Monoculture and mixed-species algal farms on a coral reef are maintained through intensive and extensive management by damselfishes

Algal farms maintained by different species of territorial herbivorous damselfishes vary in size, algal biomass, and species composition. To determine the factors that affect the structure of these farms, we compared farming behavior and intensity between two sympatric damselfish species, Stegastes nigricans (Lacepède) and S. obreptus (Whitley), which maintain two different types of farms in the Okinawa Islands, southern Japan. By weeding of indigestible algae and prompt exclusion of herbivorous fishes and grazing sea urchins, S. nigricans manage relatively small farms largely dominated by one algal crop species, Polysiphonia sp. In contrast, by delayed exclusion of herbivores, S. obreptus maintain larger farms invaded by diverse indigestible algae, such as Chondria sp. and Padina sp. As a result of intensive management, S. nigricans attains a higher algal biomass per area than does S. obreptus. This study provides a new perspective, as it suggests that damselfishes practise both intensive and extensive farming. The former results in small and highly productive monoculture farms of one palatable alga, while the latter leads to large, species-rich, mixed-culture farms.     

Farming behaviour of reef fishes increases the prevalence of coral disease associated microbes and black band disease

Microbial community structure on coral reefs is strongly influenced by coral–algae interactions; however, the extent to which this influence is mediated by fishes is unknown. By excluding fleshy macroalgae, cultivating palatable filamentous algae and engaging in frequent aggression to protect resources, territorial damselfish (f. Pomacentridae), such as Stegastes, mediate macro-benthic dynamics on coral reefs and may significantly influence microbial communities. To elucidate how Stegastes apicalis and Stegastes nigricans may alter benthic microbial assemblages and coral health, we determined the benthic community composition (epilithic algal matrix and prokaryotes) and coral disease prevalence inside and outside of damselfish territories in the Great Barrier Reef, Australia. 16S rDNA sequencing revealed distinct bacterial communities associated with turf algae and a two to three times greater relative abundance of phylotypes with high sequence similarity to potential coral pathogens inside Stegastes''s territories. These potentially pathogenic phylotypes (totalling 30.04% of the community) were found to have high sequence similarity to those amplified from black band disease (BBD) and disease affected corals worldwide. Disease surveys further revealed a significantly higher occurrence of BBD inside S. nigricans''s territories. These findings demonstrate the first link between fish behaviour, reservoirs of potential coral disease pathogens and the prevalence of coral disease.     

Geographic variation in the damselfish-red alga cultivation mutualism in the Indo-West Pacific

Background  

On coral reefs, damselfish defend their territories from invading herbivores and maintain algal turfs, from which they harvest filamentous algae. In southern Japan, intensive weeding of indigestible algae by Stegastes nigricans results in overgrowth by one filamentous alga, Polysiphonia sp. 1. Because this alga is highly susceptible to grazing and is competitively inferior to other algae, it survives only within the protective territories of this fish species, suggesting an obligate mutualism between damselfish and their cultivated alga. The wide distribution of damselfish species through the Indo-Central Pacific raises the question of whether this species-specific mutualism is maintained throughout the geographic range of the fish. To address this question, from all 18 damselfish species we conducted comprehensive surveys of algal flora within their territories throughout the Indo-West Pacific, and identified species of Polysiphonia using morphological examination and gene sequencing data.     

Effects of territorial damselfish on cryptic bioeroding organisms on dead Acropora formosa

