Diet affects male gonad maturation,female fecundity,and larval development in the granivorous ground beetle Anisodactylus punctatipennis

Abstract 1. Granivory is a specialised food habit in the Carabidae and is considered to have evolved from ancestral carnivory. Despite recent investigations, the mechanisms underlying this peculiar feeding shift have not been sufficiently elucidated. In particular, no studies have examined the effects of diet composition on male reproductive traits. 2. This study examined male gonad maturation, female fecundity, and larval development under different diets (insect larvae, mixed seeds, and insect larvae + mixed seeds) for Anisodactylus punctatipennis (Coleoptera: Carabidae). 3. Whereas sperm‐bundle length, the durations of larval stages, and adult weight did not differ among diets, the weights of seminal substance‐producing organs, fecundity, and larval survival were higher with diets containing seeds compared to the pure‐animal diet. These findings provide the first conclusive evidence for granivory in Anisodactylus. 4. This study is the first that demonstrates the effects of diet on male reproductive traits in a granivorous carabid. The results were consistent with similar observations for fecundity and larval performance in this and other studies. Thus, in addition to the female and larval traits, the dietary effects on male reproductive traits may also have played an important role in the evolution of granivory in the Carabidae.     

Field observations of climbing behavior and seed predation by adult ground beetles (Coleoptera: Carabidae) in a lowland area of the temperate zone

Granivory is a specialized food habit in the predominantly carnivorous beetle family Carabidae. Most studies of carabid granivory have been conducted under laboratory conditions; thus, our knowledge of the feeding ecology of granivorous carabids in the field is insufficient. I conducted field observations of climbing behavior and seed predation by adult carabids in a lowland area of eastern Japan, from early October to late November in 2008. This is the first systematic field observation of the feeding ecology of granivorous carabids in the temperate zone. In total, 176 carabid individuals of 11 species were observed, with 108 individuals feeding on plant seeds/flowers. Each carabid species was primarily observed feeding on a particular plant species. Frequently observed combinations were: Amara gigantea Motschulsky on Humulus scandens (Loureiro) Merrill (Moraceae) seed, Amara lucens Baliani on Artemisia indica Willdenow (Asteraceae) flower, and Amara macronota (Solsky) and Harpalus (Pseudoophonus) spp. on Digitaria ciliaris (Retzius) Koeler (Poaceae) seed. In all but one species, the sex ratio of individuals observed feeding was female-biased. In Am. gigantea and Am. macronota, a larger proportion of females than males ate seeds. In the three Amara species, copulations on plants, with the female feeding on its seeds/flowers, were often observed. These observations may indicate that, whereas females climb onto plants to feed on seeds, males climb to seek females for copulation rather than forage. Because granivorous carabids play important roles as weed-control agents in temperate agro-ecosystems, the present results would provide valuable basic information for future studies on this subject.     

Carabid larvae as predators of weed seeds: granivory in larvae of Amara eurynota (Coleoptera: Carabidae)

Up to date we do not have much information about predation on seeds by larvae of ground beetles. One of the reasons why such knowledge is important is that granivorous larvae contribute to predation of weed seeds. In this study, the food requirements of larvae of autumn breeding carabid species Amara eurynota (Panzer) were investigated in the laboratory and a hypothesis, that they are granivorous was tested. Insect diet (Tenebrio molitor larvae), three seed diets (seeds of Artemisia vulgaris, Tripleurospermum inodorum or Urtica dioica or a mixed diet (T. molitor + A. uulgaris) were used as food. For larvae of A. eurynota, seeds are essential for successful completion of development, because all those fed pure insect diet died before pupation. However, differences in suitability were observed between pure seed diets. Larvae fed seeds of A. vulgaris had the lowest mortality and fastest development of the seed diets. Those fed seeds of T. inodorum had also low mortality, but the development was prolonged in the third instar. In contrast, development of larvae reared on seeds of U. dioica was slowest of the tested diets and could not be completed, as all individuals died before pupation. When insects were included to seed diet of A. vulgaris (mixed diet), the duration of development shortened, but mortality remained the same when compared to seed diet of A. vulgaris. According to the results it was concluded that larvae of A. eurynota are granivorous. A mixed diet and seed diets of A. uulgaris and T. inodorum were suitable and insect diet and seeds of U. dioica were unsuitable diets in this experiment.     

