The impact of intralacustrine introductions with regard to space utilization and competition for territories to a cichlid fish community in Lake Malawi,Africa

At Thumbi Island West in Lake Malawi, a number of cichlid species was introduced from elsewhere in the lake some 20 years ago. These are now, in part, well established and could have an effect on the indigenous fish fauna. Repeated removals of the males of the introduced species from their territories were undertaken and artificial miniature reefs were established to find out whether the emptied existing territories or the newly established potential territories would be occupied by indigenous species. The results show that the emptied territories were almost invariably re-occupied by either a male of the introduced species conspecific to the previous territory holder or they remained empty. The newly established miniature reefs were not occupied by territorial males of either introduced or indigenous species but by young fish of a variety of species.     

Coexistence of permanently territorial cichlids of the genus Petrochromis through male-mating attack

Coexistence mechanisms of permanently territorial fishes proposed hitherto have been mainly based on interspecific competition. To test the hypothesis of the coexistence through male-mating aggression (Kohda, 1995a), spacing patterns of feeding territories of three coexisting herbivorous cichlids, Petrochromis polyodon, P. trewavasae and P. famula, were investigated in a homogeneous habitat at a shallow rocky shore in Lake Tanganyika. All three species maintained individual feeding territories that were defended against both conspecifics and congenerics and rarely overlapped either intra- or interspecifically. Territories of three species were scattered and covered almost all rocky bottoms. Territorial attacks usually occurred near the border of feeding territories. But male P. polyodon, the largest and socially most dominant fish, attacked conspecific males far beyond their feeding territories, which were widely separated. Removal of some territory owners suggested that such attacks result in the separate distribution of male P. polyodon territories. The interstitial space between P. polyodon feeding territories resulting from their mating aggression can be occupied by subordinate congeneric fishes. Male-mating attack of dominant males of Petrochromis will facilitate the coexistence of other congeners.     

Predation risk determines breeding territory choice in a Mediterranean cavity-nesting bird community

Non-direct effects of predation can be an important component of the total effect of predation, modulating animal population and community dynamics. The isolated effects of predation risk on the spatial organisation of the breeding bird community, however, remains poorly studied. We investigated whether an experimentally increased predation risk prior to reproduction affected breeding territory selection and subsequent reproductive strategies in three Mediterranean cavity-nesting birds, i.e., the little owl Athene noctua, European roller Coracias garrulus and scops owl Otus scops. We found that territories used the previous year were more likely to be re-occupied when they belonged to the safe treatment rather than to the risky treatment. The first choice of breeders of all three species was for safe territories over risky ones. When all breeding attempts in the season (i.e., final occupation) were considered, breeders also preferred safe to risky sites. In addition, little owls laid larger eggs in risky territories than in safe territories. Our study provides experimental evidence of a rapid preventive response of the three most abundant species in a cavity-nesting bird community to a short-term manipulation of predation risk. This response highlights the key role of the non-direct effects of predation in modulating avian community organisation.     

Modification of benthic communities by territorial damselfish: a multi-species comparison

The effects of territorial damselfish on coral reef benthos have been well-studied for a few relatively large-bodied species with visually distinct territories. Despite a growing body of research demonstrating their abundance, and their effects on algae, corals and other grazers, there has been little research on the effects of the territorial damselfish community as a whole. This study investigated the space occupation, territory composition, and diet of ten damselfish species at three locations: Magnetic and Orpheus Islands (Great Barrier Reef), and Kimbe Bay (Papua New Guinea). Territories were measured, and the composition of benthic communities inside and outside territories was assessed both in situ and from algal collections. The stomach contents of territorial damselfishes were also quantified. Although the larger, previously well-studied species had the most visible effect on the benthic community in their territories, all the smaller species also significantly affected the algal composition, normally with an increase of palatable algae. However, the composition of algal assemblages inside the territories of different species varied considerably. Damselfish territories were highly individual, not just among species, but also among locations. Diets were diverse and indicated a greater degree of omnivory and detritivory than previously thought. At all locations, territories occupied a substantial proportion of the substratum: >25% on Magnetic Island, >50% at Orpheus Island, and ∼50% in Kimbe Bay. Within individual zones, this figure was as high as 70%. The contribution of territorial damselfishes to a range of benthic patterns and processes is considerable, and future benthic studies may need to distinguish more closely between territory and non-territory areas. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Landmark Territoriality in the Neotropical Paper Wasps Polistes canadensis (L.) and P. carnifex (F.) (Hymenoptera: Vespidae)

