青海湖流域具鳞水柏枝植物水分利用氢同位素示踪研究

具鳞水柏枝(Myricaria squamosa)是我国高寒地区广泛分布的优势河谷灌木, 具有维持河谷湿地系统稳定的功能。然而, 目前国内外有关具鳞水柏枝水分利用来源的定量研究很少。该文运用氢稳定同位素示踪方法, 分析了青海湖流域具鳞水柏枝茎(木质部)水和潜在水源(地下水、河水和土壤水)的氢稳定同位素比率(δD)的季节变化, 发现具鳞水柏枝在不同水文环境下的植物水分利用来源有明显差异。研究结果表明, 生长在河岸边的具鳞水柏枝在6、7月主要利用地下水与河水, 分别占其所利用水分的89%、86%和55%、65%, 8月主要利用0-20 cm土层的土壤水, 9月水源不详。生长在离河岸约100 m处的具鳞水柏枝在6月主要利用地下水与河水(91%、70%), 在7-9月以0-60 cm土层的土壤水为主要水源。这表明生长在河岸边的具鳞水柏枝对地下水和河水的依赖程度较高, 而距离河岸约100 m时对土壤水的利用量较多, 反映出生长在不同生境中的具鳞水柏枝对特定水分条件的特殊适应结果。     

Factors Influencing Bank Geomorphology and Erosion of the Haw River,a High Order River in North Carolina,since European Settlement

The Haw River, a high order river in the southeastern United States, is characterized by severe bank erosion and geomorphic change from historical conditions of clear waters and connected floodplains. In 2014 it was named one of the 10 most threatened rivers in the United States by American Rivers. Like many developed areas, the region has a history of disturbance including extensive upland soil loss from agriculture, dams, and upstream urbanization. The primary objective of this study was to identify the mechanisms controlling channel form and erosion of the Haw River. Field measurements including bank height, bankfull height, bank angle, root depth and density, riparian land cover and slope, surface protection, river width, and bank retreat were collected at 87 sites along 43.5 km of river. A Bank Erosion Hazard Index (BEHI) was calculated for each study site. Mean bank height was 11.8 m, mean width was 84.3 m, and bank retreat for 2005/2007-2011/2013 was 2.3 m. The greatest bank heights, BEHI values, and bank retreat were adjacent to riparian areas with low slope (<2). This is in contrast to previous studies which identify high slope as a risk factor for erosion. Most of the soils in low slope riparian areas were alluvial, suggesting sediment deposition from upland row crop agriculture and/or flooding. Bank retreat was not correlated to bank heights or BEHI values. Historical dams (1.2–3 m height) were not a significant factor. Erosion of the Haw River in the study section of the river (25% of the river length) contributed 205,320 m³ of sediment and 3759 kg of P annually. Concentration of suspended solids in the river increased with discharge. In conclusion, the Haw River is an unstable system, with river bank erosion and geomodification potential influenced by riparian slope and varied flows.     

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.     

Local population success in heterogeneous habitats: reciprocal transplant experiments completed on a desert spider

Reciprocal transplant experiments were completed to test for selection against the mixing of behavioural phenotypes in a desert spider. Most Agelenopsis aperta populations experience low prey abundances and competition for web‐sites that provide shelter from thermal extremes. These conditions favour aggressiveness towards both prey and conspecifics (an ‘arid‐land behavioural phenotype’). The spider also occupies narrow stretches of riparian habitat bordering spring‐fed streams and rivers. Here it is released from competition for prey and foraging sites, but is subject to predation by birds. A less aggressive/more fearful behaviour is selected for in these riparian habitats (a ‘riparian behavioural phenotype’). Previous work with this spider indicates that there is genetic differentiation between arid‐land and riparian populations. However, the degree to which genetic differentiation is achieved may be limited by gene flow. Reciprocal sets of enclosures were established in: (1) a dry evergreen woodland site (arid‐land phenotype) and (2) a neighbouring riparian site (riparian phenotype) in south‐eastern Arizona. Equal numbers of field collected, early instar A. aperta were introduced into native and transplant enclosures in each habitat. After 6 months of site‐imposed selection, survivorship was determined and growth estimates and behavioural trials completed on spiders remaining in the different enclosures. The same behavioural test was subsequently applied to lab‐reared offspring of the spiders surviving the respective selection regimes. Riparian transplants showed both poor survival and retarded growth in the dry woodland habitat when compared with both arid‐land and riparian natives. Arid‐land transplants that survived, however, grew equally well in riparian habitat as did dry woodland and riparian natives. Behavioural assays conducted on test subjects after selection and on their offspring reared in a controlled laboratory environment indicate that phenotypes that were inappropriate to the respective habitats were selected against in the transplant experiments. The frequency distribution of transplant spider behaviour on a continuum from fearful to aggressive was intermediate between that exhibited by respective native riparian and dry woodland spiders. It is concluded that while arid‐land and riparian behavioural ecotypes do exist, directional gene flow of arid‐land phenotypes into riparian habitat limits population subdivision.     

