Consequences of competition on the reproduction and mortality of three species of terrestrial slugs

The interaction of three species of terrestrial slugs was studied in six field cages containing monocultures and 2-way combinations. A. columbianus and A. ater were non-aggressive species and L. maximus was highly aggressive during the summer. All three species were similar in body design and resource utilization. Reproduction (egg size, eggs/batch, number of batches, wet-weight productivity) and mortality were monitored through the 1977 season. L. maximus drastically reduced reproductive success of the non-aggressive slugs and increased their mortality. There were also significant interactions between the non-aggressive species. Compared to other invertebrates, the life history characteristics of A. columbianus are strongly K-selected and probably evolved through intraspecific exploitative competition in predictable, stable forest habitats. A. ater is best classified as a refuging r-selected species adapted to less predictable, unstable field environments. L. maximus appears to be α-selected and its overall design is suited to density-dependent regulation in fields.     

Alternative risk-taking styles: The case of time-budgeting strategies of terrestrial gastropods

The terrestrial slugs Arion ater, Limax maximus and Ariolimax columbianus have similar morphological designs but differ remarkably in their life history tactics and behavioural time budgets. The adaptive value of particular risk-taking styles was investigated using a comprehensive computer simulation model. The model allowed each species' success (growth rates, food acquisition) and costs (distance travelled, hydration deficits, injury from aggressive encounters) to be evaluated in various types of weather (benign, harsh and surprise). This was interpreted in terms of the species' life history design. In addition, each species was simulated with the behavioural strategy of the other two species substituted for its natural programming. The simulation experiments demonstrated how variations in a few simple rules could lead to divergent cost-benefit consequences. The model also illustrated that coarsely-tuned sensitivity to the environment may be better than finely-tuned responses, depending on the animal's ability to respond quickly and the degree of risk. The simulations suggested that competition has shaped the time-budgeting tactics of A. ater and L. maximus. Each species actually performed better in harsh weather, using the behavioural program of the other. L. maximus is an aggressive species with a narrow, nocturnal activity period. The concentrated activity of L. maximus may allow it to displace competitors more effectively, whereas the broader time span of A. ater's activity may be necessary to avoid L. maximus.     

The effects of food and density on the nocturnal behaviour ofArion ater andAriolimax columbianus (Pulmonata: Stylommatophora)

The effects of food supply and population density on the nocturnal behaviour of Arion ater and Ariolimax columbianus were investigated. Density did not significantly affect A. ater's level of activity or short-term movement, resting, or feeding. A. columbianus was more active and moved, rested, and fed more frequently when slug density was high. A. ater foraged and rested more often, but fed less when good food was unavailable. Ariolimax's only response to the food regime was to feed more when good food was available. Seasonal changes in the level of activity and behaviour of Arion were evident, whereas Ariolimax's activity and behavioural repertoire were not similarly affected. Arion ater's nightly activity appeared to be mainly food oriented, while Ariolimax columbianus seemed most responsive to slug density during its nocturnal activity periods.     

The effects of food and density of the movement ofArion ater andAriolimax columbianus (Pulmonata: Stylommatophora) between habitats

Experiments were conducted to determine if slug density, or food quantity and quality could cause dispersal by slugs. Arion ater dispersed quickly from overcrowded areas, where high mortality was incurred by this species. Poor food conditions, caused more dispersal than good food conditions, but food had no effect on the mortality of A. ater. Neither food nor density factors had significant effects on the dispersal of Ariolimax columbianus. However, more A. columbianus died in the high- than low-density treatments. A. columbianus mortality was unaffected by the food factor. Dispersing Arion were heavier than non-dispersers, but there was no difference between the weights of dispersing and non-dispersing Ariolimax. There were significant seasonal changes in the dispersal rate, mortality and weight of dispersing Arion, whereas only the mortality of Ariolimax changed significantly during the experiments. Since Arion ater did not markedly reduce its activity during unsuitable weather it was able to leave areas of poor food and high slug density fairly rapidly. Ariolimax columbianus' large size allowed it to remain inactive during unfavourable weather and permitted this species to be relatively unresponsive to the experimental treatments.     

