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.     

Scale-dependent patterns of variability of a grazing parrotfish (<Emphasis Type="Italic">Leptoscarus vaigiensis</Emphasis>) in a tropical seagrass-dominated seascape

Although herbivorous fish form critical linkages between primary producers and higher trophic levels, the major factors regulating their spatial structure in seagrass systems remain poorly understood. The present study examined the parrotfish Leptoscarus vaigiensis in seagrass meadows of a tropical embayment in the western Indian Ocean. Stomach content analysis and direct field observations showed that L. vaigiensis is an efficient grazer, feeding almost exclusively on seagrass leaves. Seagrass shoot density was highly correlated to all density variables (total, juvenile and subadult) and juvenile biomass of L. vaigiensis, while subadult biomass was predicted by distance to neighbouring coral habitat. Moreover, density and biomass of predatory fish (piscivores) were predicted by seagrass canopy height and the distribution patterns of predators followed those of L. vaigiensis. Hence, factors at local (seagrass structural complexity and feeding mode) and landscape scale levels (seascape context and distribution of piscivores) likely mutually structure herbivorous fish communities. The findings underscore the importance of incorporating multiple scale-dependent factors when managing coastal seagrass ecosystems and their associated key species.     

Effect of macrophyte secondary metabolites on feeding preferences of the herbivorous parrotfish Sparisoma radians

Experiments were conducted to test the role of secondary metabolites in determining the natural feeding preference hierarchy of the bucktooth parrotfish Sparisoma radians. The two least preferred food genera of S. radians, Halimeda and Penicillus, both contain 1,4-diacetoxy-1,3-butadiene terpenes, while the most preferred species, Thalassia testudinum, does not. Experiments with agar cylinders containing macrophyte homogenates showed that macrophyte biteability was not a factor. Instead preference could be altered by the application of the diacetoxybutadiene containing terpenes 4,9-diacetoxyudoteal and caulerpenyne or fractions or extracts containing them at naturally occurring concentrations. Concentration of the active terpenes affected the intensity of the fish's preference for the control in pairwise comparisons. Extracts and fractions which did not contain 4,9-diacetoxyudoteal or caulerpenyne did not affect fish feeding preferences at naturally occurring concentrations. Experiments in which S. radians were given no plant choice showed that coating T. testudinum with H. incrassata organic crude extract reduced the number of bites consumed and the biomass consumed to a level equivalent to that obtained for H. incrassata plants.     

Seasonal grazing of Pseudocalanus minutus on particles

Grazing by Pseudocalanus minutus on naturally occurring partieles has been investigated over a 2 year period at 5 m depth, in a small coastal embayment in Nova Scotia, Canada. Large variations in the standing stock of particulate matter in the water from 5 m depth were associated with seasonal changes in phytoplankton, and at times with runoff from rivers. Each size group displayed seasonal changes, with high amplitudes occurring in the large-particle size ranges. P. minutus consumption was associated with seasonal changes in total particle concentration as well as with the concentration in each size group. The food uptake was correlated (P>0.01) with the particle concentration in all particle categories, except in the size range below 3.57 . Frequency of positive electivity indices increased with particle sizes up to 57 and then decreased. Observations on particle spectra revealed considerable seasonal variability in both particle sizes and copepod feeding patterns. P. minutus adapted to seasonal variations within the particle spectrum by shifting its grazing pressure from one size range to another. By taking advantage of every particle peak concentration, P. minutus revealed a strong opportunistic feeding behaviour, and a very efficient utilization of the standing stock. Maximum consumption was recorded in early spring, when P. minutus fed on large-size particles. Feeding took place in the medium and small-size ranges during the summer and part of fall. Food uptake was rarely less than 2.26% of body weight during the winter, and reached up to 55% and sometimes more in spring. The unselective feeding patterus demonstrated by P. minutus suggest certain ecological implications of feeding pressure on standing stock.     

