High evolutionary constraints limited adaptive responses to past climate changes in toad skulls

Interactions among traits that build a complex structure may be represented as genetic covariation and correlation. Genetic correlations may act as constraints, deflecting the evolutionary response from the direction of natural selection. We investigated the relative importance of drift, selection, and constraints in driving skull divergence in a group of related toad species. The distributional range of these species encompasses very distinct habitats with important climatic differences and the species are primarily distinguished by differences in their skulls. Some parts of the toad skull, such as the snout, may have functional relevance in reproductive ecology, detecting water cues. Thus, we hypothesized that the species skull divergence was driven by natural selection associated with climatic variation. However, given that all species present high correlations among skull traits, our second prediction was of high constraints deflecting the response to selection. We first extracted the main morphological direction that is expected to be subjected to selection by using within- and between-species covariance matrices. We then used evolutionary regressions to investigate whether divergence along this direction is explained by climatic variation between species. We also used quantitative genetics models to test for a role of random drift versus natural selection in skull divergence and to reconstruct selection gradients along species phylogeny. Climatic variables explained high proportions of between-species variation in the most selected axis. However, most evolutionary responses were not in the direction of selection, but aligned with the direction of allometric size, the dimension of highest phenotypic variance in the ancestral population. We conclude that toad species have responded to selection related to climate in their skulls, yet high evolutionary constraints dominated species divergence and may limit species responses to future climate change.     

Apes in a changing world – the effects of global warming on the behaviour and distribution of African apes

Aim In this study we use a modelling approach to identify: (1) the factors responsible for the differences in ape biogeography, (2) the effects that global warming might have on distribution patterns of African apes, (3) the underlying mechanisms for these effects, and (4) the implications that behavioural flexibility might be expected to have for ape survival. All African apes are highly endangered, and the need for efficient conservation methods is a top priority. The expected changes in world climate are likely to further exacerbate the difficulties they face. Our study aims to further understand the mechanisms that link climatic conditions to the behaviour and biogeography of ape species. Location Africa. Method We use an existing validated time budgets model, derived from data on 20 natural populations of gorillas (Gorilla beringei and Gorilla gorilla) and chimpanzees (Pan troglodytes and Pan paniscus), which specifies the relationship between climate, group size, body weight and time available for various activities, to predict ape distribution across Africa under a uniform worst‐case climate change scenario. Results We demonstrate that a worst‐case global warming scenario is likely to alter the delicate balance between different time budget components. Our model points to the importance of annual temperature variation, which was found to have the strongest impact on ape biogeography. Our simulation indicates that rising temperatures and changes in rainfall patterns are likely to have strong effects on ape survival and distribution, particularly for gorillas. Even if they behaved with maximum flexibility, gorillas may not be able to survive in most of their present habitats if the climate was to undergo extreme changes. The survival of chimpanzees was found to be strongly dependent on the minimum viable group size required. Main conclusions Our model allows us to explore how climatic conditions, individual behaviour and morphological traits may interact to limit the biogeographical distributions of these species, thereby allowing us to predict the effects of climate change on African ape distributions under different climate change regimes. The model suggests that climate variability (i.e. seasonality) plays a more important role than the absolute magnitude of the change, but these data are not normally provided by climate models.     

Habitat size, recruitment, and longevity as factors limiting population size in stage-structured species

Surprisingly little research has evaluated how habitat size may limit the population size of species that use different habitats at different stages of their lives. Here we develop simple discrete-time models to describe the population dynamics of species that use separate juvenile and adult habitats. Analytic solutions, model simulations, and elasticity and sensitivity analyses show that adult abundance is only limited by the size of the juvenile habitat when both adult habitat size and recruitment are much larger than juvenile habitat size. Juvenile habitat plays a marginally greater role in limiting population size for species with closed populations, where recruitment is proportional to adult abundance, versus open populations. Because adult populations often accumulate pulses of juveniles, adult habitat size can strongly limit population size over a broad range of parameter values, an effect that increases as the longevity of a species increases. Limited empirical research from a range of taxa supports these model predictions, although few studies were designed to actually test the limiting role of juvenile versus adult habitat. Future research must carefully evaluate whether and how processes at the juvenile stage affect adult abundance, and conservation efforts may be able to use this model to evaluate the cost-effectiveness, vis-a-vis increasing adult abundance, of time and money allocated to protecting juvenile habitats.     

