Changes in amphipod densities among macroalgal habitats in day versus night collections along the Western Antarctic Peninsula

Amphipods along the western Antarctic Peninsula appear to gain refuge from predators by associating with chemically defended macroalgae rather than palatable macroalgae. However, nothing is known about amphipod activity at night. If foraging on non-chemically defended macroalgae regularly occurs, then nocturnal foraging seems beneficial since visual predators are disadvantaged. To test this hypothesis, we collected three common macroalgal species and affiliated mesograzers, approximately 3 h before and after sunset. All associated mesofauna were counted and densities recorded. Amphipod densities were significantly decreased during the night on the chemically defended Desmarestia menziesii, while significantly increased on the palatable Iridaea cordata. Additionally, the amphipod Gondogeneia antarctica was found in significantly higher densities at night on Palmaria decipiens, a species shown to be readily eaten by G. antarctica and omnivorous fish. We believe that chemically defended macroalgae act as a refuge for mesograzers during the day, while more widespread foraging occurs at night.     

A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment.     

Effects of predation risk and hunger on the behaviour of two species of tadpoles

Predation and hunger are threats for most organisms, and appropriate behavioural responses to both factors should be shaped by natural selection. In combination, however, the behavioural demands of predation avoidance and effective foraging often cannot be satisfied at the same time and lead to a conflict within organisms. We examined the behavioural responses of two closely-related species of tadpoles, Rana lessonae and R. esculenta, to simulated predation by fish and hunger. Tadpoles, hatched and reared in the laboratory, were tested in a three-way factorial (predation risk × hunger × species) experiment with four predation levels and four hunger levels. Both species decreased their swimming activity with increasing predation risk. Predation risk did not influence the amount of activity time invested in feeding but caused the tadpoles to spend less time in patches with food. Refuges were not used to avoid predation. R. esculenta was more sensitive to predation risk than R. lessonae. Hunger increased both the activity of tadpoles and the amount of activity time invested in feeding, thus indicating an increased energy intake. No interactions were observed between predation risk and hunger. These results show that tadpoles possess genetically-based behavioural mechanisms that allow them to respond in a graded manner to predation and hunger. However, they did not balance the two conflicting demands of predation avoidance and effective foraging; the two mechanisms appeared to act independently. Correspondence to: R.D. Semlitsch     

Contrasting home-range size and spatial partitioning in cryptic and sympatric pipistrelle bats

The extent of spatial partitioning in insectivorous bats, whose prey is patchily distributed and transient in nature, remains a contentious issue. The recent separation of a common Palaearctic bat, the pipistrelle, into Pipistrellus pipistrellus and Pipistrellus pygmaeus, which are morphologically similar and sympatric, provides an opportunity to examine this question. The present study used radio telemetry to address the spatial distribution and foraging characteristics of P. pipistrellus and P. pygmaeus in northeast Scotland, to test the hypothesis that coexistence between these species is facilitated through spatial segregation. We reveal large and significant differences in the spatial distribution and foraging characteristics of these two cryptic species. Individual P. pipistrellus home ranges were on average three times as large as that of P. pygmaeus, and they foraged for approximately an hour longer each night. Inter-specific spatial overlap was minimal (<5%) and core foraging areas of either species were essentially mutually exclusive despite the proximity of the two roosts. Inter-specific differences in range size were associated with the spatial dispersion of productive foraging sites within individual foraging ranges. P. pipistrellus foraging sites were highly dispersed, necessitating larger ranges. It is predicted that the spatial segregation revealed by the present study is a result of selection favouring the avoidance of competition in these species through differential habitat use.     

