A satiation model for the temporal organization of grazing in limpets

1. A compartmental model of the limpet alimentary system was designed to simulate the effects of functional constraints and substrate characteristics on the temporal patterning of grazing in these gastropods. Limitations are imposed by gut capacity and processing time. A maximum operating frequency is set for the radula activity, and a negative feedback on the grazing frequency is related to gut replenishment.
2. When holding constant the 'morphofunctional' parameters of the model, variations in the 'richness' of the substrate (i.e. in the amount of matter extracted per single rasp) generated different patterns of grazing activity, from constant grazing at different frequencies, to the appearance of distinct grazing–non-grazing subphases with a periodicity corresponding to the processing time. Moreover, when varying the digestibility of the matter (i.e. its processing time), temporary saturations were produced.
3. Automatic recording of radula activity in specimens of Patella caerulea L. collected from a Mediterranean boulder beach and transferred to tidal tanks showed that each activity phase (performed during the high-tide period) may be differently structured: from almost continuous grazing, with more or less pronounced and sometimes periodic oscillations, to alternation of grazing and non-grazing subphases. Continuous and non-continuous grazing patterns are shown also by the same specimen during subsequent activity phases.
4. The results of the model are consistent with empirical observations and suggest that satiation events related to the characteristics of the substrate encountered by the limpets when foraging, can produce highly variable grazing patterns with implications for the energetics of foraging in this group of grazers.     

The physiological control of feeding in corals: a review

Little is known about the factors that control food capture by the coral polyp and the way that internal physiological events may coordinate, or even modify, this behaviour. A total understanding of the nutritional dynamics of corals demands detailed study of their feeding behaviour, and recent advances in behavioural and electrophysiological techniques are starting to provide some insights into how polyps control this behaviour. This review describes our present understanding of the physiological control of feeding behaviour in corals and suggests new studies on food recognition, food capture, and ingestion.     

Territorial defense of an excess food supply by an algal grazing fish, ayu

The size of the territory defended by the grazing fish ayu, Plecoglossus altivelis, often increases with decreasing intruder pressure. This territorial extension occurs even after additional growth of the fish reaches a plateau due to a surplus supply of algal food. This raises questions about the ecological advantages of defending an apparently enlarged territory. Enclosures were used to carry out observations on fish maintained as: (i) individuals with algae attached to the whole bottom area (2.4m²); (ii) individuals with algae covering 0.8m²; and (iii) with three conspecifics (competitive use) with algae attached to 2.4m². In the competitive use enclosures, one (occasionally two) fish that exclusively foraged the area appeared in each of the groups. Algal resources were sufficiently renewable that no significant difference in growth rate was detected between the dominants of the competitive groups and the two types of solitary enclosures, indicating that the defended area supplied algal resources that exceeded the dominants ability to grow. All the territorial holders realized an increase in relative body weight, suggesting the occurrence of inhibition, which corresponds to Verners concept of superterritory. However, as inferred from the pheophytin/chlorophylla ratios, algae within a territory showed less senescence, perhaps owing to cropping by the dominant, and this was considered to be an adaptive response by the fish to an unpredictable food supply. In conclusion, the ayu conditionally defends a territory for long-term benefits, and inhibition is a consequence of facultative territory size and not a cause of it.     

Why are there few algae on snail shells? The effects of grazing,nutrients and shell chemistry on the algae on shells of Helisoma trivolvis

