Biology:Black-capped chickadee

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Short description: Species of small, non-migratory, North American songbird

Black-capped chickadee
Black Capped Chickadee (194768869).jpeg
Scientific classification edit
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Family: Paridae
Genus: Poecile
Species:
P. atricapillus
Binomial name
Poecile atricapillus
(Linnaeus, 1766)
Poecile atricapillus distribution map.png
Range of Poecile atricapillus
Synonyms

Parus atricapillus Linnaeus, 1766

The black-capped chickadee (Poecile atricapillus) is a small, nonmigratory, North American songbird that lives in deciduous and mixed forests. It is a passerine bird in the tit family, the Paridae. It is the state bird of Massachusetts and Maine in the United States, and the provincial bird of New Brunswick in Canada. It is well known for its ability to lower its body temperature during cold winter nights, its good spatial memory to relocate the caches where it stores food, and its boldness near humans (sometimes feeding from the hand).

The black-capped chickadee is widely distributed throughout North America, ranging from the northern United States to southern Canada and all the way up to Alaska and Yukon. It has a distinct appearance characterized by its black cap and "bib" with white sides. The black-capped chickadee is a social bird and forms flocks in the winter that include other bird species. The bird is well known for its vocalizations, including its fee-bee call and its chick-a-dee-dee-dee call, from which it derives its name.

Taxonomy

In 1760, French zoologist Mathurin Jacques Brisson included a description of the black-capped chickadee in his book Ornithologie based on a specimen collected in Canada. He used the French name La mésange a tête noire de Canada and the Latin Parus Canadensis Atricapillus.[2] Although Brisson gave it Latin names, these do not conform to the binomial system and are not recognised by the International Commission on Zoological Nomenclature.[3] In 1766, Swedish naturalist Carl Linnaeus published the 12th edition of his Systema Naturae, which included 240 species that had been previously described by Brisson.[3] One of these was the black-capped chickadee. Linnaeus included a brief description, coined the binomial name Parus atricapillus and cited Brisson's work.[4] The type location was subsequently restricted to Quebec, Canada.[5] The specific epithet atricapillus is Latin for "black-haired" from ater (black) and capillus (hair of the head).[6]

Though originally placed in the genus Parus with most other tits, mtDNA cytochrome b sequence data and morphology suggested that separating Poecile more adequately expressed these birds' relationships.[7] The genus Poecile had been introduced by German naturalist Johann Jakob Kaup in 1829.[8] Molecular phylogenetic studies have shown that the black-capped chickadee is sister to the mountain chickadee (Poecile gambeli).[9][10]

The following cladogram shows the relationships between the various chickadee species, a subset of genus Poecile:[11]

 
 
 
 
 

Grey-headed chickadee (Poecile cinctus)

 
 
 

Chestnut-backed chickadee (Poecile rufescens)

 
 

Boreal chickadee (Poecile hudsonicus)

 
 
 
 
 
 

Mexican chickadee (Poecile sclateri)

 
 
 

Carolina chickadee (Poecile carolinensis)

 
 
 

Black-capped chickadee (Poecile atricapillus)

 
 

Mountain chickadee (Poecile gambeli)

 
 
 
 
 
 
 
 

At one time, the black-capped chickadee was considered by some to be conspecific with the willow tit of Eurasia, due to their very similar appearance. This is reflected in an older version of the Peterson Field Guide for the Birds of Britain and Europe, which states "N Am. Black-Capped Chickadee" as an alternate name for the willow tit. In fact, the willow tit, black-capped chickadee, marsh tit, and Carolina chickadee are all very similar to one another in appearance.[citation needed]

Subspecies

Nine subspecies are currently recognized:[12]

  • Poecile atricapillus turneri (Ridgway, 1884): Found in southern Alaska and northwest Canada and has the palest colouring of all subspecies.[12][13]
  • Poecile atricapillus occidentalis (Baird, 1858): Ranges from southwest British Columbia to northwest California, and is the smallest in size and darkest in colouring of all subspecies.[12][13]
  • Poecile atricapillus fortuitus (Dawson, 1909): Ranges from inland southwest Canada to northwest U.S.[12]
  • Poecile atricapillus septentrionalis (Harris, 1846): Ranges from western and central Canada to southern U.S. It is visually similar to P. a. atricapillus but has slightly longer wings and tail.[12][13]
  • Poecile atricapillus bartletti (Alrich & Nutt, 1939): Lives only in Newfoundland, Canada, and is the only species with different mtDNA, likely due to geographic isolation.[12][13]
  • Poecile atricapillus atricapillus (Linnaeus, 1766): Nominate subspecies, which ranges from eastern and central Canada to northeastern U.S.[12]
  • Poecile atricapillus garrinus (Behle, 1951): Lives in western central U.S.[12]
  • Poecile atricapillus nevadensis (Linsdale, 1938): Lives in western U.S.[12]
  • Poecile atricapillus practicus (Oberholser, 1937): Lives in the Appalachian mountains in the eastern U.S.[12]

