Influences of microhabitat use and foraging mode similarities on intra- and interspecific aggressive interactions in a size-structured stream fish assemblage

Aggressive interactions, foraging behaviour and microhabitat use were observed among four sympatric stream fishes inhabiting the water column: ayu (Plecoglossus altivelis), white-spotted charr (Salvelinus leucomaenis), masu salmon (Oncorhynchus masou) and Japanese dace (Tribolodon hakonensis), each species being categorised into five body-size classes (species-size groups; SSG's). Aggressive interactions were observed between most pairs of SSG's, an almost linear dominance order being apparent throughout the three-month study period. Ayu were relatively subordinate in June, but became the second most dominant in July and the most dominant in August, as a consequence of a reversal in dominance order with salmon. In contrast, smaller-sized dace, which continually suffered from intra- and interspecific aggression, occupied the most subordinate ranks throughout the study period. Intensive aggression was observed among various SSG's, exhibiting same microhabitat propensity throughout the three months. The direction and frequency of aggressive interactions varied month by month due to a reversal in dominance order between ayu and masu salmon, and/or changes in density, body size and resource use of the component members. Opponent selectivity was higher within SSG's, where resource use was assumed to be highly overlapping, rather than among SSG's throughout the study period. Correlation analysis indicated that opponent selectivity in aggressive interactions among SSG's was positively correlated with similarity in microhabitat selectivity in June, but not in other months or with that in foraging habits, suggesting that intensive aggressive behaviour reflected overlapping habitat use among assemblage members during a certain period.     

Intraguild indirect effects through trophic cascades between stream-dwelling fishes

1. To examine the strength and role of indirect effects through trophic cascades by omnivorous fish on algal biomass in streams, we introduced one of four fish species (ayu Plecoglossus altivelis altivelis, pike gudgeon Pseudogobio esocinus esocinus, Japanese dace Tribolodon hakonensis and pale chub Zacco platypus) in experimental pools. We also investigated the indirect effects of gudgeon, dace and chub on the growth of ayu. 2. We chose the four fish species across a continuum of omnivory. Ayu fed mainly on benthic algae, and gudgeon predominantly on invertebrates. Dace and chub fed on both algae and invertebrates, but dace preyed on invertebrates more than chub. 3. The presence of gudgeon, dace and chub reduced the number of algal-feeding invertebrates and increased the algal biomass through trophic cascades. Consequently, ayu growth rate over the experimental period in pools with one of the three fish species was 25.9-44.1 times greater than the growth rate in pools with only ayu. 4. The positive indirect effect on ayu growth was large for gudgeon and dace and small for chub, whereas the addition of ayu reduced ayu growth considerably due to intraspecific competition. 5. The relative intensity of indirect effects on ayu growth through trophic cascades was predictable from the food overlap between ayu and the other fishes. However, the similar strength of indirect effects by gudgeon and dace that fed differently on algae and invertebrates suggests that feeding behaviour, prey preference and trait-mediated indirect interactions were also important in the prediction.     

Effects of experimental duration and density of Japanese dace <Emphasis Type="Italic">Tribolodon hakonensis</Emphasis> on the strength of trophic cascades on benthic algae

To analyze density-mediated indirect effects through trophic cascades caused by Japanese dace Tribolodon hakonensis on the algal biomass and growth of ayu Plecoglossus altivelis altivelis, (freshwater fish belonging to Salmoniformes), I introduced zero, five, ten, and 20 dace with and without ayu into experimental pools. Ayu fed predominantly on benthic algae, whereas dace fed on both invertebrates and algae. The percentage of algae in the food content of dace was correlated with the number of dace in a pool. In the experiment lasting 8 days, trophic cascades on algae were not clear. In contrast, when the experimental duration was extended to 20 days, the number of gastropods and ephemeropteran nymphs was negatively correlated with the number of dace, consequently increasing the algal biomass and ayu growth through trophic cascades. Compared with pools with five dace, the presence of ten and 20 dace had similar gross effects on the number of algivorous invertebrates and algal biomass and greater gross effects on the ayu growth. The significant increase in ayu growth in pools with ten and 20 dace strongly suggests that small positive effects on algal growth by dace accumulated and extended to ayu during a 20-day experimental period. These results indicate the importance of analyzing interactions between the density and behavior of the top predator and the experimental duration on the intensity of trophic cascades.     

