139:, in a hypothesis known as the optimal-inbreeding hypothesis. He argued that, since philopatry leads to the concentration of related individuals in their birth areas, and thus reduced genetic diversity, there must be some advantage to inbreeding – otherwise the process would have been evolutionary detrimental and would not be so prevalent. The major beneficial outcome under this hypothesis is the protection of a local gene complex that is finely adapted to the local environment. Another proposed benefit is the reduction of the cost of meiosis and recombination events. Under this hypothesis, non-philopatric individuals would be maladapted and over multi-generational time, philopatry within a species could become fixed. Evidence for the optimal-inbreeding hypothesis is found in
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breeding-site philopatry in males than females among birds, and the opposite bias among mammals. Many possible explanations for this sex bias have been posited, with the earliest accepted hypothesis attributing the bias to intrasexual competition, and territory choice. The most widely accepted hypothesis is that proposed by
Greenwood (1980). Among birds, males invest highly in protecting resources – a territory – against other males. Over consecutive seasons, a male that returns to the same territory has higher fitness than one that is not philopatric. Females are free to disperse, and assess males. Conversely, in mammals, the predominant mating system is one of
262:, of which cooperative breeding is a form, explains how individual offspring provide care for further offspring produced by their relatives. Animals that are philopatric to birthsites have increased association with family members, and, in situations where inclusive fitness is increased through cooperative breeding, may evolve such behaviour, as it will incur evolutionary benefits to families that do. Inclusive fitness is the sum of all direct and indirect fitness, where direct fitness is defined as the amount of fitness gained through producing offspring. Indirect fitness is defined as the amount of fitness gained through aiding related individuals offspring.
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whether due to a precisely adapted genome or not – mean that individuals that return to a site are more familiar with it, and may have more success in either defending it, or locating mates. This hypothesis does not justify whether philopatry is due to an innate behaviour in each individual, or to learning; however it has been shown that, in most species, older individuals show higher site fidelity. Neither of these hypotheses is as widely accepted as the optimal-inbreeding or dispersal hypotheses, but their existence indicates that the evolutionary causes of natal philopatry have still not been conclusively demonstrated.
83:) construct a large mound of vegetation and soil or sand to lay their eggs in. Megapodes often reuse the same mound for many years, only abandoning it when it is damaged beyond repair, or due to disturbance. Nest fidelity is highly beneficial as reproducing is time and energy consuming (malleefowl will tend a mound for five to six months per year). In colonial seabirds, it has been shown that nest fidelity depends on multi-scale information, including the breeding success of the focal breeding pair, the average breeding success of the rest of the colony, and the interaction of these two scales.
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not return to a location in following years if a breeding attempt is unsuccessful. The evolutionary benefits of such learning are evident: individuals that risk searching for a better site will not have lower fitness than those that persist with a poor site. Philopatry is not homogenous within a species, with individuals far more likely to exhibit philopatry if the breeding habitat is isolated. Similarly, non-migratory populations are more likely to be philopatric that those that migrate.
143:. Outbreeding depression involves reduced fitness as a result of random mating, which occurs due to the breakdown of coadapted gene complexes by combining allele that do not cross well with those from a different subpopulation. However, it is important to note that outbreeding depression becomes more detrimental the longer (temporally) that subpopulations have been separated, and that this does hypothesis does not provide an initial mechanism for the evolution of natal philopatry.
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philopatric. This hypothesis also applies to natal philopatry, but is primarily concerned with breeding-site fidelity. A more recent hypothesis builds on
Greenwood’s findings, suggesting that parental influence may play a large role. Because birds lay eggs, adult females are at risk of being cuckolded by their daughters, and thus would drive them out. On the other hand, young male mammals pose a threat to their dominant father, and so are driven to disperse while young.
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and more likely to have difficulty finding resources and food. Therefore, living in groups increases a species' chances of survival, which correlates to finding resources and reproducing. Again, depending on the species, returning to their birthplace where that particular species occupies that territory is the more favorable option. The birthplaces for these animals serve as a territory for them to return for feeding and refuge, like fish from a
188:. Animals that spend much of their time at sea, but which return to land to breed exhibit high levels of natal philopatry and subsequent genetic drift between populations. Many species of albatross do not breed until 6–16 years of age. Between leaving their birth island, and their return, they fly hundreds of thousands of kilometres. High levels of natal philopatry have been demonstrated via mark-recapture data. For example, more than 99% of
177:. Such speciation is most evident on islands. For mobile island-breeding animals, finding a new breeding location may be beyond their means. In combination with a small population, as may occur due to recent colonisation, or simply restricted space, genetic drift can occur on shorter timescales than is observable in mainland species. The high levels of endemism on islands have been attributed to these factors.
66:, and involves an individual, pair, or colony returning to the same location to breed, year after year . The animal can live in that area and reproduce although animals can reproduce anywhere but it can have a higher lifespan in their birth area. Among animals that are largely sedentary, breeding-site philopatry is common. It is advantageous to reuse a breeding site, as there may be
46:. In an animal behavior study conducted by Paul Greenwood, overall female mammals are more likely to be philopatric, while male mammals are more likely to disperse. Male birds are more likely to be philopatric, while females are more likely to disperse. Philopatry will favor the evolution of cooperative traits because the direction of sex has consequences from the particular
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Small mutational changes in non-nuclear DNA that become fixed in small populations are likely to be the major driver of speciation. That there is minimal structural morphological difference between the genetically distinct populations is evidence for random genetic drift, rather than directional evolution due to natural selective pressure.