Description

Black-capped chickadee clinging to a wire

The black-capped chickadee has a black cap and "bib" with white cheeks. Its underparts are white with buff-colored flanks. Its back is unstreaked and greenish gray, and the tail and wings are slate gray. Males and females are visually similar, however they can be distinguished based on a combination of weight, tail length, and tarsus length. Males have a wing length of 63.5–67.5 mm (2.50–2.66 in), a tail length of 58–63 mm (2.3–2.5 in), a bill length of 8–9.5 mm (0.31–0.37 in), and a tarsus length of 16–17 mm (0.63–0.67 in). Females are on average slightly smaller, with a wing length of 60.5–66.5 mm (2.38–2.62 in), a tail length of 56.3–63 mm (2.22–2.48 in), a bill length of 9–9.5 mm (0.35–0.37 in), and a tarsus length of 16–17 mm (0.63–0.67 in). Both sexes weigh 10–14 g (0.35–0.49 oz).[13]

Although range can generally be used to separate them, the black-capped chickadee is very similar in appearance to the Carolina chickadee. The edge of the black-capped chickadee's wings are somewhat paler than the Carolina chickadee, and the bib of the black-capped chickadee has a "messier" border than the Caroline chickadee. The most reliable way to distinguish the two species is by vocalizations. Black-capped chickadees have a slower, hoarser, two-part song, whereas Carolina chickadees have a three-part song.[13]

Black-capped chickadees are also somewhat similar to mountain chickadees and boreal chickadees. Mountain chickadees can be distinguished by a distinct white stripe above their eyes, and boreal chickadees have a brown cap as opposed to the black cap of the black-capped chickadee.[14]

Vocalization

Black-capped chickadee, Iona Beach Regional Park

The vocalizations of the black-capped chickadee are highly complex, with 16 distinct types of vocalizations being used to convey an array of information.[15] These vocalizations are likely an evolutionary adaptation to their habitat; they live and feed in dense vegetation, and even when the flock is close together, individual birds tend to be out of each other's visual range.[16]

One of the most recognizable sounds produced, particularly by the males, is the two-note fee-bee song. It is a simple, clear whistle of two notes, identical in rhythm, the first roughly a whole-step above the second.[17] The range of frequencies at which this song starts from varies; the complete frequency range spans roughly 1 kHz. Within this range, male chickadees can sing at various tones. The average starting frequency is around 4000 Hz. A decrease of roughly 200 Hz occurs when the first note (fee) is sung, and then another decrease around 400 Hz takes place between the end of fee and the beginning of bee. In spite of these multiple changes in frequency, though, anybody listening to the song only hears a pure, high-frequency tone.[18] This is distinguished from the Carolina chickadee's four-note call fee-bee fee-bay; the lower notes are nearly identical but the higher fee notes are omitted, making the black-capped song like bee bay. The males sing the song only in relative isolation from other chickadees (including their mates). In late summer, some young birds sing only a single note.[19]

Both sexes sometimes make a faint version of the fee-bee song, and this appears to be used when feeding young.[19] When males are out in the wild, they sing this song to defend their territory or attract a mate.[20] There have also been a few accounts of females singing a version of the fee-bee song when alone in the wild, although its purpose in this context is unknown. The black-capped chickadee is a monomorphic species, so distinguishing males and females based solely on their singing is difficult. A bioacoustic analysis performed on both male and female songs revealed that male fee-bee singing fluctuates more, and the absolute amplitude of both sexes is the same. The fee glissando varies far more in females, which makes identifying each sex easier.[21]

The most familiar call is the chick-a-dee-dee-dee, which gave this bird its name. This simple-sounding call is astonishingly complex.[19] Scientists have been studying it since the mid-1970s. It is produced by both males and females year-round.[18] It has been observed to consist of up to four distinct units—referred to as A, B, C, and D—which can be arranged in different patterns to communicate information about threats from predators and coordination of group movement. These four notes only ever appear in this consecutive order with each preceding note blending into the next; however, not all four notes always appear in the call.[18][19][22] Like other sounds the chickadee produces, it may be heard in multiple variations. The A and B notes are almost identical to one another in both frequency and duration, though black-capped chickadees possess the ability to quickly notice the difference between these two notes. No such similarity is seen between the C and D notes.[18] The C note fluctuates from low to high then back to low, whereas the D note has a constant frequency. While not confirmed, one study found evidence of a function behind the C and D notes: the C note is used indicate the location of food, and the D note is used to distinguish between a member of the same species and a potential predator.[22]

Neither individual notes nor groups of notes have an equal probability of appearing in the chick-a-dee-dee-dee song. Its syntax form may take on several different structures, but the two most commonly heard are [A][D] and [B][C][D].[Note 1] Calls containing the D note are most frequently heard.[18] A study of the call showed that the number of dees indicates the level of threat from nearby predators. In an analysis of over 5,000 alarm calls from chickadees, alarm calls triggered by small, dangerous raptors had a shorter interval between chick and dee and tended to have extra dees, typically four instead of two. In one case, a warning call about a pygmy owl—a prime threat to chickadees—contained 23 dees. The Carolina chickadee makes a similar call, which is faster and higher-pitched.[19]