Fission–fusion social systems as a strategy for coping with ecological constraints: a primate case

Fission–fusion social systems, in which members of a social community form frequently changing subgroups, occur in a number of mammalian taxa. Such systems are assumed to be a response to the costs of grouping, but evidence to support this hypothesis is limited. We use a linear programming approach to build a time budget model that predicts the upper bound on group size in order to test the hypothesis that fission–fusion social systems are the outcome of time constraints. Comparative data from 14 wild chimpanzee (Pan spp.) populations are used to derive a set of equations defining the relationship between climatic variables and time budget components, which are then used to calculate the upper limits on group size that can be maintained in different habitats. We validate the model by showing that it correctly predicts the presence/absence of chimpanzees across sub-Saharan Africa and the group sizes observed in natural populations. The model suggests that the costs of travel are limiting for chimpanzees. Chimpanzees can reduce these costs dramatically by fissioning their bonded communities into small foraging parties. If they did not, they would be unable to live in any habitats where they currently occur.     

Troop size and human-modified habitat affect the ranging patterns of a chacma baboon population in the cape peninsula, South Africa

Differences in group size and habitat use are frequently used to explain the extensive variability in ranging patterns found across the primate order. However, with few exceptions, our understanding of primate ranging patterns stems from studies of single groups and both intra- and inter-specific meta-analyses. Studies with many groups and those that incorporate whole populations are rare but important for testing socioecological theory in primates. We quantify the ranging patterns of nine chacma baboon troops in a single population and use Spearman rank correlations and generalized linear mixed models to analyze the effects of troop size and human-modified habitat (a proxy for good quality habitat) on home range size, density (individuals/km(2) ), and daily path length. Intrapopulation variation in home range sizes (1.5-37.7 km(2) ), densities (1.3-12.1 baboons/km(2) ), and daily path lengths (1.80-6.61 km) was so vast that values were comparable to those of baboons inhabiting the climatic extremes of their current distribution. Both troop size and human-modified habitat had an effect on ranging patterns. Larger troops had larger home ranges and longer daily path lengths, while troops that spent more time in human-modified habitat had shorter daily path lengths. We found no effect of human-modified habitat on home range size or density. These results held when we controlled for the effects of both a single large outlier troop living exclusively in human-modified habitat and baboon monitors on our spatial variables. Our findings confirm the ability of baboons, as behaviorally adaptable dietary generalists, to not only survive but also to thrive in human-modified habitats with adjustments to their ranging patterns in accordance with current theory. Our findings also caution that studies focused on only a small sample of groups within a population of adaptable and generalist primate species may underestimate the variability in their respective localities.     

Changes in habitat associations during range expansion: disentangling the effects of climate and residence time

The distributions of many species are not at equilibrium with their environment. This includes spreading non-native species and species undergoing range shifts in response to climate change. The habitat associations of these species may change during range expansion as less favourable climatic conditions at expanding range margins constrain species to use only the most favourable habitats, violating the species distribution model assumption of stationarity. Alternatively, changes in habitat associations could result from density-dependent habitat selection; at range margins, population densities are initially low so species can exhibit density-independent selection of the most favourable habitats, while in the range core, where population densities are higher, species spread into less favourable habitat. We investigate if the habitat preferences of the non-native common waxbill Estrilda astrild changed as they spread in three directions (north, east and south-east) in the Iberian Peninsula. There are different degrees of climatic suitability and colonization speed across range expansion axes, allowing us to separate the effects of climate from residence time. In contrast to previous studies we find a stronger effect of residence time than climate in influencing the prevalence of common waxbills. As well as a strong additive effect of residence time, there were some changes in habitat associations, which were consistent with density-dependent habitat selection. The combination of broader habitat associations and higher prevalence in areas that have been colonised for longer means that species distribution models constructed early in the invasion process are likely to underestimate species’ potential distribution.     

Processing power limits social group size: computational evidence for the cognitive costs of sociality

Sociality is primarily a coordination problem. However, the social (or communication) complexity hypothesis suggests that the kinds of information that can be acquired and processed may limit the size and/or complexity of social groups that a species can maintain. We use an agent-based model to test the hypothesis that the complexity of information processed influences the computational demands involved. We show that successive increases in the kinds of information processed allow organisms to break through the glass ceilings that otherwise limit the size of social groups: larger groups can only be achieved at the cost of more sophisticated kinds of information processing that are disadvantageous when optimal group size is small. These results simultaneously support both the social brain and the social complexity hypotheses.     

