Staphylinidae and Carabidae overwintering in wheat and sown wildflower areas of different age

Species richness and abundance of staphylinid and carabid beetles overwintering in winter wheat fields and 1- to 3-year-old wildflower areas were investigated during 2000/2001 on 16 study sites in Switzerland. Abundance and species richness of overwintering staphylinids significantly increased with successional age of the wildflower areas and were always higher in older wildflower areas than in winter wheat. A similar but less distinct pattern was observed for the abundance and species richness of carabid beetles. The influence of habitat parameters (vegetation cover, fine sand content, organic matter, pH, soil pore volume, surrounding landscape structure, habitat area) on the staphylinid and carabid assemblages based on the number of individuals per species and site was analysed using canonical correspondence analysis. Vegetation cover was the most significant parameter significantly characterizing both staphylinid and carabid assemblages. The amount of vegetation cover explained 15.7% of the variance, fine sand content accounted for 13.3% and surrounding landscape structure for 10.9% of the variance in the staphylinid assemblage. In the carabid assemblage, vegetation cover was the only significant factor, explaining 24.7% of the variance. This study showed for the first time that the significance of wildflower areas as a reservoir for hibernation for generalist predatory beetles increases with progressing successional age.     

Carabid beetle condition,reproduction and density in winter oilseed rape affected by field and landscape parameters

The carabid beetles Amara similata and Poecilus cupreus are abundant in Central European winter oilseed rape (OSR) fields and potential antagonists of OSR pests. Therefore, they were investigated in 29 OSR fields relative to the influence of field and landscape parameters on their nutritional condition, reproductive potential and activity density. Nutritional condition was measured by a condition factor (CF). Fecundity of female beetles was expressed by the number of ripe oocytes in the ovaries. Activity density comprised the number of individuals caught with pitfall traps during their peak of reproductive period. Soil productivity and pest abundance (as a proxy of local prey availability) were considered as field parameters. Landscape parameters comprised the distance to the next fallow and the percentage of crop area around each study field. In eight of ten cases, pest abundance proved to be the most important factor explaining carabid characteristics, indicating that P. cupreus and A. similata consume insect pests in OSR fields. Pest abundance influenced the CF of both species and sexes positively. Oocyte numbers of A. similata were negatively related to the distance to the next fallow. Oocyte numbers of P. cupreus tended to increase with increasing pest abundance. Activity density of female A. similata was negatively influenced by soil productivity. Activity density of male A. similata and of female and male P. cupreus was negatively influenced by pest abundance. When analysing exclusively the influence of severe OSR pests (Meligethes aeneus, Ceutorhynchus napi and Dasineura brassicae), abundance of larval M. aeneus and C. napi was most important in explaining the CF and activity density of male A. similata, and male and female P. cupreus. M. aeneus and C. napi could be an essential prey for the carabids studied which may reduce the pests, thereby contributing to the protection of OSR from pest herbivory.     

农田及生态补偿区甲虫群落结构的差异

为了调查冬麦田和不同年龄生态补偿区甲虫群落结构和物种多样性,从2003年7到10月和2004年5到6月在瑞士农业区周围不同景观中进行了实地调查,并采用对应及典范对应分析方法,对冬麦田和1~4a不同年龄阶段的野花地中的甲虫群落结构及物种多样性进行了比较系统的研究。结果,共捕甲虫6009只,分别属于150种,25科,其中,麦田共计38种445个体,1~4a不同年龄的野花地中分别计46种,2760个体,81种,1325个体,73种,668个体,75种,811个体。根据食性,植食性种类有98种(65.33%),5189个体(86.35%)、肉食性种类有40种(26.67%),240个体(3.99%)、食菌性种类有10种(6.67%),578个体(9.62%)、食腐性种类有2种(1.33%),2个体(0.03%)。冬麦田中栖息的甲虫种类及其物种丰富度和多度与1~4a不同年龄阶段的野花地有明显差异。这主要由于冬麦田中植物种类多样性低和植被结构简单而导致的。在不同演替阶段野花地中甲虫群落物种的丰富度一般在1~3a型野花地之间有差异。1年型野花地具有比较典型的甲虫群落结构,不同演替阶段的野花地对甲虫群落的定居,扩散及其演替过程中具有非常重要的作用。对甲虫提供丰富的食物资源和栖息,繁殖场所。因此,人为在农业区建立生态补偿区,有利于提高无脊椎动物和昆虫多样性,对物种保护方面具有重要意义。     

