Editing Behavioral ecology (section)

===Conflicts in social insects===
Although eusociality has been shown to offer many benefits to the colony, there is also potential for conflict. Examples include the sex-ratio conflict and [[worker policing]] seen in certain species of social Hymenoptera such as ''[[Dolichovespula media]]'', ''[[Dolichovespula sylvestris]],'' ''[[Dolichovespula norwegica]]''<ref>{{Cite journal|title = Co-occurrence of three types of egg policing in the Norwegian wasp Dolichovespsula wasp|last1 = Bonckaert|first1 = W.|date = 2001|journal = Behavioral Ecology and Sociobiology|doi = 10.1007/s00265-010-1064-3|first6 = T.|last2 = Tofilski|first2 = A.|first3 = F.S.|last3 = Nascimento|first4 = J.|last4 = Billen|first5 = F.L.W.|last5 = Ratnieks|last6 = Wenseleers |volume=65 |issue = 4|pages=633–640|s2cid = 2186614|url = https://lirias.kuleuven.be/handle/123456789/285105|url-access = subscription}}</ref> and ''[[Vespula vulgaris]].''<ref name=ESS>{{cite journal|last=Wenseleers|first=Tom|author2=Heikki Helanterä |author3=Adam G. Hart|author4=Francis L. W. Ratnieks|title=Worker reproduction and policing in insect societies: an ESS analysis|journal=Journal of Evolutionary Biology|date=May 2004|volume=17|issue=5|pages=1035–1047 |doi=10.1111/j.1420-9101.2004.00751.x|pmid=15312076|doi-access=free}}</ref><ref name="Foster">{{cite journal|last1=Foster|first1=Kevin R. |title=Colony kin structure and male production in ''Dolichovespula'' wasps|journal=Molecular Ecology |date=2001 |volume=10 |issue=4 |pages=1003–1010|doi=10.1046/j.1365-294X.2001.01228.x|pmid=11348506 |s2cid=12009153 }}</ref> The queen and the worker wasps either indirectly kill the laying-workers' offspring by neglecting them or directly condemn them by cannibalizing and scavenging.<ref>[[Vespula vulgaris#Defensive behaviors]]</ref>

The sex-ratio conflict arises from a [[relatedness]] asymmetry, which is caused by the [[haplodiploidy]] nature of [[Hymenoptera]].  For instance, workers are most related to each other because they share half of the genes from the queen and inherit all of the father's genes.  Their total relatedness to each other would be 0.5+ (0.5 x 0.5) = 0.75. Thus, sisters are three-fourths related to each other.  On the other hand, males arise from unfertilized larva, meaning they only inherit half of the queen's genes and none from the father.  As a result, a female is related to her brother by 0.25, because 50% of her genes that come from her father have no chance of being shared with a brother.  Her relatedness to her brother would therefore be 0.5 x 0.5=0.25.<ref name=Davies/>{{rp|382}}

According to Trivers and Hare's population-level sex-investment ratio theory, the ratio of relatedness between sexes determines the sex investment ratios.<ref name=sex>{{cite journal |author=Andrew F. G. Bourke |title=Sex allocation in a facultatively polygynous ant: between-population and between-colony variation |journal=Behavioral Ecology |year=1999|volume=10|issue=4|pages=409–421 |doi=10.1093/beheco/10.4.409 |doi-access=free }}</ref> As a result, it has been observed that there is a tug-of-war between the queen and the workers, where the queen would prefer a 1:1 female to male ratio because she is equally related to her sons and daughters (r=0.5 in each case).  However, the workers would prefer a 3:1 female to male ratio because they are 0.75 related to each other and only 0.25 related to their brothers.<ref name=Davies/>{{rp|382}} [[Allozyme]] data of a colony may indicate who wins this conflict.<ref name=Colony>{{cite journal |author=Jurgen Heinze|title=Colony structure and reproduction in the ant, ''Leptothorax acervorum''|journal=Behavioral Ecology |year=1994|volume=6|issue=4|pages=359–367 |doi=10.1093/beheco/6.4.359 |last2=Lipski |first2=Norbert |last3=Schlehmeyer |first3=Kathrin |last4=Hōlldobler |first4=Bert}}</ref>

Conflict can also arise between workers in colonies of social insects. In some species, worker females retain their ability to mate and lay eggs. The colony's queen is related to her sons by half of her genes and a quarter to the sons of her worker daughters. Workers, however, are related to their sons by half of their genes and to their brothers by a quarter. Thus, the queen and her worker daughters would compete for reproduction to maximize their own reproductive fitness. Worker reproduction is limited by other workers who are more related to the queen than their sisters, a situation occurring in many polyandrous hymenopteran species. Workers police the egg-laying females by engaging in [[oophagy]] or directed acts of aggression.<ref name=Policing.Bee>{{cite journal |last=Ratnieks|first=Francis L.W.|author2=P. Kirk Visscher|title=Worker policing in the honeybee |journal=Nature |date=December 1989 |volume=342 |issue=6251|pages=796–797 |doi=10.1038/342796a0 |bibcode = 1989Natur.342..796R |s2cid=4366903}}</ref><ref>{{cite journal|last=Gobin|first=Bruno |author2=J. Billen |author3=C. Peeters |title=Policing behaviour towards virgin egg layers in a polygynous ponerine ant|journal=Animal Behaviour |date=November 1999 |volume=58|issue=5|pages=1117–1122|pmid=10564615|doi=10.1006/anbe.1999.1245|s2cid=16428974 }}</ref>