Editing Kin selection (section)

==Objections==

The theory of kin selection has been criticised by W. J. Alonso (in 1998)<ref name=Alonso1998>{{cite journal |last=Alonso |first=W. J. |year=1998 |title=The role of Kin Selection theory on the explanation of biological altruism: A critical Review |url=http://www.origem.info/KS/Alonso_1998.pdf |journal=Journal of Comparative Biology |volume=3 |issue=1 |pages=1–14 |access-date=2013-03-16 |archive-url=https://web.archive.org/web/20131109132318/http://www.origem.info/KS/Alonso_1998.pdf |archive-date=2013-11-09 |url-status=dead }}</ref> and by Alonso and C. Schuck-Paim (in 2002).<ref name=AandSP2002>{{cite journal |last1=Alonso |first1=W. J. |last2=Schuck-Paim |first2=C. |year=2002 |title=Sex-ratio conflicts, kin selection, and the evolution of altruism |journal=PNAS |volume=99 |issue=10 |pages=6843–6847 |doi=10.1073/pnas.092584299 |pmid=11997461 |pmc=124491 |bibcode=2002PNAS...99.6843A |doi-access=free }}</ref> They argue that the behaviours which kin selection attempts to explain are not altruistic (in pure Darwinian terms) because: (1) they may directly favour the performer as an individual aiming to maximise its progeny (so the behaviours can be explained as ordinary individual selection); (2) these behaviours benefit the group (so they can be explained as group selection); or (3) they are by-products of a developmental system of many "individuals" performing different tasks (like a colony of bees, or the cells of multicellular organisms, which are the focus of selection). They also argue that the genes involved in sex ratio conflicts could be treated as "parasites" of (already established) social colonies, not as their "promoters", and, therefore the sex ratio in colonies would be irrelevant to the transition to eusociality.<ref name=Alonso1998/><ref name=AandSP2002/> Those ideas were mostly ignored until they were put forward again in a series of controversial<ref name=Abbot2011/> papers by [[E. O. Wilson]], [[Bert Hölldobler]], [[Martin Nowak]] and [[Corina Tarnita]].<ref>{{cite journal |last1=Wilson |first1=Edward O. |author1-link=E. O. Wilson |last2=Hölldobler |first2=Bert |author2-link=Bert Hölldobler |date=2005 |title=Eusociality: origin and consequences |journal=PNAS |volume=102 |issue=38 |pages=13367–13371 |doi=10.1073/pnas.0505858102 |pmc=1224642 |pmid=16157878 |bibcode=2005PNAS..10213367W |doi-access=free }}</ref><ref>{{cite journal |last=Wilson |first=E. O. |author-link=E. O. Wilson |date=2008 |title=One giant leap: how insects achieved altruism and colonial life |journal=BioScience |volume=58 |pages=17–25 |doi=10.1641/b580106 |doi-access=free }}</ref><ref>{{cite journal |last1=Nowak |first1=M. A. |last2=Tarnita |first2=Corina E. |last3=Wilson |first3=E. O. |date=2010 |title=The evolution of eusociality |journal=Nature |volume=466 |issue=7310 |pages=1057–1062 |doi=10.1038/nature09205 |pmid=20740005 |pmc=3279739|bibcode=2010Natur.466.1057N }}</ref> Nowak, Tarnita and Wilson argued that

{{Blockquote|Inclusive fitness theory is not a simplification over the standard approach. It is an alternative accounting method, but one that works only in a very limited domain. Whenever inclusive fitness does work, the results are identical to those of the standard approach. Inclusive fitness theory is an unnecessary detour, which does not provide additional insight or information.|Nowak, Tarnita, and Wilson<ref name=NTW/>}}

They, like Alonso and Schuck-Paim, argue for a [[Group selection#Multilevel selection theory|multi-level selection]] model instead.<ref name=NTW/> This aroused a strong response, including a rebuttal published in ''[[Nature (magazine)|Nature]]'' from over a hundred researchers.<ref name=Abbot2011>{{cite journal |last1=Abbot |display-authors=etal |year=2011 |title=Inclusive fitness theory and eusociality |journal=Nature |volume=471 |issue=7339 |pages=E1–E4 |doi=10.1038/nature09831 |pmid=21430721 |pmc=3836173 |bibcode=2011Natur.471E...1A }}</ref>