Low propensity for aerial dispersal in specialist spiders from fragmented landscapes

Aerial dispersal by ballooning is a passive flight, by which wind drag generates an upward lift on a silk thread. It is likely to reflect an aerial lottery, in which the absence of flight direction control is a serious cost for long-distance dispersal in a fragmented landscape. For species occurring in one patchily distributed habitat type, dispersal should evolve in a different way from morphological traits, directly linked to active dispersal. Therefore, we expect that if the risk of landing in an unsuitable habitat is lower than the probability of reaching a suitable habitat, selection should benefit a well-developed ballooning behaviour. We investigated interspecific variation in the ballooning-initiating tiptoe behaviour as it is linked to spider dispersal performance. Our results indeed indicate that ballooning performance is negatively related to habitat specialization in spiders from patchy grey dunes, so habitat specialists are characterized by poorly developed dispersal behaviour. These findings are concordant with recent insights that dispersal is selected as risk spreading in generalists, while it is selected against in specialist species.     

Starvation affects pre-dispersal behaviour of Erigone spiders

Population studies often focus on demographic and genetic consequences of dispersal strategies, generally within an evolutionary framework. Adaptive investment in dispersal is generally assessed from single types of (pre-)dispersal behaviour that are presumed to reliably reflect the dispersal strategy adopted. Various spider families show a striking and quantifiable display, known as tiptoe behaviour that prepares individuals for take-off prior to (passive) aerial dispersal (ballooning). The lack of efficient control mechanisms during ballooning prevents individuals from actively selecting a suitable habitat for landing. Ballooning dispersal is therefore often regarded as a wind lottery preceded by individual-based risk assessment. Our laboratory experiments showed that the duration of tiptoe behaviour can be used as an indicator of silk thread length, which is related to the potential dispersal distance. For two related species, Erigone arctica and E. dentiplapis, tiptoe duration decreased independently of sex after starvation, while more complex reaction norms were observed for tiptoe frequency.

Because these two aspects of dispersal behaviours show different responses towards a simple stress-factor (starvation) in two related spider species, we conclude that the level of plasticity in dispersal investment can be subject to selective forces affecting different dispersal properties in different ways. Our results, hence, plead for a more holistic approach when addressing evolutionary and applied questions related to dispersal.     

Repeatability of dispersal behaviour in a common dwarf spider: evidence for different mechanisms behind short‐ and long‐distance dispersal

Abstract 1. The response of dispersal towards evolution largely depends on its heritability for which upper limits are determined by the trait’s repeatability. 2. In the Linyphiid spider E. atra, we were able to separate long‐ and short‐distance dispersal behaviours (respectively ballooning and rappelling) under laboratory conditions. By performing repeated behavioural trials for females, we show that average dispersal trait values decrease with increasing testing days. By comparing mated and unmated individuals during two periods (before and after mating for the mated group, and the same two periods for the unmated group), we show that mating has no effect on the mean displayed dispersal behaviour or its within‐individual variation. Repeatabilities were high and consistent for ballooning motivation, but not for rappelling. 3. Ballooning motivation can be regarded as highly individual‐specific behaviour, while general pre‐dispersal and rappelling behaviours showed more individual variation. Such difference in repeatability between long‐ and short‐distance dispersal suggests that short‐ and long‐distance dispersal events are triggered by different ecological and evolutionary mechanisms.     

寡居种悦目金蛛若蛛群居机制初步研究

对寡居种悦目金蛛Argiope amoena若蛛群居生活对其结网的影响、温度和种群大小对若蛛存活率的影响、若蛛对限制性空间的利用及其扩散方式进行了观察和研究.结果表明,若蛛从卵袋出蛰后不经历群居扩散即具备结网能力,可结完整网,也具备在扩散前不进食水存活的能力;在变温条件下若蛛存活率要远远高于室温条件下的存活率,而在室温条件下若蛛开始死亡和半数死亡的时间比其聚居期长得多;若蛛主要通过群体空中扩散的方式进行扩散,可以减小能耗与敌害的威胁;不同种群大小的若蛛在限制性空间的分布均从第5d左右开始发生较大变化的,这与在空间不受限的条件下若蛛在第5d开始扩散的结果是一致的.据此我们推断:个体扩散代价最小化(包括御敌和能耗)和维持种群较高的生存率可能是寡居种悦目金蛛若蛛群居生活的主要原因.     

