Effects of life history variation on size and growth in stream-dwelling Atlantic salmon

A large size variation amongst life histories for stream-dwelling Atlantic salmon Salmo salar was found and the relative effect of life histories on size varied over time. As early as December (age 0+ years), fish that later smolted at age 2+ years were significantly larger than fish that did not smolt at age 2+ years. In contrast, there were no mass differences at age 0+ years between fish that would mature or not at age 1+ years (October). The mass differences between smolts and non-smolts persisted until smolting, and differences between mature and immature fish first appeared in May (age 1+ years). Following September (age 1+ years), there was also a significant interaction between smolting and maturity. Previously mature and immature age 2+ year smolts were not significantly different in size, but immature age 2+ year non-smolts were much lighter than mature age 2+ year non-smolts. Based on mass differences, the apparent 'decision' to smolt occurred c . 5 months before (winter, age 0+ years) the decision to mature (late spring, age 1+ years). In addition to strong seasonal growth variation, sizes of freshwater Atlantic salmon were largely structured by the complex interaction between smolt-age and maturity.     

Environmental factors, regional body size distributions and spatial variation in body size of local avian assemblages

Aim To determine how well variation in median body size of avian assemblages is predicted by (1) the environmental models usually employed in analyses of Bergmann's rule and (2) random sampling from the regional body size frequency distribution. If body size frequency distributions of local assemblages represent a random sample of a regional frequency distribution, then geographical variation in body sizes of assemblages might be a consequence of the determinants of spatial variation in species richness rather than direct influences on body size per se. Location Southern Africa. Methods Median body masses (as a measure of body size) of avian assemblages were calculated for quarter‐degree grid cells across South Africa and Lesotho. The relationship between median body mass and four environmental variables (minimum and maximum monthly temperatures, precipitation and seasonality in the normalized difference vegetation index, as a measure of seasonality in productivity) was examined using general linear models first without taking spatial autocorrelation into account, and then accounting for it by fitting an exponential spatial covariance structure. Model fit was assessed using the Akaike information criterion and Akaike weights. At each species richness value, random assemblages were sampled by either drawing species randomly from the regional body mass frequency distribution, or drawing species from the regional body mass frequency distribution with a probability proportional to their geographical distribution in the area. The ability of randomizations to predict actual body masses was examined using two‐tailed Fisher exact tests. Results Seasonality in productivity was the only environmental variable that remained a significant predictor of body mass variation in spatially explicit models, though the positive relationship was weak. When species richness was included in the models it remained the only significant predictor of size variation. Randomizations predicted median body mass poorly at low species richness, but well at high richness. Main conclusions Environmental models that have previously been proposed explain little of the variation in body mass across avian assemblages in South Africa. However, much of the variation in the median mass of assemblages could be predicted by randomly drawing species from the regional body mass frequency distribution, particularly using randomizations in which all species were drawn from the regional body mass frequency distribution with equal probability and at high species richness values. This outcome emphasizes the need to consider null expectations in investigations of the geographical variation in body size together with the probable environmental mechanisms underlying spatial variation in average size. Moreover, it suggests that in the South African avifauna, spatial variation in the body sizes of assemblages may be determined indirectly by the factors that influence geographical variation in species richness.     

Maternal nutritional condition and genetic differentiation affect brood size and offspring body size in Nicrophorus

In most animal species, brood size and body size exhibit some variation within and between populations. This is also true for burying beetles (genus Nicrophorus), a group in which the body size of offspring depends critically on the number of offspring competing for food due to the discrete nature of resource used for larval nutrition (vertebrate carcasses). In one species, brood size and body size are correlated with population density, and appear to be phenotypically plastic. We investigated potential proximate causes of between-population variation in brood size and body size in two species, Nicrophorus vespilloides and Nicrophorus defodiens. Our first experiment supported the notion that brood size is phenotypically plastic, because it was affected by environmental variation in adult nutritional condition. We found that the pre-breeding nutritional status of female N. vespilloides affected the number of eggs they laid, the number of surviving larvae in their broods, and the body size of their offspring. We do not know whether this plasticity is adaptive because greater offspring body size confers an advantage in contests over breeding resources, or whether starved females are constrained to produce smaller clutches because they cannot fully compensate for their poor pre-breeding nutritional status by feeding from the carcass. Our second experiment documents that brood size, specifically the infanticidal brood-size adjustment behavior, has undergone genetic differentiation between two populations of N. defodiens. Even under identical breeding conditions with identical numbers of first-instar larvae, females descended from the two populations produced broods of different size with corresponding differences in offspring body size.     

