Seasonal Decline Research Articles

Endoscopy as a novel method for assessing endoparasite burdens in free‐ranging European shags (Phalacrocorax aristotelis)

Abstract Parasites are considered to be a key driver of evolutionary processes in wild animal populations. However, assessing host‐endoparasite burdens non‐destructively is problematic. Collection of faecal samples can be difficult, and faecal egg counts may not always be a reliable indicator of infection intensity. Here we report on endoscopy as a method for assessing natural burdens of nematode parasites Contracaecum rudolphii Hartwich, 1964, in a wild seabird, the European shag (Phalacrocorax aristotelis, L.). We aimed to measure natural individual parasite burdens and repeatability of burdens over time, and to verify that treatment with ivermectin removed parasites. Endoscopy was rapid, averaging 6 min (n = 157), with no obvious adverse effects on behaviour or breeding success compared to non‐endoscoped birds. Nematode burdens in the oesophagus and proventriculus of conscious shags were counted and classified as absent, low, medium or high using a flexible gastroscope with a camera attachment that recorded video footage. Re‐assessment of worm burdens was highly accurate, with 94% of randomly selected videos (n = 50) giving identical categorical scores, and 70% of worm counts (n = 40) giving the same total or differing by only one worm. All birds were parasitised by C. rudolphii. Natural burdens were significantly higher in males and in late breeders. Individuals had highly repeatable categorical parasite scores over time with 65% of control birds sampled more than once (n = 17; mean interval between assessments= 10·8 days) showing no change in scores. However, although the rank ordering of birds based on categorical scores remained constant, more finely resolved quantification indicated a slight seasonal decline in worm counts within individuals. Treatment with ivermectin (4 mg kg−1 of bird weight) resulted in complete removal of parasites. There was some evidence of temporal declines in worm counts with lower doses of ivermectin, including a dose (0·7 mg kg−1) previously shown to impact chick survival and growth. Endoscopy has considerable potential for investigating individual variation and temporal changes in endoparasite burdens and drug efficacy. Applicability and limitations of this method for other host‐parasite systems are discussed.

Rates and consequences of relaying in little auks Alle alle breeding in the High Arctic an experimental study with egg removal

Most seabirds have a small clutch size. Thus, replacement of a clutch after loss can make important contributions to an individual’s lifetime reproductive success. However, in the condition of short polar summer, relaying propensity may be time‐constrained. In this study, we investigated rates and consequences of relaying in a small High Arctic seabird, the little auk Alle alle. We performed an experiment in which we removed the single egg from 20 nests of early‐laying breeders. We measured relaying rates, and compared chick body mass and breeding success between the experimental and control nests. Despite the narrow window of the Arctic summer and the closely synchronized breeding, 75% of females produced a replacement egg just 2.7% smaller in volume than the first egg. This indicates that in little auks, the demographic effects of disruptions to breeding attempts (by predators, adverse weather or human activity) may be mitigated to some extent by replacement clutches. However, peak body mass and fledging body mass were lower in the experimental than the control chicks. This effect was rather a consequence of late hatching – chicks from replacement clutches followed seasonal decline in peak body mass and fledging mass. Finally, breeding success and chick survival up to 20 d in the experimental nests were respectively 34 and 37% lower than in the control nests. Thus, the quality and post‐fledging survival of chicks from the replacement clutches were probably lower compared to the chicks hatched from the first‐laid eggs.