1. Freshwater snails often lack visible growths of algae on their shells. We tested three possible mechanisms that may account for this (grazing, snail-derived nutrients and chemical defences), using the ramshorn snail Helisoma trivolvis .
2. The experiments were carried out in floating plastic enclosures in a pond and comprised seven treatments. Grazing treatments were: a lone snail (ungrazed, as self-grazing does not occur), Helisoma with conspecifics, Helisoma with the co-occurring pond snail Physa sp., empty shells with Helisoma , and ungrazed empty shells. Nutrient effects were possible in all treatments with occupied shells (lone snail; Helisoma with conspecifics, and with Physa ) versus absent in other treatments. Testing for chemical defences compared algae on fresh empty shells, weathered shells (outer organic periostracum layer absent) and boiled fresh shells (with denaturization of susceptible proteins).
3. Diatoms dominated algal assemblages on snail shells. Although the upright diatom Gomphonema gracile was abundant on all shells, it was dominant on the shells of snails housed with other snail grazers (either Helisoma or Physa ).
4. Only the lone snail (nutrients but no grazing) showed higher algal biomass, so presumably any nutrient effect in the treatments with grazers was masked. Both Helisoma and Physa were observed apparently grazing on Helisoma shells, and consequently algal biomass in multi-snail treatments was similar to that on empty shells. Scanning electron microscopy revealed that algal density was highest near the aperture of live snails, but not empty shells; this is consistent with a nutrient addition effect. There was no evidence of chemical defences against algal growth.
5. In soft-bottomed freshwater habitats with abundant snails, shells of living snails provide nutrient-augmented substrata that may indirectly boost overall snail production.     

Amphibious behaviour and feeding ecology of the four-eyed blenny (Dialommus fuscus, Labrisomidae) in the intertidal zone of the island of Santa Cruz (Galapagos, Ecuador)

The labrisomid Dialommus fuscus , endemic to the Galapagos archipelago, is a predator of crustaceans, which moves up and down the intertidal zone with the tides. Its divided eye and elongated body permit it to forage in the wet sections of the intertidal habitat during low tide, both on the rocks and below the water line and in tide pools. Compared with other intertidal fishes (e.g. Bathygobius sp.), it was present on the rock flat in significantly smaller tide pools nearer to the water line during low tide.     

Aerial and aquatic feeding in the silver arawana,Osteoglossum bicirrhosum

Synopsis The silver arawana, Osteoglossum bicirrhosum, hunts along shorelines and within flooded forests in the Amazon River basin and supplements its limited consumption of aquatic vertebrates by leaping from the water to obtain terrestrial and arboreal prey. We offered O. bicirrhosum prey both suspended above and submerged below the surface of the water. From high-speed digital recordings, we measured kinematic variables associated with the jaws, cranium, pectoral fins, and body during orientation and prey capture. Aquatic and aerial feeding events were kinematically distinct, with aerial events generally involving faster, larger movements and a distinct delay in the onset of lower jaw depression until the head had left the water. The comparatively large gape during leaping may facilitate prey capture by overcoming variability in the apparent position of the prey due to refraction, while the delayed onset of mouth opening may serve to reduce the effects of drag. This distinctive leaping behaviour allows exploitation of the terrestrial prey base, especially during seasonal inundation of the Amazon River basin when the aquatic food base is widely dispersed.     

Rocky shore turfs dominated by Corallina (Corallinales,Rhodophyta) in northern Japan†

Three intertidal sites dominated by Corallina turfs were investigated in Hokkaido, Japan. The sites (A, B and C) differed in slope, wave exposure and length of time exposed to air during tidal cycles. Monthly samples were analyzed for frond morphology and other features. Site A, the most wave-exposed site, was dominated by Corallina sp. X, an unknown species, and sites B and C by Corallina pilulifera Postels et Ruprecht. At the different sites the populations differed in conceptacle abundance, coverage by epiphytic Titanoderma corallinae (P. Crouan et H. Crouan) Woelkerling, Chamberlain et Silva, amount of contained sediment, numbers of axes per quadrat, numbers of branch fusions, branch entanglement, frond dryweight, frond length, amount of adventitious branching, numbers of epiphytes (exclusive of T. corallinae), and numbers of animal species. Ninety-one animal species were recorded from the turfs. Corallina is affected morphologically by conditions inherent in its microhabitat, including desiccation, epiphyte loading and the abundance of herbivores.     