The Role of Abandoned Channels as Refugia for Sustaining Pioneer Riparian Forest Ecosystems

In disturbance-prone ecosystems, organisms often persist in spatial refugia during stressful periods. A clear example is the colonization of abandoned river channels by pioneer riparian trees. Here, we examine the prominence of this establishment pathway for a foundation tree species (Fremont cottonwood, Populus fremontii) within the riparian corridor of a large river, the Sacramento River in central California. We quantified the total proportion of forest that initiated as a result of channel abandonment for a 160-km reach, analyzed concurrent patterns of tree establishment with floodplain accretion and sedimentation history, and developed a conceptual model of biogeomorphic evolution of abandoned channels. Historical air photo analysis indicated that stands associated with abandoned channels comprised more than 50% of the total extant cottonwood forest area. Tree-ring evidence showed that cottonwood stands commonly developed immediately following abandonment, and the recruitment window ranged from 4 to 40 years, but was less than 10 years at most sites. Rates of floodplain rise and fine sediment accumulation were high in young sites and decreased logarithmically over time. Together, these results suggest that abandoned channels are an important refuge for cottonwood recruitment, that the greatest opportunity for colonization occurs within a short period after the cutoff event, and that sedimentation processes influence the duration of the colonization window. On rivers where tree recruitment along the active channel is severely limited by hydrologic regulation and/or land management, abandoned channel refugia may play an even more important role in sustaining an ecologically functional riparian corridor. Preserving bank erosion, active meander corridors and forest regeneration zones created by cutoff events are therefore key conservation measures on shifting rivers.     

Spatial and temporal patterns of species richness in a riparian landscape

Aim To test for control of vascular plant species richness in the riparian corridor by exploring three contrasting (although not mutually exclusive) hypotheses: (1) longitudinal patterns in riparian plant species richness are governed by local, river‐related processes independent of the regional species richness, (2) riparian plant species richness is controlled by dispersal along the river (longitudinal control), and (3) the variation in riparian plant species richness mirrors variation in regional richness (lateral control). Location The riparian zones of the free‐flowing Vindel River and its surrounding river valley, northern Sweden. Methods We used data from three surveys, undertaken at 10‐year intervals, of riparian reaches (200‐m stretches of riverbank) spanning the entire river. In addition, we surveyed species richness of vascular plants in the uplands adjacent to the river in 3.75‐km² large plots along the same regional gradient. We explored the relationship between riparian and upland flora, and various environmental variables. We also evaluated temporal variation in downstream patterns of the riparian flora. Results Our results suggest that local species richness in boreal rivers is mainly a result of local, river‐related processes and dispersal along the corridor. The strongest correlation between species richness and the environment was a negative one between species number and soil pH, but pH varied within a narrow range. We did not find evidence for a correlation between species richness on regional and local scales. We found that the local patterns of species richness for naturally occurring vascular plants were temporally variable, probably in response to large‐scale disturbance caused by extreme floods. Most previous studies have found a unimodal pattern of species richness with peaks in the middle reaches of a river. In contrast, on two of three occasions corresponding to major flooding events, we found that the distribution of species richness of naturally occurring vascular plants resembled that of regional diversity: a monotonic decrease from headwater to coast. We also found high floristic similarity between the riparian corridor and the surrounding landscape. Main conclusions These results suggest that local processes control patterns of riparian species richness, but that species composition is also highly dependent on the regional species pool. We argue that inter‐annual variation in flood disturbance is probably the most important factor producing temporal variability of longitudinal species richness patterns.     

