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{{short description|Species of bird}}
{{Short description|Species of bird}}
{{Use dmy dates|date=June 2020}}
{{Use dmy dates|date=June 2020}}
{{Speciesbox
{{speciesbox
| name = Acorn woodpecker
| name = Acorn woodpecker
| status = LC
| status = LC
| status_system = IUCN3.1
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 11 November 2021">{{cite iucn |author=BirdLife International |date=2016 |title=''Melanerpes formicivorus'' |volume=2016 |page=e.T22680813A92880196 |doi=10.2305/IUCN.UK.2016-3.RLTS.T22680813A92880196.en |access-date=11 November 2021}}</ref>
| status_ref = <ref name="iucn status 11 November 2021">{{cite iucn |author=BirdLife International |date=2016 |title=''Melanerpes formicivorus'' |volume=2016 |page=e.T22680813A92880196 |doi=10.2305/IUCN.UK.2016-3.RLTS.T22680813A92880196.en |access-date=11 November 2021}}</ref>
| image =Acorn woodpecker holding a nut in its beak-0225.jpg
| image = Acorn woodpecker holding a nut in its beak-0225.jpg
| image_caption = Male in California, United States
| image_caption = Male in [[California]], [[United States]]
| image2=Melanerpes formicivorus -Madera Canyon, Arizona, USA -female-8-2c.jpg
| image2 = Melanerpes formicivorus -Madera Canyon, Arizona, USA -female-8-2c.jpg
| image2_caption= Female in Arizona, United States
| image2_caption = Female in [[Arizona]], [[United States]]
| genus = Melanerpes
| genus = Melanerpes
| species = formicivorus
| species = formicivorus
| authority = ([[William John Swainson|Swainson]], 1827)
| authority = ([[William John Swainson|Swainson]], 1827)
|range_map = Acorn_Woodpecker_Distribution_Map.png
| range_map = Acorn_Woodpecker_Distribution_Map.png
| range_map_caption = Range of ''M. formicivorus''
| range_map_caption = Range of ''M. formicivorus''
}}
}}

[[File:Acorn woodpeckers on Angel Island (40109).jpg|thumb|Three acorn woodpeckers in [[California]]. Two face opposite directions while one breaks open an acorn.]]
The '''acorn woodpecker''' (''Melanerpes formicivorus'') is a medium-sized [[woodpecker]], {{convert|21|cm|in|abbr=on}} long, with an average weight of {{convert|85|g|oz|abbr=on}}.
The '''acorn woodpecker''' ('''''Melanerpes formicivorus''''') is a medium-sized [[woodpecker]] with a length of around {{convert|20|cm|in|abbr=on|sigfig=1}},<ref>{{Cite encyclopedia |title=Acorn woodpecker {{!}} bird |author= |encyclopedia=Encyclopedia Britannica |date= |access-date=17 April 2024 |url= https://www.britannica.com/animal/acorn-woodpecker}}</ref> and an average weight of {{convert|85|g|oz|abbr=on}}.{{cn|date=April 2024}} It is found across [[Central America]], as well as North into the [[western United States]] and South into parts of [[Colombia]].<ref name="iucn status 11 November 2021"/>


==Taxonomy==
==Taxonomy==
The acorn woodpecker was [[species description|formally described]] in 1827 by the English naturalist [[William John Swainson]] under the [[binomial name]] ''Picus formicivorus'' from a specimen collected in Mexico.<ref>{{ cite journal | last=Swainson | first=William John | author-link=William John Swainson | year=1827 | title=A synopsis of the birds discovered in Mexico by W. Bullock, F.L.S. and Mr. William Bullock jun. | journal=Philosophical Magazine | series=New Series | volume=1 | pages=433–442 [439] | doi=10.1080/14786442708674357 | url=https://www.biodiversitylibrary.org/page/15530449 }}</ref> The specific epithet combines the Latin ''formica'' meaning "ant" with ''-vorus'' meaning "eating".<ref>{{cite book | last=Jobling | first=James A. | year=2010| title=The Helm Dictionary of Scientific Bird Names | publisher=Christopher Helm | location=London | isbn=978-1-4081-2501-4 | page=163 }}</ref> The [[type locality (biology)|type locality]] is [[Temascaltepec]] in Mexico.<ref>{{ cite book | editor-last=Peters | editor-first=James Lee | editor-link=James L. Peters | year=1948 | title=Check-List of Birds of the World | volume=6 | publisher=Harvard University Press | place=Cambridge, Massachusetts | page=159 | url=https://www.biodiversitylibrary.org/page/14477592 }}</ref> The acorn woodpecker is one of 24 species now placed in the [[genus]] ''[[Melanerpes]]'' that was introduced by Swainson in 1832.<ref>{{cite book | last1=Swainson | first1=William John | year=1831 | editor1-last=Richardson | editor1-first=John | editor1-link=John Richardson (naturalist) | title=Fauna boreali-americana, or, The zoology of the northern parts of British America : containing descriptions of the objects of natural history collected on the late northern land expeditions under command of Captain Sir John Franklin, R.N.: Part 2, The Birds | page=316 | url=https://www.biodiversitylibrary.org/page/41549731 }} The title page gives the date as 1831 but the volume was not actually published until the following year.</ref><ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | editor3-last=Rasmussen | editor3-first=Pamela | editor3-link=Pamela Rasmussen | year=2020 | title=Woodpeckers | work=IOC World Bird List Version 10.1 | url=http://www.worldbirdnames.org/bow/woodpeckers/ | publisher=International Ornithologists' Union | access-date=2 October 2020 }}</ref> Within ''Melanerpes'' the acorn woodpecker is [[sister taxon|sister]] to a [[clade]] containing two South American species: the [[white woodpecker]] (''Melanerpes candidus'') and the [[white-fronted woodpecker]] (''Melanerpes cactorum'').<ref>{{Cite journal | last1=Shakya | first1=S.B. | last2=Fuchs | first2=J. | last3=Pons | first3=J.-M. | last4=Sheldon | first4=F.H. | date=2017 | title=Tapping the woodpecker tree for evolutionary insight | journal=Molecular Phylogenetics and Evolution | volume=116 | pages=182–191 | doi=10.1016/j.ympev.2017.09.005 | pmid=28890006 | url=https://www.researchgate.net/publication/319596154 }}</ref>
The acorn woodpecker was [[species description|formally described]] in 1827 by the English naturalist [[William Swainson]] under the [[binomial name]] ''Picus formicivorus'' from a specimen collected in Mexico.<ref>{{ cite journal | last=Swainson | first=William John | author-link=William John Swainson | year=1827 | title=A synopsis of the birds discovered in Mexico by W. Bullock, F.L.S. and Mr. William Bullock jun. | journal=Philosophical Magazine | series=New Series | volume=1 | pages=433–442 [439] | doi=10.1080/14786442708674357 | url=https://www.biodiversitylibrary.org/page/15530449 }}</ref> The specific epithet combines the Latin ''formica'' meaning "ant" with ''-vorus'' meaning "eating".<ref>{{cite book | last=Jobling | first=James A. | year=2010| title=The Helm Dictionary of Scientific Bird Names | publisher=Christopher Helm | location=London | isbn=978-1-4081-2501-4 | page=163 }}</ref> The [[type locality (biology)|type locality]] is [[Temascaltepec]] in Mexico.<ref>{{ cite book | editor-last=Peters | editor-first=James Lee | editor-link=James L. Peters | year=1948 | title=Check-List of Birds of the World | volume=6 | publisher=Harvard University Press | place=Cambridge, Massachusetts | page=159 | url=https://www.biodiversitylibrary.org/page/14477592 }}</ref> The acorn woodpecker is one of 24 species now placed in the [[genus]] ''[[Melanerpes]]'' that was introduced by Swainson in 1832.<ref>{{cite book | last1=Swainson | first1=William John | year=1831 | editor1-last=Richardson | editor1-first=John | editor1-link=John Richardson (naturalist) | title=Fauna boreali-americana, or, The zoology of the northern parts of British America : containing descriptions of the objects of natural history collected on the late northern land expeditions under command of Captain Sir John Franklin, R.N.: Part 2, The Birds | page=316 | url=https://www.biodiversitylibrary.org/page/41549731 }} The title page gives the date as 1831 but the volume was not actually published until the following year.</ref><ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | editor3-last=Rasmussen | editor3-first=Pamela | editor3-link=Pamela Rasmussen | year=2020 | title=Woodpeckers | work=IOC World Bird List Version 10.1 | url=http://www.worldbirdnames.org/bow/woodpeckers/ | publisher=International Ornithologists' Union | access-date=2 October 2020 }}</ref> Within ''Melanerpes'' the acorn woodpecker is [[sister taxon|sister]] to a [[clade]] containing two South American species: the [[white woodpecker]] (''Melanerpes candidus'') and the [[white-fronted woodpecker]] (''Melanerpes cactorum'').<ref>{{Cite journal | last1=Shakya | first1=S.B. | last2=Fuchs | first2=J. | last3=Pons | first3=J.-M. | last4=Sheldon | first4=F.H. | date=2017 | title=Tapping the woodpecker tree for evolutionary insight | journal=Molecular Phylogenetics and Evolution | volume=116 | pages=182–191 | doi=10.1016/j.ympev.2017.09.005 | pmid=28890006 | url=https://www.researchgate.net/publication/319596154 | doi-access=free }}</ref>


