Interspecific competition between stream-dwelling brown trout and Alpine bullhead

Density and composition of benthic invertebrates and the diet of brown trout Salmo trutta and Alpine bullhead Cottus poecilopus were studied at two sites in one Norwegian stream. The sites were separated by an impassable waterfall, and brown trout density was five to 10 times higher at the upper, allopatric site than downstream where it lived in sympatry with the Alpine bullhead. Benthic invertebrate communities did not differ between sites; however, the size distribution of chironomids and trichopterans were skewed towards lighter individuals at the sympatric site. Diet composition suggested that sympatric brown trout foraged more on invertebrate drift and from the surface than allopatric brown trout. Alpine bullhead diet did not differ significantly from brown trout diet, except that the Alpine bullhead fed on heavier individual prey within a few taxa and did not consume chironomid pupae or surface insects. The collected data support the hypothesis that brown trout living in sympatry with Alpine bullhead feed at locations with higher predation risk, which is a probable explanation for their lower population density.     

Intercohort competition causes spatial segregation in brown trout in artificial streams

When together in a flume, 13-cm L _s (age-1) trout appeared to increase their activity and behaved more aggressively, whereas 10-cm L _s (age-0) trout tended to move less and were rarely aggressive in the presence of the larger size class. Both size classes were less mobile and preferred lower water velocities in winter than in summer, and increased their use of instream cover in winter. When both size classes were present, only small trout decreased their use of low water velocities and cover. The results indicate that intercohort competition may cause spatial segregation among size groups of brown trout, especially in winter when trout attempt to minimize their maintenance costs.     

Selection on Arctic charr generated by competition from brown trout

We experimentally explored population‐ and individual‐level effects on Arctic charr (Salvelinus alpinus) resulting from resource competition with its common European competitor, the brown trout (Salmo trutta). At the population level, we compared performance of the two species in their natural sympatric state with that of Arctic charr in allopatry. At the individual level, we established selection gradients for morphological traits of Arctic charr in allopatric and in sympatric conditions. We found evidence for interspecific competition likely by interference at the population level when comparing differences in average performance between treatments. The growth and feeding rates did not differ significantly between allopatric and sympatric Arctic charr despite lower charr densities (substitutive design) in sympatric enclosures indicating that inter‐ and intraspecific competition are of similar strength. The two species showed distinct niche segregation in sympatry, and brown trout grew faster than Arctic charr. Arctic charr did not expand their niche in allopatry, indicating that the two species compete to a limited degree for the same resources and that interference may suppress the growth of charr in sympatric enclosures. At the individual level, however, we found directional selection in sympatric enclosures against individual Arctic charr with large head and long fins and against individuals feeding on zoobenthos rather than zooplankton indicating competition for common resources (possibly exploitative) between trout and these charr individuals. In allopatric enclosures these relations were not significant. Diets were correlated to the morphology supporting selection against the benthic‐feeding type, i.e. individuals with morphology and feeding behaviour most similar to their competitor, the benthic feeding brown trout. Thus, this study lends support to the hypothesis that Arctic charr have evolved in competition with brown trout, and through ecological character displacement adapted to their present niche.     

Triploid progeny from a female Atlantic salmon × brown trout hybrid backcrossed to a male brown trout

A female Atlantic salmon × brown trout hybrid was backcrossed to a male brown trout. Electrophoretic analysis of diagnostic enzymes showed that the progeny were triploid. However, a few individuals were partially diploid.     

Factors influencing the spawning migration of female anadromous brown trout

Radio telemetry was employed to study movements of adult female anadromous brown trout Salmo trutta (sea trout) during upstream spawning migration and following spawning in a stream with tributaries. Sea trout were monitored by manual tracking and by automatic listening stations. The latter suggested that initiation of upstream migration was positively correlated with stream discharge. Individual sea trout performed repeated upstream migration 'initiations'(visits) to areas where they were detected by the automatic listening stations. The first and subsequent upstream migration 'initiations' occurred under conditions of similar water temperature and stream discharge. Manual tracking indicated that in the pre‐spawning state, the distance migrated over 3 days was positively correlated with stream discharge and water temperature, whereas in the post‐spawning state, the total distance migrated was not correlated with any of these two environmental variables.     

