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     

The life histories and population dynamics of two carabid species on a Dutch heathland

Summary We deal with the causes of the synchronously fluctuating numbers of subpopulations of the carabid species Calathus melanocephalus as compared with the asynchronously fluctuating numbers of subpopulations of the carabid Pterostichus versicolor. Both species continuously occupy a large heath area, Dwingelder Veld (1600 ha), in The Netherlands, and are studied there in the same localities with the same methods. Of the adults of C. melanocephalus, 90% do not cover more than 2 ha during the entire reproductive season, while 90% of adults of P. versicolor cover no more than 12 ha. In C. melanocephalus egg production in the field is usually similar to that under optimal feeding conditions in the laboratory, but in P. versicolor egg production seems to be much lower in the field. In the field 70–80% of the eggs most probably are killed by eelworms, followed by more than 90% mortality among the remaining larvae. Comparing mortality of developmental stages in laboratory experiments with that in field experiments in enclosures, it appears that mortality of larvae is not density-dependent, even when density in the experiments is much higher than it ever is in the field. Larval mortality mainly results from the poor ability of the larvae to find prey, even when in field experiments prey density is increased far above natural densities. We discuss why these poor prey-finding abilities are not improved by natural selection. In the spring breeder P. versicolor differences between localities both in abiotic factors, soil moisture and surface temperature, and biotic factors, reactions of prey species to abiotic factors, in spring and summer when the larvae are maturing contribute to the asynchronous fluctuations of numbers between subpopulations. In the autumn breeder C. melanocephalus possible differences in biotic factors between sites are outnumbered by the effects of winters with a higher or lower than normal amount of precipitation respectively. During a wet winter mortality among the larvae is much higher than during a dry winter. As these winter conditions are similar over large areas (many km²) the fluctuations of numbers between subpopulations are synchronous.Communication No. 443 of The Biological Station, Wijster     

On the relationship between reproduction,age and survival in two carbid beetles:

Summary In this study the inverse relationship between the amount of reproduction in female carabid beetles and their survival until the next breeding season, suggested by Murdoch (1966), was examined in Pterostichus coerulescens L. and Calathus melanocephalus L. This was tested with individual and differently reproducing females, kept at the same temperature, as well as with groups of females kept at different temperatures, i.e. with very different overall levels of reproduction.No relationship was found between the numbers of eggs laid by individual females and their survival until the next breeding season.The numbers of eggs laid by individual females varies greatly, even under constant and optimal conditions,. In contrast to this, an individual female continues to lay a characteristic number of eggs over several succeeding weeks. The amount of reproduction is highly correlated with temperature when food is optimal. In P. coerulescens there was a positive correlation between the amount of reproduction of the same individuals in two succeeding years, whereas in C. melanocephalus no relationship could be found.Both in the field and in our experiments individuals of P. coerulescens generally live for 3 to 4 years, longer than C. melanocephalus which usually only lives for 2 years. The 3–4 year-old individuals of P. coerulescens die during the breeding season or shortly afterwards, but also during the winter, whereas the 2–3 year-old individuals of C. melanocephalus die during the winter. Males and females show a similar mortality after breeding. The level of reproduction of a population depends on the proportion of old beetles, since especially the very old ones die before the end of the breeding season.We reject the hypothesis of Murdoch because of these data. However, in a different way from that of Murdoch, we also suggest that old beetles are of great importance for the survival of a population, namely that this form of heterogeneity of a population may enhance its stability under varying environmental conditions.Communication No. 204 of the Biological Station, Wijster     

Pattern of movement in and dispersal from a dutch forest of Carabus problematicus Hbst. (Coleoptera,Carabidae)

Summary Data in the literature that in the Netherlands Carabus problematicus is an autumn breeder with overwintering larvae and adults have been confirmed. Pitfall catches show two peaks of adult beetles. The first small peak of activity is realized in May, the significance of which is not yet understood. The second higher peak occurs from the end of August till the beginning of October: the reproduction period. In this period at least three age classes are present. Some females were shown to take part in reproduction for at least two seasons. Old females appeared to have less eggs in the ovaries than young ones. Larvae can be trapped throughout the winter. The new generation appears in July and in the beginning of August. They immediately take part in reproduction. C. problematicus shows noctural activity with a peak towards the end of the night.In field experiments the pattern of movement and the dispersal behaviour of marked individuals was investigated and related to the choice of habitat. It was shown that in the Netherlands C. problematicus prefers a forest habitat. Some individuals were found to disperse from their favoured habitat. The percentage of beetles that disperse out of the forest during the breeding season could be estimated at 4–8%. Dispersal increases in the second part of the breeding season. Females tend to disperse more frequently than males.The pattern of movement consists of two types: a random walk and a directed movement. It is suggested that the pattern of movement of individuals dispersing out of the forest is the directed movement. It could be shown that individuals released outside the forest were able to walk towards a forest silhouette. The significance of both the pattern of movement and the dispersal behaviour for the survival of the species is discuussed.Communication no. 212 of the Biological Station of the Agricultural University (Wageningen), Wijster (Drente), The Netherlands     