A multi-site experiment to test biocontrol effects of wildflower strips in different French climate zones

Sowing of wildflower strips has been integrated in agri-environment schemes of several European countries. Their beneficial effects on natural enemies of pest insects are well documented but (1) the desired spill-over into crop fields has not always been demonstrated, and (2) the need to adapt sown mixtures to regional climatic differences has been rarely addressed.We set up a multi-site experiment in different French climatic regions to compare effects of a wildflower strip with a grass mixture and spontaneous vegetation. The design included five regions, three to five fields per region and the three strip treatments being repeated in each field. We tested strip treatment effects on vegetation (plant species richness, plant and flower cover) and on natural enemies (hoverflies, ladybirds, aphid predation). In a subset, we further analysed the spill-over into winter wheat fields including natural enemies and pest insects (cereal aphids, leaf beetles).The wildflower strip mixture developed well in all regions and increased plant species richness and flower cover compared with grass strips and spontaneous vegetation. We found a corresponding higher hoverfly abundance and aphid predation in wildflower strips that were consistent in all regions, whereas ladybird abundance was not affected. A significantly higher hoverfly abundance, aphid predation and aphid parasitism in wheat fields close to wildflower strips indicated a spill-over. No corresponding margin treatment effects were observed for aphid and leaf beetle abundance in the field. A multivariate analysis comparing the influence of climate and vegetation parameters showed that floral cover better explained variation in natural enemy abundance and predation than climate. Our results demonstrated that similar mixtures of native plants can be used over large climatic gradients to improve biocontrol. Further research is needed to improve spill-over into crop fields and to obtain consistently strong effects in different climate zones.     

Influence of nutrition on male development and reproduction in Tribolium castaneum

Tribolium beetles have evolved over several thousand years to colonize and exploit various food products that vary widely in their nutritional quality. Here Tribolium castaneum was used as a model organism to explore the effect of nutritional quality on male development and reproduction. The results showed, when tested across different qualities of nutritional diets, Tribolium males developed faster and their body size was larger on a high-quality diet, and there were significant correlations between male developmental traits. However, Tribolium males fed different nutritional diets did not show significant variation in olfactory attractiveness, mating rate, insemination rate, sperm defense (P1), sperm offense (P2), and reproductive success within a population context (RSPC). Moreover, there was no significant correlation of male reproductive performances except RSPC with developmental traits, and except for P2 and RSPC, no significant correlations between male reproductive performances. Therefore, although male developmental performance was significantly influenced by diet quality, reproductive performance was not. We discussed these findings and their sexual selection implications in light of its habitat.     

Factors Influencing Bug Diversity (Insecta: Heteroptera) in Semi-Natural Habitats

We investigated the abundance and species richness of heteropteran bugs and explored environmental factors which influence bug diversity in three types of semi-natural habitats (wildflower areas, extensively used meadows, extensively grazed pastures). To cover this topic, it is essential to know how much the relatively young wildflower areas contribute to biodiversity compared with well-established extensive meadows and pastures. Total bug species richness and phytophagous bug species richness were significantly higher in wildflower areas and meadows than in pastures. In wildflower areas, we found the highest number of zoophagous bug species and species overwintering in the egg-stage. Species overwintering as adults were most abundant in meadows. Total number of bug species as well as species richness in either trophic groups and overwintering strategies were significantly positively correlated with vegetation structure. Except for overwintering strategies, the same was true to bug abundance. The bug community based on the number of individuals per species was significantly explained by flower abundance and vegetation structure, accounting for 18.4 and 16.8% of the variance, respectively. Our results indicate that vegetation structure and flower abundance are key factors for bug species richness, abundance and bug species composition. Since wildflower areas and meadows clearly increased bug species richness and contained several specialised bug species that did not occur in pastures, we recommend the promotion of wildflower areas and extensively used meadows in order to restore both high heteropteran diversity and overall insect biodiversity in agricultural landscapes.     