Integrating life history and physiology to understand latitudinal size variation in a damselfly

Our understanding of latitudinal life history patterns may benefit by jointly considering age and mass at maturity and growth rate. Additional insight may be gained by exploring potential constraints through pushing growth rates to their maximum and scoring physiological cost‐related variables. Therefore, we reared animals of a univoltine Spanish and Belgian population and of a semivoltine Swedish population of the damselfly Enallagma cyathigerum (spanning a latitude gradient of ca 2350 km) in a common environment from the eggs until adult emergence and exposed them to a transient starvation period to induce compensatory growth. Besides age and mass at maturity and growth rate we also scored investment in energy storage (i.e. triglycerides) and immune function (i.e. total activity of phenoloxidase). At emergence, body mass was greater in Spain and Sweden and lower in Belgium, suggesting a genetic component for the U‐shaped latitudinal pattern that was found also in a previous study based on field‐collected adults. The mass difference between univoltine populations can be explained by the shorter development time in the Belgian population, and this despite a higher growth rate, a pattern consistent with undercompensating countergradient variation. In line with the assumed shorter growth seasons, Belgian and Swedish animals showed higher routine growth rates and compensatory growth after transient starvation. Despite a strong link with metabolic rates (as measured by oxygen consumption) populations with higher routine growth rates had no lower fat content and had higher immune function (i.e. immune function decreased from Sweden to Spain), which was unexpected. Rapid compensatory growth did, however, result in a lowered immune function. This may contribute to the absence of perfect compensating countergradient variation in the Belgian population and the lowest routine growth rates in the Spanish population. Our results underscore the importance of integrating key life historical with physiological traits for understanding latitudinal population differentiation.     

Relationships between body size and some life history parameters

Summary Patterns in life history phenomena may be demonstrated by examining wide ranges of body weight. Positive relationships exist between adult body size and the clutch size of poikilotherms, litter weight, neonate weight life span, maturation time and, for homeotherms at least, brood or gestation time. The complex of these factors reduces r _max in larger animals or, in more physiological terms, r _max is set by individual growth rate. Comparison of neonatal production with ingestion and assimilation suggests that larger mammals put proportionately less effort into reproduction. Declining parental investment and longer development times would result if neonatal weight is scaled allometrically to adult weight and neonatal growth rate to neonatal weight. Body size relations represent general ecological theries and therefore hold considerable promise in the development of predictive ecology.     

Correlates of species richness in mammals: body size, life history, and ecology

We present the most extensive examination to date of proposed correlates of species richness. We use rigorous phylogenetic comparative techniques, data for 1,692 mammal species in four clades, and multivariate statistics to test four hypotheses about species richness and compare the evidence for each. Overall, we find strong support for the life-history model of diversification. Species richness is significantly correlated with shorter gestation period in the carnivores and large litter size in marsupials. These traits and short interbirth intervals are also associated with species richness in a pooled analysis of all four clades. Additionally, we find some support for the abundance hypotheses in different clades of mammals: abundance correlates positively with species richness in primates but negatively in microchiropterans. Our analyses provide no evidence that mammalian species richness is associated with body size or degree of sexual dimorphism.     

How does environmental variation influence body mass,body size,and body condition? Roe deer as a case study

We tested the influence of population density and of drought intensity (measured as the Gaussen Index in spring and summer of the year of birth) on winter body mass, hind foot length, and body condition of roe deer fawns. Body mass decreased with increasing density and increased with increasing Gaussen Index in summer, in a similar way for both males and females. Hind foot length of males showed the same response. On the other hand, hind foot length of females decreased with increasing density only after dry summers, hence when environmental conditions were very harsh. Body condition was affected neither by density nor by drought intensity. Our results indicate that body mass and size are much better indicators of phenotypic quality than body condition in roe deer. The sex-specific responses of body size to environmental conditions could correspond to a differential allocation in favour of daughters by heavier than average roe deer mothers.     