Male Polistes canadensis and P. carnifex aggregate along crests of prominent ridges in Santa Rosa National Park, Costa Rica. At these sites males of both species defend territories (trees and shrubs) by chasing conspecific rivals. Territories do not contain nests or resources that are collected by females. Chasing by territorial males reduces the amount of time spent by intruders in a territory. I describe and contrast male territorial behavior of both species. Some male P. canadensis are territorial while others in the same area exhibit patrolling behavior, flying from one occupied territory to another. Males of P. carnifex exhibit territoriality only. Patrolling in P. canadensis is an outcome of relatively high male density along the ridge, rendering territories in short supply, as shown by the observation that experimentally vacated territories are seized rapidly by formerly patrolling males. Due to a high intraspecific intrusion rate, territorial male P. canadensis spend less time perching and more time flying and chasing intruders from their territories than do male P. carnifex. Males of these two species also differ in the placement of their territories along the ridgeline; P. canadensis occupy territories in saddles while P. carnifex occupy those at peaktops. I show that this divergent spatial pattern is not maintained by competitive exclusion of either species by the other, and I discuss alternative explanations for their separate spatial distributions. Comparative data suggest that males are territorial because females restrict matings to within territories, and I discuss alternative hypotheses to explain this bias in female behavior.     

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.     

The territorial defense hypothesis and the ecology of insular vertebrates

Insular lizards, birds, and mammals in high-density populations often exhibit reduced situation-specific aggression toward conspecifics. This aggressive behavior can be expressed in the form of (1) reduced territory sizes, (2) increased territory overlap with neighbors, (3) acceptance of subordinates on the territory, (4) reduced aggressiveness to certain classes of conspecifics, or (5) abandonment of territorial defense. These behavioral traits can be explained by two nonexclusive hypotheses. The resource hypothesis suggests that territorial behavior is primarily adjusted to resource densities, and that resources are more abundant on islands than on the mainland (e.g., because of a lack of competing species). The defense hypothesis suggests that, in addition to any effects of resources, the costs of defense against both territorial intruders and contenders for vacant territories are higher on islands. Recent theoretical and empirical studies indicate that these behavioral changes can occur as a result of elevated defense costs, independent of resource densities. Reduced predation, more benign climates, and an absence of habitat sinks on islands would all tend to increase the density of potential intruders and contenders, and hence the costs of defense for owners of insular territories. The two hypotheses differ in their predictions about the rates of biomass production (growth or reproduction) for holders of insular territories. Reproductive and growth data from insular-mainland pairs indicate the importance of elevated defense costs, and also suggest that many insular vertebrates reallocate their breeding resources so as to produce young that are more competitive. The suite of ecological and behavioral traits exhibited by insular territorial vertebrates can best be explained by three factors operating in concert: higher available resource densities, higher defense costs, and (sometimes) a reallocation of resources to produce young that are more competitive.     