Wildflower areas within revitalized agricultural matrices boost small mammal populations but not breeding Barn Owls

Agro-ecosystems have recently experienced dramatic losses of biodiversity due to more intensive production methods. In order to increase species diversity, agri-environment schemes provide subsidies to farmers who devote a fraction of their land to ecological compensation areas (ECAs). Several studies have shown that invertebrate biodiversity is actually higher in ECAs than in nearby intensively cultivated farmland. It remains poorly understood, however, to what extent ECAs also favour vertebrates, such as small mammals and their predators, which would contribute to restoring functional food chains within revitalised agricultural matrices. We studied small mammal populations among eight habitat types—including wildflower areas, a specific ECA in Switzerland—and habitat selection (radiotracking) by the Barn Owl Tyto alba, one of their principal predators. Our prediction was that habitats with higher abundances of small mammals would be more visited by foraging Barn Owls during the period of chicks’ provisioning. Small mammal abundance tended to be higher in wildflower areas than in any other habitat type. Barn Owls, however, preferred to forage in cereal fields and grassland. They avoided all types of crops other than cereals, as well as wildflower areas, which suggests that they do not select their hunting habitat primarily with respect to prey density. Instead of prey abundance, prey accessibility may play a more crucial role: wildflower areas have a dense vegetation cover, which may impede access to prey for foraging owls. The exploitation of wildflower areas by the owls might be enhanced by creating open foraging corridors within or around wildflower areas. Wildflower areas managed in that way might contribute to restore functional links in food webs within agro-ecosystems.     

Residency Time as an Indicator of Reproductive Restraint in Male Burying Beetles

The cost of reproduction theory posits that there are trade-offs between current and future reproduction because resources that are allocated to current offspring cannot be used for future reproductive opportunities. Two adaptive reproductive strategies have been hypothesized to offset the costs of reproduction and maximize lifetime fitness. The terminal investment hypothesis predicts that as individuals age they will allocate more resources to current reproduction as a response to decreasing residual reproductive value. The reproductive restraint hypotheses predicts that as individuals age they will allocate fewer resources to current reproduction to increase the chance of surviving for an additional reproductive opportunity. In this study, we test for adaptive responses to advancing age in male burying beetles, Nicrophorus orbicollis. Burying beetles use facultative biparental care, but the male typically abandons the brood before the female. Previous work in male burying beetles has suggested several factors to explain variation in male residency time, but no study has observed male behavior throughout their entire reproductive lifetimes to determine whether males change residency time in an adaptive way with age. We compared residency time of males that reproduced biparentally, uniparentally, and on different-sized carcasses to determine if they used an adaptive reproductive strategy. Males did not increase residency time as they aged when reproducing biparentally, but decreased residency time with age when reproducing uniparentally. A decrease in parental care with age is consistent with a reproductive restraint strategy. When female age increased over time, males did not increase their residency time to compensate for deteriorating female condition. To our knowledge, this is the first test of adaptive reproductive allocation strategies in male burying beetles.     