228:) making attempts to build nests on a south Atlantic Island, where the species had never been previously recorded, demonstrate that range extension by roaming sub-adult birds is possible. Secondly, there may be sufficient gene exchange as to prevent divergence. For example, isolated (yet geographically close) populations of the
212:, which explains how individuals that start new populations carry the genes of their source population. If only a short (in evolutionary timescales) period of time has passed, insufficient divergence may have occurred. For example, study of mitochondrial DNA microsatellites found no significant difference between colonies of
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Cooperative breeding causes the reproductive success of all sexually mature adults to be skewed towards one mating pair. This means the reproductive fitness of the group is held within a select few breeding members and helpers have little to no reproductive fitness. With this system, breeders gain an
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Speciation through natal philopatry is a self-reinforcing process. Once genetic differences are sufficient, different species may be unable to interbreed to produce viable offspring. As a result, breeding could not occur anywhere except natal island, strengthening philopatry and ultimately leading to
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Some of the known reasons for organisms to be philopatric would be for mating (reproduction), survival, migration, parental care, resources, etc.. In most species of animals, individuals will benefit from living in groups, because depending on the species, individuals are more vulnerable to predation
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Due to the dispersal capabilities of albatross, distance between populations does not appear to be a determining factor in divergence. Actual speciation is likely to occur very slowly, as the selective pressures on the animals are the same for the vast majority of their lives, which is spent at sea.
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commonly refers to the return to the area the animal was born in, or to animals remaining in their natal territory. It is a form of breeding-site philopatry. The debate over the evolutionary causes remains unsettled. The outcomes of natal philopatry may be speciation, and, in cases of non-dispersing
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Breeding fidelity is also well documented among species that migrate or disperse after reaching maturity. Birds, in particular, that disperse as fledglings will take advantage of exceptional navigational skills to return to a previous site. Philopatric individuals exhibit learning behaviour, and do
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Cooperative breeding, like speciation, can become a self-reinforcing process for a species. If the fitness benefits result in higher inclusive fitness of a family than the fitness of a non-cooperative family, the trait will eventually become fixed in the population. Over time, this may lead to the
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A second hypothesis explains the evolution of natal philopatry as a method of reducing the high costs of dispersal among offspring. A review of records of natal philopatry among passerine birds found that migrant species showed significantly less site fidelity than sedentary birds. Among migratory
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Males generally invest little in the raising of offspring, and compete with each other for mates rather than resources. Thus, dispersing can result in reproductive enhancement, as greater access to females is available. On the other hand, the cost of dispersal to females is high, and thus they are
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A number of other hypotheses exist. One such is that philopatry is a method, in migratory species, of ensuring that the sexes interact in breeding areas, and that breeding actually occurs. A second is that philopatry provides a much higher chance of breeding success. Strict habitat requirements –
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is a phenomenon whereby deleterious alleles become fixed more easily within an inbreeding population. Inbreeding depression is demonstrably costly and accepted by most scientists as a greater cost than those of outbreeding depression. Within a species, there has also been found to be variation in
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Costs for helpers include a fitness reduction, increased territory defense, offspring guarding and an increased cost of growth. Benefits for helpers include a reduced chance of predation, increased foraging time, territory inheritance, increased environmental conditions and an inclusive fitness.
204:) was shown to have genetic differences in its microsatellites between three breeding colonies located off the coast of Tasmania. The differences are not currently sufficient to propose identifying the populations as distinct species; however divergence is likely to continue without outbreeding.
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In species that exhibit lifelong monogamous pair bonds, even outside of the breeding season, there is no bias in the sex that is philopatric. However, among polygynous species that disperse (including those that find only a single mate per breeding season), there is a much higher rate of
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The term is sometimes also applied to animals that live in nests but do not remain in them during an unfavorable season (e.g., the winter in the temperate zone, or the dry season in the tropics), and leave to find hiding places nearby to pass the inactive period (common in various
318:); this is not migration in the usual sense, as the location of the hiding place is effectively random and unique (never located or revisited except by accident), though the navigation skills required to relocate the old nest site may be similar to those of migrating animals.
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areas and wintering grounds. Philopatry is generally believed to help maintain the adaptation of a population to a very specific environment (i.e., if a set of genes has evolved in a specific area, individuals that fail to return to that area may do poorly elsewhere, so
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evolution of obligate cooperative breeding, as exhibited by the
Australian mudnesters and Australo-Papuan babblers. Obligate cooperative breeding requires natally philopatric offspring to assist in raising offspring – breeding is unsuccessful without such help.
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For the breeding pair, costs include increased mate guarding and suppression of subordinate mating. Breeders receive benefits as reductions in offspring care and territory maintenance. Their primary benefit is an increased reproductive rate and survival.
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Not all isolated populations will show evidence of genetic drift. Genetic homogeneity can be attributed to one of two explanations, both of which indicate that natal philopatry is not absolute within a species. Firstly, a lack of divergence may be due to
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animals, cooperative breeding. Natal philopatry is the most common form of philopatry in females because it decreases competition for mating and increases the rate of reproduction and a higher survival rate for offspring. Natal philopatry also leads to a
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outside of the individual’s home range, and since the area evidently meets the requirements of breeding. Such advantages are compounded among species that invest heavily in the construction of a nest or associated courtship area. For example, the
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rates of philopatry, with migratory populations exhibiting low levels of philopatry – further suggesting that the ecological cost of dispersal, rather than genetic benefits of either inbreeding or outbreeding, is the driver of natal philopatry.
236:) have been shown to be genetically similar. This evidence has only recently, for the first time, been supported by mark-recapture data, which showed one bird marked on one of the two breeding islands was nesting on the other island.
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Cooperative breeding is a hierarchical social system characterized by a dominant breeding pair surrounded by subordinate helpers. The dominant breeding pair and their helpers experience costs and benefits from using this system.
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population, which is when the population is more genetically related than less related between individuals in a species. This can also lead to inbreeding and a higher rate of natural and sexual selection within a population.
