248:(r=0.5 for 2 mates, and even lower for more). Moreover, males share only 25% of their sisters' genes, and, in cases of equal sex ratios, females are related to their siblings on average by 0.5 which is no better than raising their own offspring. However, despite the shortcomings of the haplodiploidy hypothesis, it is still considered to have some importance. For example, many bees have female-biased sex ratios and/or invest less in or kill males. Analysis has shown that in Hymenoptera, the ancestral female was monogamous in each of the eight independent cases where eusociality evolved. This indicates that the high relatedness between sisters favored the evolution of eusociality during the initial stages on several occasions. This helps explain the abundance of eusocial genera within the order Hymenoptera, including three separate origins within halictid bees alone.
20:
265:- in other words, she mates with only one individual during her entire life - her progeny will be equally related to their siblings and to their own offspring (r=0.5 in both cases - this is an average of sisters and brothers ). Thus, natural selection will favor cooperation in any situation where it is more efficient to raise siblings than offspring, and this could start paving a path towards eusociality. This higher efficiency becomes especially pronounced after group living evolves.
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186:, he described the existence of sterile worker castes in the social insects as "the one special difficulty, which at first appeared to me insuperable and actually fatal to my whole theory". In the next paragraph of his book, Darwin describes a solution. If the trait of sterility can be carried by some individuals without expression, and those individuals that do express sterility help reproductive relatives, the sterile trait can persist and evolve.
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in which the queen mates with a single male, who then dies before colony founding. This seems to be the ancestral state in all
Hymenopteran lineages that have evolved eusociality. Most termites also have a mating system in which a reproductive female (the queen) commits to a single male for life (the king), and this pattern seems to be ancestral in termites. Lastly, strict monogamy facilitated eusociality in the sponge-dwelling shrimp.
244:(they come from unfertilized eggs and thus only have one set of chromosomes), and females are produced from fertilized eggs, sisters from a singly-mated mother share (on average) 75% of their genes, whereas mothers always share only 50% of their genes with their offspring. Thus, sisters will propagate their own genes more by helping their mothers to raise more sisters, than to leave the nest and raise their own daughters.
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149:, and reproductive altruism. Overlapping generations means that multiple generations live together, and that older offspring may help the parents raise their siblings. Cooperative brood care is when individuals other than the parents assist in raising the offspring through means such as food gathering and protection. Philopatry is when individuals remain living in their birthplace.
318:
distributions throughout the environments of these animals. This means there is a high cost to dispersing (individual may not find another source before it starves), and these resources must be defended for the group to survive. These requirements make it a necessity to have high social order for the survival of the group.
205:
becoming important theories during the 20th century to help explain eusociality. Inclusive fitness is described as a combination of one's own reproductive success and the reproductive success of others that share similar genes. Animals may increase their inclusive fitness through kin selection. Kin
156:
occurs when an individual performs a behavior that benefits a recipient in some way, but at the individual's own expense. Reproductive altruism is one of the most extreme forms of altruism. This is when most members of the group give up their own breeding opportunities in order to participate in the
290:
In termites, two additional hypotheses have been proposed. The first is the
Chromosomal Linkage Hypothesis, where much of the termite genome is sex-linked. This makes sisters related somewhat above 50%, and brothers somewhat above 50%, but brother-sister relatedness less than 50%. Termite workers
352:
can act. This could have been important in other eusocial genera. Biased gene conversion rates are also higher in eusocial species. This could increase genotypic diversity, which could allow workers to meet the demands of a changing social structure more easily. Another hypothesis is that the lower
268:
In many monogamous animals, an individual's death prompts its partner to look for a new mate, which would affect relatedness and hinder the evolution of eusociality: workers would be much more related to their offspring than their siblings. However, many
Hymenoptera have a form of lifetime monogamy
370:
Eusociality appears to be maintained through manipulation of the sterile workers by the queen. The mechanisms for this include hormonal control through pheromones, restricting food to young in order to control their size, consumption of any eggs laid by females other than the queen, and behavioral
189:
Darwin was on the right track, except sterility is not a characteristic shared among all eusocial animals. Sterile workers of many eusocial species are not actually physiologically sterile. Male workers can still produce sperm, and female workers sometimes lay eggs, and in some species, become the
308:
offspring (require large amounts of parental care to reach maturity); 2. Low reproductive success rates of solitary pairs that attempt to reproduce. These pre-conditions led to the two lifestyle characteristics that are observed in all eusocial species: nest building and extensive parental care.
281:
can occur, as is the case in eusocial species. Inbreeding can mimic and even surpass the effects of haplodiploidy. Siblings may actually share greater than 75% of their genes. Like in haplodiploidy kin selection, the individuals can propagate their own genes more through the promotion of more
247:
Though
Hamilton's argument appears to work well for Hymenoptera, it excludes diploid eusocial organisms (inter-sibling relatedness ≤ parent-offspring relatedness = 0.5). Even in haplodiploid systems, the average relatedness between sisters falls off rapidly when a queen mates with multiple males
294:
The symbiont hypothesis in termites is quite different from the others. With each molt, termites lose the lining of their hindgut and the subsequent bacteria and protozoa that colonize their guts for cellulose digestion. They depend on interactions with other termites for their gut to be
317:
Ecological factors were also probably a precursor to eusociality. For example, the sponge-dwelling shrimp depend upon the sponge's feeding current for food, termites depend upon dead, decaying wood, and naked mole rats depend upon tubers in the ground. Each of these resources has patchy
678:
210:, which suggests that if the benefit of a behavior to a recipient, taking into account the genetic relatedness of the recipient to the altruist, outweighs the costs of the behavior to the altruist, then it is in the altruist's genetic advantage to perform the altruistic behavior.