Movements and flight morphology in the endangered Large Blue butterflies

Movements and flight morphology of the endangered Large Blue butterflies Phengaris teleius and P. nausithous in southern Poland were studied with mark-release-recapture surveys and GIS analyses. Most individuals moved relatively small distances (<40 metres) within their habitat patches. Distances covered by both species were positively related to season progression and wing length, and negatively related to body mass. P. teleius movement distances slightly exceeded those of P. nausithous. In addition, females moved longer distances than males, although the difference was significant only in P. teleius. Morphological traits appeared to be good indicators of the inter-specific and inter-sexual differences in mobility. While P. teleius individuals were heavier than P. nausithous ones, they had considerably longer wings, which may explain longer movements in the former species. Similarly, females were heavier than males in both species, but they invested more in wing size, which is likely to compensate for the negative impact of body mass on movement distances. Our results indicate that combination of GIS analysis of movement distances recorded with mark-release-recapture methods and morphometric measurements taken in field during non-lethal handling of captured individuals proved useful for studying the mobility potential of the endangered insect species.     

Fecundity in relation to wing-morph of three closely related species of the melanocephalus group of the genus Calathus (Coleoptera: Carabidae)

Summary In two successive years the fecundity of the carabid beetles Calathus (Neocalathus) cinctus, C. (N.) melanocephalus and C. (N.) mollis was studied in relation to wing-morph and temperature. Differences were found between the three species in both egg production and timing and length of the oviposition period. In all species the fecundity of laboratory bred beetles was significantly higher than that of females collected in the field. Long-winged females of both cinctus and melanocephalus had significantly higher egg production than short-winged females, and they also tended to produce eggs over a longer period. In mollis only the fecundity of the long-winged morph was established. The observed lower relative fitness of the short-winged morph in both cinctus and melanocephalus contradicts the supposed increase of the frequency of this morph in ageing, more or less isolated, populations of these species. The loss of long-winged genotypes, resulting from flight activities, is considered the most plausible cause of the increase of short-winged beetles in ageing populations. The higher fecundity of macropterous females makes them especially suited for (re)establishing populations.Communication No. 429 of the Biological Station WijsterPresent address and address for offprint requests: Kortenburg 31, NL-6871 ND Rentum     

Age‐ and sex‐related differences in the spatial ecology of a dichromatic tropical python (Morelia viridis)

Abstract: Despite outnumbering their temperate counterparts, tropical snake species have been poorly studied. Yet, the few tropical species that have been studied show a variety of behavioural traits beyond those described in temperate species. Here we reveal both age and sexual differences in the movements of tropical green pythons (Morelia viridis: Pythonidae). We radio‐tracked 27 individuals (17 females and 10 males) for up to 18 months, locating individuals during both the day and night. The home range size for adult females (mean ± SE of 6.21 ± 1.85 ha) was correlated with snout–vent length. Neither adult males nor juveniles had a stable home range. Adult females had stable home ranges that overlapped considerably with those of other females and yellow individuals. Multiple radio‐tracked adult males passed through the territory of radio‐tracked adult females during the study. Females of all sizes were more likely to change position each day than males. There were no differences between the sexes or size categories in the distances moved in most months, although the variation in movement distances was higher in the dry season than during the wet season. In the wet season (January–March) movement distances increased and these were size‐ and sex‐related. This increased activity may be associated with mate searching. Males of M. viridis may maximize their rate of encountering mature females by roaming rather than maintaining a home range. Juvenile green pythons moved distances equal to adult snakes in most months despite their comparatively small size.     

