225:), particular common sequences of syllables (phrases) are produced by all males established in the same location (neighbours), whereas males of different locations (strangers) share only few syllables. Playback experiments provided evidence for neighbourâstranger discrimination consistent with the dear enemy effect, indicating that shared sequences were recognised and identified as markers of the group identity. Studies have shown that the dear enemy effect changes during the breeding season of the skylark. Playbacks of neighbour and stranger songs at three periods of the breeding season show that neighbours are dear enemies in the middle of the season, when territories are stable, but not at the beginning of the breeding season, during settlement and pair formation, nor at the end, when bird density increases due to the presence of young birds becoming independent. In song sparrows, where neighbours are most often the sires of extra-pair offspring, males will alter their aggression toward neighbouring males with their female's fertility status. When presented with simulated stranger and neighbour intruders during their female's pre-fertile and post-fertile periods, males displayed the dear enemy effect. However, when presented with simulated stranger and neighbour intruders during their female's fertile period, males exhibited an equal response to both stimuli, likely in order to protect their paternity. Thus, the dear enemy relationship is not a fixed pattern but a flexible one likely to evolve with social and ecological circumstances.
403:) defend territories that consist of a breeding burrow and a display area where they wave their claw to attract females. Burrow-holding males engage in agonistic contests with both intruding males that attempt burrow take-overs and with other territory-holding neighbours that apparently attempt to limit waving or other surface activities of rivals. Contests consist of one or more behavioural elements that range from no claw contact to use of the claw to push, grip, or flip an opponent. In the field, contests with intruders begin at higher intensities and escalate more rapidly than those with neighbours. However, residentâresident contests increase in intensity when burrows are close, neighbours faced each other when exiting burrows, and neighbours were of similar size. Proximity and orientation determine the ease with which a neighbour may be engaged.
132:. Badgers show heightened behavioural responses towards unfamiliar- compared with self-group scents, but there is no difference in response to neighbour- relative to self-group scents. The relative responses towards unfamiliar-group scents are greatest during the breeding seasons, but there is no seasonal differences in the responses to neighbour-group versus self-group scents. In badger populations, levels of aggression between neighbouring territory-holders are likely to be kept relatively low through neighbour recognition. However, increased levels of aggression will be shown towards dispersing or itinerant (alien) badgers, especially during periods such as the breeding season when the potential threats to the long-term fitness of territory owners are greatest.
148:
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487:) groups vocalize more and inspect more scent samples in response to olfactory cues of neighbours than strangers. It has been suggested that increased aggression towards neighbours is more common in social species with intense competition between neighbours, as opposed to reduced aggression towards neighbours typical for most solitary species. Furthermore, animals may respond in this way when encounters with intruders from non-neighbouring colonies are rare and of little consequence.
430:
343:) have been demonstrated in the field. Playbacks of non-resident sounds from a given fish's territory elicit a greater response from its nearest neighbour than playbacks of the resident's sound. Testing also included switching the sounds of the two nearest neighbours relative to each respective male's territory. Results demonstrated that all males in the colony individually recognize the sounds of their two nearest neighbours.
262:), can individually recognize neighbours and will increase aggression towards them as the threat to territorial ownership increases. Resident males treat familiar neighbours that had been moved to the opposite boundary to the shared boundary as equally aggressive as strangers. However, residents responded more aggressively towards strangers than towards neighbours on natural territories and also in neutral arena encounters.
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aggressive individual should enjoy greater benefits than a non-aggressive individual when each is faced with a non-aggressive opponent. This stipulation is plausible, as an aggressive individual might enlarge their territory or steal food or matings from a non-aggressive individual. When cooperation involves a cost, a possible mechanism for achieving stable co-operation is
98:
332:) to examine the dear enemy effect. When faced with a familiar neighbour and an unfamiliar intruder simultaneously, residents preferentially confronted the unfamiliar opponent. That is, the establishment of dear enemy recognition between a resident and a neighbour allowed the resident to direct his aggression to the greater competitive threat, i.e. the intruder.
