Turbinaria ornata as an herbivory refuge for associate algae

Habitat associations are an integral part of coral reef community structure. Commonly, one organism lives in such close association within or near another that a spatial refuge occurs, whereby one of the organisms provides protection to the other. This is often the result of defenses of the host deterring an associate organism’s consumers. In Moorea, French Polynesia, the range and abundance of the brown macroalga, Turbinaria ornata, have increased drastically since 1980 such that dense aggregations of this macroalga are a dominant component of the backreef habitat. Turbinaria ornata is both mechanically and chemically defended from herbivores. Other species of macroalgae grow within aggregations of Turbinaria and may benefit from these defenses. This study investigates whether aggregations of Turbinaria create a refuge from herbivory for associate macroalgae. When Turbinaria aggregations were removed experimentally, there was a significant increase in the number of associate algal species. Moreover, an herbivory assay using the palatable local alga Acanthophora spicifera identified herbivory as the mechanism for lower diversity on bommies lacking Turbinaria aggregations. The local increase in algal richness due to the refuge from herbivory afforded by Turbinaria may be an important contribution to macroalgal and community dynamics on reefs in Moorea, French Polynesia.     

Taste-rejection by predators and the evolution of unpalatability in prey

Aposematic species advertise their unpalatability to potential predators using conspicuous warning colouration. The initial evolution of aposematism is thought to occur by warningly coloured mutants emerging in an already unpalatable cryptic species. However, possessing defence chemicals is often costly, and it is difficult to understand what the selective benefits might be for a mutation causing its bearer to be defended in a population of otherwise palatable cryptic prey. One solution to this problem is that chemically defended individuals are tasted and rejected by predators, and are, therefore, more likely to survive predatory attacks than undefended individuals. Using naïve domestic chicks Gallus gallus domesticus as predators and cryptic green chick crumbs as prey, we asked whether the accuracy with which birds discriminated between palatable and unpalatable prey was affected by the palatability of the unpalatable prey (moderately or highly defended), or their frequency in the population (10 or 25%). Birds could discriminate between green prey on the basis of their defences, and showed better discrimination between palatable and unpalatable prey when defended crumbs were highly unpalatable, compared to when they were moderately unpalatable. Although there was no detectable effect of the frequency of unpalatable prey in the population on predator taste-rejection behaviour in our main analysis, frequency did appear to affect the strategies that birds used in their foraging decisions when prey were only moderately unpalatable. How birds used taste to reject prey also suggests that birds may be able to monitor and regulate their chemical intake according to the frequency and defence levels of the unpalatable prey. Taken together, these results show that avian predators can generate selection for unpalatability in cryptic prey by sampling and taste-rejecting prey, but that a relatively large chemical difference between palatable and unpalatable prey may be necessary before unpalatable prey can enjoy a selective advantage. The exact nature of this evolutionary dynamic will depend on other environmental factors, such as defence costs and prey availability, but it provides a mechanism by which defences can evolve in a cryptic population.     

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.     

Population structure and feeding deterrence in three shallow-water antarctic soft corals

Alcyonium paessleri and Clavularia frankliniana are numerically abundant soft corals in the nearshore (12 to 33 m depth) benthic communities of eastern McMurdo Sound. They are much less abundant in western McMurdo Sound where a third species, Gersemia antarctica, co-occurs in low numbers. The body tissues of these three species are comprised mainly of organic material (53 to 70% dry wt), which is primarily dervied from NaOH-soluble protein and refractory material. The energetic contents of the whole-body tissues of A. paessleri, C. frankliniana and G. antarctica are 15.9, 17.3, and 14.5 kJ g^-1 dry wt, respectively. The mean biomass per individual is 1.81, 0.008, and 45 g dry wt for each respective species. Based on population densities of 7.3, 1337.3, and 0.04 soft corals m^-2 for A. paessleri, C. frankliniana and G. antarctica, respectively, the population energetic densities are estimated to be 210.1, 185.1, and 26.1 kJ m^-2. Despite the relatively rich energetic content of the tissue and apparent vulnerability to predators, very little predation occurs on these soft corals. Two potential predators, the antarctic sea stars Perknaster fuscus and Odontaster validus, exhibited significant chemotactic defensive tube-foot retractions to hexane, chloroform, methanol, and aqueous methanol extracts of each soft coral. In addition, wholebody tissue of each soft coral was rejected by the demersal fish Pseudotrematomus bernacchii and the cryopelagic fish Pagothenia borchgrevinki. In contrast, whole soft-coral tissues sequentially extracted in four increasingly polar solvents were readily ingested by these antarctic fishes, indicating that sclerites do not play a significant role in deterring predators. Our results indicate that these antarctic soft corals contain bioactive compounds which deter common predatory seastars and fishes.     

