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.     

Organic farming affects the potential of a granivorous carabid beetle to control arable weeds at local and landscape scales

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Mixed effects of landscape complexity and farming practice on weed seed removal

Agricultural intensification negatively affects farmland biodiversity, and thereby, ecosystem services such as predation of weed seeds. Landscape complexity (large and diverse areas of semi-natural habitats) and organic farming (without pesticides and mineral fertilizers) can counteract this negative trend, but little is known about their relative importance. We experimentally studied seed removal of four economically relevant weed species by excluding (i) vertebrates, (ii) vertebrates plus large invertebrates and (iii) neither vertebrates nor invertebrates (control) at the field edge and in the interior of 22 organically or conventionally managed winter wheat fields, located across a gradient of landscape complexity (41–94% arable land). Arable land (%) was used as a simple predictor of landscape complexity, as it is closely related to landscape parameters such as habitat type diversity. Seed predation and removal were mainly influenced by large invertebrates, seed species identity, with Cirsium arvense being more affected than Poa trivialis, Apera spica-venti and Galium aparine and landscape complexity in interaction with farming practice, as it increased in conventional fields, but decreased in organic fields as landscape complexity decreased. Seed predation and removal did not consistently correspond to seed predator densities, as small mammal abundances decreased in conventional fields, but increased in organic fields as landscape complexity decreased and carabid activity densities were higher in organic compared to conventional fields. Slug activity density showed same response to landscape complexity and farming practice like seed removal and appeared to additionally influence the movement of seeds. In conclusion, our assessment of interacting local and landscape effect adds to the current knowledge of seed predation and removal. Farming practice and landscape complexity affect seed predator species differentially, thereby resulting in complex pattern of seed predation and removal. The service of weed seed predation is provided by small mammals and invertebrates and the disservice of seed dispersal and movement is provided by slugs.     

Tillage compromises weed seed predator activity across developmental stages

Granivorous ground beetles (Coleoptera: Carabidae) are ubiquitous throughout temperate agricultural systems, and reduce weed seedbanks. However, trade-offs may exist between tillage frequency and ecosystem services of invertebrate seed predators, especially those in the larval stages, which have relatively poor resistance to disturbance. While much research has focused on adult activity patterns and the conservation biocontrol services they provide, almost nothing is known about carabid biology and habitat requirements during larval stages, despite the fact that adult recruitment is determined by factors that promote larval survival. We present data on larval and adult Harpalus pennsylvanicus Dej., a common weed seed predator across North America, from two experiments examining its activity density across tillage and cover-cropping treatments in organic tomato systems. Larvae emerged 4–6 weeks after the adult activity peak, and larval activity density was up to 10 times higher in no-till crop environments than in cultivated areas. After a long disturbance interval, seasonal cultivation had no effect on foraging activity of adults, but reduced larval activity density in both experiments. Additionally, larvae positively correlated with living weed biomass in no-till treatments, suggesting the importance of plant-based resources in oviposition site choice. Compared with adults, larvae are relatively immobile and vulnerable to disturbance; thus, weed management strategies that rely on frequent cultivation may undermine the ecosystem services provided by granivorous insects.     

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.     

Ground beetle (Coleoptera: Carabidae) phenology, diversity, and response to weed cover in a turfgrass ecosystem

Despite being fragmented and highly disturbed habitats, urban turfgrass ecosystems harbor a surprising diversity of arthropods. The suitability of turf as arthropod habitat, however, likely depends on the extent and types of pesticides and fertilizers used. For example, moderate levels of weed cover in low-input lawns may provide alternative food resources. We conducted a 2-yr field study to: 1) characterize the ground beetle (Carabidae) species assemblage in turfgrass, and 2) assess the direct and indirect effects of lawn management on carabid communities. Weed cover and beetle activity were compared among four lawn management programs: 1) consumer/garden center, 2) integrated pest management (IPM), 3) natural organic, and 4) no-input control. Nearly 5,000 carabid beetles across 17 species were collected with the predator Cyclotrachelus sodalis LeConte numerically dominating the trap catch (87% and 45% of individuals in 2005 and 2006, respectively). Populations of C. sodalis underwent a distinct peak in activity during the third week of June, whereas omnivorous and granivorous species tended to occur at far lower levels and were less variable over the season. We found no evidence for direct effects of lawn management on carabid species diversity; however, we detected an indirect effect mediated by variation in weed cover. Seed-feeding species were positively correlated with turf weeds early in 2006, whereas strictly predaceous species were not. Thus, turf management programs that lead to changes in plant species composition (i.e., herbicide regimes) may indirectly shape epigeal arthropod communities more strongly than the direct effects of insecticide use.     