'Back to the future': how archaeological remains can describe salmon adaptation to climate change

A strategy for species to survive climate change will be to change adaptively their way of life. Understanding rapid adaptation to climate change is therefore a priority for current research. In this issue, Turrero et al. (2012) use an original approach to unravel life history trait responses to climate change in two fish species (Salmo trutta and S. salar). Going against the flow, the authors adopt the strategy of going back to the future by investigating the responses of fish to the warming periods that followed the Last Glacial Period (approximately 30-20,000 years BP). To do this, they analysed Salmo vertebrae from well-dated archaeological sites in northern Spain in order to uncover key life history traits, which they then compared to those of contemporary specimens. They found that, as the climate got warmer, Salmo species tended to reduce the time spent in growing areas and reached spawning areas at a younger age; this tendency began approximately 15,000 years BP and accelerated in contemporary periods. The implication is a lower age at maturity and a lower reproductive success, which they tentatively related to recent declines in population growth rate. This innovative study demonstrates how changes in life history traits are linked both to the population growth rate and to the evolutionary rate under climatic constraints, which may serve as a basis for future conservation research.     

Non‐breeding range size predicts the magnitude of population trends in trans‐Saharan migratory passerine birds

Understanding why populations of some migratory species show a directional change over time, i.e. increase or decrease, while others do not, remains a challenge for ecological research. One possible explanation is that species with smaller non‐breeding ranges may have more pronounced directional population trends, and their populations are thus more sensitive to the variation in environmental conditions in their non‐breeding quarters. According to the serial residency hypothesis, this sensitivity should lead to higher magnitudes (i.e. absolute values) of population trends for species with smaller non‐breeding ranges, with the direction of trend being either positive or negative depending on the nature of the environmental change. We tested this hypothesis using population trends over 2001–2012 for 36 sub‐Saharan migratory passerine birds breeding in Europe. Namely, we related the magnitude of the species' population trends to the size of their sub‐Saharan non‐breeding grounds, whilst controlling for factors including number of migration routes, non‐breeding habitat niche and wetness, breeding habitat type and life‐history strategy. The magnitude of species' population trends grew with decreasing absolute size of sub‐Saharan non‐breeding ranges, and this result remained significant when non‐breeding range size was expressed relative to the size of the breeding range. After repeating the analysis with the trend direction, the relationship with the non‐breeding range size disappeared, indicating that both population decreases and increases are frequent amongst species with small non‐breeding range sizes. Therefore, species with small non‐breeding ranges are at a higher risk of population decline due to adverse factors such as habitat loss or climatic extremes, but their populations are also more likely to increase when suitable conditions appear. As non‐breeding ranges may originate from stochasticity of non‐breeding site selection in naive birds (‘serial‐residency’ hypothesis), it is crucial to maintain a network of stable and resilient habitats over large areas of birds’ non‐breeding quarters.     

Socioecological factors influencing population structure of gorillas and chimpanzees

Differences in distribution and density between gorillas and chimpanzees are reconsidered with special reference to population structure. Both ecological and social factors influencing population structure are compared between species and between habitats within species. Gorillas and chimpanzees respond differently to a decline in food quality, such as fruit scarcity: gorillas change diet and decrease range, while chimpanzees do not change diet but may expand range. These responses result in different effects on their grouping patterns. For gorillas the dispersed distribution and reduction of range size decreases the rate of inter-unit encounters and female transfer. The concentration of social units increases the rate of aggressive contact between units and stimulates female transfer. Social units of gorillas may crowd or disperse in order to attain the optimal density. This tendency may result in similar densities of gorillas across habitats. By contrast, the distribution patterns or range size may not affect inter-unit relationships of chimpanzees. Within a single unit-group, various reproductive strategies are adopted by both sexes. Independent travel of females and flexible grouping patterns enable them to survive at very low density in extraordinary large ranges. Density and inter-unit relationships are good criteria for a healthy population of gorillas, while the size of unit-group and inter-individual relationships are good criteria for chimpanzees. Conservation planners should consider these differences for sympatric and allopatric survival in these species.     