Many damselfishes exclude other grazers from their territories and “farm” filamentous algae within their territories. In this study the indirect effect of damselfish territories on faunal composition and abundance of internal bioeroders of dead Acropora formosa (Dana, 1846) was investigated in territories of two damselfish species, Stegastes nigricans (Lacepède, 1802) and Plectroglyphidodon lacrymatus (Quoy and Gaimard, 1825). S. nigricans tends to be more protective and defend their territories more aggressively than P. lacrymatus. Newly killed branches of A. formosa were placed inside and outside damselfish territories, for 1 or 2 years, at a coral reef near Zanzibar, Tanzania. As predicted, the coral branches became covered with more filamentous algae in the S. nigricans territories than in the controls, with intermediate levels in the P. lacrymatus territories. Among the internal bioeroding fauna, polychaetes were by far the most common group. In total, there were significantly more borers in the first year than the second, which was mainly due to a high abundance of sabellids. Furthermore, sabellids were significantly more abundant in control areas and in the P. lacrymatus territories compared to the S. nigricans territories. However, many other genera showed the opposite pattern, with more polychaetes in the fish territories compared to the controls. There was also a clear difference in assemblage structure between S. nigricans territories and controls. Thus, we found strong effects of whether a piece of coral was placed inside or outside a damselfish territory on the abundance of many of the bioeroding taxa. We discuss multiple reasons for these indirect effects of the territories, including that deposit feeding bioeroders may benefit from the dense algal turf found inside the territories whereas suspension feeding bioeroders may benefit from substrate with less filamentous algae found outside territories. Considering our results in the context of the large areas of coral reefs that typically are defended as territories by damselfishes, these fish are likely to have a considerable impact on the boring community of a coral reef.     

The food habits of an herbivorous fish (Oreochromis niloticus Linn.) in Lake Awasa,Ethiopia

The natural food ofOreochromis niloticus in Lake Awasa was studied from the stomach contents of 10 fish measuring 18–32 cm (standard length) monthly from August 1984 to April 1986.Chroococcus, Oscillatoria andBotryococcus were found to be important food items in the diet. Animal foods were observed on rare occasions and these were mainly Rotifera. Blue-green algae as a group contribute about 28.1% of the ash free dry weight (AFDW) whereas diatoms and green algae (excludingBotryococcus) make 2.31% and 2.39% of AFDW, respectively.Botryococcus, detritus and other algae comprised 67.2% of AFDW and it was suspected that more than half of this was contributed byBotryococcus. Blue-green algae, which were especially abundant in the diet during the dry season, November–February, are nutritionally most important. Seasonal variation in algal species composition could influence the quality of food ingested by the fish.     

Territorial behavior of threespot damselfish (Eupomacentrus planifrons) increases reef algal biomass and productivity

Synopsis Algal growth and damselfish (Eupomacentrus planifrons) territories were studied in two reef habitats at Discovery Bay, Jamaica. Damselfish territories were contiguous in the reef flat (0 to 2.5 m), where the algal composition and biomass varied from territory to territory. In contrast, on the lower reef terrace (22 m), damselfish territories were often spatially segregated. While the algal composition of the territories was more uniform on the reef terrace, the total algal biomass was lower than in the territories on the reef flat. Damselfish are largely herbivorous, and they defend their territories against most intruding fish, including a number of herbivorous species. Areas of the reef terrace outside of damselfish territories were heavily grazed by herbivorous fishes and contained only small quantities of non-crustose algae.The reef terrace territories were characterized by a multispecific turf of algae (greens, blue-greens, and reds) covering the Acropora cervicornis framework and by the leafy, brown alga, Lobophora variegata. A rapid reduction in the biomass of brown algae and filamentous algae was noted when damselfish were permanently removed from their territories. Only calcified, encrusting algae — plants apparently somewhat undesirable as fish food sources — would be common on the terrace zone of this reef if damselfish territories were absent. Damselfish territoriality may significantly influence the dynamics of some reefs by increasing the biomass of the algal turf thereby increasing; reef productivity. Since blue-green algae, potential nitrogen fixers, occur in these algal turfs, the fish may also be indirectly affecting reef nutrition.     

The survival of <Emphasis Type="Italic">Sicyopterus stimpsoni</Emphasis>, an endemic amphidromous Hawaiian gobiid fish,relies on the hydrological cycles of streams: evidence from changes in algal composition of diet through growth stages fish

Gut contents of larval, juvenile, and adult specimens of the Hawaiian gobiid fish Sicyopterus stimpsoni were examined to catalog the algal flora ingested by this species. The developmental stages of S. stimpsoni examined represented hallmark points in the fish’s life cycle corresponding with major migratory and metamorphic transitions. The algal flora was dominated by diatom species and shifted from taxa representative of a marine, planktonic community in larval fish to a freshwater, benthic community in juvenile and adult fish. This change in diet corresponds with the migration of larval fish to freshwater streams just prior to juvenile development in which rapid modification in mouth anatomy makes ingestion of planktonic algal species difficult. Benthic diatoms from juvenile and adult fish assemblages represented multiple genera that live in a narrow set of environmental conditions. These algae grow during a specific period in the development of the benthic algal community in Hawaiian streams. This suggests a highly specialized dietary behavior that depends heavily on continually restarting the benthic algal successional pattern, which appears to be regulated by the hydrological cycles of streams on the island.     

