Host choice and fitness correlates for conopid flies parasitising bumblebees

Two parasitoid flies,Physocephala rufipes andSicus ferrugineus (Diptera, Conopidae), and their hosts,Bombus spp., coexist at various locations in northwestern Switzerland. A detailed field study showed that both conopid species use the hostB. pascuorum to a similar degree, while the hostB. terr-luc (a pooled category ofB. terrestris andB. lucorum) is more frequently parasitised than expected byS. ferrugineus. The hostB. lapidarius in turn is exclusively used byP. rufipes. Furthermore, hosts ofB. terr-luc andB. pascuorum parasitised byS. ferrugineus were larger than hosts parasitised byP. rufipes, or than those not parasitised. The findings suggest thatS. ferrugineus selects larger hosts and may displaceP. rufipes. Pupal weight, a predictor of adult body size and parasitoid fecundity, is positively correlated with host size and larger pupae are more likely to emerge, while host species had no effect on the probability of emergence in either conopid species. Host species affected pupal weight inS. ferrugineus, but not inP. rufipes, althoughP. rufipes grew larger in hosts of a given size. Daughters emerged from larger pupae than males, but this did not correlate with larger host sizes. These observations add to the scarce knowledge of dipteran parasitoids.     

Fitness of Asian corn borer, Ostrinia furnacalis (Lepidoptera: Crambidae) reared in an artificial diet

The Asian corn borer, Ostrinia furnacalis (Guenée) (Lepidoptera: Crambidae) is an economically important pest of corn. Finding simple, cheap, and suitable rearing techniques of O. furnacalis is an urgent need to support research for management of this insect. This research aimed to determine the suitability of a read bean and rice bran-based artificial diet used for mass rearing of this insect since 2009.The tested artificial diet was compared with the natural diet (sweet corn kernel) and each diet was tested in individual rearing method (one larva in each vial). The criteria used to justify the quality of diet and mass rearing procedure were based on the fitness of O. furnacalis. The degree of fitness was based on life history, growth, and development. In general, the fitness parameteres observed from O. furnacalis reared in the artificial diet at 25.7?±?1.6?°C with 57.7?±?3.8% RH, and L12:D12 were similar than those in the natural diet.Therefore, the existing artificial diet and rearing procedure were considered suitable and qualified for O. furnacalis. It is important to periodically check the laboratory colony to ensure that they have similar fitness to those found in the natural population.     

The parasitoidOoencyrtus nezarae (Hymenoptera: Encyrtidae) prefers hosts parasitized by conspecifics over unparasitized hosts

Summary Two laboratory experiments were conducted to examine the ovipositional preferences of the egg parasitoidOoencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) for parasitized and unparasitizedMegacopta punctatissimum Montandon (Hemiptera: Plataspidae). Females that had never oviposited or that had not oviposite for 3 days preferred recently parasitized hosts more than unparasitized hosts. The preference for recently parasitized hosts appeared to be mediated by the punctures in already parasitized hosts made by the ovipositor of the first female. Survival of the parasitoid progeny was lower in recently parasitized hosts than in unparasitized hosts. However, handling time of parasitized hosts was extremely short relative to that of unparasitized hosts, because the superparasitizing female could use the punctures made by the previous females. It is concluded that the females preferred the parasitized hosts over unparasitized hosts because the benefit of saving time and energy for drilling was more than the cost of progeny survival.     