Black-capped chickadees make a number of other calls and sounds, such as a gargle noise usually used by males to indicate a threat of attacking another male, often when feeding. This call is also used in sexual contexts.[20] Black-capped chickadees learns the gargle noise soon after birth and continues to develop it through to adulthood.[18] This noise is among the most complex of the calls; in one population of chickadees, the gargle contained 2 to 9 instances of 14 distinct notes, all sung within half a second.[20]

Social learning in particular is largely influential to the development of the gargle. Beginning 30 to 35 days after birth, strings of low-amplitude precursor or sub-gargles are produced for about a minute. At this time, the young have several close interactions with their family; they learn to produce such sounds by listening to their parents and siblings. Three chickadee populations were observed at three different sites over 8 years, and all of them produced vocalizations that were very similar to one another. Strings of juvenile sub-gargles are almost perfectly continuous and both low and unstable in frequency, yet lacking multiple syllables. When their vocal abilities are fully developed, a stable frequency is produced and a variety of syllables is heard that vary in length.[18]

Black-capped chickadees in an environment with ambient noise at the same frequencies as their songs have developed an evolutionary adaptation that enables them to adjust the frequency of their songs much quicker to effectively communicate with the surrounding population. The fee-bee songs of several male black-capped chickadees were monitored to identify their particular frequency. In one study, a series of both masking[Note 2] and nonmasking[Note 3] tones was played to multiple male chickadees at various locations to observe how they responded. When interacting with other chickadees close by, the males matched their frequencies; however, when the surrounding environment was noisy with other species, the males adapted by increasing the frequency of their songs. The males responded quickly to the masking tones by raising their song frequencies. Another survey, though, showed that male chickadees sometimes intentionally match the tones of competing chickadees as a way of showing aggression. Dominant males in a population often compete with lower-ranked males, and singing contests are a way male chickadees decide who gets to mate. When a male loses a contest, particularly a higher-ranking male in the population, he will often have difficulty finding a mate.[20]

Distribution and habitat

Black-capped chickadees are non-migratory and can be found throughout much of North America. They range from western Alaska, through southern Yukon and throughout the Canadian provinces, from British Columbia in the west to the Maritimes and Newfoundland in the east. The distribution continues into the United States, with its range extending to northern California in the south-west, through northern Nevada and New Mexico, continuing through the midwestern United States to New Jersey. It can also be found at higher elevations in the Appalachian Mountains.[13] In British Columbia, the black-capped chickadee is absent on Vancouver Island, the Gulf Islands, Haida Gwaii, and parts of the Sunshine Coast, where it is replaced by the chestnut-backed chickadee.[23][24]

Black-capped chickadees inhabit wooded areas, including both coniferous and deciduous forests, urban parks, willow thickets, and suburban areas. They do not vary their habitat between breeding and non-breeding seasons, however during the winter irregular irruptions may occur.[13]

Behaviour and ecology

Diet and foraging

Chickadees will take seeds from feeders and trays over to a tree branch to hammer them open

Insects (especially caterpillars) form a large part of their diet in summer. The birds hop along tree branches searching for food, sometimes hanging upside down or hovering; they may make short flights to catch insects in the air. Seeds and berries become more important in winter, though insect eggs and pupae are eaten when available.[13] Black-capped chickadees have also been known to eat the fat off of dead mammals.[25] Sunflower seeds are readily taken from bird feeders. The birds take a seed in their beak and commonly fly from the feeder to a tree, where they proceed to hammer the seed on a branch to open it.[26]

Like many other species in the family Paridae, black-capped chickadees commonly cache food, mostly seeds, but sometimes insects, also.[27] Items are stored singly in various sites such as bark, dead leaves, clusters of conifer needles, or knothole. Memory for the location of caches can last up to 28 days.[28] Within the first 24 hours, the birds can even remember the relative quality of the stored items.[29]

This caching behavoiur has led to black-capped chickadees having larger hippocampi[Note 4] compared to other chickadees, who themselves have relatively larger hippocampi compared to others caching birds in the Paridae family.[32][33] This variation in size also exists within the black-capped chickadee population based on the region they inhabit, with those who live in harsher climates (such as Alaska) having larger hippocampi.[34] However, no variation exists between the sexes.[35] The size of the hippocampus within black-capped chickadees also varies throughout the year, being the largest in October, and the smallest in February. While the exact reason for this seasonal change is unknown, it is believed that the hippocampus grows to allowed the chickadee to remember its cache locations, and then shrinks as those caches are used up.[36][37]

Foraging behaviour in the winter tends to decrease due to the changing weather. Such behaviour is largely influenced by wind and temperature. When wind speeds are higher, black-capped chickadees avoid exposure to such conditions by flying lower where vegetation offers a degree of protection, and when the temperature decreases, they search for food less frequently.[38]

