The effects of seaweed harvesting on fishes: a response

References Rangeley (1994) has argued that sampling biases, errors in experimental design and low statistical power make the conclusion by Black & Miller (1991), that there was no large impact of an experimental harvest of the seaweedAscophyllum nodosum on fishes, unmerited and misleading. Our study was conducted mainly at two sites close to Lower Argyle, southern Nova Scotia, Canada where harvesting ofAscophyllum nodosum has been carried out continuously for 30 years along-side intense and productive shellfish and finfish fisheries. In southern Nova Scotia, on-going removal ofAscophyllum nodosum with mechanical harvesters is about 17% (Sharpe & Semple 1991); planned annual removal in New Brunswick where Rangeley has worked will be only about 5% (Anon. 1993). The context of our investigation was a small-scale (400 m²) but extreme, experimental removal (100%) of the seaweed. Our focus was on fishes that moved into the intertidal zone as the tide rose in the evening and early morning and were caught in trammel nets, and on the fishes that were retained in a fixed seine that was set at high tide in early morning and the fishes removed at the next low tide. Thus, subject to the constraint on the size of the fishes imposed by the nature of our fishing gear with its stretch mesh of 25 mm, we were concerned with the identity and abundance of fishes that moved in and out of the intertidal zone and their behavioural response to the 400 m² patches of habitat that formed our experimental units. The most striking result was the unexpected, low abundance of fishes caught in our gear.     

Trends in abundance of rocky intertidal seaweeds and filter feeders across gradients of elevation,wave exposure,and ice scour in eastern Canada

On NW Atlantic rocky shores, the main basal organisms in intertidal communities are seaweeds (Ascophyllum nodosum, Fucus spp. and Chondrus crispus) and filter feeders (barnacles, Semibalanus balanoides, and mussels, Mytilus spp.). Their ecology has been extensively studied in New England (United States), but knowledge gaps exist for northern shores, which are subjected to stronger environmental stress. Therefore, we studied the above organisms on Canadian shores. We quantified the summer abundance of these seaweeds and filter feeders across full vertical (intertidal elevation) and horizontal (wave exposure and winter ice scour) environmental gradients on the Gulf of St. Lawrence and open Atlantic coasts of Nova Scotia. At the regional scale along the open Atlantic coast, seaweeds showed similar abundances in Nova Scotia than values reported for New England. However, both filter feeders were considerably less abundant in Nova Scotia. At the local scale in Nova Scotia, intense winter ice scour (which only occurs on the Gulf of St. Lawrence coast) was associated with a very low abundance of all species except barnacles. Spatial trends in Nova Scotia were similar to patterns known for certain species elsewhere, such as A. nodosum being almost restricted to sheltered habitats, regardless of elevation, and C. crispus being almost restricted to low elevations, regardless of exposure. Other trends were, however, characteristic of Nova Scotia, such as C. crispus being frequent at low elevations in exposed habitats, unlike in New England, where mussels often predominate there because of competitive advantages. In Nova Scotia, mussels were always restricted to cracks and crevices, unlike in New England, where they form extensive intertidal beds on exposed shores. The direct effects of increased environmental stress and indirect effects through altered interspecific interactions might explain the regional differences in local species distribution, which will require experimental validation. Handling editor: K. Martens     

Alien algae <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> causes habitat loss for rocky reef fishes in north Patagonia

Since the introduction of Undaria into Nuevo Gulf, Argentina, around 1992, this alien seaweed has now colonized different sites over 700 km of coast, forming dense seasonal forests in waters from 0 to 15 meters in depth. In the spring it is common for plants of Undaria to break away from the substrate and be transported by sea currents. As Undaria gets stuck onto reefs it has the potential to reduce habitat quality for reef fish by physically obstructing refuges. This study aims to assess the impact of Undaria on the abundance of four species of rocky-reef fishes by an observational experiment. Fish abundance on reefs with and without Undaria was estimated by underwater visual census methods. Sites were classified according to their topographical relief, as this was expected to influence the effect of Undaria on the abundance of reef fishes. Fish abundance decreased markedly in low-relief reefs that had been covered by Undaria. In contrast, the drifting Undaria had no effect on the abundance of any of the fish species considered in high-relief reefs, where it tends to cover only the lowest-lying areas, leaving much of the refuges for fish unaffected. In conclusion, the presence of Undaria off the coast of Argentina results in transitory habitat loss for reef fishes inhabiting low-relief reefs during late spring and early summer. Although we do not know how much of a threat this habitat loss represents for the conservation of reef fish populations of northern Patagonia, the documented local impact of Undaria within the gulfs is strong and may affect a number of recreational and commercial activities which are centered on the reefs and their fish assemblages.     

