Age‐specific maternal effects interact with larval food supply to modulate life history in Coleomegilla maculata

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A sex-specific quantitative genetic theory for life history and development

A quantitative genetic framework is developed to examine life history evolution with sex-specific differences in morphology, demographic parameters, and selection differentials occurring. Age-specific selection differentials are partitioned according to whether mediated through differential fertility, survival, or mating success, permitting the derivation of weightings according to selection mechanism as well as individual age and sex. The relationship of present to previously gained results is then examined, with applications to the evolution of bimaturism and evolutionary conservativeness of survival rate.     

The numerical response: rate of increase and food limitation in herbivores and predators

Two types of numerical response function have evolved since Solomon first introduced the term to generalize features of Lotka-Volterra predator-prey models: (i) the demographic numerical response, which links change in consumer demographic rates to food availability; and (ii) the isocline numerical response, which links consumer abundance per se to food availability. These numerical responses are interchangeable because both recognize negative feedback loops between consumer and food abundance resulting in population regulation. We review how demographic and isocline numerical responses have been used to enhance our understanding of population regulation of kangaroos and possums, and argue that their utility may be increased by explicitly accounting for non-equilibrium dynamics (due to environmental variability and/or biological interactions) and the existence of multiple limiting factors. Interferential numerical response functions may help bridge three major historical dichotomies in population ecology (equilibrium versus non-equilibrium dynamics, extrinsic versus intrinsic regulation and demographic versus isocline numerical responses).     

Quantitative genetic analysis of responses to larval food limitation in a polyphenic butterfly indicates environment‐ and trait‐specific effects

Different components of heritability, including genetic variance (V_G), are influenced by environmental conditions. Here, we assessed phenotypic responses of life‐history traits to two different developmental conditions, temperature and food limitation. The former represents an environment that defines seasonal polyphenism in our study organism, the tropical butterfly Bicyclus anynana, whereas the latter represents a more unpredictable environment. We quantified heritabilities using restricted maximum likelihood (REML) procedures within an “Information Theoretical” framework in a full‐sib design. Whereas development time, pupal mass, and resting metabolic rate showed no genotype‐by‐environment interaction for genetic variation, for thorax ratio and fat percentage the heritability increased under the cool temperature, dry season environment. Additionally, for fat percentage heritability estimates increased under food limitation. Hence, the traits most intimately related to polyphenism in B. anynana show the most environmental‐specific heritabilities as well as some indication of cross‐environmental genetic correlations. This may reflect a footprint of natural selection and our future research is aimed to uncover the genes and processes involved in this through studying season and condition‐dependent gene expression.     

Immune deployment increases larval vulnerability to predators and inhibits adult life‐history traits in a dragonfly

While deploying immune defences early in ontogeny can trade‐off with the production and maintenance of other important traits across the entire life cycle, it remains largely unexplored how features of the environment shape the magnitude or presence of these lifetime costs. Greater predation risk during the juvenile stage may particularly influence such costs by (1) magnifying the survival costs that arise from any handicap of juvenile avoidance traits and/or (2) intensifying allocation trade‐offs with important adult traits. Here, we tested for predator‐dependent costs of immune deployment within and across life stages using the dragonfly, Pachydiplax longipennis. We first examined how larval immune deployment affected two traits associated with larval vulnerability to predators: escape distance and foraging under predation risk. Larvae that were induced to mount an immune response had shorter escape distances but lower foraging activity in the presence of predator cues. We also induced immune responses in larvae and reared them through emergence in mesocosms that differed in the presence of large predatory dragonfly larvae (Aeshnidae spp.). Immune‐challenged larvae had later emergence overall and lower survival in pools with predators. Immune‐challenged males were also smaller at emergence and developed less sexually selected melanin wing coloration, but these effects were independent of predator treatment. Overall, these results highlight how mounting an immune defence early in ontogeny can have substantial ecological and physiological costs that manifest both within and across life stages.     

A shared mechanism of defense against predators and parasites: chitin regulation and its implications for life‐history theory

Defenses against predators and parasites offer excellent illustrations of adaptive phenotypic plasticity. Despite vast knowledge about such induced defenses, they have been studied largely in isolation, which is surprising, given that predation and parasitism are ubiquitous and act simultaneously in the wild. This raises the possibility that victims must trade‐off responses to predation versus parasitism. Here, we propose that arthropod responses to predators and parasites will commonly be based on the endocrine regulation of chitin synthesis and degradation. The proposal is compelling because many inducible defenses are centered on temporal or spatial modifications of chitin‐rich structures. Moreover, we show how the chitin synthesis pathway ends in a split to carapace or gut chitin, and how this form of molecular regulation can be incorporated into theory on life‐history trade‐offs, specifically the Y‐model. Our hypothesis thus spans several biological scales to address advice from Stearns that “Endocrine mechanisms may prove to be only the tip of an iceberg of physiological mechanisms that modulate the expression of genetic covariance”.     