The life history legacy of evolutionary body size change in carnivores

We estimate the body sizes of direct ancestors of extant carnivores, and examine selected aspects of life history as a function not only of species' current size, but also of recent changes in size. Carnivore species that have undergone marked recent evolutionary size change show life history characteristics typically associated with species closer to the ancestral body size. Thus, phyletic giants tend to mature earlier and have larger litters of smaller offspring at shorter intervals than do species of the same body size that are not phyletic giants. Phyletic dwarfs, by contrast, have slower life histories than nondwarf species of the same body size. We discuss two possible mechanisms for the legacy of recent size change: lag (in which life history variables cannot evolve as quickly as body size, leading to species having the 'wrong' life history for their body size) and body size optimization (in which life history and hence body size evolve in response to changes in energy availability); at present, we cannot distinguish between these alternatives. Our finding that recent body size changes help explain residual variation around life history allometries shows that a more dynamic view of character change enables comparative studies to make more precise predictions about species traits in the context of their evolutionary background.     

A concise review of geographic variation in adult body size in anadromous masu salmon, Oncorhynchus masou

Adult body size (size at maturity) is one of the key life history traits and is well known to sometimes correlate with latitude in anadromous salmonids. However, it is poorly understood whether geographic size patterns except for latitudinal trends occur or why such patterns have been shaped. The present paper briefly reviewed body size variation between anadromous returns of masu salmon Oncorhynchus masou in the Okhotsk group (10 populations along the Sea of Okhotsk coast), the Pacific group (2 populations along the Pacific Ocean coast) and the Sea of Japan group (24 populations along the Sea of Japan coast). The Okhotsk group was smaller than the Sea of Japan group. Although the statistical analysis detected no differences among the remaining combinations, the Okhotsk group was possibly smaller than the other groups because the size of the Pacific group seemed to be within range of the Sea of Japan group but tended to be larger than that of the Okhotsk group. Future research should first test whether size at maturity genetically differs between the Okhotsk group and the other two groups to explore further evolutionary factors shaping geographic size variation.     

Climate,body condition and spleen size in birds

Climatic conditions may impact on the body condition of animals and thereby affect their survival prospects. However, climate may also impact directly on the survival prospects of animals by affecting the size of immune defence organs that are used for defence against parasites. We used a large long-term database on body condition and size of the spleen in birds to test for immediate and delayed relationships between climatic conditions as indexed by the North Atlantic Oscillation (NAO) and body condition and spleen mass, respectively. Across 14 species of birds, spleen mass was significantly positively correlated with the NAO index, while the delayed effect of NAO on spleen mass was not significant. Spleen mass was positively related to body condition, but body condition did not depend significantly on NAO or delayed NAO effects. Bird species with a strong positive effect of NAO on spleen mass tended to have small spleens for their body size, while species with a strong negative effect of NAO on spleen mass tended to have relatively large spleens. Since bird species with relatively large spleen have been shown to suffer more from the negative effects of parasites, we can infer that the effects of climate as indexed by NAO on the size of the spleen depends on the importance of parasite-mediated natural selection.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Temporal repeatability of relative standard metabolic rate in juvenile Atlantic salmon and its relation to life history variation

There was a strong correlation between the relative standard metabolic rate (rSMR) values of individual Atlantic salmon Salmo salar L. measured 5 and 22 weeks after first feeding in June and October respectively (Pearson's r =0·68, 26 d.f., P <0·001). However, this is a conservative estimate of repeatability as two separate regressions were used to calculate SMR in October due to the separation of the population into an upper modal group made up of early migrants and a lower modal group comprised of delayed migrants. SMR values in June and October were similar (paired t test, t =-0·85, 27 d.f., P >0·05) when expressed as the percentage deviation from those predicted for a fish of that size based on the body mass/SMR relations in June or October indicating that relative standard metabolic rates were stable over time when food was not limiting. rSMR status was maintained in 19 of the 28 fish (i.e. 68%) between the two measures. rSMR status was correlated with life history strategy: salmon fry with a high SMR in June were more likely to become smolts during the autumn than those with a low SMR.     

Scaling of growth and life history traits relative to body size, brain size, and metabolic rate in lorises and galagos (Lorisidae, primates)

A broad range of variation in body size, brain size, and metabolic rate occurs within the primate family Lorisidae, thus providing an opportunity to examine the relationship of these three parameters to variation in growth and life history traits. Data on adult body weight, gestation length, lactation length, age at first estrus, litter size, and growth parameters were collected from a captive colony of four lorisid species, Loris tardigradus, Nycticebus coucang, Galago crassicaudatus, and G. senegalensis. The data presented here constitute the most complete life history information available for these poorly understood prosimian species. Correlation and allometric analyses were performed to determine the relationships between variables. Among the lorisids studied, adult body weight, adult cranial capacity, and relative cranial capacity did not predict variation in life history traits. Adult basal metabolic rate predicted most of the variability in gestation length, lactation length, and growth parameters. Lorisines differ from similarly sized galagines in having lower basal metabolic rates, slower growth rates, slower developmental rates, and smaller litter sizes, resulting in reduced reproductive potential. This may be a consequence of lorisine adaptation to a diet of toxic insects. Metabolic rate and diet may be among the most important parameters to examine in any study of life history evolution.     