Feeding behaviour in Littorina littorea: a study of the effects of ingestive conditioning and previous dietary history on food preference and rates of consumption

Feeding responses of the generalist herbivore, Littorina littorea (L.), to the perceived taste of macroalgae were assessed with respect to the effects of recent dietary intake and to overlapping versus nonoverlapping distributions of winkles and algae. The extent of grazing on artificial substrates impregnated with crude algal extracts was used as a measure of rate of response to the odour of preferred algae, and of feeding preference among less preferred algae, in a variety of designs. Adult L. littorea collected from a site where a range of algae were present showed preference among extracts of fucoids, whereas adults from a nearby site showed no such preference. Juvenile L. littorea of two weight cohorts collected from the former site responded faster to Porphyra umbilicalis extract-containing substrate than similar-sized animals from the latter site. Juveniles, fed either Porphyra, Ulva lactuca, or starved for two weeks in the laboratory, responded similarly to Ulva versus Porphyra extracts in a dose-dependent manner across a range of concentrations, although the Porphyra-maintained group consumed more of each, and the starved group less over seven days. Juveniles maintained on a mixed diet of Ulva and Porphyra consumed more Porphyra extract and less Ulva extract over the same period. These results are discussed in relation to the possible role of ingestive conditioning and previous dietary history in determining the occurrence and extent of chemically-mediated feeding preference in L. littorea.     

Positioning behaviour in fish shoals: a cost–benefit analysis

Shoal position can have a strong influence on individual fitness. Individuals in front positions of shoals were observed to have higher feeding rates than individuals elsewhere. Manipulation of nutritional state showed that hungry individuals had a stronger preference for front positions and that the duration of food-deprivation was positively correlated with the degree of the position preference. On the other hand, front positions (like other peripheral positions) probably incur costs in terms of increased predation risks. Experiments with Schreckstoff showed that frightened individuals seek the central part of the shoal. This suggests that individuals rotate their shoal positions according to the tradeoff between energy intake and predation risk.     

ESTABLISHMENT,PERSISTENCE AND DOMINANCE OF CORALLINA (RHODOPHYTA) IN ALGAL TURF1

The dominance of Corallina thalli in an intertidal algal assemblage was examined by a series of algal removal experiments to test the hypothesis that other tam are competitively excluded from rock substrate. An alternate explanation, that environmental factors seasonally filter out taxa leaving Corallina as the only alga adapted for year to year survival, was considered. Development of vegetation on patches of naturally exposed bare rock was monitored and compared with manipulated surfaces. Thalli of several species were selectively removed from exposed surfaces and intact turf;. changes in tam occupying primary substrate were recorded over more than three years. No significant differences in mean percent cover for Corallina, Lithothrix, bare rock, or algal crusts were found among treatments. Except for initial growth of colonizing species, abundances of other species dad not increase in the absence of Corallina. Large amounts of rock remained bare or intermittently covered by transient populations of short-lived algae. Data from single quad-rats, where individual clumps of Lithothrix were followed from month to month, indicated that the continued presence of this co-occurring and often abundant species depended on turnover of short-lived thalli rather than persistence of the same clumps. No interactions were found among the several categories of species that appeared after rock substrate was cleared. Most species were the same ones that grew, epiphytically in intact turf at the same time. In control quadrats Corallina maintained 59-95% cover while slowly increasing on surfaces earlier exposed. I predict that Corallina species will regain their dominance in the absence of competitors for primary substrate if the slowly spreading basal crusts remain undisturbed. Morphological and life history characteristics are identified that adapt Corallina to its dominant role in this habitat.     

Epiphytes on the rocky intertidal red alga RhodomelaLarix (Turner) C. Agardh: Negative effects on the host and food for herbivores?

The diversity of epiflora and fauna associated with a dominant turf-forming alga was examined in intertidal communities on the central Oregon coast. Epiphytes associated with the red alga, Rhodomelalarix (Turner) C. Agardh, were examined by surveying intertidal areas for the presence of epiphytes, and by following changes in epiphyte cover in marked quadrats of R. larix. The alga is host for at least 17 species of sessile plants and animals. To determine the role of some of the larger epiphytes in the community, Rhodomela plants were marked and monitored and herbivore feeding was examined. Data suggest that epiphytes decrease the growth rate of their host, increase the probability of axis breakage and decrease reproductive output. Epiphytes provide food for littorine snails and gammarid amphipods that live in the beds of the plant. Amphipods were found to decrease epiphyte cover on R. larix in laboratory tanks, suggesting that these herbivores may have beneficial effects on the host plant.     

