Genetic and morphological similarity of two California gull populations with different life history traits

California Gulls nesting at Mono Lake (ML) have a modal clutch size of two eggs, while at Great Salt Lake (GSL) the mode is three. We used starch gel electrophoresis to determine whether the populations are genically different. Average individual heterozygosity at ML averages 2.67% and at GSL 2.95%, both slightly below the average for other birds. The average number of alleles per locus is 1.2 for both sites. Variation between sites is insignificant, with a genetic distance of 0.001 and an F_ST of 0.0044. Therefore, the clutch size differences are without genic correlates. In fact, at the loci studied, the two samples are indistinguishable from two random samples drawn from a panmictic unit. Five morphological characteristics also exhibited little between-site difference, in agreement with the electrophoretic analysis. A marked historical reduction (bottleneck) in the Mono Lake population seems not to have affected levels of genetic variation. We conclude that the clutch size differences are either recently evolved, and determined by loci not surveyed by us, or they are environmentally induced. At the population level, these gulls are similar to passerines in terms of heterozygosity and among population differentiation.     

Geographic variation in calcium and clutch size

For over a half century numerous hypotheses have surfaced aimed at explaining a key life history trait, the evolution of clutch size in birds. A principal goal has been to explain why clutch size generally increases with latitude both within species and among closely related species. Most hypotheses have stressed food limitation, predation, or seasonality. I present a novel hypothesis to explain geographic variation: a limitation of calcium resulting from broad scale variation in this element. Because the storage capacity of avian medullary bone is limited or nonexistent, during egg formation, female birds must intake supplemental calcium. Yet calcium and other exchangeable bases are much rarer in tropical soils. I briefly review the abundant experimental and observational evidence supporting how calcium limitation affects clutch size and other life history traits, and I present a series of predictions (and apparent support for them), stemming from the calcium variation and limitation hypothesis. The balance of evidence suggests that variability in availability of environmental calcium plays a proximate and ultimate role in the evolution of clutch size. Although this hypothesis is not mutually exclusive with others, it highlights another factor that needs to be considered in studies of the geographic variation in clutch size.     

Conservation of salt lakes

Salt lakes have a variety of important uses and values, including especially both economic and scientific ones. These uses and values have been and are increasingly subject to degradation from a variety of impacts: diversion of inflows, pollution, agricultural practices, and introduction of exotic species are among the more important. Recognition of these impacts upon salt lakes has led to some international and national measures for their conservation, but considerably more effort in this direction is needed. Against this background, Mono Lake, California, USA, and the Aral Sea, central Asia, are discussed as two localities which bring into sharp focus the various matters discussed in the paper. Finally, attention is drawn to the need to conserve the Akrotiri Salt Lake, Cyprus.     

Pre-incubation feeding activities and energy budgets of Snow Geese: can food on the breeding grounds influence fecundity?

The potential contribution of early spring feeding conditions in the Arctic to clutch size variation was examined in a population of Lesser Snow Geese Anser caerulescens caerulescens. Behavioural observations were combined with energetic analyses of food material to derive an estimate of the energy budgets of pre-laying and laying females. Food intake of females between arrival on the breeding grounds and incubation was considerable; estimated energy gains in this period were in the same magnitude as the cost of one or several eggs. The pre-laying period spent on the breeding grounds can thus be energetically beneficial rather than costly. Accumulation of resources for reproduction in Snow Geese is a continual process including the breeding grounds, and nutrient limitation after arrival in the Arctic cannot sufficiently explain the environmental component of clutch size variation. The timing of migration and follicle development is such that clutch size decisions are sometimes made during the late stages of migration and some-times after arrival. In the latter case food conditions on the breeding grounds may greatly influence clutch size; in the former case they may still influence readjustments of clutch size after the initial decision. The universal negative correlation between clutch size and laying date in Snow Geese can be explained by negative fitness consequences of late hatching, which outweigh the benefits of delayed laying and further nutrient accumulation. Food shortage on the breeding grounds may sometimes be a secondary factor contributing to seasonal clutch size decline.     