Diel,semi‐lunar and seasonal patterns in the fish community of an intertidal zone of the Yangtze estuary

The structure and temporal variations of the fish community in the intertidal estuarine zone of shallow mud areas have been poorly studied in China. This paper analyses the diel, semi‐lunar and seasonal patterns of fish assemblages in the Yangtze estuary in 2006. Fish were collected by consecutive day and night samplings using tide‐stow‐nets deployed parallel to each other in three stations. A total of 56 fish species belonging to 21 families was caught during the study period. The family Cyprinidae dominated with 25 species. Freshwater fish species were the important dominant commercial fishery species and well represented with five species (sharpbelly Hemiculter bleekeri, goldfish Carassius auratus, bream Parabramis pekinensis, likely‐bream Pseudobrama simony, and glossy yellow catfish Pelteobagrus nitidus) in the three stations. Juvenile fishes dominated the fish community, comprising 93.9% in station 1 and 96.6% in station 2 of the total abundance. The number of fish species in day tides was slightly lower than those in night tides in spring and summer, but the opposite in other seasons. In neap tides, the numbers and abundance of fish species were both lower than those in the spring tides. Fish abundance was lowest in winter, increasing during spring and summer (March–September) in both stations 1 and 2, with obviously large fluctuations in each season. The pattern of habitat selection of fishes could effectively decrease the food competition of intraspecies or interspecies and favour the growth and nursing of young fishes. These findings indicate that the intertidal zones in the estuary may serve as important nursery areas for fish communities.     

Biomass and height of Ascophyllum nodosum after two decades of continuous commercial harvesting in eastern Canada

With the increasing demand for seaweed resources worldwide, management must ensure that the harvest of wild seaweed stocks is sustainable. We evaluate the impact of over 25 years of commercial harvesting of Ascophyllum nodosum in eastern Canada by comparing the biomass and height of the seaweed in the late 1990s to the late 2010s over a broad spatial scale spanning the provinces of Nova Scotia and New Brunswick. There has been no significant decrease in the biomass of A. nodosum in either province, and biomass has increased in some regions of New Brunswick during that period. The average height of A. nodosum has decreased by 7.8 cm in Nova Scotia while it increased by 13.8 cm in New Brunswick. Biomass of A. nodosum in unharvested sectors was 7% higher than that in harvested sectors while height of A. nodosum in unharvested sectors in New Brunswick is similar to the values observed in harvested sectors. Over the same period, water temperature has increased in both provinces and, in recent years, has at times exceeded the optimal growing temperature for A. nodosum within bays in Nova Scotia. We conclude that the current management and harvest of A. nodosum in eastern Canada are sustainable and maintain the biomass and height of A. nodosum beds but that control sites are necessary to offer adequate comparisons as environmental conditions are changing.

     

A field investigation into the effects of interspecific competition on the distribution and cover of Ulothrix flacca

Ulothrix flacca is a filamentous green alga found in its greatest abundance in the littoral fringe of the rocky intertidal during the winter months. It is much less abundant below this level. This experiment was undertaken tot determine if the lower abundance below the littoral fringe is due to the effects of interspecific competition or physiological limitations. Clearing experiments were performed on the Coast of Maine, USA to eliminate competition. Belts 10 cm wide that ran from the littoral fringe downward into the eulittoral zone were cleared of all attached organisms down to bare rock. A second condition was set up in which only the brown algal canopy was removed from a 60 cm wide belt. Unaltered belts were designated as controls. Squares 0.25 m² in area were cleared in the littoral fringe and eulittoral zone. U. flacca was able to colonize the cleared areas of the eulittoral zone and reach abundances equal to those found in the unaltered control areas of the littoral fringe. This indicates that physiological limitations do not restrict the abundance of U. flacca in the eulittoral zone. However, evidence of grazing in cleared areas adjacent to high concentrations of the barnacle Semibalanus balanoides suggest that competition alone is not responsible for the limited abundance of U. flacca in the eulittoral zone. Herbivores associated with the presence of barnacles may also have an impact on the abundance of U. flacca, even during the winter months.     

