Structural resistance of algae to herbivores

Experiments were conducted to determine the forces required for two limpet species to remove tissue from three different types of algae. This was done by using the radulae of the limpets, mimicking the way they use this feeding apparatus. For both Acmaea mitra Rathke (a specialist herbivore) and Notoacmea scutum (Rathke) (a generalist herbivore), Iridaea cordata (Turn.) Bory (a thin, bladed red alga) required the most force; Hedophyllum sessile (C.Ag.) Setch. (a thick, bladed brown alga) the next; and Pseudolithophyllum whidbeyensi (a crustose, calcified alga) required the least amount of force for tissue removal. These results are the inverse of the predictions made by current functional form and functional group models which use the thallus form as a predictor of herbivore resistance. Thallus form alone is a poor predictor of structural resistance to molluscan feeding; the material properties of the alga, the feeding apparatus of the herbivore, and the mode of feeding all must be considered.     

Substratum relationships of some tropical Pacific intertidal gastropods

Among tropical Indo-West Pacific herbivorous intertidal gastropods, high intertidal neritids, and limpets and certain lower intertidal limpets are restricted to a narrow range of substratum types. Littorinids, planaxids, low intertidal neritids, most lower intertidal limpets, and the trochid Monodonta labio tolerate a wide variety of substrata. This pattern of substratum specialization is related to the overall adaptive strategies of the various groups: those with a large area of soft tissue in contact with the substratum (neritids and limpets) often specialize to a particular substratum, especially at high shore levels, while those in which the area of contact between animal and substratum is small do not. High intertidal neritids and limpets able to live on a wide range of substrata tend to be species of limited geographic distribution. It is concluded that, even within a tropically similar group of animals (raspers in this case) relevant dimensions of niche separation depend upon the structure of the environment as well as upon the structure of the animals themselves.     

Distribution and morphological variation of low-shore algal turfs

The distribution of three functional groups of algae (filamentous, corticated terete and calcareous articulated) was investigated in low-shore, turf-forming assemblages from rocky shores in the western Mediterranean Sea (Italy). Algae were sampled along shores from three different stations. Shores were either exposed to wave action or sheltered and were characterised by various inclinations of the rocky substratum (horizontal, sloping and vertical). The hypotheses tested were: that the relative abundance of the three functional groups of algae (1) varies between sheltered and exposed shores, (2) is influenced by substratum inclination and (3) that these distributions are consistent across stations. There was no clear relationship between the morphology of turf-forming algae and shore characteristics. Covers of filamentous, corticated terete and calcareous articulated algae differed among shores and stations, but patterns were not consistent across shores with similar exposure to waves and substratum inclination. Turfs often comprised algae with different morphologies, in a combination of anchor and epiphytic species. It is suggested that functional groups of algae, as currently identified, are not effective in describing changes in distribution of algae between sheltered and exposed shores, when turf morphologies are considered. Future research on relationships between morphology and ecology of algae should consider the potential importance of positive interactions among associated species, which could override responses of individual functional groups.     

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.     

Herbivore vs. nutrient control of marine primary producers: context-dependent effects

Pervasive overharvesting of consumers and anthropogenic nutrient loading are changing the strengths of top-down and bottom-up forces in ecosystems worldwide. Thus, identifying the relative and synergistic roles of these forces and how they differ across habitats, ecosystems, or primary-producer types is increasingly important for understanding how communities are structured. We used factorial meta-analysis of 54 field experiments that orthogonally manipulated herbivore pressure and nutrient loading to quantify consumer and nutrient effects on primary producers in benthic marine habitats. Across all experiments and producer types, herbivory and nutrient enrichment both significantly affected primary-producer abundance. They also interacted to create greater nutrient enrichment effects in the absence of herbivores, suggesting that loss of herbivores produces more dramatic effects of nutrient loading. Herbivores consistently had stronger effects than did nutrient enrichment for both tropical macroalgae and seagrasses. The strong effects of herbivory but limited effects of nutrient enrichment on tropical macroalgae suggest that suppression of herbivore populations has played a larger role than eutrophication in driving the phase shift from coral- to macroalgal-dominated reefs in many areas, especially the Caribbean. For temperate macroalgae and benthic microalgae, the effects of top-down and bottom-up forces varied as a function of the inherent productivity of the ecosystem. For these algal groups, nutrient enrichment appeared to have stronger effects in high- vs. low-productivity systems, while herbivores exerted a stronger top-down effect in low-productivity systems. Effects of herbivores vs. nutrients also varied among algal functional groups (crustose algae, upright macroalgae, and filamentous algae), within a functional group between temperate and tropical systems, and according to the metric used to measure producer abundance. These analyses suggest that human alteration of food webs and nutrient availability have significant effects on primary producers but that the effects vary among latitudes and primary producers, and with the inherent productivity of ecosystems.     

