RESUMO
Life history theory attempts to explain why species differ in offspring number and quality, growth rate, and parental effort. I show that unappreciated interactions of these traits in response to age-related mortality risk challenge traditional perspectives and explain life history evolution in songbirds. Counter to a long-standing paradigm, tropical songbirds grow at similar overall rates to temperate species but grow wings relatively faster. These growth tactics are favored by predation risk, both in and after leaving the nest, and are facilitated by greater provisioning of individual offspring by parents. Increased provisioning of individual offspring depends on partitioning effort among fewer young because of constraints on effort from adult and nest mortality. These growth and provisioning responses to mortality risk finally explain the conundrum of small clutch sizes of tropical birds.
Assuntos
Tamanho da Ninhada , Aves Canoras/crescimento & desenvolvimento , Aves Canoras/fisiologia , Clima Tropical , Asas de Animais/crescimento & desenvolvimento , Animais , Arizona , Malásia , Mortalidade , Comportamento de Nidação , Comportamento Predatório , VenezuelaRESUMO
Growth and development rates may result from genetic programming of intrinsic processes that yield correlated rates between life stages. These intrinsic rates are thought to affect adult mortality probability and longevity. However, if proximate extrinsic factors (e.g., temperature, food) influence development rates differently between stages and yield low covariance between stages, then development rates may not explain adult mortality probability. We examined these issues based on study of 90 songbird species on four continents to capture the diverse life-history strategies observed across geographic space. The length of the embryonic period explained little variation (ca. 13%) in nestling periods and growth rates among species. This low covariance suggests that the relative importance of intrinsic and extrinsic influences on growth and development rates differs between stages. Consequently, nestling period durations and nestling growth rates were not related to annual adult mortality probability among diverse songbird species within or among sites. The absence of a clear effect of faster growth on adult mortality when examined in an evolutionary framework across species may indicate that species that evolve faster growth also evolve physiological mechanisms for ameliorating costs on adult mortality. Instead, adult mortality rates of species in the wild may be determined more strongly by extrinsic environmental causes.
Assuntos
Mortalidade , Aves Canoras/embriologia , Aves Canoras/crescimento & desenvolvimento , Animais , Arizona , Evolução Biológica , Embrião não Mamífero/fisiologia , Desenvolvimento Embrionário , Malásia , Comportamento de Nidação/fisiologia , Comportamento Predatório/fisiologia , Análise de Regressão , África do Sul , VenezuelaRESUMO
Predation can be an important agent of natural selection shaping parental care behaviours, and can also favour behavioural plasticity. Parent birds often decrease the rate that they visit the nest to provision offspring when perceived risk is high. Yet, the plasticity of such responses may differ among species as a function of either their relative risk of predation, or the mean rate of provisioning. Here, we report parental provisioning responses to experimental increases in the perceived risk of predation. We tested responses of 10 species of bird in north temperate Arizona and subtropical Argentina that differed in their ambient risk of predation. All species decreased provisioning rates in response to the nest predator but not to a control. However, provisioning rates decreased more in species that had greater ambient risk of predation on natural nests. These results support theoretical predictions that the extent of plasticity of a trait that is sensitive to nest predation risk should vary among species in accordance with predation risk.
Assuntos
Aves/fisiologia , Cadeia Alimentar , Comportamento de Nidação , Animais , Argentina , Arizona , Especificidade da EspécieRESUMO
Intrinsic processes are assumed to underlie life history expression and trade-offs, but extrinsic inputs are theorised to shift trait expression and mask trade-offs within species. Here, we explore application of this theory across species. We do this based on parentally induced embryo temperature as an extrinsic input, and mass-specific embryo metabolism as an intrinsic process, underlying embryonic development rate. We found that embryonic metabolism followed intrinsic allometry rules among 49 songbird species from temperate and tropical sites. Extrinsic inputs via parentally induced temperatures explained the majority of variation in development rates and masked a relationship with metabolism; metabolism explained a minor proportion of the variation in development rates among species, and only after accounting for temperature effects. We discuss evidence that temperature further obscures the expected interspecific trade-off between development rate and offspring quality. These results demonstrate the importance of considering extrinsic inputs to trait expression and trade-offs across species.
