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.     

Grazing dynamics in intertidal rockpools: Connectivity of microhabitats

Differences between rockpool and emergent rock communities are often attributed to their contrasting physical conditions. However, differences in grazing pressure between rockpools and open rock could also exert an important structuring role. Greater densities and/or the lack of tidal constraints on foraging may allow grazing intensity to be greater in rockpools. Here, wax discs were deployed to compare grazing intensity between rockpool and emergent rock habitats at each of three tidal heights on a moderately exposed shore in SW England. Grazing intensity was then examined in relation to herbivore density. Grazing intensity in pools was twice that on emergent rock, despite a lower density of herbivores in the rockpools. Of these herbivores, patellid limpets are the dominant grazers on rocky shores throughout the NE Atlantic and are recognised to have a major role in structuring intertidal communities. Thus, subsequent experiments focussed on the influence of limpets in determining the differences in consumer pressure between rockpools and emergent rock. Three alternative explanations were considered: (1) the effect of continuous immersion on grazing intensity in rockpools; (2) differences in limpet species abundance between the two habitats; (3) movement of limpets from emergent rock into pools to feed. The level of grazing pressure exerted by Patella ulyssiponensis (Gmelin), the predominant species living constantly immersed in rockpools, was similar to that of P. vulgata (Linnaeus) which is predominantly found on emergent rock. P. vulgata were observed moving from emergent rock into rockpools during high tide. Manipulative experiments confirmed that these foraging excursions resulted in a 2-fold increase in grazing intensity in the pools. Grazing activity of P. vulgata in rockpools was not consistent between sites and may be influenced by differences in wave exposure and/or the abundance of microbial resources. Elevated consumer pressure in rockpools may be an important factor influencing algal assemblages and probably explains the predominance of grazer resistant-species in these pools.     

Niche partitioning among three montane ground‐dwelling pheasant species along multiple ecological dimensions

Pheasants (order Galliformes) are typical ground‐dwelling birds, having a large body size and weak flight abilities. Sympatric pheasants are expected to share narrower niche space and face more extensive interspecific competition. However, little work has been undertaken simultaneously to investigate niche partitioning among sympatric pheasant species across multiple ecological dimensions. We compared microhabitat use, activity pattern and foraging strategy of sympatric Blood Pheasant Ithaginis cruentus, Buff‐throated Partridge Tetraophasis szechenyii and White Eared‐pheasant Crossoptilon crossoptilon on the Qinghai‐Tibet Plateau, China, to identify potential interspecific niche partitioning along different ecological dimensions in the breeding season. We found that the Buff‐throated Partridge differed significantly from the other two species in microhabitat use, and the three species showed different foraging strategies. It is likely that niche partitioning reduced potential interspecific competition, thus facilitating the species’ stable coexistence. Our study provides practical evidence of multidimensional niche theory within sympatric ground‐dwelling pheasant species, emphasizing that species interactions and coexistence within a guild are often not uni‐dimensional. Given global conservation concern for maintaining bird diversity, we recommend further restriction of yak grazing in these species’ habitat.     

Modeling foraging behavior of piping plovers to evaluate habitat restoration success

Habitat restoration projects are often deemed successful based on the presence of the target species within the habitat; however, in some cases the restored habitat acts as an ecological trap and does not help to improve the reproductive success of the target species. Understanding wildlife–habitat relationships through precise measurements of animal behavior can identify critical resources that contribute to high quality habitat and improve habitat restoration practice. We evaluated the success of a restored piping plover (Charadrius melodus) breeding habitat in New Jersey, USA. We identified the major factors influencing foraging rates, compared foraging activity budgets over 3 yr at restored and natural habitats, and explored the potential of artificial tidal ponds as a viable restoration alternative. Adult foraging rates were higher in artificial pond and ephemeral pool habitats, during low tide, and after breeding activity ended. Adult foraging rates were impeded by the presence of people and vehicles within 50 m. Chick foraging rates were highest at artificial ponds and bay shores and lowest in dunes and on sand flats. Chick foraging rates were strongly hindered by the presence of corvids and the number of people within 50 m. In addition, at artificial tidal ponds, piping plovers spent more time foraging and less time engaged in defensive behaviors (vigilance, crouching, and fleeing) compared to other potential habitats. Our findings support the hypothesis that artificial tidal ponds are a valuable, perhaps superior, foraging habitat. Future beach restoration projects should include this feature to maximize habitat quality and restoration success. © 2011 The Wildlife Society.     