Does prey preference change as a result of prey species being presented together? Analysis of prey selection by the predatory mite Typhlodromus pyri (Acarina: Phytoseiidae)

Summary Prey-selection behaviour of the phytoseiid mite Typhlodromus pyri Scheuten was analysed with a Markovtype model of feeding-state dynamics and feeding-state dependent searching behaviour (Sabelis 1981, 1986, 1989; Metz and Van Batenburg 1985a, b). All behavioural characteristics of the predator which are independent of the feeding state were represented by one parameter. The remaining feeding-state dependent characteristics were represented by a function of the feeding state, with one parameter. The best parameter values to describe a predator-prey interaction were determined by fitting the model to the predation rates in monocultures. Under the assumption that the parameter values are not dependent on the composition of prey species supply, the diet of the predators in mixed cultures was predicted from parameters estimated in monoculture experiments.Two prey types, apple rust mite (Aculus schlechtendali (Nalepa)) adults and European red spider mite (Panonychus ulmi (Koch)) larvae were studied. A large discrepancy was observed between calculated and experimentally determined predation rates of T. pyri in mixed cultures: the predators actually killed 3–7 times more P. ulmi larvae than was predicted by the model.The large difference between observed and predicted predation rates in mixed cultures cannot be explained by changes in the behaviour of the prey species as a result of being together. Therefore, it seems likely that the prey selection behaviour of the predator was different when prey species were presented together than when presented singly. Apparently the predatory mite T. pyri prefers P. ulmi to S. schlechtendali.     

The adaptive significance of behavioural ontogeny in some. centrarchid fishes

Synopsis Studies on the ontogeny of behaviour in fish have seldom considered the adaptive significance of the order of appearance of the behaviours. Results of laboratory studies and field observations on the ontogeny of feeding, predator-avoidance, and agonistic behaviour in largemouth bass, Micropterus salmoides, rock bass, Ambloplites rupestris, and pumpkinseed, Lepomis gibbosus, young indicate that the order of appearance of the various behavioural systems enhances the survival of the young. In the laboratory, all species spend significantly more time feeding during the first weeks of free-swimming than the later weeks. During a corresponding period in the field the young are either occupying an offshore area low in predators (rock bass, pumpkinseed) or are being guarded by a parent (largemouth bass); thus the risk of predation is low. When the young are in a predator-rich environment (inshore) both the predator-avoidance response and agonistic behaviour are well developed. Agonistic behaviour is the last to appear and may serve to disperse the young. Dispersal may relate to the feeding mode of the various species and may also reduce the probability of predation. It is apparent that the sequential onsets of the behavioural systems are in concert with ecological events and selective pressures confronting centarchid young.     

Intrinsic and extrinsic drivers of activity budgets in sympatric grey and harbour seals

Investigation of activity budgets in relation to seasonal, intrinsic (age, sex) and extrinsic (time of day, spatial) covariates enables an understanding of how such covariates shape behavioural strategies. However, conducting such investigations in the wild is challenging, because of the required large sample size of individuals across the annual cycle, and difficulties in categorising behavioural states and analysing the resulting individual‐referenced and serially correlated data. In this study, from telemetry tags deployed on 63 grey seals Halichoerus grypus and 126 harbour seals Phoca vitulina we used behavioural data, and movement data within a Bayesian state–space model (SSM), to define population‐level activity budgets around Britain. Using generalised estimating equations (GEEs) we then examined how time spent in four states (resting on land (hauled out), resting at sea, foraging and travelling) was influenced by seasonal, intrinsic and extrinsic covariates. We present and discuss the following key findings. 1) We found no evidence that regional variation in foraging effort was linked to regional population trajectories in harbour seals. 2) Grey seals demonstrated sex‐specific seasonal differences in their activity budgets, independent from those related to reproductive costs. 3) In these sympatric species there was evidence of temporal separation in time hauled out, but not in time foraging. 4) In both species, time spent resting at sea was separated into inshore (associated with tidal haul out availability) and offshore areas. Time spent resting at sea and on land was interchangeable to some extent, suggesting a degree of overlap in their functionality. This may result in a relaxation of the constraints associated with a central place foraging strategy. More generally, we demonstrate how a large dataset, incorporating differing tag parameters, can be analysed to define activity budgets and subsequently address important ecological questions.     

