150:, 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
273:, 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.
94:) 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.
154:. 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
199:. 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
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188:. 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.
77:, 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
57:. 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.
239:) 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
223:, 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.
215:) 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
329:); 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.
247:) 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.
45:, "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)".
207:) 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.
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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
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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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37:, where animals return to their birthplace to breed, may be the most common. The term derives from the Greek roots
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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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This page discusses the evolutionary reasons for philopatry. For the mechanisms of philopatry, see
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A major outcome of multi-generational natal philopatry is genetic divergence and, ultimately,
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that demonstrate site fidelity: reusing stopovers, staging points, and wintering grounds.
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increased reproductive, while helpers gain an increased inclusive fitness.
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Philopatric species that do not migrate may evolve to breed cooperatively.
1535:"First evidence for heritable variation in cooperative breeding behaviour"
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Tendency of animals to return to an area, especially to reproduce
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2618:Spatial organization
2583:Decentralised system
2421:Sea turtle migration
2275:Swarming (honey bee)
1995:William Homan Thorpe
1760:Behavioural genetics
1720:Animal consciousness
1715:Animal communication
1620:Cockburn, A (2006).
1304:Thalassarche steadi
1053:Baker, R.R. (1978).
759:10.5253/078.095.0214
339:Cooperative breeding
265:Cooperative breeding
259:Cooperative breeding
2659:Population genetics
2593:Group size measures
2155:Biological swarming
1750:Behavioural ecology
1545:(1619): 1757–1761.
1173:2003MolEc..12.2953A
1086:2007Ecogr..30..331W
1024:1978AmSci..66..347H
475:2006Sci...313..789K
392:Colonial Waterbirds
213:Thalassarche cauta
71:breeding philopatry
2608:Predator satiation
2469:Swarm (simulation)
2464:Swarm intelligence
2439:Agent-based models
2270:Swarming behaviour
2079:Behavioral Ecology
2000:Nikolaas Tinbergen
1792:Emotion in animals
1770:Cognitive ethology
1595:Behavioral Ecology
1012:American Scientist
709:10.51812/of.133792
660:Behavioral Ecology
614:(1793): 20141692.
511:Edward O. Wilson,
245:T. bulleri bulleri
241:Buller’s albatross
2641:
2640:
2628:Military swarming
2573:Animal navigation
2492:Collective motion
2479:Collective motion
2346:reverse migration
2280:Swarming motility
2113:
2112:
2005:Jakob von Uexküll
1775:Comfort behaviour
1632:(1592): 1375–83.
1496:(1744): 4065–70.
1261:Molecular Ecology
1218:Molecular Ecology
1161:Molecular Ecology
308:natural selection
197:mitochondrial DNA
47:migratory species
16:(Redirected from
2671:
2454:Crowd simulation
2431:Swarm algorithms
2402:Insect migration
2307:Animal migration
2299:Animal migration
2292:
2217:Mobbing behavior
2140:
2133:
2126:
2117:
2103:
2102:
2065:Animal Cognition
2058:Animal Behaviour
2010:Wolfgang Wickler
1710:Animal cognition
1689:
1682:
1675:
1666:
1660:
1659:
1649:
1617:
1611:
1610:
1590:
1581:
1580:
1570:
1530:
1524:
1523:
1513:
1481:
1475:
1474:
1466:
1460:
1459:
1449:
1432:(1463): 187–96.
1417:
1411:
1410:
1387:Animal Behaviour
1382:
1373:
1372:
1370:
1368:
1347:
1338:
1332:
1331:
1299:
1293:
1292:
1256:
1250:
1249:
1212:
1201:
1200:
1156:
1150:
1149:
1121:
1115:
1114:
1106:
1100:
1099:
1097:
1065:
1059:
1058:
1050:
1044:
1043:
1007:
1001:
1000:
964:
953:
952:
944:
935:
934:
898:
892:
891:
863:
857:
856:
838:
823:Animal Behaviour
818:
807:
806:
789:(5): 1299–1310.
783:Animal Behaviour
778:
772:
771:
761:
733:
727:
726:
724:
723:
717:
711:. Archived from
694:
685:
676:
675:
655:
642:
641:
631:
599:
593:
592:
569:Animal Behaviour
564:
558:
557:
534:Animal Behaviour
529:
516:
509:
503:
502:
469:(5788): 789–91.
