Herbivorous amphipods inhabit protective microhabitats within thalli of giant kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>

Many small marine herbivores utilize specific algal hosts, but the ultimate factors that shape host selection are not well understood. For example, the use of particular microhabitats within algal hosts and the functional role of these microhabitats have received little attention, especially in large algae such as kelps. We studied microhabitat use of the herbivorous amphipod Peramphithoe femorata that inhabits nest-like domiciles on the blades of giant kelp Macrocystis pyrifera. The vertical position of nest-bearing blades along the stipe of the algal thallus and the position of the nests within the lateral blades of M. pyrifera were surveyed in two kelp forests in northern-central Chile. Additionally, we conducted laboratory and field experiments to unravel the mechanisms driving the observed distributions. Peramphithoe femorata nests were predominantly built on the distal blade tips in apical sections of the stipes. Within-blade and within-stipe feeding preferences of P. femorata did not explain the amphipod distribution. Amphipods did not consistently select distal over proximal blade sections in habitat choice experiments. Mortality of tethered amphipods without nests was higher at the seafloor than at the sea surface in the field. Nests mitigated mortality of tethered amphipods, especially at the seafloor. Thus, protective microhabitats within thalli of large kelp species can substantially enhance survival of small marine herbivores. Our results suggest that differential survival from predation might be more important than food preferences in determining the microhabitat distribution of these herbivores.     

Geographic covariation of chemical quality of the host alga<Emphasis Type="Italic"> Fucus vesiculosus</Emphasis> with fitness of the herbivorous isopod<Emphasis Type="Italic"> Idotea baltica</Emphasis>

Environmental and/or genetic among-site variation in plant quality may influence growth and fecundity of specialized herbivores inhabiting a particular site. Such variation is important as it generates spatial variation in selection for traits related to plant–herbivore interaction. Littoral macroalgae are known to respond plastically to environmental variation by modifying their chemistry or morphology. We studied geographic variation in phlorotannin, nitrogen, protein, and sugar (fucose, mannitol, and melibiose) concentrations of the brown alga Fucus vesiculosus at 12 sites separated by 0.5 to 40 km in the naturally fragmented Archipelago Sea in the northern Baltic Sea. By this regional variation in algal chemistry we attempted to explain among-population variation in size and fecundity of the crustacean herbivore Idotea baltica. We observed high spatial variation in all the measured chemical characteristics of F. vesiculosus, as well as in female size and the number of eggs produced by the herbivores. Spatial variation in nitrogen or protein contents of the alga did not explain the variation of herbivore traits. However, egg size positively covaried with spatial variation in the concentration of mannitol, the major storage carbohydrate of the alga. Such a positive relationship may arise if I. baltica can utilize the nutritive value of a mannitol-rich diet thereby being better able to provision the developing eggs with energy-rich metabolites. Unexpectedly, the concentration of phlorotannins, secondary metabolites having a putative role in defense against herbivory, positively covaried with the size of the herbivore. Among-population variation in host plant chemistry and covariation of that with herbivore growth and reproduction imply that herbivores respond to the local quality of their host plants, and that geographical structuring of populations has to be taken into account in studies of plant–herbivore interactions.Communicated by M. Kühl, Helsingør     

Competition, predation and coexistence in a three trophic system

Simple ecological models that mostly operate with population densities using continuous variables, explain quite well the behavior of real populations. In this work we propose and discuss the continuous dynamics of a system of three species, which belongs to the well-known family of Lotka–Volterra models. In particular, the proposed model includes direct effects such as predation and competition among species, and indirect effects such as refuge. The model is proposed to explain recent studies about a group of crustacean (amphipods of genus Hyallela) found in all the plain streams and shallow lakes of the American continent. The studied system includes three compartments: algae, a strictly herbivore amphipod and an omnivore (herbivore and carnivore) one. The analysis of the model shows that there are stable extinction equilibria throughout all the parameters’ space. There are also equilibria with stable coexistence of the three species and two interesting binary equilibria: one with stable coexistence of algae and herbivore and other with coexistence between algae and omnivore amphipods. The presence of Allee effect in the algae growth and the existence of refuge for the herbivore amphipod (prey) determine a bottom-up control.     

