Cold tolerance and freeze-induced glucose accumulation in three terrestrial slugs

Cold tolerance and metabolic responses to freezing of three slug species common in Scandinavia (Arion ater, Arion rufus and Arion lusitanicus) are reported. Autumn collected slugs were cold acclimated in the laboratory and subjected to freezing conditions simulating likely winter temperatures in their habitat. Slugs spontaneously froze at about − 4 °C when cooled under dry conditions, but freezing of body fluids was readily induced at − 1 °C when in contact with external ice crystals. All three species survived freezing for 2 days at − 1 °C, and some A. rufus and A. lusitanicus also survived freezing at − 2 °C. ¹H NMR spectroscopy revealed that freezing of body fluids resulted in accumulation of lactate, succinate and glucose. Accumulation of lactate and succinate indicates that ATP production occurred via fermentative pathways, which is likely a result of oxygen depletion in frozen tissues. Glucose increased from about 6 to 22 μg/mg dry tissue upon freezing in A. rufus, but less so in A. ater and A. lusitanicus. Glucose may thus act as a cryoprotectant in these slugs, although the concentrations are not as high as reported for other freeze tolerant invertebrates.     

Ranging behaviour and socio-biology of Eurasian otters (Lutra lutra) on lowland mesotrophic river systems

We examined the spatial structure and socio-biology of a native wild population of Eurasian otters (Lutra lutra) on mesotrophic rivers in a mild temperate climate. Radio-tracking of 20 individuals revealed exclusive intra-sexual adult home-ranges. Adult female home-ranges (7.5 km, SD = 1.5 km, n = 7) were inversely related to river width (, F ₆ = 13.5, P = 0.014) and so appeared to be based on food resources. The aquatic area within adult male home-ranges (30.2 ha, SD = 9.5 ha, n = 5) was greater than that within adult female’s (16.8 ha, SD = 7.0 ha) (t ₁₀ = 2.437, P = 0.035), though this result is inconclusive because some males were tracked on oligotrophic rivers. One adult male expanded its range from 10.2 km to 19.3 km within 5 days of the death of the neighbouring male, suggesting that male home-ranges were heavily influenced by conspecifics.     

Size-specific susceptibility of the pest slugs Deroceras reticulatum and Arion lusitanicus to the nematode biocontrol agent Phasmarhabditis hermaphrodita

The nematode Phasmarhabditis hermaphrodita is a commercially available biocontrol agent against slugs. This product is especially interesting for use in organic farming, where products containing metaldehyde or carbamates cannot be used for controlling pest slugs. We investigated the potential of P. hermaphrodita for the control of the pest slugs Deroceras reticulatum and Arion lusitanicus. These two species are the most harmful slug pests in Switzerland. At different times of the year, we collected slug specimens of different weight and assessed their susceptibility to P. hermaphrodita in the laboratory. Batches of five slugs were subjected to five different doses of nematodes plus an untreated control and replicated three times. During six weeks, feeding and survival of the slugs were recorded. D. reticulatum was strongly affected by increasing nematode doses, irrespective of the slugs' body weight. In small specimens of A. lusitanicus, feeding and survival were strongly affected by the nematodes, while larger specimens remained almost unaffected. Because A. lusitanicus has an asynchronous development in Switzerland, it seems difficult to control the entire population with a single nematode application. To what extent nematodes will be used in practice for slug control depends on their effectivity against the pest slugs of major importance, on the longevity of the molluscicidal effect and on the price of nematodes.     

Drought tolerance in eggs and juveniles of the Iberian slug, <Emphasis Type="Italic">Arion lusitanicus</Emphasis>

The spread of invasive species is an increasing problem world wide. The invasive slug Arion lusitanicus has spread to most parts of Europe, where it often is considered as a serious pest. There is a need for better knowledge of its ecophysiology to be able to predict the effect of climatic factors, such as temperature and humidity on the population dynamics and abundance. The aim of the present study was to assemble data on the water balance and drought tolerance of eggs and juveniles of A. lusitanicus. Both eggs and juveniles had little capacity to prevent evaporative water loss and lost water when the ambient humidity fell below 99.8 and 99.5%, respectively. The water conductance of the cuticle of juveniles was 242 μg cm⁻² h⁻¹ mmHg⁻¹ and resembles that of other slug species. Both eggs and juveniles of A. lusitanicus tolerate a substantial water loss. There was no difference in water loss resistance between eggs and juveniles, but the eggs were slightly more tolerant to water loss than the juveniles. The percent water loss causing 50% mortality was 72% for the juveniles and 81% for the eggs. Despite A. lusitanicus’ tolerance of substantial water loss, its survival depends on humid habitats.     