34:, "fatherland", although in recent years the term has been applied to more than just the animal's birthplace. Recent usage refers to animals returning to the same area to breed despite not being born there, and
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Van Bekkum, Margo; Sagar, Paul M; Stahl, Jean-Claude; Chambers, Geoffrey K (2005). "Natal philopatry does not lead to population genetic differentiation in Buller's albatross (Thalassarche bulleri bulleri)".
196:) in a study returned to exactly the same nest in consecutive years. Such site-specificity can lead to speciation, and has also been observed in the earliest stages of this process. The
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Burg, T. M; Croxall, J. P (2001). "Global relationships amongst black-browed and grey-headed albatrosses: Analysis of population structure using mitochondrial DNA and microsatellites".
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species, the cost of dispersal is paid either way. If the optimal-inbreeding hypothesis was correct, the benefits of inbreeding should result in philopatry among all species.
1374:
Lee, Jin-Won; Lee, Yun-Kyoung; Hatchwell, Ben J (2010). "Natal dispersal and philopatry in a group-living but noncooperative passerine bird, the vinous-throated parrotbill".
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The exact causes for the evolution of natal philopatry are unknown. Two major hypotheses have been proposed. Shields (1982) suggested that philopatry was a way of ensuring
1148:
Abbott, Cathryn L; Double, Michael C (2003). "Genetic structure, conservation genetics and evidence of speciation by range expansion in shy and white-capped albatrosses".
379:
Frederick, Peter C; Ogden, John C (1997). "Philopatry and
Nomadism: Contrasting Long-Term Movement Behavior and Population Dynamics of White Ibises and Wood Storks".
556:
Weathers, Wesley W; Seymour, Roger S; Baudinette, Russell V (1993). "Energetics of mound-tending behaviour in the malleefowl, Leipoa ocellata (Megapodiidae)".
2621:
2024:
1332:"Piecing together the global population puzzle of wandering albatrosses: genetic analysis of the Amsterdam albatross Diomedea amsterdamensis"
1331:
2029:
678:
2310:
890:
Liberg, Olof; von
Schantz, Torbjorn (1985). "Sex-Biased Philopatry and Dispersal in Birds and Mammals: The Oedipus Hypothesis".
220:) on the Falkland Islands and Campbell Island, despite the sites being thousands of kilometres apart. Observational evidence of
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is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but
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1582:
Gerlach, G (2002). "Reproductive skew, costs, and benefits of cooperative breeding in female wood mice (Apodemus sylvaticus)".
2229:
450:
Kokko, H; López-Sepulcre, Andrés (2006). "From
Individual Dispersal to Species Ranges: Perspectives for a Changing World".
1113:
Van Ryzin, Margaret T; Fisher, Harvey I (1976). "The Age of Laysan
Albatrosses, Diomedea immutabilis, at First Breeding".
2239:
2126:
1675:
2432:
647:
Weatherhead, P. J; Forbes, M. R. L (1994). "Natal philopatry in passerine birds: Genetic or ecological influences?".
92:
26:, where animals return to their birthplace to breed, may be the most common. The term derives from the Greek roots
677:
Shitikov, Dmitry; Fedotova, Svetlana; Gagieva, Victoria; Fedchuk, Darya; Dubkova, Elena; Vaytina, Tatiana (2012).
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Migrating animals also exhibit philopatry to certain important areas on their route; staging areas, stop-overs,
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Substantial evidence for speciation due to natal philopatry has been gathered in studies of island-nesting
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1845:
956:
Lynch, Michael (1991). "The
Genetic Interpretation of Inbreeding Depression and Outbreeding Depression".
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1938:
2334:
2305:
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This page discusses the evolutionary reasons for philopatry. For the mechanisms of philopatry, see
2566:
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725:
Tryjanowski, Piotr; Goławski, Artur; Kuźniak, Stanisław; Mokwa, Tomasz; Antczak, Marcin (2007).
2586:
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2119:
1877:
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1723:
1668:
819:
229:
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173:. Without genetic exchange, geographically and reproductively isolated populations may undergo
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A major outcome of multi-generational natal philopatry is genetic divergence and, ultimately,
2395:
2300:
148:
999:
Hasler, A; Scholz, A. T; Horrall, R. M (1978). "Olfactory imprinting and homing in salmon".
855:
Pusey, Anne E (1987). "Sex-biased dispersal and inbreeding avoidance in birds and mammals".
679:"Breeding-site fidelity and dispersal in isolated populations of three migratory passerines"
2606:
2571:
2409:
2356:
2263:
1983:
1748:
1708:
1703:
1157:
1070:
1008:
521:
Greenwood, Paul J (1980). "Mating systems, philopatry and dispersal in birds and mammals".
459:
327:
253:
38:
that demonstrate site fidelity: reusing stopovers, staging points, and wintering grounds.
1993:
8:
2616:
2591:
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1161:
1074:
1012:
727:"Disperse or Stay? Exceptionally High Breeding-Site Infidelity in the Red-Backed Shrike
463:
2596:
2457:
2452:
1988:
1780:
1758:
1738:
1635:
1610:
1556:
1523:
1499:
1474:
1435:
1410:
1391:
1312:
1291:
Phalan, Ben; Phillips, Richard A; Double, Michael C (2016). "A White-capped
Albatross,
1273:
1230:
1181:
1130:
1016:
973:
915:
907:
837:
787:
752:
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487:
467:
430:
388:
303:
189:
35:
23:
1387:
770:
Weatherhead, Patrick J; Boak, Karyn A (1986). "Site infidelity in song sparrows".