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might then bias their cooperative brood care towards their own sex. This hypothesis also mimics the effects of haplodiploidy, but proposes that males would help raise only the queen's male offspring, while females would only care for the queen's female offspring.
260:
in 2007, is currently the leading hypothesis concerning the initial evolution of eusociality in the
Hymenoptera. It uses Hamilton's kin selection approach in a way that applies to both haploid and diploid organisms. If a queen is lifetime-strictly
409:
Nowak's paper, however, received major criticisms for erroneously separating inclusive fitness theory from "standard natural selection". Over 150 authors replied arguing that Nowak, et al. misrepresent 40 years of empirical literature.
303:
Although the symbiont hypothesis serves as a pre-condition for termites to evolve into eusocial societies, scientists have found two crucial pre-conditions for the evolution of eusociality across all species. These include: 1.
95:. The fact that eusociality has evolved so often in the Hymenoptera (between 8 and 11 times), but remains rare throughout the rest of the animal kingdom, has made its evolution a topic of debate among evolutionary biologists.
398:
Natural
Selection Acts on Emergent Traits: The interactions of the individuals can be considered as part of the extended phenotype of the queen. These interactions produce emergent properties upon which natural selection can
130:
and colleagues. No single trait or model is sufficient to explain the evolution of eusociality, and most likely the pathway to eusociality involved a combination of pre-conditions, ecological factors, and genetic influences.
371:
dominance. In naked mole rats, this behavioral dominance occurs in the form of the queen facing the worker head-to-head, and shoving it throughout the tunnels of the naked mole rats' burrow for quite a distance.
395:
Mutations: Mutations will arise and be selected. Some genes are known to have been silenced in social insect history, leading to the reduction of dispersal behavior and the origin of the wingless caste.
161:
of other individuals. The individuals giving up their own reproductive success form a sterile caste of workers within the group. All species that practice reproductive altruism produce one or
206:
selection is when individuals help close relatives with their reproduction process, seemingly because relatives will propagate some of the individual's own genes. Kin selection follows
282:
siblings, rather than their own offspring. For example, the need for dispersal and aggregation of multiclonal groups may have helped to drive the evolution of eusociality in aphids.
165:
queens, the only breeding females, who are larger than the rest. The remainder of the society is composed of a few breeding males, sterile male and female workers, and the young.
389:
Formation of groups: Groups could consist of parent-offspring groups or unrelated groups (in situations where cooperation is beneficial) living in a structured nest.
348:. These mechanisms are likely important to the evolution of eusociality because high recombination rates are associated with the creation of novel genes, upon which
2800:
469:
Hughes, W. O. H.; Oldroyd, B. P.; Beekman, M.; Ratnieks, F. L. W. (2008). "Ancestral
Monogamy Shows Kin Selection is Key to the Evolution of Eusociality".
326:
Genetic constraints may have influenced the evolution of eusociality. The genome structure of the order
Hymenoptera has been found to have the highest
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1849:
295:
recolonized, thus forcing them to become social. This could be a precursor, or pre-condition for why eusociality evolved in termites.
236:
proposed that eusociality arose in social
Hymenoptera by kin selection because of their interesting genetic sex determination trait of
392:
Pre-adaptations: Pre-adaptations for social living, such as progressive provisioning, will push the group further toward eusociality.
3053:
2508:
61:). This 'true sociality' in animals, in which sterile individuals work to further the reproductive success of others, is found in
385:
Nowak, et al. (2010) outlines a path by which eusociality could evolve by means of multi-level (group) selection in five steps:
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On the Origin of
Species by Means of Natural Selection or The Preservation of Favored Races in the Struggle for Life
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Eusociality can be characterized by four main criteria: overlapping generations, cooperative brood care,
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1951:
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1163:"Recombination is associated with the evolution of genome structure and worker behavior in honey bees"
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892:"Kin structure, ecology and the evolution of social organization in shrimp: a comparative analysis"
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workers collaborating on a comb have given up their ability to reproduce, an extreme expression of
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1956:
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1071:"Ecology, Not the Genetics of Sex Determination, Determines Who Helps in Eusocial Populations"
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2008:
2003:
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336:, the honeybees, have the highest recombination rate in higher eukaryotes. Genes determining
182:
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The final category, reproductive altruism, is the most divergent from other social orders.
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1227:"Variation in genomic recombination rates among animal taxa and the case of social insects"
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In species where philopatry predominates, and there are few emigrants to the nest, intense
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Abbot, Patrick (October 2009). "On the evolution of dispersal and altruism in aphids".
