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.     

Testing abundance-range size relationships in European carabid beetles (Coleoptera, Carabidae)

Four of the eight hypotheses proposed in the literature for explaining the relationship between abundance and range size (the sampling artifact, phylogenetic non-independence, range position and resource breadth hypotheses) were tested by using atlas data for carabid beetles (Coleoptera, Carabidae) from Belgium, Denmark and the Netherlands. A positive relationship between abundance and partial range size was found in all three countries, and the variation in abundance was lower for widespread species. Analysis of the data did not support three of the proposed hypotheses, but did support the resource breadth hypothesis (species having broader environmental tolerances and being able to use a wider range or resources will have higher local densities and be more widely distributed than more specialised species). Examination of species' characteristics revealed that widespread species are generally large bodied, generalists (species with wide niche breadths occurring in a variety of habitat types) and are little influenced by human-altered landscapes, while species with restricted distributions are smaller bodied, specialists (species with small niche breadths occurring in only one or two habitat types), and favour natural habitat. Landscape alteration may be an important factor influencing carabid abundance and range size in these three countries with a long history of human-induced environmental changes.     

Long adult life, low reproduction and competition in two sub-Antarctic carabid beetles

ABSTRACT. 1. On the biologically impoverished sub-Antarctic Ile de la Possession, Iles Crozet, Amblystogenium pacificum Putzeys and A.minimum Luff (Carabidae), both opportunistic carnivores, coexist in fellfield habitat above 120 m altitude, where they hunt prey in inter-stone spaces.
2. Beetles show marked claw wear showing that many individuals survive to breed in their third or later year of adult life, and in some years, particularly in A.minimum , populations consisted largely of very old beetles. This coupled with evidence from ripe egg numbers, of rarity of tenerals and of larvae, shows that adult recruitment rates were usually very low.
3. In fellfield the coexisting species pair have non-overlapping size distributions with A.pacificum the larger. In moorland A.pacificum adults occur in isolation and are smaller and overlap the size ranges of both species in fellfield. These facts are interpreted as character displacement or release, reflecting present-day interspecific competition, probably for food. Supporting evidence was obtained in terms of apparent size-differential survival of beetles of different ages.     

Patterns in the distribution, abundance and body size of carabid beetles (Coleoptera: Caraboidea) in relation to dispersal ability

The effects of dispersal ability, measured as two wing size categories (brachypterous vs. macropterous), on the distribution, abundance and body size, and on the relationships between these variables were examined in eighty-four species of carabid beetles over twenty-two sites in the northern Iberian peninsula. Geographic ranges of species (restricted to the northern Iberian peninsula vs. widespread—European or wider range) were also taken into account in the analyses because macropterous species significantly tended to exhibit wider geographic ranges than did brachypterous species. Regional distributions were wider in brachypterous-restricted and brachypterous-widespread species than in macropterous-widespread species. The three groups did not differ in abundance. Differences in regional distributions between groups may be explained by referring to a trade-off between dispersal ability and establishment ability indicated in the literature. Macropterous species would occupy relatively few sites due to a high frequency of unsuccessful colonizations. The relationships between regional distribution and abundance were positive for all the three groups, brachypterous-restricted, brachypterous-widespread and macropterous-widespread species. The regression line for the last group showed a lower elevation than those for brachypterous-restricted and brachypterous-widespread species. This fact was probably due to differences in regional distributions between groups. No relationship between abundance and body size was significant. Regressions of regional distribution on body size were positive in brachypterous-restricted and brachypterous-widespread carabids, but the relationship was not significant in macropterous-widespread carabids. These results were interpreted in terms of differences in body size–dependency of travelling velocities between flying and running carabids.     

pH-preferences and habitat selection in carabid beetles

Summary Seven species of carabid beetles were examined in different pH-Orgeln for pH-preferences. Five of these species showed significant preferences for specific pH-fields. Presumably, this parameter of distribution found in the laboratory is also effective in the field. We caught the beetles in the field on soil of the same pH-values which they prefered in the laboratory. The pH-measurements taken by previous workers in the field match our results.The receptors for H-ions are probably located on the antennae, because the preference distribution of Pt. angustatus changed into a uniform distribution after amputation of the distal segments of the antennae. The structure of these receptors could not be identified in electron microscope pictures (REM) among the multitude of different receptors on the antennae.     

