Decline In Clutch Size Research Articles

Clutch Size, but Not Egg Volume, Increases with Rainfall in an Arid-dwelling Bird.

Understanding how animals maximize reproductive success in variable environments is important in determining how populations will respond to increasingly extreme weather events predicted in the face of changing climates. Although temperature is generally considered a key factor in reproductive decisions, rainfall is also an important predictor of prey availability in arid environments. Here, we test the impact of weather (i.e., rainfall and temperature) on female reproductive investment in an arid-dwelling bird (i.e., clutch size and egg volume) and tradeoffs between the two. We predicted that female chestnut-crowned babblers (Pomatostomus ruficeps), endemic to the arid region of Australia, would increase clutch size at the expense of egg volume in response to variation in rainfall and temperature. We found that over 14 breeding seasons, clutch size decreased with increasing temperature, but increased following more rain. Egg volume, on the other hand, became larger as temperatures increased and, although not related to the amount of rain, was related to the number of days since the last rainfall. Finally, egg size decreased as clutch size increased, indicating a tradeoff between the two reproductive parameters. Our results suggest that chestnut-crowned babblers breed reactively in response to variable environments. We expect that clutch size variation in response to rain may reflect the impact of rain on arthropod abundance, whereas the effect of temperature may represent an established decline in clutch size observed in other seasonal breeders. As the tradeoff between clutch size and egg volume was modest, and clutch sizes were modified to a greater extent than egg volumes in response to rainfall, we suggest selection is more likely to increase offspring number than quality, at least in favorable years. Our results support the idea that reproductive investment is variable in fluctuating environments, which has implications for species living in habitats experiencing more extreme and less predictable weather as the global climate changes.

Temporal constraints influence reproductive characteristics that are related to the pace‐of‐life continuum in geese

Among avian species, particularly those with altricial young, life‐history strategies are characterized by a ‘slow’ pace‐of‐life at lower latitudes, where relatively low annual investments in reproduction are traded‐off for increased survival. Evidence for this pattern in precocial species, however, is equivocal, and questions about ecological drivers of latitudinal variation in reproduction remain. To better understand spatial variation in pace‐of‐life and related reproductive traits across bird species and to test a hypothesis that might explain observed spatial patterns, we analysed breeding data from closely related Canada Geese Branta canadensis and Cackling Geese Branta hutchinsii, hereafter Canada‐type geese, comprising eight sub‐species from 16 sites across a broad gradient of latitude (32°N to 69°N) and season length. Unlike the pattern reported for many altricial species, Canada‐type geese did not show reduced annual fecundity at lower latitudes, and instead this reduced reproductive investment was at higher latitudes. For three of five reproductive traits, the relative influence of growing season length (GSL; an index of the time available to breed) was greater than that of latitude. A shorter GSL resulted in later nest initiations, shorter pre‐laying intervals and higher seasonal rates of clutch size decline. Our results suggest that these and other species of geese are able to circumvent nutritional and temporal constraints imposed by shorter GSL by storing and using nutrient reserves for egg laying and incubation. Relative flexibility in reproductive traits may permit Canada‐type geese to accommodate predicted increases in climatic variability, compared to species with more rigid reproductive strategies.

Weather and climate change drive annual variation of reproduction by an aerial insectivore

For many bird species, but especially aerial insectivores, reproduction depends on weather. Climate change is likely to intensify effects, but with uncertain consequences. We report 22 years of data on Eastern Kingbird (<em>Tyrannus tyrannus</em>) reproduction for two populations located in different hygric environments undergoing climate change; mesic central New York, USA, (NY; 12 years) and xeric southeastern Oregon, USA, (OR: 10 years). Laying date became earlier with increasing temperature in the 30-day period preceding laying in identical fashion at both sites, and in years of early laying, clutch size was larger, length of laying season increased, and failed initial nesting attempts were more often replaced. High temperature in the 10-days preceding mean laying date was associated with shorter laying seasons, while a site by 10-day temperature interaction reflected an increase and decrease of clutch size with increasing 10-day temperature in NY and OR, respectively. Seasonal rate of clutch size decline was higher when the laying season was short but also slowed in xeric OR when rain was abundant in the 10-days prior to mean laying date. Nest predation drove annual variation in young fledged/nest, but the latter also increased and decreased with increasing maximum temperature during the nestling phase in mesic NY and xeric OR, respectively. Potential effects of climate change on kingbird populations are thus high given the dependence of reproduction on weather, and climate change likely contributed to declines of kingbirds in OR. Declines of kingbirds in NY appear unrelated to warming climates because higher temperatures advanced laying dates and yielded greater nest productivity. However, length of laying season declined across years at both sites, and thus early season gains may be negated by poor conditions late in the season that may be causing shorter laying seasons. Further work is needed to identify causes for the latter changes.