Dry-season changes in macroinvertebrate assemblages of highly seasonal rivers: responses to low flow, no flow and antecedent hydrology

Highly seasonal rivers can experience extended low flow, and often dry, periods. Macroinvertebrate and flow data were used to explore hypotheses on the effects of antecedent hydrology and the low-flow, dry-season period on macroinvertebrate assemblages in northern Australia. Composition differed between early and late dry seasons. Taxa were more sensitive to water quality and more rheophilous in the early dry season when their habitats were lotic than when habitats later became lentic. As flow magnitudes in the antecedent dry season and on the sampling day increased, the habitats became more oxygenated and, in turn, macroinvertebrate richness increased. Higher wet-season flow magnitudes, flow variability and rates of fall were correlated with lower richness in the following dry season. Alteration of the flow-disturbance regime that increases the likelihood of flow cessation in macroinvertebrate habitats, or extends the duration of the dry season beyond that previously experienced in these highly seasonal systems, may alter the resistance and resilience of assemblages such that the seasonal decline and recovery of biodiversity may no longer be so reliable. Given the projected increase in low-flow incidence in many regions of the world, future research needs to examine the effects of reduced flow, flow cessation and stream drying as multiple, interacting stressors on stream biota.

Effects of Temperature and Moisture Variables on Brown Rust Epidemics in Sugarcane

AbstractEpidemics of brown rust in sugarcane, caused by Puccinia melanocephala, vary in severity between seasons. Natural epidemics were studied to determine the effects of temperature and moisture variables on epidemic onset, severity and decline. Variables were monitored with disease severity in two cultivars, each grown at a different location in Louisiana. Maximum daily temperature was the variable most correlated with seasonal epidemic development and decline. Disease severity was high during 2009 and low during 2010. This contrast allowed evaluation of the effects of conducive and limiting environmental conditions on severity. Lower severity resulted from a combination of unfavourable temperature and leaf wetness conditions that delayed onset then reduced the rate of disease increase. An accumulation of 23–25 days with leaf wetness periods of at least 7 h after the daily minimum temperature exceeded 17°C preceded the onset of disease on young leaves in both severe and mild epidemics. Severe epidemics in both cultivars declined once maximum ambient daily temperature was 32°C or higher. Low and high limiting temperatures determined the initiation and decline of an epidemic, respectively, under Louisiana climatic conditions. The availability of leaf wetness was then an important determinant of disease severity during the epidemic.

Fitness Consequences of Timing of Migration and Breeding in Cormorants

In most bird species timing of breeding affects reproductive success whereby early breeding is favoured. In migratory species migration time, especially arrival at the breeding grounds, and breeding time are expected to be correlated. Consequently, migration time should also have fitness consequences. However, in contrast to breeding time, evidence for fitness consequences of migration time is much more limited. Climate change has been shown to negatively affect the synchrony between trophic levels thereby leading to directional selection on timing but again direct evidence in avian migration time is scarce. We here analysed fitness consequences of migration and breeding time in great cormorants and tested whether climate change has led to increased selection on timing using a long-term data set from a breeding colony on the island of Vorsø (Denmark). Reproductive success, measured as number of fledglings, correlated with breeding time and arrival time at the colony and declined during the season. This seasonal decline became steeper during the study period for both migration and breeding time and was positively correlated to winter/spring climate, i.e. selection was stronger after warmer winters/springs. However, the increasing selection pressure on timing seems to be unrelated to climate change as the climatic variables that were related to selection strength did not increase during the study period. There is indirect evidence that phenology or abundances of preferred prey species have changed which could have altered selection on timing of migration and breeding.

Epidemiology and fitness effects of wood mouse herpesvirus in a natural host population

Rodent gammaherpesviruses have become important models for understanding human herpesvirus diseases. In particular, interactions between murid herpesvirus 4 and Mus musculus (a non-natural host species) have been extensively studied under controlled laboratory conditions. However, several fundamental aspects of murine gammaherpesvirus biology are not well understood, including how these viruses are transmitted from host to host, and their impacts on host fitness under natural conditions. Here, we investigate the epidemiology of a gammaherpesvirus in free-living wood mice (Apodemus sylvaticus) and bank voles (Myodes glareolus) in a 2-year longitudinal study. Wood mouse herpesvirus (WMHV) was the only herpesvirus detected and occurred frequently in wood mice and also less commonly in bank voles. Strikingly, WMHV infection probability was highest in reproductively active, heavy male mice. Infection risk also showed a repeatable seasonal pattern, peaking in spring and declining through the summer. We show that this seasonal decline can be at least partly attributed to reduced recapture of WMHV-infected adults. These results suggest that male reproductive behaviours could provide an important natural route of transmission for these viruses. They also suggest that gammaherpesvirus infection may have significant detrimental effects in wild hosts, questioning the view that these viruses have limited impacts in natural, co-evolved host species.