At-sea activity patterns of breeding and nonbreeding white-chinned petrels <Emphasis Type="Italic">Procellaria aequinoctialis</Emphasis> from South Georgia

Despite the recent burgeoning in predator tracking studies, few report on seabird activity patterns, despite the potential to provide important insights into foraging ecology and distribution. In the first year-round study for any small petrel, we examined the activity patterns of the white-chinned petrel Procellaria aequinoctialis based on data from combination geolocator-immersion loggers deployed on adults at South Georgia. The petrels were highly nocturnal, flying for greater proportions of darkness than any large procellarid studied so far, except the light-mantled albatross Phoebetria palpebrata. Flight bout durations were short compared with other species, suggesting a dominant foraging mode of small-scale searching within large prey patches. When migrating, birds reduced the proportion of time on the water and increased flight bout duration. Activity patterns changed seasonally: birds flew least during the nonbreeding period, and most frequently during chick-rearing in order to meet higher energy demands associated with provisioning offspring. The degree of their response to moonlight was also stage dependent (greatest in nonbreeding, and weakest in incubating birds), a trait potentially shared by other nocturnal petrels which will have repercussions for feeding success and prey selection. For the white-chinned petrel, which is commonly caught in longline fisheries, these results can be used to identify periods when birds are most susceptible to bycatch, and therefore when use of mitigation and checking for compliance is critical.     

Adaptive peaks and alternative foraging tactics in brook charr: evidence of short-term divergent selection for sitting-and-waiting and actively searching

Some recently emerged brook charr (Salvelinus fontinalis) inhabiting still-water pools along the sides of streams are sedentary and eat crustaceans from the lower portion of the water column. Others are more active and eat insects from the upper portion of the water column. We provide evidence that this divergent foraging behavior reflects short-term divergent selection brought about by intraspecific competition in the presence of alternative food sources. Rates of encounters and interactions between individuals were density dependent, and encounter and interaction events were closely timed with prey capture attempts. In addition, aggressive fish made more foraging attempts per minute than nonaggressive fish. Aggressive fish were also either inactive or very active, while nonaggressive fish exhibited intermediate levels of activity. Growth rate potential, an important component of fitness during the early life stages of brook charr, was assessed using tissue concentrations of RNA and found to be highest for sedentary fish and for active fish making frequent foraging attempts, and lower for fish exhibiting intermediate levels of activity. Our findings support contentions that individual behavior plays an important role during initial steps in the evolution of resource polymorphisms. Received: 27 July 1998 / Accepted after revision: 16 November 1998     

Foraging in the seed-harvester ant genus Pogonomyrmex: are energy costs important?

Energy intake and expenditure on natural foraging trips were estimated for the seed-harvester ants, Pogonomyrmex maricopa and P. rugosus. During seed collection, P. maricopa foraged individually, whereas P. rugosus employed a trunk-trail foraging system. Energy gain per trip and per minute were not significantly different between species. There was also no interspecific difference in energy cost per trip, but energy cost per minute was lower for P. maricopa foragers because they spent on average 7 min longer searching for a load on each trip. Including both unsuccessful and successful foraging trips, average energy gain per trip was more than 100 times the energy cost per trip for both species. Based on this result, we suggest that time cost incurred during individual foraging trips is much more important than energy cost in terms of maximizing net resource intake over time. In addition, because energy costs are so small relative to gains, we propose that energy costs associated with foraging may be safely ignored in future tests of foraging theory with seed-harvesting ant species.     

Primary versus secondary drivers of foraging activity in sandeel schools (<Emphasis Type="Italic">Ammodytes tobianus</Emphasis>)

The commercially and ecologically valuable sandeel (Ammodytes ssp.) make distinct vertical shifts between an inactive stage, during which they seek refuge in the sand, and a pelagic schooling stage, during which they forage. This characteristic discontinuous foraging pattern constitutes a challenge to fishery biologists and has consequences for a wide range of predators ranging from birds and mammals to commercially important species. However, experimental studies that shed light on the primary drivers of foraging activity in fish are rare. In the present study, whole schools of sandeel (A. tobianus) were caught in August in east Denmark (65°02′30N; 12°37′00E) and kept in large tanks in the laboratory. It was found that the amount of food ingested and memory of past days feeding history are primary drivers of foraging activity at the level of the entire school, whereas external factors such as prey concentration and temperature are merely secondary drivers.     

