Are the oral teeth of combtooth bennies (Salariini: Teleostei) merely combs?

Careful analysis of gut contents has resulted in the reclassification of several species of grazing fish as detritivores, shifting the trophodynamic assignment of many prominent reef grazers. Combtoothed blennies, which are among the most numerous grazing fish of the Great Barrier Reef, have been shown to target the detrital component of the epilithic algal matrix (EAM). It has been suggested that blennies have specialized dental morphology that allows them to comb through fronds of algae, collecting detritus, while leaving the algal component intact. In this study, we analysed the capability of a common reef flat blenny, Salarias fasciatus, to remove algae by (i) examining oral morphology for evidence of wear and adaptations for abrasion, and (ii) a short-term EAM feeding experiment. Examination of S. fasciatus teeth with scanning electron microscopy (SEM) showed evidence of macrowear (changes in tooth height or shape), microwear (surface chips and striae), and tooth replacement that suggests abrasion on the substrate. Energy Dispersive X-ray Spectrometry (EDS) revealed that oral teeth incorporate mineral iron. When fed artificial substrata with a developed EAM layer S. fasciatus removed 57% of photosynthetic material and 38.5% of organic biomass. Although studies have found that blenny gut contents consist predominantly of detritus, blennies are still likely to contribute to the removal of algae on coral reefs.     

Multiscale habitat associations of detrivorous blennies (Blenniidae: Salariini)

The distribution and habitat associations of detrivorous blennies on a tropical coral reef were investigated at several spatial scales and compared with other fish that feed on the epilithic algal matrix to assess density and biomass contributions of small detrivorous fishes to these assemblages. At broad spatial scales total blenny abundance and biomass were highest on the tops of reefs exposed to prevailing winds. On the finer scale of microhabitat use, all species showed a preference for non-living corals, although the type of coral utilised differed between species. The high abundance of blennies on reef tops and non-living corals may be partially related to the quality and availability of detritus in these habitats. Comparisons of total blenny abundance and biomass with other territorial detrivores found that blennies accounted for approximately 60% of this functional group's density and 21% of their biomass on exposed reef tops. Overall, territorial detrivores were found to constitute approximately 37% of the density and 26% of the biomass of the detrivorous/herbivorous fish assemblage on exposed reef tops. Small detrivorous fish therefore represent a substantial proportion of fish assemblages that feed on epilithic algae and associated detritus on coral reefs.     

Interactions Between Sabre‐Tooth Blennies and Their Reef Fish Victims: Effects Of Enforced Repeated Game Structure and Local Abundance on Victim Aggression

The conditions under which humans benefit from contributing to a public good have attracted great interest; in particular the potential role of punishment of cheaters is hotly debated. In contrast, similar studies on other animals are lacking. In this study, we describe for the first time how the course of interactions between parasitic sabre‐tooth blennies (the cheaters) and their reef fish victims can be used to study both punishment and the emergence of public goods. Sabre tooth blennies (Plagiotremus sp.) sneak up from behind to bite off small pieces of scales and/or mucus from other fish. Victims regularly show spontaneous aggression as well as aggressive responses to blenny attacks. In a between species comparison, we tested how the probability of chasing a blenny is affected by (1) the option of avoiding interactions with a blenny by avoiding its small territory, and (2) variation in local abundance of conspecifics. We found that resident victim species are more aggressive towards blennies than visiting species. This difference persisted when we controlled for victim size and territoriality, suggesting that it is the enforced repeated game structure that causes residents to chase blennies. In residents, we also found a negative correlation between aggression towards blennies and local abundance, which suggests that the benefits of chasing are diluted with increasing local abundance. We discuss the implication of these results for future studies.     