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organisms at first appear to behave in stark contrast with simple interpretations of
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Current theories propose that the evolution of eusociality occurred either due to
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overall genetic diversity as eusociality levels increase throughout the family
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119:
84:
1268:"Genomic signatures of evolutionary transitions from solitary to group living"
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1030:"Colony size, social complexity and reproductive conflict in social insects"
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considered the evolution of eusociality a major problem for his theory of
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2013:
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62:
27:
23:
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732:"Evolution of sociality in a primitively eusocial lineage of bees"
2331:
2045:
1692:
1631:
1621:
794:"Kin selection versus sexual selection: why the ends do not meet"
340:
behavior and division of labor have been found in regions of the
42:
1589:
74:
70:
38:
1434:
405:: More cooperative groups out-compete less cooperative groups.
2252:
1325:"Genes involved in convergent evolution eusociality in bees"
468:
530:(1964). "The genetical theory of social behaviour, I, II".
357:
is due to a decreased exposure to parasites and pathogens.
50:
1584:
58:
54:
690:
190:
new queen if the old one dies (observed in Hymenoptera,
1113:
Duffy, J (1996). "Eusociality in a coral reef shrimp".
896:
Proceedings of the Royal Society B: Biological Sciences
843:"Lifetime monogamy and the evolution of eusociality"
330:
of any other groups in Animalia. The eusocial genus
103:: passing on one's genes to the next generation, or
344:genome with the highest rates of recombination and
437:Andersson, M. (1984). "Evolution of eusociality".
1375:
691:Foster, K.; Wenseleers, T.; Ratnieks, F. (2006).
3094:
1329:Proceedings of the National Academy of Sciences
1167:Proceedings of the National Academy of Sciences
736:Proceedings of the National Academy of Sciences
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1218:
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218:
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1820:
1806:
1457:
1443:
1384:"Inclusive fitness theory and eusociality"
522:
520:
518:
516:
134:
37:evolved repeatedly in different orders of
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1299:
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982:
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866:
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809:
765:
755:
647:
609:
457:
436:
3054:Transgenerational epigenetic inheritance
1827:
1149:
1108:
1106:
782:
729:
648:Honeycutt, R (1992). "Naked mole-rats".
626:
526:
419:
18:
1322:
1265:
1224:
883:
840:
791:
723:
571:
569:
513:
256:The monogamy hypothesis, formulated by
91:), and many genera in the insect order
3095:
1027:
989:"Evolution of eusociality in termites"
986:
975:
890:Duff, J. E.; Macdonald, K. S. (2010).
693:"Kin selection is the key to altruism"
2817:Dialogues Concerning Natural Religion
1801:
1438:
1112:
1103:
938:
668:
575:
321:
312:
1160:
1068:
730:Danforth, Bryan (October 29, 2001).
566:
451:10.1146/annurev.es.15.110184.001121
213:
168:
13:
2222:Evolutionary developmental biology
374:
14:
3119:
697:Trends in Ecology & Evolution
298:
77:, marine sponge-dwelling shrimp (
16:Origins of cooperative brood care
3076:
3067:
3066:
1563:
1047:10.1046/j.1420-9101.1999.00028.x
953:10.1111/j.1558-5646.2009.00744.x
240:. Because males are produced by
122:, or by the competing theory of
2879:Extended evolutionary synthesis
2068:Gene-centered view of evolution
1767:Sociobiology: The New Synthesis
1464:
1382:Abbot, P.; et al. (2011).
1316:
1259:
1062:
1034:Journal of Evolutionary Biology
1021:
1005:10.1146/annurev.ecolsys.28.1.27
932:
365:
3007:Hologenome theory of evolution
2874:History of molecular evolution
2100:Evolutionarily stable strategy
1989:Last universal common ancestor
684:
578:"The evolution of eusociality"
532:Journal of Theoretical Biology
1:
2801:Renaissance and Enlightenment
1323:Woodard; et al. (2011).
1266:Kapheim; et al. (2015).
1225:Wilfert; et al. (2007).
413:
360:
272:
3012:Missing heritability problem
2639:Gamete differentiation/sexes
552:10.1016/0022-5193(64)90038-4
7:
1533:Sexual selection in insects
576:Nowak; et al. (2010).
285:
251:
219:Haplodiploidy/Kin selection
10:
3124:
2644:Life cycles/nuclear phases
2196:Trivers–Willard hypothesis
1685:Austroplatypus incompertus
1513:Identity in social insects
1161:Kent; et al. (2012).
709:10.1016/j.tree.2005.11.020
378:
222:
138:
3062:
2962:
2887:
2791:
2718:
2674:
2529:
2433:
2250:
2209:
2142:Parent–offspring conflict
2078:
1947:Earliest known life forms
1868:
1835:
1722:
1701:
1572:
1561:
1472:
1088:10.1016/j.cub.2013.10.013
811:10.1016/j.cub.2007.06.033
2995:Cultural group selection
2859:The eclipse of Darwinism
2831:On the Origin of Species
2806:Transmutation of species
1480:Evolution of eusociality
3000:Dual inheritance theory
2839:History of paleontology
1743:Charles Duncan Michener
1350:10.1073/pnas.1103457108
1292:10.1126/science.aaa4788
1188:10.1073/pnas.1208094109
491:10.1126/science.1156108
135:Overview of eusociality
107:, is a central idea in
2688:Punctuated equilibrium
2009:Non-adaptive radiation
1957:Evolutionary arms race
1244:10.1038/sj.hdy.6800950
1028:Bourke, A. F. (1999).