Collective foraging decision in a gregarious insect

Group foraging by eusocial insects implies sophisticated recruitment processes that often result in collective decisions to exploit the most profitable sources. These advanced levels of cooperation, however, remain limited to a small range of species, and we still know little about the mechanisms underlying group foraging behaviours in the great mass of animals exhibiting lower levels of social complexity. In this paper, we report, for the first time in a gregarious insect, the cockroach Blattella germanica (L.), a collective foraging decision whereby the selection of food sources is reached without requiring active recruitment. Groups of cockroaches given a binary choice between identical food sources exhibited exploitation asymmetries whose amplitude increases with group size. By coupling behavioural observations to computer simulations, we demonstrate that selection of food sources relies uniquely on a retention effect of feeding individuals on newcomers without comparison between available opportunities. This self-organised pattern presents similarities with the foraging dynamics of eusocial species, thus stressing the generic dimension of collective decision-making mechanisms based on social amplification rules despite fundamental differences in recruitment processes. We hypothesise that such parsimony could apply to a wide range of species and help understand the emergence of collective behaviours in simple social systems.     

Foraging behaviour of sympatric Antarctic and subantarctic fur seals: does their contrasting duration of lactation make a difference?

The duration of periods spent ashore versus foraging at sea, diving behaviour, and diet of lactating female Antarctic (Arctocephalus gazella, AFS) and subantarctic (A. tropicalis, SFS) fur seals were compared at Iles Crozet, where both species coexist. The large disparity in lactation duration (SFS: 10 months, AFS: 4 months), even under local sympatry, has led to the expectation that AFS should exhibit higher foraging effort or efficiency per unit time than SFS to allow them to wean their pups in a shorter period of time. Previous evidence, however, has not supported these expectations. In this study, the distribution of foraging trip durations revealed two types of trips: overnight (OFT, <1 day) and long (LFT, >1 day), in common with other results from Macquarie Island. However, diving behaviour differed significantly between foraging trip types, with greater diving effort in OFTs than in LFTs, and diving behaviour differed between fur seal species. OFTs were more frequent in SFS (48%) than in AFS (28%). SFS performed longer LFTs and maternal attendances than AFS, but spent a smaller proportion of their foraging cycle at sea (66.2 vs. 77.5%, respectively). SFS dove deeper and for longer periods than AFS, in both OFTs and LFTs, although indices of diving effort were similar between species. Diel variation in diving behaviour was lower among SFS, which foraged at greater depths during most of the night time available than AFS. The diving behaviour of AFS suggests they followed the nychthemeral migration of their prey more closely. Concomitant with the differences in diving behaviour, AFS and SFS fed on the same prey species, but in different proportions of three myctophid fish (Gymnoscopelus fraseri, G. piabilis, and G. nicholsi) that represented most of their diet. The estimated size of the most important fish consumed did not vary significantly between fur seal species, suggesting that the difference in dive depth was mostly a result of changes in the relative abundance of these myctophids. The energy content of these fish at Iles Crozet may thus influence the amount and quality of milk delivered to pups of each fur seal species. These results contrast with those found at other sites where both species coexist, and revealed a scale of variation in foraging behaviour which did not affect their effort while at sea, but that may be a major determinant of foraging efficiency and, consequently, maternal investment.     

Importance of ectoparasites and mucus in cleaning interactions in the Mediterranean cleaner wrasse Symphodus melanocercus

In fish cleaning associations, the net benefits gained by clients and cleaners from cleaning have still not been clearly evaluated. In particular, the role of ectoparasitism and the importance of client mucus characteristics remain unclear for most cleaner fish species. This paper investigates the cleaning behaviour of the Mediterranean cleaner wrasse Symphodus melanocercus, based on observations, cleaner gut contents, client ectoparasites and mucus characteristics. We showed that this fish is a specialised cleaner fish, similar to some other tropical cleaner species. Gnathiid isopod larvae and caligid copepods represented a large proportion of the items preyed on by S. melanocercus. Although their feeding activity was related to their client ectoparasite load, it was also significantly linked to client mucus load, which would indicate that the cleaning behaviour of S. melanocercus is not purely altruistic. Finally, as client visit to cleaning stations is related to their ectoparasitism, we propose that ectoparasite removal is likely to be a benefit for the client fishes of S. melanocercus. Received: 28 July 2000 / Accepted: 8 November 2000     