Diversity protects plant communities against generalist molluscan herbivores

Wildflower strips are used to increase natural enemies of crop pests and to conserve insect diversity on farmland. Mollusks, especially slugs, can affect the vegetation development in these strips considerably. Although recent theoretical work suggests that more diverse plant communities will exhibit greater resistance against herbivore pressure, empirical studies are scarce. We conducted a semi‐natural experiment in wildflower strips, manipulating trophic structure (reduction in herbivorous mollusks and reduction in major predators) and plant diversity (2, 6, 12, 20 and 24 sown species). This design allowed us to assess the effect of plant diversity, biomass and composition on mollusks, and vice versa, the effect of mollusc abundance on vegetation. Seven species of mollusks were found in the strips, with the slugs Arion lusitanicus, Deroceras reticulatum and Deroceras panormitanum being most frequent. We found a negative relationship between plant diversity and mollusk abundance, which was due predominantly to a decrease in the agricultural pest species A. lusitanicus. These results are consistent with the hypothesis that plant diversity can reduce the impact of herbivores. However, plant identity also had an effect on mollusks, and accounted for a much larger fraction of the variation in mollusk communities than biodiversity effects. While overall plant diversity decreased during the 3 years of the study, in the final year the highest plant diversity was found in the plots where mollusk populations were experimentally reduced. We conclude that selective feeding by generalist herbivores leads to changes in plant community composition and hence reduced plant diversity. Our results highlight the importance of plant biodiversity as protection against generalist herbivores, which if abundant can in the long term negatively impact plant diversity, driving the system along a “low plant diversity – high mollusk abundance” trajectory.     

Methylglyoxal upregulates Dictyostelium discoideum slug migration by triggering glutathione reductase and methylglyoxal reductase activity

Glutathione (GSH)-deprived Dictyostelium discoideum accumulates methylglyoxal (MG) and reactive oxygen species (ROS) during vegetative growth. However, the reciprocal effects of the production and regulation of these metabolites on differentiation and cell motility are unclear. Based on the inhibitory effects of γ-glutamylcysteine synthetase (gcsA) disruption and GSH reductase (gsr) overexpression on aggregation and culmination, respectively, we overexpressed GSH-related genes encoding superoxide dismutase (Sod2), catalase (CatA), and Gcs, in D. discoideum. Wild-type KAx3 and gcsA-overexpressing (gcsA^OE) slugs maintained GSH levels at levels of approximately 2.1-fold less than the reference GSH synthetase-overexpressing mutant; their GSH levels did not correlate with slug migration ability. Through prolonged KAx3 migration by treatment with MG and H₂O₂, we found that MG increased after the mound stage in this strain, with a 2.6-fold increase compared to early developmental stages; in contrast, ROS were maintained at high levels throughout development. While the migration-defective sod2- and catA-overexpressing mutant slugs (sod2^OE and catA^OE) decreased ROS levels by 50% and 53%, respectively, these slugs showed moderately decreased MG levels (36.2 ± 5.8 and 40.7 ± 1.6 nmol g⁻¹ cells wet weight, P < 0.05) compared to the parental strain (54.2 ± 3.5 nmol g⁻¹). Importantly, defects in the migration of gcsA^OE slugs decreased MG considerably (13.8 ± 4.2 nmol g⁻¹, P < 0.01) along with a slight decrease in ROS. In contrast to the increase observed in migrating sod2^OE and catA^OE slugs by treatment with MG and H₂O₂, the migration of gcsA^OE slugs appeared unaffected. This behavior was caused by MG-triggered Gsr and NADPH-linked aldolase reductase activity, suggesting that GSH biosynthesis in gcsA^OE slugs is specifically used for MG-scavenging activity. This is the first report showing that MG upregulates slug migration via MG-scavenging-mediated differentiation.     