Estimating the grazing impact of marine micro-zooplankton

This paper describes a dilution technique for estimating the micro-zooplankton grazing impact on natural communities of marine phytoplankton. Experiments performed in coastal waters off Washington, USA (October, 1980), yield estimates of micro-zooplankton impact equivalent to 6 to 24% of phytoplankton standing biomass and 17 to 52% of production per day. Indirect evidence suggests that most of this impact is due to the feeding of copepod nauplii and tintinnids; in contrast, non-loricate ciliates, comprising 80 to 90% of numerical abundance, appeared to contribute little to phytoplankton mortality.Contribution No. 1237 from the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Phylogeography of the Indo-Pacific parrotfish Scarus psittacus: isolation generates distinctive peripheral populations in two oceans

Phylogenetic, phylogeographic, population genetic and coalescence analyses were combined to examine the recent evolutionary history of the widespread Indo-Pacific parrotfish, Scarus psittacus, over a geographic range spanning three marine biogeographic realms. We sequenced 164 individuals from 12 locations spanning 17,000 km, from 55oE to 143oW, using 322 base pairs of mitochondrial control region (D-loop). S. psittacus displayed high haplotype (h = 0.83–0.98), but low nucleotide (<1%) diversity. Most (>83%) genetic variation was within populations. AMOVA revealed significant partitioning and identified five geographic groups. These included one central population and four populations peripheral to the centre. The central population occupied reefs from Western Australia to Tahiti and represented the central Indo-Pacific biogeographic realm. Cocos Keeling was distinct from central and western Indo-Pacific biogeographic realms occupying a position intermediate to these. Peripheral populations (Hawaii, Marquesas) represented the eastern Indo-Pacific biogeographic realm, while Seychelles represented the western Indo-Pacific biogeographic realm. All but the central population expanded (<163 kya). Whilst all populations experienced major sea level and SST changes associated with Pleistocene glaciation cycles, the genetic structure of the central population was relatively homogenous unlike the remaining genetically distinctive populations.     

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     

Creation and preservation of vegetation patterns by grazing

Structural patterns of tall stands (“tussock”) and short stands (“lawn”) are observed in grazed vegetation throughout the world. Such structural vegetation diversity influences plant and animal diversity. A possible mechanism for the creation and preservation of such patterns is a positive feedback between grazing and plant palatability. Although some theoretical studies have addressed this point in a non-spatial setting, the spatial consequences of this feedback mechanism on the stability and spatial characteristics of vegetation structure patterns have not been studied.We addressed this issue by analyzing a spatially explicit individual-based plant-grazer simulation model, based on published empirical relations and the assumption of optimal foraging.In the model, the selection by the grazer of short stands (that have a higher energy content and are more palatable) is affected by traveling costs and the spatial organization of swards. Nevertheless, the most selected biomass in this type of short stands was the optimal biomass predicted by cropping and digestion constraints. As a result of the optimal foraging strategy, the grazers displayed Lévy-flight traveling behavior during the simulations with characteristic exponent μ ≈ 2.Patterns of short and tall stands created by grazing were preserved for at least a decade. Even in seasonal habitat, the spatial organization of the patterns remained relatively constant, despite fluctuations in the area of short stands. Heterogeneity of initial vegetation increased heterogeneity of the grazing-induced pattern, but did not affect its stability.The area of short stands that was preserved by grazing scaled with the herbivore mass to the power 0.4 and with the carrying capacity of the vegetation to the power −0.75.Patterns of tall and short stands can be created and perpetuated by optimally grazing ruminants, irrespective of possible underlying soil patterns. The simulations generate predictions for the stability and spatial characteristics of such structural vegetation patterns.     

Mortality estimates from repeated visual censuses of a parrotfish (Sparisoma viride) population: demographic implications

