Xiphister Mucosus Research Articles

How They Eat: An Examination of Digestive Physiology of Sympatric Prickleback Fishes

How fishes thrive on different diets is not understood as well as for terrestrial animals. Thus, examining the flexibility of gut function in fishes with different diets provides insight into dietary specialization in the largest vertebrate class. In this study, we measured the amylolytic activities in the guts of closely related, sympatric species: Xiphister mucosus (herbivore), X. atropurpureus (omnivore), Phytichthys chirus (omnivore), and Anoplacharus purpurescens (carnivore). For all species, the amylase activity decreased moving distally along the intestine, but less so for the carnivorous A. purpurescens. Overall, our results—more elevated amylase activity in those fish eating more algae—agree with the Adaptive Modulation Hypothesis (AMH), which suggests that there should be a match between enzymatic activity and dietary substrate intake. Measurements of proteases and lipases are in progress, but we anticipate some support for AMH in other enzyme classes. Moreover, we are also measuring these same enzyme activities in these same species fed omnivorous or carnivorous diets in the laboratory. The flexibility of the intestines of the fish species to accommodate different food types provides insight into their roles in the ecosystems and how fixed the vertebrate digestive system can be. NSF IOS 1355224 Undergraduate Research Opportunity Program at University of California, Irvine. This is the full abstract presented at the American Physiology Summit 2024 meeting and is only available in HTML format. There are no additional versions or additional content available for this abstract. Physiology was not involved in the peer review process.

Can you stomach it? Comparative transcriptomics and biochemistry of the stomachs of prickleback fishes (Stichaeidae) consuming different diets

Despite obvious whole animal, tissue level, and biochemical differences among species with different diets, dietary specialization has molecular underpinnings that are not well understood. In this project, we used comparative transcriptomics of stomach tissues to observe how prickleback fishes (Stichaeidae) achieve dietary-driven differences in stomach function. Xiphister mucosus (herbivore), X. atropurpureus (omnivore), Phytichthys chirus (omnivore) and the carnivorous Anoplarchus purpurescens were harvested from the wild, and fed omnivore and/or carnivore diets in the laboratory. Transcriptomes were sequenced for stomach tissues from two individuals per species and diet treatment. The number of shared orthologs varied with phylogenetic relatedness of the species (i.e., sister-taxa X. mucosus and X. atropurpureus shared the most expressed genes among the species), and differential expression and selection analyses are underway, although hydrogen-potassium ATPase appears to be under positive selection in some species. All four fish species strongly expressed pepsinogen and chitinase genes in their stomachs, yet biochemical activity levels (N ≥ 6 per species and diet treatment) of these two digestive enzymes matched natural diet, with the species consuming the most protein ( A. purpurescens) and chitin ( P. chirus) having the highest pepsin and chitinase activities, respectively. These activities were relatively unaltered by the laboratory diets, suggesting that activity level cannot be changed significantly through a dietary shift. Although intestinal carbohydrases typically show support for the Adaptive Modulation Hypothesis—a match among digestive enzyme activities with concentrations of ingested substrate—this may be one of the first investigations to observe such support for gastric enzymes. Thus, beyond being an important vertebrate innovation, the stomach may also provide digestive benefit depending on natural diet. NSF Grant IOS-1355224 and NSF DBI-1852096 This is the full abstract presented at the American Physiology Summit 2023 meeting and is only available in HTML format. There are no additional versions or additional content available for this abstract. Physiology was not involved in the peer review process.