The palatability of 78 wildflower strip plants to the slug Arion lusitanicus

As Arion lusitanicus has been observed in high densities in wildflower strips, the palatability of the foliage of 78 species of sown and naturally occurring plants found in such strips was investigated. In the laboratory, the slugs were offered a surplus of fresh, mature leaves and, after 16 h, a consumption index was calculated to rank the plants according to their palatability to the slugs. Arion lusitanicus showed clear preferences for particular plant species, but more than two thirds of all plants investigated were scarcely eaten. Brassica napus was most eaten, followed by Papaver rhoeas, Capsella bursa-pastoris and Lamium purpureum. Overall, A. lusitanicus preferred sown species to naturally occurring ones and preferred annuals to perennials.     

Body size and the colonisation of cereal crops by the invasive slug Arion lusitanicus

The invasive slug Arion lusitanicus lives in undisturbed stands of grass or herbaceous plants from where it disperses into crops. Factors that determined the dispersal of slugs from a grassy orchard (O) into an adjacent cereal stand were investigated. Slug‐feeding activity, abundance and body weight were established along a 120 m transect across an orchard and a cereal crop. Slugs in the source population in the orchard were more abundant and lighter in 2008 than 2009. The slugs that left the orchard and moved into the cereal crop were heavier than those that remained in the orchard and the distance they travelled was positively correlated with their weight. The abundance of slugs in the cereal crop decreased with the increase in the distance from the source population, more so in 2008, when slugs were light in weight and accumulated near the edge of the orchard, than in 2009 when the slugs were heavier and distributed more evenly. The feeding activity of the slugs leaving the orchard and distance they dispersed also increased during the course of the season. Long distance dispersal is achieved by crawling, the speed of which increased with slug size. Seasonal and annual variation in body size thus affected the distribution of slugs in the cereal crop and the distance they dispersed.     

Low temperature survival in different life stages of the Iberian slug, Arion lusitanicus

The slug Arion lusitanicus Mabille (Gastropoda: Pulmonata: Arionidae) is an invasive species which has spread to most parts of Europe. The area of origin is unknown, but A. lusitanicus seems to cope well with the local conditions in the countries to which it has migrated. It spreads rapidly, occurs often in high densities and has become a serious pest in most European countries. Therefore there is an urgent need for better knowledge of the ecophysiology of A. lusitanicus, such as the influence of climatic conditions, in order to develop prognostic models and strategies for novel pest management practises.The aim of our study was to investigate the influence of subzero temperatures in relation to winter survival. A. lusitanicus is shown to be freeze-tolerant in some life stages. Most juveniles and some adult slugs survived being frozen at −1.3 °C for 3 days, but none of the slugs survived freezing at −3 °C. The eggs survived subzero temperatures (down to −2 °C) probably by supercooling. Juveniles and adults may also survive in a supercooled state (down to −3 °C) but are generally poor supercoolers. Therefore, the winter survival of A. lusitanicus depends to a high degree on migration to habitats protected from low winter temperatures, e.g. under plant litter, buried in the soil or in compost heaps.     

Spatial and temporal movement of the Bearded Vulture using GPS telemetry in the Himalayas of Nepal

This study addresses for the first time the movement patterns of the globally near-threatened Bearded Vulture Gypaetus barbatus in its most important stronghold, the high-altitude mountain ranges of Asia. Tracked individuals (n = 8) in the Annapurna Himalayan range (Nepal) foraged over a vast range of 60 715.9 km² and our results indicated age-class differences in the use of space. Territorial adults showed very small annual home-ranges (K90 = 150.3 km²), whereas immatures wandered extensively and covered vast ranges of the mountains (K90 = 23 930.8 km²). For adults and immatures, these values are notably larger than the other two studied populations in the world (Pyrenees and South Africa). This suggests that the studied Annapurna population might exhibit lower breeding density than in the Pyrenees or South Africa, possibly due to lower food availability.     

