Life-history evolution in uncertain environments: bet hedging in time

Many vertebrates, forest herbs, and trees exhibit both variable age at maturity and iteroparity as adaptations to uncertain environments. We analyze a stochastic model that combines these two life-history adaptations with density-dependent fertility. Results for a model with only iteroparity are consistent with previous work; environmental uncertainty favors adult survival over juvenile survival. This holds true even if there is a moderately strong convex trade-off between adult survival and fecundity, but the direction of selection can depend on which life-history trait is considered a random variable. A life history with only developmental delay favors juvenile survival in uncertain environments, consistent with previous models of seed banks. When both developmental delay and iteroparity are included in the model, both adaptations are favored in uncertain environments. Our simulations show that selection is not necessarily a runaway process in which either developmental delay or iteroparity is favored, as recently proposed by Tuljapurkar and Wiener, but rather that selection can favor both mechanisms. Invasion analysis shows that selective pressure on life-history delays increases as environmental variation increases. Reproductive delay and adult survival can be either adaptations or constraints. Natural-history studies that estimate model parameters can resolve this uncertainty.     

Geographic variation in resource use by specialist versus generalist butterflyfishes

Localised patterns of resource use can be constrained by multiple factors. Comparison of resource use at multiple locations with differing resource availability can allow fundamental specialists to be distinguished from species that simply feed predominantly on prey types that are locally abundant. This study investigates geographic variation in the feeding ecology of coral‐feeding butterflyfishes to examine whether patterns of resource use and levels of dietary specialisation vary among distinct locations, corresponding with changes in resource availability. Our specific aims were to investigate whether the dietary niche breadth of four butterflyfishes varies among five geographically separated locations and assess whether each species utilises similar resources in each location. Resource availability and dietary composition of four butterflyfishes were quantified at three sites across each of five geographic locations throughout the Pacific. Niche breadth, niche overlap, and resource selection functions were calculated for each species at each site and compared among locations. Availability of dietary resources varied significantly among locations and sites. Chaetodon vagabundus, C. citrinellus and C. lunulatus had low levels of dietary specialisation and used different resources in each location. Chaetodon trifascialis had high levels of dietary specialisation and used the same few resources in each location. Our results indicate that relative levels of dietary specialisation among different butterflyfishes do hold at larger spatial scales, however, geographical variation in the dietary composition of all butterflyfishes indicates that prey availability has a fundamental influence on dietary composition. Highly specialised species such as C. trifascialis will be highly vulnerable to coral loss as they appear to be largely inflexible in their dietary composition. However, the increased feeding plasticity observed here for C. trifascialis suggests this species may have a greater capacity to respond to coral loss than previously assumed.     

Among population variation in specialist and generalist seed predation – the importance of host plant distribution, alternative hosts and environmental variation

Pre-dispersal seed predators can have important effects on population dynamics and trait selection in their host plants. However, the factors determining spatial variation in predation intensity are poorly known. We assessed the relative importance of host plant distribution, alternative hosts and environmental factors for among-population variation in predation in a system with three host plants, a specialist and a generalist pre-dispersal seed predator.
Effects of host plant population size were relatively more important in the specialist than in the generalist seed predator. The specialist seed predator Apion opeticum , utilizing only Lathyrus vernus occurred in less than half of the patches, and specialist seed predation was influenced only by host plant population size. The generalist Bruchus atomarius was present in nearly all patches, and generalist predation was influenced by environmental factors and availability of alternative hosts. Predation on alternative hosts was not affected by L. vernus presence.
The results suggest that a wide range of factors influences the strength of plant–seed–predator interactions, and that the relative importance of different factors depend on the degree of specialization. This will result in highly complex selection mosaics and coevolutionary trajectories.     

