On the relationship between food, reproduction and survival of two carabid beetles: Calathus melanocephalus and Pterostichus versicolor

Abstract.

• 1 The relative influences of temperature and availability of food on reproduction, survival and growth of all developmental stages of two carabid beetle species are discussed with special reference to the suggested relationship between availability of food, size of egg production and survival of adults from one breeding season to the next.
• 2 Temperature as well as food supply influence the length of larval growth and adult body size. Beetles grown at low temperatures and low amounts of food are smaller than those grown at higher temperature and with more food.
• 3 The number of eggs laid per female was correlated with the amount of food gathered. There was no inverse relationship (trade-off) between reproductive output and survival in the field until the next breeding season.
• 4 In 1980 no significant relationship was found between winter mortality and the amounts of food gathered by beetles in the period after reproduction and before winter diapause. However, in 1981 in C. melanocephalus a lower number of starved beetles survived the winter than the fed ones and‘field’beetles.
• 5 Only in the first part of the feeding activity period in autumn can enough food be gathered by C.melunocephalus for successful hibernation. In the second part of this period there is not enough food to build up the fat reserves needed to survive the winter.
• 6 Difference in population fluctuations of both species are discussed in relation to their life histories.

     

Hunger, partial consumption of prey and prey size preference in a carabid beetle

ABSTRACT.

• 1 Partial consumption of prey and prey size preference were studied in females of the carabid Notiophilus biguttatus F., using different size-classes of the springtail Orchesella cincta (L.) as prey.
• 2 Time to ingest a prey increases disproportionately with prey size, mainly as a consequence of satiation.
• 3 During consumption of a prey the predator's rate of ingestion decreases, partly because of diminishing returns from the prey over feeding time.
• 4 The hypothesis that the diminishing returns from the prey induce partial consumption was refuted.
• 5 Partial consumption in the beetle is due to gut limitation; its occurrence depends on prey size.
• 6 Average weight of prey remains in four prey size classes were close to weights expected from average intercatch intervals and estimates of hunger.
• 7 Prey choice depends on level of food deprivation.
• 8 Partial consumption of prey, prey size preference and profitability of prey in relation to hunger of the beetle are discussed.

     

Limitation of reproduction by feeding condition in a carabid beetle,Carabus yaconinus

1. Effects of the amount of food consumed on reproduction of the carabid beetle, Carabus yaconinus B., were studied in the laboratory by rearing beetles at different food levels, and the feeding and oviposition rates in the field were estimated on the basis of the relationships between the amount of food consumed, body weight and egg production obtained in the experiment.
2. The maximum amount of food consumed was 150 mg of minced beef per day. The number of eggs laid per day and the mean body weight increased with an increase in the amount of food consumed. High mortality occurred only when the beetles consumed less than 25 mg of minced beef per day.
3. The ratio of current body weight to the minimum one just before death by starvation, W/W_min, was used for the estimation of the rates of food consumption and egg production. The relationships between mean W/W_min ratio, the amount of food consumed and the number of eggs laid per day were clarified.
4. The relationships between ovary states (ovary weight and the number of mature eggs in the ovary) and W/W_min ratio were examined for the females caught in the field. Females with higher values of W/W_min ratio had more mature eggs.
5. The amount of food consumed by females in the field during the reproductive period was estimated to be 50–70% of the maximum value attained in the experiment and the estimated rate of oviposition was 45–59% of the maximum rate attained in the experiment.

     

Extrinsic vs. intrinsic food shortage and the strength of feeding links: effects of density and food availability on feeding rate of Hyphydrus ovatus

Summary We use field and laboratory experiments to determine whether Hyphydrus ovatus, a predatory aquatic beetle, is food limited, and whether any food shortage results from depletion of prey by these predators (intrinsic food shortage) or is independent of predation by these beetles (extrinsic food shortage). In the laboratory, differences in feeding rate influence body fat content, thus making fat content a useful index of recent feeding history. H. ovatus collected during the breeding season have fat contents significantly greater than those of H. ovatus starved for 25 days, but not significantly different from those of H. ovatus fed ad libitum for 25 days, indicating that natural feeding rates are near the maximum possible. H. ovatus confined at a density 60 times greater than natural show reduced fat content and feeding rate relative to natural, indicating that at very high densities H. ovatus is capable of depleting its prey. Addition of supplemental natural prey (primarily Cladocera) to experimental enclosures resulted in an order of magnitude increase in prey availability, and a significant increase in fat content and feeding rate of confined H. ovatus. Adults of this species do not appear to be food limited during the breeding season, and extraordinarily high densities of adults seem to be necessary to produce intrinsic food shortage. These results suggest that feeding links between H. ovatus an its principal prey do not have major effects on population dynamics under typical field conditions, and call into question the assumption that closely coupled predator-prey interactions are the sole explanation for observed food-web patterns.     