Partitioning of arthropod species diversity in temperate meadows,wildflower areas and pastures

To counteract the rapidly ongoing species decline in modern agroecosystems, most European countries have developed and adopted agri-environment schemes. The efficiency of these schemes, however, ranges from successful to nearly ineffective. To estimate the efficiency of the Swiss agri-environment scheme in particular, we investigated the contribution of extensively managed pastures and meadows and wildflower areas to beetle, heteropteran bug and spider diversity. Extensively managed meadows were shown to contribute most to landscape-scale richness (highest species richness for beetles and bugs), followed by wildflower areas (bugs). Meadows harboured approximately twice as many beetle species unique to this habitat type than wildflower areas and pastures. Bug species unique to a specific habitat type were similar between meadows and wildflower areas, but much lower (approximately 30%) in pastures. To increase beetle species richness, extensively managed meadows should be represented more often in the landscape because meadows harboured the highest number of species and most uniquely found species. To increase species richness of true bugs, both meadows and wildflower areas should be represented more often in the landscape because they harboured more species and more uniquely found species compared to pastures. Across arthropod taxa and habitat types, β diversity was more important for γ diversity than α diversity, indicating that distinctiveness, rather than within-site diversity, was key in determining the contribution of sites to γ (landscape) diversity. This substantiates the argument that increasing the number of sites is efficient to increase landscape species richness. The turnover component of β diversity was far more important than the nestedness component for all arthropod taxa and habitat types and accounted for 79.0% to 95.4% of total β diversity. In conclusion, extensively managed meadows and wildflower areas should be particularly promoted by the Swiss agri-environment scheme, as they contributed more to landscape species richness and species uniqueness than pastures.     

Establishment and management of wildflower areas for insect pollinators in commercial orchards

Sown wildflower areas are increasingly recommended as an agri-environmental intervention measure, but evidence for their success is limited to particular insect groups or hampered by the challenges of establishing seed mixes and maintaining flower abundance over time. We conducted a replicated experiment to establish wildflower areas to support insect pollinators in apple orchards. Over three years, and across 23 commercial UK orchards with and without sown wildflowers, we conducted 828 transect surveys across various non-crop habitats. We found that the abundance of flower-visiting solitary bees, bumblebees, honeybees, and beetles was increased in sown wildflower areas, compared with existing non-crop habitats in control orchards, from the second year following floral establishment. Abundance of hoverflies and other non-syrphid flies was increased in wildflower areas from the first year. Beyond the effect of wildflower areas, solitary bee abundance was also positively related to levels of floral cover in other local habitats within orchards, but neither local nor wider landscape-scale context affected abundance of other studied insect taxa within study orchards. There was a change in plant community composition on the sown wildflower areas between years, and in patterns of flowering within and between years, showing a succession from unsown weedy species towards a dominance of sown species over time. We discuss how the successful establishment of sown wildflower areas and delivery of benefits for different insect taxa relies on appropriate and reactive management practices as a key component of any such agri-environment scheme.     

Type of intruder and reproductive phase influence male territorial defence in wild-caught Siamese fighting fish

This study investigated how parental care increases with offspring age by examining the level of male aggressiveness toward potential intruders during egg guarding in a natural population of Siamese fighting fish (Betta splendens Regan). The degree of aggressiveness was measured at two reproductive phases in response to three types of intruders: male, female and females that have laid eggs. The nest-holding males became more aggressive after their eggs hatched than after the eggs were laid. Nest-holding males displayed gill cover erection, biting, tail beating and attacking at the highest rate towards male intruders, intermediate towards female intruders and the least aggressive towards females, which have laid eggs.     

Water availability drives habitat quality for the butterfly <Emphasis Type="Italic">Plebejus argus</Emphasis> in a Mediterranean sand dune landscape