Comparative transcriptomics reveal tissue level specialization towards diet in prickleback fishes

Beyond a few obvious examples (e.g., gut length, amylase activity), digestive and metabolic specializations towards diet remain elusive in fishes. Thus, we compared gut length, δ13C and δ15N signatures of the liver, and expressed genes in the intestine and liver of wild-caught individuals of four closely-related, sympatric prickleback species (family Stichaeidae) with different diets: Xiphister mucosus (herbivore), its sister taxon X. atropurpureus (omnivore), Phytichthys chirus (omnivore) and the carnivorous Anoplarchus purpurescens. We also measured the same parameters after feeding them carnivore or omnivore diets in the laboratory for 4 weeks. Growth and isotopic signatures showed assimilation of the laboratory diets, and gut length was significantly longer in X. mucosus in comparison to the other fishes, whether in the wild, or in the lab consuming the different diets. Dozens of genes relating to digestion and metabolism were observed to be under selection in the various species, but P. chirus stood out with some genes in the liver showing strong positive selection, and these genes correlating with differing isotopic incorporation of the laboratory carnivore diet in this species. Although the intestine showed variation in the expression of hundreds of genes in response to the laboratory diets, the liver exhibited species-specific gene expression patterns that changed very little (generally <40 genes changing expression, with P. chirus providing an exception). Overall, our results suggest that the intestine is plastic in function, but the liver may be where specialization manifests since this tissue shows species-specific gene expression patterns that match with natural diet.

A Tail of Four Fishes: An Analysis of Kinematics and Material Properties of Elongate Fishes.

The elongate body plan is present in many groups of fishes, and this morphology dictates functional consequences seen in swimming behavior. Previous work has shown that increasing the number of vertebrae, or decreasing the intervertebral joint length, in a fixed length artificial system increases stiffness. Tails with increased stiffness can generate more power from tail beats, resulting in an increased mean swimming speed. This demonstrates the impacts of morphology on both material properties and kinematics, establishing mechanisms for form contributing to function. Here, we wanted to investigate relationships between form and ecological function, such as differences in dietary strategies and habitat preferences among fish species. This study aims to characterize and compare the kinematics, material properties, and vertebral morphology of four species of elongate fishes: Anoplarchus insignis, Anoplarchus purpurescens, Xiphister atropurpureus, and Xiphister mucosus. We hypothesized that these properties would differ among the four species due to their differential ecological niches. To calculate kinematic variables, we filmed these fishes swimming volitionally. We also measured body stiffness by bending the abdominal and tail regions of sacrificed individuals in different stages of dissection (whole body, removed skin, and removed muscle). Finally, we counted the number of vertebrae from CT scans of each species to quantify vertebral morphology. Principal component and linear discriminant analyses suggested that the elongate fish species can be distinguished from one another by their material properties, morphology, and swimming kinematics. With this information combined, we can draw connections between the physical properties of the fishes and their ecological niches.

Long-axis twisting during locomotion of elongate fishes.

Fish live in a complex world and must actively adapt their swimming behavior to a range of environments. Most studies of swimming kinematics focus on two-dimensional properties related to the bending wave that passes from head to tail. However, fish also twist their bodies three dimensionally around their longitudinal axis as the bending wave passes down the body. We measured and characterized this movement, which we call 'wobble', in six species of elongate fishes (Anoplarchus insignis, Xiphister mucosus, Lumpenus sagitta, Pholis laeta, Apodichthys flavidus and Ronquilus jordani) from three different habitats (intertidal, nearshore and subtidal) using custom video analysis software. Wobble and bending are synchronized, with a phase shift between the wobble wave and bending wave. We found that species from the same habitats swim in similar ways, even if they are more closely related to species from different habitats. In nearshore species, the tail wobbles the most but, in subtidal and intertidal species, the head wobbles more than or the same as the tail. We also wanted to understand the relationship between wobble and the passive mechanics of the fish bodies. Therefore, we measured torsional stiffness and modulus along the body and found that modulus increases from head to tail in all six species. As wobble does not correlate with the passive properties of the body, it may play a different role in swimming behavior of fishes from different habitats.

Morphology and ontogeny of multiple lateral-line canals in the rock prickleback, Xiphister mucosus (Cottiformes: Zoarcoidei: Stichaeidae).

