Food habits, gut morphology and pH, and assimilation efficiency of the zebraperch Hermosilla azurea, an herbivorous kyphosid fish of temperate marine waters

Adult zebraperch, Hermosilla azurea, were found to be functional herbivores in that animal matter constituted <0.01% of the total dry weight of stomach contents of fish collected off Santa Catalina Island in southern California waters. The diet of these fish consisted mainly of red algae (88.2% by dry wt) and also small amounts of brown (7.8%) and green (4.0%) algae. The most important dietary item, the filamentous red algae Polysiphonia spp., was found in >78% of the stomachs and comprised >60% of the contents by dry weight. The digestive tract was long, on average 4.0 times the standard length of the fish, and was composed of the stomach, pyloric caeca, intestine, hindgut chamber with a blind caecum, and rectum. The mean pH of the cardiac stomach was acidic (3.9), whereas that of the intestine was nearly neutral (6.9) and that of the hindgut and blind caecum slightly acidic (6.3 and 6.6, respectively). Algal foods are apparently digested by acid lysis in the stomach and by microbial fermentation in the hindgut. Zebraperch assimilated nutritional constituents from six species of algae with varying degrees of efficiency: carbon (73.7 to 89.7%), nitrogen (72.4 to 84.5%), and protein (71.9 to 94.9%). The fish assimilated these constituents as efficiently or more efficiently from three species of nondietary brown algae as from three species of dietary red and green algae. These results show that zebraperch, like their tropical and subtropical relatives (members of the genus Kyphosus), can digest a wide variety of algae including brown algae containing defensive secondary compounds. Received: 3 November 1997 / Accepted: 19 June 1998     

Gut length and mass in herbivorous and carnivorous prickleback fishes (Teleostei: Stichaeidae): ontogenetic, dietary, and phylogenetic effects

Relative gut length, Zihler’s index, and relative gut mass were measured in four species of prickleback fishes and the effects of ontogeny, diet, and phylogeny on these gut dimensions were determined. Of the four species, Cebidichthys violaceus and Xiphister mucosus shift to herbivory with growth (>45 mm SL), whereas X. atropurpureus and Anoplarchus purpurescens remain carnivores. A. purpurescens belongs to a carnivorous clade, and the three other species belong to an adjacent, herbivorous clade. Gut dimensions were compared in three feeding categories of the four species: (1) small, wild-caught juveniles representing the carnivorous condition before two species shift to herbivory; (2) larger, wild-caught juveniles representing the natural diet condition of the two carnivores and the two species that have shifted to herbivory; and (3) larger, laboratory-raised juveniles produced by feeding a high-protein artificial diet to small juveniles until they have reached the size of the larger, wild-caught juveniles. Comparisons of gut dimensions in categories (1) versus (2) tested for an ontogenetic effect, in (2) versus (3) for a dietary effect, and within each category for a phylogenetic effect. C. violaceus and X. mucosus increased gut dimensions with increase in body size and did not change ontogenetic trajectory in gut dimensions on the high-protein artificial diet, suggesting that they are genetically programmed to develop relatively large guts associated with herbivory. X. atropurpureus increased its gut dimensions with increase in size similar to its sister taxon, X. mucosus, suggesting a phylogenetic influence, but decreased gut dimensions on the high-protein artificial diet, suggesting phenotypic plasticity. Nevertheless, X. atropurpureus displayed a larger gut than A. purpurescens, further evidence that it evolved in an herbivorous clade. A. purpurescens possessed a relatively small gut that was little affected by ontogeny or diet. Ontogeny and phylogeny more than diet appear to influence gut dimensions in the four species, thus favoring genetic adaptation over phenotypic plasticity as the major force acting on digestive system features in the two prickleback clades.     

Assimilation efficiency of a temperate-zone intertidal fish (Cebidichthys violaceus) fed diets of macroalgae

We studied assimilation efficiencies of the temperate-zone intertidal fish Cebidichthys violaceus (Girard, 1854) fed in the laboratory on each of the following species of macroalgae: Spongomorpha coalita (Chlorophyta), Ulva lobata (Chlorophyta), Iridaea flaccida (Rhodophyta) and Porphyra perforata (Rhodophyta). Together, these 4 algae make up over 75% of the natural summer diet of C. violaceus. Assimilation efficiency was calculated by proximate organic analysis of food and feces; the amount of ash in food and feces was used as a standard. Depending on the algal species, the fish assimilated 43 to 81% of the protein, 21 to 44% of the lipid, 45 to 62% of the carbohydrate and 31 to 52% of all three classes of organic material combined. These data are the first results showing that a temperate-zone marine fish can assimilate macroalgal constituents. Protein, carbohydrate and total organic material were absorbed more efficiently from rhodophytes than from chlorophytes. Conversely, lipid was absorbed more efficiently from chlorophytes than from rhodophytes. These results are compared with previous work showing that C. violaceus in nature eats more chlorophytes than rhodophytes, but in laboratory preference tests prefers rhodophytes to chlorophytes.     