Ecology and phenology of cattle ticks in Zambia: Development and survival of free-living stages

A study on development and survival of free-living stages of three important cattle ticks in Zambia,Amblyomma variegatum Fabricus,Boophilus decoloratus Koch, andRhipicephalus appendiculatus Neumann, was carried out to complement studies on seasonal dynamics of parasitic stages.Different instars of engorged ticks were exposed under quasi-natural conditions according to the season in which they occur naturally. Generally, development rates of all stages of the three species were related to temperature, whilst the duration of survival was influenced mainly by rainfall and consequent relative humidity.Observations on the effect of age and climate on the behaviour of ticks on pastures were also made. BothA. variegatum andR. appendiculatus completed only one generation per year. InA. variegatum, engorged females detaching early in the adult season (August to October) undergo morphogenetic diapause. Adults ofR. appendiculatus emerging between August and October enter a period of behavioural diapause before becoming active in December. These mechanisms effectively synchronize the life-cycles of these two univoltine species. The one-host tick,B. decoloratus, is able to complete three to five generations each year with no indication of seasonal synchronization.     

Behavioural and physiological stress responses to handling in wild guanacos

Both ethology and physiology can provide insights about stress conditions in wild animals during handling. Lama guanicoe (guanaco) is the major large herbivore inhabiting arid zones in Argentina. Since 2005, the Payún Matrú Cooperative, a business established and run by native people, has captured and handled guanacos for shearing at Payunia Reserve, Mendoza Province, Argentina, to market the species’ fibre for clothing and other products. The aim of our study was to evaluate the behavioural and physiological stress responses to handling of wild guanacos during these shearing activities. We evaluated stress conditions in guanacos by monitoring the frequency of agonistic behaviour in the holding corrals, following capture, the rate of vocalisation and agonistic behaviour during handling, and by estimating cortisol levels in handled animals. Crowding increased agonistic behaviour; this appeared to be related to dominance conflicts between males in the corrals. Time of corralling increased rate of agonistic behaviour and vocalisations during handling. The rate of behavioural stress responses was negatively related to cortisol levels in males; which may be due to animal fatigue or because behavioural stress responses may help mitigate physiological stress responses. Incorporating our results into animal welfare practices could help reduce stress conditions in wild guanacos. Our results suggest that both behavioural and physiological responses are important measures of stress in handled animals, and that behavioural stress responses cannot be used as a simple surrogate for glucocorticoid levels.     

Spatial distribution, resource utilisation and intraspecific competition in the dung beetle Aphodius ater

Competitive interactions in northern temperate dung beetles are poorly understood. This investigation therefore comprises a series of field and experimental work on a dung beetle species common in northern Europe, Aphodius ater, with special focus on intraspecific competitive interactions. The between-pat distribution of adult A. ater in relation to the age of sheep dung pats was studied in the field. The distribution of both sexes was contagious in the fresh pats but became more regular with increasing pat age. The successional occurrence of males and females did not differ, but immature females tended to occur in fresh pats while mature females were mainly found in older pats. With increasing age of pats, the egg load of females also increased. Egg-laying behaviour of the beetles was studied in laboratory experiments. The mean number of eggs laid per female per dung pat decreased with increasing beetle density. Thus, density-dependent processes seem to regulate resource utilisation with regard to breeding behaviour, resulting in equal exploitation of the available pats. Survival and weight of recently hatched beetles decreased with increasing initial density of eggs. Hence, in A. ater, competition between larvae for food within pats does occur. Received: 4 February 1998 / Accepted: 20 April 1998     

Behaviour-Genetics of Lake Charr (Salvelinus namaycush) and Brook Charr (S. fontinalis): Observation of Backcross and F2 Generations

Brook charr (Salvelinus fontinalis) defend territories and show high levels of agonistic behaviour. In contrast, lake charr (S. namaycush) are non-territorial and rarely show any agonistic behaviour. The genetic determinance of behavioural differences between these species was confirmed by the behavioural similarity of brook charr backcross hybrids and brook charr and a significant maternal effect on hybrid behavioural phenotypes. Backcross and F₂ generations showed either one of two distinct behavioural strategies, one aggressive and territorial and the other non-aggressive and non-territorial. It seems most likely that these strategies are conditional on individual phenotypes, such as size of nearby conspecifics. However, precise measurements of costs and benefits of each strategy are required to distinguish between these alternate hypotheses for either species. Each action pattern was an independent pattern of variation and a distinct behavioural unit. However, patterns were coordinated with all other patterns at a higher hierarchical level (corresponding to the particular behavioural strategy). It seems most likely that these strategies would be controlled by a polygenic system, even though the nature of our data does not allow us to make firm conclusions in this regard.     