Rainfall-Driven Amplification of Seasonal Acidification in Poorly Buffered Tropical Streams

Acidification in freshwater ecosystems has important ecological and biogeochemical effects. Temperate streams affected by anthropogenic acidification have been extensively studied, but our understanding of natural acidification in tropical streams has been constrained by the lack of long-term datasets. Here, we analyze 14 years of monthly observations from 13 sampling stations in eight tropical streams in lowland Costa Rica. Stream pH increased during the 4-month dry season and declined throughout the wet season. The magnitude of the seasonal pH decline was greatest following the driest dry seasons, including the historically large El Nino Southern Oscillation event in 1998 when pH values dropped below 4.0 in some streams. Dissolved CO2 accounts for the low baseline pH in the poorly buffered study streams, and we hypothesize that an influx of soil-derived CO2 via subsurface flow paths contributes to the observed seasonal pH declines. Our results show tight coupling between rainfall, terrestrial, and aquatic ecosystems in the tropics. Predicted decreases in dry season rainfall for the tropics may lead to an increased magnitude of seasonal acidification.

Speciated mercury at marine, coastal, and inland sites in New England – Part 1: Temporal variability

Abstract. A comprehensive analysis was conducted using long-term continuous measurements of gaseous elemental mercury (Hg0), reactive gaseous mercury (RGM), and particulate phase mercury (HgP) at coastal (Thompson Farm, denoted as TF), marine (Appledore Island, denoted as AI), and elevated inland (Pac Monadnock, denoted as PM) sites from the AIRMAP Observatories in southern New Hampshire, USA. Decreasing trends in background Hg0 were identified in the 7.5- and 5.5-yr records at TF and PM with decline rates of 3.3 parts per quadrillion by volume (ppqv) yr−1 and 6.3 ppqv yr−1, respectively. Common characteristics at these sites were the reproducible annual cycle of Hg0 with its maximum in winter-spring and minimum in fall, comprised of a positive trend in the warm season (spring – early fall) and a negative one in the cool season (late fall – winter). Year-to-year variability was observed in the warm season decline in Hg0 at TF varying from a minimum total (complete) seasonal loss of 43 ppqv in 2009 to a maximum of 92 ppqv in 2005, whereas variability remained small at AI and PM. The coastal site TF differed from the other two sites with its exceptionally low levels (as low as below 50 ppqv) in the nocturnal inversion layer possibly due to dissolution in dew water. Measurements of Hg0 at PM exhibited the smallest diurnal to annual variability among the three environments, where peak levels rarely exceeded 250 ppqv and the minimum was typically 100 ppqv. It should be noted that summertime diurnal patterns at TF and AI were opposite in phase indicating strong sink(s) for Hg0 during the day in the marine boundary layer, which was consistent with the hypothesis of Hg0 oxidation by halogen radicals there. Mixing ratios of RGM in the coastal and marine boundary layers reached annual maxima in spring and minima in fall, whereas at PM levels were generally below the limit of detection (LOD) except in spring. RGM levels at AI were higher than at TF and PM indicating a stronger source strength in the marine environment. Mixing ratios of HgP at AI and TF were close in magnitude to RGM levels and were mostly below 1 ppqv. Diurnal variation in HgP was barely discernible at TF and AI in spring and summer. Higher levels of HgP were observed during the day, while values that were smaller, but above the LOD, occurred at night.

