Prior residence, territory quality and life-history strategies in juvenile Atlantic salmon (Salmo salar L.)

Three groups of juvenile salmon were introduced sequentially into an artificial stream to investigate the effects of prior residence on behaviour and territory choice. Almost half of the first group obtained and defended distinct territories, the other half being constrained to an area approximately the size of one large territory. All of the fish in the subsequent groups, bar one, were also constrained to the same site. Since the fish were of similar size, prior residence alone seemed to influence which individuals obtained territories. However, within the first group, the fish that obtained territories were larger and more aggressive. The territorial fish did not appear to choose the most profitable territories, although they had the greatest opportunity to do so. Since juvenile salmon emerge from their gravel nests fairly synchronously, a time constraint on site sampling is hypothesised: there may be a risk in taking time to sample sites, since these same sites may become occupied with conspecifics. However, fish with territories fed at faster rates than non-territorial fish, possibly because of reduced competition for prey items. Consequently, fish from the first group (containing most of the territorial fish) grew faster than the other two groups. Moreover, most of the territory holders, but only one of the non-territorial fish, reached the threshold size that increases their probability of smolting the following year. This suggests that ability to obtain a defensible territory, primarily through prior residence, influences the age at which juvenile salmon can migrate to sea.     

Stocking experiment with Atlantic salmon and sea trout parr reared on either live prey or a pellet diet

Survival rate and growth parameters of Atlantic salmon fry and sea trout fry were determined after stocking in the wild. Before release (22 May 2009) into the wild the larvae were reared for 10 weeks in the hatchery in three groups: (i) fry fed on live zooplankton , (ii) fry fed on larvae of live nekton, and (iii) fry fed on prepared pellet food. In autumn (15 September 2010) the fish were caught in the wild; the survival rate and growth parameters of both Atlantic salmon and sea trout were the highest in the zooplankton‐fed group, whilst the pellet‐fed group had the lowest survival rate and growth value parameters. Most effective food for hatchery‐reared fishes to be used as stock was the natural living zooplankton. The general conclusion is that the live diet supplied in the rearing period has a positively impact on fish survival in the wild.     

Effect of competitor abundance on feeding territoriality in a grazing fish,the ayuPlecoglossus altivelis

The grazing fish, ayu,Plecoglossus altivelis Temminck & Schlegel, establishes feeding territorialiry during the young stage. The population density fluctuates from year to year by more than a hundredfold, but the determinant of territory size is less well known. The feeding territoriality of ayu was examined under simulated habitat conditions where fish density was manipulated and food resources were renewable. Fish competed for algae attached to the substrata and were divided into residents with territories, and floaters without territories. By experimental alteration of fish density the number of residents increased with density and rerritory size decreased with density. Floaters intruded into territories in a school to feed on algae, which induced overt aggression of the resident and reduced the productivity of algae growing there. Both the intruding frequency of floaters over territorial areas and their feeding pressure on algae increased at higher floater density. Floaters functioned to shift cost-benefit relationships for various territory sizes. They acted as food competitors to restrict territory size below a maximum through competitive interference. Although the growth rate of residents was inversely related to fish density, residents grew faster than floaters in each group. Under a given set of competitor abundances, economic defensibility determined territory size.     

Dominance hierarchies in group-living cleaning gobies: causes and foraging consequences

In species living in social groups, aggression among individuals to gain access to limiting resources can lead to the formation of stable social hierarchies. We tested whether dominance rank in social groups of sponge-dwelling cleaning gobies Elacatinus prochilos in Barbados was determined by physical attributes of individuals or by prior experience of dominance, and examined the foraging consequences of dominance rank. Intraspecific aggression within groups resulted in stable dominance hierarchies that were strongly correlated with fish length. Dominant individuals maintained exclusive territories while subordinate fish occupied broader home ranges. Larger, competitively dominant fish were able to monopolize areas inside the sponge lumen with the highest abundance of the polychaete Haplosyllis spp., a favoured prey item, and achieved the highest foraging rates. The removal of a territorial individual from large groups resulted in a domino-like effect in territory relocation of the remaining fish as individuals moved to the territory previously occupied by the individual just above them in the group hierarchy. Individuals added to existing groups generally failed to gain access to territories, despite being formerly dominant in their original groups. When given the opportunity to choose a location in the absence of larger competitors, gobies frequently preferred positions that were previously defended and that had abundant food. These results suggest that intraspecific competition for resources creates the observed dominance structures and provides support for the role of individual physical attributes in the formation and maintenance of dominance hierarchies.     