Individual variation in habitat use and growth of male and female brown trout

We examined variation in growth and habitat use of individually PIT‐tagged brown trout Salmo trutta in three stream enclosures, each divided into a fine substrate, deep pool habitat and a coarse substrate, shallow habitat. Habitat use and movements of individual fish were monitored continually by placing PIT detectors between habitats. All fish were measured and weighed biweekly over a three month period. There was no significant relationship between habitat use and initial body size, nor was there a consistent relationship between habitat use and densities of benthic macroinvertebrates or abundance of drifting invertebrates in the two habitats. Most habitat changes occurred at night, with activity peaks just prior to sunrise and after sunset. Trout used pools more at night than during the day. Within any given day, diurnal and nocturnal habitat use of individual fish varied little, with variation greater at night than during the day. Partial habitat segregation by sex was observed; only males used pools extensively during daytime, whereas males and females used riffles.
Growth rate was positively related to use of pools during daytime but not at night. Growth rate was also affected by enclosure, with growth rates being highest in the most downstream enclosure, which had the deepest pool (mean of 42 cm) and lowest in the most upstream enclosure, which had the shallowest pool (mean of 28 cm). A complete exchange of trout between the most upstream and downstream enclosure indicated that the enclosure effect was due to physical differences and not to individual fish differences between enclosures. The effect appears to have been caused by differences in depth as daytime use of pools was correlated with the area of the pool ≥35 cm deep, and production of trout biomass per enclosure was directly related to mean pool depth. Our results suggest that there is a relationship between habitat use and growth of individuals that is independent of body size, but that this relationship is influenced by sex of the fish and by the physical characteristics of the environment. Further, the data indicate that short‐term behavioral decisions on habitat use by brown trout have a potential effect on longer‐term individual fitness through growth rates.     

Sexual selection in pinyon jays I: female choice and male-male competition

Although pinyon jays, Gymnorhinus cyanocephalus, mate monogamously and male parental investment is high, the mechanisms of sexual selection occur in this species. Experiments were performed using captive birds to determine which male traits are favoured by female choice and which are favoured by inter-male competition. Females mated preferentially with males that were brighter in colour and had larger testes. Females did not prefer large males, but inter-male competition favoured males with larger bills and heavier bodies. Male dominance was not the primary determinant of male mating succes, but it probably conferred an advantage.     

Relationship between oxidative stress and circulating testosterone and cortisol in pre-spawning female brown trout

Reproduction in vertebrates is an energy-demanding process that is mediated by endogenous hormones and potentially results in oxidative stress. The primary aim of this study was to quantify the relationship between oxidative stress parameters (antioxidant capacity and levels of reactive oxygen metabolites) and circulating testosterone and cortisol in a common and widespread teleost fish, the brown trout (Salmo trutta, L.). Results show that trout with higher testosterone levels prior to spawning have higher levels of oxidative damage at the time that they spawn (although by the time of spawning testosterone levels had dropped, leading to a negative relationship between testosterone and oxidative damage at that time). Cortisol levels were not directly related to oxidative damage or antioxidant capacity, but concentrations of this hormone were positively related to levels of fungal infection, which was itself associated both with lower antioxidant capacity and lower levels of oxidative damage. These results highlight the complexity of interactions between different components of the endocrine system and metabolism and suggest that caution be used in interpreting relationships between a single hormone and indicators of oxidative balance or other fitness proxies.     

Parasites, male-male competition and female mate choice in the sand goby

Male sand goby Pomatoschistus minutus dominance in competition over nest sites was associated with higher body condition but not with the intensity of infection with any individual parasite of six different species, nor with an overall index combining relative levels of infection by all parasites. Body condition was not related to the intensity of infection with any individual parasite nor with the index of total relative parasite load. In trials in which females spawned, they showed a tendency to choose dominant over subordinate males as mates, but did not consistently choose less parasitized males. Variation in the relative size of the dorsal fins of males was detected, and this related to numbers of the ectoparasitic monogenean Gyrodactylus sp., suggesting that at least some infections have phenotypic effects that could allow females to detect and avoid the most heavily infected males.     