Time of breeding and female condition affect chick-growth in the Chilean Swallow (Tachycineta meyeni)

We analysed growth of nestling Chilean Swallows (Tachycineta meyeni) over four consecutive breeding seasons and determined the factors that explain variation in growth rate, asymptotic body mass and length of the nestling period. As the breeding season advanced nestlings grew more slowly and attained lower asymptotic mass, but the length of the nestling period and nestling survival did not show any seasonal trend. Asymptotic body mass of nestlings increased with female body condition, which was negatively correlated with time of breeding, suggesting that a seasonal decline in asymptotic body mass could be the result of changes in environmental conditions or lower quality of females breeding later in the season. There was no relationship between average minimum ambient temperature and either growth rate or asymptotic mass during the nestlings first 10 days. However, short periods of poor weather and snowfall had a significant effect on nestling survival. Growth rate (0.43) was lower than that of other Tachycineta species nesting at similar latitudes in the northern hemisphere but similar to those of subtropical species of the genus, suggesting that the factors that have influenced the evolution of this life-history trait differ between Chilean Swallows and other species of Tachycineta.

Seasonality of Rotavirus in South Asia: A Meta-Analysis Approach Assessing Associations with Temperature, Precipitation, and Vegetation Index

BackgroundRotavirus infection causes a significant proportion of diarrhea in infants and young children worldwide leading to dehydration, hospitalization, and in some cases death. Rotavirus infection represents a significant burden of disease in developing countries, such as those in South Asia.MethodsWe conducted a meta-analysis to examine how patterns of rotavirus infection relate to temperature and precipitation in South Asia. Monthly rotavirus data were abstracted from 39 published epidemiological studies and related to monthly aggregated ambient temperature and cumulative precipitation for each study location using linear mixed-effects models. We also considered associations with vegetation index, gathered from remote sensing data. Finally, we assessed whether the relationship varied in tropical climates and humid mid-latitude climates.ResultsOverall, as well as in tropical and humid mid-latitude climates, low temperature and precipitation levels are significant predictors of an increased rate of rotaviral diarrhea. A 1°C decrease in monthly ambient temperature and a decrease of 10 mm in precipitation are associated with 1.3% and 0.3% increase above the annual level in rotavirus infections, respectively. When assessing lagged relationships, temperature and precipitation in the previous month remained significant predictors and the association with temperature was stronger in the tropical climate. The same association was seen for vegetation index; a seasonal decline of 0.1 units results in a 3.8% increase in rate of rotavirus.ConclusionsIn South Asia the highest rate of rotavirus was seen in the colder, drier months. Meteorological characteristics can be used to better focus and target public health prevention programs.

Wood anatomical responses of oak saplings exposed to air warming and soil drought

Water is vital for plant performance and survival. Its scarcity, induced by a seasonal decline in soil water availability or an increase of evaporative demand, can cause failures of the water conducting system. An adequate tolerance to drought and the ability to acclimate to changing hydraulic conditions are important features for the survival of long-lived woody plants in dry environments. In this study we examine secondary growth and xylem anatomical acclimation of 6 year old saplings of three European oak species (Quercus robur, Q.petraea, Q.pubescens) during the third consecutive year of exposure to soil drought and/or air warming (from 2007 to 2009). Intra-annual pinning was applied to mark the development of the formation of the annual ring 2009. Vessel size, parenchyma cell density and fiber size produced at different time of the growing season 2009 were compared between drought and warming treatments and species. Drought reduced secondary growth and induced changes in xylem structure while air warming had little effect on wood anatomical traits. Results indicate that drought-exposed saplings adjust their xylem structure to improve resistance and repairing abilities after cavitation. All species show a significant radial growth reduction, a reduced vessel size with diminished conductivity and a slightly increased density of parenchyma cells. Comparisons between species fostered our understanding of the relationship between the inter-specific xylem hydraulic plasticity and the ecological response to drought. The stronger changes observed for Q.robur and Q.petraea indicate a lower drought tolerance than Q.pubescens.