Chickadee at feeder

In parts of the black-capped chickadee's range with very cold winters, such as Minnesota, survival rates are affected by access to supplemental food. Chickadees with access to bird feeders are twice as likely to survive the winter than those without access to this supplemental food. This difference in survival rates occurs primarily during months with severe weather when the temperature drops below −18 °C (0 °F) for more than five days.[26] In Pennsylvania, with milder winters on the southern edge of their range, differences between populations with and without feeders suggest that feeders influence movements of chickadees rather than actual survival.[39]

Metabolism

On cold winter nights, these birds can reduce their body temperature by as much as 12 °C (from their normal temperature of about 42 °C) to conserve energy.[40][41] Such a capacity for torpor is not very common in birds. Other bird species capable of torpor include the common swift (Apus apus), the common poor-will (Phalaenoptilus nuttallii), the lesser nighthawk (Chordeiles acutipennis), and various species of hummingbirds.[42][43]

Movement and roosting

During the winter, chickadees often flock together. Many other species of birds – including titmice, nuthatches, and warblers – can often be found foraging in these flocks. Mixed flocks stay together because the chickadees call out whenever they find a good source of food. This calling-out forms cohesion for the group, allowing the other birds to find food more efficiently.[44][45] Black-capped chickadees sleep in thick vegetation or in cavities, usually singly, though they may occasionally roost clumped together.[46]

Their flight is slightly undulating with rapid wing beats. Flights are typically short bursts of less than 15 m (49 ft) with a speed around 20 km/h (12 mph).[47] They tend to avoid flying through large open areas and will instead be found flying along tree-lines or through forests.[48]

Molt

Chickadees molt once a year; no prenuptial molt occurs in the spring. The postjuvenal molt at the end of the first summer of life is partial, involving only the body feathers and wing coverts. Thereafter, the postnuptial molts at the end of each reproductive season are always complete, involving all feathers.

Breeding

A black-capped chickadee excavating a nest cavity in a dead tree

The black-capped chickadee nests in tree cavities 1–7 m (3.3–23.0 ft) above ground.[49] The pair either excavates the hole together, uses a natural cavity, or reuses an old woodpecker nest. This species will also nest in a nesting box. Nest sites are typically chosen by females, however excavation of the cavity is done by both sexes. The nest itself is built by the female only and consists of a base of coarse material such as moss or bark strips, and lining of finer material such as mammal hair.[50] The nesting season is from late April through June, with higher-ranking females nesting before lower ranking ones.[49]Eggs are white with fine dots of reddish brown concentrated at the larger end. On average, eggs are 1.52 cm × 1.22 cm (0.60 in × 0.48 in). Clutch size is six to eight eggs. Incubation lasts 11–14 days and is by the female only, which is fed by the male. If an unusual disturbance occurs at the nest entrance, the incubating female may utter an explosive hiss, like that of a snake, a probable adaptation to discourage nest predators.[51]

Hatchlings are altricial, naked with their eyes closed. Nestlings are fed by both sexes, but are brooded by the female only (when the male brings food to her, which she passes on to the young). Young leave the nest 12–16 days after hatching, in great part because the parents start presenting food only outside the nest hole. The young are still fed by the parents for several weeks, but are capable of catching food on their own within a week after leaving the nest.[13]

Black-capped chickadees usually breed only once a year, but second broods are possible if the first one is lost. First breeding is at one year of age. Maximum recorded lifespan is 12 years, but most individuals live only half that long.[52]

Black-capped chickadees are socially monogamous, and males contribute greatly to reproduction. During the laying and incubation periods, males feed their partners extensively. When the nestlings hatch, males are the primary providers, but as the nestlings grow, females become the main caretakers. Females prefer dominant males, and greater reproductive success is closely related to the higher ranking of the male.[53]

Black-capped chickadees may interbreed with Carolina chickadees or mountain chickadees where their ranges overlap.[54][55] Interbreeding with boreal chickadees has also been documented, though it is more rare.[56]

Dominance hierarchy

During the winter, the species forms flocks through which dominance hierarchies can be easily observed. Dominance hierarchies play an important role in determining the social behaviors among the birds in these flocks. Positive correlates to higher social rankings include territory size, body condition, singing rate, and reproductive success.[57] The hierarchies are linear and stable; once a relationship is established between two birds, it stays the same for many years. In general, older and more experienced birds are dominant over younger ones, and males are dominant over females.[53] Dominant and subordinate members differ in their foraging strategies and risk-taking behaviors. Dominant individuals control access to preferred resources and restrict subordinates to foraging in novel, riskier, or suboptimal environments. Subordinate individuals are often observed foraging in the outermost tree parts that are more prone to predators, while dominant individuals forage low and close to the tree trunk.