Female discharges are more electrifying: spontaneous preference in the electric fish,Eigenmannia (Gymnotiformes,Teleostei)

Summary The tropical South American teleost Eigenmannia lineata showed a spontaneous preference for the female type, compared with the male type, of its sexually dimorphic, weak-electric organ discharge (EOD). Female and male EODs differ in waveform and harmonic content. An isolated fish was simultaneously stimulated with digitally synthesized natural male and female EODs of equal peak-to-peak amplitudes, at ±35 Hz frequency difference centered on its stable resting discharge frequency. The stimulus dipoles were arranged symmetrically to the right and left of the fish's hiding place. All stimulus conditions were permuted at random sequence. Among 11 fish tested, 8 showed a statistically significant preference for one stimulus, the female type, as measured by the amount of time a fish spent close to a stimulus dipole (P<0.05 in each fish, two-tailed). Thus female EODs rather than male EODs were more attractive to adult and juvenile fish of both sexes. It was also concluded that E. lineata is capable of discriminating female from male EODs by a complex sensory capacity requiring neither amplitude nor frequency cues. The EOD waveform changed very little within the ecological range of water conductivities (approximately 10–100 S·cm^-1); the P/N-ratio (a waveform character based on zerocrossing intervals) depended only weakly, but significantly, on conductivity (negative correlation in all four fish). Also, the effect of temperature on EOD waveform was very weak: Q ₁₀-values of the P/N-ratio were below but close to 1 in all fish (27±5°C). Thus, it can be concluded that the EOD waveform is remarkably stable within widely changing conditions-even beyond the variation found in the field-and is therefore potentially useful as a social cue.     

Influence of season,ontogeny and tide on the diet of the temperate marine herbivorous fish Odax pullus (Odacidae)

The diet of the temperate marine herbivorous fish Odax pullus (Pisces: Odacidae) was examined using gut-content analysis followed by principal-component analysis and analysis of variance. Fish were collected near Leigh, on the northeast coast of New Zealand, from February 1983 to September 1984. The data were categorised by size of fish, season, and state of tide. A major ontogenetic shift in diet was observed. Juveniles fed on animal material (crustaceans and gastropods), epiphytic rhodophytes, and some phaeophyte material. Adults fed almost exclusively on two phaeophyte taxa, the laminarian Ecklonia radiata and the fucoid Carpophyllum spp. Dietary selection was evident at the level of particular plant components; the diet of larger fish in spring samples was dominated by fucoid reproductive receptacles. There was no direct evidence of a tidal influence on diet composition of O. pullus, although the relative proportions of E. radiata and Carpophyllum spp. ingested by adult fish appeared to vary with tidal state. The volume of gut contents varied seasonally for all size classes, although the timing of peak annual food intake varied among size classes. Tidal state did not influence the volume of the gut contents. The ontogenetic and seasonal trends evident in the diet of O. pullus appeared to be related to a number of factors including relative gut length, gonad development, ontogenetic changes in feeding anatomy and seasonal changes in algal composition.     

Connectivity between marine reserves and exploited areas in the philopatric reef fish Chrysoblephus laticeps (Teleostei: Sparidae)

‘No-take’ marine protected areas (MPAs) are successful in protecting populations of many exploited fish species, but it is often unclear whether networks of MPAs are adequately spaced to ensure connectivity among reserves, and whether there is spillover into adjacent exploited areas. Such issues are particularly important in species with low dispersal potential, many of which exist as genetically distinct regional stocks. The roman, Chrysoblephus laticeps, is an overexploited, commercially important sparid endemic to South Africa. Post-recruits display resident behavior and occupy small home ranges, making C. laticeps a suitable model species to study genetic structure in marine teleosts with potentially low dispersal ability. We used multilocus data from two types of highly variable genetic markers (mitochondrial DNA control region and seven microsatellite markers) to clarify patterns of genetic connectivity and population structure in C. laticeps using samples from two MPAs and several moderately or severely exploited regions. Despite using analytical tools that are sensitive to detect even subtle genetic structure, we found that this species exists as a single, well-mixed stock throughout its core distribution. The high levels of connectivity identified among sites support the findings of previous studies that have indicated that inshore MPAs are an adequate tool for managing overexploited temperate reef fishes. Even though dispersal of adult C. laticeps out of MPAs is limited, the fact that the large adults in these reserves produce exponentially more offspring than their smaller counterparts in exploited areas makes MPAs a rich source of recruits. We nonetheless caution against concluding that the lack of structure identified in C. laticeps and several other southern African teleosts can be considered to be representative of marine teleosts in this region in general. Many such species are represented in more than one marine biogeographic province and may be comprised of regionally adapted stocks that require individual management.     