Differences in herbivore preferences, phlorotannin production, and nutritional quality between juvenile and adult tissues from marine brown algae

 Juvenile and adult marine organisms differ in their morphology, chemistry, physiology, behavior, and ecology. Because juvenile algae are thinner, smaller, and have more delicate tissues than adults, they are often assumed to be more susceptible to grazers. We examined within-species food preferences of four common generalist herbivores for juvenile and adult tissues of eight common brown algae in two-choice laboratory food-preference experiments. Our results showed that juvenile algae did not tend to be a preferred food of herbivores. Juvenile tissues were significantly preferred over adult tissues in only four of the 32 combinations of algae and herbivores tested. In 12 experiments, adult tissues were preferred over juvenile tissues, and no choice occurred in the remaining 16 experiments. When sea urchins exhibited a preference, it was always for adult tissues. The other three herbivores, an isopod and two snails, were more variable in their choices, sometimes preferring juveniles, sometimes adults, and sometimes having no preference. We measured nitrogen and phlorotannin concentrations in adult and juvenile seaweeds to see whether these parameters were correlated with herbivore food preferences. Nitrogen levels were similar in juveniles and adults of three algal species and were higher in juveniles of two. Phlorotannin levels were higher in juveniles of four species and lower in juveniles of one. The other three species showed no differences in phlorotannin levels. Phlorotannin concentrations decreased with increasing juvenile size in three species and increased with increasing size in one species. Neither nitrogen nor phlorotannin concentrations explained overall herbivore food preferences for algae of different stages. Our results suggest that preferences of certain grazers for juvenile algae are not as strong as previously assumed and are dependent on herbivore species. Preferences between juveniles and adults are likely to be determined by a combination of morphological and chemical features of the tissues and the unique responses of herbivore species to those features. Received: 10 April 2000 / Accepted: 19 November 2000     

Herbivory in the gammarid amphipod Aora typica: relationships between consumption rates, performance and abundance across ten seaweed species

This paper reveals a substantial capacity for herbivory of seaweeds in the gammarid amphipod Aora typica, adults eating seven of ten taxonomically and morphologically diverse seaweed species offered to them in a no-choice assay. The green algae Ulva spathulata and Enteromorpha intestinalis were consumed at the highest rates in both no-choice (2.3–2.5 mg blotted weight individual⁻¹ day⁻¹) and multiple-choice assays (0.5–1.3 mg blotted weight individual⁻¹ day⁻¹). Adult A. typica collected from two different species of brown seaweeds had very similar feeding preferences to each other. Juvenile A. typica grew to reproductive maturity on the green algae E. intestinalis and U. spathulata, and the brown algae Carpophyllum maschalocarpum and Ecklonia radiata. In common with previous studies on members of other amphipod families, survivorship of juvenile amphipods was positively correlated with feeding preferences of adults across seaweed species (r ²=0.43, P=0.04). However, densities of A. typica on seaweeds in the field (excluding the intertidal E. intestinalis and U. spathulata) were not significantly correlated with feeding preferences of adults (r ²=0.07, P=0.5) or survivorship of juveniles (r ²=0.17, P=0.31). This suggests that either host seaweeds are not a major dietary component of these amphipods in nature, or that the host’s value as a food source is overridden by other properties such as the degree of shelter it affords from larger consumers. This study provides the first demonstration that a member of the cosmopolitan amphipod family Aoridae is capable of consuming a diverse range of seaweeds.     