Seaweed sex pheromones and their degradation products frequently suppress amphipod feeding but rarely suppress sea urchin feeding

Summary. A diverse group of brown seaweeds produce bouquets of C₁₁ metabolites, some of which act as pheromones that cue gamete release or attract sperm to eggs following release. We demonstrate that these C₁₁ metabolites and their degradation products also frequently and strongly deter feeding by the herbivorous amphipod Ampithoe longimana, but rarely by the herbivorous sea urchin Arbacia punctulata. Across the range of concentrations tested, seven of twelve C₁₁ metabolites or mixtures that we tested deterred feeding by the amphipod, but only two of eleven deterred the sea urchin. For those compounds where we could rigorously contrast the magnitude of deterrence against the amphipod with the magnitude of deterrence against the urchin, the amphipod was deterred significantly more than the urchin by five of six metabolites. Thus, C₁₁ compounds were more frequently and more strongly deterrent to the amphipod than to the sea urchin. These findings for C₁₁ metabolites conflict with previous investigations, where other classes of seaweed chemical defenses have been shown to deter feeding by large mobile herbivores like urchins and fishes but to be relatively ineffective against mesograzers, especially the species of amphipod that we used here. Our results suggest that C₁₁ metabolites are unusual among the known seaweed chemical defenses in that they are especially effective against mesograzers, which often consume seaweed spores, zygotes, and juveniles. The high concentrations of C₁₁ metabolites in brown algal eggs could allow these defenses to be especially important in defending gametes, zygotes, or young sporelings from herbivorous mesograzers. Received 26 February 1998; accepted 9 April 1998.     

Patterns of shell repair in articulate brachiopods indicate size constitutes a refuge from predation

The cost of overcoming prey defenses relative to the value of internal tissues is a key criterion in predator/prey interactions. Optimal foraging theory predicts: (1) specific sizes of prey will result in the best returns to predators, and (2) there will often be a size at which the cost/benefit balance is low enough to effectively exclude predation. Data presented here on styles of repaired shell damage and size at which injury had been sustained was collected from samples of terebratulide brachiopods from the Antarctic Peninisula (Liothyrella uva), Falkland Islands (Magellania venosa and Terebratella dorsata) and Chile (M. venosa). The predominant form of damage on shells was indicative of predators attacking the valve margins. The modal size for repaired damage was more than 10 mm smaller than the modal size for the overall size distribution in each species and there were no repaired attacks in the largest size classes of any species. These data suggest that size forms a refuge from predation, as would be predicted by optimal foraging theory. The optimal sizes that predators appeared to attack vary between species, as do the sizes that provided a refuge from predation. High levels of multiple repairs (19% of the M. venosa population from the Falkland Islands sampled had 2 or more repairs) suggest that the mortality following attack is low, suggesting that many predators abandon their attacks.     

Behavioral response to predators reverses the outcome of competition between prey species

Summary In a laboratory experiment it was shown that piscivorous predators reversed the outcome of competitive interactions between two fish prey species, juveniles of roach (Rutilus rutilus) and perch (Perca fluviatilis), by behaviorally affecting their use of two available habitats, an open water habitat and a structurally complex refuge. The shift in the competitive relationship was the result of predators forcing the juvenile fishes into a prey refuge with high structural complexity. While roach was competitively superior in the unstructured habitat, perch was superior in the structurally complex prey refuge. The reversal in competitive relationship was demonstrated both with respect to foraging rate and growth rate and resulted from the high structural complexity in the prey refuge interfering with the roach's swimming performance. Because survival and growth patterns through the juvenile stages have profound effects on the population/community dynamics of size-structured populations such as those of fish, behaviorally induced changes in competitive ability should have significant implications also at the population and community levels.     

Selective consumption and facilitation by mesograzers in adult and colonizing macroalgal assemblages

Grazing pressure on macroalgae in littoral communities may vary with algal species, the age of an algal individual and grazer identity. Previous studies on alga–grazer interactions have shown that grazer preference for an algal species may release another one from interspecific competition. We measured the impacts of four common grazer taxa and the natural grazer guild on macroalgal communities at both their colonization and adult stages, and compared the impacts to grazer exclosures. The grazer effects were stronger on colonizing than on adult macroalgae; grazers did not reduce the total density of adult algae. Grazers both feed on propagules and indirectly facilitate other algae, depending on the grazer or algal species. Hydrobia species increased the settlement of spores of the red alga Ceramium tenuicorne. Similarly, the gastropod Theodoxus fluviatilis tended to facilitate one crustose algal species, but decreased the propagule density of annual filamentous algae, suggesting a preference for one species to the advantage of another. Effects of crustacean mesograzers on the studied macroalgae were weak. These results indicate that northern Baltic macroalgae are limited to grazing mainly during their colonization stage.     