Seven [[subspecies]] are recognised:<ref name=ioc/>
Seven [[subspecies]] are recognised:<ref name=ioc/>
* ''M. f. bairdi'' [[John L. Ridgway|Ridgway]], 1881 – Oregon (USA) to north [[Baja California]] (Mexico)
* ''M. f. bairdi'' [[Robert Ridgway|Ridgway]], 1881 – Oregon (USA) to north [[Baja California]] (Mexico)
* ''M. f. angustifrons'' [[Spencer Fullerton Baird|Baird, SF]], 1870 – south Baja California (Mexico)
* ''M. f. angustifrons'' [[Spencer Fullerton Baird|Baird, SF]], 1870 – south Baja California (Mexico)
* ''M. f. formicivorus'' ([[William John Swainson|Swainson]], 1827) – southwest USA to southeast Mexico
* ''M. f. formicivorus'' ([[William John Swainson|Swainson]], 1827) – southwest USA to southeast Mexico
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* ''M. f. striatipectus'' Ridgway, 1874 – Nicaragua to west Panama
* ''M. f. striatipectus'' Ridgway, 1874 – Nicaragua to west Panama
* ''M. f. flavigula'' ([[Alfred Malherbe|Malherbe]], 1849) – Colombia
* ''M. f. flavigula'' ([[Alfred Malherbe|Malherbe]], 1849) – Colombia

<gallery mode = packed heights = 180px>
Acorn woodpecker (Melanerpes formicivorus lineatus) female Copan.jpg|Female ''M. f. lineatus'', Honduras
Acorn woodpecker (Melanerpes formicivorus flavigula) male Las Tangaras 2.jpg|Male ''M. f. flavigula'', Colombia
Acorn woodpecker (Melanerpes formicivorus flavigula) female Las Tangaras 2.jpg|Female ''M. f. flavigula'', Colombia
Acorn woodpecker (Melanerpes formicivorus flavigula) female Las Tangaras.jpg|Female ''M. f. flavigula'', Colombia
</gallery>


==Description==
==Description==
The adult [[acorn]] woodpecker has a brownish-black head, back, wings and tail, white forehead, throat, belly and rump. The eyes are white. There is a small part on the small of their backs where there are some green feathers. The adult male has a red cap starting at the forehead, whereas females have a black area between the forehead and the cap. The white neck, throat, and forehead patches are distinctive identifiers. When flying, they take a few flaps of their wings and drop a foot or so. White circles on their wings are visible when in flight. Acorn woodpeckers have a call that sounds almost like they are laughing.
The adult [[acorn]] woodpecker has a brownish-black head, back, wings and tail, white forehead, throat, belly and rump. The eyes are initially dark in fledglings, turning to white within a few months. There is a small part on the small of their backs where there are some greenish feathers.{{cn|date=April 2024}} The bird is mostly black, with adult males have a red cap starting at the forehead, whereas females have a black area between the forehead and the cap.<ref name="AWI"/> The white neck, throat, and forehead patches are distinctive identifiers. When flying, they take a few flaps of their wings and drop a foot or so. White circles on their wings are visible when in flight. Acorn woodpeckers have a call that sounds almost like they are laughing.


