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.     

Effects of maternal diet on fecundity and larval development in the 'primitive' granivorous carabid Amara (Curtonotus) macronota

Granivory is one of the most specialized food habits in the Carabidae and has been reported for species from the tribes Zabrini and Harpalini. Most studies of carabid granivory have been conducted using specialized granivorous species, and few have examined primitive ones. This study examined effects of maternal diet on fecundity and larval development in Amara ( Curtonotus ) macronota (Solsky) (Coleoptera: Carabidae), a member of the most basal clade of the tribe Zabrini; a previous study indicated that larvae of this species are omnivores with a tendency toward carnivory. Three diet types, Tenebrio molitor L. larvae, mixed seeds [ Bidens frondosa L. (Asteraceae), Setaria spp., and Digitaria ciliaris (Retz.) Koeler (both Poaceae)], and T. molitor larvae + mixed seeds, were used as maternal diets, and larvae were reared on T. molitor larvae + mixed seeds (optimal diet) or T. molitor larvae (suboptimal diet). Fecundity differed significantly among treatments, with individuals fed the T. molitor larvae + mixed seeds diet having the highest fecundities and the mixed seeds diet producing the lowest values. Larval development [survival and duration of development through pre-overwintering stages (first and second instars)] was not significantly affected by either maternal or larval diets, but the addition of seeds to the maternal diet had a weak negative effect on larval survival. These results are in contrast with findings from specialized granivorous carabids, in which both adults and larvae performed best with pure-seed diets. Recent molecular phylogenies indicate that these specialized granivorous carabids belong to derived lineages, while A. macronota is the most basal clade of each tribe. These results are compatible with the hypothesis that carabid granivory has evolved gradually from ancestral carnivory, with omnivorous habits occurring as a transient state.     

The biology of seed discrimination and its role in shaping the foraging ecology of carabids: A review

Species of carabid (ground) beetles are among the most important postdispersal weed seed predators in temperate arable lands. Field studies have shown that carabid beetles can remove upwards of 65%–90% of specific weed seeds shed in arable fields each year. Such data do not explain how and why carabid predators go after weed seeds, however. It remains to be proven that weed seed predation by carabids is a genuine ecological interaction driven by certain ecological factors or functional traits that determine interaction strength and power predation dynamics, bringing about therefore a natural regulation of weed populations. Along these lines, this review ties together the lines of evidence around weed seed predation by carabid predators. Chemoperception rather than vision seems to be the primary sensory mechanism guiding seed detection and seed selection decisions in carabid weed seed predators. Selection of weed seeds by carabid seed predators appears directed rather than random. Yet, the nature of the chemical cues mediating detection of different seed species and identification of the suitable seed type among them remains unknown. Selection of certain types of weed seeds cannot be predicted based on seed chemistry per se in all cases, however. Rather, seed selection decisions are ruled by sophisticated behavioral mechanisms comprising the assessment of both chemical and physical characteristics of the seed. The ultimate selection of certain weed seed types is determined by how the chemical and physical properties of the seed match with the functional traits of the predator in terms of seed handling ability. Seed density, in addition to chemical and physical seed traits, is also an important factor that is likely to shape seed selection decisions in carabid weed seed predators. Carabid responses to seed density are rather complex as they are influenced not only by seed numbers but also by trait‐based suitability ranks of the different seed types available in the environment.     