Life-history and ecological correlates of geographic variation in egg and clutch mass among passerine species

Broad geographic patterns in egg and clutch mass are poorly described, and potential causes of variation remain largely unexamined. We describe interspecific variation in avian egg and clutch mass within and among diverse geographic regions and explore hypotheses related to allometry, clutch size, nest predation, adult mortality, and parental care as correlates and possible explanations of variation. We studied 74 species of Passeriformes at four latitudes on three continents: the north temperate United States, tropical Venezuela, subtropical Argentina, and south temperate South Africa. Egg and clutch mass increased with adult body mass in all locations, but differed among locations for the same body mass, demonstrating that egg and clutch mass have evolved to some extent independent of body mass among regions. A major portion of egg mass variation was explained by an inverse relationship with clutch size within and among regions, as predicted by life-history theory. However, clutch size did not explain all geographic differences in egg mass; eggs were smallest in South Africa despite small clutch sizes. These small eggs might be explained by high nest predation rates in South Africa; life-history theory predicts reduced reproductive effort under high risk of offspring mortality. This prediction was supported for clutch mass, which was inversely related to nest predation but not for egg mass. Nevertheless, clutch mass variation was not fully explained by nest predation, possibly reflecting interacting effects of adult mortality. Tests of the possible effects of nest predation on egg mass were compromised by limited power and by counterposing direct and indirect effects. Finally, components of parental investment, defined as effort per offspring, might be expected to positively coevolve. Indeed, egg mass, but not clutch mass, was greater in species that shared incubation by males and females compared with species in which only females incubate eggs. However, egg and clutch mass were not related to effort of parental care as measured by incubation attentiveness. Ecological and life-history correlates of egg and clutch mass variation found here follow from theory, but possible evolutionary causes deserve further study.     

Evolution of clutch size in cavity-excavating birds: the nest site limitation hypothesis revisited

There are two major competing hypotheses for variation in clutch size among cavity-nesting species. The nest site limitation hypothesis postulates that nesting opportunities are more limited for weak excavators, which consequently invest more in each breeding attempt by laying larger clutches. Alternatively, clutch size may be determined by diet; the clutch sizes of strong excavators may be smaller because they are able to specialize on a more seasonally stable prey. We built a conceptual model that integrated hypotheses for interspecific variation in clutch size and tested it with comparative data on life-history traits of woodpeckers (Picidae) and nuthatches (Sittidae). In most analyses, diet explained more variation in clutch size among species than did propensity to excavate. Migratory status was positively associated with clutch size but was difficult to distinguish from diet since resident species consumed more bark beetles (a prey available in winter) and had smaller clutches than migratory species. The literature suggests that cavities are not limited in natural, old-growth forests. Although our data do not rule out nest site limitation, we conclude that annual stability of food resources has a larger impact on the evolution of clutch sizes in excavators than does limitation of nest sites.     

Reversed sexual size dimorphism in raptors: evaluation of the hypotheses in kestrels breeding in a temporally changing environment

Reversed sexual size dimorphism (RSD, females larger than males) is commonly found in birds of prey. We used kestrels (Falco tinnunculus), breeding in western Finland in a temporally varying environment of 3-year vole cycles, to assess current hypotheses for the evolution and maintenance of RSD. Our 12-year data showed only weak correlations between parental size and breeding parameters (laying date, clutch size and the number of fledglings produced). The degree of RSD per se was unrelated to breeding success, contrary to the prediction of the female dominance hypothesis. Females with small males produced larger clutches in low-vole years, independently of laying date, which supports the small male (or its equivalent inter-sexual selection) hypothesis. Small females tended to have more fledglings, particularly in low-vole years, which is inconsistent with the hypotheses for an advantage of large female size (the starvation, intra-sexual selection, reproductive effort, and supplementary feeding hypotheses). As for males, smaller females may be more efficient hunters, the importance of which should be most pronounced under harsh breeding conditions. Our results suggest that the directional selection on a particular size in kestrels may be under contrasting selection pressures by the environment, and, at least in breeding females, the advantages of large size can actually be counterbalanced during harsh environmental conditions. Received: 7 May 1999 / Accepted: 20 January 2000     

Effects of parental survival on clutch size decisions in fluctuating environments

Whereas in constant environments parental survival has no effect on optimal clutch size in the absence of trade-offs between juvenile and parental survival, the situation is drastically different in fluctuating environments. We consider a model in which, with respect to reproduction, parents and offspring are equivalent at the start of the next breeding season. When generations are non-overlapping, the clutch size maximizing geometric mean surviving number of offspring is optimal among all pure clutch size strategies. We prove that, as parental survival increases relative to that of the offspring, the optimal clutch size converges to the arithmetic mean maximizing clutch size (the so-called ‘Lack clutch size’). We also give a numerical procedure for calculating optimal mixed strategies and we show that, as environmental variance increases and/or parental survival decreases, mixed rather than pure strategies become optimal. Furthermore, we explain how to estimate fitness from empirical data under the assumptions of our model. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