The structure and ontogeny of lateral-line canals in the Rock Prickleback, Xiphister mucosus, were studied using cleared-and-stained specimens, and the distribution and morphology of neuromasts within lateral-line canals were examined using histology. X. mucosus has seven cephalic canals in a pattern that, aside from four branches of the infraorbital canals, is similar to that of most teleostean fishes. Unlike most other teleosts, however, X. mucosus features multiple trunk lateral-line canals. These include a short median posterior extension of the supratemporal canal and three paired, branching canals located on the dorsolateral, mediolateral, and ventrolateral surfaces. The ventrolateral canal (VLC) includes a loop across the ventral surface of the abdomen. All trunk canals, as well as the branches of the infraorbitals, are supported by small, dermal, ring-like ossifications that develop independently from scales. Trunk canals develop asynchronously with the mediodorsal and dorsolateral canals (DLC) developing earliest, followed by the VLC, and, finally, by the mediolateral canal (MLC). Only the mediodorsal and DLC connect to the cephalic sensory canals. Fractal analysis shows that the complexity of the trunk lateral-line canals stabilizes when all trunk canals develop and begin to branch. Histological sections show that neuromasts are present in all cephalic canals and in the DLC and MLC of the trunk. However, no neuromasts were identified in the VLC or its abdominal loop. The VLC cannot, therefore, directly function as a part of the mechanosensory system in X. mucosus. The evolution and functional role of multiple lateral-line canals are discussed.

More than one way to be an herbivore: convergent evolution of herbivory using different digestive strategies in prickleback fishes (Stichaeidae)

In fishes, the evolution of herbivory has occured within a spectrum of digestive strategies, with two extremes on opposite ends: (i) a rate-maximization strategy characterized by high intake, rapid throughput of food through the gut, and little reliance on microbial digestion or (ii) a yield-maximization strategy characterized by measured intake, slower transit of food through the gut, and more of a reliance on microbial digestion in the hindgut. One of these strategies tends to be favored within a given clade of fishes. Here, we tested the hypothesis that rate or yield digestive strategies can arise in convergently evolved herbivores within a given lineage. In the family Stichaeidae, convergent evolution of herbivory occured in Cebidichthys violaceus and Xiphister mucosus, and despite nearly identical diets, these two species have different digestive physiologies. We found that C. violaceus has more digesta in its distal intestine than other gut regions, has comparatively high concentrations (>11mM) of short-chain fatty acids (SCFA, the endpoints of microbial fermentation) in its distal intestine, and a spike in β-glucosidase activity in this gut region, findings that, when coupled to long retention times (>20h) of food in the guts of C. violaceus, suggest a yield-maximizing strategy in this species. X. mucosus showed none of these features and was more similar to its sister taxon, the omnivorous Xiphister atropurpureus, in terms of digestive enzyme activities, gut content partitioning, and concentrations of SCFA in their distal intestines. We also contrasted these herbivores and omnivores with other sympatric stichaeid fishes, Phytichthys chirus (omnivore) and Anoplarchus purpurescens (carnivore), each of which had digestive physiologies consistent with the consumption of animal material. This study shows that rate- and yield-maximizing strategies can evolve in closely related fishes and suggests that resource partitioning can play out on the level of digestive physiology in sympatric, closely related herbivores.

Osteology of the prickleback genus Xiphister (Cottiformes: Zoarcoidei: Stichaeidae)

AbstractThe two members of the genus Xiphister, Xiphister atropurpureus and Xiphister mucosus, are inter‐ and subtidal marine fishes from the west coast of North America, ranging from southern Alaska to southern California. Although both species were described over 150 years ago and portions of their anatomy have been described in several studies, a detailed description of their anatomy and morphology is lacking. In this article, we describe and illustrate the skeletal anatomy of the genus, clarifying aspects of the skull roof and neurocranium, jaws, suspensorium, opercular series, vertebral column, dorsal and anal fins, trunk lateral‐line canals and pectoral girdle. We describe for the first time the elements of the hyoid and gill arches and scales. The genus is one of several within Stichaeidae with multiple, complexly branching trunk lateral lines, a feature that is found in only fifteen families of teleostean fishes. The development of the lateral‐line canals and scales in Xiphister is discussed. Data reported herein will establish a foundation for the further anatomical and systematic studies of Stichaeidae and Zoarcoidei generally.