Assuntos
Desenvolvimento Embrionário , Aves Canoras/embriologia , Aves Canoras/metabolismo , Animais , Arizona , Temperatura Corporal , Clima , Embrião não Mamífero/metabolismo , Malásia , Comportamento de Nidação , Oxigênio/metabolismo , Análise de Regressão , Clima Tropical , VenezuelaRESUMO
Causes of interspecific variation in growth rates within and among geographic regions remain poorly understood. Passerine birds represent an intriguing case because differing theories yield the possibility of an antagonistic interaction between nest predation risk and food delivery rates on evolution of growth rates. We test this possibility among 64 Passerine species studied on three continents, including tropical and north and south temperate latitudes. Growth rates increased strongly with nestling predation rates within, but not between, sites. The importance of nest predation was further emphasized by revealing hidden allometric scaling effects. Nestling predation risk also was associated with reduced total feeding rates and per-nestling feeding rates within each site. Consequently, faster growth rates were associated with decreased per-nestling food delivery rates across species, both within and among regions. These relationships suggest that Passerines can evolve growth strategies in response to predation risk whereby food resources are not the primary limit on growth rate differences among species. In contrast, reaction norms of growth rate relative to brood size suggest that food may limit growth rates within species in temperate, but not tropical, regions. Results here provide new insight into evolution of growth strategies relative to predation risk and food within and among species.
Assuntos
Cadeia Alimentar , Comportamento de Nidação , Passeriformes/crescimento & desenvolvimento , Animais , Arizona , Evolução Biológica , Feminino , Masculino , Comportamento Materno , Passeriformes/fisiologia , Comportamento Paterno , África do Sul , Especificidade da Espécie , VenezuelaRESUMO
Long embryonic periods are assumed to reflect slower intrinsic development that are thought to trade off to allow enhanced physiological systems, such as immune function. Yet, the relatively rare studies of this trade-off in avian offspring have not found the expected trade-off. Theory and tests have not taken into account the strong extrinsic effects of temperature on embryonic periods of birds. Here, we show that length of the embryonic period did not explain variation in two measures of immune function when temperature was ignored, based on studies of 34 Passerine species in tropical Venezuela (23 species) and north temperate Arizona (11 species). Variation in immune function was explained when embryonic periods were corrected for average embryonic temperature, in order to better estimate intrinsic rates of development. Immune function of offspring trades off with intrinsic rates of embryonic development once the extrinsic effects of embryonic temperatures are taken into account.
Assuntos
Desenvolvimento Embrionário , Passeriformes/crescimento & desenvolvimento , Temperatura , Animais , Arizona , Passeriformes/imunologia , Fatores de Tempo , Clima Tropical , VenezuelaRESUMO
1. Latitudinal variation in clutch sizes of birds is a well described, but poorly understood pattern. Many hypotheses have been proposed, but few have been experimentally tested, and none have been universally accepted by researchers. 2. The nest size hypothesis posits that higher nest predation in the tropics favours selection for smaller nests and thereby constrains clutch size by shrinking available space for eggs and/or nestlings in the nest. We tested this hypothesis with an experiment in a tropical forest and a comparative study between temperate and tropical field sites. 3. Specifically, we tested if: (i) predation increased with nest size; (ii) tropical birds had smaller nests controlled for body size; and (iii) clutch size was explained by nest size controlled for body size. 4. Experimental swapping of nests of different sizes showed that nest predation increased with nest size in the tropical site. Moreover, nest predation rates were higher in species with larger nests in both sites. However, nest size, corrected for body mass and phylogeny, did not differ between sites and was not related to clutch size between sites. 5. Hence, nest predation can exert selection on nest size as predicted by the hypothesis. Nest size increased with adult body mass, such that adult size might indirectly influence reproductive success through effects on nest size and nest predation risk. Ultimately, however, selection from nest predation on nest size does not explain the smaller clutch sizes typical of the tropics.