A theoretical investigation of the effect of latitude on avian life histories

Tropical birds lay smaller clutches than birds breeding in temperate regions and care for their young for longer. We develop a model in which birds choose when and how often to breed and their clutch size, depending on their foraging ability and the food availability. The food supply is density dependent. Seasonal environments necessarily have a high food peak in summer; in winter, food levels drop below those characteristic of constant environments. A bird that cannot balance its energy needs during a week dies of starvation. If adult predation is negligible, birds in low seasonal environments are constrained by low food during breeding seasons, whereas birds in high seasonal environments die during the winter. Low food seasonality selects for small clutch sizes, long parental care times, greater age at first breeding, and high juvenile survival. The inclusion of adult predation has no major effect on any life-history variables. However, increased nest predation reduces clutch size. The same trends with seasonality are also found in a version of the model that includes a condition variable. Our results show that seasonal changes in food supply are sufficient to explain the observed trends in clutch size, care times, and age at first breeding.     

Life-history variation of a neotropical thrush challenges food limitation theory

Since David Lack first proposed that birds rear as many young as they can nourish, food limitation has been accepted as the primary explanation for variation in clutch size and other life-history traits in birds. The importance of food limitation in life-history variation, however, was recently questioned on theoretical grounds. Here, we show that clutch size differences between two populations of a neotropical thrush were contrary to expectations under Lack's food limitation hypothesis. Larger clutch sizes were found in a population with higher nestling starvation rate (i.e. greater food limitation). We experimentally equalized clutches between populations to verify this difference in food limitation. Our experiment confirmed greater food limitation in the population with larger mean clutch size. In addition, incubation bout length and nestling growth rate were also contrary to predictions of food limitation theory. Our results demonstrate the inability of food limitation to explain differences in several life-history traits: clutch size, incubation behaviour, parental feeding rate and nestling growth rate. These life-history traits were better explained by inter-population differences in nest predation rates. Food limitation may be less important to life history evolution in birds than suggested by traditional theory.     

Differential maternal investment counteracts for late breeding in magpies Pica pica: an experimental study

Reproductive success in many avian populations declines throughout the breeding season. Two hypotheses have gained attention to explain such a decline: the "timing" hypothesis proposes that deteriorating food availability causes the decline in reproductive success (causal effect of breeding time), whereas the "quality" hypothesis proposes that individuals of lower phenotypic quality reproduce at the end of the breeding season, causing the correlation between breeding time and breeding success. We tested both of these hypotheses in a monogamous single breeder, the magpie Pica pica , by experimentally inducing some pairs to lay a replacement clutch, after removal of the first clutch. The first clutch was left in the nest of another magpie pair (matched by laying date and clutch size), and incubated and raised by these foster parents. In this way we obtained two clutches from the same magpie pair with full siblings raised in conditions of the first and second reproductive attempts. High quality pairs (with laying dates in the first half of the breeding season) reached similar breeding success in replacement clutches as compared to first clutches of the same female. In addition, experimental pairs reared significantly more offspring of similar quality in their replacement clutches as compared to late-season first clutches, thereby suggesting that late season first clutches were produced by pairs of lower phenotypic quality. Results indicate that high quality pairs trade-off clutch size for larger eggs in replacement clutches, which could help magpie pairs to partly compensate for poorer environmental conditions associated with a delayed breeding attempt.     

Effects of clutch sizes and incubation stage on nest desertion in the female Common Eider Somateria mollissima nesting in the high Arctic

In bird species, one of the trade-offs between reproduction and survival appears in the parental decision to desert the nest. Nest desertion is modulated by several factors including clutch size. However, the incubation stage at which predation occurs is also an important factor. In this study, we examined whether nest desertion was linked to initial clutch size, partial clutch predation (final clutch size) and the incubation stage at which it happened in a capital breeder: the female common eider (Somateria mollissima) nesting in the high Arctic. The study was performed in Kongsfjorden in 2002 on the western coast of the Svalbard Archipelago (78°55′N, 20°07′E). We observed that nest desertion was higher when the initial clutch size was small. Also, females deserted their nests more during the first third of incubation than later. Thus, as incubation proceeded, nest desertion was less likely to occur even after egg reduction. Our results pointed out that this parental decision in female eiders seemed to depend on initial clutch size and on the date into incubation of clutch reduction. Sophie Bourgeon and Francois Criscuolo contributed equally to the work     