Atypical plant–herbivore association of algal food and a kleptoplastic sea slug (Elysia clarki) revealed by DNA barcoding and field surveys

The identity of food sources and feeding preferences of specialist herbivores have been commonly inferred from spatial associations between consumer and food items. However, such basic information for well-known marine herbivores, sacoglossans (sea slugs), and their algal diets remains disappointingly lacking, especially from field studies. The sacoglossan, Elysia clarki (Pierce et al. in Molluscan Res 26:23–38, 2006), is kleptoplastic and sequesters chloroplasts from algal food to photosynthesize, so DNA identification of sequestered chloroplasts was employed to verify the algal species fed upon by the slug across its geographic range. The molecular information on the algae consumed by E. clarki was combined with field surveys of slugs and algae in slug habitats in the Florida Keys in July and August of 2008 in order to evaluate whether the diet of this herbivore could be predicted based on its spatial association with algae in the field. A considerable mismatch between food availability and kleptoplast identity was recorded. E. clarki commonly occupied areas devoid of potential food and often contained symbiotic plastids from algal species different from those most frequently found in the surveyed habitats. In three of the four study sites, algal species present were poor predictors of slug diet. These findings suggest that the photosynthetic capability of E. clarki may release the slug from the constraint of requiring proximity to its food sources and may allow for the potential lack of spatial coupling between this herbivore and its algal food. This combination of field surveys and DNA barcoding provided critical and previously unavailable information on herbivore feeding in this marine system.     

Species-specific responses of herbivores to within-plant and environmentally mediated between-plant variability in plant chemistry

Allocation of resources to growth and defense against herbivores crucially affects plant competitiveness and survival, resulting in a specific distribution of assimilates and defense compounds within plant individuals. Additionally, plants rarely experience stable environmental conditions, and adaptations to abiotic and biotic stresses may involve shifts in resistance to herbivores. We studied the allocation of phytochemicals in Brassica oleracea (Brussels sprouts) due to leaf age, drought stress and herbivore damage and assessed effects on two lepidopteran herbivores differing in diet breadth: the generalist Spodoptera littoralis and the specialist Pieris brassicae. Glucosinolates as secondary defense compounds and total nitrogen and carbon were quantified and linked to plant palatability, i.e., herbivore feeding preference. Herbivore responses were highly species-specific and partially related to changes in phytochemicals. Spodoptera littoralis preferred middle-aged leaves with intermediate levels of glucosinolates and nitrogen over young, glucosinolate and nitrogen rich leaves, as well as over old leaves, poor in glucosinolates and nitrogen. In contrast, P. brassicae preferred young leaves. Both species preferred severely drought-stressed plants to the well-watered control, although analyzed glucosinolate concentrations did not differ. Both S. littoralis and P. brassicae feeding induced an increase of indole glucosinolate levels, which may explain a reduced consumption of damaged plants detected for S. littoralis but not for P. brassicae. By revealing distinct, sometimes contrasting responses of two insect herbivores to within-plant and stress-mediated intraspecific variation in phytochemistry of B. oleracea, this study emphasizes the need to consider specific herbivore responses to understand and predict the interactions between herbivores and variable plants.     

Do plant density, nutrient availability, and herbivore grazing interact to affect phlorotannin plasticity in the brown seaweed Ascophyllum nodosum

Plants have different strategies to cope with herbivory, including induction of chemical defences and compensatory growth. The most favourable strategy for an individual plant may depend on the density at which the plants are growing and on the availability of nutrients, but this has not been tested previously for marine plant–herbivore interactions. We investigated the separate and interactive effects of plant density, nutrient availability, and herbivore grazing on the phlorotannin (polyphenolic) production in the brown seaweed Ascophyllum nodosum. Seaweed plants grown at low or high densities were exposed either to nutrient enrichment, herbivorous littorinid gastropods (Littorina obtusata), or a combination of nutrients and herbivores in an outdoor mesocosm experiment for 2 weeks. Seaweeds grown at a low density tended to have higher tissue nitrogen content compared to plants grown at a high density when exposed to elevated nutrient levels, indicating that there was a density dependent competition for nitrogen. Herbivore grazing induced a higher phlorotannin content in plants grown under ambient, but not enriched, nutrient levels, indicting either that phlorotannin plasticity is more costly when nutrients are abundant or that plants responded to herbivory by compensatory growth. However, there were no significant interactive or main effects of plant density on the seaweed phlorotannin content. The results indicate that plants in both high and low densities induce chemical defence, and that eutrophication may have indirect effects on marine plant–herbivore interactions through alterations of plant chemical defence allocation.     