Emergence dynamics and bet hedging in a desert bee,Perdita portalis

Bees (series Apiformes, superfamily Apoidea) are most diverse in arid regions of the world. Arid regions (deserts and semi-deserts) are characterized by discrete rainy seasons and extreme temporal variability in rainfall. This paper documents several novel mechanisms by which one desert bee species (Perdita portalis) copes with harsh and unpredictable conditions in xeric habitats. These same mechanisms are likely to be present in diverse bee families. First, diapausing P. portalis follow a bet-hedging emergence pattern, such that only approximately half of the larvae pupate under optimal conditions. Second, emergence is condition dependent such that larvae with a low average body weight are significantly more likely to emerge than are larvae with a high average body weight under similar conditions. Finally, larval emergence is induced by exposure to high humidity (rainfall). The parallels between bee larvae and angiosperm seeds in arid environments are striking. In both cases there is clear evidence of bet hedging, emergence (or germination) is dependent on larval (or seed) condition and rainfall triggers emergence (or germination). These patterns of emergence are significant for understanding bee species diversity in arid regions.     

Decision making by generalist and specialist aphids with the same genotype

One idea to explain the high incidence of specialisation in phytophagous insects is that their nervous systems are simple and unable to efficiently recognise multiple host plants. Here this 'neural limitations' hypothesis is tested using a generalist (the summer winged virginopara) and specialist (the autumn gynopara) morph of a single clone of the black bean aphid, Aphis fabae, that are almost identical morphologically and have the same genotype. Electrical monitoring of stylet behaviour and non-invasive behavioural observations are used to compare initial plant recognition, first reproduction, 'first registered phloem contact' and 'phloem acceptance' of the generalist and specialist on their specific host plants (generalist--herbaceous hosts, Beta vulgaris, Papaver dubium, Rumex obtusifolius, Vicia faba; specialist--woody host, Euonymus europaeus) and on the same host (E. europaeus). Additional analyses of behaviour over the first few minutes of plant contact are carried out for three mutual non-host plants: Euonymus alatus, Prunus padus and Brassica pekinensis. Few examples are found where the host-selection behaviour of the specialist on its host plant is more efficient than the generalist across its hosts and there is little difference in the behaviour of the different forms on the same host. Indeed there are several instances where the generalist is behaviourally more efficient than the specialist. These include absolute time to first reproduction by the generalist on various herbaceous hosts and the same host, E. europaeus, absolute time to first registered phloem contact by the generalist on P. dubium, and incidence of phloem acceptance by the generalist on several of its herbaceous hosts. Rejection of non-hosts is carried out more efficiently by the specialist gynopara on P. padus and B. pekinensis but not on E. alatus. For generalist and specialist morphs of the same A. fabae genotype, therefore, broad host range does not appear to be associated with a reduced ability to make plant-use decisions.     

Responses of a specialist and a generalist seed predator to variation in their common resource

Fluctuations of resources in time and space will influence not only species abundance but also interactions among species. For plant–consumer interactions, the effects of resource variation have mostly been studied in systems with high resource variability. Systems with moderate variations are less studied, although ecological and evolutionary dynamics of plants and consumers are likely to be affected also by less extreme variability. The effects of variation in a particular resource should depend on consumer diet width.
We examined how spatial and temporal variation in seed production in the perennial herb Lathyrus vernus influenced population dynamics and resource utilization in two beetle pre-dispersal seed predators with different host ranges over six years. The monophagous Apion opeticum occupied fewer patches and had lower densities than the oligophagous Bruchus atomarius . The proportion of seeds attacked increased with increases in seed production between years for both seed predators. A possible explanation for these patterns is that population dynamics of beetles are driven largely by local factors and that the same factors influence both beetle performance and seed production. In B. atomarius , patterns may also be influenced by a more pronounced preference for L. vernus in years with a high seed production in L. vernus . We conclude that relatively modest variation in seed production may result in responses that differ in both direction and extent from those usually observed in systems with high variation in seed production.     

Habitat‐related patterns in phenotypic variation in a New Zealand freshwater generalist fish,and comparisons with a closely related specialist