The social organization of adult blue tangs, <Emphasis Type="Italic">Acanthurus coeruleus</Emphasis>, on a fringing reef,Barbados, West Indies

Blue tangs in Barbados exhibit three distinct social modes: territorial, schooling and wandering. We compared the mobility, foraging, aggression performed and received and the use of cleaning stations of adult blue tangs among modes and among habitats within a single fringing reef in Barbados. Evidence from observed switches during focal observations and multiple observations of tagged individuals indicate that fish are either territorial or non-territorial. Non-territorial fish formed schools and wandered. However, wandering can be used during solitary movements by fish in either type. Fish in the territorial mode, not previously described in adults of this species, restricted activity to a small area overlapping the territories of other tangs. They actively chased conspecifics and were chased mostly by damselfishes. They swam more slowly and fed at higher rates than other modes. Fish in the schooling mode ranged widely in compact, polarized groups of conspecifics, congeners and other species. They were not aggressive and were attacked mostly by damselfishes. They swam rapidly and fed at intermediate rates. Fish in the wanderer mode showed neither aggression nor association with other individuals. They swam rapidly, well above the substrate, fed little, were chased by conspecifics, ocean surgeonfish, A. bahianus, and damselfish and visited cleaning stations more often then other modes. All three modes were observed in all four main zones of the reef, and their behaviour changed quantitatively with habitat type. We suggest that territoriality reduces competition for algal food, schooling allows fish to overcome the food defence by damselfish, tangs and ocean surgeonfish, and wandering permits solitary movement over the reef to cleaning stations, feeding sites and other resources.     

Does interspecific competition affect territorial distribution of birds? A long‐term study on Siberian Phylloscopus warblers

Seven species of Phylloscopus warblers breed in the middle taiga subzone of Central Siberia. We studied their territorial distribution over 18 breeding seasons between 1978 and 1996. Phylloscopus inornatus , the most abundant breeding bird species in the study area, breeds in dense clusters of territories. In some habitats, these clusters are very unstable from year to year, providing an excellent opportunity to study the influence of P. inornatus on the territorial distribution of other species. We present a method for quantifying decreases in a species' breeding density caused by the presence of a competing congener. Analysing all possible combinations of species, we found reciprocal territorial avoidance between P. inornatus and P. proregulus as well as between P. inornatus and P. collybita . This is consistent with the hypothesis that territorial avoidance occurs between the species of greatest morphological similarity. The occurrence of territorial avoidance is not explained by habitat characteristics such as vegetation cover and productivity. However low productivity may be necessary for territorial avoidance to occur. An alternative explanation for the observed territorial separation of some species, derived from the Fretwell-Lucas model of habitat occupancy, is examined and rejected. Causes and implications of clustering behaviour of P. inornatus are also discussed.     

Prospecting in a solitary breeder: chick production elicits territorial intrusions in common loons

In many species, young animals learn about various breedingpatches in one year and use what they have learned to settlein a promising patch the next. Common loons (Gavia immer) seemgood candidates for such prospecting as prebreeders and displacedbreeders intrude frequently into breeding territories defendedby monogamous pairs yet engage in no extrapair copulations.We tested 3 hypotheses for prospecting in loons. The permanentattributes hypothesis gained little support as we found no consistentdifferences in quality between territories and no physical orbiotic trait that predicted reproductive success in a territory.We found some support for the conspecific attraction hypothesisas intruders were attracted to conspecifics in a lake in theshort term; however, intrusions were not more frequent in territoriesthat had experienced regular use by a pair the previous yearthan in territories that had previously been vacant. Instead,the increase in intrusion rate after a year of chick productionsupported the habitat-copying hypothesis, which states thatfloaters use the presence of chicks as a cue to target territoriesfor future attempts at territorial takeover. Despite this systemof prospecting, founding of new territories was common. Onestriking finding was the tendency of territorial breeders toconceal chicks from flying intruders, perhaps to avoid futureterritorial takeover.     

Effect of density on the reproductive behaviour of the territorial male demoiselle Chromis dispilus (Pisces: Pomacentridae)

Synopsis Characteristics of nest sites and reproductive behaviour of territorial male demoiselles Chromis dispilus were examined at sites of low, medium and high population density in northeastern New Zealand, by use of SCUBA diving and a remote underwater video system. Nest territories were closer together at high density (relative to areas of low population density) and this was accompanied by an increase in the frequency of spawning sequences and territorial defence against conspecifics. Territorial males were more aggressive during periods of display and spawning than during egg guarding. Fish from areas of low density spent more time on courtship display (signal jumps) and territorial defence against fish of other species, than demoiselles from areas of high population density. Territories and nest sizes tended to be largest in areas of medium population density. As in other damselfishes, population density is a major determinant of the frequency and intensity of reproductive behaviours.     