Weed seed availability,carabid intraspecific interference and their interaction drive weed seed consumption by carabid beetles

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Plant versus prey resources: Influence on omnivore behavior and herbivore suppression

Conservation biological control tactics, such as beetle banks, that increase habitat complexity generally increase epigeal predator abundance. Habitat complexity also increases alternative food which can attract and sustain predators but may reduce predation of target pests. Our goal was to determine how alternative food from different trophic levels (fly pupae and seeds) affects behavior and biological control efficacy of omnivorous carabid beetles. Seed subsidies increased omnivorous carabid abundance more than pupae by increasing aggregation and reducing emigration. Laboratory experiment demonstrated that both omnivorous carabid species preferred seeds and pupae over cutworms. However, in field cages seeds but not pupae resulted in greater cutworm damage to corn seedlings. Our results indicate that omnivorous carabids have a stronger behavioral response to seeds than prey such that only seeds influence aggregation, emigration, and crop damage. Interestingly, whereas seeds increased omnivorous carabid abundance, pupae had no affect on carnivore abundance. Thus, carabid guild composition is skewed in favor of omnivores when seed density increases. An important finding was that the effect of seeds on behavior, predation, and crop damage was consistent among replicate carabid species suggesting our results pertain to other omnivorous species in resource diverse habitats.Our results provide insight into the mechanisms underlying the unpredictable benefit of conservation biological control tactics that alter habitat complexity.     

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.     

The strength of seeds and their destruction by granivorous insects

The influence of seed structure and strength on their destruction by granivores is central to understanding the dynamics of granivore-plant interactions. For up to nine seed species, the effects of seed size (cm³), mass (mg), density (mg/cm³) and coat strength (MPa) on the damage inflicted by three post-dispersal granivores (Harpalus pensylvanicus, Anisodactylus sanctaecrucis, and Gryllus pennsylvanicus) were evaluated. Seed destruction rates by G. pennsylvanicus were statistically unrelated to the size and toughness of the seeds. Seed densities significantly affected their destruction by A. sanctaecrucis and H. pensylvanicus, as did seed size, mass, and strength in H. pensylvanicus under choice conditions. The carabid beetles destroyed more of the small, denser seeds with stronger seed coats. The results show that different granivores are able to distinguish the structural strength and physical density of seeds as well as seed size. The relative ability of granivores to detect these seed characteristics offers a way in which diverse communities of post-dispersal insect granivores can persist within a single habitat. The authors redefine how the strength of biological structures should be evaluated in ecological studies, using guidelines commonplace in the field of engineering. Handling Editor: Stanislav Gorb The U.S. Government’s right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged     

A chance field observation and pilor laboratory studies of predation of the New Zealand flatworm by the larvae and adults of carabid and staphylinid beetles

A chance finding in an Edinburgh allotment was made of a stained carabid beetle larva that had eaten Artioposthiu triungulata previously stained red and released into the allotment during a dispersal experiment. In a pilot laboratory study the flatworm was eaten by larval and adult carabid and staphylinid beetles whereas seven other carnivorous species from five arthropod families did not. In another feeding experiment a carabid and a staphylinid larva each ate several specimens of the flatworm and gained tissue weight over 14 days. The conclusion drawn from these preliminary studies and the field observation is that certain carabid and staphylinid beetles are potential predators of A. triangulata.     

Predator interference limits fly egg biological control by a guild of ground-active beetles

We examined whether predator interference could prevent effective conservation biological control of Delia spp. flies, important pests of cole crops, by an assemblage of carabid and staphylinid beetles. In laboratory feeding trials we found that the smaller (<1 cm) beetle species common at our site readily ate dipteran eggs, while the most common large carabid species, Pterostichus melanarius, rarely did. However, P. melanarius did eat several of the smaller beetle species. We conducted two field experiments where we manipulated immigration rates of the ground predator guild and then measured predation on fly eggs. Predation rates were consistently higher in cages where predators were added at ambient densities, compared to cages where ground predators were removed. However, in the second field experiment, when we quadrupled predator immigration rates neither beetle activity-density nor predation rate increased. High immigration rate plots had a higher proportion of P. melanarius in the predator community, compared to plots with beetles added at ambient densities, suggesting that P. melanarius was reducing activity-densities of the smaller beetles, perhaps through intraguild predation. Thus, tactics to improve the biological control of Delia spp. by conserving generalist predators, such as providing in- or extra-field refuges, could be thwarted if the primary predators of fly eggs, small carabids and staphylinids, are the targets of intraguild predation by also-conserved larger predators.     

