Pooled energy budgets: Resituating human energy ‐allocation trade‐offs

Across taxa, many life‐history traits vary as a function of differences in body size. 1 - 5 Among primates, including humans, allometric relationships explain many trends in metabolic, growth, reproductive, and mortality rates. 6 - 8 But humans also deviate from nonhuman primates with respect to other developmental, reproductive, and parenting characteristics. 9 - 13 Broad relationships between life‐history traits and body size assume that energy expended in activity (foraging effort) is proportional to body size, and that energy available for growth and reproduction are equivalent. Because human subsistence and parenting are based on food sharing, and cooperation in labor and childrearing, the ways by which energy is acquired and allocated to alternate expenditures are expanded. We present a modification of the general allocation model to include a mechanism for these energy transfers. Our goal is to develop a framework that incorporates this mechanism and can explain the human life‐history paradox; that is, slow juvenile growth and rapid reproduction. We suggest that the central characteristics of human subsistence and energy transfer need to be accounted for in order to more fully appreciate human life‐history variability.     

Why What Juveniles Do Matters in the Evolution of Cooperative Breeding

The evolution of cooperative breeding is complex, and particularly so in humans because many other life history traits likely evolved at the same time. While cooperative childrearing is often presumed ancient, the transition from maternal self-reliance to dependence on allocare leaves no known empirical record. In this paper, an exploratory model is developed that incorporates probable evolutionary changes in birth intervals, juvenile dependence, and dispersal age to predict under what life history conditions mothers are unable to raise children without adult cooperation. The model’s outcome variable (net balance) integrates dependent children’s production and consumption as a function of varying life history parameters to estimate the investment mothers or others have to spend subsidizing children. Results suggest that maternal-juvenile cooperation can support the early transition toward a reduction in birth intervals, a longer period of juvenile dependence, and having overlapping young. The need for adult cooperation is most evident when birth intervals are short and age at net production is late. Findings suggest that the needs of juveniles would not have been an early selective force for adult cooperation. Rather, an age-graded division of labor and the mutual benefits of maternal-juvenile cooperation could be an important, but overlooked step in the evolution of cooperative breeding.     

Malformed offspring,sibling matings,and selection against inbreeding in the sand lizard (Lacerta agilis)

We demonstrated that sand lizards (Lacerta agilis) are more likely to have malformed offspring when they mate with siblings. Offspring with malformations, such as deformed limbs and heads, have zero survival in a natural population. Normal-looking siblings of malformed hatchlings also had a reduced survival in the wild, compared to offspring from clutches in which all siblings appeared normal. The proportion of malformed hatchlings in the natural population was ca. 10%, in spite of differences in juvenile dispersal between males and females. Male juveniles disperse significantly further from their natal sites than do female juveniles.     

No sibling odor preference in juvenile three-spined sticklebacks

Laboratory-bred juvenile three-spined sticklebacks from 11 sibshipsdid not prefer to shoal with their siblings when they were offeredthe choice between odor from unfamiliar siblings and non-kinin a fluviarium, although the power for finding a significantpreference was very high (0.99). The test fish preferred theside where odor from the heavier shoal was supplied; this shows thatthey could appreciate odor cues from conspecifics in our apparatusand should have preferred their siblings if such a preferenceexists. Our results are compatible with theoretical predictionsbut are at variance with previous findings by other authors.We used independent replicates in a blind protocol with strictrandomization of fish and procedures.     

Food quality effects on instar‐specific life histories of a holometabolous insect

1. It is a long‐standing challenge to understand how changes in food resources impact consumer life history traits and, in turn, impact how organisms interact with their environment. To characterize food quality effects on life history, most studies follow organisms throughout their life cycle and quantify major life events, such as age at maturity or fecundity. From these studies, we know that food quality generally impacts body size, juvenile development, and life span. Importantly, throughout juvenile development, many organisms develop through several stages of growth that can have different interactions with their environment. For example, some parasitoids typically attack larger instars, whereas larval insect predators typically attack smaller instars. Interestingly, most studies lump all juvenile stages together, which ignores these ecological changes over juvenile development.
2. We combine a cross‐sectional experimental approach with a stage‐structured population model to estimate instar‐specific vital rates in the bean weevil, Callosobruchus maculatus across a food quality gradient. We characterize food quality effects on the bean weevil's life history traits throughout its juvenile ontogeny to test how food quality impacts instar‐specific vital rates.
3. Vital rates differed across food quality treatments within each instar; however, their effect differed with instar. Weevils consuming low‐quality food spent 38%, 37%, and 18% more time, and were 34%, 53%, and 63% smaller than weevils consuming high‐quality food in the second, third, and fourth instars, respectively. Overall, our results show that consuming poor food quality means slower growth, but that food quality effects on vital rates, growth and development are not equal across instars. Differences in life history traits over juvenile ontogeny in response to food quality may impact how organisms interact with their environment, including how susceptible they are to predation, parasitism, and their competitive ability.