Evolution of ontogenetic dietary shifts and associated gut features in prickleback fishes (Teleostei: Stichaeidae)

We tested the hypothesis that an ontogenetic dietary shift from carnivory to herbivory or omnivory, and concomitant changes in the gut facilitating digestion of algae, are synapomorphies of the tribes Xiphisterini and Esselenichthyini in the family Stichaeidae (pricklebacks). Previous investigations have revealed that two xiphisterine pricklebacks-Xiphister mucosus and Xiphister atropurpureus-become herbivorous or omnivorous, respectively, as their bodies grow larger, and that their guts show related changes in length and function. In this study we found that, with increase in size, the basal member of the Xiphisterini, Phytichthys chirus, showed an increased proportion of algae in its diet, increased activity of α-amylase and decreased activity of aminopeptidase, all of which support the synapomorphy hypothesis. Cebidichthys violaceus, a herbivore in the Esselenichthyini, shows similar ontogenetic changes in diet and digestive tract length and physiology, but these features were not observed in two derived carnivores, Dictyosoma burgeri and Dictyosoma rubrimaculatum, within the clade. These results suggest that herbivory is isolated to C. violaceus within the Esselenichthyini. Allometric relationships of gut length as a function of body size generally follow diet within the Xiphisterini and Esselenichthyini, with herbivores having the longest guts, which become disproportionately longer than body size as the fishes grow, omnivores intermediate gut lengths, and carnivores the shortest. A carnivore from an adjacent clade, Anoplarchus purpurescens, had the shortest gut, which did not change in length relative to body length as the fish grew. Overall, our results clarify the patterns of dietary evolution within the Stichaeidae and lay the foundation for more detailed studies of dietary and digestive specialization in fishes in the family.

Sequence and expression of an α-amylase gene in four related species of prickleback fishes (Teleostei: Stichaeidae): ontogenetic, dietary, and species-level effects

Partial α-amylase gene sequences were determined and α-amylase gene expression was quantified in four species of carnivorous, omnivorous, and herbivorous prickleback fishes (family Stichaeidae) to assess the effects of ontogeny, diet, and species on expression of this gene. Pairwise comparison of α-amylase nucleotide sequences revealed 96-98 % identity, and comparison of amino acid portions revealed 93-95 % similarity among the four prickleback species. Expression was determined using in situ hybridization and intensity of expression quantified using image analysis. Alpha-amylase expression level was compared in three feeding categories of the four species: (1) small, wild-caught carnivorous juveniles; (2) larger, wild-caught juveniles of the carnivorous species and the three that had shifted to herbivory or omnivory; and (3) larger, juveniles produced by feeding a low-starch artificial diet to small juveniles until they reached the size of the larger wild-caught juveniles. The results showed no dietary effect in any species but significant ontogenetic and species-level effects in Cebidichthys violaceus, as well as in the sister species Xiphister mucosus and X. atropurpureus. Based on a phylogeny for the Stichaeidae produced for this study using two mtDNA genes and one nuclear gene, the ontogenetic dietary shifts to herbivory/omnivory evolved independently in C. violaceus and in the clade containing the two species of Xiphister. All three of these species increased α-amylase gene expression with increase in size and had higher expression than Anoplarchus purpurescens, which is a member of a third, stichaeid clade comprising carnivores. These results show the importance of α-amylase in the herbivores and omnivores.

Aquatic and Terrestrial Locomotion of the Rock Prickleback, Xiphister mucosus (Cottiformes: Zoarcoidei: Stichaeidae)

Several taxa of marine fishes are capable of both aquatic and terrestrial locomotion, yet the terrestrial locomotion mechanics of many of these taxa have not been studied. Many species of the prickleback family Stichaeidae, including the Rock Prickleback, Xiphister mucosus, are capable of both aquatic and terrestrial locomotion. In this study, high-speed video was used to compare the mechanics of aquatic and terrestrial locomotion of X. mucosus. While swimming, tail-beats occur faster, and the velocities of the head and tail are greater than on land. On land, tail-beats are slower and cover a longer distance, yet the distance traveled by the head is similar to the distance traveled during aquatic locomotion. Froude propulsion efficiencies during swimming average 0.75 (s = 0.05) indicating efficient swimming mechanics in X. mucosus. These efficiency values are typical for other anguilliform swimmers that make terrestrial excursions. Overall, X. mucosus makes slight modifications to its efficient swimming locomotion mechanics, principally by increasing movement of the tail, to move effectively over the terrestrial environment.