The phenological window for western spruce budworm: seasonal decline in resource quality

Western spruce budworm Choristoneura occidentalis Free. (Lepidoptera: Tortricidae) emerge in the spring before budburst and then face a rapidly deteriorating host quality each season. Measures of fitness, survival and fecundity, were made on cohorts of final‐instar spruce budworms deployed on host trees at several times during the season in four field locations in coastal and interior British Columbia, Canada. Survival and fecundity were strongly correlated throughout the season and varied as much as four‐fold from maxima at mid‐season to minima at the end of the season. Fitness values overall were greatest in the coastal compared with interior locations. Among interior locations, fitness was greatest at the highest elevation and least at the lowest elevation. Both cohort and sample‐based estimates of survival of wild, final‐instar budworms were relatively high in these outbreak populations. The influence of the phenological window and degree of synchrony with the host plant on herbivore abundance often depends on other processes affecting population rates of change.

Seasonality of carotenoid-based plumage coloration: modelling wavelength-specific change through spectral reconstruction

Plumage coloration has provided important model systems for research on signal expression. Whilst it had previously been assumed that moulting provided the only mechanism to change plumage coloration, recent studies have shown plumage colours to be seasonally dynamic, with implications both for the quantification of expression and for any signalling role. However, the mechanistic processes underlying such change remain uncertain. Here, we describe within-moult shifts in expression of a carotenoid-based colour trait – the yellow ventral plumage of the great tit Parus major – over a nine-month timespan. We report that plumage chromaticity (‘colour’) – but not achromaticity (‘brightness’) – exhibits a marked seasonal decline, independent of sex, age or body condition, and at a constant rate across twelve environmentally heterogeneous plots within our study site. To gain a greater understanding of the mechanisms underlying this change we employed a spectral reconstruction approach, that generates predicted spectra for any timepoint within the sampling period. By comparing spectra for both early and late in the moult we show that the seasonal decline in chromaticity is driven by both a marked reduction in ultraviolet (UV) reflectance and, to a lesser extent, loss of active carotenoid pigments. Thus, our study shows that seasonal loss of chromaticity in the great tit is driven by altered reflectance primarily in the UV section of the spectrum, a finding made possible by the use of spectral compartmentalisation and multi-parallel modelling to produce reconstructed spectra. Whether change in plumage coloration influences signal function will depend on the dynamics of the signalling system but it could clearly inflate patterns such as assortative mating and should be considered in studies of colour expression.

Climate change, breeding date and nestling diet: how temperature differentially affects seasonal changes in pied flycatcher diet depending on habitat variation

1. Climate warming has led to shifts in the seasonal timing of species. These shifts can differ across trophic levels, and as a result, predator phenology can get out of synchrony with prey phenology. This can have major consequences for predators such as population declines owing to low reproductive success. However, such trophic interactions are likely to differ between habitats, resulting in differential susceptibility of populations to increases in spring temperatures. A mismatch between breeding phenology and food abundance might be mitigated by dietary changes, but few studies have investigated this phenomenon. Here, we present data on nestling diets of nine different populations of pied flycatchers Ficedula hypoleuca, across their breeding range. This species has been shown to adjust its breeding phenology to local climate change, but sometimes insufficiently relative to the phenology of their presumed major prey: Lepidoptera larvae. In spring, such larvae have a pronounced peak in oak habitats, but to a much lesser extent in coniferous and other deciduous habitats. 2. We found strong seasonal declines in the proportions of caterpillars in the diet only for oak habitats, and not for the other forest types. The seasonal decline in oak habitats was most strongly observed in warmer years, indicating that potential mismatches were stronger in warmer years. However, in coniferous and other habitats, no such effect of spring temperature was found. 3. Chicks reached somewhat higher weights in broods provided with higher proportions of caterpillars, supporting the notion that caterpillars are an important food source and that the temporal match with the caterpillar peak may represent an important component of reproductive success. 4. We suggest that pied flycatchers breeding in oak habitats have greater need to adjust timing of breeding to rising spring temperatures, because of the strong seasonality in their food. Such between-habitat differences can have important consequences for population dynamics and should be taken into account in studies on phenotypic plasticity and adaptation to climate change.