Prey availability and selective feeding of sea trout (Salmo trutta L., 1758) fry stocked in small forest streams

The food base and stomach contents of stocked sea trout (Salmo trutta) fry were determined in the first 4 months (April–August 2010) of life in two small lowland streams after resorption of the yolk sac. In each stream, 600 individual trout were released in a 200 m stocking‐section. The macrozoobenthos given as a food base for the fry were collected once a month from the streams using a bottom scraper. Three subsamples of macrozoobentos were considered as one sample from the stocking‐section. On the same day, 50 stocked fry were captured in each stream using electric fishing gear. Preferred food components were usually the taxa represented by number in a given stream in a particular month. Width size range of the prey in fish stomachs in May was from 0.10 to 1.47 mm, and from 0.11 to 3.78 mm in August. All food items found in both streams during the study months were also represented in intensity in the fish: Cyclopoida, and larvae of Baetidae and Chironomidae. Almost all Helodidae and Simuliidae larvae as well as the majority of Limoniidae and Nemouridae larvae were available as food each month.     

Territory quality affects the relative importance of habitat heterogeneity and interference competition in a long‐lived territorial songbird

Density‐dependent reproduction is commonly explained by either the habitat heterogeneity (HHH) or individual adjustment (IAH) hypothesis. Under the HHH, high quality territories are assumed to be occupied first. At higher density, occupation of low‐quality territories increases due to lower availability of high‐quality territories, which reduces mean reproductive success. Alternatively, the IAH assumes that increased competition at higher densities reduces reproductive success in all territories. For birds of prey, HHH plays an important role in territorial species, and IAH in socially breeding species. To test the generality of this hypothesis, we studied the mechanism behind density dependence in raven Corvus corax, a long‐lived passerine bird, using long‐term population data from a large number of territories. Population density decreased reproduction, which was explained by increased usage of low quality territories at higher density, supporting the HHH. Density reduced reproduction in low quality territories, but not in high and intermediate quality territories. We additionally compared the explanatory power of different models describing brood size, representing IAH, HHH, or a combination of both. The best model represented a combination of both hypotheses, in which the effect of density depended on territory quality. Our conclusion that both IAH and HHH are supported can be explained by the biology of ravens, where territorial adults not only experience interference competition with other territorial adults, but also with social groups of juveniles and floaters. We conclude that the relative importance of IAH and HHH may depend on variation in territory quality and social structure.     

Prey availability affects territory size,but not territorial display behavior,in green anole lizards

The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.     

Population density,resource patterning,and territoriality in the Everglades pygmy sunfish

The means by which pygmy sunfish compete for food are influenced by the density of the population and the dispersion of the prey as well as by the sex and dominance status of the individual. At all densities when the prey were predictably located in a central clump, males established territories. When prey were dispersed randomly in both the high and low density population, males abandoned territorial behaviour and, like females, swam freely about the aquaria. Only in the intermediate density populations did the males maintain territories and continue to defend resources. Development of a simple cost-benefit model shows that male territorial behaviour is governed to a large extent by economic considerations. Despite these overall patterns, differences in competitive strategies were observed within populations. Dominant individuals tended to possess territories nearest the central patch of prey, and use the most intense and physiologically exhausting displays. Only under the most stressful conditions did they acquire significantly more food than subordinates, and even then these benefits were not translated into increased growth, largely because dominant fish engaged in disproportionately more energy-consuming contests.     

The critical-period concept for juvenile survival and its relevance for population regulation in young sea trout, Salmo trutta

An example of density-dependent regulation is provided by a long-term investigation (1966-present) of a population of migratory trout (estuarine and sea trout), Salmo trutta L., in a Lake District stream. Evidence for the concept of a critical period for the survival of young fish is briefly reviewed and found to be rather equivocal. The concept is, however, relevant to the trout population. Loss rates were high before but low after a critical survival time ( t_c days after fry emergence) that varied between year-classes (range 33-70 days) and was inversely density-dependent on egg density. Survivor density and loss rates were strongly density-dependent on egg density before t _c, but proportionate survival with stable loss-rates occurred after t _c. Some trout established feeding territories soon after emergence and the number of fish without territories decreased from a high initial value to a negligible value at t _c. Fish size at t_c was not constant but increased as t _c increased. The range of t _c for the different year-classes was similar to that for survival times of unfed fry in the laboratory. A new stock-recruitment model, incorporating t _c, has been developed for the trout population and shown to be related to the model (Ricker curve) used in the long-term study. The critical time can also be regarded as the critical age for survival in young trout; this concept may be relevant to other fish species.     