Causes of seasonal decline in reproduction of the cooperatively‐breeding acorn woodpecker

Clutch size and reproductive success decline seasonally in a wide range of temperate avian taxa. Two competing hypotheses have been proposed to explain such declines: the ‘timing’ hypothesis, which states that conditions affecting reproduction decline intrinsically with date, and the ‘quality’ hypothesis, which proposes that high‐quality individuals or individuals in high‐quality situations breed earlier. We contrasted the relative importance of these two hypotheses using a long‐term dataset of the cooperatively‐breeding acorn woodpecker Melanerpes formicivorus in central coastal California (USA). This population exhibits an 11% seasonal decline in clutch size, a 60% seasonal decline in fledging success, and a 77% seasonal decline in fledgling overwinter survival. Clutch size tracks seasonal availability of flying insects, which are a likely ecological driver of the seasonal decline in reproduction and, because of the nonlinear relationship of flying insects with date, constitute a likely factor constraining even earlier nesting. By parsing lay date data into within‐female and between‐female components, we found that only the within‐female component had a statistically significant direct effect on clutch size, supporting the timing hypothesis. For both fledging success and overwinter survival, however, both within‐ and between‐female effects were highly significant, with between‐female effects (differences in female quality) being stronger than within‐female effects. These results suggest that timing per se is a key factor affecting the seasonal decline in clutch size, but that both differences in female quality and an intrinsic seasonal decline in conditions drive the even more dramatic seasonal declines in fledging success and overwinter survivorship of fledglings.

Higher rates of prebreeding condition gain positively impacts clutch size: A mechanistic test of the condition‐dependent individual optimization model

Abstract A combination of timing of and body condition (i.e., mass) at arrival on the breeding grounds interact to influence the optimal combination of the timing of reproduction and clutch size in migratory species. This relationship has been formalized by Rowe et al. in a condition‐dependent individual optimization model (American Naturalist, 1994, 143, 689‐722), which has been empirically tested and validated in avian species with a capital‐based breeding strategy. This model makes a key, but currently untested prediction; that variation in the rate of body condition gain will shift the optimal combination of laying date and clutch size. This prediction is essential because it implies that individuals can compensate for the challenges associated with late timing of arrival or poor body condition at arrival on the breeding grounds through adjustment of their life history investment decisions, in an attempt to maximize fitness. Using an 11‐year data set in arctic‐nesting common eiders (Somateria mollissima), quantification of fattening rates using plasma triglycerides (an energetic metabolite), and a path analysis approach, we test this prediction of this optimization model; controlling for arrival date and body condition, females that fatten more quickly will adjust the optimal combination of lay date and clutch size, in favour of a larger clutch size. As predicted, females fattening at higher rates initiated clutches earlier and produced larger clutch sizes, indicating that fattening rate is an important factor in addition to arrival date and body condition in predicting individual variation in reproductive investment. However, there was no direct effect of fattening rate on clutch size (i.e., birds laying on the same date had similar clutch sizes, independent of their fattening rate). Instead, fattening rate indirectly affected clutch size via earlier lay dates, thus not supporting the original predictions of the optimization model. Our results demonstrate that variation in the rate of condition gain allows individuals to shift flexibly along the seasonal decline in clutch size to presumably optimize the combination of laying date and clutch size. A plain language summary is available for this article.