Asymmetric competition via induced resistance: specialist herbivores indirectly suppress generalist preference and populations

Species may compete indirectly by altering the traits of a shared resource. For example, herbivore-induced responses in plants may make plants more resistant or susceptible to additional herbivorous insect species. Herbivore-induced plant responses can significantly affect interspecific competition and herbivore population dynamics. These herbivore-herbivore indirect interactions have been overlooked in aquatic ecosystems where previous studies used the same herbivore species to induce changes and to assess the effects of these changes. We asked whether seaweed grazing by one of two herbivorous, congeneric snail species (Littorina obtusata or Littorina littorea) with different feeding strategies and preferences would affect subsequent feeding preferences of three herbivore species (both snails and the isopod Idotea baltica) and population densities of three herbivore species (both snails and a third periwinkle snail, Lacuna vincta). In addition, we measured phlorotannin concentrations to test the hypothesis that these metabolites function as induced defenses in the Phaeophyceae. Snail herbivory induced cue-specific responses in apical tissues of the seaweed Fucus vesiculosus that affected the three herbivore species similarly. When compared to ungrazed controls, direct grazing by Littorina obtusata reduced seaweed palatability by at least 52% for both snail species and the isopod species. In contrast, direct grazing by L. littorea did not decrease seaweed palatability for any herbivore, indicating herbivore-specific responses. Previous grazing by L. obtusata reduced populations of L. littorea on outplanted seaweeds by 46% but had no effect on L. obtusata populations. Phlorotannins, a potential class of inducible chemicals in brown algae, were not more concentrated in grazed seaweed tissues, suggesting that some other trait was responsible for the induced resistance. Our results indicate that marine herbivores may compete via inducible responses in shared seaweeds. These plant-mediated interactions were asymmetric with a specialist (L. obtusata) competitively superior to a generalist (L. littorea).     

Herbivory and detritivory among gammaridean amphipods from a Florida seagrass community

The gammaridean amphipods Cymadusa compta (Smith), Gammarus mucronatus Say, Melita nitida Smith and Grandidierella bonnieroides Stephensen from a seagrass community in the Indian River estuary of Florida (USA) fed variously upon large drift algae, small algae epiphytic on seagrasses and seagrass leaf debris and detritus. Consumption was measured in the laboratory using an index (CI) equivalent to mg ingested mg^-1 amphipod day^-1. Observations revealed that the amphipods fed by macrophagy, an attack upon large algae and seagrass debris, and by microphagy, small particle detritus feeding and scraping of plant surfaces for diatoms and other epiphytic algae. C. compta was a macrophagous feeder with a generalized diet of algae and seagrass debris, preferring epiphytic algae and drift algae at mean rates of 1.10 and 0.87 CI, respectively. Gammarus mucronatus fed upon epiphytes and seagrass debris equally at mean rates of 0.90 and 0.97 CI, respectively. The diet of M. nitida condisted primarily of epiphytes, consumed at an average rate of 1.05 CI. Grandidierella bonnieroides fed in a specialized microphagous manner, grooming plant surfaces for small particle detritus and diatoms at an approximate CI rate of 1.45. Assimilation of plants ingested, as reflected by carbon-14 uptake, varied similarly among the 4 amphipods. Epiphytic algae appeared to be most useful as food, providing means of 41 to 75% carbon-14 uptake as ingesta. Drift algae and seagrass debris were of less value, with means varying between 11 and 24 % of carbon-14 uptake by the amphipods. The data show a pattern of feeding which resembles resource partitioning of food both by size and kind. Other evidence, however, including population limitation by predators and an apparent overabundance of food, indicate that resource partitioning as seen may be an artifact, and one which has no co-evolutionary basis among the present species.Contribution No. 102 of Harbor Branch Foundation, Inc.     