Optimal foraging and fitness in Columbian ground squirrels

Summary Optimal diets were determined for each of 109 individual Columbian ground squirrels (Spermophilus columbianus) at two sites in northwestern Montana. Body mass, daily activity time, and vegetation consumption rates for individuals were measured in the field, along with the average water content of vegetation at each ground squirrel colony. I also measured stomach and caecal capacity and turnover rate of plant food through the digestive tract for individuals in the laboratory to construct regressions of digestive capacity as a function of individual body mass. Finally, I obtained literature estimates of average daily energy requirements as a function of body mass and digestible energy content of vegetation. These data were used to construct a linear programming diet model for each individual. The model for each individual was used to predict the proportion of two food types (monocots and dicots) that maximized daily energy intake, given time and digestive constraints on foraging. Individuals were classified as optimal or deviating, depending on whether their observed diet was significantly different from their predicted optimal diet. I determined the consequences of selecting an optimal diet for energy intake and fitness. As expected, daily energy intake calculated for deviators (based on their observed diet proportion) was less than that for optimal foragers. Deviating foragers do not appear to compensate for their lower calculated energy intake through other factors such as body size or physiological efficiency of processing food. Growth rate, yearly survivorship, and litter size increase with calculated energy intake, and optimal foragers have six times the reproductive success of deviators by age three. Optimal foraging behavior, therefore, appears to confer a considerable fitness advantage.     

Are digestive characteristics important contributors to the profitability of prey?

Summary In the field, Spinachia fed on four types of prey; copepods, isopods, mysids and amphipods. As fish size increased, mysids gradually succeeded amphipods as the most important food type in the diet. Prey dimensions and morphometry of the fish's mouth most accurately predicted capture efficiency for amphipods, whereas for mysids capture efficiency was determined by the prey's escape response and the fish's fast-start capability. Responses to model prey revealed the ability of fish to differentiate among contrasting prey characteristics, resulting in the adoption of appropriate predatory tactics. Amphipods were associated with a shorter gut evacuation time than mysids, although approximately equal proportions of energy were absorbed from each. Similar rations of mysids and amphipods were required to satiate fish. The lower energy content per unit dry mass of amphipods was off set by their lower water content. From pre-digestive behaviour, we predicted that mysids were more profitable than amphipods, and this was reflected in the fish's choice. Conversely, incorporating the net rate of energy uptake by the gut led us to predict that amphipods were more profitable. Although physiological constraints clearly influence the net rate of energy uptake, it appears that dietary preferences are based on pre-digestive predatory behaviour and hence on time minimisation.     

A further note on the estimation of a fitness function from two samples taken from a popolation

A modification is proposed for the moments method for estimating a fitness function that was discussed in an earlier paper (Manly , 1976b). This modification ensures that fitness estimates are always positive. The theory has been worked out for a fitness function of any number of variables and a computer program has been written to carry out the iterative calculations needed for estimation.     

Wolf spider feeding strategies: optimality of prey consumption in Pardosa hortensis

Feeding behaviour of the wolf spider Pardosa hortensis Thorell (Araneae, Lycosidae) was studied in the laboratory. Characteristics of feeding were measured while prey availability was increased and the results were compared with the predictions of three models: the marginal value theorem (MVT), gut limitation theory (GLT) and the digestion rate limitation model (DRL). As a result of more frequent encounters with prey, the wolf spiders were able to modify their feeding behaviour so that their net energy intake rate increased substantially. Handling time decreased by 30%, and consumption rate increased by 40%. Partial consumption of prey did not occur until the spiders became nearly satiated. This indicated that spiders did not reach the optimum predicted by MVT. The most plausible mechanism for the increased efficiency was prey-stimulated digestive enzyme production as suggested in DRL. The predictions of GLT were not applicable for most of the feeding session, though gut satiation had an influence on the final stages of feeding. P. hortensis seemed to apply a responsive but cautious strategy: (i) spiders improved feeding efficiency on entering the higher quality habitat, but (ii) feeding times appeared to be sub-optimal and (iii) spiders were also willing to continue feeding when, as they approached satiation, the previously high efficiency could not be maintained. Such feeding behaviour optimizes long-term energy intake when food is scarce and unpredictable, which corresponds well with the known degree of natural food limitation of these animals.     