Relationships of latitude,altitude, and body size to litter size and mean annual production of offspring inPeromyscus

Litter size was positively correlated with latitude and altitude but not with production of offspring or with body size in Peromyscus. Increased litter size in northern populations probably reflects shortening the breeding season by climate and not a greater mortality rate at northern latitudes compared to southern latitudes. Production of offspring was negatively correlated with body size but not with latitude, altitude, or litter size. This is probably due to larger species living longer and taking longer to mature.     

Ecoimmunology and microbial ecology: Contributions to avian behavior,physiology, and life history

Bacteria have had a fundamental impact on vertebrate evolution not only by affecting the evolution of the immune system, but also generating complex interactions with behavior and physiology. Advances in molecular techniques have started to reveal the intricate ways in which bacteria and vertebrates have coevolved. Here, we focus on birds as an example system for understanding the fundamental impact bacteria have had on the evolution of avian immune defenses, behavior, physiology, reproduction and life histories. The avian egg has multiple characteristics that have evolved to enable effective defense against pathogenic attack. Microbial risk of pathogenic infection is hypothesized to vary with life stage, with early life risk being maximal at either hatching or fledging. For adult birds, microbial infection risk is also proposed to vary with habitat and life stage, with molt inducing a period of increased vulnerability. Bacteria not only play an important role in shaping the immune system as well as trade-offs with other physiological systems, but also for determining digestive efficiency and nutrient uptake. The relevance of avian microbiomes for avian ecology, physiology and behavior is highly topical and will likely impact on our understanding of avian welfare, conservation, captive breeding as well as for our understanding of the nature of host-microbe coevolution.     

Explaining adaptive shifts in body size on islands: a life history approach

The tendency for island populations to differ in body size from their mainland relatives has been well documented, but the mechanisms for these size changes remain speculative. Explanations have typically been based on ecological interactions that directly favor either an increase or decrease in body size. While it is clear that direct ecological interactions can influence body size, life history shifts present an alternative explanation for observed insular size trends across phylogenetic groups and trophic levels. Here I describe how decreased resource availability and reduced predation pressure, the same selective forces invoked by previous hypotheses, can operate to produce body size changes via the evolution of life history traits. This mechanism is more generally applicable than previous explanations and is consistent with much of the available data.     

Reproductive biology and its impact on body size: comparative analysis of mammalian, avian and dinosaurian reproduction

Janis and Carrano (1992) suggested that large dinosaurs might have faced a lower risk of extinction under ecological changes than similar-sized mammals because large dinosaurs had a higher potential reproductive output than similar-sized mammals (JC hypothesis). First, we tested the assumption underlying the JC hypothesis. We therefore analysed the potential reproductive output (reflected in clutch/litter size and annual offspring number) of extant terrestrial mammals and birds (as "dinosaur analogs") and of extinct dinosaurs. With the exception of rodents, the differences in the reproductive output of similar-sized birds and mammals proposed by Janis and Carrano (1992) existed even at the level of single orders. Fossil dinosaur clutches were larger than litters of similar-sized mammals, and dinosaur clutch sizes were comparable to those of similar-sized birds. Because the extinction risk of extant species often correlates with a low reproductive output, the latter difference suggests a lower risk of population extinction in dinosaurs than in mammals. Second, we present a very simple, mathematical model that demonstrates the advantage of a high reproductive output underlying the JC hypothesis. It predicts that a species with a high reproductive output that usually faces very high juvenile mortalities will benefit more strongly in terms of population size from reduced juvenile mortalities (e.g., resulting from a stochastic reduction in population size) than a species with a low reproductive output that usually comprises low juvenile mortalities. Based on our results, we suggest that reproductive strategy could have contributed to the evolution of the exceptional gigantism seen in dinosaurs that does not exist in extant terrestrial mammals. Large dinosaurs, e.g., the sauropods, may have easily sustained populations of very large-bodied species over evolutionary time.