2258:
2135:
2008:
1948:
1933:
1918:
1903:
1872:
1825:
1100:
Handbook of
Australian, New Zealand and Antarctic Birds. Vol. 1: Ratites to Ducks
810:
Moore, Jim; Ali, Rauf (1984). "Are dispersal and inbreeding avoidance related?".
209:
2601:
2535:
2512:
2346:
2341:
2324:
2317:
2210:
1973:
1928:
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332:
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174:
123:
47:
593:"Nest fidelity is driven by multi-scale information in a long-lived seabird"
471:
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2003:
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1913:
1882:
1835:
1644:
1626:
1565:
1539:
1508:
1490:
1444:
1426:
1295:, at South Georgia: first confirmed record in the south-western Atlantic".
1269:
1226:
1177:
985:
876:
626:
608:
569:
479:
337:
278:
increased reproductive, while helpers gain an increased inclusive fitness.
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Philopatric species that do not migrate may evolve to breed cooperatively.
110:
1524:"First evidence for heritable variation in cooperative breeding behaviour"
2576:
2525:
2366:
2222:
1943:
1908:
1028:
747:
726:
1522:
Charmantier, Anne; Keyser, Amber J; Promislow, Daniel E.L (2007-07-22).
1020:
2361:
1134:
977:
911:
697:
400:
342:
170:
136:
76:
43:
2611:
2251:
1475:"Life histories and the evolution of cooperative breeding in mammals"
365:(11th ed.). New York, NY: J. Wiley & Sons, Inc. p. 432.
181:
1308:
1126:
969:
392:
2540:
2502:
2234:
2176:
1815:
1810:
1785:
1684:
903:
72:
1411:"The evolution of cooperative breeding through group augmentation"
184:. Genetic difference is most often detected in microsatellites in
2161:
1850:
16:
Tendency of animals to return to an area, especially to reproduce
2195:
724:
591:
Robert, A; Paiva, V. H; Bolton, M; Jiguet, F; Bried, J (2014).
676:
2437:
1753:
1203:
2183:
2104:
1860:
1653:
315:
291:
1521:
436:
10.1642/0004-8038(2007)124[1085:parto]2.0.co;2
1611:"Prevalence of different modes of parental care in birds"
1330:
Rains, Derek; Weimerskirch, Henri; Burg, Theresa (2011).
1057:
Witt, Christopher C.; Maliakal-Witt, Satya (2007-06-05).
555:
311:
1615:
Proceedings of the Royal Society B: Biological Sciences
1528:
Proceedings of the Royal Society B: Biological Sciences
1479:
Proceedings of the Royal Society B: Biological Sciences
1415:
Proceedings of the Royal Society B: Biological Sciences
597:
Proceedings of the Royal Society B: Biological Sciences
590:
1329:
75:(large, ground-dwelling birds such as the Australian
1290:
130:
998:
449:
2622:Task allocation and partitioning of social insects
1472:
1408:
1059:"Why are diversity and endemism Linked on islands"
940:. Albany (NY): State University of New York Press.
1056:
889:
646:
360:
2634:
938:Philopatry, inbreeding, and the evolution of sex
302:The level of philopatry varies within migratory
1457:
1373:
1112:
1097:
769:
1458:Davies, N.B.; Krebs, J.R.; West, S.A. (2012).
1409:Kokko, H; Johnstone, R. A; t. h, C.-B (2001).
378:
374:
372:
2120:
2025:Association for the Study of Animal Behaviour
1669:
1199:
1197:
1195:
642:
640:
638:
636:
159:
1147:
1044:The Evolutionary Ecology of Animal Migration
1577:
1575:
1247:
718:
672:
670:
516:
514:
512:
510:
369:
53:
2127:
2113:
2030:International Society for Applied Ethology
1676:
1662:
1466:
1192:
1098:Marchant, S.; Higgins, P.J., eds. (1990).
931:
929:
633:
584:
363:Henderson's Dictionary of Biological Terms
1634:
1555:
1498:
1434:
1106:
1082:
823:
746:
616:
520:
434:
1608:
1602:
1572:
1091:
951:
949:
947:
809:
805:
803:
801:
667:
507:
412:
410:
299:will favor those who exhibit fidelity).
1581:
935:
926:
549:
354:
247:
2635:
1460:An Introduction to Behavioural Ecology
1451:
1369:
1367:
1284:
1141:
992:
883:
416:
361:Lawrence, E.; Henderson, I.F. (1995).
2230:Patterns of self-organization in ants
2108:
1657:
1402:
1102:. Melbourne: Oxford University Press.
1041:
1035:
955:
944:
854:
848:
798:
407:
1462:. West Sussex (UK): Wiley-Blackwell.
1241:
763:
1473:Lukas, D; Clutton-Brock, T (2012).
1364:
285:
102:
13:
2240:symmetry breaking of escaping ants
14:
2664:
1046:. London: Hodder & Stoughton.
857:Trends in Ecology & Evolution
419:"Philopatry: A return to origins"
244:even greater genetic divergence.
