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This is my sandbox, which is used to learn about how to edit Wikipedia.

For Week 4, I added information on foundresses and usurpers to Nest usurpation. --Leah Curnew (talk) 14:07, 28 January 2018 (UTC)

Draft - Allofeeding (Lead Section)

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Please note the draft structure is as follows:

The structure of this draft (article) will be of an upside down triangle and heads/subheads will be used in the essay. It will start with a broad overview of allofeeding (Introductory paragraph), followed by multipliable paragraphs, which will cover examples allofeeding between mates, peer allofeeding and allofeeding during conditions of environmental stress that ties this behaviour to survival and fitness.

*** Note that this structure will change as research continues and that one heading maybe be cut if the headings are denser than expected***

Draft:

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Allofeeding (heading 1- Introduction)
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  • In this section, I will improve the definition of allofeeding. I will include some basic facts about allofeeding, such as the taxonomic groups it is found in, like birds (Smith, 1980) and the different kinds of allofeeding (eg. peer, during conditions of environmental stress, between mates, between parent and non-biological offspring).
  • In this section, I will discuss allofeeding in relation to altruism –research is still in process but peer-reviewing references will be used.
  • I also plan to summarize the main points from heading 2,3 and 4.
Allofeeding between Mates (heading 2)
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  •  For this section, I will talk about the behaviour of allofeeding between mates.
  • In this section, I will explain the behaviour of allofeeding between mates during egg incubation and how this is beneficial (reduced predation and energetic stress of mates) to the fitness of the species using the paper by Halley et al., 2015. I will also discuss the cost of allofeeding between mates.
  • In this section, I will also explain the behaviour of allofeeding between mates during copulation and how this is beneficial for the female, mate bonding, and offspring survival in relation to the fitness of the species using the paper by Smith, 1980.
Peer Allofeeding (heading 3)
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  • In this section, I will also explain the behaviour of peer allofeeding and how it contributes to the survival of their siblings and how this is beneficial for the fitness of the species using the paper by Carlisle, 1986.
Allofeeding and Environmental Stress (heading 4)
[edit]
  • For this section, I will talk about the behaviour of allofeeding during conditions of environmental stress (eg. winter).
  • In this section, I will explain the behaviour of allofeeding during the winter in response to low rate parental survival and how this is beneficial (increased survival of young) to the fitness using the paper by Jouventin et al., 1994 and Lecomte et al., 2006. I will also discuss the cost of allofeeding on parents and their offspring.

References (likely to be used):

  1. Carlisle, T. R., and Zahavi, A. (1986). Helping at the nest, allofeeding and social status in immature arabian babblers. Behavioural Ecology Sociobiology, 18, 339-351.
  2. Halley, M. R., Holmes, A. L., and Robinson, W. D. (2015). Biparental incubation and allofeeding at nests of Sagebrush Brewer's Sparrows. Journal of Field Ornithology, 86(2), 153-162.
  3. Jouventin, P., Barbraud, C., and Rubin, M. (1995). Adoption in the emperor penguin, Aptenodytes fivsteri. Animal Behaviour, 50, 1023-1029.
  4. Lecomte, N., Kuntz, G., Lambert, N., Gendner, J., Handrich, Y.,  Le Maho, Y., and Bost, C. (2006) Alloparental feeding in the king penguin. Animal Behaviour, 71, 457–462.
  5. Senar, J. C. (1984). Allofeeding in Eurasian Siskins (Carduelis spinus). American Ornithological Society, 86(2), 213-214.
  6. Smith, S. M. (1980). Demand Behavior: A New Interpretation of Courtship Feeding. American Ornithological Society, 82(3), 291-295.

Note that a similar version of this draft was posted to the talk page of Allofeeding.