Male Territoriality in the Bumblebee Wolf,Philanthus bicinctus (Mickel) (Hymenoptera,Sphecidae): Observations on the Behaviour of Individual ♂♂

Observations were made on the behaviour of male Philanthus bicinctus (Mickel) which defend territories around the nesting burrows of ♀♀. ♂♂ were individually identified with small number tags and focal-animal samples were taken of the behaviour of these ♂♂. Intense interactions were observed between territorial ♂♂ and 1) conspecifics and 2) intruding insects which resembled conspecifics. Evidence is presented which suggests that ♂♂ scent mark their territories.     

Kin discrimination in salmonids

Summary The data presented here suggest that significant selection pressures towards kin discrimination behaviour patterns result from kin-biased territorial defence behaviour patterns. Salmonids employ a phenotype matching recognition mechanism allowing individuals to discriminate unfamiliar kin. Kin discrimination abilities allow individuals to reduce the levels of aggression associated with territorial defence towards related conspecifics and to defend smaller territories near kin versus non-kin. This kin-biased territorial defence behaviour is observed in at least one species under a wide range of territorial quality conditions. Within kin groups, subordinate individuals obtain a greater number of foraging attempts, resulting in kin-biased foraging within the social group. As a result of this kin-bias, individuals within kin groups show significantly higher mean weight increases (increased direct fitness benefits) and reduced variance in these increases (increased indirect benefits). Since all individuals within the kin groups obtained higher, less variable weight increases, we can argue that individuals are increasing their inclusive fitness as a result of these kin-biased behaviour patterns.Based on these results, and on what is known about the life history of a variety of salmonid and non-salmonid species, we can formulate a number of testable predictions. By testing these predictions, we may be better able to understand both the proximate and ultimate causation of kin discrimination abilities in a variety of fishes.     

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.     

Spectral Overlap in Songs and Temporal Avoidance in a Peruvian Bird Assemblage

Acoustic signals are very important in communication among conspecifics for many animals, especially in relatively dark, densely foliated habitat such as tropical rainforest. However, as many species all have to use the same acoustic space, this may lead to interference and masking. Songs will function efficiently in attracting mates or defending territories depending on the level of spectral and temporal overlap with songs of other species in the local community. As signal efficiency has direct fitness consequences, organisms may have evolved a strategy to avoid competition for acoustic space. We hypothesize that, as a consequence, species in a complex rainforest community may show spectral segregation or temporal avoidance: singing at a different frequency or at a different time. As doing both is superfluous and acoustic space is limited, we expect a negative correlation between spectral and temporal overlap. We tested this hypothesis in recordings of 20 vocally dominant bird species from 14 families in a Peruvian rainforest. Using pair‐wise comparisons between bird species, we found one significant correlation between spectral overlap and temporal overlap. All others were non‐significant. However, we did find that, at frequencies more often used by multiple bird species, there was significantly less overlap than expected if all species were just singing randomly ignorant of each other’s songs. Our analyses indicate that short‐term auditory feedback mechanisms may also operate in the biodiverse environments of the tropics and may explain part of the community dynamics of acoustic signalling in the rainforest.     

Territoriality in Redlip blennies (Ophioblennius atlanticus-Pisces: Blenniidae)

Ophioblennius atlanticus is a shallow-water, surge zone Caribbean species, invariably as adult found in permanent territories on a solid substrate. As such it controls space within a complex community. Planar dimensions of the territories of adults show a mean area of about 05 m². Antagonistic reactions of different degree are shown to both transgressing and intruding conspecifics, and to a much lesser extent to other species. It is hypothesized that interstitial territories of immature specimens provide the nuclei of future territories. Non-reproductive, extra-territorial activity is not common, but does occur. Evidence is presented to show that dispossessed adult blennies have great difficulty in establishing new territories. Territories that have been vacated are subsequently shared by neighbouring fish. Territory provides space, food and shelter for its resident, and as an occupied, but not impermeable space, it is an agent of control of distribution of Redlip blennies and other species of fish.     