Dynamics of highly repetitive DNA fraction as indicator of speciation in species of the family Poaceae

Dynamics of highly repetitive DNA fraction, that compose up to 99% of large cereal genomes, is a key for understanding mechanisms of speciation on molecular level. Components of this fraction and particulary transposable elements (TEs) wre explored in diploid ancestor of cultivated wheat — the species Sitopsis group (Aegilops, Podceae) by molecular cytogenetics and molecular genetic methods. It was discovered that TE is highly dynamic in time and space and could promote or intensify morphological and karyotypical changes, some of which may be potentially important for the process of microevolution, and allow species with plastic genomes to survive as new forms or even species in times of rapid climatic changes.     

Social transmission of migratory knowledge: quantifying the risk of losing migratory behavior

When migration is a learned behavior, small populations have a significant problem of maintaining migratory knowledge across generations. These populations risk losing migratory behavior entirely, which may exacerbate existing stressors on population size. Here we investigated the success of various behavioral, demographic, and social factors towards maintaining migration within small populations. Using a discrete-time probabilistic model to simulate repeated migrations, we found that migratory group size plays an important role in maintaining migratory knowledge within the population. Rare, large groups allow for migratory knowledge to be spread to many individuals at once. When a population learns migration information incrementally, the presence of individuals that can learn quickly, therefore transitioning rapidly into leaders, has a profound impact on migrational persistence. Furthermore, small populations are better able to maintain migratory behavior when groups rely on informed leaders as compared to using collective group knowledge, even when that collective knowledge is heavily weighted towards knowledgeable individuals. Finally, we found that both species with short lifespans and species that migrate with fixed group compositions are at especially high risk of losing their migration behavior at small population sizes.     

Comparison of the predicted impact of a screwworm fly outbreak in Australia using a growth index model and a life-cycle model

Abstract. The spatial population dynamics of an Old World screwworm fly, Chrysomya bezziana Villeneuve (OWS), outbreak in Australia have been modelled in two ways. The first model uses weekly growth indices derived from climatic data to predict the adult female population. The second is a detailed cohort life-cycle model. Due to technical and time constraints, the growth index model is preferred as the biological component of a much larger bioeconomic model because of its smaller program size and faster execution. In deciding whether adoption of the growth index model would be at the expense of scientific accuracy, the life-cycle model was developed as a yardstick. We showed that the growth index model was a practical and adequate substitution for the OWS life-cycle model and a novel spatial/temporal modelling approach with generic qualities. We elaborate on the previously reported growth index model, describe the life-cycle model and compare the results of both models. In the event of an OWS incursion in northern or eastern Australia, given average climatic conditions, both models predict that most of the suitable range (some 2.3M km²) will be colonized within 4–5 years if an eradication campaign is not attempted. Much of its permanent range would be in tropical and subtropical extensive grazing regions. Where computer or funding resources are restrictive, models incorporating growth indices may prove adequate for spatial population studies of some species.     

Climate and mammalian life histories

Mammals display considerable geographical variation in life history traits. To understand how climatic factors might influence this variation, we analysed the relationship between life history traits – adult body size, litter size, number of litters per year, gestation length, neonate body mass, weaning age and age at sexual maturity – and several environmental variables quantifying the seasonality and predictability of temperature and precipitation across the distribution range of five terrestrial mammal groups. Environmental factors correlated strongly with each other; therefore, we used principal components analysis to obtain orthogonal climatic predictors that could be used in multivariate models. We found that in bats, primates and even‐toed ungulates adult body size tends to be larger in species inhabiting cold, dry, seasonal environments, whereas in carnivores and rodents a smaller body size is characteristic of warm, dry environments, suggesting that low food availability might limit adult size. Species inhabiting cold, dry, seasonal habitats have fewer, larger litters and shorter gestation periods; however, annual fecundity in these species is not higher, implying that the large litter size of mammals living at high latitudes is probably a consequence of time constraints imposed by strong seasonality. On the other hand, the number of litters per year and annual fecundity were greater in species inhabiting environments with higher seasonality in precipitation. Lastly, we found little evidence for specific effects of environmental variability. Our results highlight the complex effects of environmental factors in the evolution of life history traits in mammals. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 719–736.     

Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

Time Constraints Do Not Limit Group Size in Arboreal Guenons but Do Explain Community Size and Distribution Patterns

To understand how species will respond to environmental changes, it is important to know how those changes will affect the ecological stress that animals experience. Time constraints can be used as indicators of ecological stress. Here we test whether time constraints can help us understand group sizes, distribution patterns, and community sizes of forest guenons (Cercopithecus/Allochrocebus). Forest guenons typically live in small to medium sized one-male–multifemale groups and often live in communities with multiple forest guenon species. We developed a time-budget model using published data on time budgets, diets, body sizes, climate, and group sizes to predict maximum ecologically tolerable group and community sizes of forest guenons across 202 sub-Saharan African locations. The model correctly predicted presence/absence at 83% of these locations. Feeding-foraging time (an indicator of competition) limited group sizes, while resting and moving time constraints shaped guenon biogeography. Predicted group sizes were greater than observed group sizes but comparable to community sizes, suggesting community sizes are set by competition among guenon individuals irrespective of species. We conclude that time constraints and intraspecific competition are unlikely to be the main determinants of relatively small group sizes in forest guenons. Body mass was negatively correlated with moving time, which may give larger bodied species an advantage over smaller bodied species under future conditions when greater fragmentation of forests is likely to lead to increased moving time. Resting time heavily depended on leaf consumption and is likely to increase under future climatic conditions when leaf quality is expected to decrease.     

Social cognition on the Internet: testing constraints on social network size

The social brain hypothesis (an explanation for the evolution of brain size in primates) predicts that humans typically cannot maintain more than 150 relationships at any one time. The constraint is partly cognitive (ultimately determined by some aspect of brain volume) and partly one of time. Friendships (but not necessarily kin relationships) are maintained by investing time in them, and failure to do so results in an inexorable deterioration in the quality of a relationship. The Internet, and in particular the rise of social networking sites (SNSs), raises the possibility that digital media might allow us to circumvent some or all of these constraints. This allows us to test the importance of these constraints in limiting human sociality. Although the recency of SNSs means that there have been relatively few studies, those that are available suggest that, in general, the ability to broadcast to many individuals at once, and the possibilities this provides in terms of continuously updating our understanding of network members' behaviour and thoughts, do not allow larger networks to be maintained. This may be because only relatively weak quality relationships can be maintained without face-to-face interaction.     

Winter and summer weather modulate the demography of wild turkeys at the northern edge of the species distribution

Temperate species occupying habitats at the northern limit of their geographical distribution are limited by weather and climatic conditions. Such conditions often directly affect population dynamics, and thus, influence shifts in distribution via changes in demographic parameters. We examined this question by following three distinct populations of wild turkeys inhabiting areas exposed to a gradient of meteorological conditions at the northern limit of the species distribution. Four years of radio-telemetry on 181 birds and monitoring of 95 nests revealed that population demographics of wild turkeys were influenced by snow depth, winter temperature and summer rainfall. During winter, survival of turkeys decreased drastically when snow depth remained >30 cm for >10 days and also decreased as temperatures got colder. In the spring, snow persistence delayed nest initiation, whereas nest survival was negatively affected by rainfall. Our findings show that the effects of critical meteorological factors such as snow and temperature can be compounded when both reach the limit of a species tolerance simultaneously.     

Temperature dependent sex determination and climate change

One possible response of species to climate change is shifting their geographical range so as to track their climatic niche. Many concerns have been raised about the species ability to disperse effectively. I argue that species may have mechanisms, like temperature-dependent sex determination (TSD), that are responsive to climate change and may facilitate an appropriate shift in their geographical range. More specifically, I hypothesize that, under stable climatic conditions, populations of some TSD species at the edge of their range are regulated by reduced growth rate (due to skewed sex ratios or due to limited availability of suitable nesting sites). Under climate change, these populations face new climatic conditions that trigger fast population growth (e.g. by more balanced sex ratio, or greater availability of nesting sites). Increased population size may lead to increased dispersal, and thus efficient colonization of the newly created habitat patches. So, the species rapidly tracks the geographical position of its climatic niche. This conceptual model is speculative but it leads to specific hypotheses, and opens up new research questions about the existence of prior adaptations that will enable the appropriate response to climate change.