For terrestrial species, habitat is frequently defined as physical patches of a certain land cover type in a matrix of non-habitat. However, the appropriate definition of habitat quality requires more detailed information on the resources and conditions that are necessary for the maintenance of the species (‘resource-based habitat approach’). In arthropods, their life cycle comprises several clearly differentiated stages that may depend on different resources and conditions for successful development; in turn, males and females may have different habitat requirements due to their different behaviours. Here we examine the habitat quality for males, females and eggs of the butterfly Plebejus argus in a continuous sand dune landscape in southern Spain. For both sexes, abundance was related to host plant density (negatively) and distance to heathland vegetation (a surrogate for higher water availability; negatively for males and hump-shaped for females); and, for males only, positively with flower presence. Egg abundance was positively related to mutualistic ant frequency and flower presence, and negatively to distance to heathland. Hence, female and male adults and eggs of P. argus partly differed in their dependence on resources and conditions, although distance to wetter areas of heathland was the common variable for all models. This suggests that soil humidity is the key variable for P. argus habitat quality, probably through both direct effects on individuals and indirect effects on interacting species (mutualistic ants and nectar sources). As a result, we found a high spatial congruence in habitat quality between males and females, and a moderate congruence between adults and eggs.     

Habitat selection and polygyny in breeding Corn Buntings Miliaria calandra

Unlike many other polygynous passerine species, female Corn Buntings Miliaria calandra apparently do not suffer costs by pairing polygynously, yet it is unclear whether this is because polygynous males hold the highest quality territories or because pairing with polygynous males is unimportant in determining female reproductive success. Male Corn Buntings on North Uist, Scotland, consistently defended territories which contained nesting habitat, and females often foraged outside male territories when provisioning nestlings. Females showed strong preferences for nesting in uncultivated land, and 80% of nests were under Hogweed Heracleum sphondylium, possibly because this provided cover against predation and the weather. When provisioning nestlings, females showed strong preferences for foraging in cereal crops, probably because this habitat provided better food resources and/or better cover from predators. Males were unpaired or paired with one to three females per breeding season, but variation in territory size or vegetation composition did not explain differences in the number of females paired with individual males. We suggest that when females neither gain benefits nor suffer costs by breeding polygynously, and males do not differ greatly in the areas of habitat selected, polygyny can arise through random female settlement within the nesting habitat.     

Myrmecofauna of lucidophyllous forests in different developmental stages in south-western Japan

Lucidophyllous forest is the climax vegetation of the lowland and foothill areas of south-western Japan. We investigated the myrmecofauna of lucidophyllous forests in different developmental stages in relation to the intensity of disturbance. The less-disturbed lucidophyllous forests contained a greater variety of myrmecofauna. This pattern was explained by the condition of the habitat. Richness in epigeal and/or hypogeal ant species is related to habitat structure, especially vegetation structure expressed as summed vegetation cover, and the proportion of evergreen trees was a better predictor of ant species richness than the depth of the soil organic layer. In disturbed stands, habitat conditions have deteriorated; subsequently, some silvicolous ant species have disappeared.     

Even the Smallest Non-Crop Habitat Islands Could Be Beneficial: Distribution of Carabid Beetles and Spiders in Agricultural Landscape

Carabid beetles and ground-dwelling spiders inhabiting agroecosystems are beneficial organisms with a potential to control pest species. Intensification of agricultural management and reduction of areas covered by non-crop vegetation during recent decades in some areas has led to many potentially serious environmental problems including a decline in the diversity and abundance of beneficial arthropods in agricultural landscapes. This study investigated carabid beetle and spider assemblages in non-crop habitat islands of various sizes (50 to 18,000 square metres) within one large field, as well as the arable land within the field, using pitfall traps in two consecutive sampling periods (spring to early summer and peak summer). The non-crop habitat islands situated inside arable land hosted many unique ground-dwelling arthropod species that were not present within the surrounding arable land. Even the smallest non-crop habitat islands with areas of tens of square metres were inhabited by assemblages substantially different from these inhabiting arable land and thus enhanced the biodiversity of agricultural landscapes. The non-crop habitat area substantially affected the activity density, recorded species richness and recorded species composition of carabid and ground-dwelling spider assemblages; however, the effects were weakened when species specialised to non-crop habitats species were analysed separately. Interestingly, recorded species richness of spiders increased with non-crop habitat area, whereas recorded species richness of carabid beetles exhibited an opposite trend. There was substantial temporal variation in the spatial distribution of ground-dwelling arthropods, and contrasting patterns were observed for particular taxa (carabid beetles and spiders). In general, local environmental conditions (i.e., non-crop habitat island tree cover, shrub cover, grass cover and litter depth) were better determinants of arthropod assemblages than non-crop habitat island size, indicating that the creation of quite small but diversified (e.g., differing in vegetation cover) non-crop habitat islands could be the most efficient tool for the maintenance and enhancement of diversity of ground-dwelling carabids and spiders in agricultural landscapes.     