Polymorphism in gigantobacterial symbionts in the guts of surgeonfish (Acanthuridae: Teleostei)

The morphology and partial cytology of a group of bacteria-related organisms from the guts of surgeonfish are described. These gigantobacteria symbionts are encountered in the guts of various species of this fish family throughout the Indo-Pacific region and the Red Sea. They have been tentatively grouped and placed in a “family”, Epulopiscidae, according to the first species described, Epulopiscium fishelsoni Montgomery and Pollak, 1988. In addition to a morphological comparison between the various forms from different species of surgeonfish, suggestions are presented with regard to their possible morphological diversification during evolutionary and host-species specificity. Received: 10 June 1998 / Accepted: 29 October 1998     

Seasonality in gonads,fat deposits and condition of tropical surgeonfishes (Teleostei: Acanthuridae)

Lipids are stored in mesenteries surrouding the gut and in paired, retroperitoneal fat bodies dorsal to the anal fin in at least 24 surgeonfishes representing 5 genera (Acanthurus, Ctenochaetus, Naso, Prionurus, Zebrasoma) from Western Mexico, Hawaii, French Polynesia, Great Barrier Reef and the Red Sea. In A. nigrofuscus from the Red Sea and C. striatus from French Polynesia, fat stores and condition index (K=100 W/L ³) increased during cool seasons, several months before gonads developed, and declined with the onset of gonadal development. Fat bodies and mesenteric fat deposits developed in all sizes of A. nigrofuscus and C. striatus, and condition paralleled seasonal changes in fat deposits. Gonads of A. nigrofuscus developed in specimens of 76 to 159 mm standard length (SL), while they developed in C. striatus only at lengths>170 mm total length (TL) for females and 195 mm TL for males. Mesenteric fats and fat bodies exhibited similar changes in size, and did not differ in occurrence of major groups of lipids or their relative prevalence. The relative proportions of the major lipid classes differed, however, between these storage sites and other tissues. Muscle, liver and gonads contained less triacylglycerol relative to cholesterol and polar lipid than did mesenteric or fat body lipids. Energy and nutrients acquired during cool saasons with short feeding days appear to support reproductive events during warm seasons, when feeding during longer days fails to meet demands. Fat stores and condition continue to decline beyond the reproductive season until winter.     

Range extensions of teredinids (shipworms) and polychaetes in the vicinity of a temperate-zone nuclear generating station

A study of marine boring and fouling organisms in the vicinity of the thermal effluent from the Oyster Creek Nuclear Generating Station, Barnegat Bay, New Jersey, USA has been conducted since 1971. Two subtropical species of shipworms have been found in significant numbers in the effluent. These are Teredo bartschi Clapp and Teredo furcifera von Martens. The shipworm Lyrodus sp. and the flatworm Taenioplana teredini Hyman, which is predatory on teredinids, were also found, but on only one occasion. The polychaete Ficopomatus enigmaticus (Fauvel) has been found for the first time on the Atlantic Coast of North America. Three other polychaetes, Potamilla (Potamethus) spathiferus (Ehlers), Loimia medusa (Savigny), and Sphaerosyllis sp., have been newly identified from Barnegat Bay, but probably are not recent range extensions. Only T. bartschi, T. furcifera, and F. enigmaticus have established breeding populations. Their distributions and abundances are probably related to the thermal effluent; this is most clear in the case of T. bartschi and least clear in the case of F. enigmaticus. T. bartschi was confined at first to the heated effluent in Oyster Creek, but has spread to Forked River. It has become a major faunal element in Oyster Creek. The impact of the other exotic species has been negligible. The primary mechanism of the range extensions is by boat transport.     