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.     

Distance to neighbours influences the trade-off between hiding after disturbance and defending food patches in convict cichlids (<Emphasis Type="Italic">Archocentrus nigrofasciatus</Emphasis>)

Territories are often aggregated. Because of this, distance to neighbours should influence how territory-holders balance safety from predators with the use and defence of resources. I examined the influence of distance to a neighbour on refuge use by pairs of convict cichlids (Archocentrus nigrofasciatus) faced with a conflict between hiding and defending food patches. Neighbours could reduce the rate of intrusions by strangers as a by-product of their own resource defence. This should allow fish with near neighbours to spend more time in the refuge. Neighbours could also steal from patches that are left undefended. This should lead to a reduction in use of the refuge. When one fish was confined to its refuge (so that its patch was undefended), theft by the other increased as inter-patch distance decreased. Distance between patches did not influence the rate of intrusion by non-territorial fish. When both fish defended patches, body mass influenced the effect of inter-patch distance on refuge use. Large fish rarely used the refuge, but small territory-holders spent more time in the refuge when patches were close together, as predicted. However, when one fish was dominant at both patches, distance between patches did not influence refuge use. These results suggest that, despite increased opportunity for theft, there is no realised foraging or defensive benefit to settling near neighbours that are of similar competitive ability.Communicated by J. Krause     

The role of the amphipod <Emphasis Type="Italic">Gammarus locusta</Emphasis> as a grazer on macroalgae in Swedish seagrass meadows

Mesograzers are thought to play a critical role in seagrass beds by preventing overgrowth of ephemeral algae. On the Swedish west coast, eelgrass Zostera marina has decreased in recent decades as a result of eutrophication and increased growth of macroalgal mats (mainly filamentous Ulva spp. and Ectocarpales), with no indication of grazer control of the algae. The aim of this study was to investigate the ability of the amphipod Gammarus locusta to control algal blooms during nutrient-enriched and ambient conditions, using a combination of laboratory, field and model studies. Laboratory experiments demonstrated that juvenile and adult G. locusta could consume both Ulva spp. and Ectocarpales, but that consumption of Ulva spp. was significantly higher. Cannibalism was common in individual treatments involving multiple size-classes of G. locusta, but only large, male gammarids consumed smaller juveniles in the presence of Ulva spp. as an alternative food source. However, no negative effects of cannibalism were found on total grazing impact. A model using size-specific grazing rates and growth rates of G. locusta and of Ulva spp. suggests that approximately 62 young juvenile, or 27 adult G. locusta are needed per gram DW of Ulva spp. to control the algal growth during ambient nutrient conditions, and approximately 2.6 times as many gammarids during enhanced nutrient conditions. On the Swedish west coast, densities and mean sizes of G. locusta in eelgrass beds are below these critical values, suggesting that the gammarids will not be able to control the growth of the filamentous macroalgae. However, in the field cage experiment, immigration of juveniles and reproduction of encaged adult G. locusta resulted in unexpectedly high densities of G. locusta (>4,000 individual m⁻²), and very low biomass of Ulva spp. in both ambient and nutrient-enriched treatments. Although the high numbers of juveniles in all cages precluded any significant treatment effects, this suggests that in the absent of predators, the population of G. locusta can grow significantly and control the biomass of Ulva spp. Furthermore, low grazing of Ectocarpales in the laboratory and high biomass of these filamentous brown algae in the field indicate a preference for the more palatable green algae Ulva spp. This study indicates that the high grazing capacity of G. locusta, in combination with high reproduction and growth rates, would allow the amphipod to play a key role in Z. marina ecosystems by controlling destructive blooms of filamentous green algae. However, high predation pressure appears to prevent large populations of G. locusta in eelgrass beds on the Swedish west coast today.     

