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.     

Temperature, food and mate limitation of copepod reproductive rates: separating the effects of multiple hypotheses

Temperature, food and the availability of mates may all limitrates of egg production is freshwater diaptomids. We have usedthe reproductive phases of Diaptomus paliidus to develop anindex of food limitation (f) and an index of mate limitation(M), each of which responds independently of temperature. Theresponse of the food limitation index to high and low temperaturesand high and low food concentrations was examined in a two-wayfactorial experiment in the laboratory. The index was highlyresponsive to a change in food concentration, stable duringa change in temperature alone at the high food level and responsiveto the synergistic interaction of food and temperature effectswhen both factors were limiting. Laboratory data indicate thatthe f index should not be biased by mate limitation unless maledensities are 6.7 x 10^–l0 males 1^–1 or less. Themate limitation index was highly responsive to mate availability,and ranged from a low of 9–19 in the presence of abundantmales to 100 in tbe absence of males. Applications of the fand M indices to a natural population of D. paliidus over 6weeks indicated that both food and mate availability were limitingthe reproductive rates of the copepods during the sampling period. ¹Present address: Institute of Animal Resource Ecology, Universityof British Columbia, Vancouver, Canada V6T 1W5     

GROWTH AND DEMOGRAPHY OF POPULATIONS OF BIOMPHALARIA PFEIFFERI (GASTROPODA, PLANORBIDAE) IN THE LABORATORY

The growth and demography of the freshwater snail Biomphalariapfeifferi, an intermediate host of the parasitic worm Schistosomamansoni in Eastern Zaire, were investigated in the laboratoryunder two conditions of crowding. Both individual and populationgrowth were depressed by crowding. Crowding reduced growth rateand all the variables and parameters related to reproduction(fecundity, instantaneous birth rate, net reproduction rate),but did not affect maximum size, survivorship, instantaneousdeath rate and life expectancy. The mechanism involved in theseeffects is likely to be exploitation competition for especiallyprofitable food items such as diatoms. (Received 10 November 1986;     

Daily body mass regulation in dominance-structured coal tit (Parus ater) flocks in response to variable food access: a laboratory study

In a dominance-structured flock, social status may determinepriority of access to food. Birds of low social status mayperceive present and future access to food as less predictable,and so have a higher risk of starvation, than birds of highsocial rank. Theoretical models predict that subordinate birdsshould carry larger fat reserves and incur higher mass-dependentcosts than dominants. However, empirical tests of the assumptionsof these models are still scarce and controversial. We investigatedthe effect of dominance rank on daily mass gain under conditionsof fluctuating food availability in a laboratory experimentusing four flocks of four coal tits (Parus ater) each. Thesame amount of food was delivered in two treatments, but inone treatment the food was offered at a constant rate betweendays (fixed treatment), while in the other treatment the dailyfood supply varied in an unpredictable sequence between days(variable treatment). All birds showed greater variance inbody mass in the variable treatment than in the fixed treatment.Body mass within birds showed the same variability at dawn thanat dusk in the fixed treatment, but less variability at dawnthan at dusk in the variable treatment. This may be a mechanismto reduce the immediate risk of starvation at the beginningof the day, when fat reserves are at their lowest and the aggressionbetween flock members when feeding highest. Subordinate birdswere excluded from the feeders by dominants more often in theearly morning than in the rest of the day, and they showedmore variability in daily mass gain and body mass at dawn thandominant birds. These results support the hypothesis that subordinatebirds have a reduced probability of surviving when food availabilitychanges unexpectedly compared to dominants.     

HUNGER OVERCOMES FEAR IN NASSARIUS FESTIVUS, A SCAVENGING GASTROPOD ON HONG KONG SHORES

On protected intertidal sand flats, virtually worldwide, nassariidsnails scavenge carrion. Typically, they lie in repose beneaththe sand surface but emerge to feed when chemoreceptors detectcarrion. Distance from the potential food influences whetherthey move towards it and, if they do, whether they obtain ameal before it is consumed by others. This study shows that the rate at which Nassarius festivus movestoward food bait is similar for starved and well-fed individuals.That is, when individuals decide to feed, the rate at whichthey move toward food is always approximately the same. Thisstudy exposes another facet of nassariid nutrition related tothe degree of hunger. Recently fed individuals, i.e., thosewhich have fed for < 14 days, will cease feeding and departpalatable food if crushed conspecifics are added to it. Individualswhich have not fed for > 14 days tend to remain feeding,despite the addition of crushed conspecifics. Another studyhas shown that N. festivus consumes sufficient energy from afed-to-satiation meal to sustain it in repose for 20 days.These experiments clarify the observation. Between 14 to 20days since its last meal, N. festivus will feed when the possibilityarises, despite potentially adverse consequences. That is, hungerovercomes the fear of being consumed at a feeding assemblagebecause, without feeding, the alternative may be death by starvation.Thus, a nassariid makes decisions about whether or not to feed.The fear of predation causes N. festivus to abandon a meal ifthe previous one was obtained within a fortnight but, thereafter,it will risk predation to avoid starvation. (Received 14 March 1994; accepted 7 July 1994)     

