Extrinsic and intrinsic determinants of winter foraging and breeding phenology in a temperate seabird

In temperate regions, winter presents animals with a number of challenges including depressed food abundance, increased daily energy requirements, higher frequency of extreme weather events and shortened day length. Overcoming these constraints is critical for overwintering survival and scheduling of future breeding of long-lived species and is likely to be state dependent, associated with intrinsic abilities such as food acquisition rates. We examined the relationship between environmental and intrinsic factors on overwintering foraging and subsequent breeding phenology of the European shag Phalacrocorax aristotelis, a diurnal marine predator. We tested a range of hypotheses relating to overwintering foraging time and location. We found that individuals greatly increased their foraging time in winter to a peak of more than 90% of available daylight at the winter solstice. The seasonal patterns of foraging time appear to be driven by a combination of light levels and weather conditions and may be linked to the availability of the shag's principal prey, the lesser sandeel Ammodytes marinus. There was no evidence that shags dispersed south in winter to increase potential foraging time. Foraging time decreased after the winter solstice and, crucially, was correlated with subsequent breeding phenology, such that individuals that spent less time foraging in February bred earlier. The relationship was much stronger in females than males, in line with their more direct control of timing of breeding. Our results demonstrate that pre-breeding intrinsic foraging ability is critical in determining breeding phenology.     

Diet of Neotropic cormorant (Phalacrocorax brasilianus) in an estuarine environment

The diet of the Neotropic cormorant (Phalacrocorax brasilianus) was studied by analysing 289 regurgitated pellets collected from a roosting site at Lagoa dos Patos estuary, southern Brazil, between November 2001 and October 2002 (except April to June). In total, 5,584 remains of prey items from 20 food types were found. Fish composed the bulk of the diet representing 99.9% by mass and 99.7% by number. The main food items were White croaker (Micropogonias furnieri) (73.7% by frequency of occurrence, 48.9% by mass and 41.2% by number), followed by Catfish (Ariidae) and anchovies (Engraulididae). In Lagoa dos Patos estuary the generalist Neotropic cormorant fed mainly on the two most abundant demersal fishes (White croaker and Catfish), which accounted for the low niche breadth calculated. The total length of all fish preyed varied from 27.2 to 318.3 mm (113.5 ± 48.0 mm), and preyed White croakers’ size differed between months. Neotropic cormorants seem to prey on most abundant class sizes of White croaker instead of selecting similar prey size throughout the time. However, temporary changes in diet in terms of food items, abundance and prey size were detected, revealing a high ecological plasticity of the species. Individual daily food intake of Neotropic cormorants estimated by pellets and metabolic equations corresponded to 23.7 and 27.1% of their body mass, falling in the range of other cormorant species. Annual food consumption of the population estimated by both methods was 73.4 and 81.9 tonnes, comprising mainly immature and subadult White croaker and Catfish which are commercially important. Temporal variations in diet composition and fish size preyed by Neotropics cormorants, a widespread and generalist species, suggest shifts according to fluctuations in the abundance of prey. The plasticity of this cormorant is also revealed by their ability to adjust feeding behaviour in response to temporal or local changes in the environment, from a generalist at the species level to a specialist at the individual or local population level.     

Repellent scent-marking of flowers by a guild of foraging bumblebees (Bombus spp.)

We have found that foraging bumblebees (Bombus hortorum, B. pascuorum, B. pratorum and B.␣terrestris) not only avoid flowers of Symphytum officinale that have recently been visited by conspecifics but also those that have been recently visited by heterospecifics. We propose that the decision whether to reject or accept a flower is influenced by a chemical odour that is left on the corolla by a forager, which temporarily repels subsequent foragers. Honeybees and carpenter bees have previously been shown to use similar repellent forage-marking scents. We found that flowers were repellent to other bumblebee foragers for approximately 20 min and also that after this time nectar levels in S. officinale flowers had largely replenished. Thus bumblebees could forage more efficiently by avoiding flowers with low rewards. Flowers to which extracts of tarsal components were applied were more often rejected by wild B. terrestris workers than flowers that had head extracts applied, which in turn were more often rejected than flowers that had body extracts applied. Extracts from four Bombus species were equally repellent to foragers. The sites of production of the repellent scent and its evolutionary origins are discussed. Received: 24 November 1997 / Accepted after revision: 8 March 1998     