An experimental analysis of territory size in juvenile steelhead trout

I experimentally manipulated levels of food abundance and density of competitors to determine how these factors influence the territory size of juvenile steelhead trout, Oncorhynchus mykiss. Steelhead trout were held in artificial stream channels and I followed cohorts that were fed at one of three levels of food abundance and stocked at one of three levels of fish density. By measuring territory size over a 2-month period, while the fish were growing, I was also able to assess the effects of body size in determining the size of a territory. Defended and foraging areas were similar in absolute size, but the frequency of space use was different for defence than for foraging. As predicted, territory size decreased with increasing levels of food abundance and increased with decreasing levels of fish density. In addition, territory size increased with increasing body size even after controlling the effects of food abundance and competitor density. In comparison to previous studies, territory size of steelhead trout changed more dramatically in response to changing levels of food and competitors. For territorial animals with indeterminate growth, territory size is not only adjusted as a trade-off between the costs and benefits of defence, but also with respect to body size due to increasing metabolic demands as individuals grow. Copyright 2000 The Association for the Study of Animal Behaviour.     

The use of territorial gardening versus kleptoparasitism by a subtropical reef fish (Kyphosus cornelii) is influenced by territory defendability

Many territorial herbivorous animals increase the productivityor quality of the prey community on their territory. However,this gardening tactic may be vulnerable to kleptoparasitismunless gardeners can exclude others from the patch. We investigatedthe influence of territory defendability on the decision ofwestern buffalo bream (Kyphosus cornelii) to defend an algalgarden or to leave their territories and invade those of conspecifics.Fish monitored while away from their territories did kleptoparasitizethe gardens of conspecifics. We manipulated the structural complexityof territories through the addition of short or tall simulatedvegetation. Increasing the height of simulated vegetation onthe territory decreased the defendability of territories butincreased their safety for fish; territories with tall vegetationwere more easily invaded by conspecifics, but fish returnedmore rapidly to these territories after being disturbed. Ina paired experiment, fish spent more time away from their territories(roving) when tall vegetation was added than when short vegetationwas added. However, this effect was influenced by depth, witha greater difference between treatments on deeper territories,and by body size, with a greater difference for larger fish.These results support the prediction that territory-holdingindividuals faced with a trade-off between defending their owngarden or kleptoparasitizing from others adjust their use ofthese tactics based on the defendability of resources.     

Territorial shift in the yellow-handed titi monkey (Callicebus torquatus)

In this study, the ranging pattern of a single group of monogamous and territorial Callicebus torquatus was recorded over a 7-year period from 1974 to 1980. Data on ranging pattern were recorded by direct observation; daily movement patterns were plotted on an accurate, scaled map of the territory. Data were obtained during 1–3-month studies each year from 1974 to 1978 and in 1980, as well as during a 12-month study in 1979-1980. These studies show that the exclusive territory maintained by the group was not spatially stable over the 7-year period. The group gradually moved its territory to the northeast, to the extent that the territory used in 1980 was completely noncoincident with the territory used in 1974 by the same group. Analysis of the ranging pattern from the 12-month study indicates that the group did not exhibit seasonal preferences for different parts of its range. Seasonal ranging patterns, therefore, are not likely to account for the observed territorial shift. It is suggested that territorial shifting may be related to the need for monogamous groups to find new territories for their offspring.     

Reduced territorial defence in rainbow trout fry exposed to a paper mill effluent: using the mirror image stimulation test as a behavioural bioassay

Rainbow trout Oncorhynchus mykiss fry exposed to a paper mill effluent showed increased EROD (ethoxyresorufin-O-deethylase) activity and reduced territorial aggression in mirror image stimulation (MIS) tests. These results suggest that the MIS test could be a useful tool to detect sublethal behavioural effects of pollution in fish fry.     