Differences in size between first and replacement clutches match the seasonal decline in single clutches in Tree Swallows Tachycineta bicolor

The seasonal decline in clutch size in birds can be a response to the environmentally conditioned decrease in prospects for offspring or a consequence of a lower physical ability of late‐breeding females. To find out which of the explanations apply in Tree Swallows Tachycineta bicolor, we assessed whether replacement clutch size in this species is affected by an individual female's ability to lay a certain number of eggs. To do this, we measured the decline in clutch size as a function of laying date between first and replacement clutches in individuals that re‐nested following natural failure, and compared this with the rate of decline in clutch size with laying date for Tree Swallows that laid only a single clutch in that season. Additionally, we assessed whether the clutch size and the rate of its seasonal decline varied across years. We accounted for the truncated and under‐dispersed nature of clutch size data by using a Bayesian approach in the analysis. We found little variation in the rate of clutch size decline across years at our breeding site. Accounting for this seasonal decline in clutch size, mean clutch size was similar between single‐time breeding females and those that laid replacement clutches, implying that the number of eggs laid on the second attempt by female Tree Swallows is determined by laying date, rather than by the female's physical ability to produce a clutch of a certain size.

Differences in size and reproductive output of loggerhead turtles Caretta caretta nesting in the eastern Mediterranean Sea are linked to foraging site

Foraging success affects reproductive output in sea turtles, and is therefore an impor- tant factor to measure in order to understand population dynamics. During 2010 and 2011, we used satellite telemetry to track the at-sea behavior of 20 post-nesting loggerhead turtles Caretta caretta from Rethymno, Crete, Greece. Nineteen transmitters provided location and dive data throughout the turtles' migration towards their foraging grounds and the transition into foraging behavior. We identified 3 foraging regions: (1) 9 turtles migrated southwest towards the North African coast, with 8 concentrated in the region of the Gulf of Gabes, Tunisia; (2) 6 turtles migrated north towards the Aegean Sea; and (3) 4 turtles did not take long-distance migrations, instead remaining resident within the waters of Crete. Two fitness proxies were associated with differ- ences in post-nesting strategies. Turtles foraging in northern waters had significantly larger curved and straight carapace lengths and clutch sizes than turtles foraging near Crete or Africa. Those differences reflect the disparity in benthic prey abundances among the 3 regions. The Aegean had a higher abundance of macrobenthic fauna than the other 2 regions, and the Gulf of Gabes had an increased level of eutrophication. Deterioration of the aquatic resources in the Gulf of Gabes region may be a contributing factor in the observed steady decline in clutch size and total nests per season in 2 critical nesting beaches for loggerheads in Greece.

Nest-Site Selection and Reproductive Success of the Little GrebeTachybaptus ruficollisin Northeast Algeria

We studied the phenology, habitat selection, and interannual and seasonal changes in breeding performance of the little grebe Tachybaptus ruficollis in relation to the spatial structure of a shallow lake during three breeding seasons (2010–2012). Nests were mostly located in shallow waters, close to the shore (98.3 ± 37.9 cm), in vegetation dominated by Scirpus lacustris. The egg-laying period started at the end of March and extended for four months until the end of July. Overall, the mean clutch size was 4.7 ± 1.1 (N = 154) with a modal clutch of 5. There was a seasonal decline in both egg volume and clutch size. The overall nesting success was 60% and the chief causes of nest failure were predation (52%) and adverse weather (20%). Breeding outcome was influenced by water depth, nest size and year of breeding. Predation and nest flooding markedly varied between years. Larger nests had a higher nesting success than smaller ones. Likewise, nests located at greater water depth (further from the shore) succeeded better than ones located in shallower water. As the little grebe forages preferentially in shallow waters, these results suggest that optimal nesting locations may be the result of a trade-off between conflicting selection pressures such as foraging efficiency (better in shallow waters) and nest predation risk (greater in shallow waters).

Phytoplankton–zooplankton dynamics vary with nutrients: a microcosm study with the cyanobacterium Coleofasciculus chthonoplastes and cladoceran Moina micrura

Anti-grazer responses of phytoplankton have been widely documented to vary with external abiotic factors (e. g. nutrient status), suggesting an environment-dependence of phytoplankton-zooplankton dynamics. Here, we examined the trophic interaction between the cyanobacteria Coleofasciculus chthonoplastes and the cladoceran Moina micrura under three nutrient levels from oligotrophic to hypereutrophic conditions (0.3, 3.0 and 100.0 mu M PO43-; molar ratio NO3- : PO43- - 16 : 1) to determine the effect of nutrients on the stability of a phytoplankton-zooplankton system. In the absence of M. micrura, cell density and filament (single-filament) length of C. chthonoplastes were lower at the low nutrient level compared with those at the medium and high nutrient levels. When M. micrura was present, cell density, filament length, colony proportion and size (the mean number of cells per colony) of C. chthonoplastes were highest under the medium nutrient level, corresponding to the lowest amplitude of density fluctuations in M. micrura. Regression analyses revealed that low cell densities of C. chthonoplastes (due to its limited ability to form colonies and long filaments and hence a high consumption rate of M. micrura) at low and high nutrient levels limited the growth of M. micrura adult females, leading to a sharp decline in embryo length, clutch size and population density. In general, our results suggest that the stability of phytoplankton-zooplankton systems varies with nutrients because of the nutrient-dependent morphology in phytoplankton, and imply that extremely low or high nutrient load may destabilize phytoplankton-zooplankton systems.