Inbreeding depresses short and long distance dispersal in three congeneric spiders

Dispersal is one of the most important precopulatory inbreeding avoidance mechanisms and subject to landscape related selection pressures. In small populations, inbreeding within and between populations may strongly affect population dynamics if it reduces fitness and gene‐flow. While inbreeding avoidance is generally considered to be a key evolutionary driver of dispersal, potential effects of inbreeding on the dispersal process, are poorly known. Here, I document how inbreeding within a population, so by mating among relatives, affects the survivorship and the dispersal behaviour of three congeneric spider Erigone species (Araneae: Linyphiidae) that differ in habitat preference and regional rarity. The three species were chosen as a model because they allow the assessment of both long and short distance dispersal motivation (respectively ballooning and rappelling) under laboratory conditions. Inbreeding reduced both long and short distance dispersal modes in the three congeneric species. Because survival was depressed after inbreeding, with a tendency of reduced survival loss in the rare and highly stenotopic species, energetic constraints are likely to be the underlying mechanism. Inbreeding consequently depresses silk‐related dispersal in three related spiders. This may induce an inbreeding depression vortex with important consequences for range expansion and metapopulation dynamics of aerially dispersing species from highly fragmented landscapes.     

蜘蛛飞航研究进展

种群扩散能够避免生物类群在同一栖息地中的资源竞争和同类相残, 促进基因流动, 具有重要的进化和生态意义。相比于昆虫的迁飞行为, 某些不具翅的节肢动物也能够借助细丝的作用而实现远距离扩散, 即飞航。蜘蛛为肉食性动物, 是陆地生态系统的重要组成部分, 在生态系统中防控害虫方面作用显著。飞航是蜘蛛的一种重要扩散方式, 在其物种形成、生态位分化及害虫防控等方面具有重要意义, 但有关其飞航扩散的报道较少。本文回顾了蜘蛛飞航的研究历程, 大致可分为三个阶段; 总结了其研究现状, 包括飞航蜘蛛的优势类群及特征、飞航过程中的三个环节(即起飞、空中航行和降落)及影响因子、两种飞航机制及四种相关模型、飞航蜘蛛采集和研究方法。同时基于过往研究, 指出了未来蜘蛛飞航研究的重点领域: 加强飞航蜘蛛类群鉴定工作, 继续深度挖掘飞航数据与环境因子间的关系, 探索蜘蛛飞航的全过程(包括起飞、空中航行和降落), 更加明确蜘蛛飞航作为一种扩散方式的意义。本综述将为蜘蛛飞航的知识普及、资源保护和利用等方面提供重要参考。     

Ballooning propensity of canopy and understorey spiders in a mature temperate hardwood forest

1. Spiders frequently disperse and colonise habitats through ballooning, a passive aerial dispersal process. Ballooning is pre‐eminent in open habitat spider communities and its propensity can be modulated by habitat conditions and availability, and by life‐history traits such as body size, degree of specialisation, and feeding behaviour. 2. Using spiders from the canopy and understorey of a north‐temperate hardwood forest as a model system, our main objectives were to detect if foliage spiders of a mature forest disperse through ballooning, and identify life‐history traits that influence ballooning propensity. 3. Our results demonstrate that foliage spiders living in the canopy and understorey of a mature forest do balloon, and in some cases have very high ballooning propensities similarly to open field spiders. Species level models showed that small body size had a strong positive effect on ballooning for juveniles of species with large‐bodied adults, while individuals of small‐bodied species initiated ballooning regardless of size, habitat or development stage. A generalised linear mixed model indicated that small size web‐building spiders from the Retro Tibial Apophysis (RTA) and Orbicularia clades had the highest propensity for foliage spiders of this north‐temperate hardwood forest. 4. In conclusion, we provide the first demonstration that forest spiders can have high ballooning propensities and that ballooning propensity is negatively affected by body size and positively affected by the prominent use of silk to catch prey. However, spiders originating from the canopy and understorey of a north‐temperate hardwood forest did not differ in their ballooning propensities.     