Territory quality determines social group composition in Ethiopian wolves Canis simensis

1. We contrast the value of four different models to predict variation in territory size as follows: resource density (the ideal free distribution), population density, group size and intruder pressure (relative resource-holding potential). In the framework of the resource dispersion hypothesis, we test the effect of resource abundance and spatial variation in resource distribution on the age/sex composition of social groups. 2. We explore these drivers of territory size and group size/composition in Ethiopian wolves Canis simensis in the Bale Mountains, Ethiopia, using fine-scale distribution maps of their major prey species based on satellite-derived vegetation maps. 3. The number of adult males is correlated with territory size, while prey density, wolf population density and intruder pressure are not associated with territory size. On average, each additional adult male increases territory size by 1.18 km(2). 4. Prey abundance increases with territory size (average biomass accumulation of 6.5 kg km(-2)), and larger territories provide greater per capita access to prime foraging habitat and prey. 5. The age/sex composition of wolf packs is more closely related to territory quality than territory size. Subordinate adult females are more likely to be present in territories with greater proportions of prime giant molerat Tachyoryctes macrocephalus habitat (i.e. >80% of Web Valley territories and >20% in Sanetti/Morebawa), and more yearlings (aged 12-23 months) occur in territories with greater overall prey biomass. 6. Wolf packs with restricted access to good foraging habitat tend to defend more exclusive territories, having a lower degree of overlap with neighbouring packs. 7. The greater per capita access to prey in large groups suggests a strong evolutionary advantage of collaborative territorial defence in this species, although the relative costs of territorial expansion vs. exclusion depend upon the spatial distribution of resources. We propose a model whereby territory size is determined by the number of adult males, with the presence of subordinate females and yearlings dependent on the quality of habitat, and the abundance and distribution of prey, incorporated within territory boundaries.     

The effect of food abundance on territory size and population density of juvenile steelhead trout (<Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis>)

Optimal territory size models predict a decrease in territory size with increasing food abundance. However, most of these models may not be applicable to juvenile salmonids in streams, because they defend contiguous territories at high densities. The optimal size of a contiguous territory is predicted to (1) be independent of food abundance when food is rare and (2) decrease only when food abundance is high enough to induce a reduction in territory size below the contiguous optimum. To test these predictions, we raised equal densities of juvenile steelhead trout in outdoor stream channels over a 32-fold range of food abundance in the absence of emigration for 25 days. Increasing competition for scarce food resulted in increasing mortality, higher willingness to emigrate, higher variance in body mass, lower growth, lower population density and lower biomass. The size of territories decreased with increasing local population density, and increased with increasing body size. However, territory size did not change with food abundance, a result consistent with the prediction of a contiguous territory size model. On average, total salmonid biomass increased 5.7 times in response to the 32-fold increase in food abundance. Our data provide strong support for an earlier quantitative relationship between the abundance of stream salmonids and their food.     

Feeding pattern and diet of first feeding brown trout fry under natural conditions

The feeding habits of brown trout Salmo trutta fry were studied during the critical first feeding period in a natural spawning and nursery stream. A low proportion of the fry initiated exogenous feeding before emergence from the gravel, and while having nearly 30% of the yolk sac remaining. This probably reflected low feeding motivation or limited feeding opportunities within the gravel environment. The majority of the fry started feeding after emergence, and after most or all yolk was absorbed. Some fry emerged with large amounts of yolk remaining, while others emerged after yolk exhaustion. The degree of stomach fullness revealed that feeding was more efficient after a territory had been acquired. The diets of the young fry were dominated by chironomid larvae, followed by zooplankton and Plecoptera larvae. Fry dispersing downstream and out of the nursery area were significantly smaller than resident fry, indicating displacement due to competition for territories. The majority of the downstream dispersing fry had initiated feeding, and there was at this point no evidence of starvation in any of the fry. It therefore appeared that the later emerging fry actively migrated out of the overpopulated nursery area to find available territories further downstream.     