Feeding ecology of sympatric European shags Phalacrocorax aristotelis and great cormorants P. carbo in Iceland

The feeding ecology of the European shag (Phalacrocorax aristotelis) and the great cormorant (P. carbo) in Iceland was studied. These bird species may affect their marine environment, for instance, by predating on several commercially important fish species in coastal waters. The shag and cormorant diets were studied throughout the year in the period 1996–2000 by analysing the content of about 300 stomachs from each species. The shag relies heavily on sandeel (Ammodytes marinus) in the breeding season, whereas bull-rout (Myoxocephalus scorpius) and gadoids (Gadidae) become increasingly important in autumn and winter. The main food of the cormorant in all areas and seasons is the bull-rout. Depending on location and season, secondary food consists mainly of butterfish (Pholis gunnellus), gadoids and flatfishes (Pleuronectiformes). Similarity in diets of these two co-existing bird species was least in the breeding season when overlap in the birds’ distribution was greatest. The results suggest that predation by shags and cormorants could sometimes affect the stocks of the commercially important saithe (Pollachius virens) and plaice (Pleuronectes platessa) in Iceland.     

Energy expenditure and food consumption of foraging Imperial cormorants in Patagonia, Argentina

Energy management during the breeding season is crucial for central place foragers since parents need to feed themselves and their offspring while being spatially and temporally constrained. In this work, we used overall dynamic body acceleration as a measure of activity and also to allude to the foraging energy expenditure of breeding Imperial cormorants Phalacrocorax atriceps. We also analyzed how changes in the time or energy allocated to different activities affected the foraging trip energy expenditure and estimated the daily food requirements of the species. Birds spent 42 % of the total energy flying to and from the feeding areas and 16 % floating at sea. The level of activity underwater was almost 1.5 times higher for females than for males. The most expensive diving phase in terms of rate of energy expenditure was descending though the water column. The total foraging trip energy expenditure was particularly sensitive to variation in the amount of time spent flying. During the breeding season, adult cormorants breeding along the Patagonian coast would consume approximately 10,000 tons of food.     

Condition of larval and juvenile red drum (Sciaenops ocellatus ) from estuarine nursery habitats

RNA:DNA ratios of larval and juvenile red drum (Sciaenops ocellatus) collected from nursery habitats in the Aransas Estuary, Texas, in 1994 were quantified using a highly sensitive ethidium-bromide fluorometric technique. RNA:DNA ratios of wild red drum were evaluated by comparing individual values to a linear regression model derived for starved laboratory-reared red drum. Wild red drum were in relatively good condition with <5% of the RNA:DNA ratios within or below the 95% prediction interval of 4 to 5 d starved red drum. A multiple-regression model explained 54% of the variability in the RNA:DNA ratio of wild red drum, and identified length and water temperature (midday) as significant factors. RNA:DNA ratios increased with fish length [≃1.1 mm⁻¹, over the size range investigated (5␣to 20 mm)]. The effect of temperature on the RNA: DNA ratio was assessed on different sampling trips, and ratios increased with increasing temperature. Abundance of larval and juvenile red drum in the Aransas Estuary varied as a function of both habitat (shoal grass Halodule wrightii, turtle grass Thalassia testudinum) and site (Aransas Bay, Redfish Bay); however, no differences in RNA:DNA ratios were detected between habitats or between sites. It is postulated that the nutritional condition of newly settled red drum from the Aransas Estuary in 1994 was relatively high, and that starvation was of minor importance. Received: 19 August 1996 / Accepted: 23 August 1996     

Risk-sensitive foraging in common shrews (Sorex araneus L.)

Summary Risk-sensitive foraging theory predicts that predators which face starvation if there is a temporary shortfall in their food supply should choose feeding sites on the basis of variation in as well as mean expected reward rate. For a given mean reward rate they should choose high variance feeding sites (be risk-prone) if they are running below energy requirement, but low variance sites (be risk-averse) if they are running above.Common shrews presented with a choice between constant and variable feeding stations were more likely to visit the variable station when they were running below energy requirement and more likely to visit the constant station when they were running above. However, the tendency towards risk-aversion above requirement was greater than that towards risk-proneness below.When all shrews were considered together, the probability of visiting the variable station correlated negatively and continuously with intake relative to requirement.     