Landscape and fine-scale movements of a leaf beetle: the importance of boundary behaviour

Movement underpins animal spatial ecology and is often modelled as habitat-dependent correlated random walks. Here, we develop such a model for the flightless tansy leaf beetle Chrysolina graminis moving within and between patches of its host plant tansy Tanacetum vulgare. To parameterize the model, beetle movement paths on timescales of minutes were observed in uniform plots of tansy and inter-patch matrix (meadow) vegetation. Movement lasted longer, covered greater distances and had narrower turning angles in the matrix. Simulations of the model emulated an independent two-season multi-patch mark–resight study at daily timescales and included variable boundary-mediated behaviour affecting the probability of leaving habitat patches. As boundaries in the model became stronger there were disproportionately large decreases in net displacements, inter-patch movements and the proportion of beetles in the matrix. The model produced realistic patterns of population-level displacement over periods up to 13 days with fully permeable boundaries for one dataset and strong boundaries for the other. This may be explained by the heights of the tansy patches in each study, as beetles will be unable to cross the boundary near the top of a patch that emerges from the matrix. The simulations demonstrate the important effects of boundary behaviour on displacement patterns and indicate temporal and spatial variability in permeability. Realistic models of movement must therefore include behaviour at habitat boundaries. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Importance of habitat patch size for occupancy and density of aspen-associated saproxylic beetles

In Fennoscandian boreal forests, aspen (Populus tremula) is one of the most important tree species for biodiversity. In this study we explore how occupancy and density of beetles associated with dead aspen are related to habitat patch size and connectedness in a 45,000 ha boreal managed forest landscape in central Sweden. Patch size was estimated as amount of breeding substrate and connectedness as crown cover of living aspen in the surrounding landscape. The beetles were sampled by sieving of bark or by inspection of species-characteristic galleries in 56 patches with dead aspen. Six of nine aspen-associated species (Xylotrechus rusticus, Ptilinus fuscus, Mycetophagus fulvicollis, Cyphaea curtula, Homalota plana and Endomychus coccineus) showed a positive significant relationship between habitat patch size and occupancy. For all these species, except C. curtula, there was also a significant positive relationship between patch size and density. Connectedness was not retained as a significant variable in the analyses. Species not defined as aspen-associated constituted a significantly larger proportion of the total density of individuals of saproxylic beetles in smaller habitat patches than in larger patches. Richness of aspen-associated species was positively related to habitat patch size. Efforts in the managed forest should be directed towards preserving and creating larger patches of living and dead aspen trees and increasing the amount of aspen at the landscape level.     

A comparison of the responses of two tropical specialist herbivores to host plant patch size

Summary The effects of host plant patch size on the abundances of two specialist herbivores (the chrysomelid beetle, Acalymma innubum and the pentatomid bug, Piezosternum subulatum) were investigated in a natural forest community in the Virgin Islands. Abundances were compared early and late in the season in different sized patches of the cucurbit host plant (Cayaponia americana) growing in open habitat (with no surrounding plant community) and forest habitat (with diverse surrounding plant community). For both herbivore species, adult abundances per patch were positively correlated with patch leaf area, but there was a significant patch size effect (i.e., correlation between herbivore density per unit plant and patch leaf area) only for beetles in the forest habitat. Both herbivore species were significantly affected by surrounding plant diversity, but in opposite ways: beetles were more abundant in open patches whereas bugs were more abundant in forest patches. Relationships between abundance and patch size in open and forest patches changed through the season for both herbivore species. These changing abundance patterns are discussed with respect to (1) increases in the diversity of the plant community surrounding host plant patches, and (2) differences in herbivore movement patterns.     

How far do adult turtles move? Home range and dispersal of Kinosternon integrum

We describe the home range and movements of a population of Kinosternon integrum in Tonatico, Estado de México, México, over 3.5 years (during rainy and dry season months) using radiotelemetry in 37 adult turtles. The results showed that the home range of K. integrum was 0.151 ± 0.051 ha using 50% kernel density estimator (KDE), and 0.657 ± 0.214 ha using 95% KDE; the home range did not vary between sexes. Kinosternon integrum showed low distances traveled 51.44 ± 4.50 m, where 87.3% (n = 373) of movements were <100 m. The distance traveled differed by season, and movement category (aquatic and terrestrial movements). The shortest distance occurred during the dry season, during which some individuals move to estivation sites, and these movements were shorter than movements to artificial ponds (cattle ponds). In this population, home range and movement are similar to other species of the genus Kinosternon. Overall, the results indicate than K. integrum are highly dependent on aquatic habitats, but also utilize the terrestrial habitats for different biological activities, and to maintain viable populations. Therefore, the conservation of the entire inhabited area is fundamental. This study highlights the need to increase the studies, in Central México, concerning habitat use of freshwater turtles in order to increase the efficiency of conservation strategies.     