Density dependence,territoriality, and divisibility of resources: from optimality models to population processes

Species differ enormously in their territorial systems. Some species defend only small areas surrounded by undefended space, while others defend large contiguous territories. Using an optimization approach, we show that this variation can be explained from the density of two types of resources: divisible and nondivisible. We assume that benefits of territories are monotonously related to the defended amount of divisible resources (hereafter called food). In contrast, no benefits are obtained without a nondivisible resource (hereafter called nest site) in the territory, while more than one nest site does not further increase the benefits. The optimal territory size depends on the relative abundance of these resources. With a low density of nest sites, the optimal territory size is small and includes only the nest site. If the density of nest sites is relatively large, the optimal territory size is high, and territories are contiguous. Competition for these different resources yields contrasting patterns of how populations are regulated. If there is mainly competition for nest sites, we expect density-dependent exclusion through territoriality and no density-dependent reproduction. When competition is mainly for food, we expect density-dependent reproduction because optimal territory size will be compressed at higher densities, resulting in lower reproductive success. These predicted patterns indeed are observed in some well-studied passerine species for which both the territorial system and the occurrence of density dependence is known.     

Causes and consequences of individual variation in territory size in the American red squirrel

For most territorial species that defend food or other resources, territory size is inversely related to resource density. However, in some food-based territorial species, larger territories are known to contain greater absolute resource availability. For these latter species, both the specific determinants of territory size, and the relationship between territory size and fitness, remain unclear. We predicted that in American red squirrels ( Tamiasciurus hudsonicus ), where larger territories tend to hold increased food resources, survival should be correlated positively with territory size and that both individual (body mass, sex, duration of territory ownership) and habitat-specific (population density) attributes would contribute to variation in territory size. We monitored survival and territory size of 58 squirrels during a two-year period using live-trapping and radio-telemetry. Survival and territory size were correlated positively, with each 0.1 ha decrease in territory size typically resulting in a 16.6% increase in mortality risk; mortality risk for squirrels with the smallest versus largest territories differed by up to 23-fold. Territory size was negatively related to local population density, but not to any attributes of individual squirrels, implying that, in this species, territory size may not be influenced strongly by individual quality. We conclude that while larger territories may confer a survival advantage to squirrels, territory size is unrelated to individual physical attributes, likely because costs associated with the acquisition of a larger or richer territory outweigh potential benefits of increased food access.     

An experimental study on the natural removal of dead trout fry in a Lake District stream

The chief objective was to discover why few moribund or dead Salmo trutta fry were observed on the stream bed when mortality within the stream was known to be high ( c . 13 000 dead fry year ⁻¹ for whole stream). Newly dead fry were placed in 20 boxes embedded in the stream bed (20 fry of known total weight per box) and arranged in pairs with one box open and one closed. One pair was removed every 2 days, the fish remains being weighed and the invertebrates in the open box being identified and counted. The experiments were performed from late April to early May in 1967, 1968, 1969 and the results were similar in each year. Both wet and dry weights of fry decreased exponentially but the rate of decrease was much higher in the open boxes; detectable fish remains were about 55% of initial weights after 20 days (end of experiment) in closed boxes but zero after 16 days in open boxes. Invertebrate scavengers were responsible for the higher loss rates in the open boxes and showed a definite succession with caddis larvae and carnivorous stonefly larvae dominant at first, but then being replaced by detritivorous stonefly larvae and freshwater shrimps. These experiments show clearly why dead fry disappear rapidly from the stream bed.     

Diet niche relationships among predator and prey fish species in their early life stages in Lake Võrtsjärv (Estonia)

In Lake Võrtsjärv pikeperch was observed not to shift to piscivory in their first autumn of life, although juvenile stages of a variety of fish species were abundant in the lake. It was hypothesized that the diets of predator and prey fish fry overlap and that coarse fish species are important food competitors for juvenile piscivores and thus, pikeperch and perch fry do not shift to piscivory during their first growing season. To discover the possible linkages in this pattern, in 2009 the feeding relationships of pikeperch, perch, ruffe and roach fry were analysed. The stomach content analyses showed that in the summer period, Mesocyclops leuckarti was the most frequent prey for perch and ruffe, pikeperch consumed Leptodora kindti in large quantities, and roach ate mostly plant material. Towards autumn, M. leuckarti was the most abundant prey for all percids. However, average stomach content weight and the number of prey items eaten by ruffe were considerably higher than for other fish fry. Since the feeding opportunities of fish fry are considered poor in the examined lake, the prey has the potential to restrict the recruitment to piscivory of their predators, as prey fish seem to have better abilities to persist in this ecosystem. Furthermore, supposed competition in the juvenile stage may result in a reduced top‐down effect on coarse fish.