Manipulating carabid beetle abundance alters prey removal rates in corn fields

Generalist natural enemies such as carabid beetles have the potential to maintain a variety of pests below outbreak levels in annual crops. To assess the relationship between carabid beetle abundance and field rates of prey removal, we created plots surrounded by different boundaries that selectively affected dispersal of edaphic arthropods, primarily carabids. Three treatments were established: (1) naturally occurring communities, (2) augmented communities using ingress boundaries, and (3) reduced communities using egress boundaries. Selective boundaries altered carabid communities with minimal habitat alteration and without use of insecticides. Three times during the growing season, a fixed number of onion fly pupae were placed in plots to evaluate the impact of carabid abundance on predation rates. A combination of vertebrate and invertebrate exclosures allowed us to evaluate prey removal by invertebrates alone. In comparison to the no boundary treatment, carabids increased 54.2% and decreased 83.1% in plots surrounded by ingress and egress boundaries respectively. Predation rates were positively correlated with carabid abundance (r² = 0.70, p < 0.0001). Significantly more pupae were removed from exclosures allowing access to invertebrates alone than from total exclosures, suggesting that invertebrates represented an important group of predators. Laboratory trials tested the feeding potential of the four most abundant carabid species and showed that they readily consumed onion fly pupae, supporting our hypothesis that carabids were the main predators in field tests. This study corroborates and extends previous observations of the importance of carabid beetles as generalist predators of insect pests in agricultural fields.     

Rubidium marking reveals different patterns of movement in four ground beetle species (Col., Carabidae) between adjacent alfalfa and maize

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Comments on relationships between native seed preferences of shrub-steppe granivores and seed nutritional characteristics

Summary Kelrick et al. (1986) argued that seed preferences of desert granivores are strongly influenced by soluble carbohydrate contents of native seed species. They assumed that bitterbrush (Purshia tridentata) seeds are eaten in their entirety by rodents, but in fact these granivores eat only embryos of bitterbrush seeds. Embryos have a much lower percentage of soluble carbohydrate than whole bitterbrush seeds, and the correlation between preference and soluble carbohydrate content of seeds for six native species becomes non-significant when embryo values are substituted for whole-seed values.     

Sabal palmetto seed size: causes of variation,choices of predators,and consequences for seedlings

High variation in seed size, as is common among angiosperms, may be maintained in a plant species when several factors select for seed size. Variation may also result from differences among adult plants, such as nutrient and water availability or the amount of photosynthetic tissue. In a study of Sabal palmetto seed ecology I found high seed size variation both within- and among-palms, and investigated possible factors maintaining this variation. Seed size was positively correlated with the number of leaves on parent palms. Larger seeds produced more vigorous seedlings that had greater leaf length, area, and mass, and greater root mass. Caryobruchus gleditsiae (Bruchidae: Coleoptera), whose larvae develop within palm seeds, preferentially oviposited on larger seeds, which in turn produced larger beetle offspring. By choosing the largest seeds available, ovipositing beetles thus affect both the quantity and the quality of seeds available for recruitment. I conclude that because beetle predation selects against large seeds, while larger seeds promote seedling vigor, the maintenance of seed size variation may be an adaptation of S. palmetto promoting both seed escape from predators and seedling vigor.     