'''Measurements''':<ref>{{Cite web|title=Acorn Woodpecker Identification, All About Birds, Cornell Lab of Ornithology|url=https://www.allaboutbirds.org/guide/Acorn_Woodpecker/id|access-date=2020-09-26|website=www.allaboutbirds.org|language=en}}</ref>
'''Measurements''':<ref name="AWI">{{Cite web|title=Acorn Woodpecker Identification, All About Birds, Cornell Lab of Ornithology|url=https://www.allaboutbirds.org/guide/Acorn_Woodpecker/id|access-date=2020-09-26|website=www.allaboutbirds.org|language=en}}</ref>


* '''Length''': {{Convert|19–23|cm|in|abbr=on}}
* '''Length''': {{Convert|19–23|cm|in|abbr=on}}
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=== Distribution and habitat ===
=== Distribution and habitat ===
The acorn woodpecker's habitat is forested areas with [[oak]]s in the coastal areas and foothills of [[Oregon]], [[California]], and the [[southwestern United States]], south through [[Central America]] to [[Colombia]].<ref>Scofield, D., Alfaro, V., Sork, V., Grivet, D., Martinez, E., Papp, J., Pluess, A., Koenig, W., and Smouse, E. (2010). [https://link.springer.com/article/10.1007/s00442-010-1828-5 Foraging patterns of acorn woodpeckers (''Melanerpes formicivorus'') on valley oak (''Quercus lobata'' Nee) in two California oak savanna-woodlands]. ''Oecologia'' 166:187–196.</ref> This species may occur at low elevations in the north of its range, but rarely below {{convert|1000|m|ft|abbr=on}} in [[Central America]], and it breeds up to the timberline. Nests are excavated in a large cavity in a dead tree or a dead part of a tree.
The acorn woodpecker's habitat is forested areas with [[oak]]s in the coastal areas and foothills of [[Oregon]], [[California]], and the [[southwestern United States]], south through [[Central America]] to [[Colombia]].<ref>{{cite journal |last1=Scofield |first1=Douglas G. |last2=Alfaro |first2=Victor Ryan |last3=Sork |first3=Victoria L. |last4=Grivet |first4=Delphine |last5=Martinez |first5=Edith |last6=Papp |first6=Jeannette |last7=Pluess |first7=Andrea R. |last8=Koenig |first8=Walter D. |last9=Smouse |first9=Peter E. |title=Foraging patterns of acorn woodpeckers (''Melanerpes formicivorus'') on valley oak (''Quercus lobata'' Née) in two California oak savanna-woodlands |journal=Oecologia |date=2011 |volume=166 |issue=1 |pages=187–196 |doi=10.1007/s00442-010-1828-5|pmid=21107869 |pmc=3074073 |bibcode=2011Oecol.166..187S |hdl=10261/291553 |hdl-access=free }}</ref> This species may occur at low elevations in the north of its range, but rarely below {{convert|1000|m|ft|abbr=on}} in [[Central America]], and it breeds up to the timberline. Nests are excavated in a cavity in a dead tree or a dead part of a tree.


==Behaviour and ecology==
==Behaviour and ecology==
===Breeding===
===Breeding===
Field studies have shown that breeding groups range from monogamous pairs to [[breeding collective]]s, sometimes called "coalitions". Both types of breeding behaviors have been observed in ''Melanerpes formicivorus'' and within same populations. This complex behavior purposes that the woodpecker may undergo social plasticity<ref>{{Cite journal |last=Stacey |first=P |last2=Bock |first2=C |date=1978 |title=Social plasticity in the acorn woodpecker |url=https://www.jstor.org/stable/pdf/1747051.pdf |journal=Science |volume=202 |pages=1298-1300 |doi=10.1126/science.202.4374.1298 |via=JSTOR}}</ref>. Cooperative breeding, defined as more than two birds taking care of nestlings in the nest, is a relatively rare evolutionary trait that is thought to occur in only nine percent of bird species.<ref>Cockburn, A. (2006). [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1560291/ Prevalence of different modes of parental care in birds]. ''Proceedings of the Royal Society B Biological Sciences'' 273:1375-1383.</ref> Cooperative breeding occurs in two ways: coalitions and family groups. Coalitions of adult acorn woodpeckers nest together, localizing to storage granaries.<ref name="Koenig">Koenig, W.D. (1981). Reproductive success, group size, and the evolution of cooperative breeding in the acorn woodpecker. ''American Naturalist'' 22:505–523.</ref> Additionally, adult offspring often stay in their parents' nest and help raise the next generation of woodpeckers.<ref name="Hatch">Hatchwell, B.J., and J. Komdeur. (2000). [http://faculty.bennington.edu/~sherman/animal%20social%20beh/coop/HatchwellKomdeur2000Coop.pdf Ecological constraints, life history traits and the evolution of cooperative breeding]. ''Animal Behaviour'' 59:1079-1086.</ref> It is generally believed that limited territories drive cooperative breeding behavior in birds, and in the case of the acorn woodpecker, this limited territory is the acorn storage granary.<ref name="Hatch" /> Social monogamous acorn woodpecker females breeding outside of their group was a rare sight as reported in the mid-1990's which gave rise to the idea that indirect fitness benefits to the cobreeders and helpers reduce if extra-pair copulations occur<ref>{{Cite journal |last=Dickinson |first=J |last2=Haydock |first2=J |last3=Koenig |first3=W |last4=Stanback |first4=M |last5=Pitelka |first5=F |date=1995 |title=Genetic monogamy in single-male groups of acorn woodpeckers, Melanerpes formicivorus |url=https://www.researchgate.net/publication/229876276_Genetic_monogamy_in_single-male_groups_of_acorn_woodpeckers_Melanerpes_formicivorus |journal=Molecular Ecology |volume=4 |pages=765-769 |doi=10.1111/j.1365-294X.1995.tb00277.x}}</ref>. This further drives the suggestion for favoring cooperative breeding.
Acorn woodpeckers are cooperative breeders, living and breeding in family groups of up to 15 individuals. Field studies have shown that within the same population, groups range from monogamous pairs to [[breeding collective|polygynandrous breeding collective]]s consisting of coalitions of up to 8 males and 4 females, along with nonbreeding "[[helpers at the nest]]" that are offspring from prior breeding events. Regardless of composition, all breeder males (who are usually brothers or fathers and their sons) compete for matings with all breeder females (who are sisters or a mother and her daughter), the latter of which lay their eggs communally in the same nest cavity. There is considerable variability within and among populations, suggesting extraordinary social plasticity.<ref>{{Cite journal |last1=Stacey |first1=P |last2=Bock |first2=C |date=1978 |title=Social plasticity in the acorn woodpecker |url=https://www.jstor.org/stable/pdf/1747051.pdf |journal=Science |volume=202 |issue=4374 |pages=1298–1300 |doi=10.1126/science.202.4374.1298 |jstor=1747051 |pmid=17750490 |bibcode=1978Sci...202.1298S |s2cid=34467844 }}</ref> Cooperative breeding, defined as more than two birds taking care of nestlings in the nest, is a relatively rare evolutionary trait that is thought to occur in only nine percent of bird species.<ref>{{cite journal |last1=Cockburn |first1=Andrew |title=Prevalence of different modes of parental care in birds |journal=Proceedings of the Royal Society B: Biological Sciences |date=2006 |volume=273 |issue=1592 |pages=1375–1383 |doi=10.1098/rspb.2005.3458 |pmid=16777726 |pmc=1560291}}</ref> Most cooperative breeding species have helpers at the nest, but acorn woodpeckers are unusual in exhibiting both helping at the nest and cooperative polygamy (polygynandry). It is generally believed that limited territories are a key driver of cooperative breeding behavior in birds, and in the case of the acorn woodpecker, the availability of acorn storage granaries (see below) is a key limited resource.<ref name="Hatch">{{cite journal |last1=Hatchwell |first1=B.J. |last2=Komdeur |first2=J. |title=Ecological constraints, life history traits and the evolution of cooperative breeding |journal=Animal Behaviour |date=2000 |volume=59 |issue=6 |pages=1079–1086 |doi=10.1006/anbe.2000.1394 |pmid=10877885 |url=http://faculty.bennington.edu/~sherman/animal%20social%20beh/coop/HatchwellKomdeur2000Coop.pdf}}</ref>