Big and aerial invaders: dominance of exotic spiders in burned New Zealand tussock grasslands

As post-disturbance community response depends on the characteristics of the ecosystem and the species composition, so does the invasion of exotic species rely on their suitability to the new environment. Here, we test two hypotheses: exotic spider species dominate the community after burning; and two traits are prevalent for their colonisation ability: ballooning and body size, the latter being correlated with their dispersal ability. We established spring burn, summer burn and unburned experimental plots in a New Zealand tussock grassland area and collected annual samples 3 and 4 years before and after the burning, respectively. Exotic spider abundance increased in the two burn treatments, driven by an increase in Linyphiidae. Indicator analysis showed that exotic and native species characterised burned and unburned plots, respectively. Generalised linear mixed-effects models indicated that ballooning had a positive effect on the post-burning establishment (density) of spiders in summer burn plots but not in spring plots. Body size had a positive effect on colonisation and establishment. The ability to balloon may partly explain the dominance of exotic Linyphiidae species. Larger spiders are better at moving into and colonising burned sites probably because of their ability to travel longer distances over land. Native species showed a low resilience to burning, and although confirmation requires longer-term data, our findings suggest that frequent fires could cause long lasting damage to the native spider fauna of tussock grasslands, and we propose limiting the use of fire to essential situations.     

Dandelion Seed Dispersal: The Horizontal Wind Speed Does Not Matter for Long-Distance Dispersal - it is Updraft!

Abstract: Long-distance dispersal of seeds (LDD) surely affects most ecological and evolutionary processes related to plant species. Hence, numerous attempts to quantify LDD have been made and, especially for wind dispersal, several simulation models have been developed. However, the mechanisms promoting LDD by wind still remain ambiguous and the effects of different weather conditions on LDD, although recognized as important, have only rarely been investigated. Here we examine the influence of wind speed and updrafts on dispersal of dandelion ( Taraxacum officinale agg.), a typical wind-dispersed herb of open habitats. We used PAPPUS, a weather-sensitive mechanistic simulation model of wind dispersal, which considers frequency distribution of weather conditions during the period the simulation refers to. A simulation for the 4-month shedding period of dandelion shows that high wind speed does not promote LDD. In contrast, vertical turbulence, especially convective updrafts, are of overwhelming importance. Mainly caused by updrafts, in the simulations more than 0.05 % of dandelion seeds were dispersed beyond 100 m, a distance commonly used to define LDD. We conclude that long-distance dispersal of seeds of herbaceous species with falling velocities < 0.5 - 1.0 ms^-1 is mainly caused by convective updrafts.     

Population genetic evidence for sex‐specific dispersal in an inbred social spider

Dispersal in most group‐living species ensures gene flow among groups, but in cooperative social spiders, juvenile dispersal is suppressed and colonies are highly inbred. It has been suggested that such inbred sociality is advantageous in the short term, but likely to lead to extinction or reduced speciation rates in the long run. In this situation, very low levels of dispersal and gene flow among colonies may have unusually important impacts on fitness and persistence of social spiders. We investigated sex‐specific differences in dispersal and gene flow among colonies, as reflected in the genetic structure within colonies and populations of the African social spider Stegodyphus dumicola Pocock, 1898 (Eresidae). We used DNA fingerprinting and mtDNA sequence data along with spatial mapping of colonies to compare male and female patterns of relatedness within and among colonies at three study sites. Samples were collected during and shortly after the mating season to detect sex‐specific dispersal. Distribution of mtDNA haplotypes was consistent with proliferation of social nests by budding and medium‐ to long‐distance dispersal by ballooning females. Analysis of molecular variance and spatial autocorrelation analyses of AFLPs showed high levels of genetic similarity within colonies, and STRUCTURE analyses revealed that the number of source populations contributing to colonies ranged from one to three. We also showed significant evidence of male dispersal among colonies at one site. These results support the hypothesis that in social spiders, genetic cohesion among populations is maintained by long‐distance dispersal of female colony founders. Genetic diversity within colonies is maintained by colony initiation by multiple dispersing females, and adult male dispersal over short distances. Male dispersal may be particularly important in maintaining gene flow among colonies in local populations.     