Phagostimulation of Lytechinus variegatus (Lamarck) (Echinodermata: Echinoidea)

Artificial agarose foods containing either starch, glycogen, carragheenan, alginate, casein, gelatin, albumin, or peptones at a concentration of 1% dry weight were not phagostimulatory. Artificial foods containing sugars, especially galactose, at a concentration of 0.005 M were phagostimulatory. Of the common amino acids, only L‐phenylalanine was phagostimulatory at this concentration. The initiation of feeding postures (movement of the spines, tube feet, peristome, and Aristotle's lantern) of Lytechinus variegatus was little affected by the concentration of plant extracts. Many aspects of this behavior were attributable to physical rather than chemical stimulation. Feeding rates of L. variegatus upon artificial agarose foods containing either Thalassia testudinum or Gracilaria verrucosa were directly dependent upon plant tissue concentration. All artificial agarose food‐items were initially moved to the mouth and fed upon, however the time spent feeding was directly dependent upon plant type and tissue concentration. Retention of the artificial food‐items upon the test after feeding was not influenced by plant type or tissue concentration. Differences in consumption of foods by L. variegatus are determined by the degree of persistence in feeding upon encountered items and not be selection before initiating feeding or by retention of food‐items for subsequent feedings.     

Tiger moths and the threat of bats: decision-making based on the activity of a single sensory neuron

Echolocating bats and eared moths are a model system of predator–prey interaction within an almost exclusively auditory world. Through selective pressures from aerial-hawking bats, noctuoid moths have evolved simple ears that contain one to two auditory neurons and function to detect bat echolocation calls and initiate defensive flight behaviours. Among these moths, some chemically defended and mimetic tiger moths also produce ultrasonic clicks in response to bat echolocation calls; these defensive signals are effective warning signals and may interfere with bats'' ability to process echoic information. Here, we demonstrate that the activity of a single auditory neuron (the A1 cell) provides sufficient information for the toxic dogbane tiger moth, Cycnia tenera, to decide when to initiate defensive sound production in the face of bats. Thus, despite previous suggestions to the contrary, these moths'' only other auditory neuron, the less sensitive A2 cell, is not necessary for initiating sound production. However, we found a positive linear relationship between combined A1 and A2 activity and the number of clicks the dogbane tiger moth produces.     

The effects of foliage damage on casebearing moth larvae, Coleophora serratella, feeding on birch

ABSTRACT.

• 1 Mechanical damage to birch (Betula pendula Roth) leaves leads to an increase in the concentration of phenolic compounds, which spreads throughout the leaf within 8 days.
• 2 Coleophora serratella L. (Lepidoptera: Coleophoridae) apparently responds to this chemical change over a similar time scale. Within 24 h of pin-pricking leaves the casebearer moves from the immediate vicinity of the damage, but is just as likely to move to an undamaged portion of the damaged leaf as to vacate the leaf entirely. After 8 days mines on undamaged portions of damaged leaves were significantly smaller than mines on undamaged leaves.
• 3 Furthermore, Coleophora serratella reared on damaged trees took an average of 3 days longer to develop than those reared on undamaged trees.
• 4 It has been suggested that increased movement in response to damage-induced chemical changes causes hyperdispersed damage on plant foliage. Both within and between-leaf casebearer damage patterns were shown to be aggregated on birch.
• 5 Thus although mechanical damage can induce chemical and behavioural changes in the field, these are not reflected in the observed damaged patterns. We speculate on several possible reasons for this.