Extreme dentition does not prevent diet and tooth diversification within combtooth blennies (Ovalentaria: Blenniidae)

The dentition of fishes can be quite striking and is often correlated with a specific diet. Combtooth blennies have long incisiform oral teeth, unlike most actinopterygians. It has been suggested that the long tooth morphology is an adaptation for detritivory, but given the diversity of diets (detritus, coral polyps, polychaetes, and pieces of other fishes), are blenny teeth indeed monomorphic? Or does tooth variation associated with diet still exist at this extreme? To explore tooth and diet diversification, we used a new phylogenetic hypothesis of Blenniidae, measured tooth shape, number, and mode of attachment, and quantified blenniid diet. The ancestral diet of blennies contained detritus and diversified into many different diets, including almost exclusively detritivory. Our results reveal a dental cline that may be constrained by tooth shape, but has not prevented diet diversification. Ancestral state reconstruction of tooth morphologies suggests that the ancestor of blennies had many unattached teeth and featured transitions to fewer attached teeth, with several transitions back to attached or unattached teeth. The dentition of blenniids is not monotypic; rather it is diverse and small changes in tooth shape are accompanied by changes in size, number, attachment, and often diet.     

Food habits of fishes in azostera marina bed at aburatsubo, central japan

To clarify the feeding habits of seagrass fishes, the gut contents of 31 fish species collected in aZostera marina bed at Aburatsubo central Japan, were examined. Ontogenetic changes in food preference were recognized in 12 species. In general, juveniles of the latter preyed mainly on small crustaceans (e.g., gammaridean amphipods) or planktonic animals (e.g., calanoid and cyclopoid copepods), subsequently changing to other prey items (e.g., hard-shelled animals) with growth. Cluster analysis based on dietary overlaps showed that the seagrass fish assemblage comprised seven feeding guilds (small-crustacean, planktonic-animal, large-crustacean, polychaete, fish, hard-shelled mollusc and detritus feeders). Of these, small-crustacean feeders and planktonic-animal feeders were the most abundantly represented, including juveniles of several species, which, when adult, transferred to other feeding guilds. On the other hand detritivores were represented by a single species.     

Trophic structure of coastal freshwater stream fishes from an Atlantic rainforest: evidence of the importance of protected and forest-covered areas to fish diet

The role of riparian forests in the functioning of aquatic ecosystems is well known, and they are recognized as an important food source for riverine fauna. This study investigates the trophic structure of coastal freshwater stream fishes from a large conservation area in an Atlantic rainforest using stomach content and food availability analyses. Four samples were collected from 19 sample sites. Fishes were caught with electrofishing. Prey were sampled with trays, Surber, traps, and electrofishing to evaluate the availability of food resources. The diets of 20 fish species were determined from the stomach contents of 1691 individuals. Terrestrial and aquatic insects and detritus were the most consumed items. Fish diet and prey availability were not seasonally dependent. A cluster analysis showed five trophic functional groups: terrestrial insectivores, aquatic insectivores, detritivores, carnivores, and omnivores. Insectivores predominated in species richness (60%), abundance (47%) and biomass (39%). Allochthonous and autochthonous items were found in similar proportions in the environment; however, allochthonous items were representative for insectivores and detritivores, whereas autochthonous items were important for primarily aquatic insectivores. The preference for certain insects by insectivorous fishes was associated with food selectivity rather than the availability of the resource and demonstrated the strong relationship between feeding behavior and food preference. The absence of seasonal variation in the diets of the fishes was possibly related to the consistent food supply. Our results confirm the role of the forest as a food provider for stream fishes, such as terrestrial insects and plant debris/detritus (also consumed by aquatic insects, which subsequently serve as food for fish), highlighting the importance of conserving the Brazilian Atlantic rainforests.     