Paleozoic shell-Boring Annelids and their Trace Fossils

Polychaete shell-borings are widespread but not often commonin Paleozoic strata of most continents, occurring in pelecypod,gastropod, cephalopod, coral, stromatoporoid, crinoid, brachiopod,ectoproct, and calcareous algal skeletons of Ordovician to Permianage. Four genera are known: Vermiforichnus (Spionidae), Ordovicianto Permian; Caulostrepsis (Spionidae) , Devonian to Oligocene;Myzoslomites (Myzostomidae), Ordovician to Jurassic: and possiblyConchotrema, Devonian to Permian. The Silurian Citonia siphois a worm-boring, not a calcareous worm tube, referable to Vermiforichnus. An exceedingly well preserved Devonian fossilized polychaete,Vermiforafacta rollhisi, with complex setigerous parapodia,dorsal cirri, peristomial cirri, prostomium, and tentacle-likepalps, occurs entombed in an agglutinated tube-lined worm-boring(Vermiforichnus) . It may have been a shell-borer consideringits many similarities to the spionid, Polydora, which apparentlyecologically replaced it. Like Polydora, Vermiforafacta probably filtered food and sedimentfrom sea water and constructed agglutinated tubes which linedits borings. In bivalves, borings parallel valves and are usuallyperpendicular to commissures; in massive skeletons, they radiateoutward and laterally. Distorted later growth and "blisters"indicate that the host was often alive. Commensal to pathogenicparasitism existed and worms benefited from their hosts's feedingcurrents and protective shells. One polychaete nestler is reported.Life positions of hosts are sometimes indicated.     

Nutmeg mannikins (<Emphasis Type="Italic">Lonchura punctulata</Emphasis>) reduce their feeding rates in response to simulated competition

Group feeding animals experience a number of competitive foraging costs that may result in a lowered feeding rate. It is important to distinguish between reductions in feeding rates that are caused by reduced food availability and physical interactions among foragers from those caused by the mere presence of foraging companions that may be self-imposed in order to obtain some benefit of group membership. Starlings (Sturnus vulgaris) reduce their feeding rates when in the company of simulated competitors located in an adjacent cage that cannot affect the food availability or interact with the forager. In the present study, we investigate whether the presence of simulated competitors in another species of passerine, nutmeg mannikins (Lonchura punctulata), can result in self-imposed reductions in feeding rates. When feeding in the company of simulated competitors, mannikins spent more non-foraging time near them, fed more slowly, reduced travel times between patches, reduced their scanning time and pecked more slowly. These results provide evidence that simulated competitors induce a reduction in pecking rate: behavioural interference. These self-imposed responses to competitors may have resulted from attempts to remain close to the non-feeding companions. Such self-imposed reductions in feeding rates may be a widespread yet generally unrecognised foraging cost to group feeding individuals.     

The nutritional ecology of the ctenophore Bolinopsis vitrea: comparisons with Mnemiopsis mccradyi from the same region

Bolinopsis vitrea is a warm water lobate ctenophore which doesnot overlap in its distribution with Mnemiopsis mccradyi incontiguous waters. We examined its feeding ecology on a seriesof cruises. B. virrea ingested increasingly more prey at higherfood concentrations (2–100 prey l^–1) but feedingeffort (clearance rate) decreased with increasing food availability.On a dry weight basis, smaller tentaculate Bolinopsis ingestedseveral times more than larger lobates, but based on carbonweight, specific ingestion was fairly uniform over the entiresize range investigated (6–60 mm total length). Bolinopsiscollected during the daytime in the Bahamas rarely had morethan three prey items in their guts. These results and laboratorymeasurements of digestion times (av. = 1.9 h) allowed computationof daily rations, which could not account for the metabolicrequirement as measured on the same cruises. Results of feedingexperiments, however, implied that prey densities in excessof 11^–1 were sufficient to sustain a growing populationof Bolinopsis. Prey concentrations about an order of magnitudehigher were required for M. mccradyi based on similar experiments.These results were in general agreement with observed densitiesand distributions of ctenophores and their zooplankton preyin the Bahamas and coastal South Florida.     