Fishes of a subtropical Australian saltmarsh and their predation upon mosquitoes

Synopsis The fishes occurring in a subtropical Australian saltmarsh were studied from January to December 1984 to determine the seasonal abundance and feeding of the most common species. Wide variations in pool water levels and salinity were recorded. Eight species of fish were collected, four of which are of economic importance to fisheries. Three taxa, Gambusia affinis, Pseudomugil signifer, and gobiids predominated in terms of abundance and occurrence. The abundance of fishes fluctuated markedly on a seasonal basis with few fish being taken in winter and spring (June–November) when pool water levels were low. Diets of the most common species were similar with terrestrial foods forming an important component of all diets. Feeding activity was highest in summer (December to February) when mosquito larvae, copepods, and larval coleopterans formed a large proportion of the fishes' diets. Results from the present study indicate that fish important to commercial fisheries rarely utilise the upper littoral saltmarsh habitat and that resident fishes are unlikely to effectively control mosquito larvae in saltmarsh.     

The spring development of phytoplankton in Lake Erken: species composition,biomass, primary production and nutrient conditions — a review

Relative abundance and within-lake distributions of three fishes, northern redbelly dace (Phoxinus eos), finescale dace (Phoxinus neogaeus), and central mudminnow (Umbra limi), were examined using minnow traps in Tuesday Lake, a small bog lake in the Upper Peninsula, Michigan. For these species, catches in minnow traps placed at the perimeter of the lake were 21 to 52 times higher than catches in midlake traps. Variance: mean ratios of perimeter trap catches indicated that both dace species were highly aggregated while the distribution of mudminnows was less aggregated or random. Over an 11 day period during which all fish caught were removed from the lake, catch per unit effort (CPUE) of both dace species declined in response to fish removal. In contrast, CPUE for mudminnows was low initially, increased to an asymptote and then declined only in the last 5 days of the fish removal. The patterns of CPUE for mudminnows indicated that mudminnow trapability and/or activity was reduced in the presence of high densities of dace. The low abundance of dace in traps with many mudminnows suggested mudminnows avoided traps already containing dace. Throughout the removal period, CPUE provided an accurate index of dace abundance, whereas this was true for mudmnnows only after dace populations had been reduced drastically. Therefore, in any use of minnow traps to estimate populations, both spatial distributions and relative species abundance of small fishes must be taken into account.     

A quantitative evaluation of the effects ofAscophyllum harvesting on the littoral ecosystem

Little is known of the ecological effects of harvesting littoral algae although this is a worldwide commercial activity. In 1976 an attempt to establish harvesting in Strangford Lough, Northern Ireland, was opposed on mainly theoretical conservation grounds. The attempt began and stopped within a single small bay leaving a sharp boundary between cut and uncut areas. A subjective survey apparently confirmed the predicted loss of cryptic fauna, decline through predation and the resorting of interboulder sediment. In April 1979 the cut and uncut areas were examined in detail to determine whether any of these effects had persisted and were demonstrable scientifically. Beach and boulder transects and various other studies showed some increases in the cut area. There was significantly moreFucus, Enteromorpha andUlva; Cirratulus (inhabitingRhodochorton-bound sediment on boulder surfaces) had a greater biomass. Some changes inLittorina colour morphs were apparent. Sediment in the cut area was coarser and had significantly more crustacean meiofauna.Ascophyllum internodal length and lateral branching were increased but it still provided 20% less shore cover than in the uncut area. There were significant decreases in the cover ofCladophora on the sides of boulders and ofHalichondria, Hymeniacodon andBalanus on undersurfaces. Indeed on the habitable underside of boulders total animal cover had been reduced by nearly two-thirds and the average number of species per boulder by one-third. It is concluded thatAscophyllum harvesting has a significant and persistent effect on shore ecology. Littoral algae are a valuable commercial asset but it is important that some fairly large intertidal areas should be left unharvested for general conservation purposes.     