Terrestrial detritus supports the food webs in lowland intermittent streams of south‐eastern Australia: a stable isotope study

• 1 Large amounts of terrestrial detritus enter many low‐order forested streams, and this organic material is often the major basal resource in the metazoan food webs of such systems. However, despite their apparently low biomass, algae are the dominant food of organisms in a number of aquatic communities which conventionally would have been presumed to be dependent on allochthonous detritus, particularly those in the tropics and also in lowland intermittent streams in arid Australia.
• 2 The dual stable isotope signatures (δ¹³C and δ¹⁵N) of potential primary food sources were compared with the isotopic signatures of common aquatic animals in lowland intermittent streams in south‐eastern Australia, in both spring and summer, to determine whether allochthonous detritus was an important nutritional resource in these systems. The isotopic signatures of the major potential allochthonous plant food sources (Eucalyptus, Phalaris and Juncus) overlapped, but were distinct from algae and the dominant macrophytes growing in the study reaches. The isotopic signatures of biofilm were more spatially and temporally variable than those of the other basal resources.
• 3 Despite allochthonous detritus having relatively high C : N ratios compared to other potential basal resources, results from isosource mixing model calculations demonstrated that this detritus, and the associated biofilm, were the major energy sources assimilated by macroinvertebrate primary consumers in both spring and summer. The importance of these energy sources was also reflected in animals higher in the food web, including predatory macroinvertebrates and fish. These resources were supplemented by autochthonous sources of higher nutritional value (i.e. filamentous algae and macrophytes, which had relatively low C : N ratios) when they became more prolific as the streams dried to disconnected pools in summer.
• 4 The results highlight the importance of allochthonous detritus (particularly from Eucalyptus) as a dependable energy source for benthic macroinvertebrates and fish in lowland intermittent streams of south‐eastern Australia. This contrasts with previous stable isotope studies conducted in lowland intermittent streams in arid Australia, which have reported that the fauna are primarily dependent on autochthonous algae.

     

Nutritional value of detritus and algae in blenny territories on the Great Barrier Reef

The salariin blenny, Salarias patzneri, primarily ingests detrital aggregates and inorganic sediment, with small quantities of filamentous algae. Samples of particulate matter <125 μm and filamentous algae were collected from the territories of S. patzneri and the nutritional value of these resources assessed using organic content and protein/energy ratios calculated from protein, carbohydrate and lipid content. Samples were collected throughout the day, during both summer and winter, to enable temporal comparisons of nutritional value. The particulates <125 μm, which were predominantly amorphic detritus, accounted for 41±2% and 44±3% of the organic matter in the epilithic algal matrix (EAM) during the summer and winter, respectively, whilst filamentous algae accounted for 37±3% and 41±3% of the organic matter in the summer and winter, respectively. Carbohydrate and lipid concentration in the algal samples was significantly greater than in particulate matter <125 μm, although no significant difference was detected in protein levels of algal samples and particulates <125 μm. Consequently, the protein/energy ratio of particulates <125 μm in the summer (11±1 mg kJ⁻¹) and winter (10±1 mg kJ⁻¹) was similar to that of filamentous algae in the summer (9±1 mg kJ⁻¹) and winter (8±1 mg kJ⁻¹). These results suggest detrital aggregates are at least comparable to filamentous algae as a nutritional resource. The large contribution of high quality detritus to organic matter in the epilithic algal matrix collected from S. patzneri territories throughout the day and between seasons provides strong evidence that detritus is a valuable component of small territorial fish diets and is an integral part of coral reef food webs.     