Effects of temperature during early life history on embryonic and larval development and growth in haddock

The effect of incubation temperature (2, 4, 6, 8 and 10° C) on haddock Melanogrammus aeglefinus development and growth during the embryonic period and in subsequent ontogeny in a common post‐hatch thermal environment (6° C) was investigated. Hatching times were inversely proportional to incubation temperature and ranged from 20·3 days at 2° C to 9·1 days at 10° C. Growth rates were directly proportional to incubation temperature during both the embryonic and larval periods. There was a significant decline in growth rates following hatch in all temperature groups. Compared to the endogenously feeding embryos, growth rates in the exogenous period declined by 4·4‐fold at 4° C to 3·9‐fold at 8° C, indicative of the demarcation between the endogenous and exogenous feeding periods. Yolk utilization varied from 17 days at 2° C to 6 days at 10° C and followed a three‐stage sigmoidal pattern with the initial lag period inversely proportional to incubation temperature. Time to 50% yolk depletion varied inversely with temperature but occurred 1–1·5 days post‐hatch at all temperatures. Additionally, the period between 10 and 90% yolk depletion also decreased with increased temperature. Overall developmental rate was sequential with and directly proportional (2·3‐fold increase) to incubation temperature while the time spent in each developmental stage was inversely proportional to temperature. Larger embryos tended to be produced at lower temperatures but this pattern reversed following hatch, as larvae from higher temperature groups grew more rapidly than those from other temperature groups. Larvae from all temperatures achieved a similar length (c.total length 4·5 mm) upon complete yolk absorption. The study demonstrated the significant impact that temperature has upon developmental and growth rates in both endogenous and exogenous feeding periods. It also illustrated that temperature changes during embryogenesis had significant and persistent effects on growth in subsequent ontogeny.     

Effects of scarcity and excess of larval food on life history traits of Aedes aegypti (Diptera: Culicidae)

Few studies have assessed the effects of food scarcity or excess on the life history traits of Aedes aegypti (L.) (Diptera: Culicidae) independently from larval density. We assessed immature survival, development time, and adult size in relation to food availability. We reared cohorts of 30 Ae. aegypti larvae from newly hatched to adult emergence with different food availability. Food conditions were kept constant by transferring larvae each day to a new food solution. Immature development was completed by some individuals in all treatments. The shortest development time, the largest adults, and the highest survival were observed at intermediate food levels. The most important effects of food scarcity were an extension in development time, a decrease in the size of adults, and a slight decrease in survival, while the most important effects of food excess were an important decrease in survival and a slight decrease in the size of adults. The variability in development time and adult size within sex and treatment increased at decreasing food availability. The results suggest that although the studied population has adapted to a wide range of food availabilities, both scarcity and excess of food have important negative impacts on fitness.     

Diel and seasonal patterns of prey available to epigeal predators: Evidence for food limitation in a linyphiid spider community

Linyphiid spiders, as part of the community of natural enemies that frequent agroecosystems, can exert significant pressure on prey populations. Many aspects of linyphiid feeding ecology remain understudied, including temporal and seasonal patterns of prey utilization. To quantify the diversity, quantity, and spatial pattern of availability over diel and seasonal gradients, we monitored prey in an alfalfa crop, using web-site specific sticky traps. Although there were no differences in total number of prey between the day and night, significantly more Collembola were intercepted nocturnally rather than diurnally. Conversely, Diptera and Aphididae were significantly more abundant diurnally. Overall there were more prey at web-sites than in the general crop environment, indicating that spiders were selectively constructing webs where prey was relatively more abundant. However, spiders still may have experienced a high degree of prey limitation because the abundance of Collembola, an important detrital prey, was low over much of the season and below levels calculated as being required to sustain growth and reproductive output in linyphiid spiders. In conclusion, our data suggests that linyphiid spiders, in order to avoid food limitation, may benefit by foraging throughout the diel cycle, consuming herbivorous prey in addition to their core diet of detrital prey. This link between above and below ground food webs indicates the importance of understanding the trophic interactions between epigean spiders and their prey if they are to be integrated into future biological control programs.     

Fungal plant endosymbionts alter life history and reproductive success of aphid predators

Endosymbionts occur in most plant species and may affect interactions among herbivores and their predators through the production of toxic alkaloids. Here, we ask whether effects of mycotoxins produced by the symbiosis of the fungal endophyte Neotyphodium lolii and the grass Lolium perenne are transmitted to the aphidophagous ladybird Coccinella septempunctata when feeding on cereal aphids Rhopalosiphum padi on infected plants. The larval development of coccinellids was extended, while their survival was reduced when feeding exclusively on aphids from infected plants. Ladybirds developing on aphids from infected plants showed reduced fecundity and impaired reproductive performance. Body size and symmetries of ladybird adults were not affected by the endophytes. Consistently strong, negative effects of endophytes on the lifetime performance of ladybirds indicates that mycotoxins are transmitted along food chains, causing significant damage for top predators. Such cascading effects will influence the population dynamics of aphid predators in the long term and could feedback to the primary plant producers.     