Multiple aspects of plasticity in clutch size vary among populations of a globally distributed songbird

Plasticity in life-history characteristics can influence many ecological and evolutionary phenomena, including how invading organisms cope with novel conditions in new locations or how environmental change affects organisms in native locations. Variation in reaction norm attributes is a critical element to understanding plasticity in life history, yet we know relatively little about the ways in which reaction norms vary within and among populations. We amassed data on clutch size from marked females in eight populations of house sparrows (Passer domesticus) from North America and Europe. We exploited repeated measures of clutch size to assess both the extent of within-individual phenotypic plasticity and among-individual variation and to test alternative hypotheses about the underlying causes of reaction norm shape, particularly the decline in clutch size with date. Across all populations, females of this multibrooded species altered their clutch size with respect to date, attempt order, and the interaction of date and order, producing a reaction norm in multidimensional environmental space. The reaction norm fits that predicted by a model in which optimal clutch size is driven by a decline with date hatched in the ability of offspring to recruit. Our results do not fit those predicted for other proposed causes of a seasonal decline in clutch size. We also found significant differences between populations in response to date and the date by attempt order interaction. We tested the prediction that the relationship with date should be increasingly negative as breeding season becomes shorter but found steeper declines in clutch size with date in populations with longer seasons, contrary to the prediction. Populations also differed in the level of among-individual variation in reaction norm intercept, but we found no evidence of among-individual variation in reaction norm slope. We show that complex reaction norms in life-history characters exhibit within- and among-population variance. The nature of this variance is only partially consistent with current life-history theory and stimulates expansions of such theory to accommodate complexities in adaptive life history.

Body condition dynamics and the cost‐of‐delay hypothesis in a temperate‐breeding duck

Pre‐breeding body condition is an important determinant of reproductive success in birds, largely through its influence on timing of breeding. Declines in clutch size and recruitment probability within breeding seasons indicate a tradeoff may exist between the number of young (clutch size) and quality of young (recruitment probability). We explored local drivers of pre‐breeding body condition and tested predictions of the cost‐of‐delay hypothesis in female lesser scaup Aythya affinis. Yearling females arrived on the study site in lower body condition than older females, but both age classes had similar rates of body condition gain on the breeding grounds prior to nesting. Rates of body condition gain were positively influenced by water temperature, a proxy for wetland phenology. The effect of water level was asymptotic and interacted with water temperature, with greater rates of gain in body condition occurring in years with low water levels. Our results supported the predicted response of clutch size to the rate of pre‐breeding body condition gain. After accounting for lay date, clutch size was positively related to the rate of body condition gain (= 0.08 ± 0.039). We did not find support for a predicted interaction between rate of body condition gain and intra‐seasonal decline in clutch size (= 0.01 ± 0.01). Our results indicate that local conditions during pre‐breeding influence body condition dynamics in female lesser scaup, which subsequently affects clutch size.

Clock gene variation in Tachycineta swallows

Many animals use photoperiod cues to synchronize reproduction with environmental conditions and thereby improve their reproductive success. The circadian clock, which creates endogenous behavioral and physiological rhythms typically entrained to photoperiod, is well characterized at the molecular level. Recent work provided evidence for an association between Clock poly-Q length polymorphism and latitude and, within a population, an association with the date of laying and the length of the incubation period. Despite relatively high overall breeding synchrony, the timing of clutch initiation has a large impact on the fitness of swallows in the genus Tachycineta. We compared length polymorphism in the Clock poly-Q region among five populations from five different Tachycineta species that breed across a hemisphere-wide latitudinal gradient (Fig. 1). Clock poly-Q variation was not associated with latitude; however, there was an association between Clock poly-Q allele diversity and the degree of clutch size decline within breeding seasons. We did not find evidence for an association between Clock poly-Q variation and date of clutch initiation in for any of the five Tachycineta species, nor did we found a relationship between incubation duration and Clock genotype. Thus, there is no general association between latitude, breeding phenology, and Clock polymorphism in this clade of closely related birds.Figure 1Photos of Tachycineta swallows that were used in this study: A) T. bicolor from Ithaca, New York, B) T. leucorrhoa from Chascomús, Argentina, C) T. albilinea from Hill Bank, Belize, D) T. meyeni from Puerto Varas, Chile, and E) T. thalassina from Mono Lake, California, Photographers: B: Valentina Ferretti; A, C-E: David Winkler.