Patterns of movement of radioactive carabid beetles

Summary Tracking of individual ¹⁹²Ir-labeled ground beetles released in the field revealed that both the day-active and night-active species studied showed periods of small distances covered per day in random directions, alternating with periods of directed movement with large distances covered per day. This pattern occurred not only in the reproductive period but outside the breeding season as well in juvenile Pterostichus versicolor and spent Calathus melanocephalus. Although mean locomotory activity increased with temperature, great daily differences occurred between individuals, pointing to asynchronous behavior. In an unfavorable habitat directed movement occurred both more frequently and more extremely, sometimes resulting in escape to more favorable areas. Most of the radioactive beetles died within 7 weeks due to radiation effects, but independent field experiments and simulations showed that the recorded patterns were valid. Simulated individuals of P. versicolor living on 1 ha spread over 49 ha, whereas simulated C. melanocephalus covered only 9 ha after one activity season. Normal locomotory activities lead to both exchange of individuals between subpopulations and dispersal out of the habitat. The significance of these phenomena for population stability and for the survival of the species is discussed.Communication no. 184 of the Biological Station of the Agricultural University (Wageningen), Wijster (Drenthe), The Netherlands     

On testing temporal niche differentiation in carabid beetles

Summary It is first shown that den Boer's (1985) claims that competition should be rejected as a factor explaining community structure are not warranted by his tests on temporal niche differentiation in carabid beetles. The power of these tests is so low that they are unlikely to detect even the most extreme pattern of niche differentiation. A major factor that affects their power is the procedure of building up an artificial species pool from the local communities to which the test is to be applied. Alternative tests are then presented, and applied to data on annual activity cycles of carabids in three forest biotopes in Belgium. Three factors are shown to strongly affect their outcome: the unit into which the time scale is divided, the criteria used to incorporate constraints, and the set of species selected. Nevertheless, they provide some evidence that there is a growing niche differentiation from the successional to the climax forest, as predicted by competition theory. Other hypotheses, however, could explain the observed pattern.     

Broad‐scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles(Coleoptera: Carabidae)

Research into large‐scale ecological rules has a long tradition but has received increasing attention over the last two decades. Whereas environmental, especially climatic, influences on the geographic distribution of species traits such as body size are well understood in mammals and birds, our knowledge of the determinants and mechanisms which shape spatial patterns in invertebrate traits is still limited. This study analyzes macroecological patterns in two traits of the highly diverse invertebrate taxon of carabid beetles: body size and hind wing development, the latter being directly linked to species’ dispersal abilities. We tested for potential impacts of environmental variables (spatial, areal, topographic and climate‐related) representing both contemporary conditions and historical processes on large‐scale patterns in the two traits. Regression models revealed hump‐shaped relationships with latitude for both traits in the categories 1) all species, 2) widespread and 3) endemic (restricted‐range) species: body size and the proportion of flightless species increased from northern towards southern Europe and then decreased towards North Africa. The shared and independent influence of environmental factors was analyzed by variation partitioning. While contemporary environmental productivity and stability (represented by measures of ambient energy and water energy balance) had strong positive relationships with carabid body size, patterns in hind wing development were most notably influenced by topography (elevation range). Regions with high elevation range and low historical climate variability (since the last ice age), which likely offer long‐term stable habitats (i.e. glacial refugia), coincide with regions with high proportions of flightless species. Thus geographic patterns in carabid traits tend to be formed not only by recent climate but also by dispersal and historical climate and processes (i.e. glaciations and postglacial colonization).     