Larval and adult seed consumption affected by the degree of food specialization in Amara (Coleoptera: Carabidae)

Many carabid beetles (Coleoptera: Carabidae) are known to feed on plant seeds, but the level of specialization on this food differs. This is the first study in which seed consumption is assessed for all larval instars and adults of ground beetles. Three species of Amara with syntopic occurrence, Amara aenea (DeGeer), Amara familiaris (Duftschmid) and Amara similata (Gyllenhal), were examined. Larvae of all three instars and adults were fed seeds of Stellaria media (L.) Vill., Capsella bursa‐pastoris (L.) Med. and Taraxacum officinale Wick. ex Wigg. in a laboratory no‐choice experiment. In general, larvae, particularly the first instar, showed greater differences in seed consumption than the adults, although the latter showed similar but less marked pattern. Amara aenea consumed all offered seed diets in all life stages. All three larval instars of granivorous A. familiaris almost exclusively fed on seeds of S. media and the adults also ate significantly more of this than other seeds. Amara similata consumed mostly seeds of C. bursa‐pastoris in the first instar and adult stages, whereas the larvae of the later instars seemed to be unspecialized on particular seed diet. Differences in seed‐specific consumption between larval instars in granivorous carabids are reported for the first time. The results provide further support for the parallel evolution of various degrees of granivory in the genus Amara, which may ultimately facilitate species coexistence. The daily seed consumption by the larvae was comparable or (in case of the third instar) even higher than that by the adults. Hence, we suggest that larvae may be the important consumers of seed in the field and should not be forgotten when seed predation is assessed.     

Laboratory studies on the larval food habits of the ground beetle Amara (Curtonotus) gigantea (Motschulsky) (Coleoptera: Carabidae: Zabrini)

Ground beetles (Coleoptera: Carabidae) are important components of soil ecosystems in temperate zones, but our knowledge regarding the ecology of many species is limited. This study examined larval food habits in the ground beetle Amara (Curtonotus) gigantea (Motschulsky) using laboratory‐rearing experiments. Because this beetle is a member of the tribe Zabrini, which includes several species that exhibit granivory during larval and/or adult stages, three diet types were tested, with starvation as a control: Tenebrio larvae, mixed seeds (Bidens pilosa (Asteraceae), Setaria spp., Digitaria ciliaris (both Poaceae), and Humulus scandens (Moraceae)), and Tenebrio larvae + mixed seeds. Because of high larval mortality during overwintering under laboratory‐rearing conditions, survival and the duration of development through pre‐overwintering stages (first and second instars) were compared. Larvae fed the diet of Tenebrio larvae + mixed seeds showed the highest survival (89%), followed by the diet of Tenebrio larvae (83%), but the difference between the two treatments was not significant; similarly, developmental durations did not differ between these two diets. Larvae fed the mixed‐seeds diet showed markedly lower survival, and no larvae reached the second instar. Therefore, for A. gigantea larvae, animal food results in high larval performance, whereas seeds provide a low value diet; animal food is essential for larval development, but seeds provide no benefit, even as a dietary supplement. Overall, the present results suggest that A. gigantea larvae are omnivorous but with a mainly carnivorous food habit.     