Dynamic spatial partitioning and coexistence among tall grass grazers in an African savanna

Competitive relationships among mobile animals may be expressed through dynamically changing spatial relationships over different time frames. Less common species that are apparently inferior competitors may be able to coexist with more abundant species by concentrating in regions of the landscape little utilized by the former at spatio‐temporal scales from annual or seasonal ranges to the specific foraging localities exploited at different stages of the annual cycle. Spatial relationships may be influenced further by dependencies on other resources, predation risks and facilitatory interactions under certain conditions. Our study aimed to determine whether competition with more abundant zebra and buffalo restricted the abundance of sable antelope in a region where these three tall‐grass grazers overlapped in their herd distributions. We tracked the simultaneous movements of animals representing herds of these species over two dry seasons and one wet season using GPS‐GSM collars, and estimated seasonal or monthly range extents and their overlap. We also compared daily separation distances between these animals against the null pattern expected if their movements had been independent, and assessed how prior grazing by buffalo influenced the subsequent use of these localities by sable. The range of the sable herd was mostly separated from the seasonal range of the buffalo herd during the late dry season of 2006 and throughout the dry season of 2007. Seasonal home ranges of zebra herds overlapped partially with the range of the sable herd during most of the year. Even during times when their ranges overlapped, sable were rarely recorded within <1 km of the buffalo herd. Prior grazing by buffalo beyond a threshold level inhibited later use of these localities by sable, but the sable were nevertheless able to exploit places that were little utilized by buffalo at that time. Sable were less able to evade overlap with the small, mobile zebra herds, and hence more vulnerable to competitive exclusion by zebra than by buffalo. Our findings demonstrate how less abundant species can restrict competition from more abundant competitors through dynamic spatial partitioning in regions where their home ranges overlap.     

The persistent spatial patchiness of limpet grazing

The characteristic variability of grazing has potential consequences for intertidal productivity and community structure, particularly as many of the underlying functional relationships are thought to be non linear. As a first approximation, it can be hypothesised that grazing is patchy over short time periods before a more uniform coverage is established over longer time scales. This prediction is supported by relatively short term observations previously made of limpet foraging. We used eight arrays of wax disks on each of four shores to test the hypothesis that grazing is patchy in the short term, but that this pattern is lost as the pattern of grazing averages out over longer time scales. Wax disk arrays were exposed for two weeks at a time for a period of six weeks in 2001 and in 2002 using the same set of disk holes each time. Grazing at the same disk location could therefore be measured over two weeks and over longer periods by averaging successive deployments. We used all three successive deployments to estimate the average grazing at each disk location over a six week period in 2001 and 2002. All six deployments were used to characterise the pattern of grazing at longer time scales. The spatial pattern of grazing in arrays was summarized using semivariogram analyses. For two-week deployments, the average standardized semivariance of grazing for disks separated by 20 cm was less than one. This pattern implies spatial autocorrelation of grazing at this scale. There was no support for the hypothesis that small scale patchiness in grazing would disappear over time. The average strength of spatial autocorrelation increased when data were integrated over longer periods. A preliminary analysis indicated that the degree of autocorrelation within arrays increased with grazing intensity at short time scales. Surface roughness disrupted autocorrelation of grazing over both short and long time scales. The persistent patchiness of grazing is likely to have implications for biofilm productivity, particularly on smoother shores.     

Incorporating tides into the acoustic ecology of humpback whales

North Pacific humpback whales (Megaptera novaeangliae) migrate annually to foraging grounds in Southeast Alaska that are characterized by semidiurnal tidal cycles. Tidal activity is an important driver of marine mammal behavior on foraging grounds, but is often omitted in studies of acoustic ecology. To better understand the role of sound in this vocal species we investigated the influence of tidal height and direction on humpback whale nonsong calling behavior in Frederick Sound and described new call types for this population. The likelihood of detecting a call from the low‐frequency‐harmonic, pulsed, or noisy‐complex call classes was independent of tidal activity. The likelihood of detecting a call from the tonal call class, and a feeding call in particular, was 2.1 times higher during flood tides than during ebb tides (95% CI 1.1–4.4). This likely reflects an indirect relationship between humpback whale foraging and tides.     