Understory vegetation should not be ignored in the estimation of forest carbon stocks in Qilian Mountains National Nature Reserve

The Qilian Mountain National Nature Reserve is an important ecological barrier in northwestern China; therefore, it is of great importance to accurately evaluate the carbon storage of its main arbor forest. In this study, the Tianlaochi Basin of Qilian Mountain Nature Reserve was used as the research area to measure the carbon storage of the main arbor forests and determine the relationship between neglect of understory vegetation and errors in estimated carbon storages. The results showed that: (1) The total carbon storages of Picea crassifolia and Sabina przewalskii in Qilian Mountain Nature Reserve were 6.05 × 10⁷ and 9.48 × 10⁶ Mg and the carbon density were 426.07 Mg/hm² and 338.72 Mg/hm², respectively. (2) The understory carbon storage of Picea crassifolia and Sabina przewalskii accounted for 7.03% and 5.5% of the total carbon storage, respectively (6.81% combined). When the understory vegetation was neglected, the carbon storage in Qilian Mountain Nature Reserve was estimated to be less than 9.72 × 10⁵ Mg. (3) There are large areas of Picea and Sabina forests worldwide; therefore, ignoring understory vegetation in these systems when estimating the carbon stock will lead to underestimations. Our research can provide a reference when estimating the carbon storage of similar tree species.     

Managing road corridors to limit fire hazard. A simulation approach in southern France

Road corridors are sources of fire ignition and fire spread in French Mediterranean areas, but little is known about the flammability of vegetation and the probability of fire ignition and spread to neighbouring forests. This study simulated fire propagation in road corridors with the help of a cellular automata (CA). We assessed the relative importance of vegetation type, fuel treatment and the spatial patterning of vegetation on the probability of fire spreading to forests. The cellular automaton simulator was implemented with different types of corridors (30 m × 40 m), on the basis of an extensive field survey of vegetation. We used data from laboratory flammability experiments to determine the probability of ignition and propagation for each of the 20 cm × 20 cm cells of the CA. The probability of a fire reaching the neighbouring forest (P_FR) indicated that certain types of road corridors represent a very high risk owing to a combination of highly flammable vegetation and high spatial connectivity. The lowest P_FR values correspond to corridors with a decreasing vegetation flammability gradient in the vicinity of the forest. A several meter-wide embankment with low-flammable and/or managed vegetation can substantially reduce P_FR. These results suggest that firewise landscaping and local vegetation management can reduce fire risk in road corridors. Each corridor type should be subjected to specific vegetation management to account for flammability, growth pattern and lifetime.     

The role of glucosinolates and the jasmonic acid pathway in resistance of Arabidopsis thaliana against molluscan herbivores

Although slugs and snails play important roles in terrestrial ecosystems and cause considerable damage on a variety of crop plants, knowledge about the mechanisms of plant immunity to molluscs is limited. We found slugs to be natural herbivores of Arabidopsis thaliana and therefore investigated possible resistance mechanisms of this species against several molluscan herbivores. Treating wounded leaves with the mucus residue (‘slime trail’) of the Spanish slug Arion lusitanicus increased wound‐induced jasmonate levels, suggesting the presence of defence elicitors in the mucus. Plants deficient in jasmonate biosynthesis and signalling suffered more damage by molluscan herbivores in the laboratory and in the field, demonstrating that JA‐mediated defences protect A. thaliana against slugs and snails. Furthermore, experiments using A. thaliana mutants with altered levels of specific glucosinolate classes revealed the importance of aliphatic glucosinolates in defending leaves and reproductive structures against molluscs. The presence in mollusc faeces of known and novel metabolites arising from glutathione conjugation with glucosinolate hydrolysis products suggests that molluscan herbivores actively detoxify glucosinolates. Higher levels of aliphatic glucosinolates were found in plants during the night compared to the day, which correlated well with the nocturnal activity rhythms of slugs and snails. Our data highlight the function of well‐known antiherbivore defence pathways in resistance against slugs and snails and suggest an important role for the diurnal regulation of defence metabolites against nocturnal molluscan herbivores.     