Competition between herbivourous fishes and urchins on Caribbean reefs

Summary When the common sea urchin Diadema antillarum was removed from a 50 m strip of reef in St. Thomas, US Virgin Islands, cover of upright algae and the grazing rates and densities of herbivorous parrotfish and surgeonfish increased significantly within 11–16 weeks when compared to immediately adjacent control areas. Sixteen months after removal, Diadema had recovered to 70% of original density, abundance of upright algae no longer differed between removal and control areas, and the abundance and grazing activity of herbivorous fish in the removal was approaching equivalence with control areas. On a patch reef in St. Croix that had been cleared of Diadema 10–11 years earlier (Ogden et al. 1973b), urchins had recovered to only 50–60% of original density. This reef still showed significantly higher rates of grazing by fish and a significantly greater density of parrotfish and surgeonfish than a nearby control reef where Diadema densities had not been altered. These results indicate that high Diadema densities (7–12/m² for this study) may suppress the densities of herbivorous fish on Caribbean reefs.     

COLONIZATION AND GROWTH OF ASCOPHYLLUM NODOSUM (PHAEOPHYTA) IN MAINE1

Colonization and growth of Ascophyllum nodosum (L.) Le Jol. were investigated in two Maine estuaries from 1972 to 1978. Following denudation of intertidal rock, substrata were initially colonized by Fucus vesiculosus L.; eventually, Ascophyllum supplanted Fucus, and became dominant in terms of percentage cover. Ascophyllum settled first and most densely in the low intertidal zone, but its fastest growth occurred in the mid-intertidal zone. Some, but not all, Ascophyllum germlings produced a vesicle within one year of colonization. The mean annual growth of A. nodosum was variable among sites, zones and years.     

Spatial variation in density of stream benthic fishes in northern Hokkaido, Japan: does riparian vegetation affect fish density via food availability?

The densities of two benthic fishes, the Siberian stone loach (Noemacheilus barbatulus) and the wrinklehead sculpin (Cottus nozawae), and the biomass of their food resources (i.e., periphyton and benthic invertebrates) were compared between forest and grassland streams in northern Hokkaido, Japan, to examine whether riparian deforestation had positive effects on the benthic fishes via enhancement of food availability. The comparisons indicated that riparian vegetation had little influence on periphyton, invertebrates, or fishes. Regression analysis indicated that spatial variations in loach and sculpin densities were explained more by substrate heterogeneity, competitor abundance, or both, rather than by food abundance. However, when the two species were combined as benthic insectivores, a strong correlation was found between total benthic fish density and invertebrate biomass. Our results suggest that, although total benthic fish abundance was food limited, riparian vegetation had no positive effects via food availability on the benthic fishes in our streams.     

New ecological information onScytalina cerdale (Pisces: Scytalinidae) from a central California rocky intertidal zone

Synopsis New information regarding the ecology ofScytalina cerdale was obtained over a four year period as a consequence of a long-term marine ecological study at the Diablo Canyon Power Plant (DCPP), San Luis Obispo County, California. Twenty intertidal fish surveys were conducted at approximately quarterly intervals, between March 1979 and June 1983, at three separate rocky shore locations (stations). During each survey, a total of 108 square meters (36 m² per station) was searched for fish during periods of low tide. A total of 280S. cerdale were collected, identified, measured, and released back into the same 4 m² area, from the same intertidal station (Diablo Cove), throughout the study period. This limited intertidal occurrence most likely reflectsS. cerdale microhabitat requirements; a combination of intertidal elevation (mean = +0.3 MLLW), substratum specificity (loose gravel, 5–10mm size range, overlying a base of sand and shell fragments), and degree of wave exposure (semi-protected). Throughout the study,S. cerdale was seasonally more abundant during summer months (June through August) and less abundant during winter months (November through February). The only exception to this abundance trend followed the 1982 winter storms, which coincided with an El Niño event, whenS. cerdale abundance uncharacteristically dropped during the subsequent 1983 spring and summer surveys. Mean total lengths did not vary greatly, reflecting the absence of early juvenile fishes, and relatively high mean fish densities (3.5 fish per m² for 20 surveys) were recorded. Qualitative comparisons betweenS. cerdale abundance and seasonal changes in water temperature indicated an inverse relationship. Gravid females, demersal egg masses, and early juvenile individuals were never observed during the four years of the study. This suggests that unlike most other intertidal fishes, particularly other blennioids,S. cerdale may not utilize the intertidal zone for reproductive and recruitment purposes; Diablo Canyon is the most southerly reported distribution for this intertidal species, and therefore, may not entirely represent the species' biology as a whole.     