Central place foraging in wheatears (Oenanthe oenanthe L.): foraging itineraries when feeding nestlings

Summary Parent wheatears (Oenanthe oenanthe L.), foraging to meet their own needs and to provide food for a central place (CP, i.e. the nest), have to make decisions with respect to the configuration of foraging itineraries during round trips in the territory and to the directionality of their movements. These problems were studied in two pairs breeding in an agricultural area in central Swecen. All birds started a round trip by hunting from perches close to the CP and then moving to perches progressively further away in a roughly straight line until the first prey item was loaded. Loaded prey were either delivered singly (single prey loading: SPL) or with other prey (multiple prey loading: MPL). When the bird decided to return with several items to the CP, it abruptly changed the direction of its movements by making a left, right or backward turn and started to visit perches progressively nearer the CP, again following a roughly straight line. The decision to return continue loading was affected by the size of the prey as shown by the fact that prey carried singly was significantly larger than the first item in an MPL for all individuals. The distance to the CP also seemed to affect this decision as prey provisioned singly on average were collected significantly closer to the nest than the first item in an MPL by one pair. Both the size of prey loaded singly and load size of MPL increased with distance from the CP in one pair. The concentrated use of the territory in the other pair made any effect of distance difficult to detect. Great flexibility in foraging decisions was observed in cases when an individual, because of the size of the last prey captured, altered its decision to deliver an MPL and transported an SPL to the nest instead. On the basis of these results we propose a set of rules followed by predators such as wheatears when making decisions about delivering prey to a CP (Fig. 4).     

Jellyfish aggregations and leatherback turtle foraging patterns in a temperate coastal environment

Leatherback turtles (Dermochelys coriacea) are obligate predators of gelatinous zooplankton. However, the spatial relationship between predator and prey remains poorly understood beyond sporadic and localized reports. To examine how jellyfish (Phylum Cnidaria: Orders Semaeostomeae and Rhizostomeae) might drive the broad-scale distribution of this wide ranging species, we employed aerial surveys to map jellyfish throughout a temperate coastal shelf area bordering the northeast Atlantic. Previously unknown, consistent aggregations of Rhizostoma octopus extending over tens of square kilometers were identified in distinct coastal "hotspots" during consecutive years (2003-2005). Examination of retrospective sightings data (>50 yr) suggested that 22.5% of leatherback distribution could be explained by these hotspots, with the inference that these coastal features may be sufficiently consistent in space and time to drive long-term foraging associations.     

Origin of the antitropical distribution pattern in marine mussels (Mytilus spp.): routes and timing of transequatorial migration

Many marine species, including mussels in the Mytilus edulis species group (i.e. M. edulis L., M. galloprovincialis Lamarck, and M. trossulus Gould), have an antitropical distribution pattern, with closely related taxa occurring in high latitudes of the northern and southern hemispheres but being absent from the tropics. We tested four hypotheses to explain the timing and route of transequatorial migration by species with antitropical distributions. These hypotheses yield different predictions for the phylogenetic relationship of southern hemisphere taxa relative to their northern counter-parts. The three Mytilus species were used to test these hypotheses since they exhibit a typical antitropical distribution and representative taxa occur in both the Pacific and Atlantic. Two types of mtDNA lineages were found among populations of mussels collected from the southern hemisphere between 1988 and 1996; over 90% of the mtDNA lineages formed a distinct subclade which, on average, had 1.4% divergence from haplotypes found exclusively in northern Atlantic populations of M. galloprovincialis. These data indicate that southern hemisphere mussels arose from a migration event from the northern hemisphere during the Pleistocene via an Atlantic route. The remainder of the southern hemisphere lineages (<10%) were very closely related to mtDNA haplotypes found in both M. edulis and M. galloprovincialis in the northern hemisphere, suggesting a second, more recent migration to the southern hemisphere. There was no evidence that southern hemisphere mussels arose from Pacific populations of mussels. Received: 8 December 1998 / Accepted: 8 November 1999     

