Food supplementation in distinct Corsican oak habitats and the timing of egg laying by Blue Tits

ABSTRACT.   Resource constraints may impose physiological limitations on egg production and influence the timing of breeding in seasonally breeding birds. Food-supplementation experiments have demonstrated that food availability may influence the timing of egg laying, but the moderate response of birds in most studies suggests that the effect of food availability may be apparent only under certain ecological conditions. Experiments conducted in habitats that differ in ecological characteristics and natural availability of food sources are, therefore, needed to effectively examine the possible role of habitat-specific physiological constraints on the timing of breeding. We compared the response of Blue Tits ( Cyanistes caeruleus ) to supplemental feeding in four oak woodlands that differed in elevation, forest type, and food availability on the island of Corsica. We found that supplemental feeding advanced the date of egg laying (by about 1 week) at only one of four sites; a site dominated by evergreen holm oak where the availability of natural food was likely lower than at the other sites. Our results suggest that the response of Blue Tits to supplemental food depends on the natural level of resource abundance, but, in addition, that the effect of supplemental feeding on the timing of breeding appears to be small compared to the typical and, for our study sites, great (>1.5 mo) between-population variation in clutch initiation dates. Physiological limitations related to the acquisition of nutrients and energy may exert only a limited effect on the onset of breeding, and behavioral flexibility in the integration of fine-scale environmental signals (e.g., temperature and phenology) that predict future breeding conditions may be more important in explaining variation among populations in the timing of breeding.     

Comparative feeding ecology of Tropocyclops prasinus mexicanus (Copepoda, Cyclopoida)

The herbivorous and carnivorous feeding biology of Tropocyclopsprasinus mexicanus, an especially small cyclopoid copepod, wasstudied under in situ food conditions in three different lakesand under experimentally augmented food supplies. The mass-specificfood uptake is compared to that of two larger species—Cyclopskolensis and Cyclops vicinus. Under in situ food conditions,T.p.mexicanus depended to a larger extent on algae than invertebrateprey and showed lower mass-specific ingestion rates than thetwo larger species. Daily mass-specific uptake rates for algaeranged between 10 and 24% of its body mass versus 0.7–7%for invertebrate prey. The larger species C.vicinus and C.kolensisingested a total dry mass equivalent to 106% or up to 143% oftheir body mass with algae contributing 66 and 81%. However,under enriched food availability, T.p.mexicanus is able to ingesta biomass equivalent to its body mass, with an algae (54%) andprey (40%) portion similar to that of the larger species. Bodysize appears to be an important factor for the relative importanceof algal versus invertebrate prey for cyclopoid copepods.     

Proximate developmental mediators of sexual dimorphism in size: case studies from squamate reptiles

Sexual dimorphism in size (sexual size dimorphism; SSD) is nearlyubiquitous, but the relative importance of genetic versus environmentalcontrol of SSD is not known for most species. We investigatedproximate determinants of SSD in several species of squamatereptiles, including three species of Sceloporus lizards andthe diamond-backed rattlesnake (Crotalus atrox). In naturalpopulations of these species, SSD is caused by sexual differencesin age-specific growth. Males and females, however, may oftenshare similar potentials for growth: growth is strongly responsiveto the availability of food, and sexual differences in growthcan be greatly suppressed or completely absent under commonenvironmental conditions in the laboratory. Sexually divergentgrowth is expressed in natural environments because of inherentecological differences between males and females and becauseof potential epigenetic effects of sex-specific growth regulators.In field-active Sceloporus, sexual differences in growth rateare associated with sexual divergence in plasma testosterone.Experiments confirm that testosterone inhibits growth in speciesin which females are larger (for example, S. undulatus and S.virgatus) and stimulates growth in those in which males arelarger (for example, S. jarrovii). Interestingly, however, sexualdivergence in plasma testosterone is not accompanied by divergencein growth in S. jarrovii or in male-larger C. atrox in the laboratory.Furthermore, experimental effects of castration and testosteronereplacement on growth are not evident in captive S. jarrovii,possibly because growth effects of testosterone are supersededby an abundant, high-quality diet. In female-larger S. undulatus,growth may be traded-off against testosterone-induced reproductivecosts of activity. In male-larger species, costs of reproductionin terms of growth are suggested by supplemental feeding ofreproductive female C. atrox in their natural environment andby experimental manipulation of reproductive cost in femaleS. jarrovii. Growth costs of reproduction, however, do not contributesubstantially to the development of SSD in male-larger S. jarrovii.We conclude that the energetic costs of testosterone-induced,male reproductive behavior may contribute substantially to thedevelopment of SSD in some female-larger species. However, despitestrong evidence that reproductive investment exacts a substantialcost in growth, we do not support the reproductive cost hypothesisas a general explanation of SSD in male-larger species.     