Breeding coalitions consist of up to eight cobreeding males and up to four joint-nesting females. However, most nests consist of only a single breeder female and 1 to 3 cobreeder males.<ref name="Koenig">{{cite journal |last1=Koenig |first1=Walter D. |title=Reproductive success, group size, and the evolution of cooperative breeding in the acorn woodpecker |journal=American Naturalist |date=1981 |volume=117 |issue=4 |pages=421–443 |doi=10.1086/283726 |jstor=10.1086/283726 }}</ref> Nesting groups can also contain up to ten offspring [[helpers at the nest|helpers]].<ref>{{cite journal |last1=Haydock |first1=Joseph |last2=Koenig |first2=Walter D. |title=Reproductive skew in the polygynandrous acorn woodpecker |journal=Proceedings of the National Academy of Sciences |date=2002 |volume=99 |issue=10 |pages=7178–7183 |doi=10.1073/pnas.102624199|doi-access=free |pmid=11997473 |pmc=124548 |bibcode=2002PNAS...99.7178H }}</ref> As mentioned above, the breeder males are often brothers, and the females are usually sisters. However, reproductive vacancies—formed when all the breeders of one sex die—are filled by unrelated birds from elsewhere, so inbreeding is rare, despite the high degree of relatedness among most group members.<ref>{{cite journal |last1=Koenig |first1=Walter D. |last2=Haydock |first2=Joseph |last3=Stanback |first3=Mark T. |title=Reproductive roles in the cooperatively breeding Acorn Woodpecker: incest avoidance versus reproductive competition |journal=American Naturalist |date=1998 |volume=151 |issue=3 |pages=243–255 |doi=10.1086/286115 |pmid=18811355 |jstor=286115 }} {{Webarchive|url=https://web.archive.org/web/20170925230401/http://pages.nbb.cornell.edu/wkoenig/wicker/nb4340/k086ta_98.pdf|date=25 September 2017}}</ref>


In groups with more than one breeding female, the females lay their eggs in a single nest cavity. A female usually destroys any eggs in the nest before she starts to lay. Once all the females start to lay, they stop removing eggs.<ref>{{cite journal |last1=Koenig |first1=Walter D. |last2=Mumme |first2=Ronald L. |last3=Stanback |first3=Mark T. |last4=Pitelka |first4=Frank A. |title=Patterns and consequences of egg destruction among joint-nesting acorn woodpeckers |journal=Animal Behaviour |date=1995 |volume=50 |issue=3 |pages=607–621 |doi=10.1016/0003-3472(95)80123-5}}</ref> Although multiple paternity and maternity are common within groups containing multiple cobreeders, no extra-group paternity has been detected.<ref>{{cite journal |last1=Joste |first1=Nancy |last2=Ligon |first2=J. David |last3=Stacey |first3=Peter B. |title=Shared paternity in the acorn woodpecker (''Melanerpes formicivorus'') |journal=Behavioral Ecology and Sociobiology |date=1985 |volume=17 |issue=1 |pages=39–41 |doi=10.1007/BF00299426 |jstor=4599801 }}</ref>

Breeding coalitions consist of up to seven co-breeding males and up to three joint-nesting females. However, most nests are made up of only three males and two females.<ref name="Koenig" /> Nesting groups can also contain up to ten offspring [[helpers at the nest|helpers]].<ref>Haydock, J., and W.D Koenig. (2002). [http://www.pnas.org/content/99/10/7178.full Reproductive skew in the polygynandrous Acorn Woodpecker]. ''American Naturalist'' 162:277–289.</ref> These breeding coalitions are typically closely related. The males are often brothers, and the females are usually sisters. Inbreeding is rare, however, meaning that co-breeders of the opposite sex are almost never related.<ref>Koenig, W.D., Haydock, J., and M. Stanbeck. (1998). [http://pages.nbb.cornell.edu/wkoenig/wicker/nb4340/k086ta_98.pdf Reproductive roles in the cooperatively breeding Acorn Woodpecker: incest avoidance versus reproductive competition] {{Webarchive|url=https://web.archive.org/web/20170925230401/http://pages.nbb.cornell.edu/wkoenig/wicker/nb4340/k086ta_98.pdf|date=25 September 2017}}. ''American Naturalist'' 151:243–255.</ref>

In groups with more than one breeding female, the females put their eggs into a single nest cavity. A female usually destroys any eggs in the nest before she starts to lay. Once all the females start to lay, they stop removing eggs.<ref>Koenig, W.D., Mumme, R.L., Stanback, M., and F.A. Pitelka. (1995). Patterns and consequences of egg destruction among joint-nesting acorn woodpeckers. ''Animal Behaviour'' 50:607–621.</ref> Young from a single brood have been found with multiple paternity.<ref>Joste, N., Ligon, D., and Stacey, P. (1985). Shared paternity in the Acorn Woodpecker. ''Behavioral Ecology and Sociobiology'' '''17'''(1):39-41.</ref>

[[File:Acorn Woodpecker with Hoard.jpg|thumb|upright|right|Male acorn woodpecker with "granary tree" full of acorns]]
[[File:Acorn.jpg|thumb|upright|right|Acorn hoarded by acorn woodpecker]]