--Leah Curnew (talk) 15:49, 11 February 2018 (UTC)

Article final draft - Allofeeding

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Introduction

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Allofeeding is a type of food sharing behaviour observed in cooperatively breeding species of birds.[1][2][3] Allofeeding refers to a parent, sibling or unrelated adult bird feeding altricial hatchlings, which are dependent on parental care for their survival.[1][4] Allofeeding also refers to food sharing between adults of the same species. Allofeeding can occur between mates during mating rituals, courtship, egg laying or incubation, between peers of the same species, or as a form of parental care.[1][2][3][4][5][6][7][8][9][10][11][12]

Allofeeding evolved for different reasons in different species of birds. Sagebrush Brewer’s Sparrows allofeed their to reduce predation during incubation, Sichuan Jays mate allofeed to increase a female’s nutritional level prior to egg laying and Chinstrap Penguins mate allofeed to strengthen the bond between the pair during chick guarding.[6][7][8] Parental allofeeding is a common form of parental care in many species of birds.[13] The Chinstrap Penguins and Sichuan Jays allofeed their chicks by the regurgitating their food.[7][8] Parental allofeeding is not restricted to biological parents in birds, as seen in King Penguins.[7][9][10] In King Penguins, non-breeders allofeeding chicks is considered an altruistic behaviour. Arabian Babblers peer allofeed to increase social rank while Barn Owls peer allofeed to reduce sibling competition during food sharing.[4][11] In Sichuan Jays, allofeeding is beneficial, as it increases nest survival during the nestling and fledging period.[7] Gray Jays also display allofeeding.[7] Although many species of birds exhibit allofeeding, there are some species that do not perform allofeeding, such as Siberian Jays.[7]

Allofeeding between mates

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Many species of migratory songbirds display allofeeding during the incubation period.[6] During the incubation period, a male songbird will feed its mate, while she sits on the eggs through beak-to-beak interaction.[6] This allofeeding behaviour is suggested to be adaptive because the male is indirectly investing in its offspring.[6] The male uses its energy to forage and retrieve food, and to feed the female.[6] This behaviour of the male reduces nutritional stress of the female because it reduces the amount of time the female spends foraging.[6] In addition, the allofeeding behaviour decreases the number of times a female has to leave the nest, which in turn extends the incubation period and reduces the risk of nest being detected by a predator.[6] Overall, allofeeding behaviour contributes to increased fitness and is therefore considered advantageous.[6] However, Nolan (1958) theorizes that allofeeding is a non-adaptive and is derived from anticipatory parental care.[6]

Allofeeding during the incubation period can also transpire through both the male and the female interchangeably feeding each other via beak-to-beak interactions, while a mate receiving the food incubates the eggs.[6] Once feeding is completed, the recipient now becomes the feeder and the mate that was just foraging incubates the eggs. For example, Sagebrush Brewer’s Sparrows (Spizella breweri breweri) allofeed in this manner.[6] A recent study by Halley et al., 2015, examined allofeeding in twenty-four nests of Sagebrush Brewer’s Sparrows.[6] This study revealed that allofeeding occurred at low frequencies (55%) in nests with biparental incubation and no allofeeding occurred in uniparental nests.[6] This study suggests that allofeeding is an intraspecific signal required to maintain social bonds between mates, in addition to increasing the nutrition levels of females and concealment of the nest.[6] Furthermore, the study found that incubation sessions per hour were higher in biparental nests with allofeeding than in biparental nests without allofeeding.[6] These findings indicate that allofeeding is beneficial because it reduces the risk of predation from visually oriented predators through increased nest concealment[6], which can maximize the fitness of males while increasing the fitness of females.[6]

There have been many documented observations of allofeeding between mates (mate allofeeding).[8] However, in penguins allofeeding between mates is rare.[8] In 2010, Chinstrap Penguins (P. antarctica) allofeeding was reported during the period of chick guarding.[8] The researchers witnessed a male feeding a female, which had a large chick in its nest.[8] The behaviour was engaged by the female frequently pecking the side of the male’s bill to stimulate the male to regurgitate its food.[8] This resulted in the male regurgitating its food into the female’s open mouth. This allofeeding behaviour was identical to the manner in which a chick begs its parent for food.[8] Notably, during this behaviour the female kept the attention of male when the chick tried to gain the male’s attention.[8] It is proposed that mate allofeeding in Chinstrap Penguins was performed to strengthen the bond between the pair.[8]