458:
452:
451:
449:
425:
416:
415:
387:
378:
377:
369:
297:Other variations
201:Laysan albatross
130:Natal philopatry
114:Natal philopatry
35:natal philopatry
21:
2679:
2678:
2674:
2673:
2672:
2670:
2669:
2668:
2644:
2643:
2642:
2637:
2556:
2518:
2473:
2425:
2293:
2284:
2149:
2144:
2114:
2109:
2091:
2045:
2024:
2020:Solly Zuckerman
1960:Karl von Frisch
1945:Richard Dawkins
1930:John B. Calhoun
1915:Patrick Bateson
1903:
1837:Pain in animals
1698:
1693:
1663:
1619:
1618:
1614:
1592:
1591:
1584:
1532:
1531:
1527:
1483:
1482:
1478:
1468:
1467:
1463:
1419:
1418:
1414:
1384:
1383:
1376:
1366:
1364:
1345:
1340:
1339:
1335:
1320:10.1071/MU03057
1301:
1300:
1296:
1267:(11): 2647–60.
1258:
1257:
1253:
1214:
1213:
1204:
1167:(11): 2953–62.
1158:
1157:
1153:
1138:10.2307/1366911
1123:
1122:
1118:
1108:
1107:
1103:
1067:
1066:
1062:
1052:
1051:
1047:
1009:
1008:
1004:
981:10.2307/2409915
966:
965:
956:
946:
945:
938:
900:
899:
895:
865:
864:
860:
836:10.1.1.584.5226
820:
819:
810:
780:
779:
775:
740:Lanius collurio
735:
734:
730:
721:
719:
715:
692:
687:
686:
679:
657:
656:
645:
601:
600:
596:
566:
565:
561:
531:
530:
519:
510:
506:
460:
459:
455:
427:
426:
419:
404:10.2307/1521699
389:
388:
381:
371:
370:
366:
362:
335:
299:
267:
261:
221:founder effects
178:
173:
144:
116:
92:Leipoa ocellata
67:
28:
23:
22:
15:
12:
11:
5:
2677:
2675:
2667:
2666:
2661:
2656:
2646:
2645:
2639:
2638:
2636:
2635:
2630:
2625:
2620:
2615:
2613:Quorum sensing
2610:
2605:
2600:
2595:
2590:
2585:
2580:
2575:
2570:
2564:
2562:
2561:Related topics
2558:
2557:
2555:
2554:
2549:
2547:Swarm robotics
2544:
2539:
2534:
2528:
2526:
2524:Swarm robotics
2520:
2519:
2517:
2516:
2511:
2506:
2505:
2504:
2494:
2489:
2483:
2481:
2475:
2474:
2472:
2471:
2466:
2461:
2456:
2451:
2446:
2441:
2435:
2433:
2427:
2426:
2424:
2423:
2418:
2417:
2416:
2415:
2414:
2399:
2398:
2397:
2392:
2382:
2381:
2380:
2375:
2370:
2365:
2358:Fish migration
2355:
2353:Cell migration
2350:
2349:
2348:
2343:
2336:Bird migration
2333:
2332:
2331:
2329:coded wire tag
2326:
2325:
2324:
2314:
2303:
2301:
2295:
2294:
2287:
2285:
2283:
2282:
2277:
2272:
2267:
2266:
2265:
2255:
2254:
2253:
2248:
2238:
2237:
2236:
2226:
2225:
2224:
2222:feeding frenzy
2214:
2209:
2204:
2203:
2202:
2192:
2191:
2190:
2185:
2175:
2170:
2165:
2159:
2157:
2151:
2150:
2145:
2143:
2142:
2135:
2128:
2120:
2111:
2110:
2108:
2107:
2096:
2093:
2092:
2090:
2089:
2082:
2075:
2072:Animal Welfare
2068:
2061:
2053:
2051:
2047:
2046:
2044:
2043:
2038:
2032:
2030:
2026:
2025:
2023:
2022:
2017:
2012:
2007:
2002:
1997:
1992:
1987:
1985:Desmond Morris
1982:
1977:
1972:
1967:
1962:
1957:
1952:
1947:
1942:
1940:Marian Dawkins
1937:
1935:Charles Darwin
1932:
1927:
1922:
1917:
1911:
1909:
1905:
1904:
1902:
1901:
1896:
1891:
1886:
1881:
1880:
1879:
1874:
1869:
1864:
1854:
1849:
1844:
1839:
1834:
1829:
1824:
1819:
1817:Human ethology
1814:
1809:
1804:
1799:
1794:
1789:
1784:
1783:
1782:
1772:
1767:
1762:
1757:
1752:
1747:
1742:
1737:
1732:
1727:
1725:Animal culture
1722:
1717:
1712:
1706:
1704:
1700:
1699:
1694:
1692:
1691:
1684:
1677:
1669:
1662:
1661:
1612:
1582:
1525:
1476:
1461:
1412:
1393:(5): 1017–23.