Suspension feeding and kleptoparasitism within the genus Trichotropis (Gastropoda: Capulidae)

The marine gastropod Trichotropis cancellata is a facultativekleptoparasite, either suspension feeding or parasitically stealingfood from tube-dwelling polychaete worms. To determine whetherconclusions drawn from long-term studies in the San Juan Islands,Washington, about the relative importance of suspension feedingand kleptoparasitism can be applied generally to T. cancellataacross its biogeographic range, I expanded earlier studies toAlaska and British Columbia. Kleptoparasitism is pervasive throughoutthe range of T. cancellata, occurring with equal frequency throughoutthe areas studied. The average density and size of worm hostsare relatively constant across this range. Snail and worm densitiesare not significantly correlated at the larger scale of site(averaged over nearby sampling locations clustered around acity), but are correlated at the smaller local scale (withina sampling location). Larger worms do not support more snails.The abundance of uninfected worm hosts is usually not limiting,except potentially in some sampling locations in southwest Alaskawhere the use of a novel host (a holothurian) may be the resultof low densities of uninfected worms. Additionally, I documentedthe feeding behaviours of other trichotropid species in theseregions. Trichotropis conica is the second confirmed kleptoparasitewithin the genus Trichotropis, with kleptoparasitism as frequentin this species as in T. cancellata. Like T. cancellata, allsizes observed of T. conica are kleptoparasites. On the otherhand, Trichotropis insignis is an obligate suspension feeder.Further studies are needed to determine exactly how many timesthis behaviour has arisen and been lost in Capulidae and relatedfamilies. (Received 31 May 2007; accepted 19 October 2007)     

La reproduction de Temora stylifera: ses implications anatomiques en relation avec le facteur 'nutrition'

The culture of copepods under different feeding regimes hasallowed us to determine the effects of food availability onegg laying. In Temora stylifera spawning appears to be modifiedby the food resources of the adult. Conditions encountered duringdevelopment have little effect on spawning. The life span offemales is reduced by high food availability. In such casesthe sex organs are hypertrophied, the oviducts being distendedby the large oocytes. The ovary is little affected by changesin food availability, while the size of the oviducts variesin relation to feeding. The size of the genital system is reducedin older females. Variations in food concentration induce variationsof several days in the time required for egg laying. A rapidmaturation of the oocytes, combined with nutrition and the availabilityof food permits Temora stylifera to optimize egg laying.     

Begging in the absence of parents by nestling tree swallows

Begging by nestling passerine birds has become a model systemfor studies in animal communication. Although most beggingoccurs when parents arrive at the nest to feed (here called"primary begging"), it also occurs between feeding visits andimmediately after parents leave the nest. Begging in thesecontexts (here called "secondary begging") may have relativelylittle influence on the probability of receiving food, but could increase the overall cost of the signal and thus influence nestlingbegging strategies. The purpose of our study was to determinehow often tree swallow (Tachycineta bicolor) nestlings begin contexts other than to parents with food and to examinewhat factors influence the frequency of this begging. Secondarybegging ranged from 7% of measured begging responses at day2 to 30% by day 8 and was more frequent when the interval betweenparental feeding visits was relatively long and when the timeto respond to the arrival of parents with food was short. Increasesin both age and intervisit interval were associated with decreasesin nestling response times, suggesting that secondary beggingmay be related to the speed with which nestlings respond to stimuli. We discuss possible functions of secondary beggingand raise the possibility that it may, in fact, be an error.     