Effect of mother’s physical condition during pregnancy and lactation on postnatal growth and reproductive success of offspring in water vole <Emphasis Type="Italic">Arvicola terrestris</Emphasis>

Postnatal growth, life span, and probability of reproduction in the adult state depended on the mother’s physical condition during pregnancy and lactation in water vole. The white fat weight in the female abdominal cavity was shown to significantly increase in pregnancy and to decrease in late lactation. As an indicators for nutritional state of females, their body weight difference after parturition (or in late lactation) and expected from the regression equation relating individual body weight at the beginning and the end of each reproductive stage were used (physical condition indexes in pregnancy or lactation). The correlation of the physical condition index in pregnancy with the storage fat weight was 0.67. The metabolic resources of the mother’s body proved to favor faster offspring development. The female offspring weight at the age of 3 and 10 weeks as well as adult ones positively correlated with the mother’s nutritional state in pregnancy, while the male offspring weight demonstrated a similar correlation at the age of 3 and 6 weeks. Increased negative energy balance during lactation proved to decrease the offspring weight in both sexes after separation from mother and at the age of 6 weeks. High nutritional state of mother in pregnancy favored both the probability of reproduction and life span of female offspring. The reproduction of male offspring did not depend on the mother’s physical condition. The life span peaked in male offspring of mothers in a nutritional state below average in pregnancy and above average in lactation. Thus, the physical condition of the mother’s body is an important sex-dependent factor of phenotypic variation in the offspring body weight, reproductive competence, and life span.     

Effect of mother's physical condition during pregnancy and lactation on postnatal growth and reproductive success of offspring in water vole Arvicola terrestris L

Postnatal growth, life span, and probability of reproduction in the adult state depended on the mother's physical condition during pregnancy and lactation in water vole. The white fat weight in the female abdominal cavity was shown to significantly increase in pregnancy and to decrease in late lactation. As an indicators for nutritional state of females, their body weight difference after parturition (or in late lactation) and expected from the regression equation relating individual body weight at the beginning and the end of each reproductive stage were used (physical condition indexes in pregnancy or lactation). The correlation of the physical condition index in pregnancy with the storage fat weight was 0.67. The metabolic resources of the mother's body proved to favor faster offspring development. The female offspring weight at the age of 3 and 10 weeks as well as adult ones positively correlated with the mother's nutritional state in pregnancy, while the male offspring weight demonstrated a similar correlation at the age of 3 and 6 weeks. Increased negative energy balance during lactation proved to decrease the offspring weight in both sexes after separation from mother and at the age of 6 weeks. High nutritional state of mother in pregnancy favored both the probability of reproduction and life span of female offspring. The reproduction of male offspring did not depend on the mother's physical condition. The life span peaked in male offspring of mothers in a nutritional state below average in pregnancy and above average in lactation. Thus, the physical condition of the mother's body is an important sex-dependent factor of phenotypic variation in the offspring body weight, reproductive competence, and life span.     