Food preferences and the value of animal food for the carabid beetle Amara similata (Gyll.) (Col., Carabidae)

Abstract:  Several studies have shown that the mainly granivorous carabid beetles, e.g. Amara spp., include animal food in their diet to a considerable extent. We therefore hypothesized that the performance of these beetles would be enhanced by dietary mixing including both seeds and animal food. In order to test this, we conducted laboratory feeding experiments with adults and larvae of Amara similata . Both adults and larvae were subjected to different diet treatments including: seeds, houseflies, grasshoppers, earthworms, slugs and snails in pure and mixed diets. Larval survival, development time, pupal and teneral weights were used as indicators of food quality for the larvae. For the adult beetles, mass change was used as an indicator of food quality. We found seeds to be high-quality food, while all pure animal diets were of low quality for both adults and larvae. Animal foods added to the seed diet had both positive and negative effects. A mixed diet of all foods enhanced the mass gain of adults compared with the seed diet, but reduced larval performance dramatically. Earthworms and grasshoppers added to seeds increased the pupal and teneral weights, while reduced larval survival. Thus, A. similata is omnivorous with a mainly granivorous feeding habit. It may gain benefits on some fitness parameters and incur costs on others from preying or scavenging on animal food. Therefore, the overall fitness consequences of a mixed seed-animal diet are uncertain.     

Contrary food requirements of the larvae of two Curtonotus (Coleoptera: Carabidae: Amara) species

The larvae of carabids that are granivorous as adults can be granivorous, omnivorous or carnivorous. The differences in larval food preferences of Amara aulica and Amara convexiuscula, two closely related species of the subgenus Curtonotus, were studied. Survival and duration of development of non‐diapausing first and second instar larvae were compared. The third instar larvae were not studied because they go into diapause prior to pupation. The larvae were fed diets consisting of insect larvae (Tenebrio molitor), seeds of Artemisia vulgaris, Tripleurospermum inodorum, Urtica dioica or Cirsium arvense and a mixed diet consisting of insect larvae and seed. Larvae of A. aulica required seeds for successful development, whereas those of A. convexiuscula developed on both an insect diet and seeds of Artemisia. The difference in larval food requirements facilitates the coexistence of these closely related species, which frequently share the same habitat.     

Feeding ecology of granivorous carabid larvae: a stable isotope analysis

Although most carabids are primarily carnivorous, some carabid species are omnivorous, with mainly granivorous feeding habits during the larval and/or adult stages (granivorous carabids). This feeding habit has been established based on laboratory and field experiments; however, our knowledge of the feeding ecology of these beetles in the field is limited owing to the lack of an appropriate methodology. In this study, we tested the utility of stable isotope analysis in investigations of the feeding ecology of granivorous carabids in the field, using two closely related syntopic species, Amara chalcites and Amara congrua. We addressed two issues concerning the feeding ecology of granivorous carabids: food niche differentiation between related syntopic species during the larval stage and the effect on adult body size of supplementing seeds with an animal diet during the larval stage. To investigate larval feeding habits, we analysed newly emerged adults, most somatic tissues of which are considered of larval origin. In the two populations examined, both δ¹⁵N and δ¹³C were significantly higher in A. chalcites than A. congrua, suggesting that the two species differentiate food niches, with A. chalcites larvae being more carnivorous than A. congrua larvae. The two isotope ratios of A. chalcites samples from one locality were positively correlated with body size, suggesting that more carnivorous larvae become larger adults. However, this relationship was not detected in other species/locality groups. Thus, our results were inconclusive on the issue of diet supplementation. Nevertheless, overall, these results are comparable with those of previous laboratory‐rearing experiments and demonstrate the potential utility of stable isotope analysis in field studies on the feeding ecology of granivorous carabids.     

Size and taxonomic constraints determine the seed preferences of Carabidae (Coleoptera)

Several species of carabid beetles are important postdispersal predators of the seeds of herbaceous plants. The preferences of carabids for particular seeds differ, but the factors that determine their choice are little studied. We tested the hypothesis that preferences are determined by taxonomic constraints (carabid species affiliation), and carabid and seed size. The preferences were determined for adults of 30 species of central European field carabids mainly belonging to the tribes Zabrini (17 species) and Harpalini (10 species) (body mass 1–36 mg). In a cafeteria experiment the beetles were offered an excess of seeds from 28 species of dicotyledoneous herbaceous plants (mass 0.1–8.7 mg). The number of seeds eaten during a 5-day experiment was used as an estimate of preference. Mass of the preferred species of seed eaten was positively related to carabid body mass in both tribes. Multivariate analysis indicated three groups of carabid species with marked preferences for particular species of seeds: (i) species of Harpalini favoured mainly the seed of Cirsium arvense and Viola arvensis, (ii) some species of Zabrini the seeds of Asteraceae (Taraxacum officinale, Tripleurospermum inodorum and Crepis biennis) and (iii) other species of Zabrini the small seeds of Brassicaceae and Caryophyllaceae. The species of Harpalini were more generalist and accepted a greater proportion of seed species than Zabrini of the same size. Preferences of carabid seed predators were thus determined by taxonomic and size constraints, as in other groups of predators.     