National-scale metacommunity dynamics of carabid beetles in UK farmland

1. Understanding the wide-scale processes controlling communities across multiple sites is a foremost challenge of modern ecology. Here, data from a nation-wide network of field sites are used to describe the metacommunity dynamics of arable carabid beetles. This is done by modelling how communities are structured at a local level, by changes in the environment of the sampled fields and, at a regional level, by fitting spatial parameters describing latitudinal and longitudinal gradients. 2. Local and regional processes demonstrated independent and significant capacities for structuring communities. Within the local environment, crop type was found to be the primary determinant of carabid community composition. The regional component included a strong response to a longitudinal gradient, with significant increases in diversity in an east-to-west direction. 3. Carabid metacommunities seem to be structured by a combination of species sorting dynamics, operating at two different, but equally important, spatial scales. At a local scale, species are sorted along a resource gradient determined by crop type. At a wider spatial scale species appear to be sorted along a longitudinal gradient. 4. Nation-wide trends in communities coincided with known gradients of increased homogeneity of habitat mosaics and agricultural intensification. However, more work is required to understand fully how communities are controlled by the interaction of crops with changes in landscape structure at different spatial scales. 5. We conclude that crop type is a powerful determinant of carabid biodiversity, but that it cannot be considered in isolation from other components of the landscape for optimal conservation policy.     

Spatial dynamics of two introduced species of carabid beetles on the sub-Antarctic island of South Georgia

On the sub-Antarctic island of South Georgia two species of predatory beetle, Trechisibus antarcticus and Oopterus soledadinus (Coleoptera, Carabidae), were accidentally introduced. The colonisation process offers unique opportunities for testing ecological hypotheses in the field. As a basis for such studies, the spatial dynamics of the two species in the coastal lowland around Stromness Bay and Cumberland Bay were monitored during the period 1988–1996. Data are presented on the expansion of the two species and on the thermal characteristics of the tussock-forming grass Parodiochloa flabellata, which dominates the coastal lowlands. The largest expansion was shown by T. antarcticus, occurring as two populations in the Stromness Bay area. The origin of one of the populations, discovered in 1982, is Husvik Harbour, from where the species has colonised the central part of Stromness Bay. The second population, discovered in 1988 at Harbour Point, is invading the northern part of the area around Stromness Bay. Up to the present, three populations of O. soledadinus have been discovered. One population was found in 1988 at Husvik Harbour, co-existing with T. antarcticus, from where it is slowly expanding its distribution into the coastal zone. A second, very small, population of O. soledadinus was found in 1996 at Jason Harbour (Cumberland West Bay). The largest population of O. soledadinus, first reported in 1963, inhabits the area around Grytviken and King Edward Point (Cumberland East Bay), where it is the sole carabid species. The tussock grass vegetation offers a highly stable thermal environment compared with the more variable surroundings. Together with an ample food supply in the form of small arthropods and beetle larvae, and a vacant niche for arthropod predators, the benign microclimate of the tussock vegetation may explain the success of these predator introductions. Results of field observations on population parameters of the carabids, and laboratory measurement of egg production indicate the potential for large-scale co-existence. Accepted: 10 November 1998     

Correlation between variability and body size in vertebrates

An analysis of allozymic variation carried out in the main groups of vertebrate animals revealed a tendency towards the increased level of genetic polymorphism in the species of small animals compared to the large ones. This tendency was clearly followed in caudate amphibians, fishes, and mammals. The data are discussed in terms of the integration of monogenic and polygenic systems in the populations. It is hypothesized that this relationship between heterozygosity and body size confirms more general regularity consisting in highly statistically significant correlation between polygenic heterozygosity, maturation rate and life span. It is suggested that high rate of development in small animals resulting in early sexual maturation, can serve as a mechanism determining correlation between heterozygosity and body size at the species level. As a result, compared to large animals, small animals display higher rates of generation change, resulting in accelerated growth of population size and faster accumulation of genetic variability.     