Differentiated egg size of the cannibalistic salamander <Emphasis Type="Italic">Hynobius retardatus</Emphasis>

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients.     

Effect of Population Density and Female Body Size on Number and Size of Offspring in a Species with Size‐Dependent Contests over Resources

When size‐dependent contests over resources influence reproductive success, the trade‐off between number and size of offspring depends on the frequency of contests. Under these circumstances, clutch size should decrease and offspring size should increase as contests become more frequent. We tested these predictions with the burying beetle Nicrophorus pustulatus through manipulation of rearing densities. Burying beetles reproduce on small vertebrate carcasses, a rare but high quality food source for the larvae. Large beetles are more likely to win contests over carcasses and gain exclusive access to a carcass. The winner of a contest kills eggs and larvae already present on a carcass. As a result of the rarity of carcasses, burying beetles are unlikely to breed more than once. As predicted, brood size of N. pustulatus decreased with increasing rearing density. Despite a negative correlation between brood size and larval mass, larval mass did not increase with increasing rearing density. This may be due to the special biology of N. pustulatus which can use snake eggs for reproduction. Potentially larger supply of resources and generally small population densities of N. pustulatus may weaken selection on body size and thus the correlation between brood size and larval mass. As size‐dependent constraints can limit reproductive phenotypes, we examined whether female size influenced reproductive phenotype. Small females produced larger broods with smaller, but more variable, offspring than large females. Mechanical constraints of egg size seem an unlikely explanation for the differences because burying beetles can compensate for small egg size through parental care. Energetic constraints may impact small females because body mass and brood size of small females decreased with increasing density. Yet, at all density levels small females produced larger, not smaller, broods than large females. The larger and more variable broods of small females seem to be in agreement with a bet‐hedging strategy.     

The effect of a foliar disease (rust) on the development of Gastrophysa viridula (Coleoptera: Chrysomelidae)

Abstract.

• 1 Gastrophysa viridula Degeer (Coleoptera: Chrysomelidae) and the pathogenic rust fungus Uromyces rumicis (Schum.) Wint. both occur on leaves of Rumex crispus L. and R.obtusifolius L. Individual stages of beetle development, and egg laying, were compared on healthy and infected leaves of each plant species in the laboratory. Oviposition choice was investigated in the field and laboratory.
• 2 Beetles reared on infected leaves of each species had greater larval mortality and slower development than those reared on healthy leaves. Although larvae feeding on infected leaves consumed up to 2.5 times more dry weight than those reared on healthy leaves, they had a lower relative growth rate and pupated at a lower weight. These changes were consistent with the reduced nutritive quality of rust-infected Rumex leaves.
• 3 Fecundity of beetles reared on infected leaves of both species was considerably reduced. Eggs laid by beetles feeding on infected R.crispus leaves also had a reduced viability.
• 4 The beetle developed consistently poorer on healthy R.crispus than on healthy R.obtusifolius throughout its life-cycle. Differences in larval performance were greater between host species than between infected and healthy leaves.
• 5 Oviposition was similar on infected and healthy R.crispus in both the laboratory and field. However, adults consumed less, and laid fewer eggs on infected than on healthy R.obtusifolius. The pattern of egg laying on different aged leaves was affected by rust infection: a greater proportion of eggs was laid on the older, infected leaves, than on the equivalent aged leaves on the healthy plants. Few larvae survived from eggs laid on rusted leaves in the field.

     

Competition during early ontogeny: Effects of native and invasive planktivores on the growth,survival, and habitat use of bluegill

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Larval population density affects female weight and fecundity in the dung beetle Aphodius ater

1. Competition for food at high densities during larval development leads to reduced adult weight in the northern temperate dung beetle Aphodius ater. 2. Analysis of female beetles caught in the field showed that numbers of eggs and total egg load per female were correlated positively with beetle size. 3. Female beetles reared at different population densities during larval development in the laboratory were analysed with regard to their lifetime fecundity and reproductive lifespan. 4. High population densities during development had a negative influence on the number of eggs per female and on reproductive lifespan. Lifetime fecundity was correlated positively with female weight. 5. It was concluded that competition during larval development in the first generation of offspring will result in a lower number of offspring in the second generation in Aphodius ater, and thereby reduce parental fitness.     