1374:
1333:
1294:
1251:
1202:
1151:
1116:
1101:
1080:(3): 331–333.
1060:
1045:
1002:
954:
936:
915:10.1086/284402
893:
858:
808:
773:
728:
677:
643:
594:
559:
540:(4): 1140–62.
517:
504:
453:
417:
379:
363:
361:
358:
357:
356:
351:
346:
341:
334:
331:
298:
295:
263:Main article:
260:
257:
229:T. melanophrys
177:
174:
172:
169:
143:
140:
135:kin-structured
127:
126:
115:
112:
66:
63:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2676:
2665:
2662:
2660:
2657:
2655:
2652:
2651:
2649:
2634:
2631:
2629:
2626:
2624:
2621:
2619:
2616:
2614:
2611:
2609:
2606:
2604:
2601:
2599:
2596:
2594:
2591:
2589:
2586:
2584:
2581:
2579:
2576:
2574:
2571:
2569:
2566:
2565:
2563:
2559:
2553:
2550:
2548:
2545:
2543:
2540:
2538:
2535:
2533:
2530:
2529:
2527:
2525:
2521:
2515:
2512:
2510:
2507:
2503:
2500:
2499:
2498:
2495:
2493:
2490:
2488:
2487:Active matter
2485:
2484:
2482:
2480:
2476:
2470:
2467:
2465:
2462:
2460:
2457:
2455:
2452:
2450:
2447:
2445:
2442:
2440:
2437:
2436:
2434:
2432:
2428:
2422:
2419:
2413:
2410:
2409:
2408:
2405:
2404:
2403:
2400:
2396:
2393:
2391:
2388:
2387:
2386:
2383:
2379:
2376:
2374:
2371:
2369:
2366:
2364:
2363:diel vertical
2361:
2360:
2359:
2356:
2354:
2351:
2347:
2344:
2342:
2339:
2338:
2337:
2334:
2330:
2327:
2323:
2320:
2319:
2318:
2315:
2313:
2310:
2309:
2308:
2305:
2304:
2302:
2300:
2296:
2291:
2281:
2278:
2276:
2273:
2271:
2268:
2264:
2261:
2260:
2259:
2256:
2252:
2249:
2247:
2244:
2243:
2242:
2239:
2235:
2232:
2231:
2230:
2227:
2223:
2220:
2219:
2218:
2215:
2213:
2210:
2208:
2205:
2201:
2200:herd behavior
2198:
2197:
2196:
2193:
2189:
2186:
2184:
2181:
2180:
2179:
2176:
2174:
2171:
2169:
2166:
2164:
2161:
2160:
2158:
2156:
2152:
2148:
2141:
2136:
2134:
2129:
2127:
2122:
2121:
2118:
2106:
2098:
2097:
2094:
2088:
2087:
2083:
2081:
2080:
2076:
2074:
2073:
2069:
2067:
2066:
2062:
2060:
2059:
2055:
2054:
2052:
2048:
2042:
2039:
2037:
2034:
2033:
2031:
2027:
2021:
2018:
2016:
2013:
2011:
2008:
2006:
2003:
2001:
1998:
1996:
1993:
1991:
1990:Thomas Sebeok
1988:
1986:
1983:
1981:
1980:Konrad Lorenz
1978:
1976:
1975:Julian Huxley
1973:
1971:
1970:Heini Hediger
1968:
1966:
1963:
1961:
1958:
1956:
1953:
1951:
1948:
1946:
1943:
1941:
1938:
1936:
1933:
1931:
1928:
1926:
1923:
1921:
1918:
1916:
1913:
1912:
1910:
1906:
1900:
1899:Zoomusicology
1897:
1895:
1892:
1890:
1887:
1885:
1882:
1878:
1875:
1873:
1870:
1868:
1865:
1863:
1860:
1859:
1858:
1855:
1853:
1850:
1848:
1845:
1843:
1840:
1838:
1835:
1833:
1832:Neuroethology
1830:
1828:
1825:
1823:
1820:
1818:
1815:
1813:
1810:
1808:
1805:
1803:
1800:
1798:
1795:
1793:
1790:
1788:
1785:
1781:
1778:
1777:
1776:
1773:
1771:
1768:
1766:
1763:
1761:
1758:
1756:
1753:
1751:
1748:
1746:
1743:
1741:
1740:Anthrozoology
1738:
1736:
1733:
1731:
1728:
1726:
1723:
1721:
1718:
1716:
1713:
1711:
1708:
1707:
1705:
1701:
1697:
1690:
1685:
1683:
1678:
1676:
1671:
1670:
1667:
1657:
1653:
1648:
1643:
1639:
1635:
1631:
1627:
1623:
1616:
1613:
1608:
1604:
1601:(3): 408–18.