Long-term patterns in egg mortality during incubation and chick mortality during rearing in three species of tits in an English woodland

Capsule Mortality from egg-laying to fledging is species-dependent with Great Tits exhibiting greater losses of chicks, Coal Tits greater losses of eggs and Blue Tits exhibiting comparable rates of mortality of eggs and chicks. Aims To determine whether reproductive losses during incubation and chick-rearing were different and to ascertain whether there were different effects in three closely related songbirds. Methods Breeding success of Great Tits Parus major, Blue Tits Cyanistes caeruleus and Coal Tits Periparus ater at Treswell Wood, Nottinghamshire, England was examined, selecting only those nests which fledged at least one chick. Mortality during incubation (i.e. loss of eggs) or rearing (i.e. loss of chicks) was recorded per nest and annual means were calculated and used in the analysis, which examined relationships between year, first-egg date and species. Key factor analysis was used to investigate the relative effects of egg or chick losses on the total breeding mortality. Results Timing of reproduction showed a long-term trend to earlier nesting, but there was considerable variation from year to year in first-egg date. In this location there has been a long-term decline in clutch size in all three species. Previously unseen differences between these three species of tit are shown for mortality at different stages of the reproductive process. Coal Tits suffer the greatest losses through unhatched eggs, whereas Great Tit failures are greater during chick-rearing. Blue Tits exhibited problems in achieving success in both incubation and rearing. Conclusion Closely related tit species living in close proximity exhibit different patterns of reproductive mortality during breeding.

Breeding season food limitation drives population decline of the Little Owl Athene noctua in Denmark

Many farmland bird species have declined markedly in Europe in recent decades because of changes in agricultural practice. The specific causes vary and are poorly known for many species. The Little Owl, which feeds extensively on large invertebrates and is strongly associated with the agricultural landscape, has declined over most of northwestern Europe, including Denmark. We investigated the likely reasons for the population decline in Denmark by identifying patterns of local extinction (scale, 5 × 5 km2) and estimating demographic parameters affecting local survival, focusing on changes over time and their relationship to habitat characteristics. The distribution of the Little Owl in Denmark contracted considerably between 1972–74 and 1993–96. The extent of contraction varied across the country, and the only habitat correlate was that local disappearance was associated with smaller amounts of agricultural land. Analyses of ring recovery data suggested a constant annual adult survival rate of 61% from 1920 to 2002, which is similar to estimates from countries with stable populations. First‐year annual survival rates were much lower than values previously reported. From the 1970s into the 21st century, the mean number of fledglings declined from around 3 to &lt; 2 young per territory, but the decline in clutch size was considerably less. Reproductive parameters were higher closer to habitat types known to be important foraging habitats for Little Owls, and were also positively correlated with the amount of seasonally changing land cover (mostly farmland) within a 1‐km radius around nests as well as temperatures before and during the breeding season. Experimental food supplementation to breeding pairs increased the proportion of eggs that resulted in fledged young from 27 to 79%, supporting the hypothesis that the main proximate reason underlying the ongoing population decline is reduced productivity induced by energetic constraints after egg‐laying. Conservation efforts should target enhancement of food availability during the breeding season. Other farmland species dependent on large invertebrates are likely to share the problems that Little Owls face in modern agricultural landscapes.