In experiments, subordinate individuals display less neophobia when approaching novel foods and objects, compared to their dominant counterparts. Subordinate individuals are also more likely to enter novel environments than their dominant counterparts. This is similar to subordinate primates, which feed on novel food more readily than the dominant individuals because they are more used to eating suboptimal and unfamiliar food. No difference is observed in ability to learn novel foraging tasks between dominant and subordinate individuals.[57]

State and provincial bird

A vehicle registration plate from Maine, with a black-capped chickadee perched on a pine branch on the left of the plate
A Maine registration plate, with a black-capped chickadee on the left

The black-capped chickadee is the state bird of Maine and Massachusetts and the provincial bird of New Brunswick.[58][59][60] In 2022 the black-capped chickadee was named the official bird of Calgary, Alberta.[61] The bird is prominently featured on the standard Maine vehicle registration plate.[62]

Conservation

The IUCN classifies the black-capped chickadee as least concern due to its wide distribution and large populations.[1] In Alaska and Washington, and parts of western Canada, black-capped chickadees are among a number of bird species affected by an unknown agent that is causing beak deformities, which may cause stress for affected species by inhibiting feeding ability, mating, and grooming. Black-capped chickadees were the first affected bird species, with reports of the deformity beginning in Alaska in the late 1990s, but more recently the deformity has been observed in close to 30 bird species in the affected areas.[63]

Notes

  1. ↑ Brackets indicate that a note may be repeated more than once.[18]
  2. ↑ A tone played at the same frequency as the calls of the chickadees.[20]
  3. ↑ A tone played well above the same frequency as the calls of the chickadees.[20]
  4. ↑ The hippocampus plays a role in converting short-term memory to long-term memory, as well as supporting spatial memory.[30][31]