Mussel and dogwhelk distribution along the north‐west Atlantic coast: testing predictions derived from the abundant‐centre model

Aim We performed the first test of predictions from the abundant‐centre model using north‐west Atlantic coastal organisms. We tested the hypotheses that the density of intertidal mussels (Mytilus edulis and M. trossulus) and dogwhelks (Nucella lapillus) and mussel age and size would peak at an intermediate location along their distribution range. We also assessed the latitudinal variation in critical aerial exposure time. Location North‐west Atlantic coast between Newfoundland (Canada) and New York (USA), covering 1800 km of shoreline. Methods Using a nested design, we measured mussel density, age and size and dogwhelk density in 60 wave‐exposed rocky intertidal sites spread evenly in six regions. Critical aerial exposure times were determined using online data. Results Mytilus edulis peaked in abundance in Maine and was much less abundant in the other regions. Mytilus trossulus peaked in abundance in southern Nova Scotia and Maine, was less abundant in the other regions to the north, and was absent in the southernmost region (New York). Both mussel species were least abundant in a northern region (Cape Breton), although not in the northernmost region (Newfoundland). Critical aerial exposure times were negatively correlated with overall mussel density. Mussel age and size were similar among regions. Dogwhelks peaked in abundance in Maine and were much less abundant in the other regions, being positively correlated with overall mussel density across regions. Main conclusions Density data for M. edulis and N. lapillus provide limited support for an abundant‐centre pattern, while M. trossulus shows a clear ramped‐south distribution. Critical aerial exposure times suggest that physiological stress during summer and winter low tides may be lowest in Maine and southern Nova Scotia, which might partially explain mussel predominance in those regions. Winter ice scour in Cape Breton may explain the abundance trough observed there. Mussel size and age may be more limited by wave exposure at our sites (as they all face open waters) than by regional differences in environmental stress. Dogwhelks, which prey on mussels, seem to respond positively to prey density at the regional scale. Our study supports the notion that, while the abundant‐centre model is a useful starting point for research, it often represents an oversimplification of reality.     

Persistence of gametophyte predominance in Chondrus crispus (Rhodophyta,Gigartinaceae) from Nova Scotia after 12 years

Gametophytes predominated clearly over tetrasporophytes in an intertidal population of Chondrus crispus at Tor Bay (Nova Scotia, Canada) in the summer of 1991. Since this species is perennial and the rocky substrate is stable at this site, we predicted that gametophyte predominance would persist after several years. We confirmed this hypothesis by re-sampling the same area in the summer of 2003. This is one of the first long-term studies of the relative abundance of life-history phases done unequivocally at the same site for the Gigartinaceae.     

IMPACT OF THE INVASIVE SEAWEED SARGASSUM MUTICUM (PHAEOPHYTA) ON AN INTERTIDAL MACROALGAL ASSEMBLAGE1

The impact of the invasive seaweed Sargassum muticum (Yendo) Fensholt on a low intertidal macroalgal assemblage was assessed at a semiexposed rocky shore in northern Spain between 2002 and 2004. Sargassum muticum plants were removed from the mature macroalgal assemblage and from those occurring along the successional process of the assemblage. Biomass, richness, diversity, and percentage cover of macroalgae in experimental plots were compared with unmanipulated controls. The effect of S. muticum removal on the macroalgal assemblage more than 2 years after the beginning of the experiment was negligible. Moreover, no differences between treatments were detected in the general patterns of succession. Only significant differences in S. muticum abundance were detected between treatments at the end of the experiment. We suggest that the low abundance of S. muticum at this intertidal level and its pseudoperennial life cycle may limit competition with native macroalgae. However, long‐term removal experiments may be a more indicator of the impact of S. muticum at the upper limit of its vertical distribution.     

Community structure of fishes attracted to shallow water fish aggregation devices off South Carolina,U.S.A.