Food habits of small fishes in a common reed <Emphasis Type="Italic">Phragmites australis</Emphasis> belt in Lake Shinji,Shimane, Japan

To clarify the feeding habits of reed fishes, the gut contents of 13 fish species collected in a Phragmites australis belt in Lake Shinji were examined. Six species showed ontogenetic and/or seasonal changes in food use patterns. Smaller individuals generally preyed on small planktonic items (e.g., calanoid and cyclopoid copepods) or small crustaceans (gammaridean amphipods), subsequently changing to other prey items (e.g., mysids and filamentous algae) with growth. The most important dietary items for the reed fish assemblage comprised planktonic copepods, gammaridean amphipods and mysids. However, the relative importance of these changed seasonally, gammaridean amphipods being the most important in autumn and winter, and planktonic copepods and mysids the most important in spring and in summer. Cluster analysis based on dietary overlaps showed that the reed fish assemblage comprised five feeding guilds (planktonic-copepod, mysid, gammaridean-amphipod, filamentous-algae, and detritus feeders). Of these, the three former guilds were the most abundantly represented, whereas detritivores were represented by a single species.     

Feeding patterns of the dominant benthic and demersal fish community in a temperate estuary

The diet and feeding relationships of the 12 most abundant benthic and demersal fish species of the Mondego Estuary, Portugal, were studied between June 2003 and May 2006. Fishes were caught during the night using a 2 m beam trawl. The stomach contents were analysed for prey identification, counting and weighing. According to prey importance in diets, three main feeding guilds were identified: (1) invertebrate feeders, (2) invertebrate and fish feeders and (3) plankton and invertebrate feeders. Besides these main feeding guilds, some fishes also presented fractions of algae and zooplankton in their stomach contents. The most abundant prey items were macroinvertebrates, with several polychaetes ( Nephtys spp., Capitellidae, Spionidae and Eunicidae), Corophium spp. and Crangon crangon among the dominant prey. Pomatoschistus spp. were the most preyed on fishes. Several fish species showed a tendency for a specialized diet, but almost all also showed some degree of opportunistic feeding by preying on other food resources. High diet overlap was found between some fish species, yet exploitative competition could not be concluded.     

Food and feeding habits of four species of juvenile mullet (Mugilidae) in a tidal lagoon in Ghana

Juvenile mugilid fishes, Liza falcipinnis, L. dumerilii, Mugil bananensis and M. curema , which enter the Elmina Lagoon in the Cape Coast District of Ghana, have a similar diet comprising mainly bacteria, diatoms, blue-green/green algae, protozoans, detritus and particulate organic matter. No seasonal changes in the diet and feeding activity were observed. The relative gut length (intestine to standard length ratio) and diet showed no significant changes with size of fish in all the species. All four species were diurnal feeders, but their peak feeding times differed. Interspecific competition for food was possibly limited by species preferences for substrate particles of particular size range and differences in feeding chronology.     

Importance of feeding strategies on the long-term success of fish invasions

This study assessed the feeding strategies of nine fish species in their native (Cuiabá River) and in an invaded basin (upper Paraná River) to identify trophic variables that may explain the success of these species in the new basin, over 30 years. The following predictions were analyzed: (i) species that display omnivorous or piscivorous diets in the native basin are favored in the invasion process over the long term, and (ii) specialist feeders are favored in the invasion process provided that their food items are highly available in the invaded area. These predictions were supported by the data; the species that were successful invaders had high trophic plasticity (omnivores), consumed a wide variety of food items from specific trophic guilds (piscivores), or if a species had a specialized diet, the resources demanded are abundant (detritivores). Thus, in a long-term perspective, the food resources used by these species are rarely limiting in aquatic ecosystems, and these feeding characteristics should be one of the key factors determining the colonization success of fishes. Understanding the factors that determine the success of invasive species in new areas is critical for developing management policies aimed at minimizing the impacts of biological invasions.     

Recruitment in the redlip blenny Ophioblennius atlanticus: is space limiting?