Repeated visual censuses of different categories (juveniles, females, territorial and group males) of the stoplight parrotfish (Sparisoma viride, a protogynous hermaphrodite) over a 3-year period indicated a relatively stable size and structure of the adult population. This allowed estimates of size-specific mortality, sex change, and territory acquisition probabilities from previously reported growth rates. Comparison of the predicted number of survivors, sex changers, and territory take overs with field observations indicates that our estimates are quite reliable. However, rather large differences in mortality are obtained for the largest three size classes (>25 cm), which may be due to reduced accuracy of length estimates of large fish. A pooled mortality for these classes is therefore suggested as a more realistic estimate. The life-history implications of our findings are investigated by comparing the predicted survival and future reproductive success of fish that change sex at different sizes. Ten percent of the adults reach an age of ca. 17 years, once a length of 20 cm is attained, as predicted from the pooled estimates. Calculations based on these estimates indicate that the predicted reproductive output of a 15-year-old fish (2500 matings) is the same for early (i.e. at 20 cm) and late (>30 cm) sex changers and for an average fish subject to the estimated daily sex change and territory acquisition probabilities. These findings suggest that S. viride individuals are able to flexibly adjust the timing of sex change in an adaptive way to unpredictable local conditions. However, independent mortality estimates are needed to corroborate our present findings. Received: 2 January 1997 / Accepted: 27 January 1997     

Herbage dynamics of some semi-arid Indian grazing lands

This paper reports a time-varying coefficient, open, linear compartment model of herbage dynamics for some semi-arid Indian grazing lands with different grazing treatments. The objectives were to arrive at suitable transfer coefficients in order to approximate the pattern of changes in the compartment values and to indicate the time intervals in which these coefficients are most likely to vary. The modelling endeavour fulfilled these objectives and indicated some underlying differences in ecosystem functioning on these grazing lands.     

Genetic connectivity and historical demography of the blue barred parrotfish (Scarus ghobban) in the western Indian Ocean

Studies on genetic connectivity are essential for the design of management strategies for coral reef fisheries. In this study we used a mitochondrial DNA marker to investigate population structure of the reef-associated parrotfish, Scarus ghobban, from four countries, Kenya, Mauritius, Seychelles and Tanzania, in the western Indian Ocean. We obtained nucleotide sequences of the mitochondrial control region for 117 individuals. Measures of haplotype diversity were relatively high. Pairwise population differentiation (F _ST) was low, but not always non-significant. Analysis of molecular variance (AMOVA) showed genetic differentiation between groups, when the data was partitioned into two groups consisting of samples from Mauritius and Tanzania in one group, and samples from Kenya and Seychelles in another group. Direction of gene flow was estimated using a Bayesian approach. Migration was sometimes asymmetric or directional, coinciding with the flow of major oceanic and coastal currents in the region. Mismatch distributions, based on the observed number of differences among haplotype pairs, produced a unimodal distribution, indicative of recent demographic expansion. Phylogenetic analyses revealed three clades without any geographic structure, suggesting recent migration between historically isolated lineages. We reconstructed the historical demography of S. ghobban and examined it in the context of Pleistocene climate stages and changes in relative sea level. Overall, these results showed that populations of S. ghobban are genetically diverse and have relatively high gene flow, with some genetic structuring in the western Indian Ocean.     

Estimating influence of stocking regimes on livestock grazing distributions

Livestock often concentrate grazing in particular regions of landscapes while partly or wholly avoiding other regions. Dispersing livestock from the heavily grazed regions is a central challenge in grazing land management. Position data gathered from GPS-collared livestock hold potential for increasing knowledge of factors driving livestock aggregation patterns, but advances in gathering the data have outpaced advancements in analyzing and learning from it. We fit a hierarchical seemingly unrelated regression (SUR) model to explore how season of stocking and the location where cattle entered a pasture influenced grazing distributions. Stocking alternated between summer on one side of the pasture one year and fall on another side of the pasture the next year for 18 years. Waypoints were recorded on cattle for 50 d each year. We focused our analysis on the pasture's 10 most heavily grazed 4-ha units, because these units were the most prone to negative grazing impacts. Though grazing of the study units was always disproportionately heavy, it was much heavier with the summer than fall stocking regime: Bayesian confidence intervals indicate summer grazing of study units was approximately double the average fall grazing value. This is our core result, and it illustrates the strong effect stocking season or date or both can have on grazing distributions. We fit three additional models to explore the relative importance of stocking season versus location. According to this analysis, stocking season played a role, but stocking location was the main driver. Ostensibly minor factors (e.g. stocking location) can greatly influence livestock distributions.     