908:10.1098/rspb.2009.1483
859:10.1098/rstb.2009.0101
847:Phil. Trans. R. Soc. B
757:10.1073/pnas.012387999
31:
2980:Evolutionary medicine
2854:Mendelian inheritance
2562:Biological complexity
2550:Programmed cell death
2242:Phenotypic plasticity
1962:Evolutionary pressure
1952:Evidence of evolution
1850:Timeline of evolution
1748:The Bees of the World
993:Annu. Rev. Ecol. Syst
439:Annu. Rev. Ecol. Syst
403:Multi-level selection
183:The Origin of Species
89:Heterocephalus glaber
22:
3108:Evolutionary biology
2954:Teleology in biology
2849:Blending inheritance
2227:Genetic assimilation
2090:Artificial selection
1829:Evolutionary biology
841:Boomsma, J. (2009).
792:Boomsma, J. (2007).
197:This insight led to
159:reproductive success
124:multilevel selection
109:evolutionary biology
3017:Molecular evolution
2975:Ecological genetics
2844:Transitional fossil
2634:Sexual reproduction
2474:endomembrane system
2403:pollinator-mediated
2359:dolphins and whales
2137:Parental investment
1783:Journey to the Ants
1408:10.1038/nature09831
1400:2011Natur.471E...1A
1341:2011PNAS..108.7472W
1284:2015Sci...348.1139K
1179:2012PNAS..10918012K
1173:(44): 18012–18017.
1127:1996Natur.381..512D
853:(1533): 3191–3208.
748:2002PNAS...99..286D
662:1992AmSci..80...43H
602:10.1038/nature09205
594:2010Natur.466.1057N
588:(7310): 1057–1062.
544:1964JThBi...7....1H
483:2008Sci...320.1213H
477:(5880): 1213–1216.
346:molecular evolution
328:recombination rates
234:William D. Hamilton
101:Darwinian evolution
2990:Cultural evolution
2105:Fisher's principle
2034:Handicap principle
2024:Parallel evolution
1888:Adaptive radiation
987:Thorne, B (1997).
650:American Scientist
322:Genetic influences
313:Ecological factors
80:Synalpheus regalis
32:
3090:
3089:
2706:Uniformitarianism
2659:Sex-determination
2164:Sexual dimorphism
2159:Natural selection
2063:Unit of selection
2029:Signalling theory
1795:
1794:
1709:Bees in mythology
1335:(18): 7472–7477.
1278:(6239): 1139–43.
1121:(6582): 512–514.
1081:(23): 2383–2387.
1069:Ross, L. (2013).
947:(10): 2687–2696.
902:(1681): 575–584.
804:(16): R673–R683.
350:natural selection
199:inclusive fitness
178:natural selection
3115:
3080:
3070:
3069:
2869:Modern synthesis
2629:Multicellularity
2624:Mosaic evolution
2509:auditory ossicle
2191:Social selection
2174:Flowering plants
2169:Sexual selection
1822:
1815:
1808:
1799:
1798:
1735:The Dancing Bees
1567:
1459:
1452:
1445:
1436:
1435:
1430:
1429:
1419:
1379:
1373:
1372:
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1320:
1314:
1313:
1303:
1263:
1257:
1256:
1246:
1222:
1211:
1210:
1200:
1190:
1158:
1147:
1146:
1135:10.1038/381512a0
1110:
1101:
1100:
1090:
1066:
1060:
1059:
1049:
1025:
1019:
1018:
1016:
984:
973:
972:
936:
930:
929:
919:
887:
881:
880:
870:
838:
832:
831:
813:
789:
780:
779:
769:
759:
727:
721:
720:
688:
682:
672:
666:
665:
645:
624:
623:
613:
573:
564:
563:
524:
511:
510:
466:
455:
454:
434:
214:Current theories
169:Early hypotheses
69:, gall-dwelling
67:ambrosia beetles
3123:
3122:
3118:
3117:
3116:
3114:
3113:
3112:
3093:
3092:
3091:
3086:
3058:
2985:Group selection
2958:
2883:
2787:
2714:
2676:Tempo and modes
2670:
2525:
2429:
2246:
2205:
2081:
2074:
2051:Species complex
1864:
1855:History of life
1831:
1826:
1796:
1791:
1756:Bert Hölldobler
1730:Karl von Frisch
1723:Pioneers, works
1718:
1697:
1568:
1559:
1553:Worker policing
1538:Social conflict
1518:Kin recognition
1503:Group selection
1489:Social insects
1468:
1463:
1433:
1394:(7339): E1–E4.
1380:
1376:
1321:
1317:
1264:
1260:
1223:
1214:
1159:
1150:
1111:
1104:
1075:Current Biology
1067:
1063:
1026:
1022:
985:
976:
937:
933:
888:
884:
839:
835:
798:Current Biology
790:
783:
728:
724:
689:
685:
673:
669:
646:
627:
574:
567:
528:Hamilton, W. D.
525:
514:
467:
458:
435:
420:
416:
383:
381:Group selection
377:
375:Group selection
368:
363:
324:
315:
301:
288:
275:
258:Jacobus Boomsma
254:
242:parthenogenesis
231:
223:Main articles:
221:
216:
208:Hamilton's Rule
194:, and shrimp).