Allofeeding between mates can also occur during courtship.[8] Courtship allofeeding occurs in half of bird subfamilies and mainly appears in monogamous bird species.[8] Courtship allofeeding is hypothesized to strengthen the bond between pairs or increase a female’s nutritional level before laying eggs.[8] For example, in the Sichuan Jay (P. internigrans), females are fed by only one male during courtship.[7] It is suggested that allofeeding the female during courtship, increases a female’s nutritional level before egg laying.[7] This behaviour was also recorded in the Gray Jays (P. canadensis).[7] Other species of birds display allofeeding during courtship. Additionally, courtship allofeeding in some birds species is hypothesized to strengthen the bond between pairs.[8]

Allofeeding and parental care

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Aptenodytes patagonicus

Figure 1: An Emperor Penguin (Aptenodytes patagonicus) feeding its offspring.

A parent feeding a non-biological chick via bill-to-bill interactions is a form of allofeeding.[10][12] Parental allofeeding occurs in all altricial species of birds to ensure proper growth, development, and to prevent starvation of their offspring.[13] This form of allofeeding has been shown to be a form of parental care in some species, such as in King Penguins (A. patagonicus).[10] King Penguins are pelagic seabirds.[10] King Penguin chicks form dense groups called crèches during winter for social thermoregulation when food availability is low.[10][12] The formation of crèches results from the parents interchangeably leaving starving chicks to gather food from the sea.[10][12] A study by Lecomte et al., 2015 examined the allofeeding behaviour in King Penguins, in which they marked the white underbellies of 74 non-breeding adults and 103 breeding pairs with a unique dark colouring called dye-mark code of Nyanzol-D.[10] Lecomte et al. (2015) found 22% of the marked adults feed 65% of the chicks in crèches, revealing that allofeeding behaviour was most common during the breeding season.[10] Lecomte et al. (2015) also identified that most of the 22% of adult King Penguins demonstrating allofeeding behaviour were unable to successfully breed along with few breeders performing the behaviour.[10] The allofeeding behaviour resulted in more feedings for chicks [10], which is consistent with study by Pierre et al. (1994) that showed that allofeeding in penguins increases the chick growth rate from 35g/day to 190g/day.[12] From these results, Lecomte et al., (2015) concluded that allofeeding increased the survival rate of chicks in crèches.[10][12] The results imply that parental allofeeding in King Penguins is an altruistic behaviour, where allofeeding benefits the young at a cost to the non-biological parent.[10] Notably, no direct benefits to the non-biological parents have been described.[10] Interestingly, the cost of allofeeding non-biological chicks resulted in no detrimental effects to the fitness of the alloparent, because the cost of allofeeding is low (several grams) when compared to the nutritional costs of an adult king penguin, which is several kilograms.[10] However, it is suggested that allofeeding many chicks may cost each chick lost meals that can cause negative effects on the fitness of allofed chicks.[10] In addition, allofeeding decreases when non-biological parents had offspring.[10]

Pseudomonas antarctica

The Chinstrap Penguin (P. antarctica) is another species of penguin that displays allofeeding.[8] Allofeeding in the Chinstrap Penguin has been observed between the female and its offspring.[8] Similar to mate allofeeding in this species, the offspring of a female will peck the side of the female’s bill to signal the female to regurgitate its food into the chick's open mouth.[8]

Perisoreus internigrans

In Sichuan Jays (P. Internigrans), females are not the only sex that participates in the allofeeding of their offspring.[7] During their first week of life, the chicks are only fed by the male.[9] The male forages and stores food in a pouch within its throat. At the nest, the male regurgitates the semi-digested food from its pouch into the mouths of its chicks.[9] After the first week, the male continues to feed the chicks while the female begins feeding the chicks.[7] The female uses the same method as the male.[9]

Peer allofeeding

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Turdoides sqamiceps

There have been many reported cases of allofeeding between siblings in several species of birds.[4] For example, the Arabian Babblers (T. sqamiceps) are a territorial, desert inhibiting species of bird that participate in cooperative breeding.[4] In this species, non-breeders show numerous types of cooperative behaviour, including allofeeding.[4] Adult and immature Babblers, as well as previous and newly fledged Babblers, will frequently engage in allofeeding behaviour.[4]

