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.     

Increased genetic differentiation in a specialist versus a generalist bee: implications for conservation

Oligolectic bees are specialists that collect pollen from one or a few closely related species of plants, while polylectic bees are generalists that collect pollen from both related and unrelated species of plants. Because of their more restricted range of floral hosts, it is expected that specialists persist in more isolated populations than do generalists. We present data on the population structure of two closely related bee species sampled from a super abundant floral host in the southern Atacama Desert. Pairwise comparisons of population subdivision over identical distances revealed that the specialist bee had significantly more differentiated populations in comparison to the generalist. Further, populations of the specialist had significantly less genetic variation, measured as observed and expected heterozgyosity, than those of the generalist. Our data support the hypothesis of decreased gene flow among populations of the specialist bee even at equivalent geographic distances. The resulting reductions in effective population size for specialists make them particularly prone to extinction due to both demographic and genetic reasons. Our findings have important implications for the conservation of bees and other specialist insects. Deceased     

The functional trophic role of lotic primary consumers: generalist versus specialist strategies

1. The relative extent of generalist or specialist resource use strategies is an important question in ecology. A community dominated by specialist strategies suggests a high level of interspecific competition for resources, resulting in the evolutionary development of isolating mechanisms between species (e.g. resource specialization to avoid and/or outcompete other species). A community dominated by generalist strategies suggests less interspecific competition for resources, allowing many taxa to utilize the same resources. In stream systems, generalist food habits are a common strategy among primary consumers, but little is known about resource assimilation strategies (resources incorporated into tissue growth). Published data indicate that generalist resource assimilation strategies may prevail in lotic systems as well.
2. Functional feeding groups (FFGs) are often used to infer resource assimilation among lotic macroinvertebrates (e.g. shredder-detritivore, scraper-herbivore). While these groupings are aptly used to describe invertebrate feeding modes and community structure, the use of FFGs to describe resource assimilation among lotic consumers is not appropriate. Sufficient data now exist to seriously question how accurately FFG assignments describe the processes of energy flow and material transfer between trophic levels in stream ecosystems.
3. Because FFGs may not accurately describe functional attributes in lotic systems, an alternative approach is needed. One approach is to determine the amount of secondary production that is derived from autochthonous (e.g. periphyton and algae) and allochthonous (e.g. detritus) resources directly. A simple model of community function based on this approach is presented. The model incorporates trophic generalists into measurement of consumer–resource energetics in lotic systems.     

Intraspecific variation for resource use in Drosophila

Individuals within natural populations of Drosophila tripunctata and D. falleni varied significantly in their responses to alternative types of food baits; no such variation was detected in D. putrida. The food type at which laboratory-reared D. melanogaster tended to settle in the field was influenced by the type of food on which they had been kept prior to release. Although flies apparently forgot their previous experience within hours, its effect on their behaviour can be manifested for at least 1 day in the field. Mark -recapture experiments on wild flies are shown to be very inefficient at detecting individual variation in behaviour.     

Dealing with stochastic environmental variation in space and time: bet hedging by generalist,specialist, and diversified strategies

Building on previous work, we derive an optimization model for a two-state stochastic environment and evaluate the fitnesses of five reproductive strategies across generations. To do this, we characterize spatiotemporal variation and define grain (=patch) size as the scale of fitness autocorrelation. Fitness functions of environmental condition are Gaussian. The strategies include two specialists on each of the environmental conditions; two generalists that each fare equally well under both conditions, but one (a conservative bet hedger) optimizes the shape of the fitness function; and a diversified bet hedger producing an optimal mix of the two specialists within individual broods. When the environment is primarily in one of the two states, the specialist on that state achieves the highest fitness. In the more interesting situation where the two environments are equally prevalent in the long term, with low-moderate environmental variation, a generalist strategy (that copes with both states well) does best. Higher variation favors diversified bet hedgers, or surprisingly, specialists, depending mainly on whether spatial or temporal variation predominates. These strategies reduce variance in fitness and optimize the distribution of offspring among patches differently: specialists by spreading offspring among many independently varying patches, while diversified bet hedgers put all offspring into a few patches or a single patch. We distinguish features consistent with strategies like diversified bet hedgers that spread risk in time from features linked to strategies like specialists that spread risk in space. Finally, we present testable hypotheses arising from this study and suggest directions for future work.     