131:Evolutionary causes of philopatry
2277:
2089:
2088:
1351:10.1111/j.1600-048X.2010.05295.x
1262:10.1046/j.0962-1083.2001.01392.x
1219:10.1111/j.1365-294x.2005.02776.x
1170:10.1046/j.1365-294x.2003.01980.x
1084:10.1111/j.0906-7590.2007.04837.x
1515:
1323:
1050:
502:Sociobiology: The New Synthesis
93:matrilineal social organisation
1734:Bee learning and communication
494:
443:
1:
1388:10.1016/j.anbehav.2010.01.015
834:10.1016/s0003-3472(84)80328-0
784:10.1016/s0003-3472(86)80201-9
535:10.1016/s0003-3472(80)80103-5
348:
164:
2201:Mixed-species foraging flock
2152:Agent-based model in biology
2134:
1683:
869:10.1016/0169-5347(87)90081-4
7:
2448:Particle swarm optimization
321:
10:
2669:
2157:Collective animal behavior
251:
160:Consequences of philopatry
58:One type of philopatry is
2549:
2511:
2466:
2418:
2286:
2275:
2142:
2084:
2038:
2017:
1896:
1791:Evolutionary neuroscience
1691:
1297:Emu - Austral Ornithology
2486:Self-propelled particles
1744:Behavioral endocrinology
1339:Journal of Avian Biology
54:Breeding-site philopatry
2567:Collective intelligence
2433:Ant colony optimization
1939:Irenäus Eibl-Eibesfeldt
1719:Animal sexual behaviour
1596:10.1093/beheco/13.3.408
892:The American Naturalist
472:10.1126/science.1128566
417:Pearce, John M (2007).
194:Phoebastria immutabilis
68:territorial competition
30:, "liking, loving" and
2587:Microbial intelligence
2247:Shoaling and schooling
1878:Tool use by non-humans
1831:Philosophical ethology
1776:Comparative psychology
1724:Animal welfare science
1627:10.1098/rspb.2005.3458
1540:10.1098/rspb.2007.0012
1491:10.1098/rspb.2012.1433
1427:10.1098/rspb.2000.1349
936:Shields, W.M. (1982).
661:10.1093/beheco/5.4.426
609:10.1098/rspb.2014.1692
570:10.1006/anbe.1993.1038
222:white-capped albatross
214:black-browed albatross
141:outbreeding depression
64:breeding-site fidelity
149:Inbreeding depression
2653:Evolutionary biology
2607:Spatial organization
2572:Decentralised system
2410:Sea turtle migration
2264:Swarming (honey bee)
1984:William Homan Thorpe
1749:Behavioural genetics
1709:Animal consciousness
1704:Animal communication
1609:Cockburn, A (2006).
1293:Thalassarche steadi
1042:Baker, R.R. (1978).
748:10.5253/078.095.0214
328:Cooperative breeding
254:Cooperative breeding
248:Cooperative breeding
2648:Population genetics
2582:Group size measures
2144:Biological swarming
1739:Behavioural ecology
1534:(1619): 1757–1761.
1162:2003MolEc..12.2953A
1075:2007Ecogr..30..331W
1013:1978AmSci..66..347H
464:2006Sci...313..789K
381:Colonial Waterbirds
202:Thalassarche cauta
60:breeding philopatry
2597:Predator satiation
2458:Swarm (simulation)
2453:Swarm intelligence
2428:Agent-based models
2259:Swarming behaviour
2068:Behavioral Ecology
1989:Nikolaas Tinbergen
1781:Emotion in animals
1759:Cognitive ethology
1584:Behavioral Ecology
1001:American Scientist
698:10.51812/of.133792
649:Behavioral Ecology
603:(1793): 20141692.
500:Edward O. Wilson,
234:T. bulleri bulleri
230:Buller’s albatross
2630:
2629:
2617:Military swarming
2562:Animal navigation
2481:Collective motion
2468:Collective motion
2335:reverse migration
2269:Swarming motility
2102:
2101:
1994:Jakob von Uexküll
1764:Comfort behaviour
1621:(1592): 1375–83.
1485:(1744): 4065–70.
1250:Molecular Ecology
1207:Molecular Ecology
1150:Molecular Ecology
297:natural selection
186:mitochondrial DNA
36:migratory species
2660:
2443:Crowd simulation
2420:Swarm algorithms
2391:Insect migration
2296:Animal migration
2288:Animal migration
2281:
2206:Mobbing behavior
2129:
2122:
2115:
2106:
2105:
2092:
2091:
2054:Animal Cognition
2047:Animal Behaviour
1999:Wolfgang Wickler
1699:Animal cognition
1678:
1671:
1664:
1655:
1654:
1649:
1648:
1638:
1606:
1600:
1599:
1579:
1570:
1569:
1559:
1519:
1513:
1512:
1502:
1470:
1464:
1463:
1455:
1449:
1448:
1438:
1421:(1463): 187–96.
1406:
1400:
1399:
1376:Animal Behaviour
1371:
1362:
1361:
1359:
1357:
1336:
1327:
1321:
1320:
1288:
1282:
1281:
1245:
1239:
1238:
1201:
1190:
1189:
1145:
1139:
1138:
1110:
1104:
1103:
1095:
1089:
1088:
1086:
1054:
1048:
1047:
1039:
1033:
1032:
996:
990:
989:
953:
942:
941:
933:
924:
923:
887:
881:
880:
852:
846:
845:
827:
812:Animal Behaviour
807:
796:
795:
778:(5): 1299–1310.
772:Animal Behaviour
767:
761:
760:
750:
722:
716:
715:
713:
712:
706:
700:. Archived from
683:
674:
665:
664:
644:
631:
630:
620:
588:
582:
581:
558:Animal Behaviour
553:
547:
546:
523:Animal Behaviour
518:
505:
498:
492:
491:
458:(5788): 789–91.
447:
441:
440:
438:
414:
405:
404:
376:
367:
366:
358:
286:Other variations
190:Laysan albatross
119:Natal philopatry
103:Natal philopatry
24:natal philopatry
2668:
2667:
2663:
2662:
2661:
2659:
2658:
2657:
2633:
2632:
2631:
2626:
2545:
2507:
2462:
2414:
2282:
2273:
2138:
2133:
2103:
2098:
2080:
2034:
2013:
2009:Solly Zuckerman
1949:Karl von Frisch
1934:Richard Dawkins
1919:John B. Calhoun
1904:Patrick Bateson
1892:
1826:Pain in animals
1687:
1682:
1652:
1607:
1603:
1580:
1573:
1520:
1516:
1471:
1467:
1456:
1452:
1407:
1403:
1372:
1365:
1355:
1353:
1334:
1328:
1324:
1309:10.1071/MU03057
1289:
1285:
1256:(11): 2647–60.