Ontogenetic variation in density‐dependent growth of brown trout through habitat competition

1. Density‐dependent growth has been widely reported in freshwater fishes, but the ontogenetic evolution of competition and its subsequent effects on growth through a life span remains unclear. 2. Patterns of competition can be described by integrating population abundance data with habitat‐modelling results. Weighted usable area (WUA; m² WUA ha^?1) curves are obtained for each flow value and are then coupled with demographic data to obtain the occupancy rates (trout m^?2 WUA, the density of a given age class related to its suitable habitat) of the WUA for every age class, year and site. 3. We examined a long‐term data series searching for temporal variation in the influence of habitat occupancy rate on the growth of brown trout Salmo trutta. We tested whether (i) mean cohort mass (mean mass of the cohort during the first 3 years of life) is affected by the occupancy rate experienced across a life span; and (ii) the occupancy rate experienced at different ages influenced mean body size. 4. We observed a consistent negative power relationship between average cohort mass and mean occupancy rate through a life span, indicating that stronger cohorts were related to a reduced growth, with likely consequences for individual fitness. 5. The effects of occupancy rate on size‐at‐age were mainly detected in the size attained at the second year of life, but they were because of the competition at different times. Thus, the level of competition varied through ontogeny, in some of the rivers affecting growth since the first year of life, whereas in most of the rivers the main effects on body size resulted from the competition during the second year of life. 6. Occupancy rate appears more appropriate than density for assessing the occurrence of habitat competition in freshwater fishes, since it encompasses the differences in quantity and quality of suitable habitat for each age class. 7. Our study highlights the importance of density‐dependent growth as a key process in the dynamics of brown trout populations, its temporal variation depending on the temporal changes of density and the variation of competition associated with the habitat capacity for each life stage.     

Polygyny in the yellow-headed blackbird: female choice versus male competition

Several assumptions and predictions of the polygyny threshold and sexy son hypotheses, which were proposed to explain the maintenance of polygyny on the basis of female choice, were examined. An alternative neutral mate choice hypothesis in which male competition is responsible for polygyny was also examined. For the yellow-headed blackbird, Xanthocephalus xanthocephalus, neither territory nor male features affected female choice of mate or female reproductive success. In the study population, polygyny occurred because males competed to hold territories in order to gain access to females. Since females settled apparently randomly, males that were more aggreessive and were therefore able to secure large territories had larger harems. Several criteria must be met in order for the neutral mate choice hypothesis to apply. Evidence from the literature suggests that for some species the criteria are not met and polygyny occurs due to female choice. However, several studies provide evidence that the neutral mate choice hypothesis may apply in some populations.     

Territorial competition among wild brown trout fry: effects of ownership and body size

The effects of ownership, energy status and relative size on the ability to defend a territory were studied in dyadic contests among wild brown trout fry. A logistic regression showed that the territory owner won 85% of the contests with opponents of equal size, and that a 30% size advantage was required to balance the advantage of ownership. Energy status did not affect the ability to win territorial contests.     

Male-male competition and large size mating advantage in European earwigs, Forficula auricularia

European earwigs are sexually dimorphic in forceps shape and length. Male forceps are thought to be weapons in male contests for access to females, but recent findings suggest that females choose males on the basis of their forceps length. I investigated sexual selection on forceps length and body size and the occurrence of male-male competition. When I controlled for forceps length experimentally and statistically, relatively heavy males had greater copulation success than relatively light males. When I controlled for body size, males with relatively longer forceps had no tendency for greater copulation success than males with shorter forceps. Relatively heavy males more often took over copulations from smaller males than vice versa. Male contests were important for the outcome of mate competition, as males commonly interrupted and took over copulations. My results therefore suggest that intrasexual selection is significant in competition for copulations in male earwigs, and acts on body size. This contrasts with previous findings, which have shown intersexual selection on forceps length to be important. However, both modes of sexual selection may be acting through a two-stage process, where male-male competition first determines which males have access to females, and then through female choice among available males. Morphological measurements supported the conclusion that forceps length and body size are male secondary sexual characters, as these characters had large variance and skewed distributions in males, but were normally distributed in females. Copyright 2000 The Association for the Study of Animal Behaviour.     