The ‘Expansion-Contraction’ model of Pleistocene biogeography: rocky shores suffer a sea change?

Approximately 20,000 years ago the last glacial maximum (LGM) radically altered the distributions of many Northern Hemisphere terrestrial organisms. Fewer studies describing the biogeographic responses of marine species to the LGM have been conducted, but existing genetic data from coastal marine species indicate that fewer taxa show clear signatures of post-LGM recolonization. We have assembled a mitochondrial DNA (mtDNA) data set for 14 co-distributed northeastern Pacific rocky-shore species from four phyla by combining new sequences from ten species with previously published sequences from eight species. Nuclear sequences from four species were retrieved from GenBank, plus we gathered new elongation factor 1-alpha sequences from the barnacle Balanus glandula. Results from demographic analyses of mtDNA for five (36%) species (Evasterias troschelii, Pisaster ochraceus, Littorina sitkana, L. scutulata, Xiphister mucosus) were consistent with large population expansions occurring near the LGM, a pattern expected if these species recently recolonized the region. However, seven (50%) species (Mytilus trossulus, M. californianus, B. glandula, S. cariosus, Patiria miniata, Katharina tunicata, X. atropurpureus) exhibited histories consistent with long-term stability in effective population size, a pattern indicative of regional persistence during the LGM. Two species of Nucella with significant mtDNA genetic structure showed spatially variable demographic histories. Multilocus analyses for five species were largely consistent with mtDNA: the majority of multilocus interpopulation divergence times significantly exceeded the LGM. Our results indicate that the LGM did not extirpate the majority of species in the northeastern Pacific; instead, regional persistence during the LGM appears a common biogeographic history for rocky-shore organisms in this region.

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.

Trypsin Gene Expression by Quantitative In Situ Hybridization in Carnivorous and Herbivorous Prickleback Fishes (Teleostei: Stichaeidae): Ontogenetic, Dietary, and Phylogenetic Effects

We determined trypsin gene sequences and compared relative levels of trypsin gene expression as influenced by ontogeny, diet, and phylogeny in four related prickleback fish species. Of these species, Cebidichthys violaceus and Xiphister mucosus shift from carnivory to herbivory at approximately 45 mm standard length [SL], whereas Xiphister atropurpureus and Anoplarchus purpurescens remain carnivores. Pairwise sequence similarities among the four species were 77%-95% for the trypsin nucleotides and 69%-94% for the amino acids. Trypsin gene expression levels in small (30-40 mm SL) and larger (60-75 mm) wild-caught juveniles and larger (60-75 mm) juveniles raised on a high-protein artificial diet increased with ontogeny in all four species but in response to the diet only in the two carnivores. The indistinguishable expression levels in the sister taxa, X. mucosus and X. atropurpureus, represented the only apparent phylogenetic effect. Xiphister atropurpureus, however, increased both trypsin gene expression and enzymatic activity (the latter from a previous study) on the high-protein artificial diet, indicating transcriptional rather than posttranscriptional (shown in X. mucosus) regulation of the activity. This study provides evidence for genetically programmed upregulation of trypsin gene expression with ontogeny in both the carnivorous and herbivorous species but in response to the high-protein artificial diet only in the carnivores.