References

  1. ↑ 1.0 1.1 BirdLife International (2017). "Poecile atricapillus". IUCN Red List of Threatened Species 2017: e.T22711716A118687681. doi:10.2305/IUCN.UK.2017-3.RLTS.T22711716A118687681.en. https://www.iucnredlist.org/species/22711716/118687681. Retrieved 12 November 2021. 
  2. ↑ Brisson, Mathurin Jacques (1760) (in fr, la). Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés. 3. Paris: Jean-Baptiste Bauche. pp. 553–555, Plate 29 fig 1. https://biodiversitylibrary.org/page/35953652.  The two stars (**) at the start of the section indicates that Brisson based his description on the examination of a specimen.
  3. ↑ 3.0 3.1 Allen, J.A. (1910). "Collation of Brisson's genera of birds with those of Linnaeus". Bulletin of the American Museum of Natural History 28: 317–335. 
  4. ↑ Linnaeus, Carl (1766) (in la). Systema naturae : per regna tria natura, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 1, Part 1 (12th ed.). Holmiae (Stockholm): Laurentii Salvii. p. 341. https://biodiversitylibrary.org/page/42946537. 
  5. ↑ Paynter, Raymond A. Jr, ed (1986). Check-list of Birds of the World. 12. Cambridge, Massachusetts: Museum of Comparative Zoology. p. 82. https://biodiversitylibrary.org/page/14482199. 
  6. ↑ Jobling, J.A. (2018). "Key to Scientific Names in Ornithology". in del Hoyo, J.; Elliott, A.; Sargatal, J. et al.. Handbook of the Birds of the World Alive. Lynx Edicions. https://www.hbw.com/dictionary/definition/atricapillus. 
  7. ↑ Gill, F. B.; Slikas, B.; Sheldon, F. H. (2005). "Phylogeny of titmice (Paridae): II. Species relationships based on sequences of the mitochondrial cytochrome-b gene". Auk 122: 121–143. doi:10.1642/0004-8038(2005)122[0121:POTPIS2.0.CO;2]. http://www2.mnhn.fr/crbpo/IMG/pdf/Gill_et_al_2005_Auk_tits.pdf. 
  8. ↑ Kaup, Johann Jakob (1829) (in de). Skizzirte Entwickelungs-Geschichte und natürliches System der europäischen Thierwelt. c. 1. Darmstadt: Carl Wilhelm Leske. p. 114. https://biodiversitylibrary.org/page/41576537. 
  9. ↑ Johansson, Ulf S.; Ekman, Jan; Bowie, Rauri C. K.; Halvarsson, Peter; Ohlson, Jan I.; Price, Trevor D.; Ericson, Per G. P. (2013). "A complete multilocus species phylogeny of the tits and chickadees (Aves: Paridae)". Molecular Phylogenetics and Evolution 69 (3): 852–860. doi:10.1016/j.ympev.2013.06.019. PMID 23831453. 
  10. ↑ Tritsch, Christian; Martens, Jochen; Sun, Yue-Hua; Heim, Wieland; Strutzenberger, Patrick; Päckert, Martin (2017). "Improved sampling at the subspecies level solves a taxonomic dilemma – A case study of two enigmatic Chinese tit species (Aves, Passeriformes, Paridae, Poecile)". Molecular Phylogenetics and Evolution 107: 538–550. doi:10.1016/j.ympev.2016.12.014. PMID 27965081. 
  11. ↑ Tritsch, Christian; Martens, Jochen; Sun, Yue-Hua; Heim, Wieland; Strutzenberger, Patrick; Päckert, Martin (2017). "Improved sampling at the subspecies level solves a taxonomic dilemma – A case study of two enigmatic Chinese tit species (Aves, Passeriformes, Paridae, Poecile)". Molecular Phylogenetics and Evolution 107: 538–550. doi:10.1016/j.ympev.2016.12.014. PMID 27965081. 
  12. ↑ 12.0 12.1 12.2 12.3 12.4 12.5 12.6 12.7 12.8 12.9 Gill, Frank; Donsker, David, eds (2018). "Waxwings and allies, tits, penduline tits". World Bird List Version 14.1. International Ornithologists' Union. http://www.worldbirdnames.org/bow/waxwings/. 
  13. ↑ 13.0 13.1 13.2 13.3 13.4 13.5 13.6 13.7 13.8 13.9 Foote, Jennifer R.; Mennill, Daniel J.; Ratcliffe, Laurene M.; Smith, Susan M. (4 March 2020). "Black-capped Chickadee (Poecile atricapillus)". Birds of the World. doi:10.2173/bow.bkcchi.01. https://birdsoftheworld-org.elibrary.calgarypubliclibrary.com/bow/species/bkcchi/cur/introduction. Retrieved 17 January 2024. 
  14. ↑ "Black-capped Chickadee - Similar Species Comparison". Cornell University. https://www.allaboutbirds.org/guide/Black-capped_Chickadee/species-compare/. 
  15. ↑ Smith, Susan M. (1991). The black capped Chickadee: behavioral ecology and natural history. Ithaca, NY: Comstock Publ. Assoc. ISBN 0-8014-2382-1. 
  16. ↑ Ficken, M. S.; Ficken, R. W.; Witkin, S. R. (1978). "Vocal repertoire of the Black-capped Chickadee". Auk 95 (1): 34–48. doi:10.2307/4085493. http://sora.unm.edu/sites/default/files/journals/auk/v095n01/p0034-p0048.pdf. 
  17. ↑ Jackson, Dave (24 March 2010). "Olympic Peninsula Audubon Society". Olympic Peninsula Audubon Society. http://www.olympicpeninsulaaudubon.org/shell.php?page=gazette_Mar_2010. 
  18. ↑ 18.0 18.1 18.2 18.3 18.4 18.5 18.6 18.7 Otter, Ken A (2007). Ecology and Behaviour of Chickadees and Titmice: An Integrated Approach. Oxford University Press. pp. 153–230. ISBN 978-0-19-856999-2. 
  19. ↑ 19.0 19.1 19.2 19.3 19.4 Templeton, C. N.; Greene, E.; Davis, K. (2005). "Allometry of alarm calls: black-capped chickadees encode information about predator size". Science 308 (5730): 1934–7. doi:10.1126/science.1108841. PMID 15976305. Bibcode: 2005Sci...308.1934T. 