The distribution and abundance of benthic cyclopoid copepods in Esthwaite Water,Cumbria

The spatial and temporal distribution of the benthic cyclopoid copepods, Acanthocyclops viridis, Acanthocyclops bicuspidatus, Eucyclops agilis, Paracyclops fimbriatus and Macrocyclops albidus in a small eutrophic lake Esthwaite Water, Cumbria was investigated in 1982 and 1983. The behavioural and physiological mechanisms by which these organisms cope with the conditions of hypoxia and anoxia prevalent during seasonal stratification of the lake waters was considered. During stratification all of these species, with the exception of resting stage A. bicuspidatus, disappeared from the profundal zone and were limited to the shallower margins of the lake. None of these copepod species entered the plankton and established planktonic populations, none appeared capable of sustained anaerobic respiration, although all can withstand some degree of hypoxia, and only A. bicuspidatus appeared capable of entering resting stages. The data suggest that in eutrophic lakes species normally found in the profundus deal with anoxia by lateral migration to shallow waters where oxygen is available.     

Behavioural adaptations of ground living bushcrickets to the properties of sound propagation in low grassland

Summary The acoustic behaviour of the closely related tetigoniid species Psorodonotus illyricus and Decticus verrucivorus have been invectigated by bioacoustical and behavioural methods. Both species show adaptations concerning the acoustic behaviour with respect to the biotope and the properties of sound propagation. These insects inhabit low grassland with an average vegetation height of about 20 cm which is also the general height for the song perches. Difficulties arise for efficient acoustic communication in such habitats. Sound propagation is influenced and limited by the strong ground attenuation and the excess damping by grass vegetation. Other limiting factors are the microclimatic conditions in the biotope. The two species counteract these difficulties by moving around in the biotope during stridulation. Both species mainly stridulate in the morning, avoiding problems of reduced sound transmission which often appear in the afternoon due to negative temperature gradients and resulting shadow zones. From the high mobility of these insects, it follows that individuals have no fixed territory and consequently no rivalry against conspecifics, which is very common among Orthopterans with a high degree of territoriality. It can be concluded that the preferred biotope influences and creates behavioural patterns in Orthopterans, especially here in the two investigated species of bushcrickets Psorodonotus illyricus and Decticus verrucivorus.     

Feeding ecology of a vertebrate assemblage inhabiting a stream of NW Spain (Riobo; Ulla basin)

The food resource use of a stream in NW Spain by fish (Salmo trutta L. and Anguilla anguilla L.), birds (Cinclus cinclus L. and Motacilla cinerea L.) and mammals (Galemys pyrenaicus G. and Neomys anomalus C.) was studied. Data on seasonal diets and stream benthos prey were used to determine prey selection patterns.Caddisfly larvae are the main resource for Cinclus and Galemys, but these predators also consumed other benthic prey. Salmo fed on a wide range of benthic invertebrates, emergent pupae and terrestrial prey, whereas Anguilla consumed primarily benthic invertebrates, especially Lumbricids. Neomys fed mainly on terrestrial prey (Gasteropods and Lumbricids), but also consumed aquatic prey. Motacilla captured aquatic insects both in larval and aerial stages, as well as terrestrial prey.Both prey availability and selection led to seasonal differences in the use of food resources. All species showed a marked prey selection of aquatic taxa. Prey size plays an important role in this selection, most species consuming the largest of available prey sizes. In spite of the fact that all species feed upon freshwater invertebrates, substantial resource partitioning was observed in all seasons. This partitioning may be attributable to morpholological and physiological differences. Nevertheless, Anguilla and Galemys, two quite different animals, did feed on the same prey much of the time.     