Use of a flooded salt marsh habitat by an endangered cyprinodontid fish (<Emphasis Type="Italic">Aphanius iberus</Emphasis>)

We report the first data on the use of occasionally inundated habitats in a Mediterranean coastal lagoon by the Spanish toothcarp (Aphanius iberus), a cyprinodontid fish in danger of extinction. During a flooding period, we sampled the fish population and the macroinvertebrate community in three contrasting habitats: mats of green algae, open water, and a habitat dominated by glasswort (Salicornia patula) that is occasionally inundated. The three habitats displayed strong variation in density and species composition of invertebrates in the water column and the benthos. In general, algal mats had higher invertebrate biomasses, but glasswort had higher diversity of organisms, in part of terrestrial origin. The density of toothcarp was very low in the open water. The habitat that is occasionally inundated (glasswort) significantly had the highest density of mature toothcarp, while immature fish were similarly abundant in the glasswort and algal mat habitats. Condition (weight–length relationship) and total food biomass in the gut contents of immature toothcarp was significantly higher in algal mats than in glasswort, whereas there were no such differences for mature fish. Therefore, the occasionally flooded habitat (glasswort) was positively selected by large mature toothcarp but seemed a disadvantageous habitat for immature individuals. Handling editor: L. Mauricio Bini.     

论白鲢的食物问题

本文系作者结合自己的实践和国内外的文献资料,对白鲢食谱、食物的选择性和消化性等问题进行探讨,指出必须从新的角度来理解白鲢的食物问题和衡量养鱼池水的肥度。       

Drifting algal mats as an alternative habitat for benthic invertebrates: Species specific responses to a transient resource

Patchy occurrences of benthic drift algae (i.e. loose lying macroalgal mats) may increase habitat complexity on normally bare soft bottoms, but at the same time, extensive amounts of drifting algal mats are known to stress the benthic fauna. This paper presents results of the first detailed study of the fauna associated with drift algal mats in the northern Baltic Sea. In order to assess the importance of drifting algae as an alternative habitat for benthic fauna, benthic drift algal mats were sampled on shallow (2-9 m) sandy soft bottoms in the outer archipelago of the ?land Islands (Finland). Species composition, abundance and biomass of the macrofauna associated with algal mats were recorded. The results show that drifting algae at times can harbour very high abundances of invertebrates (up to 1116 individuals/g algal dryweight), surpassing invertebrate densities recorded in seagrass communities. The algal fauna varied between sites and over time, and factors such as ambient benthic fauna, exposure to wind-wave disturbance, depth, and algal coverage and condition influenced the invertebrate community composition of the algal mats. Abundance increased while individual biomass of the animals decreased over time (summer season; July-October). A series of laboratory experiments were conducted in order to test the ability of a few important benthic species to move up into, and survive in a drifting algal mat. Macoma balthica, Hydrobia spp., Nereis diversicolor and Bathyporeia pilosa were used in the experiments, and significant differences in their survival and mobility within drifting algae were recorded. This study shows that benthic species differ significantly in their ability to utilise the algal mats, with mainly opportunistic and mobile taxa such as Hydrobia spp., Chironomidae and Ostracoda benefiting from the algae, whereas infaunal species such as M. balthica and B. pilosa are negatively affected. The occurrence of eutrophication induced drifting macroalgal mats has increased significantly during the last decade in the northern Baltic Sea. Hence, the importance of drifting algae as a stress factor and as an alternative habitat for benthic fauna increases.     

Effects of herbivory and the season of disturbance on algal succession in a tropical intertidal shore,Phuket, Thailand

The effects of herbivory and the season of disturbance on species composition and algal succession were experimentally tested at a tropical intertidal shore, Phuket Island, Thailand. Dead coral patches were cleared of all organisms during both the dry and rainy seasons in order to study the effects of season on algal succession and cages were set up to exclude fish herbivory. Algal succession in this intertidal habitat showed a simple pattern and took a year from the early Ulva paradoxa C. Agardh stage to the late Polysiphonia sphaerocarpa Børgesen stage. The abundance of algae during succession was under the influence of seasonal change. U. paradoxa reproduced and recruited throughout the year. Caging effects did not apparently influence algal abundance, perhaps because resident herbivorous damselfishes excluded other herbivores from their territories and maintained their algal “farms”. Unexpectedly, the percent cover of Ulva in the caged plots was lower than in uncaged plots. This pattern may indicate that caging excluded damselfishes only, but allowed small herbivores that consumed substantial amounts of soft filamentous algae in the cages.     