Geographic variation in feeding preference of a generalist herbivore: the importance of seaweed chemical defenses

The ecological impacts of generalist herbivores depend on feeding preferences, which can vary across and within herbivore species. Among mesoherbivores, geographic variation in host use can occur because host plants have a more restricted geographic distribution than does the herbivore, or there is local evolution in host preference, or both. We tested the role of local evolution using the marine amphipod Ampithoe longimana by rearing multiple amphipod populations from three regions (subtropical Florida, warm-temperate North Carolina and cold-temperate New England) and assaying their feeding preferences toward ten seaweeds that occur in some but not all regions. Six of the ten seaweeds produce anti-herbivore secondary metabolites, and we detected geographic variation in feeding preference toward five (Dictyota menstrualis, Dictyota ciliolata, Fucus distichus, Chondrus crispus and Padina gymnospora, but not Caulerpa sertularioides). Amphipod populations that co-occur with a chemically-rich seaweed tended to have stronger feeding preferences for that seaweed, relative to populations that do not co-occur with the seaweed. A direct test indicated that geographic variation in feeding preference toward one seaweed (D. ciliolata) is mediated by feeding tolerance for lipophilic secondary metabolites. Among the four seaweeds that produce no known secondary metabolites (Acanthophora, Ectocarpus, Gracilaria and Hincksia/Feldmannia spp.), we detected no geographic variation in feeding preference. Thus, populations are more likely to evolve greater feeding preferences for local hosts when those hosts produce secondary metabolites. Microevolution of feeding behaviors of generalist marine consumers likely depends on the availability and identity of local hosts and the strength of their chemical defenses.     

Differential habitat use promotes sustainable coexistence between the specialist otter and the generalist mink

Optimal foraging and habitat selection theories predict that heterogeneous environments should favour the coexistence of competitors, especially when the dominant competitor is a specialist and the sub-ordinate is a generalist. In this paper, we analysed differential habitat use as a potential mechanism for the coexistence of two competing riparian mammals, the specialist and dominant Eurasian otter ( Lutra lutra ) and the generalist and sub-ordinate American mink ( Mustela vison ). We tested three hypotheses: H1: mink coexist with otters for longer in areas with abundance of habitats hosting terrestrial prey because, by not relying on aquatic prey, mink can segregate from its competitor. H2: the characteristics of the habitat closer to the riverbank will affect the length of time the two species coexist, because mink are still tied to the water even in the presence of otters. H3: denser vegetative cover along the bank increases the duration of coexistence of mink and otters because it reduces the frequency of their encounters. The first hypothesis was supported by the data and we found that in areas where terrestrial prey was abundant mink coexisted for longer with otters. The second hypothesis was also supported by the data and the characteristics of the habitat closer to the riverbank were the most important in determining coexistence time. Finally, we did not find supporting evidence for the third hypothesis. This study provides strong evidence that habitat heterogeneity plays an important role in determining the likelihood of coexistence of American mink with Eurasian otters. This result is particularly important from a conservation standpoint. Mink are invasive and a threat to endangered species in parts of their range. The knowledge that mink have a higher chance to persist in the presence of otters when terrestrial prey is abundant should be used to target areas for preferential mink management.     

Plant neighborhood influences colonization of Brassicaceae by specialist and generalist aphids

A plant's own characteristics, but also those of its neighbors, might have an impact on its probability of being colonized by herbivorous insects. A plant might be less colonized and experience associational resistance when it grows near repellent neighbors. In contrast, it might be more colonized and experience associational susceptibility near attractive neighbors. To date, mechanisms that drive associational defense are not really understood. In order to gain insights into the occurrence of associational resistance versus associational susceptibility under field conditions, we conducted an experiment to determine the influence of neighboring plants on the colonization of a focal plant by aphids. The focal plant was always Brassica oleracea. The neighbors were B. oleracea (control), B. napus, B. nigra, or Solanum lycopersicum, which represent contrasting levels of physical and chemical defenses. The focal plant, B. oleracea, was more colonized by the specialist aphid Brevicoryne brassicae, and experienced associational susceptibility when it was surrounded by B. nigra or B. napus. In contrast, B. oleracea was less colonized by the generalist aphid Myzus persicae, and experienced associational resistance when it was surrounded by S. lycopersicum, B. nigra or B. napus. Neighboring plants had no significant impact on host plant choice by the generalist aphid Macrosiphum euphorbiae. In conclusion, attraction or repulsion of the specialist aphid B. brassicae and the generalist aphid M. persicae by B. nigra, B. napus, and S. lycopersicum resulted in associational susceptibility or associational resistance for B. oleracea.     