Storage and Absorption in the Digestive System of Carnivorous and Herbivorous Prickleback Fishes (Teleostei: Stichaeidae): Ontogenetic, Dietary, and Phylogenetic Effects

The effects of ontogeny, diet, and phylogeny on glycogen storage levels and esterase and alkaline phosphatase activities in four related prickleback fishes were determined in situ using quantitative histochemistry. Of these species, Cebidichthys violaceus and Xiphister mucosus shift from carnivory to herbivory at approximately 45 mm standard length (SL), whereas Xiphister atropurpureus and Anoplarchus purpurescens remain carnivores. Comparisons between small (30-40 mm SL) and larger (60-75 mm SL) wild-caught juveniles showed that glycogen storage levels and alkaline phosphatase activity were unchanged with ontogeny. Comparisons between the larger wild-caught juveniles and juveniles of the same size that had been raised on a high-protein animal diet revealed that glycogen storage level and alkaline phosphatase activity increased in all species in response to this diet. Esterase activity also increased in response to the high-protein animal diet in all four species but increased with ontogeny only in C. violaceus, X. mucosus, and X. atropurpureus, in the xiphisterine clade, and not in A. purpurescens, in the adjacent alectriine clade. Xiphister mucosus and X. atropurpureus showed indistinguishable responses in esterase activity to ontogeny and diet despite their divergent natural diets. Overall, glycogen storage level and alkaline phosphatase activity responded primarily to diet, whereas esterase activity was also influenced by ontogeny and phylogeny and differed between intestinal regions among the species.

Digestive Enzyme Activities in Herbivorous and Carnivorous Prickleback Fishes (Teleostei: Stichaeidae): Ontogenetic, Dietary, and Phylogenetic Effects

We measured the activities of eight digestive enzymes in four species of herbivorous and carnivorous prickleback fishes and determined the effects of ontogeny, diet, and phylogeny on these enzyme activities. Of the four species, Cebidichthys violaceus and Xiphister mucosus shift to a more herbivorous diet as they grow (> or =45 mm SL [standard length]), whereas Xiphister atropurpureus and Anoplarchus purpurescens remain carnivores throughout life. Digestive enzyme activities of small (30-40 mm SL) carnivorous juveniles were compared with those of larger (60-75 mm SL) wild-caught juveniles that had consumed a natural diet and larger (60-75 mm SL) juveniles raised on a high-protein animal diet. Cebidichthys violaceus and both species of Xiphister showed ontogenetic changes in digestive enzyme activities, whereas A. purpurescens did not. Despite dietary differences between X. atropurpureus and X. mucosus, these sister taxa displayed the most similar digestive enzyme activities from ontogenetic and dietary perspectives (high alpha-amylase and lipase and low trypsin and aminopeptidase activities), and both were more similar to C. violaceus, a member of the same largely herbivorous clade, than either was to A. purpurescens, a member of an adjacent, carnivorous clade. The results support the hypothesis that phylogeny influences digestive enzyme activities in these fishes. Anoplarchus purpurescens, a carnivore with a diverse diet, showed great plasticity in enzyme activity, especially trypsin and aminopeptidase, which were elevated in this species to the highest level among the four species after consuming the high-protein diet. These results support the hypothesis that fishes with relatively broad diets can modulate digestive enzyme activities in response to changes in dietary composition.

Digestive enzyme activities in carnivores and herbivores: comparisons among four closely related prickleback fishes (Teleostei: Stichaeidae) from a California rocky intertidal habitat

Three digestive enzymes in four species of closely related prickleback fishes (family Stichaeidae: Cebidichthys violaceus, Xiphister mucosus, Xiphister atropurpureus and Anoplarchus purpurescens) were analysed to assess whether diet or phylogeny played a larger role in influencing digestive enzyme activity. Cebidichthys violaceus and X. mucosus are primarily herbivorous, whereas X. atropurpureus and A. purpurescens are mainly carnivorous. The two Xiphister species are sister taxa, and A. purpurescens is in a clade adjacent to that of the three other species. Pepsin and trypsin specific activities did not differ significantly among the four species, but α‐amylase activity was significantly higher in the two Xiphister species, followed by C. violaceus, and then A. purpurescens. The wide disparity between the two carnivores, the striking similarity between the two sister taxa, and the significant difference between the two herbivores indicate that activity of α‐amylase follows a pattern influenced more by phylogeny than by diet in these fishes.