     

The evolution of lamprey (Petromyzontida) life history and the origin of metamorphosis

Modern lampreys (Petromyzontiformes) are one of two lineages of surviving jawless fishes (agnathans), and are thus of critical importance to understanding the evolution of the vertebrates. Although their fossil record is meager, it appears they have remained morphologically conserved for at least 360 million years, but the origin of their multi-stage life history is unclear. Unlike hagfishes, the other extant group of jawless fishes, which exhibit direct development, all modern lampreys possess a complex life cycle which includes a long-lived freshwater larval (or ammocoete) period, followed by a true metamorphosis into a sexually-immature juvenile and then mature adult which differ dramatically in their morphology and ecology from the larva. Because of their basal position, it is critical to understand when the extant lamprey life history evolved, and if such a life history was present in the last common ancestor of agnathans and gnathostomes. Recent discoveries in paleontology, genomic analyses, and developmental biology are providing insights into this problem. The current review synthesizes these findings and concludes that the ancestral lamprey life cycle followed a direct development. We suggest that the larval period was short and relatively limited if present at all, but that the juvenile included modern larval traits; over the course of evolution, differential selection pressures throughout the lifetime produced distinct larval and juvenile/adult periods. Each period required the dramatically different morphologies seen in modern lampreys, ultimately requiring a true metamorphosis to accommodate the large changes in the body plan and to maximize the efficiency of each life period. As a result, modern lamprey life histories are a patchwork of ancestral and derived characters.     

Complex offspring size effects: variations across life stages and between species

Classical optimality models of offspring size and number assume a monotonically increasing relationship between offspring size and performance. In aquatic organisms with complex life cycles, the size–performance function is particularly hard to grasp because measures of performance are varied and their relationships with size may not be consistent throughout early ontogeny. Here, we examine size effects in premetamorphic (larval) and postmetamorphic (juvenile) stages of brooding marine animals and show that they vary contextually in strength and direction during ontogeny and among species. Larger offspring of the sea anemone Urticina felina generally outperformed small siblings at the larval stage (i.e., greater settlement and survival rates under suboptimal conditions). However, results differed when analyses were conducted at the intrabrood versus across‐brood levels, suggesting that the relationship between larval size and performance is mediated by parentage. At the juvenile stage (15 months), small offspring were less susceptible than large ones to predation by subadult nudibranchs and both sizes performed similarly when facing adult nudibranchs. In a sympatric species with a different life history (Aulactinia stella), all juveniles suffered similar predation rates by subadult nudibranchs, but smaller juveniles performed better (lower mortalities) when facing adult nudibranchs. Size differences in premetamorphic performance of U. felina were linked to total lipid contents of larvae, whereas size‐specific predation of juvenile stages followed the general predictions of the optimal foraging strategy. These findings emphasize the challenge in gathering empirical support for a positive monotonic size–performance function in taxa that exhibit complex life cycles, which are dominant in the sea.     

Spatial structuring of an evolving life-history strategy under altered environmental conditions