In situ grazing rates of deep-sea benthic boundary-layer zooplankton

In situ grazing rates of mixed assemblages of deep-sea benthic boundary-layer zooplankton were measured in July 1983 at a site in the North Atlantic Ocean at 2 175 m depth using a short-term radioisotope-uptake method. Zooplankton were collected with an opening-closing net system from the bottom 1 m of the ocean and incubated in situ with mixed tritiated amino acids in special cod-end chambers. Incubations were terminated at depth by the addition of MS-222. Radioisotope uptake beyond that of dead controls was shown by both the zooplankton and particulate fractions. Grazing rates in the deep-sea experiments were surprisingly high, being comparable to wintertime Narragansett Bay zooplankton grazing rates determined in a separate series of laboratory experiments. These laboratory experiments also documented nonparticle-associated uptake of dissolved amino acids by Narragansett Bay zooplankton, but the importance of this in the deep sea is unknown. The deep-sea benthic boundary layer may be a region of elevated rates and activity because of its higher particulate concentrations, and our experiments may also have measured maximum rather than average rates.     

Selective grazing by zooplankton populations in Southampton Water

Experiments were made with natural populations of both zooplankton and phytoplankton from Southampton Water (England), using an electronic particle counter to measure grazing effects. What appear to have been large differences within and between experiments in size of particles selected are shown to have been mainly a result of variations in total amout of grazing. When this was taken into account it was found that there was no difference in size of particles selected most strongly whether the zooplankton was dominated by copepods or smaller cirripede nauplii. This can be accounted for by similarities in the setous feeding appendages of these two forms. It is suggested that similarity in particle selection would lead to competition between the two grazers if they were not separated seasonally.     

Prey risk allocation in a grazing ecosystem.

Understanding the behaviorally mediated indirect effects of predators in ecosystems requires knowledge of predator-prey behavioral interactions. In predator-ungulate-plant systems, empirical research quantifying how predators affect ungulate group sizes and distribution, in the context of other influential variables, is particularly needed. The risk allocation hypothesis proposes that prey behavioral responses to predation risk depend on background frequencies of exposure to risk, and it can be used to make predictions about predator-ungulate-plant interactions. We determined non-predation variables that affect elk (Cervus elaphus) group sizes and distribution on a winter range in the Greater Yellowstone Ecosystem (GYE) using logistic and log-linear regression on surveys of 513 1-km2 areas conducted over two years. Employing model selection techniques, we evaluated risk allocation and other a priori hypotheses of elk group size and distributional responses to wolf (Canis lupus) predation risk while accounting for influential non-wolf-predation variables. We found little evidence that wolves affect elk group sizes, which were strongly influenced by habitat type and hunting by humans. Following predictions from the risk allocation hypothesis, wolves likely created a more dynamic elk distribution in areas that they frequently hunted, as elk tended to move following wolf encounters in those areas. This response should dilute elk foraging pressure on plant communities in areas where they are frequently hunted by wolves. We predict that this should decrease the spatial heterogeneity of elk impacts on grasslands in areas that wolves frequently hunt. We also predict that this should decrease browsing pressure on heavily browsed woody plant stands in certain areas, which is supported by recent research in the GYE.     

Foraging activity and resource use by three parrotfish species at Fernando de Noronha Archipelago, tropical West Atlantic

Parrotfishes are important components of the herbivore and detritivore guilds of tropical and subtropical reefs. Most of parrotfish species are protogynous hermaphrodites that change colour and sex, from initial phase females or males (IP) to terminal phase males (TP). We studied the foraging behaviour of Sparisoma amplum, S. axillare and S. frondosum, three syntopic scarids on the rocky reefs of Fernando de Noronha Archipelago, Tropical West Atlantic. The three parrotfish species differed in food selection and preference, but IP and TP individuals of the same species preferred the same food types, except for S. amplum. Feeding rates of IP individuals were higher than those of TP individuals, but the distribution of feeding frequencies throughout the day of IP and TP individuals of the same species was similar. IP individuals had higher feeding rates than TP ones, which seems related to the fact that TP individuals spend a large amount of time patrolling their territories and chasing away conspecific individuals at the study site. The general foraging pattern we found for S. amplum, S. axillare and S. frondosum is similar to patterns found for other parrotfish species in the Western Atlantic.     