===Food and feeding===
===Food and feeding===
Acorn woodpeckers, as their name implies, depend heavily on [[acorn]]s for food. Acorns are such an important resource to the California populations that acorn woodpeckers may nest in the fall to take advantage of the fall acorn crop, a rare behavior in birds.<ref>Koenig, W.D., and Stahl, J.T. (2007). ''Condor'' '''109'''(2):334-350.</ref> Acorn storage capacity, within the various trees they visit, range from 0 to 5,000 individual holes per tree. Data has shown that access to acorn crops influences the composition of acorn woodpecker communities. In 1976 there was about 90% of non-breeding adults per social unit which was the same year there was poor acorn crop. The following year, 1977, there was a significant increase in acorn production and a correlating decrease in non-breeding adults per unit (which reduced to 20%). <ref>{{Cite journal |last=Trail |first=Pepper W |date=1980 |title=Ecological Correlates of Social Organization in a Communally Breeding Bird, the Acorn Woodpecker, Melanerpes formicivorus |url=https://link.springer.com/article/10.1007/BF00299512 |journal=Behavioral Ecology and Sociobiology |volume=7 |pages=83-92 |doi=10.1007/BF00299512 |via=Springer Link}}</ref>
Acorn woodpeckers, as their name implies, depend heavily on [[acorn]]s for food. Acorns are such an important resource to the California populations that acorn woodpeckers may nest in the fall to take advantage of the fall acorn crop, a rare behavior in birds.<ref>Koenig, W.D., and Stahl, J.T. (2007). ''Condor'' '''109'''(2):334–350.</ref> Acorns are stored in small holes drilled especially for this purpose in "granaries" or "storage trees"—usually snags, dead branches, utility poles, or wooden buildings. Storage holes—always in dead tissue such as bark or dead limbs—are used year after year, and granaries can consist of thousands of holes, each of which may be filled by an acorn in the autumn. Access to acorn crops influences the composition of acorn woodpecker communities. In one study in Old Mexico, there were about 90% of non-breeding adults per social unit in 1976, a year of a poor acorn crop. The following year, 1977, there was a significant increase in acorn production and a correlating decrease in non-breeding adults per unit.<ref>{{Cite journal |last=Trail |first=Pepper W |date=1980 |title=Ecological Correlates of Social Organization in a Communally Breeding Bird, the Acorn Woodpecker, ''Melanerpes formicivorus'' |url=https://link.springer.com/article/10.1007/BF00299512 |journal=Behavioral Ecology and Sociobiology |volume=7 |issue=2 |pages=83–92 |doi=10.1007/BF00299512 |s2cid=11335989 |via=Springer Link}}</ref>

Acorn woodpeckers also feed on insects, [[sap]], and fruit. They can be seen [[Hawking (birds)|sallying]] from tree limbs to catch insects, eating fruit and seeds, and drilling holes to drink sap.


Although acorns are an important back-up food resource, acorn woodpeckers primarily feed on insects, [[sap]], and fruit. They can be seen [[Hawking (birds)|sallying]] from tree limbs to catch insects, eating fruit and seeds, and drilling holes to drink sap.
In some parts of their range, such as California, the woodpeckers create [[granary|granaries]] or "acorn trees" by drilling holes in dead trees, dead branches, telephone poles, and wooden buildings. They also drill holes in the thick bark of mature living trees, notably the Ponderosa Pine in California. These holes, always above the snow line so that the acorns can be retrieved in winter, can be observed in the hundreds on large trees. They do not harm the tree.


The woodpeckers then collect acorns and find a hole that is just the right size for the acorn. As acorns dry out, they are moved to smaller holes and granary maintenance requires a significant amount of the bird's time. The acorns are visible, and a group defends its granary against potential cache robbers like [[Steller's jay]]s and [[western scrub jay]]s. When any protective group of woodpeckers experiences a death or disruption to the hierarchy, nearby birds rush to the area and fight for access to the trove; these fights also attract woodpecker audiences who leave their own territories to witness the battles.<ref>{{Cite journal|last1=Barve|first1=Sahas|last2=Lahey|first2=Ally S.|last3=Brunner|first3=Rebecca M.|last4=Koenig|first4=Walter D.|last5=Walters|first5=Eric L.|date=2020-09-07|title=Tracking the warriors and spectators of acorn woodpecker wars|url=https://www.cell.com/current-biology/abstract/S0960-9822(20)31098-8|journal=Current Biology|language=en|volume=30|issue=17|pages=R982–R983|doi=10.1016/j.cub.2020.07.073|issn=0960-9822|pmid=32898494|s2cid=221522122}}</ref>
The woodpeckers then collect acorns and find a hole that is just the right size for the acorn. As acorns dry out, they are moved to smaller holes and granary maintenance requires a significant amount of the bird's time. The acorns are visible, and a group defends its granary against potential cache robbers like [[Steller's jay]]s and [[western scrub jay|western scrub-jay]]s.


In parts of its range the acorn woodpecker does not construct a "granary tree", but instead stores acorns in natural holes and cracks in [[Bark (botany)|bark]]. If the stores are eaten, the woodpecker will move to another area, even going from [[Arizona]] to [[Mexico]] to spend the winter.
In some more tropical parts of its range the acorn woodpecker does not construct a "granary tree", but instead stores acorns in natural holes and cracks in [[Bark (botany)|bark]]. If the acorn crop is poor and birds cannot find enough to store, the woodpeckers will move to other areas over the winter.
<gallery mode = packed heights = 180px>
Acorn Woodpecker with Hoard.jpg|Male with "granary tree" full of acorns
Acorn.jpg|Hoarded acorn
Acorn woodpeckers on Angel Island (40109).jpg|Three woodpeckers in [[California]]. One breaks open an acorn.
</gallery>


==Threats and status==
==Threats and status==
Line 71: Line 76:


==Popular culture==
==Popular culture==
[[Walter Lantz]] is believed to have patterned the call of his cartoon character [[Woody Woodpecker]] on that of the acorn woodpecker, while patterning his appearance on that of the [[pileated woodpecker]], which has a prominent crest.<ref>{{cite web|url=https://www.npr.org/templates/story/story.php?storyId=101665227 |title=Woody The Acorn (Not Pileated) Woodpecker |publisher=NPR |access-date=2011-09-14}}</ref>
[[Walter Lantz]] is believed to have patterned the call of his cartoon character [[Woody Woodpecker]] on that of the acorn woodpecker, while patterning his appearance on that of the [[pileated woodpecker]], which has a prominent crest.<ref>{{cite web|url=https://www.npr.org/templates/story/story.php?storyId=101665227 |title=Woody The Acorn (Not Pileated) Woodpecker |publisher=[[NPR]] |access-date=2011-09-14 |date=March 10, 2009 |first=Julie |last=Zickefoose}}</ref>


==References==
==References==
Line 77: Line 82:


== Further reading ==
== Further reading ==
{{refbegin}}
{{Refbegin}}
*{{Cite journal | last1=Haydock | first1=J. | last2=Koenig | first2=W.D. | last3=Stanback | first3=M.T. | date=2001 | title=Shared parentage and incest avoidance in the cooperatively breeding acorn woodpecker | journal=Molecular Ecology | volume=10 | issue=6 | pages=1515–1525 | doi=10.1046/j.1365-294X.2001.01286.x | pmid=11412372 | s2cid=21904045 | ref=none}}
*{{Cite journal | last1=Haydock | first1=J. | last2=Koenig | first2=W.D. | last3=Stanback | first3=M.T. | date=2001 | title=Shared parentage and incest avoidance in the cooperatively breeding acorn woodpecker | journal=Molecular Ecology | volume=10 | issue=6 | pages=1515–1525 | doi=10.1046/j.1365-294X.2001.01286.x | pmid=11412372 | bibcode=2001MolEc..10.1515H | s2cid=21904045 | ref=none}}
*{{ cite book | last=Skutch | first=Alexander F. | author-link=Alexander Skutch | date=1969 | chapter=Acorn woodpecker | title=Life Histories of Central American Birds III: Families Cotingidae, Pipridae, Formicariidae, Furnariidae, Dendrocolaptidae, and Picidae | series=Pacific Coast Avifauna, Number 35 | location=Berkeley, California | publisher=Cooper Ornithological Society | chapter-url=https://sora.unm.edu/sites/default/files/journals/pca/pca_035.pdf#page=522 | pages=522–531 | ref=none }}
*{{ cite book | last=Skutch | first=Alexander F. | author-link=Alexander Skutch | date=1969 | chapter=Acorn woodpecker | title=Life Histories of Central American Birds III: Families Cotingidae, Pipridae, Formicariidae, Furnariidae, Dendrocolaptidae, and Picidae | series=Pacific Coast Avifauna, Number 35 | location=Berkeley, California | publisher=Cooper Ornithological Society | chapter-url=https://sora.unm.edu/sites/default/files/journals/pca/pca_035.pdf#page=522 | pages=522–531 | ref=none }}
*{{ cite book | last1=Stiles | first1=F. Gary | last2=Skutch | first2=Alexander F. | author2-link=Alexander Skutch | year=1989 | title=A Guide to the Birds of Costa Rica | location=Ithaca, NY | publisher=Cornell University | isbn=978-0-8014-9600-4 | page=252 | ref=none }}
*{{ cite book | last1=Stiles | first1=F. Gary | last2=Skutch | first2=Alexander F. | author2-link=Alexander Skutch | year=1989 | title=A Guide to the Birds of Costa Rica | location=Ithaca, NY | publisher=Cornell University | isbn=978-0-8014-9600-4 | page=252 | ref=none }}
{{refend}}
{{Refend}}


==External links==
==External links==
{{Commons category|Melanerpes formicivorus|the acorn woodpecker}}
{{sisterlinks|d=Q368657|c=category:Melanerpes formicivorus|species=Melanerpes formicivorus|n=no|b=no|v=no|voy=no|s=no|q=no|m=no|mw=no|f=no}}
* [https://www.allaboutbirds.org/guide/Acorn_Woodpecker/ Acorn woodpecker Species Account] – Cornell Lab of Ornithology
{{Wikispecies|Melanerpes formicivorus}}
* [http://ca.geocities.com/woodpeck2006/acorn.html Acorn woodpecker]( {{Webarchive|url=https://web.archive.org/web/20070311101427/http://ca.geocities.com/woodpeck2006/acorn.html |date=11 March 2007 }} 2009-10-24), a bibliographic resource
* [http://www.birds.cornell.edu/AllAboutBirds/BirdGuide/Acorn_Woodpecker.html Acorn woodpecker Species Account] – Cornell Lab of Ornithology
* [http://ca.geocities.com/woodpeck2006/acorn.html Acorn woodpecker] ([https://www.webcitation.org/5km8YnSYP Archived] 2009-10-24), a bibliographic resource
* [http://www.mbr-pwrc.usgs.gov/id/framlst/i4070id.html Acorn woodpecker - ''Melanerpes formicivorus''] - USGS Patuxent Bird Identification InfoCenter
* [http://www.mbr-pwrc.usgs.gov/id/framlst/i4070id.html Acorn woodpecker - ''Melanerpes formicivorus''] - USGS Patuxent Bird Identification InfoCenter
* {{InternetBirdCollection|acorn-woodpecker-melanerpes-formicivorus|Acorn woodpecker}}
* {{InternetBirdCollection|acorn-woodpecker-melanerpes-formicivorus|Acorn woodpecker}}
* [http://www.bird-stamps.org/cspecies/10403800.htm Stamps] (for [[El Salvador]], [[Mexico]]) with Range Map at bird-stamps.org
* {{usurped|1=[https://web.archive.org/web/20001215185200/http://www.bird-stamps.org/cspecies/10403800.htm Stamps]}} (for [[El Salvador]], [[Mexico]]) with Range Map at bird-stamps.org
* {{VIREO|Acorn+Woodpecker|Acorn woodpecker}}
* {{VIREO|Acorn+Woodpecker|Acorn woodpecker}}
* [http://digitalmedia.fws.gov/cdm/search/searchterm/acorn%20woodpecker/order/nosort Acorn woodpecker] at the US Fish & Wildlife Service Digital Repository
* [http://digitalmedia.fws.gov/cdm/search/searchterm/acorn%20woodpecker/order/nosort Acorn woodpecker] at the US Fish & Wildlife Service Digital Repository


{{Taxonbar|from=Q368657}}
{{Taxonbar|from=Q368657}}
{{Authority control}}


{{DEFAULTSORT:woodpecker, acorn}}
[[Category:Melanerpes|acorn woodpecker]]
[[Category:Melanerpes|acorn woodpecker]]
[[Category:Fauna of the California chaparral and woodlands]]
[[Category:Fauna of the California chaparral and woodlands]]
[[Category:Native birds of the Southwestern United States|Woodpecker, Acorn]] <!-- also endemic(a Vagrant according to "www.natureserve.org") in small areas of western Canada, Brit Col, etc -->
[[Category:Native birds of the Southwestern United States]] <!-- also endemic (a Vagrant according to "www.natureserve.org") in small areas of western Canada, Brit Col, etc -->
[[Category:Birds of Mexico]]
[[Category:Birds of Mexico]]
[[Category:Birds of the Sierra Madre Occidental]]
[[Category:Birds of the Sierra Madre Occidental]]
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[[Category:Birds of Central America]]
[[Category:Birds of Central America]]
[[Category:Birds of El Salvador]]<!-- Because of the Stamp issued-->
[[Category:Birds of El Salvador]]<!-- Because of the Stamp issued-->
[[Category:Birds of Colombia|Woodpecker, Acorn]]
[[Category:Birds of Colombia]]
[[Category:Birds described in 1827|acorn woodpecker]]
[[Category:Birds described in 1827|acorn woodpecker]]
[[Category:Taxa named by William John Swainson]]
[[Category:Acorns]]