Human disturbances to ecosystems have created challenges to populations worldwide, forcing them to respond phenotypically in ways that increase their fitness under current conditions. One approach to examining population responses to disturbance in species with complex life histories is to study species that exhibit spatial patterns in their phenotypic response across populations or demes. In this study, we investigate a threatened population of fall chinook salmon (Oncorhynchus tshawytscha) in the Snake River of Idaho, in which a significant fraction of the juvenile population have been shown to exhibit a yearling out-migration strategy which had not previously been thought to exist. It has been suggested that dam-related environmental changes may have altered the selective pressures experienced by out-migrating fall chinook, driving evolution of a later and more selectively advantageous migration strategy. Using isotopic analysis of otoliths from returning adult spawners, we reconstructed the locations of individual fish at three major juvenile life stages to determine if the representation of the yearling life history was geographically structured within the population. We reconstructed juvenile locations for natal, rearing and overwintering life stages in each of the major spawning areas in the basin. Our results indicate that the yearling life-history strategy is predominantly represented within one of the main spawning regions, the Clearwater River, rather than being distributed throughout the basin. Previous studies have shown the Clearwater River to have cooler temperatures, later hatch dates, and later outmigration of juveniles, indicating a link between environment and expression of the yearling life history. Our data suggest that this new yearling life history may be disproportionally represented in returning adult spawners, indicating selection for this life history within the population.     

Parasitism in early life: environmental conditions shape within‐brood variation in responses to infection

Parasites play key ecological and evolutionary roles through the costs they impose on their host. In wild populations, the effect of parasitism is likely to vary considerably with environmental conditions, which may affect the availability of resources to hosts for defense. However, the interaction between parasitism and prevailing conditions is rarely quantified. In addition to environmental variation acting on hosts, individuals are likely to vary in their response to parasitism, and the combined effect of both may increase heterogeneity in host responses. Offspring hierarchies, established by parents in response to uncertain rearing conditions, may be an important source of variation between individuals. Here, we use experimental antiparasite treatment across 5 years of variable conditions to test how annual population productivity (a proxy for environmental conditions) and parasitism interact to affect growth and survival of different brood members in juvenile European shags (Phalacrocorax aristotelis). In control broods, last‐hatched chicks had more plastic growth rates, growing faster in more productive years. Older siblings grew at a similar rate in all years. Treatment removed the effect of environment on last‐hatched chicks, such that all siblings in treated broods grew at a similar rate across environmental conditions. There were no differences in nematode burden between years or siblings, suggesting that variation in responses arose from intrinsic differences between chicks. Whole‐brood growth rate was not affected by treatment, indicating that within‐brood differences were driven by a change in resource allocation between siblings rather than a change in overall parental provisioning. We show that gastrointestinal parasites can be a key component of offspring's developmental environment. Our results also demonstrate the value of considering prevailing conditions for our understanding of parasite effects on host life‐history traits. Establishing how environmental conditions shape responses to parasitism is important as environmental variability is predicted to increase.     

Delayed Phenotypic Expression of Growth Hormone Transgenesis during Early Ontogeny in Atlantic Salmon (Salmo salar)?

Should growth hormone (GH) transgenic Atlantic salmon escape, there may be the potential for ecological and genetic impacts on wild populations. This study compared the developmental rate and respiratory metabolism of GH transgenic and non-transgenic full sibling Atlantic salmon during early ontogeny; a life history period of intense selection that may provide critical insight into the fitness consequences of escaped transgenics. Transgenesis did not affect the routine oxygen consumption of eyed embryos, newly hatched larvae or first-feeding juveniles. Moreover, the timing of early life history events was similar, with transgenic fish hatching less than one day earlier, on average, than their non-transgenic siblings. As the start of exogenous feeding neared, however, transgenic fish were somewhat developmentally behind, having more unused yolk and being slightly smaller than their non-transgenic siblings. Although such differences were found between transgenic and non-transgenic siblings, family differences were more important in explaining phenotypic variation. These findings suggest that biologically significant differences in fitness-related traits between GH transgenic and non-transgenic Atlantic salmon were less than family differences during the earliest life stages. The implications of these results are discussed in light of the ecological risk assessment of genetically modified animals.     

Patterns of Juvenile Habitat Use and Seasonality of Settlement by Permit, Trachinotus falcatus