Arion slugs as nest predators of small passerine species – a review

Arionid slugs have been reported to attack nestlings of some ground‐ or shrub‐nesting passerine birds, mainly in Europe. We review these reported cases and consider their effects. The slugs can cause grave or even fatal injuries to the nestlings. Surprisingly, no brood defence by the parents has been described. The information on the frequency of slug predation in bird populations is scanty, and the scale of the phenomenon is unknown. The expansion of the invasive Arion vulgaris Moquin‐Tandon, 1855 (synonymously A. lusitanicus or A. lusitanicus auct. non Mabille, 1868) in Europe may result in an increase of the negative influence of slugs on the breeding success of some passerines in the near future.     

Density and distribution of seeds in bottom sediments in Zostera marina beds in Ago Bay,central Japan

The density of Zostera marina L. seeds in bottom sediments was examined to study the reproductive patterns of the Z. marina population in Ago Bay, Mie Prefecture, central Japan.Seeds and seed coats were numerous in Tategami, where the annual type of Z. marina grows. In contrast, seeds were scarce in Hamajima, where the perennial type of Z. marina grows. Bottom sediment was sampled with sediment cores at Tategami in November 2004 and March 2005 to examine density and depth distribution of the seeds. Seeds were found as deep as 8 cm, but no deeper. On the other hand, empty seed coats were found as deep as 16 cm in both months. In the upper layers of the sediment to a depth of 8 cm, the average number of seed coats was 7960 ± 2997 m⁻² in November and 16,318 ± 2922 m⁻² in March. Deeper than 8 cm, the number of seed coats gradually decreased owing to decomposition, and none was found below 16 cm. We used the density of reproductive shoots and number of seeds per spadix in Tategami to estimate the fate of seeds and seed coats of the annual type of Z. marina in bottom sediments: out of the 6000 seeds m⁻² produced annually, 72% disappears from the stand and 28% is buried locally. The density and distribution of Z. marina seeds are among the most important factors in maintenance and propagation of the annual population at Tategami.     

The effects of fire on woody plant encroachment are exacerbated by succession of trees of decreased palatability

The ingression of woody plants into the grassy layer of savannas and grasslands has become a global concern. The increase of woody plants has been primarily attributed over grazing, fire and more recently to the increase of atmospheric CO₂. We used long-term observations and analyses to assess changes in woody vegetation in Ithala Game Reserve (IGR), South Africa. Textural analysis of aerial photographs was used to detect changes in woody vegetation, from 1943 to 2007 in Ithala Game Reserve (IGR), South Africa. Daily rainfall data from 1905 to 2009 were used in a time-series analysis to determine if rainfall patterns have changed. The time-series analysis showed that the low magnitude (0–10 mm) rainfall events decreased from 1916 to 2009 and high magnitude rainfall events increased (10–20 and >20 mm). The mean annual rainfall increased from ～700 to ～850 mm from the 1930s to the 2000s. This change in rainfall was a key factor in the increase in woody vegetation from 1943 to 2009. We also used field data from the same reserve collected over 30 years to assess the increases in tree cover. Tree cover and density increased significantly by 32.5% and 657.9 indiv ha^?1 respectively, over 64 years. Before the proclamation of IGR in 1972, increases in woody vegetation from 1943 were non-significant. After the proclamation of IGR, herbivore population numbers and spatial distribution influenced the accumulation of grassy biomass required to fuel fires. In areas with reduced fuel loads, the consequential suppression of fire accelerated the rate of woody plant invasion into savannas. The increase in woody vegetation coincided with a decrease in palatable (e.g. Acacia gerrardii and Acacia davyi) and an increase in unpalatable woody plants. The avoidance of the unpalatable trees (e.g. Euclea and Searsia species) by large mammalian herbivores has allowed these trees to increase in density relatively unhindered.     