Risk-sensitive foraging behaviour of the round-eared elephant shrew (Macroscelides proboscideus)

We examined the risk-sensitive foraging behaviour of the round-eared elephant shrew by open-economy choice experiments, in which animals were deprived of food immediately prior to experiments but given food ad libitum afterwards, to test the energy budget rule. The energy budget rule states that if an animal's (daily) energy budget is negative it should behave in a risk-prone manner. A risk-prone elephant shrew should select food from a more variable rather than a constant feeding station, although both feeding stations yield the same average return. The choice of a variable station can indicate the degree to which an animal is an energy-shortfall minimizer. Elephant shrews running below energy requirement did not choose feeding stations in accordance with the rule. Under laboratory conditions, approximating either average summer or winter temperatures, elephant shrews showed risk-averse behaviour. A polycyclic activity profile, the ability to switch the diet, and greater than expected physiological control over energy balance, may favour a continuously foraging animal such that short-term energy deficits are minimized. We argue that, under these conditions, a risk-averse response to reward-size variance is expected, because an elephant-shrew may not reliably perceive those circumstances under which risk-prone behaviour should be adopted.     

A comparison of sound propagation and song frequency in temperate marsh and grassland habitats

Summary Attenuation of pure tones was measured in marsh and grassland habitat. At surface level, in grassland, the ground effect strongly attenuated frequencies below 2.0 kHz (Fig. 2). The ground effect was reduced by increasing source-receiver elevation. In marsh habitat the ground effect did not occur, and low frequencies were optimal for sound propagation (Fig. 3). As predicted from sound propagation tests, analysis of recorded songs of seven grassland and six marsh species indicated that minimum and emphasized frequency were significantly lower in songs of marsh birds. Maximum frequency did not differ between habitats (Table 4). Buzzing songs of yellow-headed blackbirds were broadcast and re-recorded at 1 and 50 m from the speaker in marsh and grassland habitats. Low frequency components attenuated more rapidly than higher frequency components in grassland, and the reverse occurred in marsh. Results suggested that the ground effect restricts the use of low frequencies by grassland birds. This evidence is consistent with the sound window hypothesis of Morton.     

Parameters of light-dependent photosynthesis for phytoplankton size fractions in temperate estuarine and coastal environments

Photosynthetic parameters for netplankton (>22 m) and nanoplankton (<22 m) varied over similar ranges but exhibited different seasonal and geographic patterns of variation. Nanoplankton a was relatively constant (0.06 mg C [mg Chl · h]^-1 [E m^-2 s^-1]^-1), but P _m ^B (mg C [mg Chl · d]^-1) was an exponential function of temperature independent of nutrient concentration and vertical stability in the euphotic zone. The temperature function gives a P _m ^B of 24 at 25°C for nanoplankton growing in an estuarine environment characterized by high nutrient concentrations and a shallow, stratified euphotic zone. Variations in netplankton a and P _m ^B were less predictable and were not correlated with temperature, nutrients or vertical stability. Chain forming diatoms with small cells were able to achieve high (0.10 to 0.15) and P _m ^B (20 to 24) that were 3 to 5 times higher than large-celled diatoms and dinoflagellates were able to achieve.     

Seasonal changes in foraging strategies of nesting blackbirds (Turdus merula L.)

Summary Blackbirds are usually multiple-prey loaders and forage mainly on the ground. We analysed the foraging behaviour of ten males in an urban park in Budapest during the breeding season from 1984 to 1986. At the end of April and in May blackbirds fed their nestlings mainly on earthworms (load type I). In this period the average scarching time and route were shorter, and the territories of the pairs tended to be smaller, than in June, when males in addition brought a great variety of invertebrates per load (load type II) to the young. The average dry weight and energy content of the two load types did not differ significantly. However, the average energy delivery rate (energy content/intervisiting time) and rate of energy gain (energy content/searching time) were higher when males collected earthworms, because intervisiting and searching times were shorter. The frequency distribution of searching times for load type I suggests that the encounter with earthworms was random. For load type II birds seemed to employ a fixed-mass foraging strategy. In June, the drier conditions reduced the availability of earthworms, and blackbirds extended their foraging areas.     