171:
143:
137:
126:as proposed by
85:naked mole-rats
17:
12:
11:
5:
3121:
3111:
3110:
3105:
3088:
3087:
3085:
3084:
3074:
3063:
3060:
3059:
3057:
3056:
3051:
3046:
3041:
3036:
3035:
3034:
3024:
3019:
3014:
3009:
3004:
3003:
3002:
2997:
2992:
2982:
2977:
2972:
2966:
2964:
2960:
2959:
2957:
2956:
2951:
2950:
2949:
2944:
2939:
2938:
2937:
2927:
2922:
2917:
2912:
2907:
2897:
2891:
2889:
2885:
2884:
2882:
2881:
2876:
2871:
2866:
2861:
2856:
2851:
2846:
2841:
2836:
2835:
2834:
2825:Charles Darwin
2822:
2821:
2820:
2808:
2803:
2797:
2795:
2789:
2788:
2786:
2785:
2780:
2775:
2770:
2765:
2763:Non-ecological
2760:
2755:
2750:
2745:
2740:
2735:
2730:
2724:
2722:
2716:
2715:
2713:
2712:
2703:
2694:
2680:
2678:
2672:
2671:
2669:
2668:
2663:
2662:
2661:
2656:
2651:
2646:
2641:
2631:
2626:
2621:
2616:
2611:
2606:
2601:
2596:
2591:
2586:
2581:
2580:
2579:
2569:
2564:
2559:
2554:
2553:
2552:
2547:
2536:
2534:
2527:
2526:
2524:
2523:
2522:
2521:
2516:
2514:nervous system
2511:
2506:
2501:
2493:
2492:
2491:
2486:
2481:
2476:
2471:
2466:
2456:
2451:
2446:
2440:
2438:
2431:
2430:
2428:
2427:
2422:
2417:
2412:
2407:
2406:
2405:
2395:
2394:
2393:
2388:
2387:
2386:
2381:
2371:
2366:
2361:
2356:
2351:
2350:
2349:
2344:
2334:
2324:
2319:
2318:
2317:
2307:
2302:
2297:
2292:
2291:
2290:
2280:
2275:
2274:
2273:
2263:
2257:
2255:
2248:
2247:
2245:
2244:
2239:
2234:
2229:
2224:
2219:
2213:
2211:
2207:
2206:
2204:
2203:
2198:
2193:
2188:
2187:
2186:
2181:
2176:
2166:
2161:
2156:
2151:
2146:
2145:
2144:
2139:
2129:
2124:
2119:
2118:
2117:
2107:
2102:
2097:
2092:
2086:
2084:
2076:
2075:
2073:
2072:
2071:
2070:
2060:
2055:
2054:
2053:
2048:
2038:
2037:
2036:
2026:
2021:
2016:
2014:Origin of life
2011:
2006:
2001:
1999:Microevolution
1996:
1994:Macroevolution
1991:
1986:
1981:
1980:
1979:
1969:
1964:
1959:
1954:
1949:
1944:
1939:
1934:
1932:Common descent
1929:
1928:
1927:
1917:
1912:
1910:Baldwin effect
1907:
1906:
1905:
1900:
1890:
1885:
1880:
1874:
1872:
1866:
1865:
1863:
1862:
1857:
1852:
1847:
1842:
1836:
1833:
1832:
1825:
1824:
1817:
1810:
1802:
1793:
1792:
1790:
1789:
1788:
1787:
1779:
1771:
1758:
1753:
1752:
1751:
1740:
1739:
1738:
1726:
1724:
1720:
1719:
1717:
1716:
1711:
1705:
1703:
1699:
1698:
1696:
1695:
1690:
1689:
1688:
1676:
1675:
1674:
1664:
1663:
1662:
1650:
1649:
1648:
1636:
1635:
1634:
1629:
1627:Dwarf mongoose
1624:
1614:
1613:
1612:
1607:
1602:
1597:
1592:
1587:
1576:
1574:
1570:
1569:
1562:
1560:
1558:
1557:
1556:
1555:
1550:
1545:
1540:
1535:
1530:
1525:
1520:
1515:
1510:
1505:
1500:
1495:
1487:
1482:
1476:
1474:
1470:
1469:
1462:
1461:
1454:
1447:
1439:
1432:
1431:
1374:
1315:
1258:
1237:(4): 189–197.
1212:
1148:
1102:
1061:
1040:(2): 245–257.
1020:
974:
931:
882:
833:
781:
742:(1): 286–290.
722:
683:
681:. John Murray.