Role of granivory and insectivory in the life cycle of the carabid beetle Amara similata

1. The cereal field carabid beetle Amara similata was selected to elucidate the role of seeds and insects as food sources. Three experiments were performed to rank different weed seeds and insects in terms of food value: (i) fecundity in relation to adult diet, (ii) larval survival in relation to diet and (iii) larval survival in relation to parental diet.
2. Seeds were found to be of high value and insects of low value both for adults and larvae. Adding insects to a seed diet gave no significant improvement. The value of single species of seeds varied within adults and larvae, but also between adults and larvae. This is the first report that Amara larvae are granivorous.
3. The value of different insects varied by species. Cereal aphid species were of the lowest value both for adult and larval beetles. On a diet of mixed insects, the adults were able to produce a low number of eggs but the larvae all died before pupation.
4. There was no simple relation between survival of the larvae and the quality of the parents' diet.     

Spatial and temporal patterns of carabid activity-density in cereals do not explain levels of predation on weed seeds

Seed predation is an important component of seed mortality of weeds in agro-ecosystems, but the agronomic use and management of this natural weed suppression is hampered by a lack of insight in the underlying ecological processes. In this paper, we investigate whether and how spatial and temporal variation in activity-density of granivorous ground beetles (Coleoptera: Carabidae) results in a corresponding pattern of seed predation. Activity-density of carabids was measured by using pitfall traps in two organic winter wheat fields from March to July 2004. Predation of seeds (Capsella bursa-pastoris, Lamium amplexicaule, Poa annua and Stellaria media) was assessed using seed cards at the same sites and times. As measured by pitfall traps, carabids were the dominant group of insects that had access to the seed cards. In the field, predation of the four different species of seed was in the order: C. bursa-pastoris>P. annua>S. media>L. amplexicaule; and this order of preference was confirmed in the laboratory using the dominant species of carabid. On average, seed predation was higher in the field interior compared to the edge, whereas catches of carabids were highest near the edge. Weeks with elevated seed predation did not concur with high activity-density of carabids. Thus, patterns of spatial and temporal variation in seed predation were not matched by similar patterns in the abundance of granivorous carabid beetles. The lack of correspondence is ascribed to effects of confounding factors, such as weather, the background density of seeds, the composition of the carabid community, and the phenology and physiological state of the beetles. Our results show that differences in seed loss among weed species may be predicted from laboratory trials on preference. However, predator activity-density, as measured in pitfall traps, is an insufficient predictor of seed predation over time and space within a field.     

Seasonal variation in seed predation by adult carabid beetles

In many agro‐ecosystems of the temperate region, granivorous ground beetles (Coleoptera: Carabidae) are the most important invertebrate predators of seeds after dispersal. The consumption of a particular species of seed differs among carabid species, but seasonal variation in consumption has not yet been studied. Seasonal differences in the quantity of seed consumed by the carabids Harpalus affinis (Schrank) and Harpalus distinguendus (Duftschmid) were studied in a cafeteria experiment. Adults were collected on five dates between April and November and provided with seeds of 28 species of dicotyledonous herbaceous plants. Proportions eaten of the seeds offered (preferences) were nearly identical in both carabid species. Overall seed consumption increased from early spring until late June, then gradually decreased towards autumn. Despite this variation, the preferences on particular dates were essentially similar. Seasonal variation in carabid physiology and feeding experience thus influenced overall seed consumption, but not their preferences for particular seed species.     