The role of different types of detached macrophytes in the food and habitat choice of a surf-zone inhabiting amphipod

Allorchestes compressa is the dominant macroinvertebrate species in wrack accumulations on surf zones of south-western Australia. These amphipods were provided with a choice of macrophyte material representing brown and red algae and seagrass in a series of preference experiments in the laboratory and field. Feeding experiments showed that A. compressa exhibited a strong preference for particular types of macrophytes (P < 0.01). Amphipods primarily consumed brown algae, with 69–98% of the biomass of Ecklonia radiata and 64% of the biomass of Sargassum sp. lost over the experiments. This study has shown that the amphipod A. compressa exhibits a clear preference for brown algae over red algae and seagrass as food. In terms of habitat preference, tank experiments using a series of pair-wise comparisons showed that, in the absence of fish predators, A. compressa selected seagrass as its preferred habitat over the other types of wrack (P < 0.001). When satiated or starved, between 68 and 83 and 79 and 98% of amphipods were found in Amphibolis and Posidonia, respectively. In contrast, field-cage experiments revealed that A. compressa preferred either mixed wrack, brown algae or red algae over seagrass as a habitat (P < 0.01). The contrasts between results from the laboratory and field suggest that other factors such as the presence of predators, water flow and light could influence habitat choice in the surf zone. This study shows that allochthonous material transported to surf zones from other habitats therefore play different roles in driving secondary production in this shoreline habitat.     

Differential responses of vertebrate and invertebrate herbivores to traits of New Zealand subalpine shrubs

Plant traits are influenced by herbivore diet selection, but little is known about how traits are affected by different types of herbivores. We related eight traits of 27 subalpine shrub species in South Island, New Zealand, to damage of these shrubs by introduced red deer (Cervus elaphus) and native invertebrate herbivores using phylogenetically explicit modeling. Deer preferentially consumed species that grew quickly, were low in foliar tannins, or had high leaf area per unit mass. However, these traits did not trade off against each other; rather, they could be related to different multivariate defense strategies. Although the proportion of leaves damaged by leaf-chewing invertebrates also increased with stem growth, invertebrates did not damage the same fast growing species as those preferred by deer. Other traits may also be important in determining herbivore preferences, as suggested by the high proportion of variation in herbivory explained by phylogeny. Last, we found that the composition of invertebrate herbivore communities was more similar among closely related shrubs, and consequently, the range of invertebrate-plant associations may change if introduced deer shift plant composition toward slow-growing species. Overall, our results demonstrate the importance of herbivore type and coevolved interactions for the adaptive significance of plant traits.     

Species-specific defense strategies of vegetative versus reproductive blades of the Pacific kelps <Emphasis Type="Italic">Lessonia nigrescens</Emphasis> and <Emphasis Type="Italic">Macrocystis integrifolia</Emphasis>

Chemical defense is assumed to be costly and therefore algae should allocate defense investments in a way to reduce costs and optimize their overall fitness. Thus, lifetime expectation of particular tissues and their contribution to the fitness of the alga may affect defense allocation. Two brown algae common to the SE Pacific coasts, Lessonia nigrescens Bory and Macrocystis integrifolia Bory, feature important ontogenetic differences in the development of reproductive structures; in L. nigrescens blade tissues pass from a vegetative stage to a reproductive stage, while in M. integrifolia reproductive and vegetative functions are spatially separated on different blades. We hypothesized that vegetative blades of L. nigrescens with important future functions are more (or equally) defended than reproductive blades, whereas in M. integrifolia defense should be mainly allocated to reproductive blades (sporophylls), which are considered to make a higher contribution to fitness. Herein, within-plant variation in susceptibility of reproductive and vegetative tissues to herbivory and in allocation of phlorotannins (phenolics) and N-compounds was compared. The results show that phlorotannin and N-concentrations were higher in reproductive blade tissues for both investigated algae. However, preferences by amphipod grazers (Parhyalella penai) for either tissue type differed between the two algal species. Fresh reproductive tissue of L. nigrescens was more consumed than vegetative tissue, while the reverse was found in M. integrifolia, thus confirming the original hypothesis. This suggests that future fitness function might indeed be a useful predictor of anti-herbivore defense in large, perennial kelps. Results from feeding assays with artificial pellets that were made with air-dried material and extract-treated Ulva powder indicated that defenses in live algae are probably not based on chemicals that can be extracted or remain intact after air-drying and grinding up algal tissues. Instead, anti-herbivore defense against amphipod mesograzers seems to depend on structural traits of living algae.     