Demography of the Heike firefly Luciola lateralis (Coleoptera: Lampyridae), a representative species of Japan’s traditional agricultural landscape

Populations of the Heike firefly, Luciola lateralis, a representative species of Japan’s traditional agricultural landscape (known as satoyama), have recently experienced rapid declines in many areas of Japan. Owing to the popularity of this firefly, many local communities have increased conservation efforts through the restoration of aquatic habitat complexes in satoyama. To provide fundamental parameters to predict population dynamics of the firefly, we conducted a mark–recapture study in restored paddy fields, and we estimated adult population parameters such as population size, survival, recruitment, sex ratio, and body size. We found that capture probability generally decreased as the season advanced, probably because of seasonal changes in detectability and/or firefly behavior. The daily survival rate of adults decreased over the season and may be related to a seasonal decline in adult body size. Adult population exhibited a highly male-biased sex ratio. Firefly abundance in the restored paddy fields doubled during the 4-year study period. Our analysis showed that adult detectability, recruitment, and survival rate are seasonally variable and could affect population size estimates obtained by a simple flash census. The mark–recapture technique can provide precise estimates of adult L. lateralis population characteristics and, thus, is a valuable method for predicting firefly populations and assessing the success of the restoration program.

The ostrich (Struthio camelus) ejaculate – effects of the method of collection, male age, month of the season, and daily frequency

1. Over three breeding seasons on a farm in Poland, semen was collected from 11 ostriches using the dummy and the teaser method to study the effects of the method of collection, male age, month in the breeding season, and daily collection frequency on ejaculate characteristics. 2. A total of 259 ejaculates were collected, with an average volume of 1·28 ± 0·6 (±SEM) ml. Sperm concentration was 3·34 ± 0·08 × 109/ml, the total number of spermatozoa 4·32 ± 0·22 × 109, and motility 4·56 ± 0·04. 3. There was no difference in ejaculates collected by the dummy and teaser methods, but the between-individual variation was considerable. Ejaculate characteristics increased with male age and varied between months, with little evidence for seasonal decline. Daily collections for 10 days did not affect sperm output. 4. The results open up avenues for further research on development of a viable protocol for artificial insemination in ostriches and efficient semen storage. 5. The between-male variation suggests that the ejaculate output could be maximized through selection.

Seasonal decline of offspring quality in the European starling Sturnus vulgaris: an immune challenge experiment

In seasonally fluctuating environments, timing of reproduction is a crucial determinant of fitness. Studies of birds show that late breeding attempts generally result in offspring of lower reproductive value, with lower recruitment and long-term survival prospects. Several proximate mechanisms, including a seasonal decline of immune system functioning, may lead to a seasonal decline of offspring fitness. We investigated seasonal variation in offspring quality by subjecting first- and second-brood chicks of a sexually size dimorphic species, the European starling Sturnus vulgaris, to an immune challenge with a bacterial endotoxin (LPS), and evaluated their growth and physiological response in terms of total plasma antioxidant capacity (TAC), concentration of reactive oxygen metabolites and hematocrit. LPS challenge did not affect chick growth or oxidative status. However, hematocrit of second-brood chicks was higher in LPS chicks compared to controls. Body mass halfway through the rearing period (days 8–9 post-hatching), TAC and hematocrit were lower among second- vs. first-brood chicks. Interestingly, sexual dimorphism in body mass at days 8–9 post-hatching markedly differed between broods, first-brood males being 4.7% and second-brood males 22.7% heavier than their sisters, respectively. Pre-fledging mortality occurred among second-brood chicks only and was strongly female-biased. Our findings suggest that starling chicks, even if in poor conditions, are little affected by a bacterial challenge, at least in the short-term. Moreover, our study indicates that sex differences in body size, possibly mediated by sex-specific maternal investment in egg size, may heavily impact on pre-fledging survival in a different way in the course of the breeding season, resulting in sex-specific seasonal decline of offspring fitness. Finally, we suggest that levels of circulating antioxidants should be regarded among the proximate causes of the association between timing of fledging and long-term survival in avian species.

