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Regulation of territorial behavior in the sedentary song sparrow, Melospiza melodia morphna.

Male song sparrows (Melospiza melodia morphna) of western Washington State show year-round territoriality. Although territorial aggression during the breeding season was accompanied by high circulating levels of luteinizing hormone (LH) and testosterone (T), similar aggression in the non-breeding season occurred when gonadal hormone levels were basal. Experimental removal of territorial males in autumn resulted in new males taking over the territory within a few days. These males had basal levels of LH and T despite establishing a new territory and the accompanying high level of aggression. There was also no difference in plasma levels of corticosterone in replacement versus established territorial males suggesting that social stress was not a factor. To test the possibility that low levels of gonadal steroids, or potentially some gonadal hormone not measured here, might regulate territorial aggression in the non-breeding season, free-living male song sparrows were castrated in early autumn and their aggressive territorial behavior was quantified by a simulated territorial intrusion (STI). Castrates showed the same aggressive responses to STI as controls indicating that establishment and maintenance of a territory was not dependent upon hormones of gonadal origin. Castrated males retained territories throughout the autumn and winter and even into the following breeding season. Further experiments were conducted to establish a role, if any, for T in territorial behavior in this species. During the breeding season when males had high levels of T, the responses to STI continued after the stimuli (caged conspecific male and tape-recorded songs) were withdrawn. Some males continued to patrol the territory and sang for long periods after STI. In contrast, males exposed to STI in autumn showed a strong aggressive response during the challenge, but then territorial aggression abated rapidly when the stimuli were withdrawn. Implants of T into males in autumn reinstated the high level of aggression after withdrawal of STI in a manner similar to that seen during the breeding season. These data suggest that in M. m. morphna T does not activate territorial aggression per se, but may increase the intensity of an aggressive response to STI, and persistence of the behavior after STI is withdrawn.

Aggression↗

Evidence that helping at the nest does not result in territory inheritance in the Seychelles warbler.

In an environment that has a shortage of territories, helping to rear younger siblings ('alloparenting') is proposed to facilitate territory acquisition in two ways: (i) through group augmentation that leads to an increase of the territory with subsequent partial inheritance (budding); and (ii) through site dominance that leads to greater success when competing for the natal or a nearby territory after the death of the territory owner (complete territory inheritance). Most young Seychelles warbler (Acrocephalus sechellensis) males either show alloparenting or budding behaviour. Future budders had significantly more aggressive interactions with neighbours and assisted their parents more with territory defence than similarly aged future alloparents or non-helpers. This led to an increase of the natal territory of future budders before actual budding took place, whereas the natal territories of future alloparents remained constant in size. Alloparents never became budders and vice versa, refuting partial inheritance as an advantage of alloparenting. Natural male breeding vacancies were never inherited by alloparents born on vacant or other territories, but were inherited by budders born on the vacant territory or, if these were absent, predominantly by budders from neighbouring territories. We offer explicit experimental evidence against the 'helping at the nest to inherit' hypothesis. Experimentally created male breeding vacancies, with both a male alloparent and a similarly aged sibling budder present simultaneously in the vacant territory, were filled by budders only. Site dominance over territory inheritance is linked to budding and not to alloparenting.

Animals↗

Territory acquisition in loons: the importance of take-over.

We examined patterns of territory acquisition and reconnaissance in common loons, Gavia immer, from northern Wisconsin. Among all territory acquisitions, 41.5% occurred through passive occupation of territories left vacant after the death or desertion of a previous resident, 17% constituted founding of new territories and the remaining 41.5% came about through take-over: either usurpation of defended territories or appropriation of territories before the seasonal return of previous owners. Take-overs occurred in both sexes, but individuals acted alone, never in pairs. Displaced breeders usually took refuge on undefended lakes near their former territories; about half of these loons later regained former territories through passive occupation or took possession of new territories elsewhere. As predicted by the reconnaissance hypothesis, usurpations occurred most often in territories that had produced chicks during the previous 12 months, suggesting that loons use the presence or absence of chicks as a cue for territorial usurpation. Large individuals of both sexes held onto territories longer than small individuals, an indication that body size might be correlated with fighting ability. In terms of life history, loons appear to locate good territories through reconnaissance, usurp them in a subsequent year and recover from displacements by reclaiming their original territories or new ones. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Interactions between cleaning gobies and territorial damselfish on coral reefs.