1600:
1596:
1589:
1587:
1583:
1578:
1574:
1569:
1564:
1560:
1556:
1552:
1548:
1544:
1540:
1536:
1529:
1526:
1521:
1517:
1512:
1507:
1503:
1499:
1495:
1491:
1487:
1480:
1477:
1472:
1465:
1462:
1457:
1453:
1448:
1443:
1439:
1435:
1431:
1427:
1423:
1416:
1413:
1408:
1404:
1400:
1396:
1392:
1388:
1381:
1379:
1375:
1363:
1359:
1355:
1351:
1344:
1337:
1334:
1329:
1325:
1321:
1317:
1314:(4): 359–61.
1313:
1309:
1305:
1298:
1295:
1290:
1286:
1282:
1278:
1274:
1270:
1266:
1262:
1255:
1252:
1247:
1243:
1239:
1235:
1231:
1227:
1223:
1219:
1211:
1209:
1207:
1203:
1198:
1194:
1190:
1186:
1182:
1178:
1174:
1170:
1166:
1162:
1155:
1152:
1147:
1143:
1139:
1135:
1131:
1127:
1120:
1117:
1112:
1105:
1102:
1096:
1091:
1087:
1083:
1079:
1075:
1071:
1064:
1061:
1056:
1049:
1046:
1041:
1037:
1033:
1029:
1025:
1021:
1018:(3): 347–55.
1017:
1013:
1006:
1003:
998:
994:
990:
986:
982:
978:
974:
970:
963:
961:
959:
955:
950:
943:
941:
937:
932:
928:
924:
920:
916:
912:
909:(1): 129–35.
908:
904:
897:
894:
889:
885:
881:
877:
874:(10): 295–9.
873:
869:
862:
859:
854:
850:
846:
842:
837:
832:
829:(1): 94–112.
828:
824:
817:
815:
813:
809:
804:
800:
796:
792:
788:
784:
777:
774:
769:
765:
760:
755:
752:(2): 316–20.
751:
747:
743:
741:
732:
729:
718:on 2017-08-09
714:
710:
706:
702:
698:
697:Ornis Fennica
691:
684:
682:
678:
673:
669:
666:(4): 426–33.
665:
661:
654:
652:
650:
648:
644:
639:
635:
630:
625:
621:
617:
613:
609:
605:
598:
595:
590:
586:
582:
578:
575:(2): 333–41.
574:
570:
563:
560:
555:
551:
547:
543:
539:
535:
528:
526:
524:
522:
518:
514:
508:
505:
500:
496:
492:
488:
484:
480:
476:
472:
468:
464:
457:
454:
448:
443:
440:(3): 1085–7.
439:
435:
431:
424:
422:
418:
413:
409:
405:
401:
398:(2): 316–23.
397:
393:
386:
384:
380:
375:
368:
365:
359:
355:
352:
350:
347:
345:
344:Kin selection
342:
340:
337:
336:
332:
330:
328:
324:
318:
317:and species.
316:
311:
309:
304:
296:
294:
290:
286:
282:
278:
274:
272:
271:Kin selection
266:
258:
256:
252:
248:
246:
242:
238:
234:
230:
226:
222:
216:
214:
210:
209:shy albatross
206:
202:
198:
194:
189:
187:
186:genetic drift
183:
175:
170:
168:
164:
161:
155:
153:
149:
141:
139:
136:
131:
125:
123:
118:
117:
113:
111:
107:
105:
99:
95:
93:
89:
85:
80:
76:
72:
64:
62:
60:
59:mating system
56:
50:
48:
44:
40:
36:
32:
19:
18:Site fidelity
2568:Allee effect
2542:Nanorobotics
2532:Ant robotics
2509:Vicsek model
2394:
2084:
2077:
2070:
2063:
2056:
2015:E. O. Wilson
1965:Jane Goodall
1925:Donald Broom
1894:Zoosemiotics
1847:Sociobiology
1629:
1625:
1615:
1598:
1594:
1542:
1538:
1528:
1493:
1489:
1479:
1470:
1464:
1429:
1425:
1415:
1390:
1386:
1365:. Retrieved
1353:
1349:
1336:
1311:
1307:
1303:
1297:
1264:
1260:
1254:
1221:
1217:
1164:
1160:
1154:
1129:
1125:
1119:
1110:
1104:
1077:
1073:
1063:
1054:
1048:
1015:
1011:
1005:
975:(3): 622–9.