Quantitative analysis of foraging behaviour in a field population of the South American sun-star Heliaster helianthus

Foraging behaviour in the South American sunstar Heliaster helianthus (Lamarck) was investigated quantitatively on a subtropical rocky shore in central Peru. H. helianthus feeds mainly on two mussel species, Semimytilus algosus (Gould) and Perumytilus purpuratus (Lamarck). A sequence of foraging behaviour was described and observations were made of the timing of attacks on mussel beds by sun-stars. Although H. helianthus is capable of foraging out of water, an unusual trait for asteroids, the timing of such foraging appears to be well adjusted to avoid the risk of prolonged heat and desiccation. Foraging activity began 4 h 52 min before high tide and 3 h 19 min after high tide, with a mode between 2 h 30 min and 2 h before high tide. Fifty-two percent of all foraging activity began between 3 h 30 min and 2 h before high tide, while only 12.4% began after high tide. This suggests that H. helianthus mainly relies upon changes in the rate of tidal increase as a cue to begin foraging. Foraging activity ceased between 3 h 40 min before and 4 h 29 min after high tide, with 48% ending between 2 and 0.5 h before high tide. The duration of foraging ranged from 19 to 190 min, with values between 30 and 80 min accounting for 67.4% of all observations. The median duration was 62 min. No significant correlation was detected between the time when foraging activity commenced and its duration. The intensity of foraging activity varied on consecutive days, with a general pattern of decreasing intensity after a day of relatively high activity. Foraging location in relation to a mussel bed was analysed on a marked, 8 m stretch of rocky shore. The numbers of foraging H. helianthus observed on different sections of the shore were related neither to the width of the mussel zone nor to the vertical position of the lower edge of the mussel zone, indicating that sun-stars do not rely upon these factors to assess prey availability and that ideal free distribution with regard to prey abundance does not occur on the spatial scale examined.     

Ontogeny of predator-sensitive foraging and routine metabolism in larval shorthorn sculpin, <Emphasis Type="Italic">Myoxocephalus scorpius</Emphasis>

Most animals will reduce foraging activity in the presence of a predatory threat. However, little is known about the onset of this decision-making ability during the early life stages of fishes, and how the trade-off between foraging and predator-avoidance may be affected by changes in metabolic demand during ontogeny. To examine these issues, the foraging behaviour of larval shorthorn sculpin Myoxocephalus scorpius was monitored during visual exposure to a predatory threat (juvenile Atlantic cod, Gadus morhua) throughout development at 3°C (March–April, 2004). Larvae did not respond to predatory exposure during the first week post-hatch, but thereafter showed drastic reductions in foraging activity when exposed to predators. During early development, the mass-specific routine metabolism of shorthorn sculpin larvae displayed a triphasic ontogeny and peaked during metamorphosis. This high mass-specific metabolic demand could make reduced foraging under predation threat very costly during this stage of development. To further investigate this possibility, additional experiments were performed (March–April, 2005) where larvae were reared with visual exposure to predators for 6 h day⁻¹ during the feeding period. At 7-week post-hatch, larvae exposed to predators were smaller (wet mass and SL), showed decreased levels of whole-body lipids and certain fatty acids, and experienced higher rates of mortality as compared to control larvae. In environments where abundant predators cause larval fish to reduce their foraging rate, growth and survival of larvae may be negatively affected. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Age-related shifts in the diet composition of southern elephant seals expand overall foraging niche

Southern elephant seals are important apex predators in a highly variable and unpredictable marine environment. In the presence of resource limitation, foraging behaviours evolve to reduce intra-specific competition increasing a species’ overall probability of successful foraging. We examined the diet of 141 (aged 1–3 years) juvenile southern elephant seals to test the hypotheses that differences between ages, sexes and seasons in diet structure occur. We described prey species composition for common squid and fish species and the mean size of cephalopod prey items for these age groups. Three cephalopod species dominated the stomach samples, Alluroteuthis antarcticus, Histioteuthis eltaninae and Slosarczykovia circumantarcticus. We found age-related differences in both species composition and size of larger prey species that probably relate to ontogenetic changes in diving ability and haul-out behaviour and prey availability. These changes in foraging behaviour and diet are hypothesised to reduce intra-specific food competition concomitant with the increase in foraging niche of growing juveniles.     