Male competition and female choice interact to determine mating success in the bluefin killifish

Whether male competition and female choice act in concert, independently,or in opposition is a critical issue for understanding sexualselection. In complex social systems, the outcomes of pairwiseinteractions may not be accurate indicators of how sexual selectionemerges. We investigated how female choice and male competitioninteract in the bluefin killifish, Lucania goodei, in a 3-stagedexperiment where 1) females could choose between 2 males, 2)those males could interact in the presence of that female, and3) females and males could freely interact and spawn. In thepairwise stages (1 and 2), females displayed pronounced preferencesbetween males and male competition produced a distinctly dominantindividual. None of the morphological traits, including color,measured in males were associated with either female preferenceor male dominance. When all 3 fish interacted (stage 3), maleactivity level was the sole predictor of spawning success. Maleswith elevated activity levels were more aggressive toward malesand females, exhibited intensified courtship, and obtained morespawns. Female preference did not predict the number of spawnswith a male, but it did predict her latency to spawn; femalesspawned more quickly with preferred males. Thus, male competitionand female choice interact to determine reproductive success,but there is evidence for conflict and a cost to females ofassociating with dominant males. Reproductive success in thisspecies is not easily predicted from simple measures of morphologyor female preference and is influenced by complex social interactions,both between males, and between males and females.     

Male-male competition and repeated evolution of terrestrial breeding in Atlantic Coastal Forest frogs*

Terrestrial breeding is a derived condition in frogs, with multiple transitions from an aquatic ancestor. Shifts in reproductive mode often involve changes in habitat use, and these are typically associated with diversification in body plans, with repeated transitions imposing similar selective pressures. We examine the diversification of reproductive modes, male and female body sizes, and sexual size dimorphism (SSD) in the Neotropical frog genera Cycloramphus and Zachaenus, both endemic to the Atlantic rainforest of Brazil. Species in this clade either breed in rocky streams (saxicolous) or in terrestrial environments, allowing us to investigate reproductive habitat shifts. We constructed a multilocus molecular phylogeny and inferred evolutionary histories of reproductive habitats, body sizes, and SSD. The common ancestor was small, saxicolous, and had low SSD. Terrestrial breeding evolved independently three times and we found a significant association between reproductive habitat and SSD, with shifts to terrestrial breeding evolving in correlation with decreases in male body size, but not female body size. Terrestrial breeding increases the availability of breeding sites and results in concealment of amplexus, egg-laying, and parental care, therefore reducing male-male competition at all stages of reproduction. We conclude that correlated evolution of terrestrial reproduction and small males is due to release from intense male-male competition that is typical of exposed saxicolous breeding.     

The volitional travel speed varies with reproductive state in mature female brown trout Salmo trutta

This study tested the effect of reproduction on the volitional travel speed of mature female brown trout Salmo trutta L. The downstream travel speed in the pre-spawning state was 0·25 m s⁻¹ (95% CI : 0·19, 0·34) while it increased significantly to 0·65 m s⁻¹ (95% CI: 0·49, 0·87) in the post-spawning state. The results suggest state-dependent travel speed in S. trutta .     

Prey orientation in piscivorous brown trout

Piscivorous brown trout Salmo trutta change their feeding behaviour depending on prey species, prey size and number of prey eaten. In trout which had eaten fish recently, most had one fish in their stomach, but up to 16 prey fish were found. Individuals of the small-sized minnow Phoxinus phoxinus were swallowed chiefly tail first, whereas individuals of the larger Arctic charr Salvelinus alpinus were taken both head and tail first. The largest charr were swallowed head first. In stomachs containing more than one fish prey, prey orientation was likely to be mixed. For all three types of prey orientation (i.e. tail first, head first and mixed), significant and positive correlations existed between prey length and predator length. The maximum prey size eaten tail first or in mixed orientation was about 70–85% of the size of prey eaten head first, indicating morphological advantages in eating the prey head first.