Carlisle et al. (1986) revealed that Arabian Babblers participate in peer allofeeding in order to increase social rank, which increases fitness.[4] This allofeeding behaviour is supported by the Zahavi's hypothesis.[4] An individual (a dominate Babbler) can increase in social rank by allofed a subordinate Babbler.[4] On the contrary, an individual (a subordinate Babbler) can decrease in social rank when they are allofed by a dominate Babbler.[4] Subordinate Babblers have been observed refusing to be allofeed by dominate Babbler.[4] The refusal resulted in the dominate Babbler becoming aggressive with the subordinate - hitting or chasing the subordinate.[4] Dominate Babblers also showed the same aggressive behaviours when a subordinate tried to allofeed it.[4]

Tyto alba

Another species that exhibits peer allofeeding is the Barn Owl (T. alba).[11] Barn Owl nest siblings establish a hierarchy for the sharing of food resources via vocal negotiations.[11] It is suggested that Barn Owl nest siblings show peer allofeeding to reduce sibling competition in the food sharing hierarchy.[11] Peer allofeeding observed in 60 nestling pairs revealed that the behaviour occurs by a donor sibling placing the food item on the ground in front of the receiver sibling, which transfers the food item to its bill using its talons.[11] Alternatively, the donor sibling uses its bill to place the food item in the bill of the receiver sibling.[11] As well, peer allofeeding behaviour in Barn Owl siblings is proposed to increase the inclusive fitness of the peer performing the allofeeding.[11]

Intraspecific allofeeding

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Perisoreus Internigrans

Figure 2: Gray Jays (Perisoreus canadensis) feeding offspring at the nest.

Intraspecific allofeeding in Sichuan Jays (P. internigrans) is performed during the nestling and fledging period by non-breeders.[7] One to three non-breeders support a nesting breeding pair by feeding the chicks, regurgitating the food from their throat pouches.[7][9] In 2009, Jing et al. determined that two non-breeders contributed to 44% of feedings in the nest of two Sichuan Jays.[7] Jing et al. (2009) suggest that allofeeding behaviour contributes to increased nest survival.[7]

In Sichuan Jays, allofeeding is hypothesized to be evolutionary selected.[7] In this species, two to three non-breeders will allofeed the young of a breeding pair.[7] The large body mass (21% heavier than Siberian Jays) of the Sichuan Jays, and the presence of more than one non-breeder, allows one non-breeder to ward off a predator while another non-breeder protects the young from additional predators.[7] Both the large body mass of the non-breeders and the presence of multiple non-breeders reduces the risk of injury or death when predators are encountered.[7] This reveals that the benefits of allofeeding the young outweighs the cost of injury or death of the non-breeder and/or chick when more than one non-breeder is present.[7] Peer allofeeding has also been observed in the Gray Jay (P. canadensis).[7] In the Gray Jay, non-breeders allofeed the young after fledging.[7]

Interestingly, not all species of Jay allofeed their young, such as the Siberian Jay (P. infaustus).[7] In the Siberian Jays, non-breeders do not allofeed the chicks of a breeding pair.[7] It is predicted that Siberian Jays do not display allofeeding because the cost of predation is too high.[7] Siberian Jays are continuously threatened by a more dangerous predators, the Eurasian Sparrowhawk and Northern Goshawk, then Gray Jays, which are threatened by Eurasian red squirrel.[7] One non-breeder per breeding pair and the small size (21% less body mass than the Sichuan Jays) of the Siberian Jays, in combination with continuous risk of predation, reveals that the non-breeding Siberian Jays are unable to confront predators and protect the nest simultaneously.[7] This indicates that the cost of allofeeding the young is more costly to the non-breeder when one non-breeder is present because an encounter with a predator increases the risk of injury and death of both the young and the non-breeder.[7] This reveals that the benefit of allofeeding the young does not exceed the cost of injury or death of non-breeder and/or young.[7] This is consistent with the predator-avoidance hypothesis, suggesting that the cost of allofeeding in the Siberian Jays prevents allofeeding from being evolutionary selected.[7]