The lateral extent of the subsidy from an upland stream to riparian lycosid spiders

Adult aquatic insects emerging from streams can subsidize riparian food webs, but little is known of the spatial extent of these subsidies. Stable isotope (¹⁵N) enrichment of aquatic insects, principally a species of stonefly (Plecoptera: Leuctridae), emerging from an upland stream was used to trace the subsidy from the stream ecosystem to riparian spiders (Lycosidae). The downstream profile of spider δ¹⁵N correlated closely with that of adult stoneflies, indicating that they were deriving nutrition from aquatic sources. The contribution of adult aquatic insects to spider diets was determined using a two-source mixing model. Adult aquatic insects made up over 40% of spider diets adjacent to the stream, but <1% at 20 m from the stream. Enrichment of riparian spiders declined exponentially with distance from the stream channel. Aquatic-terrestrial subsidies were spatially restricted, but locally important, to riparian lycosid spiders at the study site.     

Sex‐Specific Response of Pardosa milvina (Araneae: Lycosidae) to Experience with a Chemotactile Predation Cue

Predators frequently leave behind chemical information (i.e., semiochemicals such as pheromones or kairomones) that can be detected by their prey and used to avoid areas where predators are likely present. Prey that have interacted indirectly with predators via chemical information thus may gain insight into their risk of being consumed that naïve individuals lack. Pardosa milvina (Araneae: Lycosidae) is a chemosensitive wolf spider that shows adaptive responses to chemotactile cues deposited by the larger wolf spider Tigrosa helluo. We raised offspring from P. milvina to examine the effect of experience with a predation cue on activity, foraging, and antipredator behavior. Spiders differed in activity and foraging behavior across ontogeny and between sexes, but there was no effect of experience with a predation cue. However, a sex‐specific effect of experience was found in antipredator behavior. Male spiders, but not females, used experience with a predator cue to increase their survival in the presence of a live predator. Specifically, naïve males were attacked sooner than experienced males, indicating that prior exposure to predator cues can modify Pardosa antipredator behavior. Intersexual differences in how spiders respond to experience with a predation cue likely reflect the risk of predation faced by males and females in nature.     

Burrow-blocking behaviour in Geolycosa wrightii (Araneae: Lycosidae)

Patterns of behaviour associated with preventing wind-blown sand from entering the burrows of the wolf spider Geolycosa wrightii were investigated in a laboratory study. The method of closing the burrow depended upon the rate of sand entry. The spider employed a web block at low rates and used its body to block at high rates.     

Low river flow alters the biomass and population structure of a riparian predatory invertebrate

1. Low flows in rivers are predicted to increase in extent and severity in many areas in the future, yet the consequent impacts of river drying on terrestrial communities via (i) changes to riparian microclimatic conditions and (ii) the identity and abundance of emerging aquatic insects available to riparian predators have not been quantified. 2. We investigated the influence of low river flow on a riparian fishing spider, Dolomedes aquaticus, in five New Zealand rivers containing permanently flowing and drying reaches and, in one river, along a longitudinal drying gradient. 3. The biomass of aquatic insects, potential prey for D. aquaticus, declined with low river flows while the abundance of potential terrestrial prey remained similar at all sites. In the replicate rivers, and along the longitudinal drying gradient, spider biomass was lower, and size classes were skewed towards more small and fewer large spiders, in drying sites. A desiccation experiment in the laboratory indicated high sensitivity of the spiders, with prey presence increasing spider survival. 4. Differences in the spatial distribution, biomass and population size structure of spiders were observed along the longitudinal drying gradient and disappeared within 16 days of the water returning to all sites. 5. In total, low river flow affected the biomass of D. aquaticus, as well as their size class structure and spatial distribution. This indicates that low river flows have the potential to affect adjacent terrestrial ecosystems.     