Activity areas and daily movements of an arboreal monitor lizard,Varanus tristis (Squamata: Varanidae) during the breeding season

During the breeding season, in the western Great Victoria Desert, Western Australia, male Varanus tristis move greater daily distances (186.5 m) than females (99.7 m) and have larger activity areas (40.3 ha vs 3.7 ha). V. tristis retreat predominantly to live and dead upright Eucalyptus gongylocarpa (marble gum) trees that have hollows in their trunks and limbs. There is no observable difference between males and females in their preferred habitat, but females are found more frequently in a lesser number of preferred retreats during the breeding season.     

Limited dispersal and its effect on population structure in the milkweed beetle Tetraopes tetraophthalmus

Summary The movement patterns of adult milkweed beetles, Tetraopes tetraphthalmus, were monitored via a mark-recapture technique. Movement or dispersal patterns were studied in two natural populations, one in which the host plant, Asclepias syriaca, was nearly continuously distributed over a 250×90 m area and another where Asclepias was distributed in 17 small discrete patches. In both populations dispersal distances resulting from the flight patterns of the adult beetles were quite short, averaging less than 40 m from the point of first encounter 10 days after marking. Males were shown to be more vagile than females. The distribution of dispersal distances collected from one of the populations was fit to three statistical distributions cited in the literature as expected from dispersal by many small-scale movements or observed in other species. It was found that an equation describing an exponential decay gave the best statistical fit to the data collected here for milkweed beetles. The data is discussed in the context of the effects of the limited dispersal power of the beetles and the distribution of suitable habitat on the population structure of Tetraopes.     

Random movement pattern of fed and unfed adult Colorado potato beetles in bare‐ground habitat

• 1 Although the successful management of the Colorado potato beetle Leptinotarsa decemlineata (Say) depends on the prevention of its dispersal, its walking pattern in the landscape remains poorly understood. In the present study, post‐diapause, early summer, late summer and colony adult beetles, both fed and unfed before release, were tracked with a harmonic radar to establish their walking movement pattern in a bare‐ground field.
• 2 The random walk model successfully described the dispersal of all beetle types, whether fed or unfed.
• 3 The diverse life history of this species was manifested by an increased distance travelled and deviations of individual paths from the random model. Starved post‐diapause beetles travelled furthest and individual paths deviating from random were both local and directed, probably aiming to maximize opportunities for host colonization. Starved early summer beetles also travelled further than fed beetles but relied more on random movement to disperse in the habitat. Starving had little impact on the distance travelled or the path deviations of late summer beetles that are searching for overwintering site rather than hosts.
• 4 The increased displacement of starving beetles over fed beetles corresponded with an increased walking step and index of straightness.
• 5 The impact of starvation on travel distance was greater than expected from laboratory tests.
• 6 In conclusion, the results obtained in the present study suggest a random walking pattern to search arable land until host volatile or visual impulses trigger a more directed walk or flight.

     

Habitat use and dispersal characteristic by <Emphasis Type="Italic">Stethophyma grossum</Emphasis>: the role of habitat isolation and stable habitat conditions towards low dispersal

Species in a highly fragmented environment, such as the intensively used agricultural landscapes of Europe, are expected to be in danger of extinction. We hypothesize according to Kisdi’s theory (Am Nat 159:579–596, 2002) that species in fragmented landscapes with isolated habitats in general tend to possess low dispersal. In order to verify this hypothesis we studied the movement patterns of Stethophyma grossum, a hygrophilous species of wetlands, by mark–release–recapture techniques in a landscape with scattered suitable habitats over 3 years. The study focused on the major population in this landscape (site #1) as dispersal behaviour was assumed to be greatest. Actually, marked individuals of S. grossum were never found in any further suitable habitats in close vicinity to site #1. Despite that the peatland meadow of study site #1 was all over covered with homogenous vegetation only 6% (1.8 ha) of the whole area (30 ha) were occupied by S. grossum. The mean recapture rate over 3 years amounted to 39% with no significant differences between males and females. Both covered little distances within their mean range size of 1.8 ha; the median distances were 36.91 m for males and 26.65 m for females. We confirm the hypothesis that sub-populations of species in longstanding naturally isolated habitats, which habitat conditions have been stable; evolved low dispersal with little movements which are routine movements to find mating partners or food.     