Experimentally induced foraging mode shift by sympatric charrs in a Japanese mountain stream

Foraging mode shifts may allow animals flexibility to adaptto a variety of ecological conditions. Theory holds that ectothermssuch as fishes should shift from ambush to active search modeswhen prey density declines, to maintain a minimum encounterrate. Salvelinus malma (Dolly Varden) sympatric withS. leucomaenis(white-spottedchirr) in a northern Japan mountain stream were observed toshift from ambushing drifting invertebrates at fixed focal pointsto picking benthic invertebrates from the substrate along circuitousroutes throughout pools, as drift declined throughout summer.Experimentally reducing drift in two pools during early summerwhen all charr were drift foragers caused subordinate malmato make marked shifts from drift to benthic foraging within1 h, whereas nearly all leucomaenis remained drift foragers.Some charr of both species responded to reduced drift by emigratingfrom pools. Combined results of three replicate experimentsrevealed that malma shifted to benthic foraging when their rateof drift forays was reduced below a specific threshold, therebyupholding predictions of the theory. Adaptive foraging modeshirts may promote coexistence of these congeneric charrs inreaches of Japanese mountain streams     

The Importance of Egg Size and Social Effects for Behaviour of Arctic Charr Juveniles

Early behaviour can determine food intake and growth rate with important consequences for life history and survival in fishes. Egg size is known to affect growth rate of young Arctic charr but its influence on the development of behaviour is poorly documented. It is believed that egg size influence on growth and potentially on the behaviour of young fish decreases over time, minimized by the effects of social factors. Shortly after first feeding, we examined differences in mobility and foraging of Arctic charr in relation to egg size and social environment. The behaviour of juveniles from small and large eggs was compared five times over the course of development and in three different experimental settings: long‐term isolation (isolation before hatching), short‐term isolation vs. group rearing and mixed size group vs. homogeneous size groups. Egg size affected foraging behaviour and mobility of fish: fish coming from large eggs were more mobile and foraged more than fish coming from small eggs. Social environment affected foraging behaviour, mobility and space use: fish in a group were more mobile, foraged more and responded faster to food delivery than isolated fish. The interaction of egg size and social effects was seen primarily in foraging activities but did not affect mobility or space use. Large fish in groups foraged more than the three other groups: large fish in isolation, small fish in groups and small fish in isolation. Agonistic behaviour was rarely observed and there was no significant effect of group composition on agonistic behaviour. We discuss the importance of egg size and social effects at early stages of development with a focus on the evolutionary ecology of Arctic charr.     

Intra- and interspecific dominance hierarchies and variation in foraging tactics of two species of stream-dwelling chars

Aggressive interactions, foraging behavior, habitat use and diet were studied in sympatric populations of white-sported char,Salvelinus leucomaenis, and Dolly Varden,Salvelinus malma, in a Japanese mountain stream. Underwater observations on individuals of both species revealed two distinct behavioral regimes: aggressive drift foragers and non-aggressive benthos foragers. Aggressive drift foragers defended partial territories around focal points from which they made forays to capture invertebrates drifting in the water column. Non-aggressive benthos foragers cruised around and beneath cobble in large foraging ranges that overlapped each other. Intra- and interspecific, size-dependent dominance hierarchies were recognized among aggressive drift foragers, whereas non-aggressive benthos foragers showed no such relationships. Terrestrial invertebrates were the most abundant prey in the diets of drift foragers, whereas a very small proportion of the diet of benthos foragers was made up of these taxa. Benthos foragers showed more complex diet composition than drift foragers. These results suggest that non-aggressive benthos foragers may avoid not only interference but also exploitative competition by using alternative foraging tactics. The proportion of drift foragers to benthos foragers among white-spotted char was more than 35 times that among Dolly Varden. The significant difference in the proportion of each species using the two types of foraging strategy results in interspecific food segregation in sympatric populations.     

Resource utilization by bull char and cutthroat trout in a mountain stream in Montana,U.S.A.