1. Within‐species phenotypic variation is hugely variable and may play a role in determining the range of habitats a species can exploit. Our study addressed two main questions: 1. does phenotypic variation allow some species (i.e. habitat‐generalists) to use heterogeneous habitats and 2. are habitat‐generalists more variable than species occupying relatively homogeneous environments (i.e. habitat‐specialists)? 2. We examined the morphology of the common bully (Gobiomorphus cotidianus), a habitat‐generalist eleotrid fish found in lakes and rivers throughout New Zealand. We also compared the level of morphological variability in common bullies with that in the closely related redfin bully (Gobiomorphus huttoni), a habitat‐specialist of moderate‐ and fast‐flowing rivers. 3. Common and redfin bullies were collected from the South Island of New Zealand. A series of body and fin measurements were made, and cephalic dorsal head pores of the mechanosensory lateral‐line system were counted. The pores and associated canal neuromasts are important for prey detection and predator avoidance in other species, particularly, in turbulent conditions where the effectiveness of superficial neuromasts may be compromised. 4. The common bully had more dorsal head pores in fish from rivers than in those from lakes. This pattern was apparent only in adults, suggesting that selective pressures associated with adult habitat, be it rivers or lakes, are responsible. 5. As expected, there was greater phenotypic variability in the generalist common bully than in the specialist redfin bully, particularly with regard to the sensory pores, suggesting their importance for survival in turbulence. 6. We identified habitat‐related patterns in phenotypic variability in a generalist species and demonstrated a link between phenotypic variability and habitat breadth. Variation in the common bully may explain its ability to occupy a range of habitats.     

Primate population dynamics: variation in abundance over space and time

The rapid disappearance of tropical forests, the potential impacts of climate change, and the increasing threats of bushmeat hunting to wildlife, makes it imperative that we understand wildlife population dynamics. With long-lived animals this requires extensive, long-term data, but such data is often lacking. Here we present longitudinal data documenting changes in primate abundance over 45 years at eight sites in Kibale National Park, Uganda. Complex patterns of change in primate abundance were dependent on site, sampling year, and species, but all species, except blue monkeys, colonized regenerating forest, indicating that park-wide populations are increasing. At two paired sites, we found that while the primate populations in the regenerating forests had increased from nothing to a substantial size, there was little evidence of a decline in the source populations in old-growth forest, with the possible exception of mangabeys at one of the paired sites. Censuses conducted in logged forest since 1970 demonstrated that for all species, except black-and-white colobus, the encounter rate was higher in the old-growth and lightly-logged forest than in heavily-logged forest. Black-and-white colobus generally showed the opposite trend and were most common in the heavily-logged forest in all but the first year of monitoring after logging, when they were most common in the lightly-logged forest. Overall, except for blue monkey populations which are declining, primate populations in Kibale National Park are growing; in fact the endangered red colobus populations have an annual growth rate of 3%. These finding present a positive conservation message and indicate that the Uganda Wildlife Authority is being effective in managing its biodiversity; however, with constant poaching pressure and changes such as the exponential growth of elephant populations that could cause forest degradation, continued monitoring and modification of conservation plans are needed.     

Intra-population variation in the diet of an avian top predator: generalist and specialist foraging in Great Black-backed Gulls Larus marinus

ABSTRACT

Capsule: Great Black-backed Gulls Larus marinus breeding on Skokholm, UK, fed predominantly on seabirds, rabbits, refuse, and marine prey, with the majority of pairs being dietary generalists, but with some specialist pairs.

Aims: To understand the significance of Great Black-backed Gulls as top predators on a small offshore island with internationally important numbers of breeding seabirds (Skokholm, UK) by quantifying their diet and to determine how this varies within the breeding season, to test for pair-level dietary specialization and to examine the consequences of dietary differences for reproductive performance.

Methods: Regurgitated pellets were collected and analysed from 26 breeding pairs on Skokholm during 2017 and related to breeding success.

Results: Analysis of 1035 pellets revealed that, overall, Great Black-backed Gulls fed on seabirds (48% – mostly Manx Shearwaters Puffinus puffinus), mammals (38% – mostly European Rabbits Oryctolagus cuniculus), anthropogenic waste (7%), and marine prey (7%). Diet varied among pairs with 18 (73%) generalist pairs and 7 (27%) specialist pairs (of which, 5 were bird specialists and 2 were mammal specialists). Diet also varied seasonally, but pair-level dietary diversity was repeatable through the breeding season. Dietary diversity did not covary with breeding success.

Conclusion: Great Black-backed Gulls are top predators on Skokholm. Variation in diet among pairs emphasizes that not all individuals contribute equally in terms of predation. Understanding the incidence of this variation has important ecological implications, particularly where apex predators may exert a strong top-down influence.     