Aerial dispersal plasticity under different wind velocities in a salt marsh wolf spider

Dispersal can be regarded as a process operating both betweenand within patches of suitable habitat. For uncontrolled dispersalprocesses, the risk of crossing the borders of the habitat patchand arriving in the unsuitable landscape matrix will increasewith decreasing patch area, in particular when the distancebetween isolated habitat patches is larger than the species'average dispersal capacity. Ballooning dispersal in spiderscan be considered as a passive dispersal process, in which disperseddistances depend on the prevalent wind velocity. We executeda reaction norm analysis to analyze how dispersal propensityof the salt marsh wolf spider Pardosa purbeckensis dependedon population characteristics (patch size) and the environment(wind velocity). Dispersal propensity was affected by the interactionbetween wind velocity and maternal patch size. Ballooning propensitiesdecreased with decreasing salt marsh size. Interestingly, genotypesfrom large salt marshes show higher ballooning propensitiesunder higher wind velocities, whereas those from small habitatpatches show their highest dispersal propensity under low windvelocities. Crossing reaction norms and subsequently stronggenotype x environment interaction variation was observed inall populations but tended to be lower in genotypes from largesalt marshes. It is likely that this pattern results from differencesin wind velocity–related costs of within-habitat dispersalin salt marshes of different sizes.     

Ballooning dispersal using silk: world fauna, phylogenies, genetics and models

Aerial dispersal using silk ('ballooning') has evolved in spiders (Araneae), spider mites (Acari) and in the larvae of moths (Lepidoptera). Since the 17th century, over 500 observations of ballooning behaviours have been published, yet there is an absence of any evolutionary synthesis of these data. In this paper the literature is reviewed, extensively documenting the known world fauna that balloon and the principal behaviours involved. This knowledge is then incorporated into the current evolutionary phylogenies to examine how ballooning might have arisen. Whilst it is possible that ballooning co-evolved with silk and emerged as early as the Devonian (410-355 mya), it is arguably more likely that ballooning evolved in parallel with deciduous trees, herbaceous annuals and grasses in the Cretaceous (135-65 mya). During this period, temporal (e.g. bud burst, chlorophyll thresholds) and spatial (e.g. herbivory, trampling) heterogeneities in habitat structuring predominated and intensified into the Cenozoic (65 mya to the present). It is hypothesized that from the ancestral launch mechanism known as 'suspended ballooning', widely used by individuals in plant canopies, 'tip-toe' and 'rearing' take-off behaviours were strongly selected for as habitats changed. It is contended that ballooning behaviour in all three orders can be described as a mixed Evolutionary Stable Strategy. This comprises individual bet-hedging due to habitat unpredictability, giving an underlying randomness to individual ballooning, with adjustments to the individual ballooning probability being conferred by more predictable habitat changes or colonization strategies. Finally, current methods used to study ballooning, including modelling and genetic research, are illustrated and an indication of future prospects given.     

Levels of gene flow among populations of a wolf spider in a recently fragmented habitat: current versus historical rates

Gene flow among populations is important for countering the deleterious effects of random genetic drift and inbreeding, as well as spreading beneficial mutations. Wind-driven aerial dispersal is known to occur in numerous plants and invertebrates. Its evolution suggests that historically, suitable habitat patches were dense enough to make such undirected dispersal evolutionarily advantageous. Using microsatellite markers we assessed the population genetic structure of seven populations of a wolf spider (Rabidosa rabida) capable of ballooning. Historically, each spider population received a mean of 1.5 migrants per generation from the other six populations. Over the past several generations the number of migrants reaching a population is only 0.2. This statistically significant reduction in gene flow coincides with high levels of habitat fragmentation and suggests that undirected aerial dispersal is ineffective in this fragmented landscape. Further, individuals within populations showed signficantly elevated levels of homozygosity relative to Hardy–Weinberg expectations, suggesting that cursorial dispersal may be very limited and genetic structure within populations exists. Inbreeding coefficients averaged 0.18 over all seven populations with very little variation among populations (s = 0.02). Fitness was lower in smaller populations relative to larger ones. Altered landscapes pose evolutionary dilemmas for many metapopulations and species that depend on undirected movement for dispersal may be particularly vulnerable to habitat fragmentation.     