Revision as of 02:02, 3 January 2025

Acorn woodpecker
Male in California, United States
Female in Arizona, United States
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Piciformes
Family: Picidae
Genus: Melanerpes
Species:
M. formicivorus
Binomial name
Melanerpes formicivorus
(Swainson, 1827)
Range of M. formicivorus

The acorn woodpecker (Melanerpes formicivorus) is a medium-sized woodpecker with a length of around 20 cm (8 in),[2] and an average weight of 85 g (3.0 oz).[citation needed] It is found across Central America, as well as North into the western United States and South into parts of Colombia.[1]

Taxonomy

The acorn woodpecker was formally described in 1827 by the English naturalist William Swainson under the binomial name Picus formicivorus from a specimen collected in Mexico.[3] The specific epithet combines the Latin formica meaning "ant" with -vorus meaning "eating".[4] The type locality is Temascaltepec in Mexico.[5] The acorn woodpecker is one of 24 species now placed in the genus Melanerpes that was introduced by Swainson in 1832.[6][7] Within Melanerpes the acorn woodpecker is sister to a clade containing two South American species: the white woodpecker (Melanerpes candidus) and the white-fronted woodpecker (Melanerpes cactorum).[8]

Seven subspecies are recognised:[7]

  • M. f. bairdi Ridgway, 1881 – Oregon (USA) to north Baja California (Mexico)
  • M. f. angustifrons Baird, SF, 1870 – south Baja California (Mexico)
  • M. f. formicivorus (Swainson, 1827) – southwest USA to southeast Mexico
  • M. f. albeolus Todd, 1910 – east Chiapas (southeast Mexico) to Belize and northeast Guatemala
  • M. f. lineatus (Dickey & Van Rossem, 1927) – Chiapas (south Mexico) to north Nicaragua
  • M. f. striatipectus Ridgway, 1874 – Nicaragua to west Panama
  • M. f. flavigula (Malherbe, 1849) – Colombia

Description

The adult acorn woodpecker has a brownish-black head, back, wings and tail, white forehead, throat, belly and rump. The eyes are initially dark in fledglings, turning to white within a few months. There is a small part on the small of their backs where there are some greenish feathers.[citation needed] The bird is mostly black, with adult males have a red cap starting at the forehead, whereas females have a black area between the forehead and the cap.[9] The white neck, throat, and forehead patches are distinctive identifiers. When flying, they take a few flaps of their wings and drop a foot or so. White circles on their wings are visible when in flight. Acorn woodpeckers have a call that sounds almost like they are laughing.

Measurements:[9]

  • Length: 19–23 cm (7.5–9.1 in)
  • Weight: 65–90 g (2.3–3.2 oz)
  • Wingspan: 35–43 cm (14–17 in)

Distribution and habitat

The acorn woodpecker's habitat is forested areas with oaks in the coastal areas and foothills of Oregon, California, and the southwestern United States, south through Central America to Colombia.[10] This species may occur at low elevations in the north of its range, but rarely below 1,000 m (3,300 ft) in Central America, and it breeds up to the timberline. Nests are excavated in a cavity in a dead tree or a dead part of a tree.

Behaviour and ecology

Breeding

Acorn woodpeckers are cooperative breeders, living and breeding in family groups of up to 15 individuals. Field studies have shown that within the same population, groups range from monogamous pairs to polygynandrous breeding collectives consisting of coalitions of up to 8 males and 4 females, along with nonbreeding "helpers at the nest" that are offspring from prior breeding events. Regardless of composition, all breeder males (who are usually brothers or fathers and their sons) compete for matings with all breeder females (who are sisters or a mother and her daughter), the latter of which lay their eggs communally in the same nest cavity. There is considerable variability within and among populations, suggesting extraordinary social plasticity.[11] Cooperative breeding, defined as more than two birds taking care of nestlings in the nest, is a relatively rare evolutionary trait that is thought to occur in only nine percent of bird species.[12] Most cooperative breeding species have helpers at the nest, but acorn woodpeckers are unusual in exhibiting both helping at the nest and cooperative polygamy (polygynandry). It is generally believed that limited territories are a key driver of cooperative breeding behavior in birds, and in the case of the acorn woodpecker, the availability of acorn storage granaries (see below) is a key limited resource.[13]

Breeding coalitions consist of up to eight cobreeding males and up to four joint-nesting females. However, most nests consist of only a single breeder female and 1 to 3 cobreeder males.[14] Nesting groups can also contain up to ten offspring helpers.[15] As mentioned above, the breeder males are often brothers, and the females are usually sisters. However, reproductive vacancies—formed when all the breeders of one sex die—are filled by unrelated birds from elsewhere, so inbreeding is rare, despite the high degree of relatedness among most group members.[16]

In groups with more than one breeding female, the females lay their eggs in a single nest cavity. A female usually destroys any eggs in the nest before she starts to lay. Once all the females start to lay, they stop removing eggs.[17] Although multiple paternity and maternity are common within groups containing multiple cobreeders, no extra-group paternity has been detected.[18]

Food and feeding

Acorn woodpeckers, as their name implies, depend heavily on acorns for food. Acorns are such an important resource to the California populations that acorn woodpeckers may nest in the fall to take advantage of the fall acorn crop, a rare behavior in birds.[19] Acorns are stored in small holes drilled especially for this purpose in "granaries" or "storage trees"—usually snags, dead branches, utility poles, or wooden buildings. Storage holes—always in dead tissue such as bark or dead limbs—are used year after year, and granaries can consist of thousands of holes, each of which may be filled by an acorn in the autumn. Access to acorn crops influences the composition of acorn woodpecker communities. In one study in Old Mexico, there were about 90% of non-breeding adults per social unit in 1976, a year of a poor acorn crop. The following year, 1977, there was a significant increase in acorn production and a correlating decrease in non-breeding adults per unit.[20]

Although acorns are an important back-up food resource, acorn woodpeckers primarily feed on insects, sap, and fruit. They can be seen sallying from tree limbs to catch insects, eating fruit and seeds, and drilling holes to drink sap.

The woodpeckers then collect acorns and find a hole that is just the right size for the acorn. As acorns dry out, they are moved to smaller holes and granary maintenance requires a significant amount of the bird's time. The acorns are visible, and a group defends its granary against potential cache robbers like Steller's jays and western scrub-jays.