Predation by juvenile Platichthys flesus (L.) on shelled prey species in a bare sand and a drift algae habitat

Due to increasing eutrophication of the coastal Baltic waters, drifting algae are a common phenomenon. Drifting algal mats accumulate on shallow sandy bottoms in late summer and autumn, and affect the ambient fauna. Juvenile flounder, Platichthys flesus, utilize these habitats during their first few years. They feed on benthic meio- and macrofauna; part of their diet consists of shelled species, such as Ostracods, and juvenile Hydrobia spp. and Macoma balthica. Earlier studies have shown that up to 75% of ostracods and 92% of hydrobiids survive the gut passage of juvenile flounder, while all M. balthica are digested by the fish. We conducted laboratory experiments to study how the shelled prey responded to a drift algal mat, and the predation efficiency of juvenile P. flesus on these prey species on bare sand and with drifting algae (50% coverage). Hydrobia spp. utilized the drift algae as a habitat and, after 1 h, 50% had moved into the algae; ostracods and M. balthica were more stationary and, after 96 h, only 23 and 12%, respectively, were found in the algae. For the predation efficiency of P. flesus, a two-way ANOVA with habitat (algae, bare sand) and predation (fish, no fish) as factors revealed that both algae and predation affected negatively the survival of all three prey species. The algae, thus, affected the predation efficiency of juvenile P. flesus and the consumption of prey was much reduced in the algal treatments compared to the bare sand. This was due probably to increased habitat complexity and the ability of prey, especially hydrobiids, to use the algal mat as a refuge. Altered habitat structure due to drift algae, together with the resultant changes in habitat (refuge) value for different prey species, may profoundly change the structure of benthic communities.     

Effects of territorial damselfish on an algal-dominated coastal coral reef

Territorial damselfish are important herbivores on coral reefs because they can occupy a large proportion of the substratum and modify the benthic community to promote the cover of food algae. However, on coastal coral reefs damselfish occupy habitats that are often dominated by unpalatable macroalgae. The aim of this study was to examine whether damselfish can maintain distinctive algal assemblages on a coastal reef that is seasonally dominated by Sargassum (Magnetic Island, Great Barrier Reef). Here, three abundant species (Pomacentrus tripunctatus, P. wardi and Stegastes apicalis) occupied up to 60% of the reef substrata. All three species promoted the abundance of food algae in their territories. The magnitudes of the effects varied among reef zones, but patterns were relatively stable over time. Damselfish appear to readily co-exist with large unpalatable macroalgae as they can use it as a substratum for promoting the growth of palatable epiphytes. Damselfish territories represent patches of increased epiphyte load on macroalgae, decreased sediment cover, and enhanced cover of palatable algal turf.     

Influence of diet on growth,food retention time,and gill ventilation rate of nymphs of Cloeon sp. (Ephemeroptera: Baetidae)

Early instar nymphs of the mayfly, Cloeon sp. were kept on an algal or a detrital diet at 20 ± 1 °C in the laboratory. A control group was not given any food. Nymphs kept on algae showed significantly (P = 0.001) higher growth both in terms of length and biomass than for those kept on detritus. None of the nymphs given algae or detritus died during the experiment, and 40 per cent of those fed algae and 20 per cent fed detritus reached maturity by the termination of the experiment. Food retention time as well as gill ventilation rate of nymphs feeding on algae were significantly (P = 0.001) higher than those feeding on detritus. It is probable that these two factors largely enabled the nymphs to survive as well as to grow and reach maturity, though relatively slowly, solely on a detrital diet. However, algae was clearly shown to be a superior food resource for Cloeon sp. nymphs.     