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.     

Individual variation in resource use by opossums leading to nested fruit consumption

Despite recent findings on the ecological relevance of within population diet variation far less attention has been devoted to the role diet variation for ecological services. Seed dispersal is a key ecological service, affecting plant fitness and regeneration based on foraging by fruit‐eating vertebrates. Here we used a network approach, widely used to understand how seed‐dispersal is organized at the species level, to gain insights into the patterns that emerge at the individual‐level. We studied the individual fruit consumption behavior of a South American didelphid Didelphis albiventris, during the cool–dry and warm–wet seasons. In species–species networks the heterogeneity in specialization levels generates patterns such as nestedness and asymmetry. Because generalist populations may be comprised of specialized individuals, we hypo thesized that network structural properties, such as nestedness, should also emerge at the individual level. We detected variation in fruit consumption that was not related to resource availability, ontogenetic or sexual factors or sampling biases. Such variation resulted in the structural patterns often found in species–species seed‐dispersal networks: low connectance, a high degree of nestedness and the absence of modules. Moreover structure varied between the warm–wet and cool–dry seasons, presumably as a consequence of seasonal fluctuation in fruit availability. Our findings suggest individuals may differ in selectivity causing asymmetries in seed dispersal efficiency within the population. In this sense the realized dispersal would differ from the expected dispersal estimated from their average dispersal potential. Additionally the results suggest possible frequency‐dependent effects on seed dispersal that might affect individual plant performance and plant community composition.     

Effects of oil droplets by Pieris caterpillars against generalist and specialist carnivores

Pieris rapae larvae secrete small oil droplets from their dorsal setae, which adhere to objects that touch them. The function of the droplets was studied in terms of both generalist and specialist predators. We tested the function of the droplets against ants under laboratory and field conditions. In the laboratory observation, Formica japonica ants that touched the larvae with their antennae initiated antennal cleaning and did not prey on the larvae. In the field, predation pressure by ants on larvae with oil droplets was significantly lower than on larvae from which oil droplets were removed. Thus, we concluded that the droplets had a defensive function. By contrast, oviposition by Cotesia glomerata, the specialist parasitic wasp of P. rapae larvae, was not affected by the presence of oil droplets. Furthermore, the wasps exhibited searching behavior and oviposition behavior towards filter paper which had been impregnated with the droplets substance, suggesting that the oil functions as a host-searching cue and an oviposition stimulant for C. glomerata. According to these results, the functions of the droplets are discussed with regard to the prey–predator interaction.     

Comparison of learning in related generalist and specialist eucoilid parasitoids

Effects of learning in two microhabitat specialists, Leptopilina boulardia Barbotin et al. and L. fimbriata Kieffer were compared to previous and new results of learning in the microhabitat generalist L. heterotoma Thomson. Females were given one or more oviposition experiences on hosts in different types of substrate. In all species oviposition experience affected the choice for a substrate, although this effect of learning was considerably less in L. fimbriata compared to the other two species. Patch times, known to be highly determined by experience in the generalist L. heterotoma, were much less flexible in the specialists. L. boulardi and L. fimbriata have fixed patch times on their natural substrate and have variable patch times on other substrates only. In all three species one oviposition affected the choice for a substrate. Additional ovipositions showed no different effect. An accumulative effect of the number of ovipositions on patch times was found in L. heterotoma only. Retention of the learning effect was only studied in L. boulardi, and was shown to be similar to that reported for L. heterotoma, i.e. two to three days. Although learning was found in both the generalist and the specialist species studied, it seems to affect their foraging behaviour differently.     