Coping with an unpredictable and stressful environment: the life history and metabolic response to variable food and host availability in a polyphagous tephritid fly

The way energy resources are used under variable environmental conditions lies at the heart of our understanding of resource management and opportunism in many organisms. Here we sought to determine how a time-limited, synovigenic and polyphagous insect with a high reproductive-potential (Anastrephaludens), copes behaviourally and metabolically with environmental unpredictability represented by constant and variable regimes of host availability and variation in food quality. We hypothesized that an adaptive response to a windfall of nutritious food would be the rapid accumulation of energy metabolites (whole body lipids, glycogen and proteins) in the female. We also studied patterns of oogenesis as an indicator of egg-reabsorption under stressful environmental conditions. As predicted, patterns of energy metabolites were mainly driven by the quality and temporal pattern of food availability. In contrast, patterns of host availability had a lower impact upon metabolites. When given constant access to high quality nutrients, after an initial increase early in life, whole body lipids and glycogen were regulated downward to a steady-state level and somatic protein levels did not vary. In contrast, when food uncertainty was introduced, whole body lipid, glycogen and protein oscillated sharply with peaks associated with pulses of high-quality food. Production of eggs was highest when offered continuous access to hosts and high quality food. Importantly, females fully recovered their reproductive capacity when fruit became available following a period of host deprivation. With no evidence of egg resorption and high levels of egg dumping, it appears that egg dumping may favour the continuous production of eggs such that the female’s reproductive tissues are ready to respond to rapid changes in the availability of hosts. Our results exemplify the capacity of insects to maximize reproduction under variable and stressful environmental conditions.     

The non-lethal effects of predators and the influence of food availability on life history of adult Chironomus tentans (Diptera: Chironomidae)

1. We used a laboratory experiment to determine effects of a predator (other than straightforward prey consumption) and food availability, during the larval stage, on adult size, age at emergence and fecundity of Chironomus tentans. 2. Predator presence and decreased food availability resulted in reduced adult emergence. 3. Predator presence and food availability resulted in smaller size and greater age at emergence of male and female chironomids. There were no significant interactions between effects of predator presence and food availability. 4. Predator presence had no significant effect on fecundity. 5. Smaller size and greater age at emergence can have important implications for adult survival, size and age at first reproduction and, thus, intrinsic rate of population growth.     

The adaptive value of gluttony: predators mediate the life history trade‐offs of satiation threshold

Animals vary greatly in their tendency to consume large meals. Yet, whether or how meal size influences fitness in wild populations is infrequently considered. Using a predator exclusion, mark–recapture experiment, we estimated selection on the amount of food accepted during an ad libitum feeding bout (hereafter termed ‘satiation threshold’) in the wolf spider Schizocosa ocreata. Individually marked, size‐matched females of known satiation threshold were assigned to predator exclusion and predator inclusion treatments and tracked for a 40‐day period. We also estimated the narrow‐sense heritability of satiation threshold using dam‐on‐female‐offspring regression. In the absence of predation, high satiation threshold was positively associated with larger and faster egg case production. However, these selective advantages were lost when predators were present. We estimated the heritability of satiation threshold to be 0.56. Taken together, our results suggest that satiation threshold can respond to selection and begets a life history trade‐off in this system: high satiation threshold individuals tend to produce larger egg cases but also suffer increased susceptibility to predation.     

Life‐history traits of Tubastraea coccinea: Reproduction,development, and larval competence

The sun coral Tubastraea coccinea Lesson, 1829 (Dendrophylliidae) is a widely distributed shallow‐water scleractinian that has extended its range to non‐native habitats in recent decades. With its rapid spread, this coral is now one of the main invasive species in Brazil. Its high invasive capability is related to opportunistic characteristics, including several reproductive strategies that have allowed it to disperse rapidly and widely. To better understand the reproductive biology of T. coccinea and aid in developing management strategies for invaded areas, we investigated aspects of its reproductive performance and life cycle, including the effects of colony size, seawater temperature and salinity, and lunar periodicity on offspring production and larval metamorphosis competence. A total of 18,139 offspring were released in different developmental stages, mainly from the larger colonies, which also produced larvae with longer competence periods. The main reproductive peak occurred during the First Quarter and New Moon phases and was highest in water temperatures around 26°C. Together, these results help to explain the rapid expansion of T. coccinea into non‐native habitats such as the Caribbean and southwestern Atlantic, and will inform actions of the recent Brazilian National Plan for the prevention, eradication, control, and monitoring of sun corals.     