Comparisons in water relations of plants between newly formed riparian and non-riparian habitats along the bank of Three Gorges Reservoir,China

The water table in the Three Gorges Reservoir (TGR) rose significantly since the construction of the Three Gorges Dam across the Yangtze River. Little is known about how such a change in local hydrological condition will affect the ecophysiology of plants along the bank of the world’s largest reservoir. In this study, water relations of the dominant plants were investigated over an entire year period by comparing stable isotope compositions of xylem water, leaf water potentials and foliar carbon isotope ratios at a newly-formed riparian site near the river bank and two non-riparian sites at higher elevations. The isotopic compositions of xylem water indicated that the plants in the newly-formed riparian zone acquired water mainly from the soil previously infiltrated by local rain rather than the Yangtze River water. Predawn and midday water potentials of the riparian trees were also similar to those at the non-riparian sites. Leaf δ¹³C values of plants across the three sites also showed no difference, even though there was a significant difference among species. Our results indicate that plants at the newly formed transient riparian area along the bank of TGR do not depend on the Yangtze River water for their water sources and that rising water levels in this reservoir may not lead to immediate changes in water relations of the plants along the bank. Communicated by K. Winter.     

How within-group behavioural variation and task efficiency enhance fitness in a social group

How task specialization, individual task performance and within-group behavioural variation affects fitness is a longstanding and unresolved problem in our understanding of animal societies. In the temperate social spider, Anelosimus studiosus, colony members exhibit a behavioural polymorphism; females either exhibit an aggressive 'asocial' or docile 'social' phenotype. We assessed individual prey-capture success for both phenotypes, and the role of phenotypic composition on group-level prey-capture success for three prey size classes. We then estimated the effect of group phenotypic composition on fitness in a common garden, as inferred from individual egg-case masses. On average, asocial females were more successful than social females at capturing large prey, and colony-level prey-capture success was positively associated with the frequency of the asocial phenotype. Asocial colony members were also more likely to engage in prey-capture behaviour in group-foraging situations. Interestingly, our fitness estimates indicate females of both phenotypes experience increased fitness when occupying colonies containing unlike individuals. These results imply a reciprocal fitness benefit of within-colony behavioural variation, and perhaps division of labour in a spider society.     

Changes in wolf spider (Araneae) assemblages across woodland–pasture boundaries in the central wheat‐belt of New South Wales,Australia

The abundance of wolf spiders (Lycosidae) was measured across woodland–pasture boundaries in the wheat‐belt of New South Wales, Australia, to determine the nature and magnitude of any edge effect. Spiders were collected by spotlighting along sample plots in woodlands located at distances of 5, 20, 35 and 200 m from the edge, and along sample plots in paddocks located at distances of 5 and 20 m from the edge. The wolf spider assemblage changed significantly across the edge, but the difference could be accounted for only by a change between the woodland and the paddock and not by any changes within the woodland at different distances from the edge. Ground cover (wolf spider microhabitat) changed significantly between the paddock and the woodland, but there were no consistent differences in microhabitat with distance from edge within either paddocks or woodlands. There was a significant correlation between an ordination of sites based on spider species abundance and an ordination based on microhabitat variables, suggesting that the wolf spider assemblage was responding to differences in microhabitat. Fine‐scale selection of microhabitat by most wolf spider species was non‐random, with most species preferring locations with grass cover, rather than more open locations. The present study indicates that wolf spiders are mostly unaffected by edge conditions at the woodland–paddock boundary. Accordingly, small and/or linear remnants with high edge‐to‐area ratios may constitute suitable faunal habitat for wolf spiders and perhaps other terrestrial arthropod species, despite the fact that this configuration is unsuitable for many vertebrate species.     