References

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  1. ^ a b c Senar, Juan Carlos (1984). "Allofeeding in Eurasian Siskins (Carduelis spinus)". The Condor. 86 (2): 213–214. doi:10.2307/1367046. JSTOR 1367046.
  2. ^ a b de Kort, Selvino R.; Emery, Nathan J.; Clayton, Nicola S. (2006). "Food sharing in jackdaws, Corvus monedula: what, why and with whom?". Animal Behaviour. 72 (2): 297–304. doi:10.1016/j.anbehav.2005.10.016. S2CID 53160564.
  3. ^ a b Kort, Selvino R. de; Emery, Nathan J.; Clayton, Nicola S. (2003-05-01). "Food offering in jackdaws (Corvus monedula)". Naturwissenschaften. 90 (5): 238–240. doi:10.1007/s00114-003-0419-2. ISSN 0028-1042. PMID 12743708. S2CID 2404878.
  4. ^ a b c d e f g h i j k l m n Carlisle, Tamsie R.; Zahavi, Amotz (1986-04-01). "Helping at the nest, allofeeding and social status in immature arabian babblers". Behavioral Ecology and Sociobiology. 18 (5): 339–351. doi:10.1007/BF00299665. ISSN 0340-5443. S2CID 39397574.
  5. ^ Smith, Susan M. (1980). "Demand Behavior: A New Interpretation of Courtship Feeding". The Condor. 82 (3): 291–295. doi:10.2307/1367395. JSTOR 1367395.
  6. ^ a b c d e f g h i j k l m n o p q r Halley, Matthew R.; Holmes, Aaron L.; Robinson, W. Douglas (2015-06-01). "Biparental incubation and allofeeding at nests of Sagebrush Brewer's Sparrows". Journal of Field Ornithology. 86 (2): 153–162. doi:10.1111/jofo.12098. ISSN 1557-9263.
  7. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae Jing, Yu; Fang, Yun; Strickland, Dan; Lu, Nan; Sun, Yue-Hua (2009). "Alloparenting in the Rare Sichuan Jay (Perisoreus Internigrans) - Cuidado Aloparental en la Especie Rara Perisoreus internigrans". The Condor. 111 (4): 662–667. doi:10.1525/cond.2009.080114. JSTOR 10.1525/cond.2009.080114. S2CID 85099807.
  8. ^ a b c d e f g h i j k l m n o p q r Mori, Yoshihisa; Kokubun, Nobuo; Shin, Hyoung-Chul; Takahashi, Akinori (2010-10-01). "An observation of between-mates feeding behaviour in chick-guarding chinstrap penguins". Polar Biology. 33 (10): 1437–1438. doi:10.1007/s00300-010-0842-8. ISSN 0722-4060. S2CID 39865776.
  9. ^ a b c d e f Blomgren, Arne Blomgren. "Studies of less Familiar birds 162 Siberian Jay - British Birds". British Birds. Retrieved 2018-03-12.
  10. ^ a b c d e f g h i j k l m n o p q Lecomte, Nicolas; Kuntz, Gregoire; Lambert, Nicolas; Gendner, Jean-Paul; Handrich, Yves; Maho, Yvon Le; Bost, Charles-André (2006). "Alloparental feeding in the king penguin". Animal Behaviour. 71 (2): 457–462. doi:10.1016/j.anbehav.2005.07.007. S2CID 53203003.
  11. ^ a b c d e f g h Roulin, A.; Monstiers, B. Des; Ifrid, E.; Silva, A. Da; Genzoni, E.; Dreiss, A. N. (2016-02-01). "Reciprocal preening and food sharing in colour-polymorphic nestling barn owls". Journal of Evolutionary Biology. 29 (2): 380–394. doi:10.1111/jeb.12793. ISSN 1420-9101. PMID 26563617. S2CID 29861997.
  12. ^ a b c d e f Jouventin, Pierre; Barbraud, Christophe; Rubin, Michel (1995). "Adoption in the emperor penguin, Aptenodytes forsteri". Animal Behaviour. 50 (4): 1023–1029. doi:10.1016/0003-3472(95)80102-2. S2CID 53198665.
  13. ^ a b Eggers, Sönke; Griesser, Michael; Ekman, Jan (2005-01-01). "Predator-induced plasticity in nest visitation rates in the Siberian jay (Perisoreus infaustus)". Behavioral Ecology. 16 (1): 309–315. doi:10.1093/beheco/arh163. ISSN 1045-2249.
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