Mathematical modeling of colony formation in algal blooms: phenotypic plasticity in cyanobacteria

In this paper, we analyzed a mathematical model of algal-grazer dynamics, including the effect of colony formation, which is an example of phenotypic plasticity. The model consists of three variables, which correspond to the biomasses of unicellular algae, colonial algae, and herbivorous zooplankton. Among these organisms, colonial algae are the main components of algal blooms. This aquatic system has two stable attractors, which can be identified as a zooplankton-dominated (ZD) state and an algal-dominated (AD) state, respectively. Assuming that the handling time of zooplankton on colonial algae increases with the colonial algae biomass, we discovered that bistability can occur within the model system. The applicability of alternative stable states in algae-grazer dynamics as a framework for explaining the algal blooms in real lake ecosystems, thus, seems to depend on whether the assumption mentioned above is met in natural circumstances.     

Fish-induced variation in abdominal spine length of Leucorrhinia dubia (Odonata) larvae?

The lengths of dorsal and lateral spines on abdominal segments 4, 6 and 9 were measured on last-instar larvae of Leucorrhinia dubia (Odonata) from seven natural lentic systems containing fish and nine systems lacking fish. Larvae from systems with fish had significantly longer spines than larvae from systems without fish. In contrast, lake/pond area and pH had no effect on the length of spines. The length of the spines was not correlated with larval size, but there was a high correlation between the length of the three spines measured. Also, abundances of L. dubia larvae differed between systems, being significantly lower in systems with fish. Laboratory experiments showed that perch (Perca fluviatilis) handled long-spined larvae for significantly longer times than short-spined larvae. The results suggest that fish predators may induce morphological defences in insects.     

Insect herbivores as potential causes of mortality and adaptation in gallforming insects

Summary Recent studies have suggested that plant galls benefit only the insects living in them and not the host plants, and that galls are induced by insects primarily to improve the plant as a microenvironment or a food source. The potential advantage to insects of protection from their predators and parasitoids has been considered unclear and perhaps minor in importance. However, the potential threat to gallforming insects from other insect herbivores has usually been relatively neglected. This paper notes literature and observations which suggest that herbivores may either consume or be deterred by galls. Even soft leaf galls produced by Hormaphis and Phylloxera aphids appeared to deter some herbivores in the field. The threat of herbivory to galls might help explain general patterns of gall ecology and morphology, and deserves closer attention.     

Biological control through provision of additional food to predators: a theoretical study

In this paper we analyze a variation of a standard predator-prey model with type II functional response which represents predator-prey dynamics in the presence of some additional food to the predator. The aim is to study the consequences of providing additional food on the system dynamics. We conclude that handling times for the available foods play a key role in determining the eventual state of the ecosystem. It is interesting to observe that by varying the quality and quantity of additional food we can not only control and limit the prey, but also limit and eradicate the predators. In the context of biological pest control, the results caution the manager on the choice of quality and quantity of the additional food used for this purpose. An arbitrary choice may have opposite effects leading to increase in pest concentration and eradication of the predator. This study offers insight into the possible management strategies that involve manipulation of quality and supply level of additional food to predators, for the benefit of biological control. The theoretical conclusions agree with results of some practical biological control experiments.     

Efficiency and establishment of three introduced parasitoids of the mealybug Paracoccus marginatus (Hemiptera: Pseudococcidae)

Three introduced parasitoids Acerophagus papayae, Anagyrus loecki, and Pseudleptomastix mexicana of the mealybug Paracoccus marginatus have been released in Miami-Dade and Broward counties (Florida) in 2003. Efficiency and establishment of these previously introduced parasitoids to control P. marginatus were measured in 2005 and 2006, at three locations in Homestead (Miami-Dade). Mealybug populations were initially established on three branches (per plant) of 10 hibiscus plants inside closed-sleeve cages. The three treatments, closed, open, and no-cage environments were applied to the three established mealybug populations on each plant. The number of mealybugs and natural enemies was monitored in all treatments. After 72 h, the number of P. marginatus in the open-sleeve cage was higher than in the no-cage treatment but both were lower than the number in the closed-sleeve cage. Efficiency of parasitoids was evaluated by their percentage parasitism. Percentage parasitism (=recovery) was also used as an indicator for parasitoid establishment. Two of the three previously released parasitoids (A. papayae and A. loecki) were recovered in this study. Acerophagus papayae had a higher percentage parasitism than A. loecki in both the open-sleeve cage (31.0% vs 2.3%) and the no-cage treatments (21.4% vs 1.6%) and caused the most mortality of P. marginatus. There was no recovery of P. mexicana in either of the treatments. Although both A. papayae and A. loecki were established in tested areas, A. papayae was more efficient in controlling P. marginatus than A. loecki. The reasons for not recovering P. mexicana in release areas is yet unknown.     