Long adult life spain and polyphagy of a carabid beetle,Leptocarabus kumagaii in relation to reproduction and survival

The reproductive patterns of two different age classes of Leptocarabus kumagaii adults, i.e. young beetles which were in the first year of adult stage and old ones which survived for more than two years, were studied in the field. The numbers of eggs laid were compared between young and old females and the effects of food quality and quantity were studied on egg production and maintenance in the laboratory.

1. Both of the young and old beetles reproduced in the field population. The number of eggs laid did not significantly differ between the two.
2. The number of eggs laid by female increased with an increase in the amount of miced beef eaten. Plant materials were ineffective for egg production but sufficient for the survival of both male and female beetles.
3. The two life historical aspects, i.e. repeated reproduction and polyphagy were discussed in relation to environmental fluctuation and limitation of food resources.

     

A wide window of migration phenology captures inter‐annual variability of favourable conditions: Results of a whole‐lake experiment with juvenile Pacific salmon

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Laboratory study of cannibalism and interspecific predation in ladybirds

Abstract.

• 1 In the absence of aphids, adult females of Adalia bipunctata (L.) showed a greater reluctance to eat eggs than males.
• 2 Eggs and young larvae were more vulnerable to cannibalism than older larvae and starved larvae were more vulnerable than well-fed larvae.
• 3 Both egg and larval cannibalism is inversely related to the abundance of aphids.
• 4 Eggs are a better food, in terms of larval growth and survival, than aphids.
• 5 In the absence of aphids interspecific predation occurred, but not equally, between the coccinellids A.bipunctata, A.decempunctata (L.), Coccinella septempunctata L. and C.undecempunctata L.
• 6 Larvae and adults of A. bipunctata and C.septempunctata were reluctant to eat conspecific eggs painted with a water extract of the other species' eggs and larvae of C. septempunctata were more likely to die after eating a few eggs of A.bipunctata than vice versa.
• 7 These results indicate that cannibalism occurs mainly when aphid prey is scarce and is adaptive in that it improves the chances of survival, and coccinellids, to varying degrees, are defended against interspecific predation.

     

How predacious are predators? A study on Ocypus similis,a rove beetle of cereal fields

It is attempted determine the diet breadth and to quantify the predator pressure by Ocypus similis (F.) (Coleoptera: Staphylinidae), a dominant species of the rove beetle fauna in a winter wheat field. Laboratory and field experiments were conducted to determine the range of prey taxa and the consumption of a whole predator generation (including larvae) during its activity in the field. In laboratory experiments, adults and larvae of O. similis fed on a variety of prey except spiders and larvae of soldier beetles (Cantharidae), but they preferred millipedes as food (Diplopoda: Julidae, Polydesmidae). This preference could also be observed in the field. Although the predator population reached its maximum density in April, highest predator pressure occurred in June because of high consumption rates during the second larval stage. Adult beetles and the individuals of the second larval stage contributed both about 1/3 to total population consumption. Predation by adult beetles was mainly restricted to early spring and summer while larval consumption was highest from April to July. The staphylinids reduced the population density of diplopods by 7–35%. Because millipedes are an important part of the decomposer community, predator activity may lead to a decrease of decomposition rates.     

Observaciones autoecológicas en Liolaemus nitidus (Lacertilia: Iguanidae)

In an attempt to provide background information that may lead to a more comprehensive understanding of the genus Liolaemus, the paper documents some basic features of the autecology of Liolaemus nitidus. The study was carried out at two typical localities where L. nitidus is found: at the coast north of Santiago de Chile, and at the Andes ranges east of Santiago. The following parameters were investigated: microhabitat utilization, preferred body temperatures, diet, weight of gonads and of fat bodies, and frequency of regenerated tails. The main findings reported are:

1. Liolaemus nitidus is preferentially a rock‐dweller, although, when available, it can also be found perching on branches of the bromeliad Puya chilensis.

2. Preferred field body temperatures, evaluated with a quick‐registering Schultheis thermometer, indicate that L. nitidus has a preferendum of about 35.4°C. This mean preferred body temperature is not statistically different from the laboratory preferendum reported in the literature.

3. The diet of L. nitidus was found to vary with body size. Small individuals tend to eat mostly insects, whereas larger individuals tend to have more plant material in their guts. Total gut content is nevertheless relatively heavier for small than for large individuals. These facts are interpreted in the light of previous results obtained with Liolaemus species, and in relation to food availability. Regarding food availability, the above results are consistent with the hypothesis that arthropods are relatively scarce in the xerophytic areas where L. nitidus lives, and therefore these lizards complement their diet with plant material. Arthropod content of the guts was analyzed by size and taxonomic order. The findings are that mean prey size, and size of the largest and of the smallest prey in each stomach tend to be positively correlated with body size of the lizard, thus suggesting prey selectivity. On the other side, the orders most preyed upon by L. nitidus are Coleopterans, Hymenopterans (fundamentally ants), and Homopterans.