Unexpected prey of juvenile spotted scat (Scatophagus argus) near a wharf: The prevalence of fouling organisms in stomach contents

A knowledge of fish diets can contribute to revealing the trophic role and ecological function of species in aquatic ecosystems. At present, however, there are no efficient or comprehensive methods for analyzing fish diets. In this study, we investigated the diets of juvenile Scatophagus argus collected near a wharf in Daya Bay, China, by dissection and high‐throughput sequencing (HTS) using the 18S rDNA V4 region. Microscopy disclosed large amounts of bryozoans and unrecognizable detritus. In contrast, HTS analysis indicated that the fish diets were considerably more diverse than visual inspection suggested. After eliminating fish sequences, approximately 17,000 sequences from taxa in nine phyla (Ciliophora, Bryozoa, Annelida, Bacillariophyta, Chlorophyta, Arthropoda, Dinoflagellata, Tunicata, and Phaeophyta) were identified from the analysis of stomach contents. Twenty‐one food categories were identified, most of which (95.2%) were benthic fouling organisms that could easily be collected around wharfs. These consisted of bryozoans (31.9%), ciliates (45.7%), polychaetes (14.6%), and green algae (3.0%). Therefore, to adapt to anthropogenic habitat modification, the fish had probably shifted from planktonic to benthic feeding. The prevalence of fouling organisms in the stomachs of juvenile S. argus indicates that the fish have responded to habitat changes by widening their food spectrum. This adaptation may have increased their chances of survival. The fouling organisms that inhabit highly perturbed coastal ecosystems could represent a food source for animals at higher trophic levels. Our results accordingly suggest that human activity might significantly influence fish feeding behavior and material transfer along the food chain.     

The distribution and food habits of the dwarf African cichlid,Pelvicachromis pulcher in the River Sombreiro,Nigeria

In the River Sombreiro, Nigeria, the dwarf cichlid, Pelvicachromis pulcher was restricted to an upstream zone of cold, soft, acid freshwater. The sites nearest the source had smaller fish. Sex ratio (M:F) was 1:1 except in the most upstream site. Benthic algae, higher plant remains, detritus and copepods were the main food items.     

Food of an endangered cyprinodont (Aphanius iberus): ontogenetic diet shift and prey electivity

Synopsis We studied the ontogenetic diet shift and prey electivity of an endangered cyprinodontid fish endemic to the Iberian Peninsula, the Spanish toothcarp (Aphanius iberus). The toothcarp’s diet was omnivorous, dominated by harpacticoid copepods (Mesochra lilljeborgi and Tisbe longicornis), copepod nauplii and detritus. Diet composition varied greatly among habitats, depending on prey availability. In a rarely inundated habitat (glasswort), there was more consumption of the isopod Protracheoniscus occidentalis and the harpacticoid copepod Mesochra lilljeborgi, while in algal mats another harpacticoid (Tisbe longicornis), chironomid dipterans and invertebrate eggs were more important in diet. Although a benthic feeding habitat has previously been suggested, in our study the diet was based rather on water column organisms for both glasswort and algal mat habitats. There was also an ontogenetic diet shift, with an increase of mean prey length with fish length, clearly linked to a microhabitat change. Smaller fish showed positive electivity and greater reliance on planktonic prey (e.g. copepod nauplii, the harpacticoid copepods Mesochra lilljeborgi and Tisbe longicornis, the rotifer Brachionus plicatilis, and ostracods), while larger fish elected and preyed on more benthic organisms (e.g. Canuella perplexa, Mesochra rapiens, and ephydrid dipterans).     

Group foraging by the herbivorous cichlid fish,Petrochromis fasciolatus,in Lake Tanganyika

Group foraging of the algae eating cichlid,Petrochromis fasciolatus, was studied along a rocky shore in Lake Tanganyika, in an area occupied by the feeding territories of several herbivorous cichlid species. SolitaryP. fasciolatus or schools comprising a few members only were nearly always repelled from such territories, although larger schools (40–250 individuals) ofP. fasciolatus were able to overwhelm the defenses of territory owners and enable subsequent foraging. The frequency and duration of individual foraging sessions on the algal mats were positively correlated with group size, probably owing to individuals in large schools suffering fewer attacks from territory owners. Almost all of the foraging sites ofP. fasciolatus were inside the territories of other herbivorous cichlids, especially those ofNeolamprologus moorii (more than 80% of total), in which the algal density was more than 10 times that in territories of the other species.N. moorii territories included only 4% rocky substrate, indicating that schools ofP. fasciolatus selectively chose territories containing rich resources.