Lipid storage and composition in tropical surgeonfishes (Teleostei: Acanthuridae)

The lipid composition of tropical marine reef fishes is poorly known, despite their use as food by local human populations and recent interest in health-related benefits of fish lipids. We examined the composition of lipids from epaxial muscle, liver, and two storage sites [mesenteries surrounding the gut (intraperitoneal fat, IPFs) and retroperitoneal fat bodies (FBs) posterior to the peritoneal cavity] in three species of surgeonfishes from Ishigaki Island, Japan: Naso lituratus (Bloch and Schneider, 1801), Acanthurus lineatus (Linnaeus, 1758), and A. bariene (Lesson, 1830). Triacylglycerols dominated all samples of neutral lipid and constituted ≥ 99% of FBs and IPFs. Polar lipids generally contained large fractions of phosphatidylethanolamine and phosphatidylcholine. Quantified fatty acids ranged in length from C₁₄ to C₂₄. C₁₆ fatty acids prevailed (>35% of neutral fatty acids, >23% of polar fatty acids), although C₁₈ (>16 and >14%, respectively) and C₂₀ acids (>8 and >19%, respectively) were also common. Saturated fatty acids, dominated by palmitic acid (16:0), comprised 38.7 to 50.7% of acids from neutral lipids and 30.8 to 41.1% from polar lipids. The most common monounsaturated acids were 18:1n9 and 20:1n9. Polyunsaturated acids were prevalent in polar lipids (especially 20:4n6, 20:5n3, 22:2n3, 22:5n3, 22:5n6 and 22:6n3). Common polyunsaturated acids of neutral lipids were 18:2n6, 18:4n3, several n-3 and n-6 C₂₀ acids, 22:2n3 and 22:5n3. IPF and FB were almost identical across species, and lipids of fat bodies (IPFs, FBs) were more similar to those of muscle than those of liver for all three species. The FBs appear to constitute an accessory storage site, which overcomes constraints on lipid storage imposed by a small, inflexible abdominal cavity that contains both viscera and consistently voluminous gut contents. Fatty acid signatures indicate that largely overlooked epiphytic or epilithic diatoms contribute significantly to lipid acquisition. The combination of large quantities of both saturated and n-3 and other polyunsaturated fatty acids in surgeonfishes, in contrast to low saturates and high polyunsaturated acids in lipids of commercially important cool-water fishes, suggests that a study of dietary effects of fish lipids on human inhabitants of the tropics may be instructive insofar as human health and nutrition are concerned. Received: 16 March 1998 / Accepted: 6 August 1998     

Histology and ultrastructure of the skin of Lepadichthys lineatus (Gobiesocidae: Teleostei)

The skin of the Red Sea clingfish Lepadichthys lineatus Briggs, 1966, which lives on shallow-water crinoids, consists of a continuous layer of giant mucus-producing cells. Upon stimulation, these cells are able, in seconds, to envelope the fish entirely with a thick layer of their secretion. The cells are 75 to 290 high, and are enveloped by septa of compressed epithelium. They rest on a strong collagenous layer, and extend to the skin surface, where they open. The nuclei of the giant cells are large, and irregularly shaped; their cytoplasm has a very extensive endoplasmic reticulum and undergoes deep structural changes during maturation. Their ability to produce large amounts of mucus seems to be an adaptive device to protect the fish from contact with the rough surface of the crinoid host.     

Quantitative distribution of herbivorous reef fishes in the Gulf of Aqaba (Red Sea)

The distribution of the main herbivorous fishes (Acanthuridae, Scaridae, Siganidae) was studied across a coral reef of the Jordanian coast in the Gulf of Aqaba (Red Sea). Visual counts were realized by diving along transects (200 m long and 5 m wide), parallel to the shore, at 10 stations located from the lagoon to 40 m deep on the outer reef slope. Herbivorous reef fishes reach their highest abundance on the reef front, where 234 fishes were counted per 1,000 m². Their density decreases on the reef flat, with an average of 150 fish 1,000 m^-2, and is lowest on the outer reef slope (69 fish 1,000 m^-2). Surgeonfishes form 63% of the herbivorous ichthyofauna, parrotfishes 35%, and rabbitfishes 2%. Families and species display different distributions according to biota. The Acanthuridae dominate on the reef flat, whereas the Scaridae are more numerous on the outer reef slope. The evolution of the social structure of the main species was observed: the adults generally school in the lagoon and on the reef flat, but are mainly solitary on the reef slope. The distribution of juvenile individuals is more restricted: they are concentrated on the reef front and on the upper part of the reef slope.This study is part of a cooperation programme between the University of Nice (France) and the University of Jordan, to study the ecology of the coral reefs and the surrounding waters of the Jordanian coast (Gulf of Aqaba, Red Sea)     