Search strategies of ants in landmark-rich habitats

Search is an important tool in an ant’s navigational toolbox to relocate food sources and find the inconspicuous nest entrance. In habitats where landmark information is sparse, homing ants travel their entire home vector before searching systematically with ever increasing loops. Search strategies have not been previously investigated in ants that inhabit landmark-rich habitats where they typically establish stereotypical routes. Here we examine the search strategy in one such ant, Melophorus bagoti, by confining their foraging in one-dimensional channels to determine if their search pattern changes with experience, location of distant cues and altered distance on the homebound journey. Irrespective of conditions, we found ants exhibit a progressive search that drifted towards the fictive nest and beyond. Segments moving away from the start of the homeward journey were longer than segments heading back towards the start. The right tail distribution of segment lengths was well fitted by a power function, but slopes less than −3 on a log-log plot indicate that the process cannot be characterized as Lévy searches that have optimal slopes near –2. A double exponential function fits the distribution of segment lengths better, supporting another theoretically optimal search pattern, the composite Brownian walk.     

Investigating foraging utilization distribution of female New Zealand sea lions, Auckland Islands

Detrimental interactions between marine mammals and fisheries are increasing worldwide. The ability to manage these interactions requires the knowledge of where and how interactions occur and the effects they have on species. Many pinnipeds are central place foraging colonial breeders who are restricted in foraging range during breeding. Here, we use a utilization distribution approach to examine the foraging habitats of lactating New Zealand (NZ) sea lions (Phocarctos hookeri) from Dundas and Enderby Islands, Auckland Islands. Annually, the NZ sea lions which breed on these two islands produce 83% of this Nationally Critical species’ pups. Satellite transmitters were attached to 55 females during 2001–2007. Data showed that NZ sea lions utilize the entire Auckland Island shelf with partial habitat partitioning between females from the two breeding islands. This habitat partitioning results in differing degrees of overlap with fisheries and therefore possible differing fishery-related impacts on breeding areas.     

Foraging movements and homing in the limpet Patella vulgata L

In two populations of Patella vulgata in the Isle of Man foraging activity is restricted to daytime high waters, with about 75 per cent of the population foraging on any one day. Movement commences as the tide covers the limpets, and they travel an average of 0·4 m from their home sites to which they return at least an hour before the falling tide exposes them again. The mean speed on the outward and return journey is around 0·60 cm min^?, and during the intervening browsing period about 0·08 cm min^?. This behaviour contrasts with that of the same species in Alderney, where foraging is limited to nocturnal low waters: there is possibly a geographical trend in behaviour. P. vulgata, especially in the Isle of Man, is one of the more mobile limpets.     

TEMPORAL VARIATION IN FORAGING BEHAVIOUR OF PATELLA GRANULARIS (PATELLOGASTROPODA) AND SIPHONARIA CONCINNA (BASOMMATOPHORA) ON A SOUTH AFRICAN SHORE

Foraging activity of two mid- to low- shore species of limpet,Patella granulans (Prosobranchia) and Siphonaria concinna (Pulmonata)from an exposed shore on the Eastern Cape coast of South Africawas monitored. In both species, activity was compared duringspring and neap tides and, in P. granularis between summer andwinter. Rhythms of activity of the two species were similar,with foraging excursions being mainly associated with nocturnallow tide times, although some P. granularis foraged during daytimelow tides. It is suggested that foraging excursions in P. granularisare triggered by wave action. Both species foraged further onspring tides than on neap tides and this is suggested to bea result of the limited time limpets have to forage. P. granulariswas also found to forage further during summer when comparedto winter and the possibility that seasonal micro-algal productivityinfluences foraging distances in limpets is discussed. The foraging activity of both species could be divided into3 distinct phases, a relatively rapid outward phase, a muchslower foraging phase and a rapid homeward phase. Whether ornot these limpets graze throughout an excursion is not known.S. concinna was found to home to a fixed scar, although duringthe experiment some scar-swapping occurred. P. granularis didnot home to a fixed scar but possessed a ‘home range’(approx. 5 cm²) to which it returned after each excursion. Patella granularis was found to move randomly during foraging,whilst S. concinna foraged in a non-random direction -whichtook individuals upshore. No tidal-influence is thought to bepresent in this case and the possibility of a learning componentin the foraging behaviour of certain limpet species in relationto the return to optimal feeding patches is discussed. (Received 18 May 1996; accepted 2 September 1996)     