667:
625:
565:
512:
456:
417:
415:
412:
407:
406:
400:
396:
393:
390:
379:Main article:
376:
373:
367:
364:
362:
359:
323:
320:
314:
311:
300:
299:Pre-conditions
297:
287:
284:
274:
271:
253:
250:
220:
217:
215:
212:
174:Charles Darwin
170:
167:
139:Main article:
136:
133:
120:W. D. Hamilton
118:, proposed by
15:
9:
6:
4:
3:
2:
3120:
3109:
3106:
3104:
3101:
3100:
3098:
3083:
3079:
3075:
3073:
3065:
3064:
3061:
3055:
3052:
3050:
3047:
3045:
3042:
3040:
3037:
3033:
3030:
3029:
3028:
3027:Phylogenetics
3025:
3023:
3020:
3018:
3015:
3013:
3010:
3008:
3005:
3001:
2998:
2996:
2993:
2991:
2988:
2987:
2986:
2983:
2981:
2978:
2976:
2973:
2971:
2968:
2967:
2965:
2961:
2955:
2952:
2948:
2945:
2943:
2940:
2936:
2933:
2932:
2931:
2930:Structuralism
2928:
2926:
2923:
2921:
2918:
2916:
2913:
2911:
2908:
2906:
2905:Catastrophism
2903:
2902:
2901:
2898:
2896:
2893:
2892:
2890:
2886:
2880:
2877:
2875:
2872:
2870:
2867:
2865:
2864:Neo-Darwinism
2862:
2860:
2857:
2855:
2852:
2850:
2847:
2845:
2842:
2840:
2837:
2833:
2832:
2828:
2827:
2826:
2823:
2819:
2818:
2814:
2813:
2812:
2809:
2807:
2804:
2802:
2799:
2798:
2796:
2794:
2790:
2784:
2781:
2779:
2778:Reinforcement
2776:
2774:
2771:
2769:
2766:
2764:
2761:
2759:
2756:
2754:
2751:
2749:
2746:
2744:
2741:
2739:
2736:
2734:
2731:
2729:
2726:
2725:
2723:
2721:
2717:
2711:
2710:Catastrophism
2707:
2704:
2702:
2701:Macromutation
2698:
2697:Micromutation
2695:
2693:
2689:
2685:
2682:
2681:
2679:
2677:
2673:
2667:
2664:
2660:
2657:
2655:
2652:
2650:
2647:
2645:
2642:
2640:
2637:
2636:
2635:
2632:
2630:
2627:
2625:
2622:
2620:
2617:
2615:
2612:
2610:
2607:
2605:
2604:Immune system
2602:
2600:
2597:
2595:
2592:
2590:
2587:
2585:
2582:
2578:
2575:
2574:
2573:
2570:
2568:
2565:
2563:
2560:
2558:
2555:
2551:
2548:
2546:
2543:
2542:
2541:
2538:
2537:
2535:
2533:
2528:
2520:
2517:
2515:
2512:
2510:
2507:
2505:
2502:
2500:
2497:
2496:
2494:
2490:
2487:
2485:
2482:
2480:
2477:
2475:
2472:
2470:
2467:
2465:
2464:symbiogenesis
2462:
2461:
2460:
2457:
2455:
2452:
2450:
2447:
2445:
2442:
2441:
2439:
2437:
2432:
2426:
2423:
2421:
2418:
2416:
2413:
2411:
2408:
2404:
2401:
2400:
2399:
2396:
2392:
2389:
2385:
2382:
2380:
2377:
2376:
2375:
2372:
2370:
2367:
2365:
2362:
2360:
2357:
2355:
2352:
2348:
2345:
2343:
2340:
2339:
2338:
2335:
2333:
2330:
2329:
2328:
2325:
2323:
2320:
2316:
2313:
2312:
2311:
2308:
2306:
2303:
2301:
2298:
2296:
2293:
2289:
2286:
2285:
2284:
2281:
2279:
2276:
2272:
2269:
2268:
2267:
2264:
2262:
2259:
2258:
2256:
2254:
2249:
2243:
2240:
2238:
2235:
2233:
2230:
2228:
2225:
2223:
2220:
2218:
2215:
2214:
2212:
2208:
2202:
2199:
2197:
2194:
2192:
2189:
2185:
2182:
2180:
2177:
2175:
2172:
2171:
2170:
2167:
2165:
2162:
2160:
2157:
2155:
2152:
2150:
2147:
2143:
2140:
2138:
2135:
2134:
2133:
2132:Kin selection
2130:
2128:
2127:Genetic drift
2125:
2123:
2120:
2116:
2113:
2112:
2111:
2108:
2106:
2103:
2101:
2098:
2096:
2093:
2091:
2088:
2087:
2085:
2083:
2077:
2069:
2066:
2065:
2064:
2061:
2059:
2056:
2052:
2049:
2047:
2044:
2043:
2042:
2039:
2035:
2032:
2031:
2030:
2027:
2025:
2022:
2020:
2017:
2015:
2012:
2010:
2007:
2005:
2002:
2000:
1997:
1995:
1992:
1990:
1987:
1985:
1982:
1978:
1975:
1974:
1973:
1970:
1968:
1965:
1963:
1960:
1958:
1955:
1953:
1950:
1948:
1945:
1943:
1940:
1938:
1935:
1933:
1930:
1926:
1923:
1922:
1921:
1918:
1916:
1913:
1911:
1908:
1904:
1901:
1899:
1896:
1895:
1894:
1891:
1889:
1886:
1884:
1881:
1879:
1876:
1875:
1873:
1871:
1867:
1861:
1858:
1856:
1853:
1851:
1848:
1846:
1843:
1841:
1838:
1837:
1834:
1830:
1823:
1818:
1816:
1811:
1809:
1804:
1803:
1800:
1785:
1784:
1780:
1777:
1776:
1772:
1769:
1768:
1764:
1763:
1762:
1759:
1757:
1754:
1749:
1746:
1745:
1744:
1741:
1736:
1733:
1732:
1731:
1728:
1727:
1725:
1721:
1715:
1712:
1710:
1707:
1706:
1704:
1700:
1694:
1691:
1687:
1686:
1682:
1681:
1680:
1677:
1673:
1670:
1669:
1668:
1665:
1661:
1660:
1656:
1655:
1654:
1651:
1647:
1646:
1642:
1641:
1640:
1637:
1633:
1630:
1628:
1625:
1623:
1620:
1619:
1618:
1615:
1611:
1608:
1606:
1603:
1601:
1598:
1596:
1593:
1591:
1588:
1586:
1583:
1582:
1581:
1578:
1577:
1575:
1571:
1566:
1554:
1551:
1549:
1546:
1544:
1541:
1539:
1536:
1534:
1531:
1529:
1526:
1524:
1523:Kin selection
1521:
1519:
1516:
1514:
1511:
1509:
1508:Haplodiploidy
1506:
1504:
1501:
1499:
1496:
1494:
1491:
1490:
1488:
1486:
1483:
1481:
1478:
1477:
1475:
1471:
1467:
1460:
1455:
1453:
1448:
1446:
1441:
1440:
1437:
1427:
1423:
1418:
1413:
1409:
1405:
1401:
1397:
1393:
1389:
1385:
1378:
1370:
1366:
1361:
1356:
1351:
1346:
1342:
1338:
1334:
1330:
1326:
1319:
1311:
1307:
1302:
1297:
1293:
1289:
1285:
1281:
1277:
1273:
1269:
1262:
1254:
1250:
1245:
1240:
1236:
1232:
1228:
1221:
1219:
1217:
1208:
1204:
1199:
1194:
1189:
1184:
1180:
1176:
1172:
1168:
1164:
1157:
1155:
1153:
1144:
1140:
1136:
1132:
1128:
1124:
1120:
1116:
1109:
1107:
1098:
1094:
1089:
1084:
1080:
1076:
1072:
1065:
1057:
1053:
1048:
1043:
1039:
1035:
1031:
1024:
1015:
1010:
1006:
1002:
999:(11): 27–54.
998:
994:
990:
983:
981:
979:
970:
966:
962:
958:
954:
950:
946:
942:
935:
927:
923:
918:
913:
909:
905:
901:
897:
893:
886:
878:
874:
869:
864:
860:
856:
852:
848:
844:
837:
829:
825:
821:
817:
812:
807:
803:
799:
795:
788:
786:
777:
773:
768:
763:
758:
753:
749:
745:
741:
737:
733:
726:
718:
714:
710:
706:
702:
698:
694:
687:
680:
676:
671:
663:
659:
655:
651:
644:
642:
640:
638:
636:
634:
632:
630:
621:
617:
612:
607:
603:
599:
595:
591:
587:
583:
579:
572:
570:
561:
557:
553:
549:
545:
541:
537:
533:
529:
523:
521:
519:
517:
508:
504:
500:
496:
492:
488:
484:
480:
476:
472:
465:
463:
461:
452:
448:
444:
440:
433:
431:
429:
427:
425:
423:
418:
411:
404:
401:
397:
394:
391:
388:
387:
386:
382:
372:
358:
356:
351:
347:
343:
339:
335:
334:
329:
319:
310:
307:
296:
292:
283:
280:
270:
266:
264:
259:
249:
245:
243:
239:
238:haplodiploidy
235:
230:
229:Kin selection
226:
225:Haplodiploidy
211:
209:
204:
203:kin selection
200:
195:
193:
187:
185:
184:
179:
175:
166:
164:
160:
155:
150:
148:
142:
132:
129:
125:
121:
117:
116:kin selection
112:
110:
106:
102:
98:
94:
90:
86:
82:
81:
76:
72:
68:
64:
60:
56:
52:
48:
44:
40:
36:
29:
25:
21:
3103:Sociobiology
3039:Polymorphism
3022:Astrobiology
2970:Biogeography
2925:Saltationism
2915:Orthogenesis
2900:Alternatives
2829:
2815:
2748:Cospeciation
2743:Cladogenesis
2692:Saltationism
2649:Mating types
2598:
2572:Color vision
2557:Avian flight
2479:mitochondria
2217:Canalisation
2095:Biodiversity
1840:Introduction
1781:
1773:
1765:
1761:E. O. Wilson
1747:
1734:
1683:
1657:
1653:Thysanoptera
1643:
1485:Presociality
1479:
1391:
1387:
1377:
1332:
1328:
1318:
1275:
1271:
1261:
1234:
1230:
1170:
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1114:
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1074:
1064:
1037:
1033:
1023:
996:
992:
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934:
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885:
850:
846:
836:
801:
797:
739:
735:
725:
703:(2): 57–60.
700:
696:
686:
670:
656:(1): 43–53.
653:
649:
585:
581:
535:
531:
474:
470:
442:
438:
408:
384:
369:
366:Manipulation
354:
341:
331:
325:
316:
302:
293:
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276:
267:
255:
246:
232:
196:
188:
181:
172:
151:
144:
113:
88:
78:
33:
3049:Systematics
2920:Mutationism
2738:Catagenesis
2666:Snake venom
2599:Eusociality
2577:in primates
2567:Cooperation
2495:In animals
2315:butterflies
2288:Cephalopods
2278:Brachiopods
2210:Development
2184:Mate choice
1937:Convergence
1920:Coevolution
1878:Abiogenesis
1659:Kladothrips
1595:Crabronidae
1580:Hymenoptera
1493:Agriculture
1466:Eusociality
538:(1): 1–52.