Life history variation in space and time: environmental and seasonal responses of a parthenogenetic invasive freshwater snail in northern Germany

Hydrobiologia - The processes that lead to a successful invasion are complex. Here, we investigated life history characteristics potentially explaining the invasion success of Potamopyrgus...     

River regulation and fish larvae: variation through space and time

1. Patterns in abundance and distribution of larval fish in a heavily regulated and a mildly regulated Australian lowland river were compared over four breeding seasons to gain some insight into how river regulation affects fish populations.
2. Larvae from a total of 13 species from nine families were recorded from the two rivers. The mildly regulated Broken River supported twice as many species as the heavily regulated Campaspe River. The two rivers shared three introduced species but only two native species. The dominant species in the Campaspe was not found in the Broken River.
3. The two most abundant species in the Campaspe were classified as `opportunists'. They are small, short-lived species, which spawn for up to 9 months, encompassing extremes in temperature and flow. The extended spawning period may place a subset of larvae in optimal conditions for recruitment and is hypothesised as being the key to the success of these species.
4. Most species spawned each year, despite large interannual variation in flow and temperature conditions. Poor recruitment over several decades, rather than a failure to spawn, is considered the most likely explanation for differences in the larval fish faunas between the two rivers.
5. The highly regulated section of the Campaspe River downstream of the regulating impoundment is thought to provide suboptimal habitat conditions for larvae relative to the less regulated downstream sections.
6. The timing of occurrence of larvae of the dominant species varied by breeding season and may be the result of flexibility in the timing of spawning.     

Population genomics of marine fishes: identifying adaptive variation in space and time

Studies of adaptive evolution have experienced a recent revival in population genetics of natural populations and there is currently much focus on identifying genomic signatures of selection in space and time. Insights into local adaptation, adaptive response to global change and evolutionary consequences of selective harvesting can be generated through population genomics studies, allowing the separation of the effects invoked by neutral processes (drift-migration) from those due to selection. Such knowledge is important not only for improving our basic understanding of natural as well as human-induced evolutionary processes, but also for predicting future trajectories of biodiversity and for setting conservation priorities. Marine fishes possess a number of features rendering them well suited for providing general insights into adaptive genomic evolution in natural populations. These include well-described population structures, substantial and rapidly developing genomic resources and abundant archived samples enabling temporal studies. Furthermore, superior possibilities for conducting large-scale experiments under controlled conditions, due to the economic resources provided by the large and growing aquaculture industry, hold great promise for utilizing recent technological developments. Here, we review achievements in marine fish genomics to date and highlight potential avenues for future research, which will provide both general insights into evolution in high gene flow species, as well as specific knowledge which can lead to improved management of marine organisms.     

Use of novel pollen species by specialist and generalist solitary bees (Hymenoptera: Megachilidae)

If trade-offs between flexibility to use a range of host species and efficiency on a limited set underlie the evolution of diet breadth, one resulting prediction is that specialists ought to be more restricted than generalists in their ability to use novel resource species. I used foraging tests and feeding trials to compare the ability of a generalist and a specialist solitary mason bee species to collect and develop on two pollen species that are not normally used in natural populations (novel pollens). Osmia lignaria (Hymenoptera: Megachilidae) is a generalist pollen feeder; O. californica, is more specialized. Adults of the specialist were more limited in use of novel hosts, but only in some contexts. Both bee species refused to collect one novel pollen. The specialist accepted a second novel pollen only when it was presented along with its normal pollen, whereas the generalist collected novel pollen whether presented alone or with normal pollen. Surprisingly, larvae of the specialist were more flexible than were generalists. The specialist grew well on mixtures of normal and novel pollen species, in some cases better than on its normal host alone. Larvae of the generalist grew more poorly on all diets containing novel pollens than on their normal host. Data on these two species of bees suggest that specialization by itself need not reduce flexibility on novel hosts. The findings also provide information about mechanisms of specialization in bees. Similar to some folivores, specific cues of the pollen host and the bee's interpretation of these contribute, along with foraging economics, to pollen choice by adults. The ability of the larvae to cope with specific components of one pollen species need not interfere with its ability to use others.     