Interactions between rate processes with different timescales explain counterintuitive foraging patterns of arctic wintering eiders

To maximize fitness, animals must respond to a variety of processes that operate at different rates or timescales. Appropriate decisions could therefore involve complex interactions among these processes. For example, eiders wintering in the arctic sea ice must consider locomotion and physiology of diving for benthic invertebrates, digestive processing rate and a nonlinear decrease in profitability of diving as currents increase over the tidal cycle. Using a multi-scale dynamic modelling approach and continuous field observations of individuals, we demonstrate that the strategy that maximizes long-term energy gain involves resting during the most profitable foraging period (slack currents). These counterintuitive foraging patterns are an adaptive trade-off between multiple overlapping rate processes and cannot be explained by classical rate-maximizing optimization theory, which only considers a single timescale and predicts a constant rate of foraging. By reducing foraging and instead digesting during slack currents, eiders structure their activity in order to maximize long-term energetic gain over an entire tide cycle. This study reveals how counterintuitive patterns and a complex functional response can result from a simple trade-off among several overlapping rate processes, emphasizing the necessity of a multi-scale approach for understanding adaptive routines in the wild and evaluating mechanisms in ecological time series.     

Riding the tide: intriguing observations of gulls resting at sea during breeding

Adult Lesser Black‐backed Gulls Larus fuscus were followed during the breeding season using a high‐resolution GPS tracking system. Little is known about the foraging movements of these birds and what they do when they are away from the colony. The study revealed intriguing yet infrequent behavioural patterns showing that birds would sit on the sea surface drifting passively with the tidal current for several hours, resting at sea rather than in the breeding colony. The potential function of the observed patterns is discussed as well as the relevance of these observations to other species.     

Absence of a tidal component in the diel pattern of locomotory activity of sea catfish,Arius felis

Synopsis Sea catfish, tested individually for 3-day periods in an automated data acquisition system, displayed a nocturnal pattern of locomotor activity with 62% of the activity occurring during scotophase. In addition, there were two relative maxima of activity at 0700–0900 and 1300–1500 C.S.T. This diel pattern of activity could not be correlated with tidal phenomena in the fish's natural environment and is virtually identical to that exhibited by several freshwater catfish species. Thus, even though sea catfish are a primarily estuarine fish and subject to daily tidal fluctuations, tidal rhythmicity is apparently not used as a cue for diel activity patterns.     

Experimental evidence for the interplay of exogenous and endogenous factors on the movement ecology of a migrating songbird

Movement patterns during songbird migration remain poorly understood despite their expected fitness consequences in terms of survival, energetic condition and timing of migration that will carry over to subsequent phases of the annual cycle. We took an experimental approach to test hypotheses regarding the influence of habitat, energetic condition, time of season and sex on the hour-by-hour, local movement decisions of a songbird during spring stopover. To simulate arrival of nocturnal migrants at unfamiliar stopover sites, we translocated and continuously tracked migratory red-eyed vireos (Vireo olivaceus) throughout spring stopover with and without energetic reserves that were released in two replicates of three forested habitat types. Migrants moved the most upon release, during which time they selected habitat characterized by greater food abundance and higher foraging attack rates. Presumably under pressure to replenish fuel stores necessary to continue migration in a timely fashion, migrants released in poorer energetic condition moved faster and further than migrants in better condition and the same pattern was true for migrants released late in spring relative to those released earlier. However, a migrant's energetic condition had less influence on their behavior when they were in poor quality habitat. Movement did not differ between sexes. Our study illustrates the importance of quickly finding suitable habitat at each stopover site, especially for energetically constrained migrants later in the season. If an initial period prior to foraging were necessary at each stop along a migrant's journey, non-foraging periods would cumulatively result in a significant energetic and time cost to migration. However, we suggest behavior during stopover is not solely a function of underlying resource distributions but is a complex response to a combination of endogenous and exogenous factors.     