Small-scale vegetation patterns in the parental environment influence the phase state of hatchlings of the desert locust

Desert locusts (Schistocerca gregaria Forskål (Orthoptera: Acrididae)) change phase in response to population density. Solitarious insects avoid one another; when crowded, they shift to the gregarious phase and aggregate. Laboratory experiments and individual‐based modelling have shown that small‐scale resource distribution can affect locust phase state via an influence on crowding. Laboratory work has also shown that parental phase state is transmitted to offspring via maternal inheritance. These effects had not been investigated in the field previously. We maintained small populations of adult desert locusts in semi‐field enclosures with different distribution patterns of a single plant species (Hyoscyamus muticus L. (Solanaceae)). The offspring of locusts exposed to more clumped patterns of vegetation exhibited more gregarious behaviour when tested in a behavioural phase assay than did progeny from parents left in enclosures with more scattered vegetation. These effects on nymphal behaviour appeared to be mediated by influences of resource distribution on adult phase state. Phase state in small semi‐field populations was influenced by small‐scale vegetation distribution. Phase differences engendered by environmental structure were maintained in time and transmitted to progeny.     

Surrounding species diversity improves subtropical seedlings’ carbon dynamics

Increasing biodiversity has been linked to higher primary productivity in terrestrial ecosystems. However, the underlying ecophysiological mechanisms remain poorly understood. We investigated the effects of surrounding species richness (monoculture, two‐ and four‐species mixtures) on the ecophysiology of Lithocarpus glaber seedlings in experimental plots in subtropical China. A natural rain event isotopically labelled both the water uptaken by the L. glaber seedlings and the carbon in new photoassimilates through changes of photosynthetic discrimination. We followed the labelled carbon (C) and oxygen (O) in the plant–soil–atmosphere continuum. We measured gas‐exchange variables (C assimilation, transpiration and above‐ and belowground respiration) and δ¹³C in leaf biomass, phloem, soil microbial biomass, leaf‐ and soil‐respired CO₂ as well as δ¹⁸O in leaf and xylem water. The ¹³C signal in phloem and respired CO₂ in L. glaber in monoculture lagged behind those in species mixture, showing a slower transport of new photoassimilates to and through the phloem in monoculture. Furthermore, leaf‐water ¹⁸O enrichment above the xylem water in L. glaber increased after the rain in lower diversity plots suggesting a lower ability to compensate for increased transpiration. Lithocarpus glaber in monoculture showed higher C assimilation rate and water‐use efficiency. However, these increased C resources did not translate in higher growth of L. glaber in monoculture suggesting the existence of larger nongrowth‐related C sinks in monoculture. These ecophysiological responses of L. glaber, in agreement with current understanding of phloem transport are consistent with a stronger competition for water resources in monoculture than in species mixtures. Therefore, increasing species diversity in the close vicinity of the studied plants appears to alleviate physiological stress induced by water competition and to counterbalance the negative effects of interspecific competition on assimilation rates for L. glaber by allowing a higher fraction of the C assimilated to be allocated to growth in species mixture than in monoculture.     

Temporal plasticity in thermal‐habitat selection of burbot Lota lota a diel‐migrating winter‐specialist

In this study, animal‐borne telemetry with temperature sensors was coupled with extensive habitat temperature monitoring in a dimictic reservoir, to test the following hypotheses: behavioural thermoregulation occurs throughout the year and temperature selection varies on a diel and seasonal basis, in a winter‐specialist diel‐migrating fish. Burbot Lota lota demonstrated nightly behavioural thermoregulation throughout the year, with a large seasonal shift between selection for very cold temperatures (<2° C) optimal for reproduction during the spawning period and selection for warmer temperatures (12–14° C) optimal for hunting and feeding during non‐reproductive periods. During daylight hours, while L. lota avoided habitats warmer than optimal for reproduction and feeding during the spawning and non‐reproductive periods, respectively, active selection was limited to selection for 4–6° C habitat during the prespawning period. Although behavioural thermoregulation explained the night‐time migration, behavioural thermoregulation only partially explained daytime behaviour, indicating that diel migration is best explained by a combination of factors. Thus, thermal‐habitat selection was a good predictor of night‐time habitat occupancy in a diel‐migrating species. Together, these results show that thermal‐habitat selection by fishes may be important throughout the year and a more seasonally plastic behaviour than previously recognized.     