Assuntos
Aves/fisiologia , Tamanho da Ninhada , Comportamento de Nidação/fisiologia , Animais , Arizona , Tamanho Corporal , Demografia , Ecossistema , VenezuelaRESUMO
Species with "slow" life history strategies (long life, low fecundity) are thought to produce high-quality offspring by investing in larger, but fewer, young. Larger eggs are indeed associated with fewer eggs across taxa and can yield higher-quality offspring. Tropical passerines appear to follow theory because they commonly exhibit slow life history strategies and produce larger, but fewer, eggs compared with northern species. Yet, I show here that relative egg mass (corrected for adult mass) varies extensively in the tropics and subtropics for the same clutch size, and this variation is unexplained. I propose a hypothesis to explain egg size variation both within the tropics and between latitudes: Relative egg mass increases in species with cooler egg temperatures and longer embryonic periods to offset associated increases in energetic requirements of embryos. Egg temperatures of birds are determined by parental incubation behavior and are often cooler among tropical passerines because of reduced parental attentiveness of eggs. Here, I show that cooler egg temperatures and longer embryonic periods explained the enigmatic variation in egg mass within and among regions, based on field studies in tropical Venezuela (36 species), subtropical Argentina (16 species), and north temperate Arizona (20 species). Alternative explanations are not supported. Thus, large egg sizes may reflect compensation for increased energetic requirements of cool egg temperatures and long embryonic periods that result from reduced parental attentiveness in tropical birds.
Assuntos
Temperatura Corporal , Tamanho Celular , Embrião não Mamífero/fisiologia , Modelos Biológicos , Óvulo/citologia , Aves Canoras/fisiologia , Clima Tropical , Animais , Argentina , Arizona , Tamanho da Ninhada , Especificidade da Espécie , VenezuelaRESUMO
Embryonic development rates are reflected by the length of incubation period in birds, and these vary substantially among species within and among geographical regions. The incubation periods are consistently shorter in North America (Arizona study site) than in tropical (Venezuela) and subtropical (Argentina) South America based on the study of 83 passerine species in 17 clades. Parents, mothers in particular, may influence incubation periods and resulting offspring quality through proximate pathways, while variation in maternal strategies among species can result from selection by adult and offspring mortality. Parents of long-lived species, as is common in the tropics and subtropics, may be under selection to minimize costs to themselves during incubation. Indeed, time spent incubating is often lower in the tropical and subtropical species than the related north temperate species, causing cooler average egg temperatures in the southern regions. Decreased egg temperatures result in longer incubation periods and reflect a cost imposed on offspring by parents because energy cost to the embryo and risk of offspring predation are both increased. Mothers may adjust egg size and constituents as a means to partially offset such costs. For example, reduced androgen concentrations in egg yolks may slow development rates, but may enhance offspring quality through physiological trade-offs that may be particularly beneficial in longer-lived species, as in the tropics and subtropics. We provide initial data to show that yolks of tropical birds contain substantially lower concentrations of growth-promoting androgens than north temperate relatives. Thus, maternal (and parental) effects on embryonic development rates may include contrasting and complementary proximate influences on offspring quality and deserve further field study among species.
Assuntos
Desenvolvimento Embrionário/fisiologia , Comportamento de Nidação/fisiologia , Passeriformes/embriologia , Passeriformes/metabolismo , Análise de Variância , Androgênios/análise , Animais , Proteínas do Ovo/química , Embrião não Mamífero/metabolismo , Geografia , América do Norte , América do Sul , Temperatura , Fatores de TempoRESUMO
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.
Assuntos
Adaptação Fisiológica , Fenômenos Fisiológicos da Nutrição Animal , Reprodução/fisiologia , Aves Canoras/fisiologia , Análise de Variância , Animais , Argentina , Pesos e Medidas Corporais , Comportamento Alimentar/fisiologia , Aves Canoras/crescimento & desenvolvimento , Gravação em VídeoRESUMO
Viewing life-history evolution in birds based on an age-specific mortality framework can explain broad life-history patterns, including the long incubation periods in southern latitudes documented here. I show that incubation periods of species that are matched phylogenetically and ecologically between Argentina and Arizona are longer in Argentina. Long incubation periods have mystified scientists because they increase the accumulated risk of time-dependent mortality to young without providing a clear benefit. I hypothesize that parents of species with low adult mortality accept increased risk of mortality to their young from longer incubation if this allows reduced risk of mortality to themselves. During incubation, songbird parents can reduce risk of mortality to themselves by reducing nest attentiveness (percentage of time on the nest). Here I show that parents of species with lower adult mortality exhibit reduced nest attentiveness and that lower attentiveness is associated with longer incubation periods. However, the incubation period is also modified by juvenile mortality. Clutch size variation is also strongly correlated with age-specific mortality. Ultimately, adult and juvenile mortality explain variation in incubation and other life-history traits better than the historical paradigm.