Resource utilization of sympatric populations of bull char,Salvelinus confluentus, and west-slope cutthroat trout,Oncorhynchus clarki lewisi, were studied by underwater observations of foraging behaviour and microhabitat use, and dietary analysis in a mountain stream of the Flathead River Basin, northwest Montana, U.S.A. Nearly 70% of bull char were categorized as benthic foragers, which moved constantly and captured prey primarily from the streambed, while all cutthroat trout were drift foragers, which held relatively fixed focal points in the midwater layers of pools during foraging. The composition of stomach contents was markedly different between the two species. Bull char fed primarily on baetid mayflies captured from the benthos or drift, whereas cutthroat trout ate primarily terrestrial invertebrates. The species also used different microhabitats. Bull char held positions close to the streambed and rarely strayed far from overhead cover, whereas cutthroat trout held focal points farther above the bed and far from overhead cover. Dietary segregation between these two salmonids appeared to result not only from differences in foraging tactics but also in the foraging microhabitats. Resource partitioning is considered to be one of important mechanisms allowing coexistence of these two stream salmonids.     

华南地区桑园和荒地两种生境中红火蚁觅食工蚁的食物组成

【目的】红火蚁Solenopsis invicta Buren是世界最危险的有害入侵生物之一,2004年入侵我国华南地区,并给农林业安全、生态安全带来极大危害。调查并明确田间红火蚁觅食工蚁的食物种类及数量,不仅为评估红火蚁猎食对生态环境中节肢动物群落的影响,而且为红火蚁防治饵剂的改进提供科学依据。【方法】本研究采用蚁道剖析法,对华南地区桑园和荒地两种生境中红火蚁觅食工蚁搬运的固体残片取样和鉴定;依搬运工蚁及其固体残片尺寸进行测量分级并称重,以分析工蚁多态性与固体食物大小的关系;采用薄层色谱法和氨基酸分析法测定嗉囊液体样品的营养成分及含量。【结果】在华南地区桑园和荒地中,红火蚁觅食工蚁搬运回巢的固体食物包括固体的动物残片和植物种子,其中动物残片属于3门7纲21个类群(包括14个昆虫目),昆虫纲动物所占比例为总固体食物的45.53%~46.10%。工蚁偏好搬运长1.400±0.043 mm~2.306±0.063 mm和宽0.723±0.028 mm~1.261±0.051 mm的固体残片,其重量在0.203~0.413 mg之间。红火蚁觅食工蚁嗉囊液体由多种氨基酸、果糖和葡萄糖组成,在桑园和荒地采集到的红火蚁嗉囊液体样本中的氨基酸总含量分别为1 544.31 mg/L和861.48 mg/L,氨基酸种类分别为33种和32种,其中31种氨基酸为共有的。【结论】华南地区桑园和荒地中,红火蚁固体食物组成均以昆虫纲动物为主;参与固体残片搬运觅食蚁的80%属于中型工蚁,搬运较大型固体残片的大型工蚁仅占5%;红火蚁工蚁嗉囊液体含有丰富的氨基酸和单糖。     

Seasonal changes in stream salmonid population densities in two tributaries of a boreal river in northern Japan

We examined seasonal changes in population densities of stream salmonids (masu salmon Oncorhynchus masou, white-spotted charr Salvelinus leucomaenis, and rainbow trout O. mykiss) in two tributaries of the Shoro River, eastern Hokkaido, Japan. In one small tributary, water temperature was relatively high during the winter, and populations of salmon and trout increased through immigration at this time of the year, becoming dominant components of the salmonid assemblage; the density of charr in this stream decreased during the winter, but charr was dominant during the summer. In another medium-sized tributary, the water temperature fell to close to 0°C during the winter, and densities of salmon and charr decreased in this season, through emigration; trout were very rare in this stream. Seasonal patterns of stream salmonid densities vary among species and between localities, resulting in seasonal changes in species composition. For a comprehensive understanding of population processes, a whole-river survey across seasons will be necessary.     

