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The Pay-to-Stay Hypothesis suggests that provisioning behavior done by helpers is rewarded by dominant breeders. The dominant breeders reward them by allowing them to stay within the community /family structure longer. The helpers receive nepotistic (hierarchal power) benefits because they are favored by relatives.

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Kin selection is an evolutionary strategy that favors the reproductive success of an organisms relatives. Kin selection occurs when an animal engages in self-sacrificial behavior that benefits the genetic fitness of its relatives. The elements of kin selection lead directly to the concept now known as Hamilton's rule, which states that aid-giving behavior can evolve when the indirect fitness benefits of helping relatives compensate the aid giver for any losses in personal reproduction it incurs by helping.

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Honeybees are great image of how working together can be very beneficial. Kin selection theory thus explains altruistic behavior as a function of improving one’s own genetic legacy, or why some individuals help others at a personal cost: their genes win out in the bargain. Honeybees all work to protect the queen who is the mother of all in the hive. So promoting the survival and increase of population is in your genetic and physical well-being.

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Alloparenting is the care given by individuals other than parents. The term babysitting is a resemblance of this behavior. Alloparents are referred to as "helpers" and aid biological mothers in caring for her offspring. From an evolutionary perspective, alloparents may have offered critical advantages for survival and reproductive success.

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How does parental investment play into Pay-To-Stay?  

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How does parental investment play into Pay-To-Stay?  

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Dominants may respond to increased effort by rent-paying subordinates by reducing their own effort. If ‘‘lazy’’ dominants survive better, selfish dominants will increase their standards of what they're willing to accept from the subordinates. Rent is paid if the maximum effort that subordinates are willing to pay, equals or exceeds the minimum that dominants require. This means a selfish dominant does not create a mutually beneficial situation, so a subordinate is less likely to participate. Both the dominant
and the subordinate benefit more from group formation than
from the dispersal of the subordinate.

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Why do individuals participate in Pay-to-Stay?

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Why do individuals participate in Pay-to-Stay?

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There are two reasons why individuals may participate in Pay-to-Stay that we are going to cover. The first is due to kin selection and the second is due to ecological constraints.

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Individuals may participate in pay-to-stay due to kin selection. This is because it will benefit subordinates more if they help raise kin rather than trying to reproduce on their own. For example, cooperatively breeding acorn woodpeckers gain indirect fitness by helping in provisioning related offspring (Koenig & Walters, 2011). Provisioning behavior is the submissive behaviors helpers perform that help dominants raise their young.

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Start video at 0:55.00

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The Anomalies: The Acorn Woodpecker | bioGraphic

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Evidence for Pay-to-stay due to kin selection

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Evidence for Pay-to-stay due to kin selection

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Koenig and Walters analyzed the relationship between provisioning by helpers and the length of time they remain in the natal group. The pay-to-stay hypothesis states that there should be a positive relationship between the two; when provisioning goes up, time in the natal community does as well. The research found no such relationship in their population (Koenig & Walters, 2011). This is because individuals participate in Pay-to -to receive indirect fitness via kin selection, not to stay in the community longer.

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Pay-to-stay due to ecological constraints

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Pay-to-stay due to ecological constraints

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Ecological constraints leading to cooperation in the form of Pay-to-stay is evident because helpers in a community (those who do not breed, but support others who do) will leave to bred independently if the resources out weigh the costs.

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Evidence for Pay-to-stay due to ecological constraints

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Evidence for Pay-to-stay due to ecological constraints

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Helpers reduced obedient behavior in the presence of independent breeding options (black circles) (Bergmüller et al, 2005).

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Evidence for Pay-to-stay due to ecological constraints

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Evidence for Pay-to-stay due to ecological constraints

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One study found that all helpers invested less in costly submissive behaviors towards group members. They also found that large helpers (those who helped significantly) reduced help, which supports the Pay-to-Stay hypothesis (Bergmüller et al, 2005). They choose to pay-to -stay when the benefits of staying outweigh the payment to stay. If they cannot breed independently due to ecological constraints then an individual would benefit more by staying in a community and helping raise kin.

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Annotation for "When Should Helpers Pay-To-Stay?" 

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Annotation for "When Should Helpers Pay-To-Stay?" 

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Ecological constraints evidence

Bergmüller, R., Heg, D., & Taborsky, M. (2005). Helpers in a cooperatively breeding cichlid stay and pay or disperse and breed, depending on ecological constraints. Proceedings. Biological sciences, 272(1560), 325–331. https://doi.org/10.1098/rspb.2004.2960

pictures
https://www.audubon.org/field-guide/bird/acorn-woodpecker
https://www.atlasobscura.com/articles/acorn-woodpeckers-hoard-thousands-of-acorns-in-a-single-tree

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Erin Korte
Alexis Huskey