Aposematism in a soft-bodied insect: a case for kin selection

Summary This paper describes the influence on predator behaviour, and the survival of an aposematic aphid, Aphis nerii, in comparison with a palatable, cryptic aphid, Acyrthosiphon pisum, when offered to two predators with different foraging tactics. The experiments were designed to test Fisher's (1930) suggestion that aposematism could evolve by kin selection, since aposematic animals often occur in aggregations of relatives. Initially, spiders (Zygiella x-notata) and birds (Parus major) killed high proportions of distasteful A. nerii (60% and 54% respectively). With experience, the predators killed and ate fewer A. nerii. The decreasing mortality of A. nerii after initial encounters with predators, coupled with its apparently obligate parthenogenesis, indicate that the evolution of aposematism in this soft-bodied insect is consistent with kin selection.     

Survival trade-offs between two predator-induced phenotypes in Pacific treefrogs (Pseudacris regilla)

In many organisms, specific predator species induce defensive phenotypes that are qualitatively different from the phenotypes induced by other predator species. This differential induction implies that there is no optimal phenotype that works best against all predators. However, few studies have actually tested the hypothesis that each predator-induced phenotype provides the highest survival rate in encounters with the predator that induced that phenotype. In this experiment, I reared Pacific treefrog (Pseudacris regilla) larvae with chemical cues from two different predators (bluegill sunfish and predaceous diving-beetle larvae), and without predator cues. The Pacific treefrog larvae in the three treatments differed in their morphology and foraging behavior. I then exposed tadpoles from each treatment to free-foraging predaceous diving beetles and bluegill sunfish. Tadpoles survived best when exposed to the predator whose cues they were reared with, and worst when exposed to the other predator. In both predator environments, the tadpoles reared in the nonpredator control treatment had intermediate survival between the two predator-induced groups. Thus, there is no generalized "antipredator" response to these predators; rather, there was a clear trade-off in survival abilities between the predators.     

Vigilance behaviour and fitness consequences: comparing a solitary foraging and an obligate group-foraging mammal

Vigilance behaviour in gregarious species has been studied extensively, especially the relationship between individual vigilance and group size, which is often negative. Relatively little is known about the effect of conspecifics on vigilance in non-obligate social species or the influence of sociality itself on antipredator tactics. We investigated predator avoidance behaviour in the yellow mongoose, Cynictis penicillata, a group-living solitary forager, and compared it with a sympatric group-living, group-foraging herpestid, the meerkat, Suricata suricatta. In yellow mongooses, the presence of conspecifics during foraging—an infrequent occurrence—reduced their foraging time and success and increased individual vigilance, contrary to the classical group-size effect. Comparing the two herpestids, sociality did not appear to affect overt vigilance or survival rates but influenced general patterns of predator avoidance. Whereas meerkats relied on communal vigilance, costly vigilance postures, and auditory warnings against danger, yellow mongooses avoided predator detection by remaining close to safe refuges and increasing “low-cost” vigilance, which did not interfere with foraging. We suggest that foraging group size in herpestids is constrained by species-distinct vigilance patterns, in addition to habitat and prey preference.     

Shoal composition, body size and foraging in sticklebacks

Individuals which deviate from the majority in groups are likely to be most vulnerable to predation. This oddity effect, by definition, is frequency dependent, eventually fading at equal frequencies of the phenotypes in a group. It has been hypothesized that the increased predation risk of odd individuals may play an important role in the formation of phenotypically uniform shoals of fish. However, recent work has indicated that individuals may experience, or value, their predation hazard differently depending on their own size in relation to that of other group members: single large fish, but not small ones, appear concerned about their oddity in a shoal. Here I show that the apparent wariness of large fish is also expressed in a frequency-dependent manner, closely conforming to what is predicted if the oddity effect is responsible for their behavior. Using foraging activity of individuals as a means to evaluate their predation risk, I demonstrate with shoals comprising 12 threespine sticklebacks (Gasterosteus aculeatus) that large fish forage least actively when in a shoal consisting of 2 large and 10 small fish. An increase in the number of large fish to 4 among 8 small individuals clearly results in an increase in their foraging activity. However, having reached an equal frequency with small fish in a shoal, large fish do not seem to change their foraging activity much even when their number in a shoal increases further. In contrast, foraging activity of small sticklebacks remains fairly constant throughout the entire range of tested shoal compositions, providing further evidence that small and large fish respond to their oddity differently. Received: 12 February 1998 / Accepted after revision: 7 May 1998     