Mapping, characterisation, and comparison of the spatio-temporal distributions of cabbage stem flea beetle (Psylliodes chrysocephala), carabids, and Collembola in a crop of winter oilseed rape (Brassica napus)

The spatio‐temporal distribution of Psylliodes chrysocephala (L.) (Coleoptera: Chrysomelidae), a pest of oilseed rape (Brassica napus) (L.) (Cruciferae) and its potential predators, carabid beetles, within a crop of winter oilseed rape is described. The distribution of Collembola, a potential alternative food source for the predators, is also investigated. Insects were collected from spatially referenced sampling points across the crop and the counts mapped, analysed, and the degree of spatial association between the distributions determined using Spatial Analysis by Distance IndicEs (SADIE). Immigration into the crop by adult P. chrysocephala occurred from two edges and resulted in a non‐uniform distribution of the pest within the crop. Infestation of rape plants by P. chrysocephala larvae was greatest within the central area of the crop. Significant spatial association between adult female P. chrysocephala and the larval infestation of plants occurred throughout October. Three carabid species were active and abundant during peak pest immigration into the crop, viz., Trechus quadristriatus (Schrank) (Coleoptera: Carabidae), Pterostichus madidus (Fabricius) (Coleoptera: Carabidae), and Nebria brevicollis (Fabricius) (Coleoptera: Carabidae). Two of these species, T. quadristriatus and P. madidus, showed significant spatial association with the larvae of P. chrysocephala during October. All three carabid species showed a significant spatial association with Collembola during mid‐September, indicating that the latter may be an important food source for carabids during this period. In laboratory feeding experiments, only T. quadristriatus consumed the eggs of P. chrysocephala suggesting that, in the adult stage, this species may be the most important of the naturally occurring carabids as a predator of P. chrysocephala in the field. Adult T. quadristriatus may be a valuable component of an Integrated Pest Management strategy for winter oilseed rape, and the conservation of this species could be beneficial.     

Effects of diet on development of Podisus nigrispinus (Dallas) (Het., Pentatomidae), a predator of the cotton leafworm

Abstract:  The effects of diet on development of Podisus nigrispinus (Dallas) (Het., Pentatomidae) were studied at 25°C, relative humidity of 60 ± 10%, and photoperiod of 14 h light : 10 h dark. Development times of P. nigrispinus nymphs were similar when fed with third or fifth instar larvae of cotton leafworm ( Alabama argillacea Hübner) (Lep., Noctuidae) or Tenebrio molitor L. (Col., Tenebrionidae). When fed with housefly larvae ( Musca domestica L.) (Dipt., Muscidae) or artificial diet, the predator had a longer development time. Independent of diet, instar or sex, the females of P. nigrispinus showed a longer longevity than the males. The total survival of the nymphal stage, on the different diets, varied from 22.46 (fed with housefly) to 77.33% (fed with T. molitor larvae). P. nigrispinus males were heavier when fed with third or fifth instar cotton leafworm larvae than when fed with artificial diet. The weight of the females varied from 37.91 (with artificial diet) to 64.68 mg (with fifth instar cotton leafworm larvae). Independently of the diet, newly emerged females of P. nigrispinus were heavier than the males. Females of P. nigrispinus which were fed with fifth instar cotton leafworm larvae had heavier ovaries than those fed other diets.     

Effect of diet on the fecundity of three carabid beetles

Abstract In the present study, the importance of diet in terms of fecundity is compared for three species of the carabid genus Amara (Coleoptera: Carabidae), using an insect diet, two types of seed diet (Capsella bursa‐pastoris, Stellaria media) and a mixed diet. It is expected that the species of carabid studied have different food requirements for reproduction. Diet affects reproduction performance and egg production significantly. A mixed diet and both single‐seed diets are suitable for reproduction in Amara aenea (DeGeer) because a higher proportion of the females reproduce and lay significantly more eggs than on a purely insect diet. Females of Amara familiaris (Duftschmid) do not reproduce unless provided with seeds of S. media. Seeds of C. bursa‐pastoris or a mixed diet are equally suitable diets for reproduction of Amara similata (Gyllenhal); a diet of insects or seeds of S. media is unsuitable. The results support the hypothesis that the species under investigation have specific food requirements, suggesting that seed feeding has evolved to different degrees in particular species: A. aenea is omnivorous, whereas A. familiaris and A. similata specialize on the seed of a particular plant species or family. This resource partitioning facilitates co‐occurrence of carabid species.     