4. Correlations between either relative gonad weight or relative weight of the fat bodies, versus dry weight of the lizards are significant and positive. These results suggest that L. nitidus is a relatively K‐selected species, where there is a premium for somewhat delayed reproduction and perhaps larger clutches. Heavier fat bodies in the mountain population are interpreted in terms of length of the growing season and different needs to accumulate energy reserves for the unfavorable season.

5. Frequency of regenerated tails is greater in adults than in juveniles, but does not differ between sexes, suggesting that risk of predation would be more associated with the span of time the lizard is exposed to the environment than with differential behavior of males and females.

Results are interpreted and compared, when available, with data for other Liolaemus species.

These comparisons suggest that L. nigromaculatus and L. chiliensis, two species of similar size and ecological requirements as to L. nitidus, could be among the potentially severest competitors in limiting the geographic distribution of this species.     

Diving beetle offspring oviposited in amphibian spawn prey on the tadpoles upon hatching

In highly ephemeral freshwater habitats, predatory vertebrates are typically unable to become established, leaving an open niche often filled by macroinvertebrate predators. However, these predators are faced with the challenge of finding sufficient food sources as the rapid rate of desiccation prevents the establishment of extended food chains and limits the number of prey species present. It could therefore be advantageous for predators to synchronize their phenology with that of their prey within sites of extreme ephemerality. We report the first case of adult diving beetles (Hydaticus parallelus) ovipositing their eggs within spawn of an amphibian species (sandpaper frog, Lechriodus fletcheri). This behavior was found among several pools used by L. fletcheri for reproduction. Beetle eggs oviposited in frog spawn were found to hatch within 24 h of the surrounding L. fletcheri eggs, with the larvae becoming voracious consumers of the hatched tadpoles. Although it has yet to be established experimentally whether this is an adaptive behavior, the laying of eggs among potential future tadpole prey in this instance should confer significant fitness benefits for the offspring upon hatching, ensuring that they are provided an immediate source of food at the start of their development and potentially throughout. This oviposition behavior might be common among diving beetles and could form a significant predatory threat for amphibians with a free‐swimming larval stage in ephemeral freshwater habitats.     

Diet, fecundity and egg size in some polyphagous predatory carabid beetles

Egg size was measured at different rates of egg laying in three polyphagous carabid species, known to be useful predators of cereal aphids; the small Bembidion lampros Herbst and the medium-sized Pterostichus cupreus L. and P. melanarius Illiger. Variations in fecundity, as well as the ability of the medium-sized species to also build up fat reserves, were obtained when beetles were subjected to different dietary regimes consisting of aphids, or foods with a lower or higher protein content. Egg size was found to be dependent on the rate of egg laying within a species. A diet of cereal aphids appeared to be adequate for egg production in these polyphagous carabids, but female P. cupreus were unable to build up fat reserves when they ingested aphids contaminated with the aphicide pirimicarb. Beetles were able to devote resources to more and larger eggs (B. lampros), or to larger eggs and/or fat reserves (P. melanarius/P. cupreus) when given access to a carbohydrate-rich food with low protein content. The highest rate of egg laying was obtained when female P. cupreus and P. melanarius were given a more varied diet at frequent intervals; including regular shifts between unsprayed aphids, carbohydrate-rich food and protein-rich maggots. Within the varied diet treatment a negative relationship was obtained between egg size and egg number among similar-sized individuals of P. cupreus and P. melanarius; females producing the largest number also laid the smallest eggs. Egg size affected larval survival, since first instars hatching from large eggs were found to survive longer than those hatching from small eggs. The influence of differences in food intake on reproduction, maintenance metabolism, and survival of fieldinhabiting carabids is discussed.     

To hatch or not to hatch? Egg hatch response to larval density and to larval contact in a treehole mosquito

Abstract.

• 1 We investigated the effect on egg hatch of exposure to: (1) varying larval density, and (2) larval contact in Aedes triseriatus Say (Diptera: Culicidae). For 2 days in the laboratory we submerged eggs into a treehole water medium containing 0 (control), 4, 12 or 24 larvae that could either contact the eggs directly or were separated from them by a screen. Following treatment, abundance of microorganisms on the egg surfaces, a food source for newly hatched larvae and a proposed hatching stimulus, was assessed by counts made from serial dilutions of samples.
• 2 We discovered a complex hatching response to larval contact and to larval density, and an interaction between these two factors in their effect on microbial growth. Hatching was inhibited in the 0-larva control, even though microorganisms grew abundantly on the eggs. Hatch rate, as well as microbial counts, were high for eggs in direct contact with 4 larvae. As density increased in the larval contact treatment, microorganisms disappeared from the egg surfaces and hatch rate declined.
• 3 When protected from larval grazing, eggs supported numerous microbial colonies irrespective of larval density. In contrast to the contact treatment group, egg hatch increased with increasing larval density. These observations suggest that the combination of microbial growth and a larval factor stimulates hatch. This hatching response may have evolved because both abundant microorganisms and numerous larvae reflect a habitat of good quality.