1246:
1242:
1202:
1193:
1156:(11): 2953–62.
1146:
1142:
1127:10.2307/1366911
1111:
1107:
1096:
1092:
1055:
1051:
1040:
1036:
997:
993:
970:10.2307/2409915
954:
945:
934:
927:
888:
884:
853:
849:
825:10.1.1.584.5226
808:
799:
768:
764:
729:Lanius collurio
723:
719:
710:
708:
704:
681:
675:
668:
645:
634:
589:
585:
554:
550:
519:
508:
499:
495:
448:
444:
415:
408:
393:10.2307/1521699
377:
370:
359:
355:
351:
324:
288:
256:
250:
210:founder effects
167:
162:
133:
105:
81:Leipoa ocellata
56:
17:
12:
11:
5:
2666:
2656:
2655:
2650:
2645:
2628:
2627:
2625:
2624:
2619:
2614:
2609:
2604:
2602:Quorum sensing
2599:
2594:
2589:
2584:
2579:
2574:
2569:
2564:
2559:
2553:
2551:
2550:Related topics
2547:
2546:
2544:
2543:
2538:
2536:Swarm robotics
2533:
2528:
2523:
2517:
2515:
2513:Swarm robotics
2509:
2508:
2506:
2505:
2500:
2495:
2494:
2493:
2483:
2478:
2472:
2470:
2464:
2463:
2461:
2460:
2455:
2450:
2445:
2440:
2435:
2430:
2424:
2422:
2416:
2415:
2413:
2412:
2407:
2406:
2405:
2404:
2403:
2388:
2387:
2386:
2381:
2371:
2370:
2369:
2364:
2359:
2354:
2347:Fish migration
2344:
2342:Cell migration
2339:
2338:
2337:
2332:
2325:Bird migration
2322:
2321:
2320:
2318:coded wire tag
2315:
2314:
2313:
2303:
2292:
2290:
2284:
2283:
2276:
2274:
2272:
2271:
2266:
2261:
2256:
2255:
2254:
2244:
2243:
2242:
2237:
2227:
2226:
2225:
2215:
2214:
2213:
2211:feeding frenzy
2203:
2198:
2193:
2192:
2191:
2181:
2180:
2179:
2174:
2164:
2159:
2154:
2148:
2146:
2140:
2139:
2132:
2131:
2124:
2117:
2109:
2100:
2099:
2097:
2096:
2085:
2082:
2081:
2079:
2078:
2071:
2064:
2061:Animal Welfare
2057:
2050:
2042:
2040:
2036:
2035:
2033:
2032:
2027:
2021:
2019:
2015:
2014:
2012:
2011:
2006:
2001:
1996:
1991:
1986:
1981:
1976:
1974:Desmond Morris
1971:
1966:
1961:
1956:
1951:
1946:
1941:
1936:
1931:
1929:Marian Dawkins
1926:
1924:Charles Darwin
1921:
1916:
1911:
1906:
1900:
1898:
1894:
1893:
1891:
1890:
1885:
1880:
1875:
1870:
1869:
1868:
1863:
1858:
1853:
1843:
1838:
1833:
1828:
1823:
1818:
1813:
1808:
1806:Human ethology
1803:
1798:
1793:
1788:
1783:
1778:
1773:
1772:
1771:
1761:
1756:
1751:
1746:
1741:
1736:
1731:
1726:
1721:
1716:
1714:Animal culture
1711:
1706:
1701:
1695:
1693:
1689:
1688:
1681:
1680:
1673:
1666:
1658:
1651:
1650:
1601:
1571:
1514:
1465:
1450:
1401:
1382:(5): 1017–23.
1363:
1322:
1283:
1240:
1191:
1140:
1105:
1090:
1069:(3): 331–333.
1049:
1034:
991:
943:
925:
904:10.1086/284402
882:
847:
797:
762:
717:
666:
632:
583:
548:
529:(4): 1140–62.