Trait‐mediated effects of predation across life‐history stages in container mosquitoes

1. It was determined if the predatory midge Corethrella appendiculata Grabham imposes a fitness cost in a native mosquito, Ochlerotatus triseriatus Say, and an invasive mosquito, Aedes albopictus Skuse. The hypothesis that decreased activity of immature prey in the presence of predator cues is associated with life history costs through all life cycle stages was tested. 2. In experiment 1, individual larvae of O. triseriatus or A. albopictus were raised in the presence or absence of predation cues at two resource levels. Prey were video recorded to detect behavioural responses and to measure development time, size at emergence, and adult longevity. In experiment 2, prey populations were reared in similar environments and the frequency of predator cue additions was varied. 3. Only O. triseriatus reduced its activity in the presence of predation cues. Predation cues were associated with longer immature development times and shorter adult life spans in O. triseriatus, whereas in A. albopictus, the cues were associated with a larger size of emerging adults. 4. In the present study, it was found that behavioural modifications during the larval stage can affect mosquitoes through multiple stages of their complex life cycle. The species‐specific behavioural differences are probably attributable to the longer evolutionary history O. triseriatus has with predators, relative to the invasive A. albopictus.     

Host plant evaluation behaviour of Bemisia tabaci and its modification by external or internal uptake of imidacloprid

The behaviour of adult Bemisia tabaci Gennadius (Homoptera: Aleyrodidae) was observed on leaves of cotton, Gossypium hirsutum L., to study the process of host plant evaluation. Seven distinct behavioural elements were identified and recorded for a period of 20 min after initial contact with leaves. The behaviour of the whiteflies was observed on untreated leaves or on leaves treated with either foliar or systemically applied imidacloprid. Thus, contact with imidacloprid by whiteflies was either via the external cuticle or by ingestion, allowing determination of the effect of uptake route on host evaluation behaviour. Analysis of behavioural sequences and comparison of latencies, frequencies and durations of elements of behaviour between treatments revealed that host evaluation behaviour was similar on untreated and foliar treated leaves, but markedly different on those treated systemically. After their first probe, whiteflies on systemically treated leaves were much more active, spending significantly greater time walking and dabbing the labium tip onto the leaf surface. There was also a 50% reduction in the total time spent probing into systemically treated leaves, compared to the other treatments. This effect was the result of a shorter mean probe duration, and no effect of treatment was found on the frequency of probing. Pre-oviposition behaviour was significantly more common on the systemically treated leaves, although the number of eggs laid was similar on all treatments. These results also show that the host evaluation phase of adult female B. tabaci was dominated by probing. The behavioural sequence that leads to probing was affected significantly by imidacloprid when ingested, but not by direct contact with this compound on the leaf surface. The wider implications for the mechanisms of whitefly–host plant interactions are discussed.     

Immunocytochemical Localisation of Glycosaminoglycan-Like Activity in the Mucus and Associated Tissues of Terrestrial Slug Species (Gastropoda: Pulmonata)

Indirect immunofluorescence assays were conducted on cryotome sections of four terrestrial slug species from three distinct phylogenetic groups, Arion ater (L.), Arion hortensis (Férussac), Tandonia (Milax) budapestensis (Hazay), and Deroceras reticulatum (Müller) using monoclonal antibodies for two glycosaminoglycans (GAGs), heparan sulphate, and chondroitin sulphate. Specific staining for a heparan sulphate-like component was demonstrated in the foot and tail regions of each species and was particularly intense in A. ater and A. hortensis, notably in the epidermis and associated mucus-like material, and in mucus-like material from the pedal gland region of the latter species. Subsequent studies with A. ater confirmed the presence of heparan-sulphate–like activity in the caudal gland duct region. No evidence of specific staining for chondroitin sulphate-like activity was found in any of the slug species. This work suggests that a specific GAG, or a group of closely related GAGs, is a common component of mucus in a range of slug species and of different types of mucus, including trail (pedal) mucus and the more viscous mucus produced by the caudal gland.