Long-term nitrate removal in a stream riparian zone

The long-term capacity of riparian zones in regulating groundwater nitrate fluxes is not well understood. This study analyses patterns of nitrate removal for the period 1994–2012 at two sites in a river floodplain that have received high groundwater nitrate loading from a large upland aquifer for over 32 years. During the study, mean NO₃ ^?–N concentrations entering the riparian zone varied between 20–30 and 30–42 mg/L at the upstream and downstream sites respectively, but did not show any clear inter-annual trend. A permeable sand layer in the riparian zone is underlain by a regional aquitard at a depth of 5–6 m and 4 m at the upstream site and downstream site respectively. Denitrification resulted in a decline in nitrate concentrations as lateral groundwater flow in the sand layer interacted with buried peat and channel bar deposits that range up to 3 m in depth at both riparian sites. This interaction was greater at the downstream site where the organic deposits extend down to <1 m from the aquitard in some locations. At the upstream site nitrate removal efficiency in the sand layer, at depths of 3–4 m ~20 m from the river bank, declined from 68 % in 1996–1998 to 42 % in 2009–2012. A smaller decline from 92 to 82 % occurred in the sand layer 10 m from the river bank during the study. In contrast, no clear pattern of change was evident at the downstream site where a nitrate removal efficiency of 98–100 % occurred at the river bank in most years between 1994 and 2012. These data suggest that the long-term nitrate removal performance of some riparian zones may decline if carbon availability for denitrification becomes limited as a result of variations in the quantity, quality and location of subsurface organic deposits that interact with deeper groundwater flowpaths.     

Spider diversity responds strongly to edge effects but weakly to vegetation structure in riparian forests of Southern Brazil

Riparian forests bordering open terrestrial environments may have three microhabitats differing in structure and conditions: a grassland/pasture-forest edge (GE), a forest interior (FI) and a river–forest edge. The influence of such edge effects and vegetation characteristics on spider diversity of riparian forests was evaluated in Southern Brazil. Four different rivers were sampled on the tree–shrub strata with a beating tray, twice per season for 2 years. There were six transects per river, two per microhabitat. We compared spider abundance, species richness and composition. Vegetation variables sampled were vertical structure and (horizontal) density, canopy height and cover. Overall 42,057 spiders were sampled, 28 spider families and 440 species. The FI had higher spider abundance than the edges. Average species richness differed among rivers. Microhabitats did not differ in average richness, although overall richness (from sample-based rarefaction) was higher for GE than FI. High abundances in FI may result from lowered stress due to abiotic conditions, while higher GE richness may result from a faunal superposition between forest species and those from the grassland/pasture. Only canopy cover returns a positive relationship with spider diversity (richness and adult abundance). This might result from more spider species preferring to build webs or hunt under low-light environments. Rivers had spider faunas differing in composition but among microhabitats species composition was the same. Vegetation structure has been hypothesized to affect spiders, but this impact might be best seen in specific subgroups or guilds within spiders, not in the whole assemblage.     

Discrimination of Variation in a Male Signaling Trait Affects Detection Time in Visual Predators

In many species, expression of elaborate male characteristics likely represents a balance between sexual selection and natural selection via predation, as male traits selected to elicit rapid detection or response on the part of females also increase detection by predators. Predation costs are frequently inferred, but the underlying mechanisms associated with specific traits have rarely been directly explored. Males of the wolf spider Schizocosa ocreata (Araneae: Lycosidae) exhibit a sexually selected signaling trait (dark tufts of bristles) on their forelegs, and are sympatric with a number of visually hunting generalist predators, including cannibalistic conspecifics, that may impact spider survival. Here we use latency of orientation response of the American toad, Bufo americanus (Anura: Bufonidae), to video, 'virtual' courting S. ocreata male stimuli as an index of predator detection, and latency of orientation response of female S. ocreata to the same stimuli as an index of conspecific detection. When compared with stimuli representing the population average, elimination of the signal trait had no significant effect on predator detection but did increase latency to orient in conspecifics. Increasing the size of the signal trait had no effect on conspecific detection but did significantly reduce latency to orient for predatory toads. Results clearly indicate that for a courting male spider of a given size and vigor level, variation in the expression of a secondary sexual characteristic alone can incur differential direct costs and benefits by influencing latency of orientation to visual signals by predators and conspecifics.     