Drift dispersal of mayfly nymphs in the presence of chemical and visual cues from diurnal drift- and nocturnal benthic-foraging fishes

1. The drift of Baetis thermicus nymphs in the presence of chemical, visual and hydrodynamic cues, considered individually and in combination, produced by different predatory fishes was examined experimentally in laboratory streams. Masu salmon ( Oncorhynchus masou ) and freshwater sculpin ( Cottus nozawae ) are typical drift- and benthic-foraging fishes, respectively.
2. Observations of fish swimming in the streams revealed differing diel periodicity between the species; sculpin were nocturnal foragers and salmon diurnal.
3. The drift rate of Baetis by night increased in the presence of chemical cues from sculpin, with other cues having no interactive effects. In contrast, the drift rate increased primarily in the presence of both chemical and, particularly, visual cues from salmon, although no additional effects were found for any non-visual cues. Visual cues could enable Baetis to assess precisely the predation risk from foraging salmon by day, whereas Baetis could not use visual cues to detect sculpin either at night, because of the low light intensity, or during the day, because of the low activity of sculpin at that time.
4. In natural streams, which are often inhabited by several predatory fish employing different modes of foraging, invertebrates may be able to precisely assess the risk and effectively to avoid predators by using cues unique to each.     

Individuals in foraging groups may use vocal cues when assessing their need for anti-predator vigilance

Many studies of social species have reported variation in the anti-predator vigilance behaviour of foraging individuals depending on the presence and relative position of other group members. However, little attention has focused on how foragers assess these variables. It is commonly assumed that they do so visually, but many social species produce frequent calls while foraging, and these 'close' calls might provide valuable spatial information. Here, we show that foraging pied babblers (Turdoides bicolor) are less vigilant when in larger groups, in the centre of a group and in closer proximity to another group member. We then show that foragers are less vigilant during playbacks of close calling by more individuals and individuals on either side of them when compared with calls of fewer individuals and calls on one side of them. These results suggest that foragers can use vocal cues to gain information on group size and their spatial position within a group. Future studies of anti-predator vigilance should consider the relative importance of both visual and vocal monitoring of group members.     

Resource patch size and flexible foraging in white-faced capuchins (Cebus capucinus)

White-faced capuchins (Cebus capucinus)on Barro Colorado Island, Panama, have a flexible foraging strategy. Typically, foraging party size is small and individuals feed dispersed from one another. When seasonal fruiting of large volume trees occurs, the majority of the group forages simultaneously. As C. capucinusdo not display a rigorous dominance structure and there are few indications that individuals or coalitions monopolize food patches,individuals are expected to display scramble strategies instead of high frequencies of contest competition. I recorded foraging party size (simultaneous foragers), the total number of animals to feed successively, and the diameter at breast height (DBH) of fruit trees used in two habituated troops. Individuals in each group spent a substantial amount of time — 65 and 48% of foraging time for each group — foraging in party sizes of one. Monkeys predominantly foraged alone in small trees (0- to 20- cm DBH), successively in medium trees (21- to 60- cm DBH), and simultaneously in large trees (>61- cm DBH). They used small trees more frequently than all other tree classes. In medium-sized trees, although fruit was plentiful, space was limited. In these trees Cebusforaged successively. In large-volume trees, space and fruit were abundant and several individuals fed together. As the DBH of fruiting trees increased, the average foraging party size increased exponentially. Cebus capucinusat Barro Colorado Island modify their foraging party size to adapt to the seasonal patterns of fruit production.     

Intraspecific aggression in rosyside dace, a drift-feeding stream cyprinid

Individual rosyside dace Clinostomus funduloides in a semi-natural, artificial stream displayed substantial differences in their aggressiveness and could be classified as: (1) non-aggressive (NA, 18 of 30 rosyside dace), (2) moderately aggressive (MA, 9 of 30) and (3) highly aggressive (HA, 3 of 30). Rosyside dace groups, however, did not exhibit linear dominance hierarchies and fish size was only weakly correlated with the number of aggressive acts performed per individual. Small rosyside dace (<56 mm L_F) were always non-aggressive, but larger fish were present in all three aggression classes. The difference in size between the contestants was significantly, although not very strongly, correlated with the probability of winning an agonistic interaction (r ²=0·39). Aggressive rosyside dace may have ultimately gained higher fitness than less aggressive ones. HA individuals occupied the upstream-most position within foraging groups significantly more often than other rosyside dace. This location should be the most profitable one because its occupant will be the first to encounter prey. HA rosyside dace also occupied significantly higher focal velocities that were closer to energetic optima than MA and NA ones. They also had greater foraging rates and were less solitary than less aggressive fish, but these differences only were significant at the P=0·066 and P=0·081 level, respectively. Finally, HA fish performed significantly more aggressive acts and feedings backwards than other individuals. Despite these differences, the effects of intraspecific aggression in rosyside dace appeared less substantial than those that have been observed in stream salmonids.     