Morphological patterns in high-intertidal gastropods: Adaptive strategies and their limitations

An analysis of shell form in several families of high-intertidal herbivorous gastropods has revealed the existence of morphological gradients. Among tropical littorinids, relative spire height and degree of development of external shell sculpture generally increase interspecifically, and often intraspecifically, from low to high shore levels. This gradient is also evident from temperate to tropical latitudes, and may be modified or overridden by exposure to wave action and other factors. Among acmaeid, patellid, and siphonariid limpets, there is a well-marked interspecific increase in relative shell height with increasing shore level, but latitudinal trends in morphology cannot at present be distinguished. Evidence from both tropical and temperate coasts indicates that limpets found in shaded places tend to have smoother shells than those often exposed to direct sunlight. In the Neritidae, relative shell globosity tends to increase interspecifically from low to high shore levels; most high-shore species have strong shell sculpture, with the base of the shell only slightly excavated. A decrease in the number of neritid species towards higher latitudes is accompanied by selective deletion of globose, strongly sculptured types living at high shore levels. Neritids and some limpets can employ evaporative cooling as a mechanism for temperature regulation. The morphological gradients in these groups reflect a relative increase in the volume of the extravisceral cavity (water reservoir) and a decrease in the area in contact with the substratum, as well as a relative decrease in the rate of water loss, in an upshore direction. Neritids are better adapted than limpets to high temperature and desiccation stresses because of their ability to regulate rate of evaporation and contact between soft parts and substratum. These differences are reflected in the complementary patterns of latitudinal diversity in the two groups. In Littorinidae, which attach to the substratum by a mucus film, the morphological gradients reflect a minimization of water loss and substratum contaet, and a maximization of reflective relative to absorptive surface area.This paper is an expanded version of part of a Ph. D. dissertation presented to Yale University.     

Functional identity and functional structure change through succession in a rocky intertidal marine herbivore assemblage

Despite the great interest in characterizing the functional structure and resilience of functional groups in natural communities, few studies have examined in which way the roles and relationships of coexisting species change during community succession, a fundamental and natural process that follows the release of new resources in terrestrial and aquatic ecosystems. Variation in algal traits that characterize different phases and stages of community succession on rocky shores are likely to influence the magnitude, direction of effects, and the level of redundancy and complementarity in the diverse assemblage of herbivores. Two separate field experiments were conducted to quantify per capita and population effects and the functional relationship (i.e., redundancy or complementarity) of four herbivore species found in central Chile during early and late algal succession. The first experiment examined grazer effects on the colonization and establishment of early-succession algal species. The second experiment examined effects on the late-successional, dominant corticated alga Mazzaella laminarioides. Complementary laboratory experiments with all species and under natural environmental conditions allowed us to further characterize the collective effects of these species. We found that, during early community succession, all herbivore species had similar effects on the ephemeral algae, ulvoids, but only during the phase of colonization. Once these algae were established, only a subset of the species was able to control their abundance. During late succession, only the keyhole limpet Fissurella crassa could control corticated Mazzaella. The functional relationships among these species changed dramatically from redundant effects on ephemeral algae during early colonization, to a more complementary role on established early-successional algae, to a dominant (i.e., keystone) effect on late succession. This study highlights that functional relationship within consumer assemblages can vary at different phases and times of community succession. Differentiation in herbivore roles emphasizes the need to evaluate consumer's impacts through different times of community succession, and through experimental manipulations to make even broad predictions about the resilience or vulnerability of diverse intertidal assemblages to human disturbances.     