We studied the interactions between cleaning gobies, Elacatinus spp., at cleaning stations and territorial dusky damselfish, Stegastes fuscus, on a Barbadian fringing reef to enhance our understanding of the behavioural impact of cleaning stations on members of a coral reef community. Cleaning stations within damselfish territories were visited by significantly fewer species and fewer individuals than those outside damselfish territories. At cleaning stations within damselfish territories, the main client was the territory holder itself. Client behaviour did not differ between the two types of cleaning station; however, cleaning gobies at stations within damselfish territories spent less time cleaning clients, their cleaning bouts were shorter, and their feeding rate on clients was slower. These effects appear to be due to the repeated aggression of territorial damselfish towards fish intruding on to their territories to visit cleaning stations. The overlap between a territorial damselfish and a cleaning station therefore appears to have detrimental impacts on cleaning goby foraging. The presence of cleaning stations within damselfish territories exerted some effects on the territory owners. Although the rates of intrusions were similar on territories with and without cleaning stations, damselfish with cleaning stations tended to chase intruders more often, had a lower foraging rate, and were at higher risk of egg predation because of increased intrusions by egg predators. A potential benefit to territorial damselfish is regular access to a cleaning station; however, the magnitude of this benefit is unknown. This study reveals that damselfish, which are ubiquitous on coral reefs, can generate significant variation in levels of use of cleaning stations, which leads to new questions relating to the settlement behaviour of both cleaning gobies and damselfish. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Scent marking in a territorial African antelope: I. The maintenance of borders between male oribi.

Scent marking is ubiquitous among the dwarf antelope and gazelles of Africa, but its function has been the subject of debate. This study examined preorbital gland scent marking in the oribi, Ourebia ourebi, a territorial African antelope. Several hypotheses for the function of scent marking by territorial antelope were tested with observational data. Of these, the hypotheses that scent marking is driven by intrasexual competition between neighbouring males, and that marks serve as an honest advertisement of a male's ability to defend his territory from rivals, were supported best. Thirty-three territorial male oribi on 23 territories marked most at borders shared with other territorial males, and territorial males marked more often at borders shared with multimale groups than at borders shared with a single male. This suggests that males perceived neighbouring male groups as a greater threat to territory ownership than neighbouring males that defended their territories without the aid of adult subordinates. Marking rate was unrelated to territory size or the number of females on adjacent territories, but males with many male neighbours marked at higher rates than those with fewer male neighbours. These results suggest that the presence of male neighbours has a greater effect on the scent marking behaviour of territorial antelope than has been considered previously. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Scent marking in a territorial African antelope: II. The economics of marking with faeces.

Faeces may be ideal substances for scent marking because they have a minimal energetic cost to the signaller. However, marking with faeces is also constrained by the animal's ability to produce faeces. This study examined whether limits on the volume of faeces produced by oribi Ourebia ourebi, in Serengeti National Park, Tanzania, caused territorial males to regulate their output and prioritize the placement of faecal marks. Territorial males marked with faeces more often, and with a smaller volume per defecation, than did juvenile males and females. Territorial males also defecated only on established dung middens along borders shared with other territorial males or on top of a female's urine and faeces. In contrast, juvenile males and females defecated randomly with regard to their location in territories. Territorial males with larger harems marked with faeces at higher rates and less volume than males with few or no females. This difference suggests that when males overmark female excretions they reduce the amount of faeces available for marking other preferred sites, such as along territory borders shared with other males. Dominant males with adult subordinates marked with faeces less often, and with a greater volume per mark, than males that defended territories without the aid of subordinates. Dominant males also reduced the volume of marks less as the number of females on their territory increased than did males without subordinates. Territories occupied by more than one adult male also were marked with faeces at higher rates, and with marks of greater volume, than territories held by single males. These results suggest that the presence of subordinate males reduced the demand on dominant males to regulate the volume and placement of faecal marks. Overall, these results suggest that territorial male oribi regulate their faecal marking behaviour in response to a limited supply of faeces. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