Resource assurance predicts specialist and generalist bee activity in drought

Many short-lived desert organisms remain in diapause during drought. Theoretically, the cues desert species use to continue diapause through drought should differ depending on the availability of critical resources, but the unpredictability and infrequent occurrence of climate extremes and reduced insect activity during such events make empirical tests of this prediction difficult. An intensive study of a diverse bee–plant community through a drought event found that bee specialists of a drought-sensitive host plant were absent in the drought year in contrast to generalist bees and to specialist bees of a drought-insensitive host plant. Different responses of bee species to drought indicate that the diapause cues used by bee species allow them to reliably predict host availability. Species composition of the bee community in drought shifted towards mostly generalist species. However, we predict that more frequent and extended drought, predicted by climate change models for southwest North America, will result in bee communities that are species-poor and dominated by specialist species, as found today in the most arid desert region of North America.     

Testing assumptions of the enemy release hypothesis: generalist versus specialist enemies of the grass Brachypodium sylvaticum

The enemy release hypothesis (ERH) suggests greater success of species in an invaded range due to release from natural enemies. The ERH assumes there will be more specialist enemies in the native range and that generalists will have an equal effect in both ranges. We tested these assumptions with the grass Brachypodium sylvaticum in the native range (Switzerland) and invaded range (Oregon, USA). We assessed all the kinds of damage present (caused by fungi, insects, mollusk and deer) on both leaves and seeds at 10 sites in each range and correlated damage with host fitness. Only two of the 20 fungi found on leaves were specialist pathogens, and these were more frequent in the native range. Conversely there was more insect herbivory on leaves in the invaded range. All fungi and insects found on seeds were generalists. More species of fungi were found on seeds in the native range, and a higher proportion of them were pathogenic than in the invaded range. There were more kinds of enemies in the native range, where the plants had lower fitness, in accordance with the ERH. However, contrary to assumptions of the ERH, generalists appear to be equally or more important than specialists in reducing host fitness.     

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.     

An ecological genomic approach challenging the paradigm of differential plant responses to specialist versus generalist insect herbivores

A general prediction of the specialist/generalist paradigm indicates that plant responses to insect herbivores may depend on the degree of ecological specialization of the insect attacker. However, results from a single greenhouse experiment evaluating the responses of the model plant Arabidopsis thaliana to three specialist (Plutella xylostella, Pieris rapae, and Brevicoryne brassicae) and three generalist (Trichoplusia ni, Spodoptera exigua, and Myzus persicae) insect species did not support the previous prediction. Using an ecological genomic approach, we assessed plant responses in terms of herbivore-induced changes in genome-wide gene expression, defense-related pathways, and concentrations of glucosinolates (i.e., secondary metabolites that are ubiquitously present in cruciferous plants). Our results showed that plant responses were not influenced by the degree of specialization of insect herbivores. In contrast, responses were more strongly shaped by insect taxa (i.e., aphid vs. lepidopteran species), likely due to their different feeding modes. Interestingly, similar patterns of plant responses were induced by the same insect herbivore species in terms of defense signaling (jasmonic acid pathway), aliphatic glucosinolate metabolism (at both the gene expression and phenotypic levels) and genome-wide responses. Furthermore, plant responses to insect herbivores belonging to the same taxon (i.e., four lepidopteran species) were not explained by herbivore specialization or phylogenetic history. Overall, this study suggests that different feeding modes of insect taxa as well as herbivore-specific plant responses, which may result from distinct ecological/evolutionary interactions between A. thaliana (or a close relative) and each of the lepidopteran species, may explain why observed responses deviate from those predicted by the specialist/generalist paradigm.     

Geographic variation in sperm production by Trinidadian guppies

Population comparisons yield important insights into adaptive differentiation. However, despite the current interest in sperm competition and spermatogenesis, geographic variation in these traits has received little attention. We tested the hypothesis that sperm production covaries with risk by comparing five natural populations of Trinidadian guppies (Poecilia reticulata): two that inhabited dangerous Crenicichla localities, two from low-risk Rivulus sites and a ''transplant'' population comprised of the descendants of guppies moved from one of the high-risk sites to a low-predation environment. As predicted, males from the three low-risk sites performed significantly more courtship displays and had larger sperm reserves than their high-risk counterparts. This result implies higher rates of sperm competition in the low-risk sites.     

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.