972:
968:
948:
906:
902:
896:
871:
867:
861:
826:
822:
786:
782:
776:
749:
745:
739:
731:
720:. Retrieved
713:the original
700:
696:
663:
659:
611:
607:
597:
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568:
562:
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533:
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507:
466:
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367:
349:Natal homing
319:
312:
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291:
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122:Natal homing
119:
108:
100:
96:
91:
74:
70:
68:
51:
42:
38:
30:
29:
2588:Eusociality
2537:Microbotics
2407:butterflies
2378:sardine run
2312:altitudinal
2234:pack hunter
1955:Dian Fossey
1920:Marc Bekoff
1908:Ethologists
1224:(1): 73–9.
2648:Categories
2502:clustering
2395:philopatry
2373:salmon run
2368:Lessepsian
1857:Structures
1852:Stereotypy
1132:(1): 1–9.
1126:The Condor
722:2017-12-08
360:References
354:Salmon run
237:T. steadi
182:speciation
176:Speciation
148:inbreeding
88:malleefowl
55:coral reef
31:Philopatry
2623:Stigmergy
2603:Mutualism
2263:bait ball
2086:Behaviour
2029:Societies
1867:Honeycomb
1559:0962-8452
1356:: 69–79.
1074:Ecography
969:Evolution
831:CiteSeerX
703:: 53–62.
193:albatross
84:megapodes
2654:Ethology
2552:Symbrion
2514:BIO-LGCA
2317:tracking
2246:ant mill
2188:sort sol
2183:flocking
2147:Swarming
2105:Category
2050:Journals
1877:Instinct
1827:Learning
1822:Instinct
1797:Ethogram
1780:Grooming
1703:Branches
1696:Ethology
1656:16777726
1577:17490945
1520:22874752
1456:11209890
1407:53152484
1367:17 March
1328:83493045
1281:11883879
1246:27492523
1238:16367831
1197:22369696
1189:14629376
1032:27848646
997:28568822
931:84154558
888:21227869
803:53146722
768:85395481
638:25209940
589:53154413
554:53178299
491:16902127
333:See also
315:families
2412:monarch
2341:flyways
2322:history
2173:Droving
1807:Feeding
1647:1560291
1568:2493572
1511:3427589
1447:1088590
1289:3026130
1169:Bibcode
1146:1366911
1082:Bibcode
1020:Bibcode
989:2409915
923:2461568
853:7674864
629:4173691
499:9523058
471:Bibcode
463:Science
434:The Auk
412:1521699
303:molting
2385:Homing
2207:Locust
1654:
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1518:
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552:
515:, 1975
497:
489:
410:
2449:Boids
2390:natal
2178:Flock
1884:Swarm
1812:Hover
1765:Breed
1403:S2CID
1346:(PDF)
1324:S2CID
1285:S2CID
1242:S2CID
1193:S2CID
1142:JSTOR
1028:JSTOR
985:JSTOR
927:S2CID
919:JSTOR
849:S2CID
799:S2CID
764:S2CID
746:Ardea
716:(PDF)
693:(PDF)
585:S2CID
550:S2CID
495:S2CID
408:JSTOR
327:wasps
73:, or
43:patra
39:philo
2229:Pack
2195:Herd
1872:Nest
1862:Hive
1652:PMID
1573:PMID
1555:ISSN
1516:PMID
1452:PMID
1369:2021
1277:PMID
1234:PMID
1185:PMID
1036:PMID
993:PMID
884:PMID
634:PMID
487:PMID
325:and
323:bees
1642:PMC
1634:doi
1630:273
1603:doi
1563:PMC
1547:doi
1543:274
1506:PMC
1498:doi
1494:279
1442:PMC
1434:doi
1430:268
1395:doi
1358:doi
1316:doi
1312:104
1269:doi
1226:doi
1177:doi
1134:doi
1090:doi
977:doi
911:doi
907:126
876:doi
841:doi
791:doi
754:doi
705:doi
668:doi
624:PMC
616:doi
612:281
577:doi
542:doi
479:doi
467:313
442:doi
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400:doi
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