The energetic equivalence rule rejected because of a potentially common sampling error: evidence from carabid beetles

Abundance data from pitfall traps are widely used to estimate the relationship between beetle body size and abundance. Such data probably overestimate densities of large bodied species and may overestimate slopes of size‐abundance relationships. Here, we test this idea by comparing size‐abundance patterns found using data from pitfall trapping with those found with data from a quantitative method of estimating abundance, quadrat sampling. We use data from a total of 47 communities. As expected, slopes of size‐abundance relationships are significantly more positive when estimated using data from pitfall traps compared to when using quadrat sampling data. This was seen when looking across different communities, within communities sampled by both methods and when focusing on the set of species found by both methods within a community. These results were also generally found regardless of method of analysis, which were done using regression with species values as independent data points and using the independent contrast method, and with slopes estimated using ordinary least square regression or the structural relation. Most important, slopes of size‐abundance relationships based on data from pitfall traps were on average significantly more positive than ?0.75 on log–log scales, and thus inconsistent with the energetic equivalence rule. Slopes based on quadrat sampling, on the other hand, were on average not significantly different from ?0.75. The rejection of the energetic equivalence rule based on data from pitfall traps here is therefore a sampling artefact. Similar problems may apply to abundance data from virtually all insect trapping methods, and should make us consider re‐examining many of the size‐abundance patterns documented so far. As a large proportion of all animal species are insects, and traps are widely used to estimate abundance, this is a potentially important problem for our general understanding of the relationship between species body size and abundance.     

Catches in pitfall traps in relation to mean densities of carabid beetles

Summary Based on continuous pitfall catches summed over the whole activity period for the carabid beetles Pterostichus versicolor Sturm and Calathus melanocephalus L., a satisfactorily linear relationship is presented between the mean densities in several habitats and in different years and the numbers of beetles trapped. This means that with the help of continuous pitfall sampling a reliable relative measure of the sizes of carabid populations can be obtained.Simulation experiments show that for individuals of P. versicolor the distances covered each day should be represented by strips of some width instead of by lines to obtain numbers trapped that equal the field data. The influence on numbers trapped of the high mobility of beetles in unfavorable habitats is probably not significant. Simulation experiments with C. melanocephalus indicate that a much lower capture efficiency than for P. versicolor has to be programmed to fit the simulation results with the field data.The catch from different types of pitfalls is not always proportional to trap perimeter due to different vegetation cover of the immediate trap surroundings and to different kinds of roofs placed over the pitfalls. For the same species the capture efficiency of a pitfall can also change throughout the season.Simulation experiments show that the number and shape of the pitfalls need not influence the catches, provided that the traps keep the same total perimeter and are broadly spaced.Remodelling of some data from the literature reveals that the linear relationship between mean densities and year-catches in pitfalls is probably a general one among carabid species. This leads to the hypothesis that carabid beetles have to work through a fixed reproductive program each year by means of an almost constant total amount of locomotory activity which is realized step by step during spells of suitable weather.Some recommendations are given for intraspecific and interspecific comparisons of pitfall catches between sites and years.Communication no. 196 of the Biological Station of the Agricultural University (Wageningen), Wijster (Drenthe), The Netherlands     

Clutch size manipulations in two seed beetles: consequences for progeny fitness

Seed beetles (Coleoptera: Bruchidae) lay their eggs on discrete resource patches, such that competition among larvae for food is an important component of their biology. Most seed beetles, including Stator limbatus, lay eggs singly on individual seeds and avoid superparasitism except when seeds are limiting. In contrast, S. beali, a closely related congener, lays eggs in clutches on a single seed. We tested the hypothesis that natural selection on larval life history characters favors small clutches (selection against large clutches) in S. limbatus, but that selection against large clutches is relaxed in S. beali because of the large size of its host's seeds. We manipulated clutch size and examined its relationship to offspring fitness. Clutch size affected the survivorship of S. limbatus larvae(r ²=0.14), but had no detectable effect on the survivorship of S. beali larvae (r ²=0.04). Also, clutch size had a large effect on development time and body weight of S. limbatus, but not of S. beali. We discuss the implications of this result for the evolution of clutch size in S. limbatus and S. beali.