  20. ↑ 20.0 20.1 20.2 20.3 20.4 20.5 Goodwin, Sarah E.; Podos, Jeffrey (9 January 2013). "Shift of song frequencies in response to masking tones". Animal Behaviour 85 (2): 435–440. doi:10.1016/j.anbehav.2012.12.003. 
  21. ↑ Hahn, Allison H.; Krysler, Amanda; Sturdy, Christopher B. (11 May 2013). "Female song in black-capped chickadees (Poecile atricapillus): Acoustic song features that contain individual identity information and sex differences". Behavioural Processes 98: 98–105. doi:10.1016/j.beproc.2013.05.006. PMID 23694740. 
  22. ↑ 22.0 22.1 Dawson, Michael R. W.; Charrier, Isabelle; Sturdy, Christopher B. (May 2006). "Using an artificial neural network to classify black-capped chickadee (Poecile atricapillus) call note types". The Journal of the Acoustical Society of America 119 (5): 3161–3172. doi:10.1121/1.2189028. PMID 16708971. Bibcode: 2006ASAJ..119.3161D. 
  23. ↑ Wright, K. G.. "Black-capped Chickadee". The Atlas of the Breeding Birds of British Columbia. https://www.birdatlas.bc.ca/accounts/speciesaccount.jsp?sp=BCCH&lang=en. 
  24. ↑ Dahlsten, Donald L.; Brennan, Leonard A.; McCallum, D. Archibald; Gaunt, Sandra L. (4 March 2020). "Chestnut-backed Chickadee (Poecile rufescens)". Birds of the World. doi:10.2173/bow.chbchi.01. 
  25. ↑ Hammerstrom, Frances (1942). "Dominance in Winter Flocks of Chickadees". The Wilson Bulletin 54 (1): 32-42. https://sora.unm.edu/sites/default/files/journals/wilson/v054n01/p0032-p0042.pdf. Retrieved 27 January 2024. 
  26. ↑ 26.0 26.1 Brittingham, M.C.; Temple, S.A. (1988). "Impacts of Supplemental Feeding on Survival Rates of Black-capped Chickadees". Ecology 69 (3): 581. doi:10.2307/1941007. Bibcode: 1988Ecol...69..581B. 
  27. ↑ Heinrich, Bernd; Collins, Scott L. (June 1983). "Caterpillar Leaf Damage, and the Game of Hide-and-seek with Birds". Ecology 64 (3): 592–602. doi:10.2307/1939978. Bibcode: 1983Ecol...64..592H. 
  28. ↑ Hitchcock, C. L.; Sherry, D. F. (1990). "Long-term memory for cache sites in the Black-capped Chickadee". Animal Behaviour 40 (4): 701. doi:10.1016/S0003-3472(05)80699-2. 
  29. ↑ Sherry, D. F. (1984). "Food storage by the Black-capped Chickadee: memory for the location and contents of caches". Animal Behaviour 32 (2): 451. doi:10.1016/S0003-3472(84)80281-X. 
  30. ↑ Squire, Larry R. (1992). "Memory and the hippocampus: A synthesis from findings with rats, monkeys, and humans.". Psychological Review 99 (2): 195–231. doi:10.1037/0033-295X.99.2.195. 
  31. ↑ O'Keefe, John M.; Nadel, Lynn; O'Keefe, John (1978). The hippocampus as a cognitive map. Oxford: Clarendon Press. ISBN 0-19-857206-9. https://www.cmor-faculty.rice.edu/~cox/neuro/HCMComplete.pdf. Retrieved 28 January 2024. 
  32. ↑ Sherry, David F. (1989). "Food Storing in the Paridae". The Wilson Bulletin 101 (2): 289–304. ISSN 0043-5643. https://sora.unm.edu/sites/default/files/journals/wilson/v101n02/p0289-p0304.pdf. Retrieved 28 January 2024. 
  33. ↑ Hampton, Robert R.; Sherry, David F.; Shettleworth, Sara J.; Khurgel, Moshe; Ivy, Gwen (1995). "Hippocampal Volume and Food-Storing Behavior Are Related in Parids". Brain, Behavior and Evolution 45 (1): 54–61. doi:10.1159/000113385. 
  34. ↑ Roth, Timothy C; Pravosudov, Vladimir V (7 February 2009). "Hippocampal volumes and neuron numbers increase along a gradient of environmental harshness: a large-scale comparison". Proceedings of the Royal Society B: Biological Sciences 276 (1656): 401–405. doi:10.1098/rspb.2008.1184. 
  35. ↑ Petersen, Karin; Sherry, David F. (September 1996). "No sex difference occurs in hippocampus, food-storing, or memory for food caches in black-capped chickadees". Behavioural Brain Research 79 (1-2): 15–22. doi:10.1016/0166-4328(95)00257-X. 
  36. ↑ Smulders, T. V.; Sasson, A. D.; DeVoogd, T. J. (May 1995). "Seasonal variation in hippocampal volume in a food‐storing bird, the black‐capped chickadee". Journal of Neurobiology 27 (1): 15–25. doi:10.1002/neu.480270103. 
  37. ↑ Phillmore, Leslie S.; Hoshooley, Jennifer S.; Sherry, David F.; MacDougall‐Shackleton, Scott A. (August 2006). "Annual cycle of the black‐capped chickadee: Seasonality of singing rates and vocal‐control brain regions". Journal of Neurobiology 66 (9): 1002–1010. doi:10.1002/neu.20282. 
  38. ↑ Otter, Ken A. (2007). Ecology and Behaviour of Chickadees and Titmice. New York, NY: Oxford University Press. p. 268. ISBN 978-0-19-856999-2. 
  39. ↑ Egan, E.S.; Brittingham, M.C. (1994). "Winter Survival Rates of a Southern Population of Black-capped Chickadees". Wilson Bulletin 106 (3): 514. 
  40. ↑ Chaplin, S. B. (1974). "Daily energetics of the Black-capped Chickadee, Parus atricapillus, in winter". Journal of Comparative Physiology 89 (4): 321–330. doi:10.1007/BF00695350. 
  41. ↑ Chaplin, S. B. (1976). "The physiology of hypothermia in the Black-capped Chickadee Parus atricapillus". Journal of Comparative Physiology B 112 (3): 335–344. doi:10.1007/BF00692303. 
  42. ↑ Hainsworth, F. R.; Wolf, L. L. (17 April 1970). "Regulation of Oxygen Consumption and Body Temperature during Torpor in a Hummingbird, Eulampis jugularls". Science 168 (3929): 368–369. doi:10.1126/science.168.3929.368. PMID 5435893. Bibcode: 1970Sci...168..368R. 