This study investigates pre-and post-recruitment characteristics of a population of the redlip blenny (Ophioblennius atlanticus) on a fringing reef in Barbados, W.I. Recruits were observed in 6 of the 8 months of the study, but 90% of all recruitment occurred within a single month. Monthly recruitment rate was not correlated with the monthly mortality of residents, suggesting that the rate at which space becomes available on the reef does not control the timing of blenny recruitment. Most recruitment occurred when adult blenny density was low, i.e. when most total space was available on the reef. Postrecruitment territory size of resident blennies was half that of pre-recruitment territory size, and was better predicted by fish size than it was before recruitment. Aggression by resident blennies increased with blenny density, and was primarily directed at recruiting conspecifics. These results suggest increased competition for territorial space after recruitment. Following the major recruitment pulse, the monthly percentage of recruits dying was correlated with density. This may indicate density-dependent mortality, but could result from the covariance of density with age. We suggest that whether reef fish populations are space-limited or recruitment-limited may vary between species and within species between locations.     

Multivariate analysis of feeding relationships of fishes in blackwater streams

Synopsis The autumnal diets of 23 fish species in 9 families from South Carolina blackwater streams were characterized by rank order statistics and analyzed by detrended correspondence analysis (DCA). DCA ordinations of fishes and their food items indicate two primary gradients. The first axis contrasts fine particle feeders and omnivores (Catostomidae, some cyprinids) eating algae, detritus, microcrustacea and bivalves, with predators(Anguilla, Esox, largeAmeiurus) on decapods and vertebrates. The second is a gradient from benthic feeders (percid darters,Noturus catfishes) with varied invertebrate diets, to taxa (Gambusia, some cyprinids and centrarchids) feeding at the surface. With increasing size, fishes shift to larger prey and increased surface feeding. Much of the trophic differentiation within the assemblage reflects diversification at generic and family levels. DCA is an effective method for summarizing trophic relations in diverse assemblages.     

Aggressive mimicry of the cleaner wrasse by Aspidontus taeniatus functions mainly for small blennies

The mimic blenny Aspidontus taeniatus Quoy & Gaimard is well known for its resemblance to the juvenile and adult cleaner wrasse Labroides dimidiatus (Valenciennes) in colour and shape. As various reef fishes including piscivores actively approach the cleaner wrasse to solicit cleaning by posing, two types of benefits have been suggested for this resemblance, that is, protective mimicry and aggressive mimicry. In aggressive mimicry, the mimic blenny is supposed to have considerable opportunities to bite the fin of deceived fishes when they pose, but some studies have confirmed that fin biting does not seem to be the main feeding tactic in the blenny in nature. Here, we examined the feeding tactics including fin biting by the mimic blenny in relation to its body size in a field observational survey in the coral reefs of Sesoko Island, Okinawa, Japan. The blenny was observed feeding mainly on four food items: the tentacles of Christmas tree worms, the mantle edges of boring clams, the demersal eggs in damselfishes’ nests and the fins of fishes. The feeding frequency by fin biting significantly decreased with body size, while that by egg predation significantly increased with body size of the blenny. When predating on eggs, the blenny was vigorously attacked by egg‐guarding fish, but often succeeded in raiding their nests by forming a feeding group. When feeding by fin biting, the blenny attacked prey fish without performing any cleaning. The ratio of fin biting was considerably higher in small‐sized blennies, suggesting reliance on this feeding tactic because of a difficulty in conducting a risky egg predation. Thus, our results suggest that the mimic blenny utilizes aggressive mimicry only when it is small as an alternative feeding tactic.     