298:), have a large variability in call characteristics and are able to discriminate between neighbouring and unfamiliar conspecifics. Calling is of the longest duration in response to an unfamiliar acoustic stimulus; in contrast, the response to a familiar conspecific call does not show any difference from solitary vocalisations. Terrestrial red-backed salamanders,
379:. Behavioural tests with workers reveal no alarm behaviour or mortality in pairings of workers from the same colony but a full range from no alarm to overt aggression, with associated death, when individuals were paired from different colonies. The level of mortality increases with differences in the composition of
351:
The home ranges of colony living ants often overlap the ranges of other conspecific colonies and colonies of other species. In laboratory experiments, the frequency and severity of agonistic interactions among workers from different colonies increases with the distance between their nests; this has
62:
The ultimate function of the dear enemy effect is to increase the individual fitness of the animal expressing the behaviour. This increase in fitness is achieved by reducing the time, energy or risk of injury unnecessarily incurred by defending a territory or its resources (e.g. mate, food, space)
42:
phenomenon in which two neighbouring territorial animals become less aggressive toward one another once territorial borders are well established. As territory owners become accustomed to their neighbours, they expend less time and energy on defensive behaviors directed toward one another. However,
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sniffed both castoreum and anal gland secretion from a stranger longer than from a neighbour. Furthermore, beavers responded aggressivelyâstanding on the mound on their hind feet, pawing, overmarking, or a combination of theseâlonger to castoreum, but not to anal gland secretion, from a stranger
75:
game. In this view, a territory owner that acts non-aggressively towards a neighbour can be thought of as cooperating, while a territory owner that acts aggressively towards its neighbour can be considered to have defected. A necessary condition for the prisonerâs dilemma game to hold is that an
321:) is dependent on the presence of females. Reduced aggression consistent with dear enemy recognition occurs between conspecific neighbours in the absence of females, but the presence of a female in a male's territory instigates comparably greater aggression between the neighbours.
214:), hoots to defend its territory. Male little owls respond less to their neighbour's hoots played back from the usual location. However, responses to playback of a neighbour from an unusual location are similar to responses to playback of a stranger's hoots from either location.
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than from a neighbour. When the mounds containing the scents were allowed to remain overnight and the beavers' responses measured the following morning, the beavers' responses were stronger to both castoreum and anal gland secretion from a stranger than from a neighbour.
46:
The dear enemy effect has been observed in a wide range of animals including mammals, birds, reptiles, amphibians, fish and invertebrates. It can be modulated by factors such as the location of the familiar and unfamiliar animal, the season, and the presence of females.
43:
aggression toward unfamiliar neighbours remains the same. Some authors have suggested the dear enemy effect is territory residents displaying lower levels of aggression toward familiar neighbours compared to unfamiliar individuals who are non-territorial "floaters".
527:) males which live in gangs do not differ in their response behaviour toward neighbouring and stranger males and largely ignore any non-gang member, irrespective of familiarity; that is, they neither show a "dear enemy" nor "nasty neighbour" effect.
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Lesbarrèresa, D. and LodĂŠa, T., (2002). Variations in male calls and responses to an unfamiliar advertisement call in a territorial breeding anuran, Rana dalmatina: evidence for a âdear enemyâ effect. Ethology, Ecology & Evolution, 14: 287-295.
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Audio playback studies are often used to test the dear enemy effect in birds. These studies have demonstrated several bird species respond more aggressively to played back songs of strangers than to songs of neighbours; such species include the
251:), reduced their aggression levels in repeat interactions with familiar rivals and increased their aggression levels towards unfamiliar males. The time taken for interactions to be settled was also lower towards familiar than unfamiliar males.
195:) differ individually in their aggressiveness. Increased aggression by residents towards intruders indicates that residents not only respond to intrinsic aggressiveness of their neighbours, but also to short-term changes in aggression levels.
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Brunton, D.H., Evans, B., Cope, T. and Ji, W. (2008). A test of the dear enemy hypothesis in female New
Zealand bellbirds (Anthornis melanura): female neighbors as threats. Behavioral Ecology, 19 (4): 791-798. DOI: 10.1093/beheco/arn027
896:
Hkinzk, J., Foitzik, S., Hippert, A. and HĂślldobler, B., (1996). Apparent dear-enemy phenomenon and environment-based recognition cues in the ant
Leptothorax nylanderi. Ethology, 102: 510â522. DOI: 10.1111/j.1439-0310.1996.tb01143.x
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against a familiar animal with its own territory; the territory-holder already knows about the abilities of the neighbour, and also knows that the neighbour is unlikely to try to take over the territory because it already has one.
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Leiser, J.K., (2003). When are neighbours âdear enemiesâ and when are they not? The responses of territorial male variegated pupfish, Cyprinodon variegatus, to neighbours, strangers and heterospecifics. Animal
Behaviour, 65:
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are able to discriminate the odours of familiar neighbours and strangers. It has been suggested that this discrimination may be used by males to avoid unnecessary chases and fights by becoming known to their neighbours.