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.     

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.     

Unravelling the process of weed seed predation: Developing options for better weed control

Seed predation can cause significant losses of weed seeds in agricultural systems and can, thus, contribute to weed control. The removal of Lolium multiflorum and Vicia villosa seeds by harvester ants, Messor barbarus, and granivorous rodents, Mus spretus, in six cereal fields in NE Spain was separated into three sequential processes, namely (1) the probability of finding a seed cache (cache encounter rate), (2) the percentage of seeds utilized once a seed cache has been found (seed exploitation rate) and (3) seed selection if multiple species are present (preference). Identifying the most important behavioural component and factors that drive it may help to better understand and manage seed predation.Seed cache encounter rate correlated well with overall seed removal rate caused by harvester ants (r² = 0.91), or rodents (r² = 0.82). Once found, seed exploitation rates were high and fairly constant from spring to autumn for harvester ants, and low throughout the season for rodents. Harvester ants removed almost all L. multiflorum seeds from caches found, while the exploitation of V. villosa seeds varied across the season. In the case of rodents, cache encounter rate, but not exploitation rate, could be explained by canopy cover provided by the crop. L. multiflorum seemed to be preferred in early 2007, whereas V. villosa was in 2008.The adoption of no-till or minimum tillage systems together with the establishment of field edge vegetation are likely to encourage seed cache encounter and exploitation rates by both harvester ants and rodents, thus leading to increased weed control in semi-arid cereals.     

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

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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.     

Habitat effects on second-order predation of the seed predator Harpalus rufipes and implications for weed seedbank management

Seed predators provide a valuable ecosystem service to farmers by reducing densities of weed seeds, and, in turn, densities of weed seedlings they must manage. The predominant invertebrate weed seed predator in Maine, USA, agroecosystems is the carabid beetle Harpalus rufipes DeGeer. Pitfall trapping has shown that H. rufipes prefers sites with vegetative cover to fallow sites, preference speculated to be driven by predator avoidance behavior. To test this hypothesis, ‘second-order predation assays’ were developed, in which live H. rufipes prey were presented to second-order predators. Field experiments were conducted to determine foremost if H. rufipes was subject to second-order predation, and secondly, whether (a) vegetative cover affords H. rufipes protection from second-order predators, and (b) high rates of second-order predation correspond with decreased invertebrate seed predation rates. Two 72-h experiments were conducted (mid August and September 2012) at crop and non-crop sites across a 28 ha diversified farm in Stillwater, ME, USA.Second-order predation was 2.8% per day. Based on images from motion-sensing cameras, H. rufipes’ predators included birds and small mammals. Neither a relationship between second-order predation and vegetative treatment, nor an empirical relationship between second-order predation and invertebrate seed predation were detected. However, a simulation model predicted that 2.8% per day second-order predation could increase the number of seeds entering the seedbank by more than 17% annually. Additionally, complex habitats supported higher rates of second-order predation than did simple habitats.     

Annual variation of oilseed rape habitat quality and role of grassy field margins for seed eating carabids in arable mosaics

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The fear diet: Risk,refuge, and biological control by omnivorous weed seed predators

Weed seed biocontrol by omnivorous mice and insects can limit weed seedbanks, but this ecosystem service can be difficult to predict given the broad diet breadth of seed predators and their potential for intraguild predation. Seed foraging behavior is further modified by fluctuating cues of predation risk from higher trophic levels and the availability of refuge habitat. Uncertainty about whether co-occurring insects and mice additively contribute to weed biocontrol or interfere with each other via intraguild predation limits our ability to recommend habitat management strategies that reliably promote seed destruction. Using seed removal assays, fluorescent powder tracking, and stable isotope analyses, we assessed effects of a predation risk cue (moonlight) on mouse foraging patterns in a patchwork of vegetated and exposed plots in a cultivated field. Mouse foraging activity decreased on exposed ground during the full moon, compared to dark nights, yet foraging movements were unaffected by moon cycle within refuge patches. Weed seed consumption was more than three times higher in cover than exposed soil, and 78% of that difference was attributable to invertebrate granivores. Mice and invertebrate granivores both exhibited higher foraging activity in cover, indicating co-occurrence of intraguild predators and prey. However, stable isotope analyses of fecal samples revealed that mice captured in refuge habitats fed at slightly lower trophic levels than those in exposed habitats (suggesting minimal intraguild predation in refuge habitat), and mouse diet was unaffected by moonlight. Despite increased availability of invertebrate prey in cover patches, mice do not appear to preferentially exploit prey when avoiding their own predators or interfere with weed seed predation. Therefore, functional redundancy of mice and invertebrate seed predators in cover crops and other refuge habitats may strengthen and stabilize weed seed biocontrol.     