Day and night feeding territoriality in Willets Catoptrophorus semipalmatus and Whimbrel Numenius phaeopus during the non-breeding season in the tropics

This study was carried out to determine whether or not, and under what conditions, Willets Catoptrophorus semipalmatus and Whimbrel Numenius phaeopus continue to defend and use their daytime feeding territories at night in a tropical environment. The study was conducted in coastal Venezuela by registering, by night and by day, the behaviour and the position of colour-marked and radio-tagged birds. Night observations were made with the use of a light intensifier.
In Willets, both territorial and non-territorial birds were observed. The use and defence of territories was observed only on sandy mud areas where Fiddler Crabs Uca cumulanta were plentiful; the soft mud sites were used only by non-territorial birds. All Whimbrel were territorial. Individuals of both species defending a given space during daytime continued to occupy and defend the same area during the night. The proportion of birds foraging at night was higher on moonlit than on moonless nights. Territorial defence involved alert postures, parallel walks, calls and the chasing of intruders. Agonistic encounters between Willets and Whimbrel and other species were rare. Willets were territorial at all states of the tide except when high tides flooded the territories. Tide had no effect on the time of feeding in Whimbrel. Moonlight was clearly the factor conditioning the occurrence of nocturnal foraging on territories by both species. Foraging strategies and the type of prey and substrata explain why the incidence of night foraging varied with moonlight in territorial Willets and Whimbrels and not in non-territorial individuals.     

Harem structure and female territoriality in the dwarf hawkfish <Emphasis Type="Italic">Cirrhitichthys falco</Emphasis> (Cirrhitidae)

We investigated the territoriality and the spatial and mating relationships of the haremic hawkfish, Cirrhitichthys falco, on a reef off Kuchierabu-jima Island in southern Japan. Each individual maintained a territorial home range which was defended against same-sex conspecifics at the boundary of the home range. The territory of each male encompassed the territories of 2–3 females, allowing the male to completely monopolize mating opportunities with those females. Based on our observations, we classified the harem type of C. falco as a territorial female type. Large juveniles maintained independent home ranges outside the female territories. In contrast, small juveniles were allowed to cohabit within the territory of an adult female. Stomach contents analysis revealed that the smallest size class of C. falco fed primarily on copepods. In contrast, all other size classes fed primarily on decapods. Together, these results suggest that female territoriality plays an important role in defending food resources.     

Sex Differences in Territorial Behavior Exhibited by the Spotted Hyena (Hyaenidae, Crocuta crocuta)

Spotted hyenas ( Crocuta crocuta ) are gregarious carnivores that defend group territories against encroachment by neighboring conspecifics. Here we monitored the behavior of members of one clan of free-ranging spotted hyenas during border patrols, 'wars' with neighboring clans, and other interactions with alien intruders, to document differences between the sexes in territorial behavior in this species. We also examined the possibility that the probability or rate of attack on alien hyenas encountered within the clan's territory would vary with the sex of the intruders. Initiation and leadership of most cooperative territorial behaviors were by adult female clan members, although border patrols were occasionally conducted by groups composed exclusively of resident males. The vast majority of alien intruders into the territory of the study clan were males. Resident females were more likely to attack intruding females than intruding males, but hourly rates of aggression directed by females towards aliens did not vary with intruder sex. Resident males were more likely than resident females to attack alien males, and resident males directed significantly higher hourly rates of aggression towards intruding males than females. Although female leadership in most cooperative territorial behaviors distinguishes spotted hyenas from many mammalian carnivores, other sex differences in the territorial behavior of spotted hyenas resemble those documented in other gregarious predators. Sex differences observed in hyena territoriality are consistent with the hypothesis that male and female clan members derive different selective benefits from advertisement and defense of group territories.     

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.