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Kaib1, M., Franke, S., Francke, W. and Brand, R., (2002). Cuticular hydrocarbons in a termite: phenotypes and a neighbourâstranger effect. Physiological
Entomology, 27, 189â198. DOI: 10.1046/j.1365-3032.2002.00292.x
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Maciej, P., Patzelt, A., Ndao, I., Hammerschmidt, K. and Julia
Fischer, J., (2013). Social monitoring in a multilevel society: a playback study with male Guinea baboons. Behav. Ecol. Sociobiol., 67(1): 61â68. DOI:
479:) are able to recognize a greater proportion of workers from neighbouring colonies as non-colony members. When recognized as non-colony members, more aggression is exhibited toward neighbours than non-neighbours.
80:, where pairs of individuals trade bouts of cooperative behaviour with one another. Dear enemy cooperation could be explained by reciprocal altruism if territorial neighbours use conditional strategies such as
273:), dyads of males behave differently depending on whether the lizards are prior neighbours, with prior neighbours exhibiting less bobbing relative to nodding forms of headbob displays than non-neighbours.
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Mollesf, L.E. and
Vehrencamp, S.L., (2001). Neighbour recognition by resident males in the banded wren, Thryothorus pleurostictus, a tropical songbird with high song type sharing. Animal Behaviour, 61:
498:) are more aggressive toward the songs of neighbouring females. This is opposite to the dear enemy phenomenon and suggests that neighbouring females pose a greater threat than strangers in this species.
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McMann, S. and
Paterson, A.V., (2012). Display behavior of resident brown anoles (Anolis sagrei) during close encounters with neighbors and nonneighbors. Herpetological Conservation and Biology, 7(1):
639:
VachĂŠ, M., Ferron, J. and Gouat, P., (2001). The ability of red squirrels (Tamiasciurus hudsonicus) to discriminate conspecific olfactory signatures. Canadian
Journal of Zoology, 79: 1296-1300
517:) do not discriminate behaviourally between the calls of neighbours and strangers, and female collared lizards show no difference in their behaviour to neighbouring or unfamiliar females.
302:, defend territories under rocks and logs on the forest floor in the eastern United States. Individuals are more aggressive to unfamiliar salamanders than to familiar individuals. In the
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Moser-Purdy, C; MacDougall-Shackleton, E; Mennill, D. J. (2017). "Enemies are not always dear: male song sparrows adjust dear enemy effect expression in response to female fertility".
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Pratt, A.E. and McLain, D.K., (2006). How dear is my enemy: Intruder-resident and resident-resident encounters in male sand fiddler crabs (Uca pugilator). Behaviour, 143: 597-617
812:
Briefer, E., Rybak, F. and Aubin, T., (2008). When to be a dear enemy: flexible acoustic relationships of neighbouring skylarks, Alauda arvensis. Animal
Behaviour, 76: 1319â1325
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Leiser, J.K. and
Itzkowitz, M., (1989). The benefits of dear enemy recognition in three-contender convict cichlid (Cichlasoma nigrofasciatum) contests. Behaviour, 136: 983-1003
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A range of studies have found evidence of an effect opposite to the dear enemy effect, i.e. more aggression is shown toward neighbours than strangers. This has been termed the
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Husakf, J.F. and Fox, S.F., (2003). Adult male collared lizards, Crotaphytus collaris, increase aggression towards displaced neighbours. Animal Behaviour, 65: 391â396
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Hyman, J., (2002). Conditional strategies in territorial defense: do Carolina wrens play tit-for-tat? Behavioral Ecology, 13: 664-669. DOI: 10.1093/beheco/13.5.664
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Bard, S., Hau, M., Wikelski, M. and Wingfield, J.C. (2002). Vocal distinctiveness and response to conspecific playback in the spotted antbird. Condor, 104: 387-394
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Newey, P.S., Robson, S.K. and Crozier, R.H., (2010). Weaver ants Oecophylla smaragdina encounter nasty neighbors rather than dear enemies. Ecology, 91(8):2366-72
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have been known to slowly fly into the territory of an adjacent male territory holder in order to test and establish the mutual boundary of their two territories.
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Myrberg, A.A. and Riggio, R.J., (1985). Acoustically mediated individual recognition by a coral reef fish (Pomacentrus partitus). Animal Behaviour, 33: 411â416
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Husak, J.F. and Fox, S.F., (2003). Spatial organisation and the dear enemy phenomenon in adult female collared lizards., Journal of Herpetology, 37, 211-215
306:, there is variability in the nature of the call and the frog presents a more aggressive call to strangers in comparison to the response to its neighbors.