     

Impacts of silica-based defences in grasses on the feeding preferences of sheep

Grasses, which dominate many terrestrial ecosystems, sustain high densities of grazing mammals, so are of great economic and ecological importance. Traditionally, grasses are thought to be adapted to tolerate grazing rather than defend against it; however, silica deposited in the leaves of grasses has recently been shown to act as a feeding deterrent to invertebrate herbivores and small mammals. This study assesses whether silica is effective as a feeding deterrent to larger mammalian herbivores. We assess the impact of manipulated silica levels in five grass species on the feeding preferences of sheep both within and between grass species.Sheep feeding behaviour was driven by between-species differences in palatability. Hence, within a single species silica addition did not cause significant changes in feeding preference. However, there were significant differences in both the feeding preferences and bite rates between grass species, and these differences were much more marked when the grasses had been exposed to high levels of silica. The impacts that silica had on preference were least pronounced in palatable species (e.g. Poa annua) compared with less-palatable species (e.g. Brachypodium pinnatum and Festuca ovina). Sheep fed for longer, took more bites and had a higher bite rate on the grass species with the lowest leaf silica concentrations, namely P. annua.Sheep were less affected by silica defences than smaller, non-ruminant herbivores, but the changes in species preference rankings caused by silica suggest it may lead to changes in sward composition. Further, in species that are already relatively low in palatability, silica-induced reductions in bite rate could potentially reduce forage intake rates, with consequences for sheep performance.     

Reproduction and feeding ethology of a tropical,intertidal sand-dwelling anemone (Actinoporus elongatus,Carlgren, 1900)

Actinoporus elongatus is a widely distributed, tropical, intertidal anemone from the Indo-Pacific and Caribbean regions. Its recorded occurrences are disjunct and its feeding and reproductive ecology have not previously been described in detail. A population of this sand-dwelling species was studied for one year at Shelly Bay, Queensland, Australia. Individuals were found to be randomly distributed within the study site at a density of 42 anemones per hectare. No immigration, juvenile recruitment or mortality was observed. Histological analysis indicated a protracted period of gamete maturity during the summer months, and that the sex ratio was 1:1. A. elongatus is an opportunistic feeder with two distinct feeding methods — actinopharyngeal eversion and a newly described trapdoor response. Ciliary currents on the tentacular crown and oral disk were used mainly for cleaning and sand removal.Department of Zoology, James Cook University of North QueenslandDepartment of Marine Biology, James Cook University of North QueenslandDepartment of Marine Biology, James Cook University of North Queensland     

Low nutritive quality as a plant defence: Effects of herbivore-mediated interactions

Summary A plant may lower its nutritive quality, for herbivores, by using secondary compounds, morphological characters and/or having a lowered nutrient content. If such traits decrease the amount of resources lost through herbivory, then they act as antiherbivore defences. However, if herbivores compensate for the lowered nutrient availability, by increasing their intake rates or by prolonging their feeding periods, then this may render the defence useless. I analyse the conditions for evolution of this type of plant defences in a game theoretical model. The predictions of the model depend on the amount of compensatory feeding performed by the herbivores and on the herbivores' mobility in relation to the spatial structure of the plant population. When herbivores cannot compensate for a lowered nutritive quality, the defence can evolve irrespective of the type of herbivore. When herbivores can compensate for such defences, the outcome depends on how the herbivores compensate. In situations where herbivores compensate only on defended plants, which could correspond to immobile herbivores, this type of defence can evolve only if the level of compensation is lower than a certain critical value. When herbivores compensate more on defended than on undefended plants, e.g. because of low mobility, the outcome depends on the level of compensation performed on defended plants. If this level of compensation is high, then the model predicts a stable coexistence of defended and undefended plants and, if it is low, then the populations can consist of only defended plants. When herbivores compensate more on undefended plants than on defended ones, e.g. highly mobile herbivores, the result is populations consisting of either only defended plants, or only undefended plants. Consequently, the fact that herbivores may compensate for lowered nutrient quality does not, as such, nullify the notion of low nutrient quality as a plant defence. However, compensatory feeding may restrict the conditions for the evolution of such defences.