Synopsis Permit, Trachinotus falcatus are economically and ecologically important throughout their range of the Caribbean, subtropical and tropical western Atlantic, and Gulf of Mexico. Despite their economic importance, little is known about the biology and ecology of permit, and most existing information is from Florida. While sufficient information is available to paint a general picture of permit life history, details are lacking for most life stages. For the juvenile life stage, nursery habitats and size and age at settlement have not yet been defined. Although six distinct habitat types (medium energy and low energy windward beaches, leeward beaches, and windward, leeward, and lagoon interior mangrove shorelines) were sampled to determine spatial patterns of habitat use by early juvenile permit at Turneffe Atoll, Belize, Central America, and the Florida Keys, USA, 99% of juvenile permit were found along medium energy windward beaches, indicating their role as nursery habitat for this species. A sub-sample of juvenile permit from Florida was examined to estimate spawning date and age at settlement from otoliths. Size-frequency distributions and otolith age analysis indicate that larval duration is approximately 15 – 20 days, and settlement occurs year-round. Since permit in Florida spawn March through July, from March through September in Cuba, and from February through October in Belize, year-round settlement indicates population connectivity via larval transport. These results lay the foundation for future research on larval supply, population connectivity, and juvenile ecology, and will aid in the ongoing formulation of a conservation plan toward a sustainable fishery for permit.     

Diversity of juvenile Chinook salmon life history pathways

Life history variability includes phenotypic variation in morphology, age, and size at key stage transitions and arises from genotypic, environmental, and genotype-by-environment effects. Life history variation contributes to population abundance, productivity, and resilience, and management units often reflect life history classes. Recent evidence suggests that past Chinook salmon (Oncorhynchus tshawytscha) classifications (e.g., ‘stream’ and ‘ocean’ types) are not distinct evolutionary lineages, do not capture the phenotypic variation present within or among populations, and are poorly aligned with underlying ecological and developmental processes. Here we review recently reported variation in juvenile Chinook salmon life history traits and provide a refined conceptual framework for understanding the causes and consequences of the observed variability. The review reveals a broad continuum of individual juvenile life history pathways, defined primarily by transitions among developmental stages and habitat types used during freshwater rearing and emigration. Life history types emerge from discontinuities in expressed pathways when viewed at the population scale. We synthesize recent research that examines how genetic, conditional, and environmental mechanisms likely influence Chinook salmon life history pathways. We suggest that threshold models hold promise for understanding how genetic and environmental factors influence juvenile salmon life history transitions. Operational life history classifications will likely differ regionally, but should benefit from an expanded lexicon that captures the temporally variable, multi-stage life history pathways that occur in many Chinook salmon populations. An increased mechanistic awareness of life history diversity, and how it affects population fitness and resilience, should improve management, conservation, and restoration of this iconic species.     

Pleiotropic Effects of methoprene-tolerant(Met), a Gene Involved in Juvenile Hormone Metabolism, on Life History Traits in Drosophila melanogaster

Life history theory assumes that there are alleles with pleiotropic effects on fitness components. Although quantitative genetic data are often consistent with pleiotropy, there are few explicit examples of pleiotropic loci. The Drosophila melanogaster gene Methoprene-tolerant (Met) may be such a locus. The Met gene product, a putative juvenile hormone receptor, facilitates the action of juvenile hormone (JH) and JH analogs; JH affects many life history traits in arthropods. Here we use quantitative complementation to investigate effects of Met mutant and wildtype alleles on female developmental time, onset of reproduction, and fecundity. Whereas the alleles did not differ in their effects on developmental time, we detected allelic variation for the onset of reproduction and for age-specific fecundity. Alleles influenced phenotypic co-variances among traits (developmental time and onset of reproduction; onset of reproduction and both early and late fecundity; early and late fecundity), suggesting that alleles of Met vary in their pleiotropic effects upon life history. Furthermore, the genetic covariance between developmental time and early fecundity attributed to alleles of Met was negative, indicating consistent pleiotropic effects among alleles on these traits. The allelic effects of Met support genetic models where pleiotropy at genes associated with hormone regulation can contribute to the evolution of life history traits.     

Self-grooming and sibling recognition in meadow voles,Microtus pennsylvanicus,and prairie voles,M. ochrogaster