Soil seed bank dynamics in response to an extreme flood event in a riparian habitat

A significantly increased water regime can lead to inundation of rivers, creeks and surrounding floodplains- and thus impact on the temporal dynamics of both the extant vegetation and the dormant, but viable soil-seed bank of riparian corridors. The study documented changes in the soil seed-bank along riparian corridors before and after a major flood event in January 2011 in southeast Queensland, Australia. The study site was a major river (the Mooleyember creek) near Roma, Central Queensland impacted by the extreme flood event and where baseline ecological data on riparian seed-bank populations have previously been collected in 2007, 2008 and 2009. After the major flood event, we collected further soil samples from the same locations in spring/summer (November–December 2011) and in early autumn (March 2012). Thereafter, the soils were exposed to adequate warmth and moisture under glasshouse conditions, and emerged seedlings identified taxonomically. Flooding increased seed-bank abundance but decreased its species richness and diversity. However, flood impact was less than that of yearly effect but greater than that of seasonal variation. Seeds of trees and shrubs were few in the soil, and were negatively affected by the flood; those of herbaceous and graminoids were numerous and proliferate after the flood. Seed-banks of weedy and/or exotic species were no more affected by the flood than those of native and/or non-invasive species. Overall, the studied riparian zone showed evidence of a quick recovery of its seed-bank over time, and can be considered to be resilient to an extreme flood event.     

Body condition helps to explain metabolic rate variation in wolf spiders

1. Metabolism is the fundamental process that powers life. Understanding what drives metabolism is therefore critical to our understanding of the ecology and behaviour of organisms in nature. 2. Metabolic rate generally scales with body size according to a power law. However, considerable unexplained variation in metabolic rate remains after accounting for body mass with scaling functions. 3. We measured resting metabolic rates (oxygen consumption) of 227 field‐caught wolf spiders. Then, we tested for effects of body mass, species, and body condition on metabolic rate. 4. Metabolic rate scales with body mass to the 0.85 power in these wolf spiders, and there are metabolic rate differences between species. After accounting for these factors, residual variation in metabolic rate is related to spider body condition (abdomen:cephalothorax ratio). Spiders with better body condition consume more oxygen. 5. These results indicate that recent foraging history is an important determinant of metabolic rate, suggesting that although body mass and taxonomic identity are important, other factors can provide helpful insights into metabolic rate variation in ecological communities.     

Managing riparian zones for river health improvement: an integrated approach

Riparian zones are among the most valuable ecosystems on the earth. They act as the ecological engineers that improve river health through delivering a range of ecosystem functions. Stream bank stabilization, pollutant and sediment buffering, temperature regulation, provision of energy to river food webs and communities, groundwater recharge and provision of ecological corridors and habitat for wildlife, are among major ecosystem functions of riparian zones that play a great role in river health. Besides these ecosystem functions, riparian zones also provide various ecosystem goods and services for human well-being. But in the current scenario, riparian zones are under severe threat due to agricultural activities, urbanization, river flow alteration, overexploitation, climate change, pollution, and biological invasion. In the present and probable future scenarios of declining river health and global environmental changes, there is a pressing need of an integrated approach for managing riparian zones. This review article aims to advocate an integrated approach for riparian zone management based on various components such as riparian condition assessment, policy framework, stakeholder’s participation, management practices, legislation, and awareness. Authors also discussed riparian zones in context of their concepts, features, functions, and threats.

     

Honesty of agonistic signalling and effects of size and motivation asymmetry in contests

Game theoretical models predict that the main function of fighting behaviour is to assess the relative fighting ability of opponents. The sequential assessment game has often been used to investigate contests, while honest signalling theory has received much less attention. With the wolf spider Hygrolycosa rubrofasciata we investigated whether male agonistic signalling can reveal honest information about fighting ability, and how size and motivation asymmetries affect male fighting behaviour. We also determined whether male–male competition affects the courtship behaviour of the males. We found that agonistic drumming activity is an honest indicator of male fighting ability, and that relative size asymmetry and motivation to fight both contribute to the fighting ability. We also found that male–male competition decreases the courtship drumming rate of subdominant males, suggesting that male–male competition limits the opportunities for female choice. We conclude that sequential assessment and honest signalling may both be used in settling contests, and that they may be used simultaneously. Received: 10 December 1998 / Received in revised form: 23 June 1999 / Accepted: 5 July 1999