Environmental and genetic background of tiptoe-initiating behaviour in the dwarfspider Erigone atra

Ballooning is a widespread behavioural trait in invertebrates, which enhances passive aerial dispersal. We investigated the influence of common lineage (family effect) and postnatal environmental conditions on latency to initiate preballooning tiptoe behaviour (ballooning latency) in the dwarfspider Erigone atra (Blackwall, 1841). The ballooning latency of individual spiders was tested in a specially designed test chamber. In a 3-week experiment, in which the spiders were fed only during the first day of each week, ballooning latency had low repeatability at weekly intervals. Ballooning latency declined with increasing food deprivation during the first week but not during the second and the third weeks. At intervals of less than 1 h, however, ballooning latency showed high repeatability. We also investigated whether maternal and postnatal environmental conditions (i.e. during juvenile development) influence phenotypic variation in ballooning latency. We reared offspring of several families under two feeding and two temperature conditions. Environmental conditions explained more variation in ballooning latency than family. Ballooning latency was lower in spiders reared at 20°C than in those reared at 15°C. In addition, spiderlings fed four prey per 3 days were faster ballooners than those fed only four prey per week. An interaction between factors was present, indicating the existence of different reaction norms between the two environmental conditions. The expression of ballooning latency behaviour thus strongly depends on current nutrition, feeding history and the feeding and temperature conditions during juvenile development. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

SPATIALLY CORRELATED EXTINCTIONS SELECT FOR LESS EMIGRATION BUT LARGER DISPERSAL DISTANCES IN THE SPIDER MITE TETRANYCHUS URTICAE

Dispersal is a central process to almost all species on earth, as it connects spatially structured populations and thereby increases population persistence. Dispersal is subject to (rapid) evolution and local patch extinctions are an important selective force in this context. In contrast to the randomly distributed local extinctions considered in most theoretical studies, habitat fragmentation or other anthropogenic interventions will lead to spatially correlated extinction patterns. Under such conditions natural selection is thought to lead to more long‐distance dispersal, but this theoretical prediction has not yet been verified empirically. We test this prediction in experimental spatially structured populations of the spider mite Tetranychus urticae and supplement these empirical results with insights from an individual‐based evolutionary model. We demonstrate that the spatial correlation of local extinctions changes the entire distribution of dispersal distances (dispersal kernel) and selects for overall less emigration but more long‐distance dispersal.     

Short-term response of ground-dwelling arthropods to storm-related disturbances is mediated by topography and dispersal

Wind disturbances and consequent salvage logging lead to drastic changes in forest soil conditions, vegetation and microclimate, potentially affecting arthropod communities. In mountain regions, topography is expected to be particularly important to modulate the effect of canopy removal and soil disturbance potentially amplifying the ecological contrast between forest and disturbed areas. Here, we studied the short-term response of ground beetles (Carabidae), spiders (Araneae), and harvestmen (Opiliones) in wind-damaged spruce forests along statistically orthogonal gradients in elevation, slope, and aspect. We addressed three main ecological questions: (i) Does the effect of wind disturbance on diversity depend on topography? (ii) Are there specific taxon-related responses to disturbances?, and (iii) What is the role of dispersal in shaping species assembly dynamics? We generally observed that increasing slope and elevation amplified the differences between undisturbed forest and windfall areas. On the one hand, the diversity of ground beetles and harvestmen seemed to be negatively affected by wind disturbance, causing a loss of specialized forest species with a low rate of colonization of species typical of open habitats. On the other hand, several novel spider species were able to rapidly colonize windfalls and community composition strongly shifted from forest to disturbed areas. Species with long-range dispersal strategies (e.g. flying and ballooning) were those more likely to colonize windfalls. Our findings suggest that disturbance effects on ground-dwelling organisms were modulated by underlying environmental gradients and that short-term response of different taxa was dependent on their dispersal ability.     