Foraging by the herbivorous parrotfish Sparisoma radians

The foraging behavior of the bucktooth parrotfish Sparisoma radians was studied in seagrass beds off St. Croix, US Virgin Islands and in laboratory preference tests. Thalassia testudinum was the dominant item in the field diet with the epiphytized distal portion of the blades most favored. Other seagrasses, Syringodium filiforme and Halodule wrightii, were taken in relation to their abundance. Several algae were also eaten, particularly Halimeda spp. and Penicillus spp. Although abundances of the algae varied, the fish maintained a steady mixture of plant species in their diet. Laboratory feeding tests for various plants presented in pairs showed clear preferences in nearly all cases. The preference hierarchy was (1) T. testudinum with epiphytes, (2) H. wrightii, (3) T. testudinum without epiphytes, (4) S. filiforme, and the algae, (5) Dictyota divaricata, (6) Enteromorpha flexuosa, (7) Caulerpa mexicana, (8) Halimeda incrassata and (9) Penicillis pyriformis. Preference did not vary significantly with satiation. The catch per unit effort (kilocalories absorbed per bite) was calculated for each plant, using (a) absorption values calculated for S. radians fed the different plants, (b) the calorific values for tested plants, and (c) an estimate of the amount of material taken per bite. The ranking of catch per unit effort closely paralleled the preference hierarchy with the exception of C. mexicana which has a toxin. Fish fed diets of single plants, mixed plant diet, and starved controls showed differential survival which paralleled the preference hierarchy, the most preferred plants leading to longest survival. Comparison of laboratory results with field feeding behavior shows that inclusion of plants in the diet is not related directly to preference rank, availability or survival value, but that the fish deliberately eat a variety of plants presumably which maintains balanced diet. This result indicates that models of optimal foraging for herbivores should include nutrient constraints and avoidance of toxins in order to predict accurately the behavior of an animal in the field.     

Trophic ecology of coral reef gobies: interspecific,ontogenetic, and seasonal comparison of diet and feeding intensity

In fishes, a small body size may facilitate cost-effective exploitation of various primary and secondary food resources, but may pose difficulties associated with digestion of plant material and finding sufficient food in a foraging area potentially restricted by a high risk of predation. We examined the trophic ecology of five common, small-bodied coral reef fish from the family Gobiidae. For each species, we determined diet composition, feeding bite rate, foraging substrate, and feeding behaviour, and examined whether diet composition and feeding bite rate changed ontogenetically and seasonally. The five species showed a diverse range of trophic modes: Amblygobius bynoensis and Amblygobius phalaena were herbivores, Valenciennea muralis was a carnivore, Asterropteryx semipunctatus a detritivore, and Istigobius goldmanni an omnivore. Both the herbivores and detritivore supplemented their diet with animal material. The consumption of a wide range of dietary resources by the two smallest species with the most restricted mobility (A. semipunctatus and I. goldmanni) may ensure energy requirements are met within a restricted foraging area. There was a significant difference in mean feeding bite rate among species, with carnivore > herbivore > omnivore > detritivore. None of the species exhibited an ontogenetic shift in diet composition or increase in feeding bite rate, indicating that (1) postmaturation growth is not facilitated by a higher quality diet or increased feeding intensity following maturation, and (2) their small body size does not preclude herbivory. The herbivores had the highest gut:fish length ratio, which may facilitate plant digestion. While diet did not change seasonally, the mean feeding bite rate was significantly lower in winter than summer for four of the study species.