Food habit divergence between white-spotted charr and masu salmon in Japanese mountain streams: circumstantial evidence for competition

 The diet and foraging microhabitat of white-spotted charr, Salvelinus leucomaenis, were compared between mountain stream reaches where it occurs with (sympatric) and without (allopatric) masu salmon, Oncorhynchus masou masou, a potential competitor, to examine the evidence for interspecific competition between these fish, which commonly co-occur in Japanese mountain streams. In three streams examined, the similarity between the diets of allopatric charr and salmon was much greater than that between the diets of sympatric charr and salmon. Both allopatric charr and sympatric salmon intensively utilized terrestrial invertebrates among stream drifts (52%–65% of the diet), whereas this prey category made up only an intermediate portion of sympatric charr diets (11%–29%). Examination of available prey composition in stream drifts showed that the consumption of terrestrial invertebrates by allopatric charr and sympatric salmon was approximately twice as much as that by sympatric charr. The presence of salmon, a potential competitor, may alter the diet of charr in the sympatric reaches. Charr holding focal points closer to the streambed were considered less efficient than sympatric salmon in their utilization of terrestrial invertebrates drifting primarily on the stream surface, although the foraging microhabitat of the charr was not influenced by the salmon. The mechanisms responsible for the dietary divergence between sympatric charr and salmon are probably the consequence of scramble competition over terrestrial invertebrates drifting on the stream surface. Received: January 21, 2002 / Accepted: November 19, 2002 Acknowledgments We thank Y. Tokuda, T. Takasu, Y. Kaneda, H. Jyoya, and H. Aoe for their assistance. This work was partly supported by funding through the Takara Harmonist Fund by the Takara Syuzo Co. Ltd. and the Japan Ministry of Education, Science, Sport and Culture (grants 09NP1501 and 11440224). Correspondence to:H. Miyasaka     

Competition in a group of equal foragers

Abstract Using techniques from renewal process theory, we build a stochastic model for gain accumulation in a group of equal competitors foraging in a patchy environment. The model for gain of the individuals is based on the waiting times between subsequent prey encounters by the group. These waiting times depend on the number of foragers in the group. A single parameter of this dependency encompasses a variety of foraging scenarios, from co-operation to scramble. With constant patch size, correlations between gains of any pair of foragers are negative. This dependency is most intense in small groups. Increased variation in patch size makes correlations in gains between group members positive irrespective of the group size. For a solitary forager, variance in gain approaches zero with increasing time in the patch. For an individual member in a group, variance grows monotonically. Thus, depending on the patch departure rule controlling the time to be spent in the patch, solitary foragers may have a smaller variance in gain than members in a group. As solitary foragers also potentially harvest all prey in the patch, it is hard to believe that grouping behavior would evolve solely on the basis of foraging.     

Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

Tolerance of understory plants subject to herbivory by roe deer

Classical foraging theory states that animals feeding in a patchy environment can maximise their long term prey capture rates by quitting food patches when they have depleted prey to a certain threshold level. Theory suggests that social foragers may be better able to do this if all individuals in a group have access to the prey capture information of all other group members. This will allow all foragers to make a more accurate estimation of the patch quality over time and hence enable them to quit patches closer to the optimal prey threshold level. We develop a model to examine the foraging efficiency of three strategies that could be used by a cohesive foraging group to initiate quitting a patch, where foragers do not use such information, and compare these with a fourth strategy in which foragers use public information of all prey capture events made by the group. We carried out simulations in six different prey environments, in which we varied the mean number of prey per patch and the variance of prey number between patches. Groups sharing public information were able to consistently quit patches close to the optimal prey threshold level, and obtained constant prey capture rates, in groups of all sizes. In contrast all groups not sharing public information quit patches progressively earlier than the optimal prey threshold value, and experienced decreasing prey capture rates, as group size increased. This is more apparent as the variance in prey number between patches increases. Thus in a patchy environment, where uncertainty is high, although public information use does not increase the foraging efficiency of groups over that of a lone forager, it certainly offers benefits over groups which do not, and particularly where group size is large.     