Variability in grazer-mediated defensive responses of green and red macroalgae on the south coast of South Africa

Variabilities in the responses of several South African red and green macroalgae to direct grazing and the responses of one green alga to cues from grazers were tested. We used two feeding experiments: (1) testing the induced responses of three red and one green algae to direct grazing by mesograzers and (2) a multi-treatment experiment, in which the direct and indirect effects of one macrograzer species on the green alga Codium platylobium were assessed. Consumption rates were assessed in feeding assays with intact algal pieces and with agar pellets containing non-polar extracts of the test algae. Defensive responses were induced for intact pieces of Galaxaura diessingiana, but were not induced in pellets, suggesting either morphological defence or chemical defence using polar compounds other than polyphenols. In contrast, exposure to grazing stimulated consumption of Gracilaria capensis and Hypnea spicifera by another grazing species. In the multi-treatment experiment, waterborne cues from both grazing and non-grazing snails induced defensive algal traits in C. platylobium. We suggest that inducible defences among macroalgae are not restricted to brown algae, but that both the responses of algae to grazers and of grazers to the defences of macroalgae are intrinsically variable and complex.     

Going deep: common murres dive into frigid water for aggregated,persistent and slow-moving capelin

Owing to the necessity of delivering food to offspring at colonies, breeding seabirds are highly constrained in their foraging options. To minimize constraints imposed by central-place foraging and to optimize foraging behavior, many species exhibit flexible foraging tactics. Here we document the behavioral flexibility of pursuit-diving common murres Uria aalge when foraging on female capelin Mallotus villosus in the northwest Atlantic. Quite unexpectedly, being visual foragers, we found that common murres dived throughout the day and night. Twenty-one percent of recorded dives (n = 272 of 1,308 dives) were deep (≥50 m; maximum depth = 152 m, maximum duration = 212 s), bringing murres into sub-0°C water in the Cold Intermediate Layer (CIL; 40–180 m) of the Labrador Current. Deep dives occurred almost exclusively during the day when murres would have encountered spatially predictable aggregations of capelin between 100 and 150 m in the water column. Temperatures within the CIL shaped trophic interactions and involved trade-offs for both predators and prey. Sub-0°C temperatures limit a fish’s ability to escape from endothermic predators by reducing burst/escape speeds and also lengthening the time needed to recover from burst-type activity. Thus, while deep diving may be energetically costly, it likely increases certainty of prey capture. Decreased murre foraging efficiency at night (indicated by an increase in the number of dives per bout) reflects both lower light conditions and changing prey behavior, as capelin migrate to warmer surface waters at night where their potential to escape from avian predators could increase.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Consequences of behavioral vs. numerical dominance on foraging activity of desert seed-eating ants

Dominance relationships among species play a major role in the structure of animal communities. Yet, dominant species with different trade-offs in resource exploitation and monopolization could affect community structure in variable ways. In ants, dominant species could be classified into either behavioral dominants that exhibit territorial aggression or numerical dominants that exhibit high biomass or frequency of occurrence. While each class of dominance has generally been found to negatively affect the foraging activity of species in ant communities, the concurrent effect of both classes of species has never been tested. Here, we examined the effects of two behaviorally dominant species, Crematogaster inermis and Monomorium salomonis, and a numerically dominant species, Messor arenarius, on the foraging behavior of seed-eating species in a desert ant assemblage. In a 1-year study, the foraging activity of the ant species was assessed using seed baits, which were sampled during night and day. While the numerically dominant species exhibited high foraging efficiency and negatively affected the ability of other seed-eating species to obtain seeds, significantly more seeds remained at baits that were occupied the previous night by each of the two behaviorally dominant species, possibly due to aggressive exclusion of M. arenarius foragers from the baits. This exclusion also facilitated greater foraging activity of the seed-eating species. Our results demonstrate how these two types of dominance could differently affect the foraging activity of ant species in the community.     

Defensive secretion of a flightless grasshopper: Failure to prevent lizard attack

Summary We tested the role of a grasshopper defensive secretion in deterring lizard predation. Adults, but not young larvae, of the chemically defended lubber grasshopperRomalea guttata (=microptera) froth a volatile secretion when attacked by predators. The lizardAnolis carolinensis failed to strike juvenile lubbers (which lack secretion) in laboratory trials. Survivorship of palatable crickets loaded with secretion offered toA. carolinensis was not significantly different from survivorship of control crickets. In experiments designed to investigate if lizards learn an aversion to the secretion, striking times forSceloporus undulatus fed wax worms coated with secretion were not significantly different over three days of trials. Three primary conclusions are drawn from these data. First, the secretion may not be necessary for lubber protection from lizards. Second, lubber secretion does not appear to deter lizards from attacking or eating prey items. Third, lizards do not appear to develop an aversion to the secretion.     