半自然农田边界与相邻农田步甲和蜘蛛的时空分布

分别于小麦、玉米收获前后,采用陷阱法调查了华北地区典型农业景观中具有不同植被结构的农田边界及其相邻农田中两类重要天敌类群步甲和蜘蛛的多样性.通过比较农田生境及相邻农田边界间两类天敌群落的时空分布格局及其与相邻半自然生境植被群落的相关关系,探讨半自然农田边界对两类天敌类群的保护作用.结果表明: 整个取样季节农田边界处蜘蛛的多度显著高于农田内部;而步甲多样性在农田与边界间无显著性差异,仅呈现不同的群落结构;作物收获后蜘蛛分科数在边界处的增加以及在农田的减少,显示了蜘蛛在农田和边界之间的迁移活动.边界植被结构对蜘蛛和步甲多样性有不同影响:边界较高的草本层盖度和较低的乔木层盖度有利于增加农田中某些步甲优势种的多样性;而较高的草本层盖度有利于增加皿蛛科蜘蛛的多样性.因此,半自然生境的存在可以通过天敌在农田和边界之间的迁移运动促进农田天敌多样性的维持;但不同类型半自然生境植被群落结构可能影响其对不同天敌群落多样性的维持和保护作用.为促进农业景观对天敌的保护作用,提高其害虫控制功能,需要深入了解不同天敌的生境需求及食物需求,精心设计有利于天敌多样性维持的半自然生境.     

Increasing habitat connectivity in agricultural landscapes as a weed management strategy reconciling ecology and agronomy

While studies have explored how habitat amount drives weed assemblages in agroecosystems, knowledge remains limited of the effects of habitat connectivity. The response-effect trait framework provides insights into the mechanisms underpinning the relationship between landscape structure and the taxonomic diversity and abundance of weed assemblages. This study evaluated how habitat connectivity and habitat amount affect weed diversity and abundance in winter cereal fields, and whether these effects are driven by the functional composition of weed assemblages. We sampled weeds in 27 winter cereal fields. We measured habitat connectivity and habitat amount provided by wooded, grassland and cropland elements. We selected five traits related to the dispersal, establishment, and competitive abilities of weed species likely to respond to landscape structure: seed number per plant, type of reproduction, seed dry mass, plant vegetative height and seed germination rate. The functional composition of weed assemblages was assessed using community weighted mean trait values. Weed diversity and abundance were used as proxies of weed management. The taxonomic approach did not reveal any effect of landscape structure on weed diversity and abundance. Only the grassland elements that contributed to habitat connectivity, and to a lesser extent to habitat amount, drove the functional composition of weed assemblages. High habitat amount favoured species with many seeds, while high habitat connectivity favoured species with fewer seeds, a higher ability to reproduce vegetatively and higher seed germination rates. In turn, higher seed germination rates increased weed evenness and reduced weed abundance. Some of these relationships were influenced by the presence of rare species. Overall, high connectivity provided by grassland elements increases weed evenness and reduces weed abundance by shaping weed functional composition. Our study suggests that land-use planning policies that enhance the connectivity provided by grassland elements could be considered as a weed management strategy reconciling ecology and agronomy.     

Insect pests in winter oilseed rape affected by field and landscape characteristics

Winter oilseed rape (OSR, Brassica napus) cropping is often associated with an intensive use of pesticides. The transformation of structurally rich landscapes into more monotonous landscapes may be partly responsible for this, because non-crop habitats believed to benefit natural enemies have been eliminated. We examined the influence of field (soil quality, nitrogen fertilization) and landscape characteristics (OSR area and isolation, non-crop area, landscape diversity, proportions of grassy fallows and woody areas) on three major European OSR pest groups: pollen beetles, stem weevils, and brassica pod midges. Twenty-nine landscape sectors ranging from structurally poor to complex were studied at eight spatial scales (radii 250–2000 m) centered in the studied OSR fields. Abundances of pollen beetles and stem weevils were significantly positively correlated with soil quality and negatively related to OSR area in the surroundings. Generally, abundances of all groups were positively related to woody areas, but not related to grassy fallow area. Pod midges and stem weevils tended to respond primarily to landscape variables at small (250–500 m) and medium (1000–1250 m) scales, while pollen beetles responded at medium to large (1000–2000 m) scales. The results are discussed in relation to differences in overwintering strategies and mobility of pest insects. Strategies at the field and landscape level, aiming to reduce pest pressure in OSR fields, are also discussed.     