The distribution of herbivorous fishes on the Great Barrier Reef

The composition and functionality of ecologically important herbivorous fish assemblages were examined throughout much of Australia’s Great Barrier Reef (GBR). Diversity and abundance of surgeonfishes (Acanthuridae), parrotfishes (Labridae) and rabbitfishes (Siganidae) were strongly associated with position on the continental shelf, whilst effects of latitude were weaker and inconsistent. Species distributions varied considerably amongst taxonomic groups; parrotfishes were mostly widespread whilst distributions of surgeonfishes were often restricted. Most inshore environments supported depauperate herbivore assemblages dominated by different taxa and functional groups compared with assemblages in offshore environments. There were also strong cross-shelf transitions in the main taxa performing each functional role. Overall, this study show that the functional contributions of herbivorous fish assemblages to important ecosystem processes and the contributing taxa vary considerably amongst different GBR environments. Additionally, the two most numerically dominant herbivores actively select detritus, not algae, supporting increasing evidence for the importance of detritus in coral reef ecology.     

Chemoautotrophic symbiosis in the tropical clamSolemya occidentalis (Bivalvia: Protobranchia): ultrastructural and phylogenetic analysis

Patterns of algal seasonality, and their effect on the diet and feeding preferences of the herbivorous crab Grapsus albolineatus were investigated over an 18-mo period from March 1993 to August 1994 on an exposed tropical rocky shore (Hok Tsui Peninsula at Cape d' Aguilar, Hong Kong). Algal cover was greatest in the winter months, and lowest in the summer. Foliose algae such as Ulva fasciata, Porphyra suborbiculata, and Dermonema frappieri were dominant in the winter, but died off in the summer. During the hot summer months, perennial encrusting algae e.g., Ralfsia expansa, Hildenbrandia rubra, H. occidentalis, coralline crusts and the encrusting cyanobacteria Kyrtuthrix maculans, were the dominant algal species. Seasonal variation in algal abundance influenced the dietary selectivity of the herbivorous crab G. albolineatus. In the winter, the crab fed selectively on filamentous algae (e.g. Hincksia spp., Cladophora spp., Enteromorpha spp., and the cyanobacteria Lyngbya sp.). Foliose algae (e.g. U. fasciata, P. suborbiculata, Pterocladia tenuis) formed a small part of the diet, despite being the dominant species on the shore. Foliose and filamentous algae were virtually absent from the shore in the summer, and the crabs switched to feeding solely on encrusting algae. Electivity indices revealed preferences for green and brown turf species, and avoidance of foliose algae. Faecal analysis revealed that a greater proportion of the food is digested in the winter, suggesting that G. albolineatus is able to digest filamentous algae more efficiently than encrusting algae. Feeding preferences of G. albolineatus appear to be influenced by a number of factors, including the availability, digestibility and morphology of algae. The foraging behaviour and cheliped morphology of the crab also affect food choice. The monsoonal nature of Hong Kong's climate controls the diversity and abundance of intertidal algae and, therefore, indirectly influences the diet and subsequent growth and reproductive success of the herbivorous crab G. albolineatus.     

A re-evaluation of the diel feeding hypothesis for marine herbivorous fishes

We re-evaluated the "diel feeding hypothesis" by measuring diel variation in starch, protein, and floridoside in three algal "types" collected from a fringing coral reef at Lizard Island, Great Barrier Reef, Australia. Samples of two species of rhodophyte algae, Gracilaria arcuata and Acanthophora spicifera, and the turf assemblage from the territories of the herbivorous pomacentrid Stegastes nigricans were collected at four time periods through the day: 0630-0730, 1000-1100, 1330-1430, and 1630-1730 hours. We also measured the ability of several species of marine fish (the herbivores Acanthurus nigricans, A. lineatus, A. olivaceus, and Parma alboscapularis and the detritivore Ctenochaetus striatus) to hydrolyse floridoside by estimating !-galactosidase activity in tissue from the anterior intestine. We detected no diel pattern in protein content of the algae but found a significant steady increase in starch content throughout the day. Floridoside content increased in the morning and decreased in the afternoon, a pattern that may be driven by midday photoinhibition of the algae. All the fishes tested could utilise floridoside. Our results support the diel feeding hypothesis. Although floridoside content decreased in the afternoon, our results suggest floridoside was used during the day by the algae to synthesise starch. Thus the algae increased in nutritional value until photoinhibition occurred at midday then subsequently maintained their nutritional value during the afternoon. This pattern of algal nutrients increasing to a midday peak and remaining relatively constant throughout the afternoon correlates well with the diel feeding pattern in many species of marine herbivorous fish.     