Effects of imposed salinity gradients on dissimilatory arsenate reduction, sulfate reduction, and other microbial processes in sediments from two California soda lakes

Salinity effects on microbial community structure and on potential rates of arsenate reduction, arsenite oxidation, sulfate reduction, denitrification, and methanogenesis were examined in sediment slurries from two California soda lakes. We conducted experiments with Mono Lake and Searles Lake sediments over a wide range of salt concentrations (25 to 346 g liter(-1)). With the exception of sulfate reduction, rates of all processes demonstrated an inverse relationship to total salinity. However, each of these processes persisted at low but detectable rates at salt saturation. Denaturing gradient gel electrophoresis analysis of partial 16S rRNA genes amplified from As(V) reduction slurries revealed that distinct microbial populations grew at low (25 to 50 g liter(-1)), intermediate (100 to 200 g liter(-1)), and high (>300 g liter(-1)) salinity. At intermediate and high salinities, a close relative of a cultivated As-respiring halophile was present. These results suggest that organisms adapted to more dilute conditions can remain viable at high salinity and rapidly repopulate the lake during periods of rising lake level. In contrast to As reduction, sulfate reduction in Mono Lake slurries was undetectable at salt saturation. Furthermore, sulfate reduction was excluded from Searles Lake sediments at any salinity despite the presence of abundant sulfate. Sulfate reduction occurred in Searles Lake sediment slurries only following inoculation with Mono Lake sediment, indicating the absence of sulfate-reducing flora. Experiments with borate-amended Mono Lake slurries suggest that the notably high (0.46 molal) concentration of borate in the Searles Lake brine was responsible for the exclusion of sulfate reducers from that ecosystem.     

Variation and trade-off in reproductive output of the Andean frog<Emphasis Type="Italic"> Hyla labialis</Emphasis>

I analysed variation in number and size of eggs in association with variation in body size of Hyla labialis females from three altitudinally widely separated populations in the Colombian Andes. Females at 3,500 m were significantly larger and heavier than those at 2,700 m, which in turn were larger and heavier than those at 2,000 m. After accounting for interpopulational differences in body size, clutch size (i.e. number of eggs) and egg size differed significantly between populations, but clutch mass did not. At 3,500 m, large relative clutch mass tended to be associated with larger eggs, whereas at lower altitudes it was associated with relatively larger clutch size. Within each population, females showed a significant trade-off between clutch size and egg size, even though relative clutch mass varied substantially and a clutch rarely used up the carrying capacity of their body cavity. At 3,500 m, comparisons between the early and late breeding season within a year suggested that climate fluctuations may play a role in reproductive-output variation within a population. Laboratory-kept females in a high somatic condition after spawning laid larger clutches in the next breeding event, but the trade-off persisted despite ad libitum food supply. Comparisons between field and laboratory suggest that limits to reproductive output were due to physiological, but not morphological restrictions. Therefore, the trade-off is better explained by energy limitation for egg growth depending on clutch size, than by the body cavity limitation hypothesis.     

Time-dependent reproductive decisions in the blue tit

Many breeding attempts in birds do not result in any fledged young due to predation on eggs or young. Consequently, the influence of time constraints on reproductive decisions are integrated parts of the reproductive behaviour of birds breeding within short, seasonal climate zones. In this study, I mimicked nest predation by removing blue tit (Parus caeruleus) clutches shortly after completion. Around 75% of the removed clutches were followed by a repeat clutch. Females producing their first clutch early in the season and females with an early onset of incubation in the laying sequence (an indication of high parental or territory quality) were most likely to initiate a repeat clutch. A trade‐off between the benefits of a repeat clutch and survival likely stopped late females in bad condition from investing more in the current reproductive season. Females producing a repeat clutch laid fewer eggs, had an earlier onset of incubation in the laying sequence and produced larger eggs than they did when producing their original first clutch. Eggs produced after the onset of incubation were especially large in the repeat clutches. Since food availability was presumably higher when the female produced her repeat clutch compared with her first clutch, females made a strategical decision when reducing clutch size, whereas onset of incubation and egg size may have been energetically constrained when producing the first clutch. Females that produced a relatively large clutch, had a relatively early onset of incubation, and laid relatively large eggs in their first clutch also did so when producing a repeat clutch, indicating that some of the variation in breeding parameters are due to differences in parental or territory quality. Differences between years in the temperature‐dependent development rate of caterpillars seem to affect the time constraints on breeding. A year with a predicted early seasonal decline in caterpillars resulted in short intervals between removal and relaying, small clutches and an early onset of incubation.     