Understanding the impact of flooding on trait-displacements and shifts in assemblage structure of predatory arthropods on river banks

1. Species assemblages of naturally disturbed habitats are governed by the prevailing disturbance regime. Consequently, stochastic flood events affect river banks and the inhabiting biota. Predatory arthropods occupy predominantly river banks in relation to specific habitat conditions. Therefore, species sorting and stochastic processes as induced by flooding are supposed to play important roles in structuring riparian arthropod assemblages in relation to their habitat preference and dispersal ability. 2. To ascertain whether assemblages of spiders and carabid beetles from disturbed river banks are structured by stochastic or sorting mechanisms, diversity patterns and assemblage-wide trait-displacements were assessed based on pitfall sampling data. We tested if flooding disturbance within a lowland river reach affects diversity patterns and trait distribution in both groups. 3. Whereas the number of riparian spider species decreased considerably with increased flooding, carabid beetle diversity benefited from intermediate degrees of flooding. Moreover, regression analyses revealed trait-displacements, reflecting sorting mechanisms particularly for spiders. Increased flooding disturbance was associated with assemblage-wide increases of niche breadth, shading and hygrophilic preference and ballooning propensity for spider (sub)families. Trait patterns were comparable for Bembidiini carabids, but were less univocal for Pterostichini species. Body size decreased for lycosid spiders and Bembidiini carabids with increased flooding, but increased in linyphiid spiders and Pterostichini carabids. 4. Our results indicate that mainly riparian species are disfavoured by either too high or too low degrees of disturbance, whereas eurytopic species benefit from increased flooding. Anthropogenic alterations of flooding disturbance constrain the distribution of common hygrophilous species and/or species with high dispersal ability, inducing shifts towards less specialized arthropod assemblages. River banks with divergent degrees of flooding impact should be maintained throughout dynamic lowland river reaches in order to preserve typical riparian arthropod assemblages.     

Density and genetic relatedness increase dispersal distance in a subsocial organism

Although dispersal distance plays a major role in determining whether organisms will reach new habitats, empirical data on the environmental factors that affect dispersal distance are lacking. Population density and kin competition are two factors theorised to increase dispersal distance. Using the two‐spotted spider mite as a model species, we altered these two environmental conditions and measured the mean dispersal distance of individuals, as well as other attributes of the dispersal kernel. We find that both density and relatedness in the release patch increase dispersal distance. Relatedness, but not density, changes the shape of the dispersal kernel towards a more skewed and leptokurtic shape including a longer ‘fat‐tail’. This is the first experimental demonstration that kin competition can shape the whole distribution of dispersal distances in a population, and thus affect the geographical spread of dispersal phenotypes.     

Increased propensity for aerial dispersal in disturbed habitats due to intraspecific variation and species turnover

Animal dispersal depends on multiple factors, such as habitat features and life‐history traits of the species. We studied the propensity for ballooning dispersal in spiders under standardized laboratory conditions. The 1269 tested individuals belonged to 124 species and originated from 16 sites with wide variation in habitat type. Spiders from disturbed habitats ballooned 5.5 times more than spiders from stable habitats. In Meioneta rurestris , for which we had enough data for a single‐species analysis, individuals were most dispersive if they originated from highly disturbed habitats. While the data for the other species were not sufficient for single‐species analyses, a hierarchical model that included the data simultaneously on all species suggested that dispersal propensity generally increases within species with the level of habitat disturbance. Dispersal probability showed a trend to increase with niche width, but the higher commonness of species with wide niches provides an alternative explanation for this pattern. As the prevalence of especially dispersive species was highest in disturbed habitats, variation in dispersal propensity was influenced by both inter‐ and intraspecific factors. We conclude that the positive correlation between niche width and dispersal propensity enables generalist species to utilize highly disturbed habitats, whereas the persistence of specialist species with restricted dispersal ability requires the conservation of stable habitats.