Foraging strategy and predator avoidance behaviour: an intraspecific approach

Relationships between predator avoidance behaviour (scanning and flocking) and foraging were studied in Calidris alpina, to test predictions regarding the effect of foraging techniques on such behaviours. The scanning hypothesis predicts that individuals with a tactile hunting technique and individuals with a visual hunting technique (both continuous searchers) do not differ in any variable related to scanning behaviour. The flocking hypothesis predicts that visually hunting individuals witl tend to form smaller flocks than tactile-foraging individuals. The two continuous feeding strategies did not differ among individuals in vigilance rate, nor in vigilance time or mean scan duration. However, with respect to flocking behaviour, visual foragers differed from tactile foragers in foraging flock size. The relationships between flocking behaviour and foraging strategy are discussed. The pattern found at the intraspecific level are the same as those found at interspecific level.     

Task-specific expression of the foraging gene in harvester ants

In social insects, groups of workers perform various tasks such as brood care and foraging. Transitions in workers from one task to another are important in the organization and ecological success of colonies. Regulation of genetic pathways can lead to plasticity in social insect task behaviour. The colony organization of advanced eusocial insects evolved independently in ants, bees, and wasps and it is not known whether the genetic mechanisms that influence behavioural plasticity are conserved across species. Here we show that a gene associated with foraging behaviour is conserved across social insect species, but the expression patterns of this gene are not. We cloned the red harvester ant (Pogonomyrmex barbatus) ortholog (Pbfor) to foraging, one of few genes implicated in social organization, and found that foraging behaviour in harvester ants is associated with the expression of this gene; young (callow) worker brains have significantly higher levels of Pbfor mRNA than foragers. Levels of Pbfor mRNA in other worker task groups vary among harvester ant colonies. However, foragers always have the lowest expression levels compared to other task groups. The association between foraging behaviour and the foraging gene is conserved across social insects but ants and bees have an inverse relationship between foraging expression and behaviour.     

Effects of Chiton granosus (Frembly, 1827) and other molluscan grazers on algal succession in wave exposed mid-intertidal rocky shores of central Chile

Molluscan grazers can have important effects on the abundance, colonization rates, and successional pathways of algal assemblages and the entire intertidal community. In general, early successional algae are more readily consumed than corticated algae and kelps, which usually get established later in the community succession. To generalize, however, the effect of different grazers on algal assemblages must be examined on different coasts and under different scenarios. This information could help us understand the mechanisms of ecosystem processes and situations in which general models do not apply. Along the coast of Chile, humans harvest large keyhole limpets, which seem to be the only invertebrate grazers capable of controlling the dominant corticated alga Mazzaella laminarioides, a canopy-forming species that can cover extensive areas of the mid intertidal zone. In this scenario, where large limpets are harvested, the overall effects of the diverse molluscan assemblage of limpets, chitons and snails on algal succession and on corticated algae in particular are not clear. We conducted a 26-month-long experiment to evaluate the effects of molluscan grazers on mid-intertidal algal succession and to isolate the effects of Chiton granosus, the most conspicuous member of the assemblage at these tidal elevations. At sites heavily impacted by humans the molluscan grazer assemblage had strong negative effects on colonization and abundance of green algae such as ulvoids and Blidingia minima. In doing so, the grazer assemblage had a strong negative indirect effect on the establishments of chironomid fly larvae, which were only observed on green algal mats and rarely on bare rock. No significant effects were detected on epilithic microalgae, and effects on sessile invertebrates were highly variable over space and time. C. granosus also had significant negative effects on green algae but did not account for the total grazing pressure exerted by the guild. Limited foraging excursions (ca. 35 cm) from refuges and moderate site (crevice) fidelity in this species may contribute to the patchiness in green algal distribution observed in the field. Nearly 13 months after rock surface were experimentally cleared, M. laminarioides appeared in all experimental plots, but increased over three times faster in enclosures containing C. granosus than in exclosures plots or controls, suggesting that moderate levels of herbivory could actually facilitate the establishment of this alga in the succession and that the green algal cover found in the absence of grazers may delay its establishment.