Laboratory studies on larval food habits of two syntopic,related, granivorous ground beetles <Emphasis Type="Italic">Amara chalcites</Emphasis> and <Emphasis Type="Italic">A. congrua</Emphasis> (Coleoptera: Carabidae): a comparison with stable isotope analysis

This study examined larval food habits of two syntopic, related, carabids Amara chalcites Dejean and A. congrua Morawitz using a laboratory-rearing experiment, and the results were compared with those of a previous stable-isotope analyses of the same species. Larvae were reared on six different diets (seeds of Capsella bursa-pastoris (Linnaeus) Medikus, Stellaria media (Linnaeus) Villars, Taraxacum officinale Weber ex Wiggers; mixed seeds; Tenebrio molitor Linnaeus larvae; and Tenebrio larvae + mixed seeds). For both species, beetles fed seed diets exhibited moderate to high survival rates, with the exception of those fed Stellaria seeds, on which A. congrua showed low survival rates. The pure-animal diet resulted in high survival rates of A. chalcites but low survival rates of A. congrua. These results were consistent with those of the isotope analysis, in which A. chalcites larvae were more carnivorous than were A. congrua larvae, and the larvae of the two species appeared to prefer different types of seeds. In contrast, the effect of diet on adult weight differed between the two methods. In the laboratory experiments, supplementing seeds with animal food during the larval stage positively affected adult weight for both species, whereas in the isotope analysis, the positive effect only occurred in A. chalcites. Possible explanations for the discrepancy between the two methods and suggestions for future research are provided.     

Impact of predatory carabids on below- and above-ground pests and yield in strawberry

The impact of adult carabid beetles on below- and above-ground pests and fruit yield was examined in the laboratory and a two-year strawberry field study. In the laboratory, adults of Carabus nemoralis Muller, Nebria brevicollis (F.), Pterostichus algidus LeConte, Pterostichus melanarius (Illiger), and Scaphinotus marginatus Fischer (Coleoptera: Carabidae) consumed black vine weevil, Otiorhynchus sulcatus (F.) (Coleoptera: Curculionidae) eggs, larvae and/or pupae placed on the surface. The same five carabid species showed no impact or low removal rates of O. sulcatus larvae that had burrowed into the root of potted strawberry plants. In an assay with only P. melanarius, adults consumed O. sulcatus larvae placed on the soil surface more frequently than larvae buried 1.3 or 5?cm below. In a field study, the density of adult carabids, predominantly P. melanarius, was manipulated with augmented, exclusion, and open control plots (2?×?2?m). Manipulating carabid density had no impact on the removal of sentinel O. sulcatus larvae and pupae that were buried belowground which is consistent with laboratory observations. Increasing carabid density within augmented plots led to greater removal of red clover seeds, Trifolium pratense L., placed on the soil surface in the first?year. Decreasing carabid density within exclusion plots resulted in fewer marketable fruits compared to control plots in both years. These results suggest that certain adult carabids may have limited impact belowground, and some beneficial impacts above-ground with pest control and crop protection.     

Invertebrate biodiversity affects predator fitness and hence potential to control pests in crops