Territoriality and nutritional condition in Cynotilapia afra (Gunther) and Pseudotropheus zebra (Boulenger), cichlidae in Lake Malawi National Park, Malawi

The standard length, diurnal activities, territory sizes, and areas over which individuals foraged and the nutritional condition of territorial and non-territorial Cynotilapia afra and Pseudotropheus zebra were compared. Results show that territorial tenureship in these fishes does not depend on the male size, implying that aggressiveness, experience and motivation are more important in the maintenance of territory. However, terri-toriality in C. afra (t = 10.93, P < 0.05) and P. zebra (t = 3.31, P < 0.05) manifests itself in the reduction of nutritional condition because territorial males engage in energetically more demanding activities, e.g. courtship displays, fertilizing ripe females and chasing intruders that trespass into their territories. Territorial C. afra (t = 4.77; P < 0.05) and P. zebra (t = 5.89; P < 0.05) also fed over significantly smaller areas and spent significantly less time feeding than did non-territorial males. The biological significance of territoriality in these fish species is therefore not food intake, but mate attraction and reproductive success for which they trade off their nutritional condition. However, there might be a nutritional threshold below which the cost out-weighs the benefit. Hence, territorial males in poorer health abandon their territories in order to regain their condition.     

Importance of closed landfills as green space in urbanized areas: ecological assessment using carabid beetles

In a closed landfill, we investigated the diversity and ecological characters of carabid beetles to understand the ecological importance of closed landfills that have the potential as a multi-functional habitat for improving biodiversity in urbanized areas. In addition, we studied the influence of environmental factors (vegetation structure, soil) on distribution and diversity of carabid beetles. A total of 92,495 individuals representing 15 carabid species were collected from the closed landfill. Although the species richness of carabid beetles recorded in the closed landfill was not higher than the other green spaces in the city, the closed landfill could sufficiently provides a stable habitat as a semi-natural area for carabid beetles. Soil pH, Na, and tall grass plant cover influenced carabid assemblage in the closed landfill. However, other environmental variables (e.g., K⁺, Na⁺, Mg²⁺, bare land cover, weedy cover, and tree cover) were not correlated with carabid species composition. It is implied that in the closed landfill, which is a highly modified engineered environment, other abiotic environmental (e.g., drainage, soil texture, leachate, and landscape context, etc.) and biotic factors (e.g., intra- and interspecific competition) may have affected carabid assemblage. Although artificial drainages are essential facilities for landfill management, they are a critical factor that affects the species inhabiting the landfill. However, carabid beetles seemed to randomly fall into the artificial drainage. For successful management of closed landfills, it is very important that minimize the intervention and that develop the ecological sensitively management method.     

Behavioral Tactics of Male Sockeye Salmon (Oncorhynchus nerka) under Varying Operational Sex Ratios

Previous studies have demonstrated several reproductive-behavior patterns in male salmon, including competitive and sneaking tactics, the formation of hierarchies, and non-hierarchical aggregations around ripe females. Through behavioral observations at varying spatial and temporal scales, we examined the hypothesis that operational sex ratio (OSR) determines male sockeye salmon (Oncorhynchus nerka) distribution and breeding tactics. Patterns of male distribution and behavior varied over both coarse and fine scales, associated with apparent shifts in reproductive opportunities, the physical characteristics of the breeding sites, and the deterioration of the fish as they approached death. Females spawned completely within a few days of arriving on the spawning grounds, whereas males courted the available ripe females from the date of their arrival on the spawning ground until their death. This difference in reproductive lifespans tended to elevate late-season OSRs but was partially counterbalanced by male departures and the arrival of other ripe females. The proportion of males able to dominate access to ripe females decreased and the number of large courting groups increased over the course of the season, apparently related to both increasing OSR and the deteriorating physical condition of males. However, great variation in OSR was observed within the spawning sites on a given day. OSRs were generally higher in shallow than in deep water, perhaps because larger females or more desirable breeding sites were concentrated in shallow water. The aggregations of males courting females were not stable (i.e. many arrivals and departures took place) and male aggression varied with group size. Aggression was most frequent at low OSRs and in groups of intermediate size (2–4 males per female), and much less frequent in larger groups, consistent with the needs of maximizing reproductive opportunities while minimizing unproductive energy expenditure. These results indicate that, while OSR strongly influences male distribution and behavior, other factors such as physical condition, limited movement and habitat choice may also affect male reproductive opportunities.