Latitudinal variation in top-down and bottom-up control of a salt marsh food web

The shrub Iva frutescens, which occupies the terrestrial border of U.S. Atlantic Coast salt marshes, supports a food web that varies strongly across latitude. We tested whether latitudinal variation in plant quality (higher at high latitudes), consumption by omnivores (a crab, present only at low latitudes), consumption by mesopredators (ladybugs, present at all latitudes), or the life history stage of an herbivorous beetle could explain continental-scale field patterns of herbivore density. In a mesocosm experiment, crabs exerted strong top-down control on herbivorous beetles, ladybugs exerted strong top-down control on aphids, and both predators benefited plants through trophic cascades. Latitude of plant origin had no effect on consumers. Herbivorous beetle density was greater if mesocosms were stocked with beetle adults rather than larvae, and aphid densities were reduced in the "adult beetle" treatment. Treatment combinations representing high and low latitudes produced patterns of herbivore density similar to those in the field. We conclude that latitudinal variation in plant quality is less important than latitudinal variation in top consumers and competition in mediating food web structure. Climate may also play a strong role in structuring high-latitude salt marshes by limiting the number of herbivore generations per growing season and causing high overwintering mortality.     

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).     

The effect of ocean acidification on early algal colonization stages at natural CO2 vents

Marine algae exhibit different responses to ocean acidification, suggesting that a decrease in pH does not always favour marine photosynthetic organisms. In order to understand the effect of acidification on algal community development, early colonization stages were investigated using carbon dioxide vents around the Castello Aragonese (Ischia, Italy) as a natural laboratory. Settlement tiles were placed in zones with different pH (normal, medium and low), and species composition and coverage measured after 2, 3 and 4 months of deployment. The number of species decreased by 4 and 18 % at medium and low pH zones, respectively (P < 0.05). The structure of the algal assemblage differed between pH zones during the 4 months of the experiment, due to the addition and/or replacement of new species. This leads to a change in the succession of morphological forms as soft crustose algae replaced calcareous species, and turf species were dominant in cover; more complex thalli started to occur only at medium pH. These results support previous findings that ocean acidification will induce changes in benthic algal communities.     

The trophic status of herbivorous fishes on coral reefs

Estimates of feeding rates, alimentary tract structure and temporal patterns of food processing obtained from twelve species of nominally herbivorous fishes on the northern Great Barrier Reef were compared. These included members of the families Acanthuridae, Scaridae and Kyphosidae. Based on an analysis of diet and short-chain fatty acid (SCFA) profiles from a previous study we initially partitioned the twelve species into four dietary categories, as follows: (a) Category 1: herbivores with a diet of macroscopic brown algae and high SCFA profiles in the hindgut region (Naso unicornis, Kyphosus vaigiensis); (b) Category 2: herbivores feeding on turfing and filamentous red and green algae with moderate SCFA profiles in the hindgut region (N. tonganus, K. cinerascens, Zebrasoma scopas, Acanthurus lineatus); (c) Category 3: zooplankton feeders with moderate SCFA profiles (N. vlamingii, N. brevirostris); (d) Category 4: species feeding on detrital and sedimentary materials with low levels of SCFA (Chlorurus microrhinos, Scarus schlegeli, Ctenochaetus striatus, A. olivaceus). The purpose of this comparison was to determine whether measures of feeding activity, alimentary tract structure, and food processing were concordant with diet. A dichotomy in feeding rates was observed. Species with a diet of algae and zooplankton (categories 1–3) had slower feeding rates than those feeding on detrital aggregates and sediment (category 4). The pattern of food processing also followed the same dichotomy with species of categories 1–3 retaining food in the alimentary tract overnight and commencing the feeding day with substantial amounts of food in the intestine and hindgut. Category-4 species commenced the feeding day with empty alimentary tracts suggesting a rapid turnover of gut contents. Within the herbivorous and zooplankton-feeding species neither alimentary tract structure nor food processing mode were predicted by diet or SCFA profiles. A hindgut fermentation chamber was present in K. vaigiensis but not in N. unicornis, a species with high levels of SCFA in the hindgut region and a diet of brown macroscopic algae. In contrast N. vlamingii, with a diet dominated by animal matter, retained large amounts of food material in a hindgut chamber over the entire feeding cycle. In tropical perciform fishes, herbivory and fermentation are not associated with the alimentary tract structures that characterise herbivorous terrestrial vertebrates. Estimates of the abundance of the different groupings of nominally herbivorous fishes indicated that the dominant elements in the reef grazing and browsing fauna were consumers of detrital and sedimentary materials. These could not be classified as herbivores. Members of this group were dominant in all habitats investigated. Explicitly herbivorous taxa were a minority component in all habitats investigated.Communicated by G.F. Humphrey, Sydney     