Self-grooming in response to the odours of an opposite-sex conspecific may reflect sexual motivation on the part of the actor. We tested the hypothesis that meadow voles, Microtus pennsylvanicus, and prairie voles,M. ochrogaster , both self-groom at different rates when exposed to the odours of their siblings as compared to those of nonsiblings. This prediction was studied under the context of social memory for siblings and the effects of isolation on memory for siblings. The hypothesis explains the self-grooming responses of meadow voles isolated from their siblings for 10 days and of prairie voles isolated from their siblings for 20 days. However, 20 days of isolation for male and female meadow voles, and 30 days of isolation for male prairie voles were sufficient to induce these animals to self-groom at a similar rate to both sibling and nonsibling odours, suggesting that after isolation these animals no longer recognized their opposite-sex siblings. Female prairie voles isolated for 30 days self-groomed more in response to the odours of male nonsiblings than to those of male siblings, suggesting that female prairie voles still recognize their male siblings after isolation. This study is the first to provide empirical evidence that, in the context of sibling recognition, self-grooming behaviour is directed at unfamiliar opposite-sex conspecifics, and that the communicative function of self-grooming (sexual motivation) is associated with social memory for siblings. Differences in the self-grooming behaviour of meadow voles and prairie voles may be associated with several aspects of their life history characteristics.     

Age-dependency in hunting ability among the Ache of eastern Paraguay

This paper examines changes in hunting ability across the lifespan for the Ache of eastern Paraguay. Hunting ability is decomposed into two components-finding prey and probability of kill upon encounter- and analyzed for important prey species. Results support the argument that skill acquisition is an important aspect of the human foraging niche with hunting outcome variables reaching peaks surprisingly late in life, significantly after peaks in strength. The implications of this study are important for modeling the role of the human foraging niche in the co-evolution of various outstanding human life history characteristics such as large brains, long lifespans, and extended juvenile periods.     

Contrasting effects of tropical cyclones on the annual survival of a pelagic seabird in the Indian Ocean

Tropical cyclones are renowned for their destructive nature and are an important feature of marine and coastal tropical ecosystems. Over the last 40 years, their intensity, frequency and tracks have changed, partly in response to ocean warming, and future predictions indicate that these trends are likely to continue with potential consequences for human populations and coastal ecosystems. However, our understanding of how tropical cyclones currently affect marine biodiversity, and pelagic species in particular, is limited. For seabirds, the impacts of cyclones are known to be detrimental at breeding colonies, but impacts on the annual survival of pelagic adults and juveniles remain largely unexplored and no study has simultaneously explored the direct impacts of cyclones on different life‐history stages across the annual life cycle. We used a 20‐year data set on tropical cyclones in the Indian Ocean, tracking data from 122 Round Island petrels and long‐term capture–mark–recapture data to explore the impacts of tropical cyclones on the survival of adult and juvenile (first year) petrels during both the breeding and migration periods. The tracking data showed that juvenile and adult Round Island petrels utilize the three cyclone regions of the Indian Ocean and were potentially exposed to cyclones for a substantial part of their annual cycle. However, only juvenile petrel survival was affected by cyclone activity; negatively by a strong cyclone in the vicinity of the breeding colony and positively by increasing cyclone activity in the Northern Indian Ocean where they spend the majority of their first year at sea. These contrasting effects raise the intriguing prospect that the projected changes in cyclones under current climate change scenarios may have positive as well as the more commonly perceived negative impacts on marine biodiversity.     

Acoustic signals of baby black caimans

In spite of the importance of crocodilian vocalizations for the understanding of the evolution of sound communication in Archosauria and due to the small number of experimental investigations, information concerning the vocal world of crocodilians is limited. By studying black caimans Melanosuchus niger in their natural habitat, here we supply the experimental evidence that juvenile crocodilians can use a graded sound system in order to elicit adapted behavioral responses from their mother and siblings. By analyzing the acoustic structure of calls emitted in two different situations (‘undisturbed context’, during which spontaneous calls of juvenile caimans were recorded without perturbing the group, and a simulated ‘predator attack’, during which calls were recorded while shaking juveniles) and by testing their biological relevance through playback experiments, we reveal the existence of two functionally different types of juvenile calls that produce a different response from the mother and other siblings. Young black caimans can thus modulate the structure of their vocalizations along an acoustic continuum as a function of the emission context. Playback experiments show that both mother and juveniles discriminate between these ‘distress’ and ‘contact’ calls. Acoustic communication is thus an important component mediating relationships within family groups in caimans as it is in birds, their archosaurian relatives. Although probably limited, the vocal repertoire of young crocodilians is capable of transmitting the information necessary for allowing siblings and mother to modulate their behavior.     