The effect of grazing on biodiversity in coastal dune heathlands

The effect of sheep grazing on species richness, higher order diversity measures, inequality, species composition, functional diversity and allometric relationships at a coastal dune heathland site was investigated. After a prescribed fire in 2002, the site was divided into two parts, where one of the parts was unmanaged and the other part was fenced and grazed by sheep. Sheep grazing had a positive effect on species richness as well as a significant positive effect on the functional diversity at the coastal dune heathland site. Generally, the cover of dwarf shrubs was negatively affected by grazing, whereas the cover of sedges and grasses was positively affected by grazing. There is a need for comparative investigations of the effect of different management methods not only on floristic biodiversity, but on all relevant kinds of biodiversity as well as on soil structure, soil chemistry and habitat micro- and macrostructure. Consequently, we advocate the initiation of an international systematic investigation of the effect of different management methods.     

Nutrition and grazing behavior of the green turtle Chelonia mydas

The apparent digestibility coefficients for 4 size classes of the green turtle Chelonia mydas feeding on the seagrass Thalassia testudinum were measured in Union Creek, Great Inagua, Bahamas, from September 1975 to August 1976. The values ranged from 32.6 to 73.9% for organic matter; from 21.5 to 70.7% for energy; from 71.5 to 93.7% for cellulose; from 40.3 to 90.8% for hemicellulose; and from 14.4 to 56.6% for protein. Digestive efficiency increased with increases in water temperature and body size. There was no seasonal variation in the nutrient composition of T. testudinum blades. Grazing on T. testudinum may be limited by its low quality as a forage, a result of its high fiber content and possible low protein availability. Turtles did not graze at random over the extensive beds of T. testudinum, but maintained grazing plots of young leaves by consistent recropping. They thus consumed a more digestible forage-higher in protein and lower in lignin-than the ungrazed, older leaves of T. testudinum. The selectivity of green turtles for either a seagrass or algal diet may reflect the specificity of their intestinal microflora.     

Impact of copepod grazing on the red-tide flagellate Chattonella antiqua

Although planktonic copepods are major suspension feeders in the sea, the impact of their grazing pressure upon red-tide flagellates has not been fully investigated. In the present study, the grazing of adult females of several copepod species is examined using three food types: viz. natural suspended particles, natural suspended particles mixed with cultured Chattonella antiqua, and cultured C. antiqua. The functional response on C. antiqua was investigated for five species of copepods (Acartia erythraea, Calanus sinicus, Centropages yamadai, Paracalanus parvus and Pseudodiaptomus marinus). Ingestion rates increased linearly with increasing cell concentrations until a maximum level was reached, beyond which the rates were constant. This cell concentration was higher for larger copepods. The weight-specific maximum ingestion rates were higher in the small species. In general, copepods tended to feed selectively on larger particles when feeding on natural particles. This tendency was strongest in a simulated red-tide environment. Thus, it can be surmised that copepods may selectively graze on C. antiqua during the outbreak of a red tide. Grazing pressure by the natural copepod community in Harima Nada, the Inland Sea of Japan, was calculated by integration of the laboratory determined feeding rates and field measurements of zooplankton biomass. The daily removal rate was 3.4 to 30.8% (mean: 12.3%) of C. antiqua biomass at 20 cells ml^-1 and decreased to 0.6–4.3% (mean: 1.8%) at 500 cells ml^-1. Therefore, the grazing pressure by the copepod community is important at the initial stage of the red tide.     

Calculation and use of selectivity coefficients of feeding: Zooplankton grazing

A straightforward method for calculating selectivity coefficients (W_ij) of predation from raw data, mortality rates of prey, filtering rates, feeding rates and electivity indices is derived. Results from a comparison of selectivity coefficients for the copepod Diaptomus oregonensis grazing under a number of experimental conditions suggest that W_ij's for size-selective feeding are invariant, a conclusion also supported by the leaky-sieve model. Recommendations are made on how to use W_ij's in linear and nonlinear feeding constructs for zooplankton and other animals.