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.     

Growth response of an early successional assemblage of coralline algae and benthic diatoms to ocean acidification

The sustained absorption of anthropogenically released atmospheric CO₂ by the oceans is modifying seawater carbonate chemistry, a process termed ocean acidification (OA). By the year 2100, the worst case scenario is a decline in the average oceanic surface seawater pH by 0.3 units to 7.75. The changing seawater carbonate chemistry is predicted to negatively affect many marine species, particularly calcifying organisms such as coralline algae, while species such as diatoms and fleshy seaweed are predicted to be little affected or may even benefit from OA. It has been hypothesized in previous work that the direct negative effects imposed on coralline algae, and the direct positive effects on fleshy seaweeds and diatoms under a future high CO₂ ocean could result in a reduced ability of corallines to compete with diatoms and fleshy seaweed for space in the future. In a 6-week laboratory experiment, we examined the effect of pH 7.60 (pH predicted to occur due to ocean acidification just beyond the year 2100) compared to pH 8.05 (present day) on the lateral growth rates of an early successional, cold-temperate species assemblage dominated by crustose coralline algae and benthic diatoms. Crustose coralline algae and benthic diatoms maintained positive growth rates in both pH treatments. The growth rates of coralline algae were three times lower at pH 7.60, and a non-significant decline in diatom growth meant that proportions of the two functional groups remained similar over the course of the experiment. Our results do not support our hypothesis that benthic diatoms will outcompete crustose coralline algae under future pH conditions. However, while crustose coralline algae were able to maintain their presence in this benthic rocky reef species assemblage, the reduced growth rates suggest that they will be less capable of recolonizing after disturbance events, which could result in reduced coralline cover under OA conditions.     

Mass mortality ofDiadema antillarum

It has been hypothesized that herbivorous fishes and the regular echinoidDiadema antillarum Philippi compete for benthic algae as their major food resource. Mass mortality ofD. antillarum in February 1984 provided the opportunity to test the hypothesis that herbivorous fishes and sea urchins were competing previously. Visual censuses of herbivorous fishes conducted over 4 yr in four reef zones on Tague Bay Reef, St. Croix, U.S. Virgin Islands, before and after the mass mortality indicated that population densities increased approximately three-fold in backreef and shallow (2m) forereef zones and two-fold, and four-fold in mid (5m) and deep (10m) forereef zones, respectively. Juvenile parrotfishes constituted the major component of these increases, except in the shallow forereef where acanthurids became most abundant. Grazing intensity by herbivorous fishes increased in three of the four reef zones immediately following the mass mortality. These data support the hypothesis that exploitative competition for algal resources was occurring prior to the sea urchin mass-mortality, although alternative hypotheses cannot be discounted completely. Despite the increases in the abundances of, and grazing by, herbivorous fishes, the algal community continued to increase in percent cover and biomass, indicating that increased grazing by fishes does not compensate for the loss of grazing byD. antillarum in controlling algal abundance and community structure.     