Synopsis Twenty-two fish aggregation devices were deployed in 14 m of water off South Carolina. Species composition and abundance were determined by diver visual census on eight occasions from May through November, 1985. A total of 21 families and 36 species of fishes was observed at 121 stations. Pelagic fishes dominated the fauna with a 99.3% relative abundance, and Decapterus punctatus accounted for 97.6% of the individuals. Caranx crysos, Diplectrum formosum, Decapterus punctatus, Centropristis striata and Monacanthus hispidus were the most frequent species. Total fish abundance, number of species and abundance of four of the six most common species were significantly affected by season. Hurricane activity may have caused a significant drop in pelagic fish abundance at the FADs in July. No significant correlations among species abundances were found after removal of season and FAD type effects. Spatial zonation and seasonal occurrence patterns suggest some competition among pelagic fishes. Several factors that regulate FAD faunal abundance and composition are hypothesized, including: juvenile fish availability, availability of shelter, availability of adequate food resources, interspecific and intraspecific competition, severe sea conditions, and sporadic intrusions of large predatory fishes. It is hypothesized that the abundances of benthic and pelagic FAD fishes are correlated and that there is a direct or indirect energetic link between shallow water pelagic and benthic fish assemblages near FADs.     

Grazing by the intertidal fish Anoplarchus purpurescens upon a distasteful polychaete worm

Synopsis Cirratulid worms are common inhabitants of the central California rocky intertidal zone and appear to offer a potentially rich source of food for intertidal fishes. However, analyses of stomach contents revealed that they do not appear in the diets of the commonest intertidal fishes. Apparently only one species, the eel blenny Anoplarchus purpurescens, feeds on cirratulids in significant amounts.Feeding experiments employing local intertidal fishes showed that the common intertidal cirratulid, Cirriformia luxuriosa, is distasteful to most of the fishes. Only two species ate its tentacles, and only A. purpurescens consistently ate large quantities. We propose that C. luxuriosa possesses a predation-deterring chemical similar to that reported in C. spirabrancha, although A. purpurescens apparently has been able to circumvent this anti-predator mechanism.The ability of A. purpurescens to eat Cirriformia tentacles allows it to tap a seemingly little-used food source, and thus may decrease competition between A. purpurescens and other intertidal fishes. This feeding relationship also represents a possible example of coevolution between a predator and its prey.     

Morphological variation in populations of the diatom Asterionella ralfsii W. Smith from Nova Scotia,Canada

Preliminary investigations of the diatom genus Asterionella ralfsii W. Smith from Kejimkujik National Park, Nova Scotia indicate that its morphology differs from other reported forms. Mean cell length increased between the spring and the fall. Bimodal distribution of length classes occurred in several lakes and was not related to measured environmental variables. The need for further work on this species in this and other geographical areas is considered.     

Why do anemonefishes inhabit only some host actinians?

Synopsis The 25 species ofAmphiprion and one ofPremnas (family Pomacentridae) are obligate symbionts of 10 species of facultatively symbiotic sea anemones. Throughout the tropical Indo-West Pacific range of the relationship, a fish species inhabits only certain of the hosts potentially available to it. This specificity is due to the fishes. Five fishes occupy six sea anemone species at Lizard Island, Great Barrier Reef, Australia.Entacmaea quadricolor harborsP. biaculeatus, A. melanopus andA. akindynos. Adults ofPremnas occur deeper than about 3 m in large, primarily solitary actinians; juveniles may occupy peripheral members ofEntacmaea clones in shallow water. Specimens ofA. melanopus live exclusively in clonal anemones, which are found no deeper than 3 m. Most individuals ofA. akindynos inEntacmaea are juveniles, occurring shallow and deep, in solitary anemones or at the margins of clones. Interspecific as well as intraspecific social control of growth may be responsible for keeping fish small at clone fringes. Conspicuous specimens ofE. quadricolor depend upon their anemonefish to survive. Actinians cleared of symbionts disappeared within 24 h, probably having been eaten by reef fishes.Entacmaea, the most abundant and widespread host actinian at Lizard Island and throughout the range of the association, is also arguably the most attractive to anemonefishes. I believe its vulnerability to predation was a factor in its evolving whatever makes it desirable to fishes. Experimental transfers pitted fish of one species against those of another, controlling for ecophenotype of host, and sex, size and number of fish. Competitive superiority was in the same order as abundance and over-all host specificity:P. biaculeatus, A. melanopus, A. akindynos. At least three factors are necessary to explain patterns of species specificity - innate or learned host preference, competition, and stochastic processes.