In some more tropical parts of its range the acorn woodpecker does not construct a "granary tree", but instead stores acorns in natural holes and cracks in bark. If the acorn crop is poor and birds cannot find enough to store, the woodpeckers will move to other areas over the winter.

Threats and status

Acorn woodpeckers, like many other species, are threatened by habitat loss and degradation. Competition for nest cavities by non-native species is an ongoing threat in urbanized areas. Conservation of this species is dependent on the maintenance of functional ecosystems that provide the full range of resources upon which the species depends. These include mature forests with oaks capable of producing large mast crops and places for the woodpeckers to nest, roost, and store mast. Residents are encouraged to preserve mature oak and pine-oak stands of trees and to provide dead limbs and snags for nesting, roosting, and granary sites to help preserve the acorn woodpecker's population.

Walter Lantz is believed to have patterned the call of his cartoon character Woody Woodpecker on that of the acorn woodpecker, while patterning his appearance on that of the pileated woodpecker, which has a prominent crest.[21]

References

  1. ^ a b BirdLife International (2016). "Melanerpes formicivorus". IUCN Red List of Threatened Species. 2016: e.T22680813A92880196. doi:10.2305/IUCN.UK.2016-3.RLTS.T22680813A92880196.en. Retrieved 11 November 2021.
  2. ^ "Acorn woodpecker | bird". Encyclopedia Britannica. Retrieved 17 April 2024.
  3. ^ Swainson, William John (1827). "A synopsis of the birds discovered in Mexico by W. Bullock, F.L.S. and Mr. William Bullock jun". Philosophical Magazine. New Series. 1: 433–442 [439]. doi:10.1080/14786442708674357.
  4. ^ Jobling, James A. (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. p. 163. ISBN 978-1-4081-2501-4.
  5. ^ Peters, James Lee, ed. (1948). Check-List of Birds of the World. Vol. 6. Cambridge, Massachusetts: Harvard University Press. p. 159.
  6. ^ Swainson, William John (1831). Richardson, John (ed.). Fauna boreali-americana, or, The zoology of the northern parts of British America : containing descriptions of the objects of natural history collected on the late northern land expeditions under command of Captain Sir John Franklin, R.N.: Part 2, The Birds. p. 316. The title page gives the date as 1831 but the volume was not actually published until the following year.
  7. ^ a b Gill, Frank; Donsker, David; Rasmussen, Pamela, eds. (2020). "Woodpeckers". IOC World Bird List Version 10.1. International Ornithologists' Union. Retrieved 2 October 2020.
  8. ^ Shakya, S.B.; Fuchs, J.; Pons, J.-M.; Sheldon, F.H. (2017). "Tapping the woodpecker tree for evolutionary insight". Molecular Phylogenetics and Evolution. 116: 182–191. doi:10.1016/j.ympev.2017.09.005. PMID 28890006.
  9. ^ a b "Acorn Woodpecker Identification, All About Birds, Cornell Lab of Ornithology". www.allaboutbirds.org. Retrieved 26 September 2020.
  10. ^ Scofield, Douglas G.; Alfaro, Victor Ryan; Sork, Victoria L.; Grivet, Delphine; Martinez, Edith; Papp, Jeannette; Pluess, Andrea R.; Koenig, Walter D.; Smouse, Peter E. (2011). "Foraging patterns of acorn woodpeckers (Melanerpes formicivorus) on valley oak (Quercus lobata Née) in two California oak savanna-woodlands". Oecologia. 166 (1): 187–196. Bibcode:2011Oecol.166..187S. doi:10.1007/s00442-010-1828-5. hdl:10261/291553. PMC 3074073. PMID 21107869.
  11. ^ Stacey, P; Bock, C (1978). "Social plasticity in the acorn woodpecker" (PDF). Science. 202 (4374): 1298–1300. Bibcode:1978Sci...202.1298S. doi:10.1126/science.202.4374.1298. JSTOR 1747051. PMID 17750490. S2CID 34467844.
  12. ^ Cockburn, Andrew (2006). "Prevalence of different modes of parental care in birds". Proceedings of the Royal Society B: Biological Sciences. 273 (1592): 1375–1383. doi:10.1098/rspb.2005.3458. PMC 1560291. PMID 16777726.
  13. ^ Hatchwell, B.J.; Komdeur, J. (2000). "Ecological constraints, life history traits and the evolution of cooperative breeding" (PDF). Animal Behaviour. 59 (6): 1079–1086. doi:10.1006/anbe.2000.1394. PMID 10877885.
  14. ^ Koenig, Walter D. (1981). "Reproductive success, group size, and the evolution of cooperative breeding in the acorn woodpecker". American Naturalist. 117 (4): 421–443. doi:10.1086/283726. JSTOR 10.1086/283726.
  15. ^ Haydock, Joseph; Koenig, Walter D. (2002). "Reproductive skew in the polygynandrous acorn woodpecker". Proceedings of the National Academy of Sciences. 99 (10): 7178–7183. Bibcode:2002PNAS...99.7178H. doi:10.1073/pnas.102624199. PMC 124548. PMID 11997473.
  16. ^ Koenig, Walter D.; Haydock, Joseph; Stanback, Mark T. (1998). "Reproductive roles in the cooperatively breeding Acorn Woodpecker: incest avoidance versus reproductive competition". American Naturalist. 151 (3): 243–255. doi:10.1086/286115. JSTOR 286115. PMID 18811355. Archived 25 September 2017 at the Wayback Machine
  17. ^ Koenig, Walter D.; Mumme, Ronald L.; Stanback, Mark T.; Pitelka, Frank A. (1995). "Patterns and consequences of egg destruction among joint-nesting acorn woodpeckers". Animal Behaviour. 50 (3): 607–621. doi:10.1016/0003-3472(95)80123-5.
  18. ^ Joste, Nancy; Ligon, J. David; Stacey, Peter B. (1985). "Shared paternity in the acorn woodpecker (Melanerpes formicivorus)". Behavioral Ecology and Sociobiology. 17 (1): 39–41. doi:10.1007/BF00299426. JSTOR 4599801.
  19. ^ Koenig, W.D., and Stahl, J.T. (2007). Condor 109(2):334–350.
  20. ^ Trail, Pepper W (1980). "Ecological Correlates of Social Organization in a Communally Breeding Bird, the Acorn Woodpecker, Melanerpes formicivorus". Behavioral Ecology and Sociobiology. 7 (2): 83–92. doi:10.1007/BF00299512. S2CID 11335989 – via Springer Link.
  21. ^ Zickefoose, Julie (10 March 2009). "Woody The Acorn (Not Pileated) Woodpecker". NPR. Retrieved 14 September 2011.

Further reading