Natural enemies that control pests usually allow farmers to avoid, or reduce, the use of pesticides. However, modern farming practices, that maximize yields, are resulting in loss of biodiversity, particularly prey diversity. Does this matter? Pests continue to thrive, and without alternative prey the predators should, perforce, concentrate their attentions upon the pests.We showed that a diverse diet significantly enhances predator fecundity and survival. Experiments were conducted using common generalist predators found in arable fields in Europe, the carabid beetle Pterostichus melanarius (Coleoptera: Carabidae) and the linyphiid spider Erigone atra (Araneae: Linyphiidae). We tested the hypothesis that mixed species diets were optimal, compared with restricted diets, with respect to parameters such as predator weights, egg weights, numbers of eggs laid, egg development times, egg hatching rates and predator survival. In carabids, an exclusive earthworm diet was as good as mixed diets containing earthworms for egg production and hatching, but less good than such mixed diets for increase in beetle mass and sustained egg laying. For spiders, aphids alone (Sitobion avenae) or with the Collembola Folsomia candida, drastically reduced survival. Aphids plus the Collembola Isotoma anglicana improved survival but only aphids with a mixed Collembola diet maximized numbers of hatching eggs.Predators offered only pests (slugs or aphids) had lowest growth rates and fecundity. We therefore demonstrated that conservation of a diversity of prey species within farmland, allowing predators to exploit a diverse diet, is essential if predators are to continue to thrive in crops and regulate agricultural pests.     

Invertebrate biodiversity affects predator fitness and hence potential to control pests in crops

Natural enemies that control pests usually allow farmers to avoid, or reduce, the use of pesticides. However, modern farming practices, that maximize yields, are resulting in loss of biodiversity, particularly prey diversity. Does this matter? Pests continue to thrive, and without alternative prey the predators should, perforce, concentrate their attentions upon the pests.We showed that a diverse diet significantly enhances predator fecundity and survival. Experiments were conducted using common generalist predators found in arable fields in Europe, the carabid beetle Pterostichus melanarius (Coleoptera: Carabidae) and the linyphiid spider Erigone atra (Araneae: Linyphiidae). We tested the hypothesis that mixed species diets were optimal, compared with restricted diets, with respect to parameters such as predator weights, egg weights, numbers of eggs laid, egg development times, egg hatching rates and predator survival. In carabids, an exclusive earthworm diet was as good as mixed diets containing earthworms for egg production and hatching, but less good than such mixed diets for increase in beetle mass and sustained egg laying. For spiders, aphids alone (Sitobion avenae) or with the Collembola Folsomia candida, drastically reduced survival. Aphids plus the Collembola Isotoma anglicana improved survival but only aphids with a mixed Collembola diet maximized numbers of hatching eggs.Predators offered only pests (slugs or aphids) had lowest growth rates and fecundity. We therefore demonstrated that conservation of a diversity of prey species within farmland, allowing predators to exploit a diverse diet, is essential if predators are to continue to thrive in crops and regulate agricultural pests.     

Effects of maternal diet quality on offspring performance in the rove beetle Tachyporus hypnorum

Abstract.  1. The reproductive output of an individual is known to be influenced by diet quality, but the quality of the parent's diet can also influence the performance of the offspring. Dietary maternal effects may interact with the effects of the offspring's diet to produce a variety of response patterns.
2. Maternal effects were investigated in a polyphagous predator, the rove beetle Tachyporus hypnorum , using three single-species diets: two low-quality diets consisting of the aphids Sitobion avenae and Rhopalosiphum padi , and the high-quality control Drosophila melanogaster diet. Offspring of females fed these diets were raised on the same monotypic diets and allowed to reproduce. Several fitness parameters were measured to indicate possible maternal effects in both F1 and F2 generations.
3. Maternal diet effects in F1 were found in egg size, hatching success, time to hatching, larval development time, larval survival, and sex ratio. Both aphid diets resulted in smaller eggs. A diet of R. padi resulted in reduced hatching success, longer time spent in the egg stage, and a female-biased sex ratio. A maternal diet of R. padi also prolonged larval development on S. avenae diet, while a maternal diet of S. avenae decreased survival on the R. padi diet. These effects were independent of egg size.
4. A maternal diet of R. padi enhanced the survival of F1 larvae raised on the same diet. Developmental selection operating through a high egg mortality may be the explanation for this seemingly positive effect.
5.  Sitobion avenae alone caused a significant reduction in the hatching success of F2 eggs, thus revealing grandmaternal effects.
6. The prediction that polyphagous predators are less likely to evolve adaptive maternal effects is supported by the fact that none of the documented maternal effects could be interpreted as adaptive.