The effect of translocation on movement behaviour—A test of the assumptions of behavioural studies

Animal movement behaviour is intensively investigated with capture-mark-recapture studies. For the analysis of such experiments, the influence of marking technique, handling and translocation of marked animals on movement pattern is of crucial importance since it may mask or overrule the effects of the main research question. Here we present a capture-mark-recapture experiment on the movement behaviour of the blue-winged grasshopper Oedipoda caerulescens. We analyzed the influence of translocation of individuals from familiar to unfamiliar sites. Our study clearly demonstrates a significant influence of translocation to unfamiliar sites on the movement behaviour of O. caerulescens. Translocated individuals moved longer distances, showed smaller daily turning angles, and thus movements were more directed than those of resident individuals. The effect of translocation on daily moved distances was most pronounced on the first day of the experiment. We thus conclude that at least for the first day after translocation, movement behaviour is significantly influenced by translocation itself. Data ignoring this influence will be unsuitable for the prediction of dispersal behaviour, habitat detection capability or habitat preference.     

Mobility and metapopulation structure of Geocrinia alba and Geocrinia vitellina,two endangered frog species from southwestern Australia

Abstract Dispersal rates can have an important impact on many population processes. Dispersal can lead to population regulation and regional stability in the face of local instability through the formation of a metapopulation. This may be particularly true for frogs because they often have patchy distributions. In this paper I investigate dispersal by male Geocrinia alba and Geocrinia vitellina, using a mark-recapture study. Pit traps were used to determine whether frogs move out of their breeding habitat. I found that both species were very philopatric. Between the 1993 and 1994 breeding seasons, 76–86% of individuals were displaced less than 10m and 90–97% were displaced less than 20m. Dispersal was even lower within each breeding season, with 92–95% of individuals being displaced less than 5 m. Pit trapping indicated that some individuals of both species move out of the swamps in late autumn and return at the beginning of the breeding season, in late winter and early spring. Therefore, although displacement may be very restricted, the frogs are likely to move greater distances. The extreme breeding-site philopatry exhibited by G. alba and G. vitellina suggests that movement between disjunct populations is unlikely and that populations in continuous habitat may be relatively isolated from one another by distance alone. This is consistent with previous predictions based on genetic studies, which suggested there was very little migration between populations. These data support the contention that neither species is likely to exist as a metapopulation because recolonization of vacant habitat is improbable.     

Defining core habitat of local populations of the gray treefrog (Hyla versicolor) based on choice of oviposition site

Concern over amphibian population declines and loss of terrestrial and aquatic habitat have emphasized the need to define habitat requirements for each stage in a species' life history. The realization that pond-breeding amphibians spend most of their lives in the terrestrial environment suggests the need to protect terrestrial as well as aquatic habitat. Many studies on amphibian populations have focused on emigration from breeding sites to define habitat use; however these studies do not typically elucidate terrestrial activities of adults within the breeding season. We measured colonization rates of artificial pools by gray treefrogs (Hyla versicolor) at multiple distances from natural breeding ponds. We found a non-random distribution of egg deposition among distances, with 95% of eggs deposited within 15 m of the breeding pond. Additionally, we found that the time to first colonization of artificial pools increased with respect to distance. Our results indicate that adult gray treefrogs may travel up to 200 m within a breeding season, and that multiple breeding ponds may be considered part of a single population. We suggest that a minimum core terrestrial habitat of 60 m surrounding breeding sites is appropriate for protection of local populations of gray treefrogs.     

Movement patterns and habitat use of three declining littoral fish species in a north-temperate mesotrophic lake

We explored patterns of habitat use and movement of three declining fish species intolerant to eutrophication in a north-temperate (Minnesota, USA) glacial lake: the blackchin shiner Notropis heterodon, blacknose shiner Notropis heterolepis, and banded killifish Fundulus diaphanus. We marked individuals with elastomer tags and estimated movement distances of recaptured individuals. Estimated home ranges for all species ranged from 3,264 to 19,525 m², which covered 0.8 and 5.0% of our study lake’s total littoral area. Individuals of all species traveled to opposite ends of the lake over periods of time as short as 24 h. Using Geographic Information System (GIS) overlays and generalized additive models, we found fish species occurrences to be positively associated with macrophyte biovolume greater than 20% and with a high probability of occurrence of Chara. The magnitude of main and interaction effects varied among years and species. Overall, blackchin shiner occurrence was most strongly associated with biovolume. In other species by year combinations, biovolume and Chara explained varying degrees of variance in fish probability of occurrence. Our results suggest that controlling lake eutrophication and protecting of refuge areas of dense macrophytes and Chara may be needed to conserve these species.     

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

Abstract.

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