Disappearing refuges in time and space: how environmental change threatens species coexistence

Understanding the impacts of environmental changes on species survival is a major challenge in ecological research, especially when shifting from single- to multispecies foci. Here, we apply a spatially explicit two-species simulation model to analyze the effects of geographic range shifting and habitat isolation on different coexistence mechanisms. The model explicitly considers dispersal, local competition, and growth on a single resource. Results highlight that both range shifting and habitat isolation severely impact coexistence. However, the strength of these impacts depends on the underlying coexistence mechanisms. Neutrally coexisting species are particularly sensitive to habitat isolation, while stabilized coexistence through overcompensatory density regulation is much more sensitive to range shifting. We conclude that, at the community level, the response to environmental change sensitively depends on the underlying coexistence mechanisms. This suggests that predictions and management recommendations should consider differences between neutral versus stabilized community structures whenever possible.     

Distinctiveness in the face of gene flow: hybridization between specialist and generalist gartersnakes

Patterns of divergence and polymorphism across hybrid zones can provide important clues as to their origin and maintenance. Unimodal hybrid zones or hybrid swarms are composed predominantly of recombinant individuals whose genomes are patchworks of alleles derived from each parental lineage. In contrast, bimodal hybrid zones contain few identifiable hybrids; most individuals fall within distinct genetic clusters. Distinguishing between hybrid swarms and bimodal hybrid zones can be important for taxonomic and conservation decisions regarding the status and value of hybrid populations. In addition, the causes of bimodality are important in understanding the generation and maintenance of biological diversity. For example, are distinct clusters mostly reproductively isolated and co‐adapted gene complexes, or can distinctiveness be maintained by a few ‘genomic islands’ despite rampant gene flow across much of the genome? Here we focus on three patterns of distinctiveness in the face of gene flow between gartersnake taxa in the Great Lakes region of North America. Bimodality, the persistence of distinct clusters of genotypes, requires strong barriers to gene flow and supports recognition of distinct specialist (Thamnophis butleri) and generalist (Thamnophis radix) taxa. Concordance of DNA‐based clusters with morphometrics supports the hypothesis that trophic morphology is a key component of divergence. Finally, disparity in the level of differentiation across molecular markers (amplified fragment length polymorphisms) indicates that distinctiveness is maintained by strong selection on a few traits despite high gene flow currently or in the recent past.     

Claw morphology, prey size selection and foraging efficiency in generalist and specialist shell-breaking crabs

Claw morphology, and claw-closing forces of four species of intertidal crabs from San Juan Island, Washington were compared and related these findings were related to prey size selection, shell breaking times and total handling times on their snail prey, Littorina sitkana Philippi. Two functional groups of crabs emerged: generalists and specialists on hard-shelled prey. The generalist, Hemigrapsus nudus (Dana), has an omnivorous diet and possesses weak claws with small, fine denticles and mechanical advantage (MA) of the claw's lever system <0.3, while the specialists, Lophopanopeus bellus (Stimpson), Cancer oregonensis (Dana) and C. productus (Randall), consume hard-shelled prey and possess large, powerful claws with broad, blunt molars and MA>0.3. The claws of the generalist, H. nudus, exhibited weaker claw closing forces (5 N) than those of similar sized specialists (>12 N). When crabs of similar weight were offered four size categories of Littorina sitkana, the generalist, Hemigrapsus nudus, exhibited a consistent preference for the smallest size categories, while the three specialists attacked all size classes offered. Hemigrapsus nudus took significantly longer (134 s) than the specialists (30–52 s) to break open a 4 mm L. sitkana. This difference in shell-breaking time between the generalist and the specialists increased with increasing prey size. The rate of successful attacks on increasingly larger L. sitkana decreased with prey size in the generalist (70% on 4 mm, 37% on 6 mm, and 0% on 8 mm snails), but remained high in the specialists (70–100%). Strength limitation of the claws is the best hypothesis to explain the avoidance of large snails by the generalist, H. nudus.     