Seasonal clutch size decline and individual variation in the timing of breeding are related to female body condition in a non-migratory species, the Upland Goose Chloephaga picta leucoptera

Several studies have shown that waterfowl females in good condition lay larger clutches and start egg-laying earlier in a breeding season. However, most of these studies lack corresponding data on individual condition and timing of breeding in different years. We analysed data on clutch size, egg size, hatching date and female body condition of a non-migratory species, the Upland Goose Chloephaga picta leucoptera, recorded in three seasons on New Island, Falkand Islands. We found a strong seasonal decline in both egg and clutch size. Egg size increased with clutch size. The mean egg volume and total clutch volume increased with female body condition and hatching date was earlier for females in higher body condition. Chicks hatched from nests with higher mean egg volume had higher early body condition. We also compared individual females between different study years and found that individual body condition showed a significant repeatability between years, whereas hatching date did not. While female aging did not systematically affect hatching date, females produced clutches that hatched earlier in years of higher body condition. We could thus show for the first time that the frequently reported negative relationship between female condition and hatching date also holds true on an individual basis. To our knowledge, this is the first study on seasonal clutch size decline in sheldgeese and one of a few in waterfowl comparing condition and timing of breeding within individuals between different years.

No experimental evidence for local competition in the nestling phase as a driving force for density‐dependent avian clutch size

1. In birds, local competition for food between pairs during the nestling phase may affect nestling growth and survival. A decrease in clutch size with an increase in breeding density could be an adaptive response to this competition. To investigate whether breeding density causally affected the clutch size of great tits (Parus major), we manipulated breeding density in three out of eight study plots by increasing nest-box densities. We expected clutch size in these plots to be reduced compared to that in control plots. 2. We analysed both the effects of variation in annual mean density (between-year comparisons) and experimental density (within-year comparison between plots) on clutch size variation, the occurrence of second broods and nestling growth. We examined within-female variation in clutch size to determine whether individual responses explain the variation over years. 3. Over the 11 years, population breeding density increased (from 0.33 to 0.50 pairs ha(-1)) while clutch size and the occurrence of second broods decreased (respectively from 10.0 to 8.5 eggs and from 0.39 to 0.05), consistent with a negative density-dependent effect for the whole population. Nestling growth showed a declining but nonsignificant trend over years. 4. The decline in population clutch size over years was primarily explained by changes occurring within individuals rather than selective disappearance of individuals laying large clutches. 5. Within years, breeding density differed significantly between manipulated plots (0.16 pairs ha(-1) vs. 0.77 pairs ha(-1)) but clutch size, occurrence of second broods and nestling growth were not affected by the experimental treatment, resulting in a discrepancy between the effects of experimental and annual variation in density on reproduction. 6. We discuss two hypotheses that could explain this discrepancy: (i) the decline in breeding performance over time was not due to density, but resulted from other, unknown factors. (ii) Density did cause the decline in breeding performance, but this was not due to local competition in the nestling phase. Instead, we suggest that competition acting in a different phase (e.g. before egg laying or after fledgling) was responsible for the density effect on clutch size among years.

Variation in the Relationship between Lay Date and Clutch Size in Three Cavity-Nesting Woodland Passerines

Clutch sizes of many single-brooded birds decrease as the breeding season progresses. This decrease is usually quantified using data from several years, an approach that would mask any annual variation. We used 15 years of data from 295 nestboxes occupied by Blue Tit, Great Tit and Pied Flycatcher to determine whether the strength of the relationship between lay date and clutch size is consistent, or whether it varies annually. Both lay date and year were strong predictors of clutch size for all species. However, Generalised Linear Modelling revealed an interaction between lay date and year in the prediction of clutch size, indicating that the strength of the relationship between lay date and clutch size varied between years. Multilevel modelling was used to establish proximate factors that may be responsible for annual variability. Factors affecting the relationship between lay date and clutch size were species-specific. For Blue Tits, seasonal decline in clutch size was steepest when the density of all cavity-nesting species was high (47% variance explained). For Great Tits, decline was steepest in “early” seasons, particularly when density was high (32% variance explained), and for Pied Flycatchers, decline was steepest in warmer years (33% variance explained). Thus annually variable factors appear to influence not only breeding phenology and clutch size individually, but also the relationship between these variables.