Accelerated growth rates in late‐hatched Rhinoceros AukletCerorhinca monoceratachicks depend on food conditions and growth stage: an experimental approach

In some bird species, the survival of chicks hatching later in the season is lower than those hatched earlier due to increased risk of predation and a seasonal decline in feeding conditions. To reduce these risks, it might be advantageous for late‐hatched chicks to grow faster and hence fledge at younger age. In this experimental study, the growth rates of early‐ and late‐hatched Rhinoceros AukletCerorhinca monoceratachicks were compared under average and poor food supplies in captivity. Controlling for potentially confounding effects of chick mass at 10 days old, chick age and nest‐chamber temperature, late‐hatched chicks had higher wing growth rate than early‐hatched chicks before attaining the minimum wing length required for fledgling under both average and poor food supplies. After attaining the minimum wing length, however, late‐hatched chicks had a lower fledging mass, indicating a potential cost that could diminish the early advantage of fast wing growth.

Parental Investment Theory and Nest Defence by Imperial Shags: Effects of Offspring Number, Offspring Age, Laying Date and Parent Sex

AbstractNest defence is a common form of parental care employed by birds to improve the survival of their offspring. Theory predicts that parents should adjust their nest defence according to the value of the brood at stake, defending more intensively broods with high survival and reproductive prospects. We evaluated the influence of offspring number, offspring age, laying date and parent sex on nest‐defence intensity (NDI) of the Imperial Shag Phalacrocorax atriceps, a sexually dimorphic seabird with seasonal decline in offspring survival and very limited renesting potential. We also evaluated whether NDI was correlated within pairs and whether NDI of both members of the pair was correlated with incubation and breeding success. To elicit defensive behaviour, we simulated predation attempts using a Kelp Gull Larus dominicanus model. As predicted by theory, NDI was positively correlated with the number of offspring in the nest and offspring age. NDI during chick rearing was higher than that at early and late incubation, while no differences were found between incubation stages. Contrary to our prediction, we did not find differences in NDI according to laying date. NDI for males was higher than females, while NDI was also positively correlated within pairs. NDI was not statistically related to incubation or breeding success. These results suggest that other factors, such as laying date or parental quality and age, play a much larger role in determining the outcome and productivity of a nesting attempt. Our results provide partial support for parental investment theory; while parental defence increased with brood value according to offspring number and age, parental defence was not related to laying date, a factor strongly affecting offspring survival and recruitment probabilities in this species.

Seasonal Decline of Investment in Egg Production with Increasing Food Abundance on the Great Cormorant in a Riverfront Colony

Abstract To understand seasonal declines of investment in egg production, we studied the relationships among laying date, clutch, and egg sizes in a Great Cormorant Phalacrocorax carbo population. Seasonal changes in the amount and number of fish caught per casting net were used as an index of food abundance for Great Cormorants. Clutch size and egg volume were positively correlated and decreased as the breeding season progressed. Clutch volume (i.e. total egg volume within clutches) was explained by clutch size and laying date. However, the amount and number of fish caught per casting net increased through the season. We conclude that the observed seasonal decline in egg production investment was not a result of declining food abundance during the breeding season in Great Cormorant. The amount of egg investment appeared to depend on the fishing ability of the individual rather than fish availability during the breeding season. Early in the breeding season, only those individuals with strong fishing abili...