A field experiment on the effectiveness of spiders and carabid beetles as biocontrol agents in soybean

1 Spiders and carabid beetles are abundant generalist predators that prey upon insect pests of soybean. A field experiment was conducted to determine the impact of spiders and carabids on soybean yield. Prior to planting, three 7 × 7 m plots were fenced in order to reduce spider and carabid immigration. Carabids that emerged within the plots were not removed, but spiders that ballooned into these predator‐reduction plots or that entered by climbing the fence were removed by pitfall trapping and searching the vegetation. Three unmanipulated, unfenced plots served as the control treatment. 2 Densities of spiders on soybean vegetation, and activity‐densities of spiders and carabids determined by pitfall trapping, were c. 75% lower in the spider‐carabid reduction treatment than in control plots. Despite clear differences between treatments in numbers and activity of these major generalist predators, the weight of soybeans harvested did not differ between control and spider‐carabid reduction plots. 3 Paralleling the absence of an effect of predator reduction on soybean yield was the absence of any significant difference between treatments in densities of whiteflies (Aleyrodidae), leafhoppers (Cicadellidae), thrips (Thysanoptera), Lepidoptera larvae and herbivorous Coleoptera. 4 Our experiment provides no evidence that spiders and carabid beetles at ambient densities affect soybean yield. Low populations of pest species or low predation pressure on soybean pests by spiders and carabids at the ambient densities of this experiment could be responsible for this result.     

Ground beetle (Coleoptera: Carabidae) assemblages in conventional and diversified crop rotation systems

Ground beetles (Coleoptera: Carabidae) are important in agro-ecosystems as generalist predators of invertebrate pests and weed seeds and as prey for larger animals. However, it is not well understood how cropping systems affect ground beetles. Over a 2-yr period, carabids were monitored two times per month using pitfall traps in a conventional chemical input, 2-yr, corn/soybean rotation system and a low input, 4-yr, corn/soybean/triticale-alfalfa/alfalfa rotation system. Carabid assemblages were largely dominated by a few species across all cropping treatments with Poecilus chalcites Say comprising >70% of pitfall catches in both years of study. Overall carabid activity density and species richness were higher in the low input, 4-yr rotation compared with the conventionally managed, 2-yr rotation. There were greater differences in the temporal activity density and species richness of carabids among crops than within corn and soybean treatments managed with different agrichemical inputs and soil disturbance regimes. Detrended correspondence analysis showed strong yearly variation in carabid assemblages in all cropping treatments. The increase in carabid activity density and species richness observed in the 4-yr crop rotation highlights the potential benefits of diverse crop habitats for carabids and the possibility for managing natural enemies by manipulating crop rotations.     

Interference competition,not predation,explains the negative association between wood ants (Formica rufa) and abundance of ground beetles (Coleoptera: Carabidae)

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Wild flower seed predation by Pterostichus madidus (Carabidae: Coleoptera)

The objective of the study was to investigate the ability of Pterostichus madidus, one of the most abundant carabid beetles in the United Kingdom, to consume wildflower seeds. The plant species chosen are commonly included in wildflower seed mixes sown to enhance the biodiversity of arable field margins, and hence any seed predation could potentially reduce successful seedling establishment. The study combined cafeteria style laboratory feeding experiments with predation studies in an established field margin. In laboratory studies the Pterostichus madidus consumed both unimbibed and imbibed seeds of Centaurea scabiosa, Galium verum, Leucanthemum vulgare and Primula veris. Seed consumption was still evident when Drosophila pupae were also supplied, although there was significant preference for the pupae. Placement of seeds in the field showed high levels of seed consumption, although part of this was attributed to slugs. Carabid beetles may have a significant role as wild flower seed predators and thus may affect establishment of re‐seeded flower‐rich meadows.     

Post-dispersal predation on Pinus sylvestris seeds by Fringilla spp: ground substrate affects selection for seed color

