in salinity as a consequence of climate change may have negative impact on the parasite’s abundance.     

Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Correlations between weather and recruitment of Nassarius pauperatus,an intertidal gastropod with a planktotrophic larval stage

Recruitment of the intertidal gastropod Nassarius pauperatus (Lamarck) was estimated annually for more than a decade at three sites in South Australia. Recruitment varied temporally, but was relatively synchronous amongst sites, even though these were at least 10 km apart. The density of recruits at each site was negatively correlated with evaporation and also with several indices of temperature for February. We postulate that either high temperatures or desiccation cause the death of recently settled individuals, which may be unable to avoid these extremes by burrowing deeply into the substratum.     

Effects of settlement density on spatial arrangement in four intertidal barnacles

Balanus amphitrite was studied in an estuary (Halifax River) in central Florida from October to November 1990, while three other barnacle species were studied on the central coast (Monterey Bay) of California from April 1988 to July 1989. Mean nearest-neighbor distances indicated that in the majority of cases the spatial arrangement of settlers was random for three different balanomorph barnacles-B. amphitrite, B. glandula, and Chthamalus dalli. In Pollicipes polymerus, a lepidomorph species, strong aggregation among settlers was almost always observed. In the three balanomorph species there was no apparent relationship between settler density and the degree of aggregation. In P. polymerus there was a significant positive correlation between settler density and the degree of aggregation. Morphological differences between balanomorph and lepidomorph barnacles may influence the evolution of settlement behavior.     

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.     

Community succession following massive ice-scour on a rocky intertidal shore: recruitment,competition and predation during early,primary succession

The importance of species interactions and recruitment variability was examined during the first year and a half of primary succession (1988–1989) on an exposed rocky seashore near Halifax, Nova Scotia. Previous work suggested that emergent rock on these shores is normally dominated by fucoid rockweeds because predatory whelks control the sessile animal competitors, mussels and barnacles, and because herbivorous littorinids control ephemeral algal competitors. Abundances of all species except seasonal ephemeral algae were very small throughout this experiment and we found no significant effects of carnivory, herbivory, plant-animal competition or plant-plant competition. A slight facilitation of Fucus recruitment is attributed to a thin mat of ephemeral, blue-green algae. Very few other studies have directly manipulated intertidal ephemeral algae. As primary succession may be very rare in this assemblage, these results may be specific to these circumstances, but they highlight the varying importance of species interactions with variable recruitment. In particular, it appears that variations in recruitment success may be important to community structure, even when recruitment is not limited by propagule supply. The scale of the study also provides insight into successional processes occurring after the recent, extensive ice-scour of exposed seashores in this region.     

Effects of physical disturbance on an estuarine intertidal community: field and mesocosm results compared

The response of an estuarine macrobenthic community to different intensities of physical disturbance was examined using comparable field and mesocosm experiments. In the field study, areas of sediment (2500 cm²) were disturbed by surface-raking at a range of disturbance intensities: no disturbance, once every 2 mo, once every month, once every 2 wk, once or three times every week. These disturbance regimes were also imposed on buckets of sediment (855 cm²) collected from the field study site and maintained within a mesocosm system. In the univariate data analysis, neither field nor mesocosm communities displayed any trends in relation to the frequency of disturbance. Diversity indices were generally reduced in the mesocosm communities compared with the field communities. Multivariate analysis indicated significant differences between the field communities subjected to the most intense disturbance and those communities disturbed every ±2 wk or less. These differences were attributed to reductions in the five numerically dominant species in the high- disturbance treatments. More statistically significant differences were observed between the different disturbance intensities in the mesocosm experiment than in the field study. The implications of differences between field and mesocosm experiments results are discussed. Received: 14 June 1999 / Accepted: 3 December 1999     

Effects of detrital non-native and native macroalgae on the nitrogen and carbon cycling in intertidal sediments

The decay of non-native and native seaweed mixing may modify sediment biogeochemistry and organic matter transfers within benthic food webs according to their composition and biomass. The non-native species Sargassum muticum was deliberately added to the sediment of an intertidal sandflat at different biomass and mixed to the native species Ulva sp. and Fucus vesiculosus. The sediment porewater was then ¹³C and ¹⁵N enriched to test whether both detrital diversity and biomass influenced the transfer of porewater carbon and nitrogen to the sediment and to the macrofauna consumers. More ¹⁵N-nitrogen was mobilized to sediments and macrofauna when the 3-species detrital mixing was buried, probably because this mixing provided species-specific compounds such as polyphenols due to the presence of S. muticum and F. vesiculosus, as well as large amounts of nitrogen due to the presence of Ulva. Our study revealed the importance of detrital diversity and non-native seaweeds for the nitrogen cycling in the benthic food web.     

Effects of a complex archipelago on genetic subdivision of the intertidal limpet Siphonaria kurracheensis

Allozyme variation in the intertidal limpet Siphonaria kurracheensis was examined in 11 populations from the Houtman Abrolhos Islands, Western Australia, and four from the adjacent mainland, to test if the effects of these islands on genetic subdivision should include species with high dispersal. Consistent with extensive planktonic dispersal, genetic subdivision was low, with an average standardized variance in allelic frequencies (F_ST) of 0.010 over 400 km along the mainland, and 0.009 over 70 km in the Abrolhos Islands. Nevertheless, subdivision was statistically significant at the smallest scale examined (about 10 km), showing a pattern of local heterogeneity and large-scale homogeneity, which is common in planktonic dispersers. Among the island populations, heterozygosity was positively correlated with a multivariate index of exposure. The level of heterozygosity at the most exposed island sites is equivalent to that at the (also exposed) mainland sites, whereas it is lower in the most sheltered island sites. Although this indicates that some island populations are less well connected to the major pool of larvae, the overall effect of the archipelago on genetic subdivision is no greater than occurs on the mainland coast. This is contrary to findings in previously studied species with less potential for dispersal, suggesting that the local impediments to dispersal are selective in their effects, and are unlikely to be broadly important for genetic divergence in widely dispersing species.     