Role of Polyphenolic Molecular Size in Reduction of Assimilation Efficiency in Xiphister Mucosus

Feeding deterrent effects of polyphenolic compounds from both terrestrial and marine environments have been well studied. However, the effects of this compound class on herbivore digestion and assimilation are less well known, particularly with respect to marine herbivores. This study found that the reduction of assimilation efficiency in Xiphister mucosus, a marine herbivorous fish, fed algae treated with marine polyphenolics (phlorotannins) was affected by phlorotannin molecular size (mass). In general, phlorotannins &gt; 16.60 x 10—21 g(= &gt; 10 kDa) significantly decreased the assimilation of X. mucosus; those &lt;8.30 x 10—21 g (= &lt;5 kDa) rarely, if ever, had an effect. To determine the ecological relevance of these findings, phlorotannin molecular size profiles were completed for several temperate and tropical brown algal species. In all but one of the algal species examined, the phlorotannins were concentrated in the &gt; 16.60 x 10—21 size range. Thus, the size range of phlorotannins that significantly reduced assimilation was also the size range in which they were most concentrated. This work supports the idea that polyphenolics as a chemical class do not all have the same bioactivity, but differ in their activity in a size—dependent manner. It also addresses some of the discrepancies seen in feeding—preference studies, providing an alternative explanation for results.

Herbivore responses to a seasonally fluctuating food supply: Growth potential of two temperate intertidal fishes based on the protein and energy assimilated from their macroalgal diets

The protein and energy assimilated from eight main dietary algae by the stichaeid fishes Cebidichthys violaceus (Girard) and Xiphister mucosus (Girard) were calculated using bimonthly data on the chemical compositions (carbohydrate, protein, lipid) of these algae and the stomach contents and assimilation efficiencies of the two fish species. The effects of food choice on diet quality were estimated by comparing the amounts of energy and protein determined to be assimilated from the actual (selective) diets with those that would be assimilated from hypothetical (nonselective) diets based on algal availability. Both fishes consumed carbohydrate-rich, protein-poor algae in the winter and protein-rich, carbohydrate-poor algae during the rest of the year. Thus, contrary to an expectation from optimal diet theory, energy intake was increased by selection only during winter, whereas protein intake was increased at the expense of energy intake at all other times. Growth rates of the two fishes predicted by a growth curve based on the relative amounts of protein and energy assimilated from their diets were at or just above maintenance levels in winter and higher during other times of the year. The additional species of algae and the small amounts of animal material in the diets of the two fishes appeared not to alter substantially the predicted annual growth pattern. Although Cebidichthys violaceus and Xiphister mucosus seem to be able to grow during most times of the year on a completely algal diet, their rates of energy consumption, which in turn regulate protein intake, are unknown and must be examined before more definite conclusions can be drawn about the growth of these fishes.

Dietary selectivity in the field and food preferences in the laboratory for two herbivorous fishes (Cebidichthys violaceus and Xiphister mucosus) from a temperate intertidal zone

Diets and food selectivity of two stichaeid fishes (Cebidichthys violaceus and Xiphister mucosus) from the rocky intertidal zone of the central California coast (USA) were studied at each season of the year by gut content analysis and abundance measurements of potential macrophyte food items. Both fishes, after reaching a standard length of about 44 mm, were almost exclusively herbivorous. The bulk of the diet consisted of 8 to 10 species of chlorophytes and rhodophytes. These main dietary components were chiefly annual seaweeds with high surfaceto-weight ratios (sheetlike forms or small, highly branched forms). Perennial seaweeds were eaten in relatively large amounts only during the winter. Macrophytes eaten in only trace amounts included about 20 species of chlorophytes, chrysophytes, phaeophytes, rhodophytes and a spermatophyte. The small amount of animal material in the diet (never more than 2% by weight) could well have been ingested incidentally while the fishes were feeding on seaweeds. Food preference tests with up to 19 macrophyte species in the laboratory revealed that both fishes chose to eat three annual rhodophytes (Smithora naiadum, Porphyra perforata and Microcladia coulteri) in preference to Ulva lobata, an annual chlorophyte that was more abundant in the diets of field-caught specimens.