Respiratory metabolism of crabs from marine and estuarine habitats: an interspecific comparison

The gill-surface area and R-T (metabolic rate-temperature) response during aquatic and aerial respiration of three different species of crabs, the subtidal Scylla serrata, the intertidal Sesarma quadratum, and the supratidal Ocypoda platytarsis, were compared. Scylla serrata has the highest, Sesarma quadratum the second highest, and O.platytarsis the lowest gill-surface area. Oxygen-consumption values in water under different ambient partial pressures of oxygen indicate that all three species display equally efficient aquatic respiration despite variation in gill area and number. There is no coherent R-T response either inter- or intra-specifically. However, the ability to extract oxygen from the surrounding medium (respiratory efficiency) conditions the R-T response in the different species. The R-T trends in aquatic respiration of Scylla serrata and aerial respiration of O.platytarsis reveal a normal response to temperature. The proportion of aquaticoxygen uptake to total respiration in all sizes in both Scylla serrata and Sesarma quadratum remains more or less constant (between 88 and 92%), whereas in the supratidal O.platytarsis, this proportion decreases with increasing weight (from 88 to 64 %), indicating a progressive loss of gill respiration to total respiration. All three species are metabolically equally efficient; what one lacks in respiratory surface it compensates by respiratory efficiency.     

The loading effect in central place foraging wheatears (Oenanthe oenanthe L.)

Summary The behaviour of five adult wheatears (Oe. oenanthe) delivering prey (maggots and mealworms) from artificial patches to the nestlings was studied. The existence of a so-called loading effect was confirmed. Handling times for subsequent prey items increased with the number held in the beak, which led to a decrease in collecting rates with patch time and, in most cases, to positively accelerating loading functions when collecting times for prey items are plotted as a function of load size. All birds seemed to become more efficient at loading prey in the experimental patches during the course of the nestling period. Loading functions for maggots and mealworms were slightly different. When forcing the birds to visit several cups (only one item in each cup) and remove a layer of moss before reaching the prey (low-density patches), all birds took fewer prey and two of them stayed longer in them than in highdensity patches (one cup filled with prey items). This was due to an increase in search times with the number of prey held in the beak.From knowledge of the loading functions and travel times to the nest, it is possible to predict the optimal load sizes according to a mathematical solution of the delivery rate model of Orians and Pearson (1979). By transforming collecting and travel times to energy expenditures, it is also possible to derive predictions from an energy efficiency model (maximizing energy delivery per unit energy expended in a round-trip). The observed average load sizes did not differ significantly from those predicted by the delivery rate model, but they were significantly smaller in all cases than those predicted by the energy efficiency model. For birds feeding nestlings, it may be more important to sacrifice efficiency in energy expenditure in favour of greater delivery rates, thereby maximizing the growth rate of the young.     

Prey ecology and behaviour affect foraging strategies in the Great Cormorant

The fine link between a particular dive pattern and a specific prey item represents a challenging task in the analysis of marine predator–prey relationships. There is growing evidence that prey type affects diving seabirds’ foraging strategies, dive shapes and underwater activity costs. This study investigates whether a generalist diver, the Great Cormorant Phalacrocorax carbo, modifies the time budget allocated to prey-capture behaviour and breathing strategies (reactive vs. anticipatory) with respect to the prey type (pelagic vs. benthic). Video recordings of 91 Great Cormorants show how the ecology and behaviour of their main prey, Mullets (Mugilidae) and Flounders Platichthys flesus, affect dive/surface durations and the diving pattern. The demersal habit and the low mobility of Flounders leads to an easy access to prey with an anticipatory strategy. Moreover, the patchy distribution of this fish species increases prey-capture rates. Conversely, Mullets exploit the whole water column and are highly mobile, and this is reflected in the need of performing two sequential dives to capture a prey, both longer and likely more expensive, with a consequent switch of strategy from reactive in the searching phase to anticipatory breathing during prey-capture events. This study provides evidence that a generalist diver may switch between different foraging strategies, and it shows how each of them may be optimal under particular ecological conditions. These constraints influence the dynamics that operate within the marine food chains and have relevant implications in managing lagoon areas, including fish ponds.