     

The larval diet of three anisopteran (Odonata) species

SUMMARY.

• 1 Comparisons between larval diets of three anisopteran species (Anax imperator, Aeshna cyanea and Libellula depressa) showed that their food intake changed depending on: (1) predator species, (2) time of year, and (3) developmental stage.
• 2 Although this last factor is not so important, the mean size of prey items and the range of prey species eaten by Auax imperator and Aeshna cyanea larvae increased with predator size.
• 3 Comparisons between prey availability and diets indicated differential selectivity by these predators.

     

Pollen as an alternative food for Harmonia axyridis

The current study examines the potential of the multicoloured Asian lady beetle Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) to use pollen as a food to sustain development and reproduction in the absence of insect prey. Three populations of H. axyridis were used in this study: a long-term laboratory population (since 1998) and a melanic and non-melanic population originating from field collected individuals in Belgium. The insects were allowed to develop and reproduce on frozen eggs of Ephestia kuehniella Zeller (Lepidoptera: Phycitidae), frozen moist bee pollen or an even mixture of the two. Females of the field population offered the mixed diet initiated oviposition sooner than those fed only E. kuehniella eggs, but other developmental and reproductive traits were similar on these diets. A diet of pollen alone allowed 35–48% of the larvae of the field population of H. axyridis to successfully reach adulthood. However, developmental time for these individuals was prolonged by 31–49% and adult body weight was reduced by 37–68%, compared to individuals offered the diets containing E. kuehniella eggs. When fed exclusively on pollen in their larval and adult life, about 40% of the adult females of either field population were able to produce a small number of viable eggs. The laboratory and field strains differed in their response to diet for a number of developmental and reproductive traits. The exploitation of pollen and other plant foods at times when insect prey is scarce, may offer a further competitive advantage to the non-indigenous coccinellid H. axyridis over native European predatory lady beetles that share the same niche and are less capable of using pollen as an alternative food.     

A bioenergetic assessment of the influence of stocking practices on rainbow trout (Oncorhynchus mykiss) growth and consumption in a New Zealand lake

Summary

• 1 To investigate the carrying capacity and factors affecting growth of rainbow trout in Lake Rotoiti, we employed a bioenergetics model to assess the influence of stocking rates, timing of releases and prey abundance on growth and prey consumption. We hypothesised that stocking rates and prey abundance would affect growth and prey consumption by influencing per‐capita prey availability, and that the environmental conditions encountered by fish at the time of stocking would affect growth and consumption.
• 2 Prey consumption of stocked rainbow trout was calculated with the Wisconsin bioenergetics model. We calculated growth trajectories of released trout based on data from stocked trout that were released in spring and autumn from 1993 to 2009 and then re‐captured by anglers. Diet, prey energy density, body mass lost during spawning and lake temperature were measured locally.
• 3 Stocking timing had no effect on return rates to anglers or length or weight of caught fish. Although trout released in autumn were smaller than those released in spring, autumn‐released trout grew at a faster rate and had similar lengths and weights to spring cohorts after 2 years of growth in the lake. Modelled consumption parameters were negatively correlated with trout population size, suggesting that stocking rates (347–809 fish ha^?1 year^?1) caused density‐dependent effects on growth. Although common smelt (Retropinna retropinna) accounted for 85% of total prey consumption, no significant relationship was found between prey consumption by individual trout and adult smelt abundance, possibly because trout are targeting smaller smelt that our abundance estimate did not account for.
• 4 Releasing trout in autumn appears to be advantageous for growth, possibly because (i) temperature is more suitable for growth in autumn–winter than in spring–summer and (ii) prey for small trout is abundant in autumn. Mild winter conditions appear to enhance overwinter survival and growth of rainbow trout in warm‐temperate lakes compared to higher latitudes. This implies that moderately productive warm‐temperate lake ecosystems are highly suitable for trout growth in winter, but less so in summer, when lake stratification and high nutrient levels may create conditions suitable for algal blooms and hypolimnetic deoxygenation. High growth rates of trout in warm‐temperate lakes can therefore be supported by timing releases to coincide with favourable winter conditions.