506:
493:
442:
406:
368:
352:
350:
347:
346:
345:
340:
335:
330:
323:
320:
287:
284:
252:Main article:
249:
246:
218:T. melanophrys
166:
163:
161:
158:
132:
129:
124:kin-structured
116:
115:
104:
101:
55:
52:
15:
9:
6:
4:
3:
2:
2665:
2654:
2651:
2649:
2646:
2644:
2641:
2640:
2638:
2623:
2620:
2618:
2615:
2613:
2610:
2608:
2605:
2603:
2600:
2598:
2595:
2593:
2590:
2588:
2585:
2583:
2580:
2578:
2575:
2573:
2570:
2568:
2565:
2563:
2560:
2558:
2555:
2554:
2552:
2548:
2542:
2539:
2537:
2534:
2532:
2529:
2527:
2524:
2522:
2519:
2518:
2516:
2514:
2510:
2504:
2501:
2499:
2496:
2492:
2489:
2488:
2487:
2484:
2482:
2479:
2477:
2476:Active matter
2474:
2473:
2471:
2469:
2465:
2459:
2456:
2454:
2451:
2449:
2446:
2444:
2441:
2439:
2436:
2434:
2431:
2429:
2426:
2425:
2423:
2421:
2417:
2411:
2408:
2402:
2399:
2398:
2397:
2394:
2393:
2392:
2389:
2385:
2382:
2380:
2377:
2376:
2375:
2372:
2368:
2365:
2363:
2360:
2358:
2355:
2353:
2352:diel vertical
2350:
2349:
2348:
2345:
2343:
2340:
2336:
2333:
2331:
2328:
2327:
2326:
2323:
2319:
2316:
2312:
2309:
2308:
2307:
2304:
2302:
2299:
2298:
2297:
2294:
2293:
2291:
2289:
2285:
2280:
2270:
2267:
2265:
2262:
2260:
2257:
2253:
2250:
2249:
2248:
2245:
2241:
2238:
2236:
2233:
2232:
2231:
2228:
2224:
2221:
2220:
2219:
2216:
2212:
2209:
2208:
2207:
2204:
2202:
2199:
2197:
2194:
2190:
2189:herd behavior
2187:
2186:
2185:
2182:
2178:
2175:
2173:
2170:
2169:
2168:
2165:
2163:
2160:
2158:
2155:
2153:
2150:
2149:
2147:
2145:
2141:
2137:
2130:
2125:
2123:
2118:
2116:
2111:
2110:
2107:
2095:
2087:
2086:
2083:
2077:
2076:
2072:
2070:
2069:
2065:
2063:
2062:
2058:
2056:
2055:
2051:
2049:
2048:
2044:
2043:
2041:
2037:
2031:
2028:
2026:
2023:
2022:
2020:
2016:
2010:
2007:
2005:
2002:
2000:
1997:
1995:
1992:
1990:
1987:
1985:
1982:
1980:
1979:Thomas Sebeok
1977:
1975:
1972:
1970:
1969:Konrad Lorenz
1967:
1965:
1964:Julian Huxley
1962:
1960:
1959:Heini Hediger
1957:
1955:
1952:
1950:
1947:
1945:
1942:
1940:
1937:
1935:
1932:
1930:
1927:
1925:
1922:
1920:
1917:
1915:
1912:
1910:
1907:
1905:
1902:
1901:
1899:
1895:
1889:
1888:Zoomusicology
1886:
1884:
1881:
1879:
1876:
1874:
1871:
1867:
1864:
1862:
1859:
1857:
1854:
1852:
1849:
1848:
1847:
1844:
1842:
1839:
1837:
1834:
1832:
1829:
1827:
1824:
1822:
1821:Neuroethology
1819:
1817:
1814:
1812:
1809:
1807:
1804:
1802:
1799:
1797:
1794:
1792:
1789:
1787:
1784:
1782:
1779:
1777:
1774:
1770:
1767:
1766:
1765:
1762:
1760:
1757:
1755:
1752:
1750:
1747:
1745:
1742:
1740:
1737:
1735:
1732:
1730:
1729:Anthrozoology
1727:
1725:
1722:
1720:
1717:
1715:
1712:
1710:
1707:
1705:
1702:
1700:
1697:
1696:
1694:
1690:
1686:
1679:
1674:
1672:
1667:
1665:
1660:
1659:
1656:
1646:
1642:
1637:
1632:
1628:
1624:
1620:
1616:
1612:
1605:
1597:
1593:
1590:(3): 408–18.
1589:
1585:
1578:
1576:
1567:
1563:
1558:
1553:
1549:
1545:
1541:
1537:
1533:
1529:
1525:
1518:
1510:
1506:
1501:
1496:
1492:
1488:
1484:
1480:
1476:
1469:
1461:
1454:
1446:
1442:
1437:
1432:
1428:
1424:
1420:
1416:
1412:
1405:
1397:
1393:
1389:
1385:
1381:
1377:
1370:
1368:
1352:
1348:
1344:
1340:
1333:
1326:
1318:
1314:
1310:
1306:
1303:(4): 359–61.
1302:
1298:
1294:
1287:
1279:
1275:
1271:
1267:
1263:
1259:
1255:
1251:
1244:
1236:
1232:
1228:
1224:
1220:
1216:
1212:
1208:
1200:
1198:
1196:
1187:
1183:
1179:
1175:
1171:
1167:
1163:
1159:
1155:
1151:
1144:
1136:
1132:
1128:
1124:
1120:
1116:
1109:
1101:
1094:
1085:
1080:
1076:
1072:
1068:
1064:
1060:
1053:
1045:
1038:
1030:
1026:
1022:
1018:
1014:
1010:
1007:(3): 347–55.
1006:
1002:
995:
987:
983:
979:
975:
971:
967:
963:
959:
952:
950:
948:
939:
932:
930:
921:
917:
913:
909:
905:
901:
898:(1): 129–35.
897:
893:
886:
878:
874:
870:
866:
863:(10): 295–9.
862:
858:
851:
843:
839:
835:
831:
826:
821:
818:(1): 94–112.
817:
813:
806:
804:
802:
793:
789:
785:
781:
777:
773:
766:
758:
754:
749:
744:
741:(2): 316–20.
740:
736:
732:
730:
721:
707:on 2017-08-09
703:
699:
695:
691:
687:
686:Ornis Fennica
680:
673:
671:
662:
658:
655:(4): 426–33.
654:
650:
643:
641:
639:
637:
628:
624:
619:
614:
610:
606:
602:
598:
594:
587:
579:
575:
571:
567:
564:(2): 333–41.
563:
559:
552:
544:
540:
536:
532:
528:
524:
517:
515:
513:
511:
503:
497:
489:
485:
481:
477:
473:
469:
465:
461:
457:
453:
446:
437:
432:
429:(3): 1085–7.
428:
424:
420:
413:
411:
402:
398:
394:
390:
387:(2): 316–23.
386:
382:
375:
373:
364:
357:
353:
344:
341:
339:
336:
334:
333:Kin selection
331:
329:
326:
325:
319:
317:
313:
307:
306:and species.