Turning the game around: toxicity in a nudibranch-sponge predator–prey association

Escalation theory proposes enemy-related selection as the most relevant factor of natural selection among individual organisms. When hazardous to predators, prey might be considered enemies that influence predator evolution. Opisthobranch molluscs that prey on chemically defended prey are an interesting study case on this subject. Predation on chemically defended species paved the way for opisthobranchs to enter in an arms race, developing means to detoxify and/or excrete harmful compounds, which led to the sequestration of those compounds and their self-defensive use, an escalation of defenses. Here we aim to understand whether the opisthobranch predator is better protected than its chemically defended prey, using as predator–prey model, a nudibranch (Hypselodoris cantabrica) and the sponge it preys upon (Dysidea fragilis), and from which it obtains deterrent chemical compounds. Specimens of both species were collected on the Portuguese coast, and their crude extracts were analyzed and used in palatability tests. Nudibranchs revealed a higher natural concentration of crude extract, probably due to a progressive accumulation of the compounds. Both predator and prey extracts revealed similar mixtures of deterrent metabolites (furanosesquiterpenes). Palatability tests revealed a more effective deterrence in the nudibranch extracts because significant rejection rates were observed at lower concentrations than those necessary for the sponge extracts to have the same effect. We concluded that the predator is chemically better protected than its prey, which suggests that its acquisition of chemical defenses reveals a defensive escalation.     

Processing of allochthonous macrophyte subsidies by sandy beach consumers: estimates of feeding rates and impacts on food resources

Allochthonous subsidies of organic material can profoundly influence population and community structure; however, the role of consumers in the processing of these inputs is less understood but may be closely linked to community and ecosystem function. Inputs of drift macrophytes subsidize sandy beach communities and food webs in many regions. We estimated feeding rates of dominant sandy beach consumers, the talitrid amphipods (Megalorchestia corniculata, in southern California, USA, and Talitrus saltator, in southern Galicia, Spain), and their impacts on drift macrophyte subsidies in field and laboratory experiments. Feeding rate varied with macrophyte type and, for T. saltator, air temperature. Size-specific feeding rates of talitrid amphipods were greatest on brown macroalgae (Macrocystis, Egregia, Saccorhiza and Fucus). Rates for large individuals of both species ranged from ∼40 mg wet wt individual⁻¹12 h⁻¹ on brown macroalgae to negligible feeding by M. corniculata on a vascular plant (surfgrass). Amphipod growth rates were also greatest on Macrocystis and lowest on surfgrass, Phyllospadix. For a Californian beach with substantial inputs of macrophyte wrack (>70 kg wet wt m⁻¹ month⁻¹ in summer), we estimated that the population of talitrid amphipods could process an average of 55% of the palatable Macrocystis input. Our results indicate that talitrid amphipod populations can have a significant impact on drift macrophyte processing and fate and that the quantity and composition of drift macrophytes could, in turn, limit populations of beach consumers.     

Behavioral responses to a novel predator and competitor of invasive mosquitofish and their non-invasive relatives (Gambusia sp.)

Attributes of the recipient community may affect the invasion success of arriving non-indigenous organisms. In particular, biotic interactions may enhance the resistance of communities to invasion. Invading organisms typically encounter a novel suite of competitors and predators, and thus their invasiveness may be affected by how they cope with these interactions. Behavioral plasticity may help invaders to respond appropriately to novelty. We examined the behavioral responses of highly invasive mosquitofish to representative novel competitors and predators they might encounter as they spread through North America. We compared the behavior of invasive Gambusia holbrooki and G. affinis to that of two close relatives of lower invasive potential (G. geiseri and G. hispaniolae) in order to elucidate whether responses to novelty related to invasiveness. In short-term assays, female Gambusia were paired with a novel competitor, Pimephales promelas, and a novel predator, Micropterus dolomieu. Behavioral responses were measured in terms of foraging success and efficiency, activity, refuge use, predator inspections, and interspecific aggression. Contrary to a priori predictions, invasive and non-invasive responses to novel interactions did not differ consistently. In response to novel competition, both invasive species increased foraging efficiency, but so did G. geiseri. In response to novel predation, only G. holbrooki decreased consumption and activity and increased refuge use. No antipredator response was observed in G. affinis. We found consistent differences, however, between invasives and non-invasives in foraging behavior. Both in the presence and absence of the competitor and the predator, invasives foraged more efficiently and consumed more prey than non-invasives.Communicated by P. Bednekoff     

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.