Size-related consumption of Scirpus maritimus tubers by greylag geese Anser anser explained by their functional response

To explain the preference of wintering greylag geese Anser anser for small Scirpus maritimus tubers (<10mm) over larger ones, our hypothesis was that the former would provide higher intake rates. This 'consumption rate hypothesis' was tested experimentally by deriving the functional responses of geese feeding on tubers of three contrasting sizes. Goose consumption rates were measured as: (i) feeding rate (tubers/min) and (ii) instantaneous intake rate (g fresh weight/min) on various tuber densities (5-200 tubers/tray). Geese had linear functional responses over the range of tuber densities offered, and tuber size affected their consumption rate. The results were then used to re-examine intake rates according to relative biomass of tubers found in the wild. In support of our hypothesis, the larger tubers allowed the lower intake rates. Foraging mechanisms that possibly control the tuber ingestion rate of geese were explored. After controlling (through chemical analyses) that nutritional components of tubers do not differ between sizes, it was concluded that geese preference is mainly due to size-related constraints (i.e. handling time) on their ingestion rates. Additional limiting factors (tuber extraction, digestion capacity) which are likely to constrain food intake of wild geese are also discussed.     

Climate impacts on blacklegged tick host-seeking behavior

The nymph of the blacklegged tick (Ixodes scapularis), the primary North American vector of the causative agent of Lyme disease, must attach to a host by the end of its questing season in order to feed and subsequently molt into an adult. The proper timing of this behavior is critical both for the tick’s survival and for perpetuating the transmission of tick-borne pathogens. Questing also depletes limited nymphal lipid reserves and increases desiccation risk. Given this tradeoff, questing behavior and its environmental influences can be expressed in a dynamic state variable model. We develop what we believe to be the first such model for a tick, and investigate the influence of climate on nymph fitness predictions. We apply these results to the hypothesized inland migration of I. scapularis from island refugia, evaluating fitness under suboptimal questing strategies and uncertain environmental conditions.     

Behavioural fitness component effects of the alba polymorphism of Colias (Lepidoptera, Pieridae): resource and time budget analysis

1. Alba, a white wing colour morph of butterflies of the genus Colias , makes fitness-related changes at several levels of phenotypic organization: physiology, development and behaviour. Two sympatric species of Colias are studied, which differ greatly in their frequency of alba vs its sister yellow/orange morph. How resource and time constraints on Colias interact with the morphs' different patterns of pupal resource allocation to alter the balance of the morphs' fitness components in the two species is discussed.
2. These species, C. alexandra and C. scudderi , differ in melanin-based solar energy absorption, in larval dietary richness and in local adult nectar resources. The two female morphs determine alternate thermal balance, internal resource allocation and behavioural effects in each species, thus changing the morphs' time budgets and fitness-component impacts between the species. In particular, female egg output differences between the morphs appears to reverse between species: alba fecundity is greater than yellow fecundity in C. scudderi , but alba is less than yellow in C. alexandra .
3. These differences are consistent with the large observed differences in alba frequency between the species. Some important questions about the selective regime maintaining a polymorphism here, rather than an alternation of monomorphisms, remain.     

Effects of patch scale on density-dependence and species-dependence in two host-parasitoid systems

Summary Data from two host-parasitoid communities were analyzed to ascertain whether patch scale affected the kinds of correlations existing between 1) spatial differences in host density and the intensity of parasitism (density-dependence) and 2) number of species of parasitoids and the intensity of parasitism (species-dependence). We concluded that parasitization rates are usually independent of both host density and number of parasitoid species present regardless of patch scale. Therefore, the responses of parasitoids to host density and the addition of parasitoid species to a community are equally unpredictable in outcome.