Sources of variation in herbivore preference: among-individual and past diet effects on amphipod host choice

Understanding which factors affect the feeding preferences of herbivores is essential for predicting the effects of herbivores on plant assemblages and the evolution of plant–herbivore interactions. Most studies of marine herbivory have focussed on the plant traits that determine preferences (especially secondary metabolites), while few studies have considered how preferences may vary among individual herbivores due to genetic or environmental sources of variation. Such intraspecific variation is essential for evolutionary change in preference behaviour and may alter the outcome of plant–herbivore interactions. In an abundant marine herbivore, we determined the relative importance of among-individual and environmental effects on preferences for three host algae of varying quality. Repeated preference assays were conducted with the amphipod Peramphithoe parmerong and three of its brown algal hosts: Sargassum linearifolium, S. vestitum and Padina crassa. We found no evidence that preference varied among individuals, thus constraining the ability of natural selection to promote increased specialisation on high-quality S. linearifolium. Most of the variation in preference occurred within individuals, with amphipod preferences strongly influenced by past diet. The increased tendency for amphipods to select alternate hosts to that on which they had been recently feeding indicates that amphipods are actively seeking mixed diets. Such a feeding strategy provides an explanation for the persistence of this herbivore on hosts in the field that support poor growth and survival if consumed alone. The effects of past diet indicate that herbivore preferences are a function of herbivore history in addition to plant traits and are likely to vary with the availability of algae in space and time.     

Avian herbivory: an experiment, a field test, and an allometric comparison with mammals

Mechanistic studies on herbivore functional responses have largely taken place in mammals; very little has been done in herbivorous birds so far. Here we aim to fill that gap by experimentally quantifying the (short-term) functional response of a large avian herbivore, the Bewick's Swan (Cygnus columbianus bewickii). We explicitly distinguish between encounter-limited and handling-limited foraging by analyzing the results in the framework of the models of D. E. Spalinger and N. T. Hobbs, originally developed for mammalian herbivory. Bite size in captive swans was experimentally manipulated by varying sward height. The time interval between two bites increased with bite size, which supports the handling-limited model (process 3) and rejects the encounter-limited models (processes 1 and 2). Subsequently, we took the obtained functional response parameters into the field in order to predict, from measurements of sward height, (1) bite sizes, (2) handling times, and (3) short-term intake rates in free-ranging swans. Indeed, for all three variables, the observed values closely matched the experimentally based predictions. Finally, we review functional response parameters available in the literature on avian herbivores and scale them allometrically in relation to mammals. This analysis revealed that maximum bite sizes, and therefore maximum intake rates, in herbivorous birds are smaller than in herbivorous mammals. We hypothesize and provide evidence that birds compensate by longer daily foraging times.     

Settlement and recruitment of sea urchins (Strongylocentrotus spp.) in a sea-urchin barren ground and a kelp bed: are populations regulated by settlement or post-settlement processes?

I sampled recruitment of very small sea urchins (Strongylocentrotus spp.) by using the anesthetic magnesium chloride to remove individuals from substrata collected in sea-urchin barren grounds (barrens) and kelp beds at Naples Reef near Santa Barbara, California, USA. Preliminary sampling found low numbers of newly settled individuals(<0.6 mm test diam) from April–July in 1984 and 1985, and in April, 1986. In early May, 1986, I found many newly settled seaurchins (0.3 to 0.6 mm, 5 to 17 d old), and I compared the densities of the cohort on several types of natural substrata in barrens and kelp-bed habitats. Newly settled individuals of both purple sea urchins (S. purpuratus) and red sea urchins (S. franciscanus) were present in similar, high densities (1 000 S. purpuratus m^-2) on foliose red algal turf, a dominant substratum ofthe kelp bed, and on crustose coralline algae, the dominant substratum of an adjacent barrens. Larvae of S. purpuratus reared and tested in the laboratory showed high rates of settlement on both red algal turf and on crustose coralline algae, but significantly lower rates on rock. Larvae also settled in response to a partiallypurified extract of coralline algae. The reduced settlement on natural rock surfaces relative to either algal treatment and the significant settlement in response to the extract of coralline algae indicate that larvae discriminate between natural substrata and probably respond to a settlement cue other than, or in addition to, a simple microbial (bacterial) film. The similar densities of young recruits of S. purpuratus on dominant substrata of barrens and kelp bed show that, at least in this case, differential settlement cannot explain the high densities of sea urchins in the barrens habitat. Movement between barrens and kelp bed is unlikely given the small sizes of the newly recruited sea urchins relative to the large distances often involved. Reduced post-settlement mortality of newly settled individuals in the barrens remains the most likely mechanism leading to the higher densities of sea urchins in barrens relative to kelp-bed habitats.