Restricted host use by the herbivorous amphipod <Emphasis Type="Italic">Peramphithoe tea</Emphasis> is motivated by food quality and abiotic refuge

There is a growing list of marine invertebrate herbivores known to restrict their host choices to a subset of available species, yet the relative importance of the evolutionary forces that select for specialized feeding habits remain unclear. One such specialist is the gammaridean amphipod Peramphithoe tea (F. Ampithoidae) that restricts its distribution to the brown laminarian seaweed Egregia menziesii in Oregon. A field survey indicated that among available seaweeds in the low intertidal zone of Boiler Bay, Oregon, Egregia housed greater than 90% of P. tea individuals. A set of laboratory-based habitat and feeding choice assays revealed that this specialized host distribution is likely the consequence of choices made by adult P. tea. The restricted host choice is apparently maintained by at least two evolutionary forces. First, a juvenile performance assay indicates that both Egregia and the co-occurring seaweed Alaria marginata, provide high food quality relative to other seaweeds available in the low-intertidal zone. Second, a field transplantation experiment revealed that Egregia protects adult amphipods from becoming dislodged with wave energy more readily than did Alaria. Thus, Egregia’s value as good quality food and refuge from abiotic stress together explain the restricted host use of P. tea. A comparison with previous studies suggests that use of Egregia is not consistent across the geographic range of P. tea, suggesting the possibility that the host preferences of local populations may respond evolutionarily to geographic shifts in seaweed communities.     

Preliminary evidence that the feeding rates of generalist marine herbivores are limited by detoxification rates

Herbivores tend to increase feeding rate and fitness when consuming a mixed diet relative to a single diet. According to the detoxification limitation hypothesis (DLH), feeding choices and rates when confronted with chemically rich plants are determined by herbivore physiology, and specifically by the metabolic pathways that herbivores use to manipulate secondary metabolites. We tested two predictions of the DLH using two generalist herbivores, the urchin Arbacia punctulata and amphipod Ampithoe longimana. These herbivores have geographic ranges which overlap with brown seaweeds that produce diterpenes (Dictyota menstrualis, D. ciliolata) and a green seaweed that produces sesquiterpenes and diterpenes (Caulerpa sertularioides). As predicted by the DLH, herbivore consumption rates in no-choice feeding assays were limited by extract intake rates. This suggests an upper limit in the herbivores’ abilities to physiologically manipulate seaweed metabolites. Contrary to a second prediction of the DLH, urchins consumed equal amounts of foods coated with limiting concentrations of two seaweed extracts offered singly, as a mixture, or as a pairwise choice. This result suggests that secondary metabolites of these seaweeds are manipulated by a linked set of detoxification pathways. Improving our understanding of the mechanisms that underlie diet mixing depends on greater attention to the physiology of herbivore resistance to secondary metabolites.     

Influence of dung volatiles on the process of resource selection by coprophagous beetles

Summary. Most dung beetles colonize the faeces of several vertebrate species without much discrimination, and are thus often considered as polyphagous. Recent studies have provided evidence for clear feeding preferences in scarab beetles colonizing dung of herbivore species, but little is known about these insects’ abilities to discriminate among odours from faeces of various herbivores. In this study, trophic preferences were examined using blocks of pitfall traps baited with dung from four different herbivore species, i.e., sheep, cattle, horse, and red deer, in a mountainous area of south-central France. 4941 coprophagous scarabs, belonging to 27 species, were captured. Beetles were more attracted to dung of sheep (2257 individuals) than that of cattle (1294 individuals), followed by deer dung (768 individuals) and horse dung (622 individuals). Eleven of the 27 beetle species collected had significant feeding preferences for one of the four dung types. For each insect species, trophic habits did not vary between the two different sites of trapping, an open pasture and a wooded habitat. In laboratory olfactometer bioassays, scarab beetles orientated preferentially towards the dung volatiles from the dung type they preferred in the field. Trypocopris pyrenaeus, Anoplotrupes stercorosus, and Aphodius rufipes were more attracted to volatile compounds from sheep dung, Onthophagus fracticornis significantly preferred horse dung volatiles, and Aphodius haemorrhoidalis responded positively to deer dung odours. The role of dung olfactory cues in the process of resource selection by dung beetles is discussed.     