Nutrient limitation and soil development: Experimental test of a biogeochemical theory

Walker & Syers (1976) proposed a conceptual model that describesthe pattern and regulation of soil nutrient pools and availability during long-term soil and ecosystem development. Their model implies that plantproduction generally should be limited by N on young soils and by P on oldsoils; N and P supply should more or less equilibrate onintermediate-aged soils. We tested the application of this model to nutrientlimitation, using a well characterized substrate age sequence in Hawaiianmontane rain forest. Earlier experiments had evaluated nutrient limitationin forests on a young (300 y) and an old (4,100,000 y) substrate on the samedevelopmental sequence; N alone limited tree growth on the youngsubstrate, while P alone did so on the old one. An additional fertilizerexperiment based on replicated treatments with N, P, and all othernutrients combined, applied in individually and in all factorialcombinations, was established in an intermediate-aged site in theLaupahoehoe Forest Reserve, Hawaii. Here, diameter increments of thedominant tree Metrosideros polymorpha increased slightly with Nadditions, and nearly doubled when N and P were added together.Additions of elements other than N and P had no significant effecton growth. These results show that N and P had equilibrated (relativeto plant requirements) in the intermediate aged site. Together withthe earlier experiments, these results suggest that the Walker and Syersmodel provides a useful starting point for explaining the nature anddistribution of nutrient limitation in forest ecosystems.     

Food limitation in a wild cyclopoid copepod population: direct and indirect life history responses

We studied the effects of food limitation on the population dynamics of the freshwater cyclopoid copepod Diacyclops thomasi in Oneida Lake, New York. In the field population, maximum juvenile abundance coincided seasonally with high phytoflagellate concentration. During the clear-water phase (a seasonal period of low algal density), D. thomasi disappeared from the water column, but fourth-instar copepodids (CIV) were found encysted in developmental arrest in the sediment. Laboratory assays of the effect of the density of two types of food on copepod life history parameters showed that temporal variation in the concentration of relatively small phytoflagellates significantly affected stage-specific development times. This food limitation was most pronounced during the clear-water phase, and supplementation of the diet with a laboratory-cultured phytoflagellate, Chlamydomonas, prevented food limitation. Although developmental arrest appears to be controlled primarily by photoperiod, availability of the larger, more mobile food, Euglena, also influenced the percentage of individuals entering developmental arrest in the laboratory. An investigation of the spatial and temporal emergence pattern in the field revealed that CIV copepodids started to emerge in late autumn and that emergence rates were significantly greater at deep-water locations (9–12 m water depth) compared with shallow-water locations (5–7 m). The clear-water phase in Oneida Lake is an annual event, probably produced by intense grazing by Daphnia pulicaria and Daphnia galeata. Food limitation is thus very likely a recurrent phenomenon for D. thomasi. This apparent seasonal competitive impact of Daphnia on Diacyclops affects both nauplii and immature copepodids. Diacyclops shows two types of responses to the food limitation: (1) the physiological response of slowed active development, and (2) the adaptive response of developmental arrest. Received: 3 November 1997 / Accepted: 1 March 1998     

Microhabitat selection by larval Chironomus tentans (Diptera: Chironomidae): effects of predators, food, cover and light

1. As part of a study designed to estimate the developmental costs of antipredator behaviour of larval chironomids, we used laboratory experiments to study effects of food and factors that could influence predation risk [presence of fish, cover from fish (simulated debris) and light level], on microhabitat selection by Chironomus tentans larvae in the third and fourth instar. 2. Larvae were more likely to build tubes where there was more food although their ability to move far to find food appeared limited. 3. Larvae did not avoid areas with fish and the presence of fish did not alter larval response to food. 4. Larvae avoided areas of cover (simulated debris) but cover did not alter larval response to food. 5. When provided with a choice between light and dark areas, larvae initially without tubes were found more often in the dark areas. Light level had no effect on location of larvae that had begun the experiment with tubes. 6. Results suggest the tubicolous life-style of larval Chironomus tentans limits their ability to select microhabitats that could alter their risk of predation.     

Phenotypic plasticity for plant development, function and life history

A single genotype can produce different phenotypes in different environments. This fundamental property of organisms is known as phenotypic plasticity. Recently, intensive study has shown that plants are plastic for a remarkable array of ecologically important traits, ranging from diverse aspects of morphology and physiology to anatomy, developmental and reproductive timing, breeding system, and offspring developmental patterns. Comparative, quantitative genetics and molecular approaches are leading to new insights into the adaptive nature of plasticity, its underlying mechanisms and its role in the ecological distribution and evolutionary diversification of plants.