The benefits of stealing from a predator: foraging rates, predation risk, and intraspecific aggression in the kleptoparasitic spider Argyrodes antipodiana

In the trade-off between food and safety, the role of aggressiveintraspecific interactions has not been extensively examined.Here I present information on this system using a kleptoparasiticspider, Argyrodts antipodiana, and its host spider and potentialpredator, Eriophora pustulosa. A. antipodiana can feed eitherat a potentially dangerous site (the hub of its host's web withthe host), or at a relatively safe site (on food bundles aroundthe edge of the host's web). I found that A. antipodiana cangain food very quickly when feeding with the host, apparentlyby exploiting the host's ability to digest the prey. Thus A.antipodiana follows predictions based on foraging models inthat it accepts a higher predation risk at the hub because ofthe higher food payoff. A. antipodiana also aggressively competesfor access to more food. However, aggressive competition increasesthe predation risk from the host, especially at tile hub wherethe host is very close. Consequently, A. antipodiana modifiesits level of intraspecific aggressiveness in accordance withits position on the web: at the hub, where the cost of aggressionis high (due to predation risk), A. antipodiana reduces itsaggressiveness, but it is aggressive away from the hub whencompeting for food bundles. The ability of A. antipodiana tochange interaction intensity as a function of its position onthe web enables it to exploit a rich, but risky, food sourceand provides a new angle for examining food and safety tradeoffs in light of intraspecific competition for food     

Feeding behavior of the cirratulid Cirriformia filigera (Delle Chiaje, 1825) (Annelida: Polychaeta).

Observations of the feeding behavior of Cirriformia filigera (Delle Chiaje, 1825) (Annelida: Polychaeta) from the intertidal zone of S?o Francisco and Engenho D'água beaches (S?o Sebasti?o, State of S?o Paulo) were made in the laboratory. This species, like other cirratulids, is a deposit feeder, feeding mainly on sediment surface with the aid of its grooved and ciliated palps, which are used to capture food particles. The worm lies just beneath the substrate surface in a J-shaped tube. When feeding, it extends up to 4 palps over the sediment surface, capturing food particles which pass down the groove of each palp directly to the mouth. Only fine sand grains are ingested. The worm frequently extends 4 branchial filaments into the overlying water for aeration. When it moves with the prostomium sideways, it collects and transports sand grains that pass backwards along its ventral region until reaching the middle part of its body. Next, the parapodia and palps move the sand grains to the dorsal posterior end of the animal, covering this area with sand. Some sand grains are also ingested as the worm moves.     

FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL HELIX ASPERSA MULLER

Feeding of the land snail Helix aspersa (Müller) was observedat monthly intervals. Three natural populations in Galicia (NW-Spain)were studied. At two sites only a few plants constituted thebulk of the diet and in spring the snails' diet had the highestdiversity (H'). In the third population feeding and distributionof Helix aspersa (Müller) were observed in a small plotwith permanent patches of Urtica dioica. Nearly one half offeeding snails fed upon Urtica dioica. Most of the other observations wereon Mentha suaveolens, Ranunculus repens and Gramineae. The diversity ofthe snails' diet showed seasonal variation with the maximumin the autumn months. Comparison between the availability ofthe different plant species and their contribution to the snails'diet showed that the snails did not eat at random; Urtica dioicawas eaten much more than expected from its occurrence and grasseswere strongly under-represented in the snails' diet. Temporalchanges of availability were significantly correlated with the amountseaten in the case of Urtica, but not for the other food plants.The distribution of the snails in the plot was significantlycorrelated with that of Urtica. Chemical analyses of the foodplants revealed Urtica as the species with the higher protein,ash and calcium contents. The strong preference of Helix aspersafor Urtica dioica could be explained by the value of Urticaas food or by its suitability as habitat for the snails. The largestproportions of green material in the snails' diet occurred inthe spring and juveniles ate more green material than adultsin the three populations. (Received 16 March 1998; accepted 30 November 1998)     

Daily foraging routines and feeding effort of a small bird feeding on a predictable resource

According to theory, a small diurnal bird living in a predictable environment should have the highest feeding effort as late as possible in the day in order to minimize the time it carries large and costly reserves. The feeding effort should also decline with increasing food availability. We tested both these ideas with the lesser spotted woodpecker (Dendrocopos minor). For most of the year, this bird feeds on wood-living insects in dead tree branches. This food supply is likely to be highly predictable on a daily scale. Our results corroborated the theory. We found that the proportion of time spent actively feeding was lower in the mornings (before noon) than in the afternoons. We also found that woodpeckers spent less time feeding the higher their food availability However, for a given food availability they spent more time feeding in the afternoons. This supports the idea that feeding is less and other activities are more valuable in the mornings given a predictable food resource. This is the first demonstration of daily routines in small birds concordant with a predictable environment. In spring, males but not females reduced their feeding time. This difference between the sexes may be related to their sex-specific reproductive effort.