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Palphramand1, K.L. and White, P.C.L., (2007). Badgers, Meles meles, discriminate between neighbour, alien and self scent. Animal Behaviour, 74: 429â436
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Briefer, E., Aubin, T., Lehongre, K. and Rybak, F., (2008). How to identify dear enemies: the group signature in the complex song of the skylark
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Falls, J.B. and McNicholl, M.K., (1979). Neighbor-stranger discrimination by song in male blue grouse. Canadian Journal of Zoology, 57: 457-462
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Frankie, Gordon W.; Vinson, S. B.; Lewis, Alcinda (1979-04-01). "Territorial Behavior in Male Xylocopa micans (Hymenoptera: Anthophoridae)".
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Brindley, E.L., (1991). Response of European robins to playback of song: neighbor recognition and overlapping. Animal Behaviour, 41: 503-512
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MĂźller, C.A. and Manser, M.B., (2007). âNasty neighboursâ rather than âdear enemiesâ in a social carnivore. Proc. R. Soc. B., 274: 959-965
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Akçay, C. et al., (2009). Good neighbour, bad neighbour: song sparrows retaliate against aggressive rivals. Animal Behaviour, 78: 97â102
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282:) has been observed to be less aggressive towards conspecifics. It also exhibits headbob activity similar to that of the brown anole.
84:. In the tit-for-tat strategy, a subject will cooperate when its partner (neighbour) cooperates and defect when the partner defects.
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Jaeger, R. G., (1981). Dear enemy recognition and the costs of aggression between salamanders. American Naturalist 117: 964-972.
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Langen, T.A., Tripet, F. and Nonacs, P., (2000). The red and the black: habituation and the dear-enemy phenomenon in two desert
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A range of studies have found no evidence of the dear enemy effect showing the effect is not universal. Territorial males of the
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Rosell, F. and Bjørkøyli, T. (2002). A test of the dear enemy phenomenon in the Eurasian beaver. Animal Behaviour, 63: 1073â1078
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Osborne, L., (2005). Rival recognition in the territorial tawny dragon (Ctenophorus decresii). Acta Ethologica, 8: 45-50
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Hardouin, L.A., Tabel, P. and Bretagnolle, V., (2006). Neighbourâstranger discrimination in the little owl,
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Lovell, S.F. (2004). Neighbor-stranger discrimination by song in a suboscine bird, the alder flycatcher,
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birds, it has seldom been investigated in territorial non-passerine species. The nocturnal
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Bee, M.A., (2003). A test of the "dear enemy effect" in the strawberry dart-poison frog (
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The dear enemy effect has been reported in colonies of the fungus-growing termite
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547:(Huxley, J., Hardy, A. and Ford, E., eds). London, Allen and Unwin. pp. 71-83
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The little owl hoots less intensively at familiar neighbours than unfamiliar
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732:. Journal of Experimental Biology, 211: 317-326. DOI: 10.1242/jeb.013359
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The interaction between two neighbours can be modelled as an iterated
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Although neighbourâstranger discrimination has been reported in many
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Individual recognition of noises produced males of the bicolor
678:. Behavioral Ecology, 15: 799-804 DOI: 10.1093/beheco/arh082
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Some researchers have staged three-way contests between male
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ants. Behavioral Ecology and Sociobiology, 48: 285-292
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Fisher, J., {1954}. Evolution and bird sociality. In:
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588:). Behavioral Ecology and Sociobiology, 54: 601-610
314:The dear enemy effect in male variegated pupfish (
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393:Male sand fiddler crabs attract mates by waving
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561:. Sinauer Associates. pp. 281â282.
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254:Another territorial lizard, the
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113:) presented with a two-way
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1328:Jane Goodall
1288:Donald Broom
1257:Zoosemiotics
1210:Sociobiology
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1283:Marc Bekoff
1271:Ethologists
525:Papio papio
267:brown anole
247:Ctenophorus
229:In reptiles
178:banded wren
166:blue grouse
126:Meles meles
82:tit for tat
40:ethological
1483:Categories
1220:Structures
1215:Stereotypy
531:References
509:) and the
473:weaver ant
464:media help
337:damselfish
319:variegatus
317:Cyprinodon
292:agile frog
208:little owl
93:In mammals
88:Occurrence
1449:Behaviour
1392:Societies
1230:Honeycomb
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381:cuticular
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67:Mechanism
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1468:Category
1413:Journals
1240:Instinct
1190:Learning
1185:Instinct
1160:Ethogram
1143:Grooming
1066:Branches
1059:Ethology
986:Archived
933:25083909
907:Pheidole
800:53273443
660:June 15,
361:Pheidole
269:lizard (
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1128:Breed
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840:27â37
796:S2CID
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