445:: 165–189.
141:Eusociality
128:E.O. Wilson
93:Hymenoptera
47:Hymenoptera
35:Eusociality
3097:Categories
2910:Lamarckism
2888:Philosophy
2811:David Hume
2773:Peripatric
2768:Parapatric
2753:Ecological
2733:Anagenesis
2728:Allopatric
2720:Speciation
2684:Gradualism
2609:Metabolism
2469:chromosome
2459:Eukaryotes
2237:Modularity
2154:Population
2080:Population
2041:Speciation
2019:Panspermia
1972:Extinction
1967:Exaptation
1942:Divergence
1915:Cladistics
1903:Reciprocal
1883:Adaptation
1714:Coalescent
1702:In culture
1679:Coleoptera
1645:Synalpheus
1639:Crustacean
1600:Halictidae
1528:Polyethism
675:Darwin, C.
414:References
361:Mechanisms
279:inbreeding
273:Inbreeding
263:monogamous
147:philopatry
41:, notably
3044:Protocell
2895:Darwinism
2783:Sympatric
2532:processes
2420:Tetrapods
2369:Kangaroos
2295:Dinosaurs
2232:Inversion
2201:Variation
2122:Gene flow
2115:Inclusive
1925:Mutualism
1870:Evolution
1672:Aphididae
1667:Hemiptera
1605:Honey bee
1543:Thelytoky
1498:Gamergate
941:Evolution
306:Altricial
30:behavior.
3072:Category
2947:Vitalism
2942:Theistic
2935:Spandrel
2619:Morality
2614:Monogamy
2489:plastids
2454:Flagella
2410:Reptiles
2391:sea cows
2374:primates
2283:Molluscs
2261:Bacteria
2149:Mutation
2082:genetics
2058:Taxonomy
2004:Mismatch
1984:Homology
1898:Cheating
1893:Altruism
1775:The Ants
1693:Isoptera
1617:Mammalia
1610:Vespidae
1426:21430721
1369:21482769
1310:25977371
1253:17389895
1231:Heredity
1207:23071321
1143:33166806
1097:24268409
1056:85187599
969:12865398
961:19500147
926:19889706
877:19805427
820:17714661
776:11782550
717:16701471
620:20740005
507:20388889
499:18511689
286:Termites
252:Monogamy
192:termites
154:Altruism
97:Eusocial
63:termites
45:and the
43:termites
28:eusocial
24:Honeybee
2963:Related
2793:History
2654:Meiosis
2589:Empathy
2584:Emotion
2484:nucleus
2425:Viruses
2415:Spiders
2327:Mammals
2310:Insects
2110:Fitness
2046:Species
1845:Outline
1632:Meerkat
1622:Blesmol
1417:3836173
1396:Bibcode
1360:3088614
1337:Bibcode
1301:5471836
1280:Bibcode
1272:Science
1198:3497793
1175:Bibcode
1123:Bibcode
917:2842683
868:2781870
744:Bibcode
677:(1859)
658:Bibcode
611:3279739
590:Bibcode
560:5875341
540:Bibcode
479:Bibcode
471:Science
105:fitness
39:animals
3082:Portal
2758:Hybrid
2594:Ethics
2436:organs
2398:Plants
2384:lemurs
2379:humans
2364:horses
2354:hyenas
2342:wolves
2337:canids
2271:origin
1590:Apidae
1573:Groups
1473:Topics
1424:
1414:
1388:Nature
1367:
1357:
1308:
1298:
1251:
1205:
1195:
1141:
1115:Nature
1095:
1054:
1014:349550
1011:
967:
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914:
875:
865:
828:886746
826:
818:
774:
767:117553
764:
715:
618:
608:
582:Nature
558:
505:
497:
355:Apidae
338:worker
75:thrips
71:aphids
57:, and
2545:Death
2540:Aging
2519:brain
2305:Fungi
2266:Birds
2179:Fungi
1977:Event
1860:Index
1139:S2CID
1052:S2CID
965:S2CID
824:S2CID
503:S2CID
180:. In
51:wasps
49:(the
3032:Tree
2504:hair
2444:Cell
2347:dogs
2332:cats
2322:Life
2300:Fish
2253:taxa
1786:1994
1778:1990
1770:1975
1750:2000
1737:1927
1422:PMID
1365:PMID
1306:PMID
1249:PMID
1203:PMID
1093:PMID
957:PMID
922:PMID
873:PMID
816:PMID
772:PMID
713:PMID
616:PMID
556:PMID
495:PMID
399:act.
342:Apis
333:Apis
227:and
201:and
163:more
59:ants
55:bees
2530:Of
2499:eye
2449:DNA
2434:Of
2251:Of
1585:Ant
1548:War
1412:PMC
1404:doi
1392:471
1355:PMC
1345:doi
1333:108
1296:PMC
1288:doi
1276:348
1239:doi
1193:PMC
1183:doi
1171:109
1131:doi
1119:381
1083:doi
1042:doi
1009:PMC
1001:doi
949:doi
912:PMC
904:doi
900:277
863:PMC
855:doi
851:364
806:doi
762:PMC
752:doi
705:doi
606:PMC
598:doi
586:466
548:doi
487:doi
475:320
447:doi
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