The effect of forest fragments on abundance,diversity and species composition of predatory ants in sugarcane fields

Habitat loss and fragmentation have gradually caused loss of diversity and consequently the decline of ecological services. This study aimed to evaluate the effect of tropical forest fragments as natural habitats (river valley fragments and plateau fragments) on the community of predatory and omnivorous ants in nearby sugarcane fields. Twenty fields adjacent to these fragments were selected and evaluated one (dry season) and four months (rainy season) after harvest. In each field, ants were sampled in five linear plots (10 m inside the fragment, 0 m (field path between field and fragment), 5 m, 50 m and 100 m inside the crop fields). Each plot comprised ten sardine baits in a row parallel to the field edge. Species richness and frequency of ant species decreased with increasing distance from the forest fragments. Inside fields, species richness and frequency were higher during the period of vegetative growth (rainy season) than after harvest (dry season). Ant communities of sugarcane fields and forest fragments were more similar later in the season than directly after sugarcane harvest suggesting recolonization of the fields from the fragments. Several ant species were limited to forest fragments after harvest but occurred later in the season also in sugarcane fields confirming the potential contribution of fragments to the recolonization process and therefore to biological control of sugarcane-dominated pest insects.     

Arable weeds in organically managed wheat fields foster carabid beetles by resource- and structure-mediated effects

Arable weeds in organically managed fields may foster arthropod generalist predators by the provision of shelter and favorable microclimate (structure-mediated effects) and the provision of additional animal and floral food resources (resource-mediated effects). In three organically managed winter wheat fields in Central Germany, we investigated the impact of weed removal and introduction of artificial weed-like structure on the activity density and species richness of carabid beetles with respect to trophic groups, microclimatic conditions, and densities of potential prey. Removal of weeds reduced both carabid activity density and species richness but did not affect trophic group composition. The decline in carabid activity density was dampened by the addition of artificial structure. Mean daily surface temperature and light intensity were significantly lower under weeds and artificial plants than under wheat plants alone. Weed removal reduced the abundance of leafhoppers and true bugs, but the response was inconsistent across fields. We conclude that the presence of arable weeds in organically managed wheat fields fosters carabid activity density and species richness via resource-mediated effects, such as a higher availability of weed-borne resources (e.g. seeds and pollen) and herbivorous prey. Structure-mediated effects (altering the microclimate) add to this positive effect. The presence of weeds in organically managed wheat fields enhances carabid activity density and diversity and needs to be integrated into future management strategies for natural enemy conservation.     

Temperature and rate of seed consumption by ground beetles (Carabidae)

Predation is a key source of seed mortality in many weed species and thus is a part of natural control. In the field, the intensity of seed predation by invertebrates varies during the course of a year. One source of this variation is fluctuations in ambient temperature. Here, the effect of temperature on seed consumption is investigated for the first time, using two abundant carabid seed predators, Pseudoophonus rufipes and Harpalus affinis (Coleoptera: Carabidae), and dandelion (Taraxacum officinale) as a model system. Field collected individuals were sexed, kept at one of six constant temperatures between 10 and 28 °C and provided with a surplus of seed. Seed consumption was recorded over a period of 4 days. Averaged over all the temperatures, the smaller H. affinis consumed 12.2 seeds day^?1 and larger P. rufipes 29 seeds day^?1. On average, females consumed more seeds than males. Seed consumption by both species increased with temperature. In H. affinis the increase was linear and different for males and females. In P. rufipes the consumption was similar in both sexes but curvilinear because there was no further increase in consumption above 20 °C. Assuming a linear relationship between temperature and consumption at up to 20 °C we calculated the temperature at which seed consumption ceased (?0.1 to 0.3 °C in H. affinis and 6.3–6.9 °C in P. rufipes) and the increment in seed consumption per 1 °C increase in temperature above this threshold (0.4–1.0 and 1.5–4.2 seeds individual^?1 day^?1, respectively) for the two species. Thus, it is possible to calculate the average daily consumption of each species over a range of temperatures up to 20 °C.