Background matching might lower the risk of seeds being eaten by seed predators that search visually. In aviary experiments, we analyzed the selection of diff erent-colored seeds by ground-feeding finches (Fringillacoelebs and F.montifringilla) against four naturally occurring forest soil substrates. The substrates were fresh burn (black), 6-year-old burn (brown), mineral soil (pale yellow) and Pleuroziumschreberi feather moss (green). We used color-sorted seeds of Pinussylvestris, a species with a large natural variation in seed color, ranging from pale yellow to black. Although seeds were scattered on the substrates at a density of only 91 seeds m⁻², birds removed seeds effectively. Both bird species found more pale than dark seeds on the fresh burn substrate. F. montifringilla also recovered more pale than dark seeds on the old burn, and more dark than pale seeds on mineral soil. In moss, the birds found very few seeds compared to the other substrates, and there was no color discrimination. P.sylvestris is frequently regenerating after fire, suggesting that dark seeds would be favored under selection from visually searching predators. Fire-adapted conifers with serotinous cones, e.g., Pinuscontorta ssp. latifolia, which spread their seeds primarily on freshly burnt surfaces, produce uniformly black or dark brown seeds. However, regeneration of the non-serotinous P.sylvestris is often extended for several years after a fire, during which substrate color and structure change. This may have helped to maintain variation in seed color. When regeneration of a plant species occurs on a substrate of uniform color, we believe that selection by visually searching seed predators will result in the evolution of cryptic seed color. Received: 16 August 1996 / Accepted: 30 November 1996     

Effect of beetle size on predation of cabbage root fly eggs by ground beetles

Laboratory studies were done to determine the numbers of cabbage root fly (Delia radicum L.) (Diptera: Anthomyiidae) eggs eaten by sixty species of ground beetles (Coleoptera: Carabidae) collected from a range of habitats. For 35 species that were between 2.7 mm and 10 mm long, there was a linear relationship between the numbers of eggs eaten and beetle length. For each 1 mm increase in length above 2.7 mm, an additional 18 eggs were eaten/beetle/day. Eight further species within the above size range did not eat any fly eggs, as these species are known to be either phytophagous or to feed on moving prey. Large numbers of eggs were eaten by beetles in 13 of the 25 genera tested. Based on size, maximum numbers of eggs were eaten by 7 species of Agonum Bonelli, 5 species of Amara Bonelli and seven species of Bembidion Latreille. Only one of the 6 species of Pterostichus Bonelli tested, P. strenuus (Panz.), ate the expected numbers of eggs, the other species being too large to feed from such small prey items as cabbage root fly eggs. The numbers of beetles larger than 10 mm that ate eggs was highly variable and so 10 mm was considered the upper size limit of carabid predators of cabbage root fly eggs.     

Diversity of foraging strategies and responses to predator interference in seed-eating carabid beetles

The prediction of pest regulation by multi-predator communities often remains challenging because of variable and opposite effects of niche complementarity and predator interference. Carabid communities are regulating weeds in arable fields and include a mix of species ranging from granivores to predators that are obligate omnivores. It is not clear from field studies whether granivore and obligate omnivore species either contribute equally or are complementary in the process of weed suppression, and little is known about the impact of potential predator interference within carabid communities on weed suppression. We compared the weed seed foraging strategy of the granivore Harpalus affinis and the obligate omnivore Poecilus cupreus. Using no-choice test experiments, we compared their activity and seed acceptance for four weed species through a scoring of the proportion of tested individuals consuming weeds, their latency before the consumption of the first seed and the total number of seeds consumed. We then evaluated their seed acceptance for dandelion seed Taraxacum officinale under predator interference by using chemical cues of carabids and tested the impact of three treatments, namely cues of intraspecific competition, interspecific competition and intraguild predation. We found that the obligate omnivore P. cupreus was highly active, had a low latency before consuming its first seed but had an interest in only two of the four weed species. P. cupreus seed acceptance remained unchanged in the presence of predator cues. By contrast, H. affinis was slow to start its seed consumption, accepted equally seeds of the four weed species and significantly increased its seed consumption in the presence of cues mimicking intraguild predation. These findings indicate that the two species differ in their foraging strategies, and as such, could have different contributions to weed seed suppression. This novel result calls for further studies documenting the foraging strategy of carabid species that thrive in arable fields as this could significantly improve our understanding of the delivery of weed seed regulation.