Spatial Group Structure as Potential Mechanism to Maintain Cooperation in Fish Shoals of Unrelated Individuals

In public goods games, group members share the benefits created through individual investments. If benefits are shared equally, individual contributions readily become altruistic, and hence, a social dilemma arises in which group interests and individual interests oppose each other. However, contributions to public goods can be self‐serving if each investor gains a disproportionate benefit from its own contribution. This scenario may hold for our study system, the interactions of shoaling‐unrelated scalefin anthias Pseudanthias squamipinnis and the ectoparasitic blenny Plagiotremus tapeinosoma. The blenny bites off pieces of skin, mucus and scales from anthias that in return may chase the blenny. Chasing the blenny represents a public good as it makes the parasite change victim species for its next attack. Laboratory experiments using artificial Plexiglas hosts suggest that one reason why individuals contribute to the public good is that the blenny may specialise on non‐punishing ‘free‐riders’. Here, we investigated how far a spatial structure within the shoal and limited space use by the blenny may contribute to punishment being self‐serving. Field observation reveals that anthias indeed live in spatially structured groups and that blennies have preferred areas for attacks. Thus, some anthias individuals are consistently more exposed to blennies than others and hence may gain disproportional benefits from their punishment. In conclusion, spatial structure may contribute to the maintenance of punishment in blenny–anthias interactions even when groups are large.     

Food habits of fishes in the surf zone of a sandy beach at Sanrimatsubara, Fukuoka Prefecture, Japan

To clarify the feeding habits of fishes in surf zones, the gut contents of 19 fish species collected in the surf zone of a sandy beach at Sanrimatsubara, western Japan, were examined. Ontogenetic changes in food preference were recognized in seven species (Mugil cephalus cephalus, Lateolabrax latus, Sillago japonica, Paralichthys olivaceus, Paraplagusia japonica, Takifugu poecilonotus, and Takifugu niphobles). A cluster analysis based on dietary overlaps showed that the surf zone fish assemblage comprised six trophic groups (zooplankton, benthic and epiphytic crustacean, detritus, polychaete, fish, and insect feeders). Of these, the most abundant trophic group was zooplankton feeders, along with benthic and epiphytic crustacean feeders.     

Effects of diet on female fecundity and larval development in the carrion beetle Necrophila japonica

The carrion beetle subfamily Silphinae (Coleoptera: Silphidae) contains dominant macroinvertebrates of soil ecosystems in temperate zones. However, their feeding habits, which determine the role of each species in the ecosystem, have not been sufficiently studied. Moreover, although a diet shift from necrophagy on vertebrate carcasses to predatory feeding on invertebrates is known to occur in this subfamily, the processes and mechanisms of this shift have also been inadequately addressed. We examined female fecundity and larval development on various diets in a Silphinae species, Necrophila (Eusilpha) japonica (Motschulsky). The experimental diets included a meat diet and various invertebrate diets, which reflect the ‘ancestral’ feeding habit in Silphinae, necrophagy, and the ‘derivative’ feeding habit, predatory feeding. Female fecundity was significantly higher on the meat diet (minced beef) than on an insect larvae diet (mealworms and dipteran larvae) but did not significantly differ from that on an earthworm diet. Larval developmental performance was significantly higher on the earthworm diet than on the meat and insect larvae diets. Our results for larval development were consistent with those of previous stable isotope analyses of the same species, in which isotopic values of larval samples agree with those of hypothetical consumers that utilize earthworms. The consistency of results among different methods indicates that N. japonica larvae are most likely earthworm feeders. In contrast, our results for the female fecundity experiment differed from those of previous stable isotope analyses, in which vertebrate carcasses unlikely serve as the staple diet of adults in the field; thus, the feeding habits of N. japonica adults remain unresolved. Our observations that females and larvae performed best on the meat and earthworm diets, respectively, may indicate that, in Silphinae, the diet shift from necrophagy to predatory habits occurs earlier in larvae than in adults.     