From breeding pairs to fox towns: the social organisation of arctic fox populations with stable and fluctuating availability of food

Food availability can impact group formation in Carnivora. Specifically, it has been suggested that temporal variation in food availability may allow a breeding pair to tolerate additional adults in their territory at times when food abundance is high. We investigate group occurrence and intraspecific tolerance during breeding in a socially flexible canid, the arctic fox (Vulpes lagopus). We compare Iceland and Sweden where resource conditions differ considerably. A breeding pair was the most common social unit in both populations, but as predicted, groups were more frequent where food abundance varied substantially between years (Sweden: 6 %) than where food availability was stable (Iceland: ≤2 %). Within Sweden, supplemental feeding increased group occurrence from 6 to 21 %, but there was no effect of natural variation in lemming (Lemmus lemmus) availability since group formation was rare also at lemming highs. Thus, additional factors appeared to influence the trade-off between intraspecific territoriality and tolerance. We report two cases where related females showed enduring social relationships with good-neighbour strategies. Related females also engaged in alloparental behaviour in a ‘fox town’ with 31 foxes (4 adults, 3 litters). In contrast, when unrelated foxes bred close to each other, they moved or split their litters during summer, presumably because of territorial conflict. We suggest that fluctuating food availability is linked to group formation in this Arctic carnivore, but also when food availability increases, additional factors such as relatedness, alloparental benefits, competition and predator defence appear necessary to explain group formation.     

Effects of Daphnia longispina on rotifer populations in a natural environment: relative importance of food limitation and interference competition

To test the relative importance of exploitative and interferencecompetition from Daphnia limiting rotifer populations in Bermejalesreservoir, birth rates of rotifers, food abundance, zooplanktonbiomass and size distribution of Daphnia were analysed fromhigh frequency sampling at two depths. A depletion in food abundanceby Daphnia feeding should be reflected in the birth rates ofrotifers susceptible to exploitative competition, while interferenceshould produce direct negative relationships between the numberof individuals of the interacting species. Results show thatsignificant relationships between food (chlorophyll-a) and birthrates of rotifers are found when the ratio chlorophyll-a/herbivorouszooplankton biomass is used as a measure of food availability.In our study, Anuraeopsis fissa was only affected through exploitativecompetition by Daphnia, while only Synchaeta oblonga showedevidence of being suppressed through interference competitionby this cladoceran.     

Endocrine Bases of Spatial and Temporal Opportunism in Arctic-Breeding Birds

The primary objectives of this paper are to define, explorethe environmental factors favoring, and discuss hypotheses concerningthe endocrine bases of two important arctic breeding strategiesthat we call spatial and temporal opportunism. We identify severalspecies that display spatial opportunism in the Arctic, andone that displays temporal opportunism. In spatial opportunism,breeding may be highly seasonal but the locality where individualsbreed may change from year to year as a result of unpredictablespatial distribution of food, nest site availability, or otherfactors such as predator abundance. We suggest that flexibilityof the transition from migration to settlement distinguishesspatial opportunists from site-faithful migrants. Thus far,data are available for only two hypotheses regarding the endocrinebasis of this flexibility. Circulating patterns of testosterone(associated with territory establishment) and corticosterone(associated with migratory activity) appear not to be involvedin the regulation of spatial opportunism in white-crowned sparrows(Zonotrichia leucophrys gambelii), but more detailed study isnecessary. In temporal opportunism (that may occur simultaneouslywith spatial opportunism), temporally variable food availabilityapparently selects for the capacity to initiate reproductionacross a wide portion of the year. Tonic activity of the hypothalamo-pituitary-gonad(HPG) axis could provide a mechanism to minimize the delay betweendiscovery of abundant food and onset of nesting in any season.However, the evidence for one arctic temporal opportunist, thewhite-winged crossbill (Loxia leucoptera), indicates that theHPG axis is not tonically active, but probably switches off(i.e., becomes photorefractory) in autumn, as for other birdsbreeding at these latitudes. Opportunistic breeding very earlyin the year (e.g., March near Fairbanks, Alaska) is associatedwith increased luteinizing hormone secretion, probably in responseto a combination of abundant food and social stimuli, afterrefractoriness dissipates. We have taken the first step of identifyingspatial and temporal opportunism as important phenomena in theArctic, and discussing hypotheses related to endocrine mechanisms.Future research should identify specific environmental cuesinvolved, and elucidate the neuroendocrine and endocrine mechanismsunderlying these two reproductive strategies.