Allelopathic inhibition on red tide microalgae by five macroalgal extracts

This study aims to identify effective antialgal allelochemicals from marine macroalgae that inhibit the growth of red tide microalgae. Practically, new algicidal agents were developed to control red tide. The growth inhibitory effects of 5 marine macroalgae Porphyra tenera, Laminaria japonica, Ulva pertusa, Enteromorpha clathrata, and Undaria pinnatifida on Skeletonema costatum were evaluated by adding crude seawater extracts of macroalgal dry tissue into the culture medium containing S. costatum. The half-effective concentrations at 120 h (EC_{50, 120 h}) of the seawater extracts were 0.6, 0.9, 1.0, 1.0, and 4.7 g/L for the five macroalgae above, respectively. E. clathrata, L. japonica and U. pertusa showed strong allelopathic effect on the growth of S. costatum. There have been no previous reports with regard to the allelopathic effects of the former two macroalgae so far. The possible allelochemicals of 21 compounds of the E. clathrata were detected using Gas chromatography-mass spectrometry (GC-MS) analysis. Unsaturated fatty acids, acrylic acid (C₃H₄O₂), and linolenic acid (C₁₈H₃₀O₂) were the most likely allelochemicals in E. clathrata.     

Effects of age and sex ratios on offspring recruitment rates in translocated black rhinoceros

Success of animal translocations depends on improving postrelease demographic rates toward establishment and subsequent growth of released populations. Short‐term metrics for evaluating translocation success and its drivers, like postrelease survival and fecundity, are unlikely to represent longer‐term outcomes. We used information theory to investigate 25 years of data on black rhinoceros (Diceros bicornis) translocations. We used the offspring recruitment rate (ORR) of translocated females—a metric integrating survival, fecundity, and offspring recruitment at sexual maturity—to detect determinants of success. Our unambiguously best model (AICω = 0.986) predicted that ORR increases with female age at release as a function of lower postrelease adult rhinoceros sex ratio (males:females). Delay of first postrelease reproduction and failure of some females to recruit any calves to sexual maturity most influenced the pattern of ORRs, and the leading causes of recruitment failure were postrelease female death (23% of all females) and failure to calve (24% of surviving females). We recommend translocating older females (≥6 years old) because they do not exhibit the reproductive delay and low ORRs of juveniles (<4 years old) or the higher rates of recruitment failure of juveniles and young adults (4–5.9 years old). Where translocation of juveniles is necessary, they should be released into female‐biased populations, where they have higher ORRs. Our study offers the unique advantage of a long‐term analysis across a large number of replicate populations—a science‐by‐management experiment as a proxy for a manipulative experiment, and a rare opportunity, particularly for a large, critically endangered taxon such as the black rhinoceros. Our findings differ from previous recommendations, reinforce the importance of long‐term data sets and comprehensive metrics of translocation success, and suggest attention be shifted from ecological to social constraints on population growth and species recovery, particularly when translocating species with polygynous breeding systems.     

Hull fouling on commercial ships as a vector of macroalgal introduction

Hull fouling is thought to have been the vector of introduction for many algal species. We studied ships arriving at a Mediterranean harbour to clarify the present role of commercial cargo shipping in algal introductions. A total of 31 macroalgal taxa were identified from 22 sampled hulls. The majority of records (58%) were of species with a known cosmopolitan geographical distribution. Due to a prevalence of cosmopolitan species and a high turnover of fouling communities, species composition of assemblages did not appear to be influenced by the area of origin, length of ship or age of coating. In the light of the present results, hull fouling on standard trading commercial vessels does not seem to pose a significant risk for new macroalgal species introductions. However, a high proportion of non-cosmopolitan species found on a ship with non-toxic coating may modify this assessment, especially in the light of the increasing use of such coatings and the potential future changes in shipping routes.     

Effects of life history and reproduction on recruitment time lags in reintroductions of rare plants

Reintroductions are important components of conservation and recovery programs for rare plant species, but their long-term success rates are poorly understood. Previous reviews of plant reintroductions focused on short-term (e.g., ≤3 years) survival and flowering of founder individuals rather than on benchmarks of intergenerational persistence, such as seedling recruitment. However, short-term metrics may obscure outcomes because the unique demographic properties of reintroductions, including small size and unstable stage structure, could create lags in population growth. We used time-to-event analysis on a database of unusually well-monitored and long-term (4–28 years) reintroductions of 27 rare plant species to test whether life-history traits and population characteristics of reintroductions create time-lagged responses in seedling recruitment (i.e., recruitment time lags [RTLs]), an important benchmark of success and indicator of persistence in reintroduced populations. Recruitment time lags were highly variable among reintroductions, ranging from <1 to 17 years after installation. Recruitment patterns matched predictions from life-history theory with short-lived species (fast species) exhibiting consistently shorter and less variable RTLs than long-lived species (slow species). Long RTLs occurred in long-lived herbs, especially in grasslands, whereas short RTLs occurred in short-lived subtropical woody plants and annual herbs. Across plant life histories, as reproductive adult abundance increased, RTLs decreased. Highly variable RTLs were observed in species with multiple reintroduction events, suggesting local processes are just as important as life-history strategy in determining reintroduction outcomes. Time lags in restoration outcomes highlight the need to scale success benchmarks in reintroduction monitoring programs with plant life-history strategies and the unique demographic properties of restored populations. Drawing conclusions on the long-term success of plant reintroduction programs is premature given that demographic processes in species with slow life-histories take decades to unfold.