Early Social Networks Predict Survival in Wild Bottlenose Dolphins

A fundamental question concerning group-living species is what factors influence the evolution of sociality. Although several studies link adult social bonds to fitness, social patterns and relationships are often formed early in life and are also likely to have fitness consequences, particularly in species with lengthy developmental periods, extensive social learning, and early social bond-formation. In a longitudinal study of bottlenose dolphins (Tursiops sp.), calf social network structure, specifically the metric eigenvector centrality, predicted juvenile survival in males. Additionally, male calves that died post-weaning had stronger ties to juvenile males than surviving male calves, suggesting that juvenile males impose fitness costs on their younger counterparts. Our study indicates that selection is acting on social traits early in life and highlights the need to examine the costs and benefits of social bonds during formative life history stages.     

Trade-offs in skillacquisition and time allocation among juvenile chacma baboons

We hypothesize that juvenile baboons are less efficient foragers than adult baboons owing to their small size, lower level of knowledge and skill, and/or lesser ability to maintain access to resources. We predict that as resources are more difficult to extract, juvenile baboons will demonstrate lower efficiency than adults will because of their lower levels of experience. In addition, we hypothesize that juvenile baboons will be more likely to allocate foraging time to easier-to-extract resources owing to their greater efficiency in acquiring those resources. We use feeding efficiency and time allocation data collected on a wild, free-ranging, non-provisioned population of chacma baboons (Papio hamadryas ursinus) in the Moremi Wildlife Reserve, Okavango Delta, Botswana to test these hypotheses. The major findings of this study are: 1. Juvenile baboons are significantly less efficient foragers than adult baboons primarily for difficult-to-extract resources. We propose that this age-dependent variation in efficiency is due to differences in memory and other cognitive functions related to locating food resources, as is indicated by the greater amount of time juvenile baboons spend searching for food. There is no evidence that smaller body size or competitive disruption influences the differences in return rates found between adult and juvenile baboons in this study. 2. An individual baboon’s feeding efficiency for a given resource can be used to predict the duration of its foraging bouts for that resource. These results contribute both to our understanding of the ontogeny of behavioral development in nonhuman primates, especially regarding foraging ability, and to current debate within the field of human behavioral ecology regarding the evolution of the juvenile period in primates and humans. Sara E. Johnson is Assistant Professor of Anthropology at California State University, Fullerton. She received her Ph.D. in Anthropology (Human Evolutionary Ecology) from the University of New Mexico in 2001. She uses behavioral ecology and life history theory to address her research interests in the evolution of primate and human growth; ecological variation and phenotypic plasticity in growth and development; ecological variation in life course trajectories, including fertility, health, morbidity, and mortality differentials; food acquisition and production related to nutrition; societal transofmration and roles of the elderly among indigenous peoples; and women’s reproductive and productive roles in both traditional and nontraditional societies. For the past decade she has conducted research on these issues in several different populations, including chacma baboons in the Okavango Delta of Botswana, two multiethnic communities of forager/agropastoralists in the Okavango Delta of Botswana, and among New Mexican men. John Bock is Associate Professor of Anthropology at California State University at Fullerton and is Associate Editor of Human Nature. He received a Ph.D. in Anthropology (Human Evolutionary EcologY) from the University of New Mexico in 1995, and from 1995 to 1998 was an Andrew W. Mellon Foundation postdoctoral fellow in demography and epidemiology at the National Centre for Epidemiology and Population Health at Australian National University. His recent research has focused on applying life history theory to understanding the evolution of the primate and human juvenile period. Bock has been conducting research among the Okavango Delta peoples of Botswana since 1992, and his current research there is an examination of child development and family demography in relation to socioecology and the HIV/AIDS epidemic. Other research is focused on health disparties among minorities and indigenous peoples in Botswana and the United States related to differential access to health care.     

The importance of kinship and familiarity in social interactions between mice

We investigated a range of social interactions between mice which differed in their degree of relatedness and familiarity. Unfamiliar half siblings (sharing paternity only) differed significantly from unfamiliar non-siblings (sharing neither mother nor father) in their tendency to perform aggression-related interactions and in the amount of passive body contact they showed. Differences between half and full siblings in patterns of interaction appeared to be due to differing degrees of familiarity with companions. Kinship effects disappeared completely when animals were allowed to become familiar. We discuss the functional significance of the familiarity and kinship effects we found, including differences between the sexes in the types of interaction showing kinship effects. Differences between adult and juvenile mice are also briefly discussed.