Screening for induced herbivore resistance in Swedish intertidal seaweeds

Terrestrial plants have long been known to induce resistance towards herbivores in response to direct grazing, and strong evidence of inter-plant information transfer through volatile signals has been reported recently. Still, little is known about information exchange in aquatic plant–herbivore interactions. In this study, 12 Swedish seaweed species were exposed either to direct grazing by a generalist crustacean herbivore (Idotea granulosa), or to waterborne signals produced by actively feeding herbivores for 1 week. In order to test for the presence of induced chemical resistance in the different seaweed species, the dried and homogenized seaweed tissues were incorporated into an agar matrix, and herbivores were allowed to choose between the resulting control and induced artificial diets in two different two-choice feeding trials. The herbivores were actively feeding from all seaweed species in the induction experiments, although the amount of seaweed tissue consumed differed significantly between species. A chemically based induced herbivore resistance was found in response to direct grazing in four of the tested seaweed species (two red, one brown, and one green seaweed species). Furthermore, four seaweeds (one red, two brown, and one green seaweed species) induced resistance towards further grazing in response to waterborne chemical signals. Several seaweed species responded differently when exposed to different herbivore-related cues, indicating that both cues and responses can be highly specific. The results show that herbivore-induced resistance is present in 7 of 12 of the tested Swedish seaweed species, but that different signals (i.e., direct grazing and waterborne cues) elicit complex responses in the seaweeds.     

Feeding habits and food requirements of some amphipods in the Black Sea

Food composition and food requirements of four amphipod species of the Black Sea — Dexamine spinosa (Mont.), Amphithoe vaillanti Lucas, Gammarellus carinatus (Rathke), Gammarus locusta L. — were studied and an attempt made to assess the quantitative composition of the food, daily rhythms of feeding, and daily food rations. The composition of the food of the above-named amphipods is similar and consists primarily of seaweeds. Daily feeding intensity is characterized by a distinct increase during the dark period (D. spinosa, A. vaillanti); however, in G. locusta feeding intensity increases in the morning. The daily food rations of amphipods vary greatly (from 1.7 to 360% of the specimens' body weight), depending upon factors such as the kind of food, age, sex, the physiology of specimens, and the water temperature. The total quantity of food eaten by the populations of these amphipods in the coastal zone of the Black Sea amounts to 4000 g organic matter per square metre within 1 year.     

The effect of micrograzers on algal community structure in a coral reef microcosm

The effect of amphipod grazing on algal community structure was studied within a 75 l refuge tank connected to a 6500 l closed-system, coral reef microcosm. When amphipods (Ampithoe ramondi) were absent or present in low numbers, a high biomass of mostly filamentous algal species resulted, including Bryopsis hypnoides, Centroceras clavulatum, Ceramium flaccidum, Derbesia vaucheriaeformis, Enteromorpha prolifera, Giffordia rallsiae, and Polysiphonia havanensis. These microalgae disappeared when amphipod density increase beyond approximately 1 individual cm^-2 of tank surface. The macroalga Hypnea spinella germinated in the system in association with amphipod tube sites. H. spinella plants remained rare until filamentous species were eliminated by amphipod grazing. Feeding trials confirmed that H. spinella was protected from grazing by its size rather than a chemical defense strategy. The H. spinella community we observed is similar to the flora described on algal ridges where physical conditions exclude fish grazing. We suggest that amphipods and similar micrograzers are responsible for the algal community structure of these ridges. Caging experiments may be subject to similar effects from increased amphipod grazing on the algae. Introduction of fish that are amphipod predators into the refuge tank caused an increase in algal species diversity but total H. spinella growth rates fell from 25 g dry wt month^-1 to less than 8 g dry wt month^-1. We describe amphipod behavior in relation to changes in population density and food supply, and we stress the potential for increasing the productivity of commercial seaweeds through maintenance of appropriate amphipod species in mariculture facilities.     

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.