阿拉善荒漠连续放牧对三趾跳鼠巢区的影响

三趾跳鼠(Dipus sagitta)是内蒙古西部荒漠草原和沙地的优势害鼠,巢区研究对了解探讨其繁殖策略和种群动态规律有着重要的理论和实践意义。本研究通过2012~2014年的野外标志重捕实验,对三趾跳鼠的巢区在连续放牧和禁牧条件下的变动特征进行了分析。结果表明,连续放牧区雌性三趾跳鼠在6月的巢区范围显著大于禁牧区(P0.05);禁牧区雄性三趾跳鼠的巢区仅在7月份显著大于连续放牧区(P0.05),其他季节没有显著差异(P0.05)。放牧对雄性巢区无显著影响(P0.05),但连续放牧使雌性三趾跳鼠巢区范围显著大于禁牧区。雌性与雄性三趾跳鼠巢区均与一年生草本生物量存在正相关关系。这些结果表明,相对于雄性三趾跳鼠来说,雌性对由放牧导致的食物资源变化比较敏感,这种现象在哺乳期更为显著;繁殖是影响雄性三趾跳鼠巢区的因素之一。因此,保卫资源假说的理论可能是放牧导致三趾跳鼠巢区变化的重要原因之一。     

Experimental manipulation of predation risk and food quality: effect on grazing behaviour in a central-place foraging herbivore

The relative importance of predation risk and food quality on spatial grazing pressure and activity patterns was tested in a central-place foraging herbivore: the European rabbit. Rabbits grazed less with increasing distance from their burrows into adjacent grassland, thereby creating a gradient of increasing vegetation height and plant biomass and decreasing plant nutrient concentration. When nitrogen concentration was experimentally increased by 150% through fertilizing and mowing, rabbits visited these plots four times more frequently than the untreated control plots. Addition of predator scent (mink pellets) did not result in different patch use by rabbits. The combined addition of fertilizer and mink pellets had the same effect as addition of fertilizer alone. However, the mink pellets changed the temporal activity pattern of rabbits as measured with infrared detectors. Rabbits were predominantly nocturnal but shifted their activities to the day when mink pellets were added, resulting in equal activities during night and day. We conclude that rabbits are sensitive to perceived predation risk, but that this does not influence their spatial grazing pressure. A selection for the highest food quality on the other hand can explain the observed natural rabbit grazing gradient. Food quality was highest close to the burrows, therefore rabbits selecting for high quality food should forage most intensely close to the burrows and only move further away for higher quality items or when the vegetation close to their burrows is depleted. Through intensive grazing close to the burrows rabbits facilitated for themselves either through stimulating fresh protein rich re-growth or the return of nutrients through faeces or both. This is in contrast with central-place foraging theory where intense feeding close to the burrow is assumed to lead to reduced food resources.     

Temporal pattern of foraging and microhabitat use by Galápagos marine iguanas,Amblyrhynchus cristatus

We observed a colony of marine iguanas (Amblyrhynchus cristatus) on Isla Fernandina, Galápagos, Ecuador, while measuring local micrometeorological and tidal conditions. We found size-related differences in foraging mode, with smaller iguanas feeding intertidally during daytime low tides and larger iguanas feeding subtidally. Despite having greater opportunity, subtidal foragers did not time their foraging bouts or exploit their environment in ways that optimized their period at high body temperature. Instead, the foraging schedule of these iguanas served to maximize their rate of rewarming following emergence from the cool sea. Intertidal feeders, by contrast, showed much greater behavioral flexibility in attempting to exploit their thermal environment. We suggest that size-ordered differences in marine iguana thermoregulatory behavior reflect underlying ontogenetic changes in costs and benefits of thermoregulation due to differences in predator pressure, quantity of food and electrolytes taken at each feeding, mode of foraging, and agonistic tendencies.     

Foraging strategies of Great Cormorants Phalacrocorax carbo carbo wintering north of the Arctic Circle

This study describes how 30 Great Cormorants Phalacrocorax carbo carbo managed to catch sufficient food for their daily energetic needs under conditions of reduced daylight and cold while wintering north of the Arctic Circle. Activity observations showed that the Great Cormorants' daily foraging pattern was generally bimodal, with morning and evening feeding peaks. They compensated for shorter daylengths in midwinter by starting to forage later and ending progressively earlier at lower light intensities. Fishing constituted only a minor part of their time–activity budget, and was one of the most efficient reported in marine birds. The Great Cormorants spent less than 60 minutes a day fishing in midwinter. Although subzero ambient temperatures and blizzards contributed to increased heat loss in midwinter, this potential energy loss did not seem to be compensated for by an increase in fish intake. Instead the Great Cormorants seemed to economize energy expenditure by halving the time spent at sea, and halving the number but doubling the mass of each fish taken.