Nest-building activity and laying date influence female reproductive investment in magpies: an experimental study

Nest size or nest-building activity has recently been hypothesized to be a postmating sexually selected signal in monogamous birds: females may assess a male's parental quality and willingness to invest in reproduction by his participation in nest building. Females may thus adjust their reproductive effort (i.e. clutch size) not only to their own abilities but also to those of their mates. We investigated whether female magpies, Pica pica, use nest-building activity rather than nest size to adjust their reproductive effort during replacement breeding attempts. After we removed their first clutch, high-quality pairs that built a large nest for the first clutch were more capable of building a replacement nest and females adjusted their clutch size in relation to the time it took to build the nest rather than nest size. We also found support for the hypothesized trade-off between clutch size and egg size in magpies. In replacement clutches females decreased clutch size and increased egg volume, thereby probably improving the survival probability of their offspring in less favourable conditions.Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Broodmate aggression and life history variation in accipitrid birds of prey

Aggressive sibling competition for parental food resources is relatively infrequent in animals but highly prevalent and extreme among certain bird families, particularly accipitrid raptors (Accipitriformes). Intense broodmate aggression within this group is associated with a suite of traits including a large adult size, small broods, low provisioning rates, and slow development. In this study, we apply phylogenetic comparative analyses to assess the relative importance of several behavioral, morphological, life history, and ecological variables as predictors of the intensity of broodmate aggression in 65 species of accipitrid raptors. We show that intensity of aggression increases in species with lower parental effort (small clutch size and low provisioning rates), while size effects (adult body mass and length of nestling period) are unimportant. Intense aggression is more closely related to a slow life history pace (high adult survival coupled with a restrained parental effort), rather than a by‐product of allometry or food limitation. Consideration of several ecological variables affecting prey abundance and availability reveals that certain lifestyles (e.g., breeding in aseasonal habitats or hunting for more agile prey) may slow a species’ life history pace and favor the evolution of intense broodmate aggression.     

Energetic constraints on clutch size and time of breeding in temperate zone birds

Summary A model for the energy budget of a breeding pair of Great Tits is synthesized from published data. The model is used to investigate the relationship between temperature, clutch size, onset of laying, and male feeding on breeding success. Two periods of energetic stress are found, one at the end of the incubation period which is independent of the clutch size, and one at the end of the fledging period which does depend upon the clutch size. This energetic bottleneck theory is easily testable and is in agreement with existing data. The model also provides a consistent explanation of the effects of reduced male feeding, and changes in clutch size and nest box insulation on breeding success.     

Parental investment decision rules in tree swallows: parental defense, abandonment, and the so-called Concorde Fallacy

I conducted an experiment to detect the importance of past investmentand expected benefits to reproductive "decision making" by treeswallows (Tachycineta bicolor). Three experimental groups, balancedfor measurable indicators of parental quality, were created:one had its clutch reduced in size near the beginning of incubation,the second had its brood reduced near chickfledging, and thethird had no brood or clutch reduction. All experimental groupswere tested for differences in 19 measures of parental defensebehavior at the same stage in the nesting cycle by exposingthem on successive days to two different predators, a ferretand a rat snake, at the nest box. There were no detectable effectsof experimental treatment on parental defense; however, therewere differences in abandonment frequency related to the severityof experimental clutch reduction. These results suggest thattree swallows respond to differences in expected benefits in"deciding" whether or not to abandon a nesting attempt. Becausepast investment can affect the prospective costs of abandonmentand renesting, the most natural currency for comparing costsand benefits is prospective. Adopting such a currency for parental"decisions" lessens concern over the so-called Concorde Fallacy,but it highlights our relative ignorance of how past reproductiveeffort influences the costs of future reproduction. [Behav Ecol1991,2:133–142]