305:
300:
298:
293:
283:
279:
275:
271:
267:
263:
261:
260:Kin selection
255:
245:
241:
237:
235:
231:
227:
223:
219:
215:
211:
205:
203:
199:
198:shy albatross
195:
191:
187:
183:
178:
176:
175:genetic drift
172:
157:
153:
150:
144:
142:
138:
128:
125:
120:
114:
112:
107:
106:
100:
96:
94:
88:
84:
82:
78:
74:
69:
65:
61:
51:
49:
48:mating system
45:
39:
37:
33:
29:
25:
21:
2557:Allee effect
2531:Nanorobotics
2521:Ant robotics
2498:Vicsek model
2383:
2073:
2066:
2059:
2052:
2045:
2004:E. O. Wilson
1954:Jane Goodall
1914:Donald Broom
1883:Zoosemiotics
1836:Sociobiology
1618:
1614:
1604:
1587:
1583:
1531:
1527:
1517:
1482:
1478:
1468:
1459:
1453:
1418:
1414:
1404:
1379:
1375:
1354:. Retrieved
1342:
1338:
1325:
1300:
1296:
1292:
1286:
1253:
1249:
1243:
1210:
1206:
1153:
1149:
1143:
1118:
1114:
1108:
1099:
1093:
1066:
1062:
1052:
1043:
1037:
1004:
1000:
994:
964:(3): 622–9.
961:
957:
937:
895:
891:
885:
860:
856:
850:
815:
811:
775:
771:
765:
738:
734:
728:
720:
709:. Retrieved
702:the original
689:
685:
652:
648:
600:
596:
586:
561:
557:
551:
526:
522:
501:
496:
455:
451:
445:
426:
422:
384:
380:
362:
356:
338:Natal homing
308:
301:
289:
280:
276:
272:
268:
264:
257:
242:
238:
233:
225:
217:
206:
201:
193:
179:
168:
154:
145:
134:
118:
117:
111:Natal homing
108:
97:
89:
85:
80:
63:
59:
57:
40:
31:
27:
19:
18:
2577:Eusociality
2526:Microbotics
2396:butterflies
2367:sardine run
2301:altitudinal
2223:pack hunter
1944:Dian Fossey
1909:Marc Bekoff
1897:Ethologists
1213:(1): 73–9.
2637:Categories
2491:clustering
2384:philopatry
2362:salmon run
2357:Lessepsian
1846:Structures
1841:Stereotypy
1121:(1): 1–9.
1115:The Condor
711:2017-12-08
349:References
343:Salmon run
226:T. steadi
171:speciation
165:Speciation
137:inbreeding
77:malleefowl
44:coral reef
20:Philopatry
2612:Stigmergy
2592:Mutualism
2252:bait ball
2075:Behaviour
2018:Societies
1856:Honeycomb
1548:0962-8452
1345:: 69–79.
1063:Ecography
958:Evolution
820:CiteSeerX
692:: 53–62.
182:albatross
73:megapodes
2643:Ethology
2541:Symbrion
2503:BIO-LGCA
2306:tracking
2235:ant mill
2177:sort sol
2172:flocking
2136:Swarming
2094:Category
2039:Journals
1866:Instinct
1816:Learning
1811:Instinct
1786:Ethogram
1769:Grooming
1692:Branches
1685:Ethology
1645:16777726
1566:17490945
1509:22874752
1445:11209890
1396:53152484
1356:17 March
1317:83493045
1270:11883879
1235:27492523
1227:16367831
1186:22369696
1178:14629376
1021:27848646
986:28568822
920:84154558
877:21227869
792:53146722
757:85395481
627:25209940
578:53154413
543:53178299
480:16902127
322:See also
304:families
2401:monarch
2330:flyways
2311:history
2162:Droving
1796:Feeding
1636:1560291
1557:2493572
1500:3427589
1436:1088590
1278:3026130
1158:Bibcode
1135:1366911
1071:Bibcode
1009:Bibcode
978:2409915
912:2461568
842:7674864
618:4173691
488:9523058
460:Bibcode
452:Science
423:The Auk
401:1521699
292:molting
2374:Homing
2196:Locust
1643:
1633:
1564:
1554:
1546:
1507:
1497:
1443:
1433:
1394:
1315:
1276:
1268:
1233:
1225:
1184:
1176:
1133:
1029:677550
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1019:
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910:
875:
840:
822:
790:
755:
625:
615:
576:
541:
504:, 1975
486:
478:
399:
2438:Boids
2379:natal
2167:Flock
1873:Swarm
1801:Hover
1754:Breed
1392:S2CID
1335:(PDF)
1313:S2CID
1274:S2CID
1231:S2CID
1182:S2CID
1131:JSTOR
1017:JSTOR
974:JSTOR
916:S2CID
908:JSTOR
838:S2CID
788:S2CID
753:S2CID
735:Ardea
705:(PDF)
682:(PDF)
574:S2CID
539:S2CID
484:S2CID
397:JSTOR
316:wasps
62:, or
32:patra
28:philo
2218:Pack
2184:Herd
1861:Nest
1851:Hive
1641:PMID
1562:PMID
1544:ISSN
1505:PMID
1441:PMID
1358:2021
1266:PMID
1223:PMID
1174:PMID
1025:PMID
982:PMID
873:PMID
623:PMID
476:PMID
314:and
312:bees
1631:PMC
1623:doi
1619:273
1592:doi
1552:PMC
1536:doi
1532:274
1495:PMC
1487:doi
1483:279
1431:PMC
1423:doi
1419:268
1384:doi
1347:doi
1305:doi
1301:104
1258:doi
1215:doi
1166:doi
1123:doi
1079:doi
966:doi
900:doi
896:126
865:doi
830:doi
780:doi
743:doi
694:doi
657:doi
613:PMC
605:doi
601:281
566:doi
531:doi
468:doi
456:313
431:doi
427:124
389:doi
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