Latitudinal and seasonal patterns in clutch size of some single‐brooded British birds

Capsule Daylength, rather than latitude, was found to be an important determinant of variation in clutch size. Aims To describe the nature of spatial and temporal variation in clutch size, and explore the ecological correlates of these patterns. Methods We tested the prediction that seasonal declines in clutch size will be greater at higher latitudes. The environmental variables focused on were the influence of daylength, plant productivity, seasonality (i.e. Ashmole's hypothesis) and physiological mechanisms that relate clutch size to ambient temperature. We used data from 1980 to 2003 on spatial variation in clutch size across Britain for single‐brooded species, in which clutch size can be taken as a measure of annual reproductive investment. We included all seven species, from five families, with sufficient data in the British Trust for Ornithology's Nest Record Scheme. Results There are strong seasonal declines in clutch size but little evidence for latitudinal gradients in clutch size or in latitudinal gradients in the rate of seasonal clutch size decline. Of the environmental variables investigated, daylength had the most marked effect on clutch size; this was positive in diurnal species and negative in the one nocturnal species. Conclusions Although this study was confined to a relatively small latitudinal range of 8°, we found marked latitudinal gradients in a number of factors thought to drive spatial patterns in clutch size. Moreover, such variation is of sufficient magnitude to generate spatial patterns in other ecological variables in Britain. There is thus no simple explanation for the lack of a latitudinal gradient in clutch size. The results concerning daylength indicate that the time available for foraging is an important determinant of variation in clutch size.

Breeding Ecology of Montagu'S Harrier (Circus Pygargus) in Marshes of Eastern Poland: Importance of Aggregated Nesting

Breeding success in Montagu’s Harrier (Circus pygargus) was studied in peat bogs of eastern Poland during the period 1996–2002. Harriers nested only in the area covered with great fen sedge (Cladium mariscus). The studied Harriers laid from 1 to 6 eggs, on average 4.09 eggs (n = 118). With the progression of the season, no significant decline in clutch size was recorded. The highest values of clutch size were found for pairs nesting in five nest aggregations. Harriers fledged on average 2.49 young per nest and 1.46 young per breeding pair. Early clutches in semicolonies were larger than later ones. Aggregations of five and four nests were found optimal in terms of clutch size, total loss and fledging success. The results of the study agree with the hypothesis that reproductive success may be highest at intermediate colony size.

Orchestration of avian reproductive effort: an integration of the ultimate and proximate bases for flexibility in clutch size, incubation behaviour, and yolk androgen deposition

How much effort to expend in any one bout of reproduction is among the most important decisions made by an individual that breeds more than once. According to life-history theory, reproduction is costly, and individuals that invest too much in a given reproductive bout pay with reduced reproductive output in the future. Likewise, investing too little does not maximize reproductive potential. Because reproductive effort relative to output can vary with predictable and unpredictable challenges and opportunities, no single level of reproductive effort maximizes fitness. This leads to the prediction that individuals possessing behavioural mechanisms to buffer challenges and take advantage of opportunities would incur fitness benefits. Here, we review evidence in birds, primarily of altricial species, for the presence of at least two such mechanisms and evidence for and against the seasonal coordination of these mechanisms through seasonal changes in plasma concentrations of the pituitary hormone prolactin. First, the seasonal decline in clutch size of most bird species may partially offset a predictable seasonal decline in the reproductive value of offspring. Second, establishing a developmental sibling-hierarchy among offspring may hedge against unpredictable changes in resource availability and offspring viability or quality, and minimize energy expenditure in raising a brood. The hierarchy may be a product, in part, of the timing of incubation onset relative to clutch completion and the rate of yolk androgen deposition during the laying cycle. Because clutch size should influence the effects of both these traits on the developmental hierarchy, we predicted and describe evidence in some species that females adjust the timing of incubation onset and rate of yolk androgen deposition to match clutch size. Studies on domesticated precocial species reveal an inhibitory effect of the pituitary hormone prolactin on egg laying, suggesting a possible hormonal basis for the regulation of clutch size. Studies on the American kestrel (Falco sparverius) and other species suggest that the seasonal increase in plasma concentrations of prolactin may regulate both a seasonal advance in the timing of incubation onset and a seasonal increase in the rate of yolk androgen deposition. These observations, together with strong conceptual arguments published previously, raise the possibility that a single hormone, prolactin, functions as the basis of a common mechanism for the seasonal adjustment of reproductive effort. However, a role for prolactin in regulating clutch size in any species is not firmly established, and evidence from some species indicates that clutch size may not be coupled to the timing of incubation onset and rate of yolk androgen deposition. A dissociation between the regulation of clutch size and the regulation of incubation onset and yolk androgen deposition may enable an independent response to the predictable and unpredictable challenges and opportunities faced during reproduction.