The seed bank in a subtropical freshwater marsh: implications for wetland restoration

Seed bank samples were collected from Huli Marsh, a subtropical shallow water mountainous marsh in Hunan Province, South China. Core samples were divided into upper and lower layers (each 5 cm in depth) and allowed to germinate in three water levels (0, 5 and 10 cm) over a 4-month period. A total of 51 species germinated and the mean density was 9211 ± 7188 seedlings m⁻². In the top 5 cm 41 species and 5747 ± 5111 seedlings m⁻² germinated, whereas 40 species and 3464 ± 3363 seedlings m⁻² did so from 5–10 cm. Germinated seedling density was significantly higher in the upper layer, largely due to differences in eight species. With increasing experimental water depth, less seedlings germinated: respectively, 9788 ± 7157 m⁻², 2050 ± 2412 m⁻² and 1978 ± 2616 m⁻², of 44, 21 and 19 species, submerged under 0, 5 or 10 cm. Seven species could emerge only in 0 water level. Vallisneria natans occurred only in 5 cm water, whereas Ottelia alismoides occurred in 10 cm water. In the vegetation survey of the marsh, 25 species were recorded, which was less than half of the species recorded in the seed bank. The top 10 dominants in the standing vegetation, accounting for 89% of vegetation abundance, represented only 10% in the seed bank. Twenty germinated species that also occurred in the standing vegetation accounted for 56% of the total seed bank. Our observed number of species germinating from a Chinese wetland seed bank is within the range observed elsewhere in the northern hemisphere (15–113 species).     

A putative Ariadne-like ubiquitin ligase is required for Dictyostelium discoideum development

The Dictyostelium rbrA gene encodes a putative Ariadne ubiquitin ligase. rbrA⁻ cells form defective slugs that cannot phototax. Prestalk cell numbers are reduced in rbrA⁻ slugs, and these prestalk cells do not localize to the tip of slugs. Chimeric slugs containing wild-type cells could phototax and form fruiting bodies.     

The influence of weather upon the activity of slugs

Summary The activity of slugs was measured by a catch per unit effort sampling system based upon night searching, and was related to the microclimatological conditions in the habitat by regression analysis. The activity of Arion hortensis Fér., Arion subfuscus (Drap.), Arion lusitanicus Mab., Milax budapestensis (Hazay), and of all slugs irrespective of species, was found to be best related to temperature and vapour pressure deficit. The relevance of these results to the application of effective methods for the control of slugs is discussed.     

Effects of climate change,coal mining and grazing on vegetation dynamics in the mountain permafrost regions

The ecological environment in alpine regions is fragile and sensitive to land-use and land-cover (LULC) change and climate warming. However, there are limited studies on the response of LULC and vegetation activity to climate change and human interference in mountainous permafrost regions. Based on in-situ meteorological and multi-source remote sensing data, we performed time trend and partial correlation analyses to investigate the spatial and temporal variation of LULC, landscape pattern, and vegetation growth under the impact of climate change and human activities in the source region of the Datong River from 2000 to 2019. Our results showed that the alpine desert area decreased significantly at a rate of −13.1 km² yr⁻¹ (p < 0.05), while the alpine meadow area increased at a rate of 8.3 km² yr⁻¹ (p < 0.1). Mining and road areas showed a significant increasing trend at a rate of 3.2 km² yr⁻¹ and 1.2 km² yr⁻¹, respectively. The increasing alpine meadow and mining areas were mainly derived from alpine deserts and alpine wetlands, respectively. The number of alpine wetland patches increased significantly along with a significant decrease in the landscape shape index of the rivers. Vegetation growth, as indicated by the enhanced vegetation index (EVI) was positively correlated with temperature but negatively correlated with precipitation and solar radiation in 59.6%, 52.3%, and 56.5% of the vegetated areas, respectively (p < 0.05). Temperature was the dominant climate factor controlling vegetation dynamics, and the recent warming hiatus resulted in a significant increase in EVI for alpine deserts, but no significant changes in EVI for alpine wetlands and alpine meadows. Increasing risk of negative impacts from human activities, including mineral exploration and grazing, on vegetation distribution and growth was observed. This study provides clear evidence of the upward invasion of alpine meadows into alpine desert areas under warm and humid climatic conditions. As climate warming intensifies, alpine meadow expansion may be impeded by extreme precipitation and permafrost thawing.