Feeding habits and seasonal trophic guilds structuring fish community in the bay mouth region of a tropical estuarine habitat

Understanding trophic relationships of fish in estuarine ecosystem is an important element for sustainable resource management. This study examined the feeding habits of 29 dominant fish species, characterized the trophic guilds, assessed the impact of season and clarified the role of diets in structuring the fish community in the mouth region of Pattani Bay, Thailand. Samples of 5792 fishes collected monthly by gillnets from March 2019 to February 2020 were used for stomach content analyses. It was found that the number of food types and fullness index differed between fish taxa (P < 0.001). Most fishes were specialist feeders feeding on specific food components and were categorized into five trophic guilds: piscivore, shrimp-fish feeder, polychaete feeder, zooplanktivore and planktivore. Six species were piscivorous, considered as apex predators, that fed almost entirely on fishes. High diet overlaps among some species (>0.6) were recorded. Not much variation in seasonal guilds was observed: four guilds in the dry season, three in the moderate rainy season and four in the rainy season. Some species remained in the same guild the whole year round, but some fishes changed seasonally. Two fish communities from different regions of the bay were segregated based on feeding habits. The inner bay community comprised mainly copepod and plankton feeders, but there were more piscivores in the deeper bay mouth area. Results from this study help us to understand the feeding habits and trophic guilds of dominant fish species at the mouth of this tropical estuarine bay.     

The relationship of diet to growth and ammonium excretion in salt marsh fish

Synopsis Detritus is an abundant but poor quality food source for consumers in salt marsh ecosystems. Here we present results of feeding experiments to determine the ability of Fundulus heteroclitus, Cyprinodon variegatus, and Mugil cephalus, three major detritivores in Great Sippewissett Marsh, Massachussetts, to assimilate detritus and use it for growth. C. variegatus, the sheepshead minnow, gained weight on a detrital diet, but F. heteroclitus, the marsh killifish, and juvenile M. cephalus, the striped mullet, lost weight and suffered high mortality on detrital diets. C. variegatus is a herbivore with morphological adaptations for ingesting plant material. F. heteroclitus is a carnivore poorly suited to effectively assimilate detritus from the diet. Although adult M. cephalus are adapted for ingesting and assimilating detrital material, the young may lack these adaptations and thus do not assimilate detritus. C. variegatus excretes ammonium at a lower rate than F. heteroclitus when fed animal food, or when starved for short periods of time. This protein sparing effect could be crucial for survival when detritus is the only food available at certain times of the year. While both C. variegatus and M. cephalus are visitors in the marsh and may leave to exploit food sources elsewhere, F. heteroclitus is a year-round resident and is the most abundant species of fish in the marsh. Yet when high quality food sources become scarce in late summer and fall, detritus, although plentiful, is apparently not a suitable alternative.To whom all correspondences should be addressed.     

Nitrogen,phenolic acids,and other feeding cues for salt marsh detritivores

Summary A number of feeding cues determine the palatability of detritus to detritivorous invertebrates. In salt marsh detritus the feeding cues include flavor provided by phenolics such as ferulic acid, pH, salinity, and nitrogen content. We examined the feeding responses to each cue by using experimental manipulations where we changed concentrations of these chemical cues. The palatability of detritus of the salt marsh grass Spartina alterniflora to the detritivorous snail Melampus bidentatus is reduced by increases in ferulic acid in the detritus. The acidity of the acid is partly responsible for inhibition of feeding, but other flavors of the ferulic acid are the major factor. Changes in salinity makes detritus more or less palatable to different species of detritivores. Available nitrogen confers greater palatability to detritus.In the field the feeding cues are all present simultaneously, and detritivores feed based on a hierarchy of cues. For M. bidentatus the presence of sufficient available nitrogen overwhelms the response to ferulic acid, the second most important cue. Salinity and pH, although used as discriminatory cues by themselves, are located lower in the hierarchy than nitrogen and ferulic acid.As detritus ages in the marsh, nitrogen content changes. Similarly, detritus from eutrophied environments shows enhanced nitrogen content. The differences in palatability of new and old detritus, and from eutrophic and non-eutrophic environments, suggests that detritivores respond not to total nitrogen but rather to the availabel nitrogen fractions. The increase in lignin found in old and eutrophic detritus may increase the binding of available nitrogen, and hence reduce the palatability of detritus.