Comparison of environmental and mutational variation in flowering time in Arabidopsis

Developmental dynamics can be influenced by external and endogenousfactors in a more or less analogous manner. To compare the phenotypiceffects of (i) environmental [i.e. standard (stPhP) and extended(exPhP) photoperiods] changes in Arabidopsis wild types and(ii) endogenous genetic variation in eav1–eav61 earlyflowering mutants, two temporal indicators were analysed, thetime to bolting (DtB) and the number of leaves (TLN). It wasfound that DtB and TLN are differentially affected in differentenvironmental and genetic contexts, and some factors of dynamicconvergence were identified. The quantitative response to photoperiodis markedly contingent on the phototrophic input for DtB, butless so for TLN. To discriminate the light quantity and periodcomponents in DtB, two novel temporal indicators were determined,LtB (photosynthetic time to bolting) and PChron (DtB h^–1of photoperiod), respectively. The use of PChron results ina coincidence of the variation profiles across stPhP and exPhP,interpreted as a buffering of the trophic response. Unlike naturalaccessions and later flowering mutants, the variation profilesacross stPhP and eav mutants are significantly divergent, pointingto differences in environmental and genetic variation in floweringtime. Yet, phenocopy effects and dynamic convergence betweenwild-type and mutant profiles are detected by using exPhP andthe LtB indicator. Additional analyses of the cauline leaf number(CLN) show that the apical and basal boundaries of the primaryinflorescence vary co-ordinately. The finding that the correlativitybetween CLN and TLN changes across photoperiods suggests thatdifferent states of intra-connectedness are involved in ontogeneticspecification of flowering time and embodied in the primaryinflorescence. Key words: Arabidopsis, bolting, correlativity, developmental dynamics, flowering time, early flowering mutant, phase change, phenocopy, phenotypic plasticity, photoperiodic response     

Mitochondrial DNA variation in space and time in the northeastern Pacific gastropod, Littorina keenae

The present population structure of a species reflects the influence of population history as well as contemporary processes. To examine the relative importance of these factors in shaping the current population structure of Littorina keenae, we sequenced 762 base pairs of the mitochondrial ND6 and cytochrome b genes in 584 snails from 13 sites along the northeastern Pacific coast. Haplotype network analysis revealed a 'star-like' genealogy indicative of a recent population expansion. Nested clade and mismatch analyses also supported the hypothesis of sudden population expansion following a population bottleneck during the Last Glacial Maximum. Analysis of molecular variance and pairwise Phi(ST) showed no significant spatial population differentiation from Mexico to Oregon - not even across the recognized biogeographic boundary at Point Conception. This is probably due to high contemporary gene flow during the free-swimming larval stage of this snail. Surprisingly, we found a highly significant temporal population differentiation between a San Pedro sample from 1996 and one from 2005, which gave an estimate of effective population size (N(e)) of only 135. Nearly statistically significant changes in the frequency of a particular haplotype in three other populations over 2-3 years further support Hedgecock's 'sweepstakes' hypothesis. When by chance nearly all of the progeny from an aggregation of highly fecund sisters that possess a rare haplotype successfully recruit to become the next generation, the rare haplotype can become temporarily common across the entire species' range. This modification of the sweepstakes hypothesis can explain why the temporal variation that we observed was much greater than the spatial variation.     

Patterns, mechanisms and spatial scale of aggregation in generalist and specialist predatory mites (Acari: Phytoseiidae)

Prey species often distribute themselves patchily in their habitats. In response to this spatial variation in prey density, some predator species aggregate in patches of higher prey density. This paper reviews a series of laboratory experiments to demonstrate the patterns of responses by phytoseiid predators (Phytoseiulus persimilis, Typhlodromus occidentalis and Amblyseius andersoni) to spatial variation in the density of their spider mite prey (Tetranychus urticae) and reveal the behavioural mechanisms underlying the observed patterns. In addition, patterns of aggregation were examined at a variety of spatial scales on plants in greenhouses. The patterns, mechanisms and spatial scale of aggregation in three predatory species are discussed in relation to their varying degrees of polyphagy. The results show that a specialist predator species (1) aggregates more strongly than generalist predators, (2) does so not because it finds prey patches of high density more easily but because it remains in these patches longer than generalist predators and (3) tends to aggregate more often at lower levels of spatial scale than generalist predators. It is suggested that these conclusions, based mainly on laboratory studies of a small sample of species, should be tested in the future on a wider selection of specialist and generalist species at different scales in the field. This revised version was published online in November 2006 with corrections to the Cover Date.