Identification of tenuis of four French Polynesian Carapini (Carapidae: Teleostei)

Four species of adult Carapini (Carapidae) occur on Polynesian coral reefs: Encheliophis gracilis, Carapus boraborensis, C. homei and C. mourlani. Samples collected in Rangiroa and Moorea allowed us to obtain different tenuis (larvae) during their settlement phases or directly inside their hosts. These were separated into four lots on the basis of a combination of pigmentation, meristic, morphological, dental and otolith (sagittae) features. Comparison of these characters with those of the adults allows, for the first time, taxonomic identification of these tenuis-stage larvae.     

Sexual maturity and hermaphroditism of the red porgy Pagrus pagrus (Teleostei: Sparidae)

Gonadal changes related to sexual maturity were examined by histology in reared populations of the red porgy Pagrus pagrus L., aged 0.5 to 6.5 years. More than 50% of red porgies were mature at the age of 4 years although some individuals were already mature at the age of 3, as indicated by changes in the gonadosomatic index, plasma levels of vitellogenin, 17β-estradiol and testosterone in females and males. Sex frequency distribution analysis in relation to the age and histological analysis of the gonads indicated the presence of males originated from the sex change of adult functional females. P. pagrus is a protogynous species. Some females had not changed sex up to the age of 6.5 years. Received: 16 July 1998 / Accepted: 29 April 1999     

A new fat body associated with the gonad of surgeonfishes (Acanthuridae: Teleostei)

Adults of nine species of surgeonfishes (Acanthuridae) from the Red Sea, Israel and the Eastern Tropical Pacific off Mexico possess previously undescribed fat bodies associated with both male and female gonads. These structures are unique in that they are external to the abdominal cavity, are composed almost entirely of highly-saturated triacylglycerols, and appear to have close circulatory connection with the gonads. In all species except one, the fat bodies rest in triangular pockets between the trunk musculature and skin, which extend posteriorly just above the anal fin base. The composition, arrangement and dynamics of the fat bodies suggest that they support the active gonad during a spawning season that lasts, in one species, from May to September.     

Social organization and sexual pattern in the Mediterranean parrotfish Sparisoma cretense (Teleostei: Scaridae)

We examined the social organization and reproductive pattern of a population of Sparisoma cretense L. at Lampedusa Island (Italy). During the breeding season (July to September) individuals occur either in territorial or in non-territorial groups, which quantitatively differ with respect to use of space, sex-ratio and sexual activity. Territorial groups consist of one male plus one to three females, whereas in non-territorial groups as many as 54 fish share a common area. Among non-territorial groups, variations, in both sex-ratio and behavior, suggest they could represent either feeding aggregations or reproductive aggregations. Spawning, always in pairs, occurs daily in a short period of time before dusk and has been observed only in territorial groups. Group spawning has never been observed, but another alternative mating tactic, streaking on pair spawning, was recorded. In the non-breeding season fish do not aggregate in organized social units. Histological examination of gonads showed that adults are larger than 12 cm total length. Females have an asynchronous ovary, typical of species spawning several times during the breeding season. Males show secondary testes, indicating that their gonads develop ovaries as juveniles, which are later redifferentiated into functional testes. Histological and demographic data seem to indicate that, as in other species of this genus, prematurational sex-change occurs. The sexual pattern appears to be essentially gonochoristic, but the potential for sex-change is not excluded. Received: 5 January 1999 / Accepted: 22 June 1999     

Biology of Serranicotyle labracis,ectoparasite of Dicentrarchus labrax (Teleostei): contribution to the study of its populations

Serranicotyle labracis (Van Beneden and Hesse, 1863) Maillard, Euzet and Silan, 1988 (synonym: Microcotyle labracis Van Beneden and Hesse, 1863) is a monogenean Polyopisthocotylea and a gill parasite of the bass Dicentrarchus labrax (Linné, 1758). Analysis of a western Mediterranean population of this parasite (Gulf of Lions, Languedoc, France) clarified several aspects of its biology. Its location in the gill biotope (Silan et al. 1987) was not fundamentally different from that found by Oliver (1977) and Winch (1983), but its distribution in host populations differed from that reported in the earlier studies; it displayed a negative binomial distribution in the study zone. At a finer level of observation, the behaviour of the hosts, as a function of degree of maturity and sex, appeared to determine more than any other factor the extent of infestation by the parasite. Previous suggestions in the literature to account for these differences in population dynamics are discussed.