  43. ↑ Ruf, Thomas; Geiser, Fritz (August 2015). "Daily torpor and hibernation in birds and mammals". Biological Reviews 90 (3): 891–926. doi:10.1111/brv.12137. https://onlinelibrary.wiley.com/doi/full/10.1111/brv.12137. Retrieved 30 January 2024. 
  44. ↑ Freeberg, Todd M. (7 February 2023). "Chickadees, titmice and nuthatches flocking together benefit from a diversity bonus – so do other animals, including humans". https://theconversation.com/chickadees-titmice-and-nuthatches-flocking-together-benefit-from-a-diversity-bonus-so-do-other-animals-including-humans-192319. 
  45. ↑ Thompson, Craig (Spring 1983). "The dominance hierarchy of the Black-capped Chickadee and its relation to breeding territory and frequency of visitation to an artificial food source". Field Station Bulletin 16: 14–20. http://dc.uwm.edu/cgi/viewcontent.cgi?article=1095&context=fieldstation_bulletins. 
  46. ↑ Loery, G.; Nichols, J. D. (1985). "Dynamics of a Black-capped Chickadee population, 1958–1983". Ecology 66 (4): 1195–1203. doi:10.2307/1939172. 
  47. ↑ Greenewalt, C. H. (1955). "The flight of the Black-capped Chickadee and the White-breated Nuthatch". Auk 72 (1): 1–5. doi:10.2307/4081384. http://sora.unm.edu/node/20304. 
  48. ↑ St. Clair, Colleen Cassady; Bélisle, Marc; Desrochers, André; Hannon, Susan (1998). "Winter Responses of Forest Birds to Habitat Corridors and Gaps". Conservation Ecology 2 (2). ISSN 1195-5449. https://www.jstor.org/stable/26271681. Retrieved 30 January 2024. 
  49. ↑ 49.0 49.1 Cite error: Invalid <ref> tag; no text was provided for refs named Smith1991
  50. ↑ Aitken, Kathryn E. H.; Martin, Kathy (December 2007). "The importance of excavators in hole-nesting communities: availability and use of natural tree holes in old mixed forests of western Canada". Journal of Ornithology 148 (S2): 425–434. doi:10.1007/s10336-007-0166-9. https://link.springer.com/article/10.1007/s10336-007-0166-9#citeas. Retrieved 30 January 2024. 
  51. ↑ Forbush, E.H. (1925-29) Birds of Massachusetts and other New England states. Mass. Dept. of Agriculture, Boston.
  52. ↑ Löf, R. A. (1967). "Ten years of banding black-capped chickadees". EBBA News 30: 195–198. 
  53. ↑ 53.0 53.1 Oort, Harry Van; Otter, Kenneth A.; Fort, Kevin T.; Mcdonell, Zoe (2007). "Habitat, Dominance, And The Phenotypic Quality of Male Black-Capped Chickadees". The Condor 109 (1): 88. doi:10.1650/0010-5422(2007)109[88:hdatpq2.0.co;2]. 
  54. ↑ Martin, Stephen G.; Martin, KathyAnne (April 1996). "Hybridization between a Mountain Chickadee and Black-capped Chickadee in Colorado". C.F.O Journal 30 (2): 60-65. https://cobirds.org/wp-content/uploads/2023/08/CB_1996_30_2_April-1996.pdf#page=18. Retrieved 30 January 2024. 
  55. ↑ Bronson, C. L.; Grubb, Thomas C.; Sattler, Gene D.; Braun, Michael J. (1 July 2005). "Reproductive Success Across The Black-Capped Chickadee (Poecile Atricapillus) and Carolina Chickadee (P. Carolinensis) Hybrid Zone in Ohio". The Auk 122 (3): 759–772. doi:10.1093/auk/122.3.759. 
  56. ↑ Lait, Linda; Lauff, R. F.; Burg, T. M. (2012). "Genetic evidence supports Boreal Chickadee (Poecile hudsonicus) x Black-capped Chickadee (Poecile atricapillus) hybridization in Atlantic Canada". The Canadian Field-Naturalist 126: 143. doi:10.22621/cfn.v126i2.1330. 
  57. ↑ 57.0 57.1 An, Yong Seok; Kriengwatana, Buddhamas; Newman, Amy E.; Macdougall-Shackleton, Elizabeth A.; Macdougall-Shackleton, Scott A. (2011). "Social Rank, Neophobia and Observational Learning in Black-capped Chickadees". Behaviour 148 (1): 55–69. doi:10.1163/000579510x545829. 
  58. ↑ "The problem with naming 'the chickadee' as Maine's state bird". Bangor Daily News. 28 February 2019. https://bangordailynews.com/2019/02/28/news/state/a-nest-divided-maines-great-chickadee-debate/. 
  59. ↑ "Massachusetts State Bird". SHG Resources. http://www.shgresources.com/ma/symbols/bird/. 
  60. ↑ Government of New Brunswick. "Symbols of New Brunswick". http://www2.gnb.ca/content/gnb/en/departments/government_services/symbols.html. 
  61. ↑ Lirette, Dominika. "The votes are in. Calgary's official bird is...". Canadian Broadcasting Corporation. https://www.cbc.ca/news/canada/calgary/calgary-vote-official-bird-chickadee-1.6453753. 
  62. ↑ "Maine License Plates". Maine Secretary of State. https://www.maine.gov/sos/kids/government/license-plates. 
  63. ↑ Beak Deformities . Alaska Science Center of the United States Geological Survey. Alaska.usgs.gov (15 February 2013). Retrieved on 2013-03-23.

Further reading

  • Smith, S.M. (1991). The black-capped Chickadee: Behavioural Ecology and Natural History. Cornell University Press. ISBN:0-8014-2382-1 (1991 reprint).
  • Smith, S.M. (1993). Black-capped Chickadee. In The Birds of North America, no. 39. (A. Poole, P. Stettenheim and F. Gill, eds.) Philadelphia: The Academy of Natural Sciences.
  • Otter, K.A. (ed) (2007). "Ecology and behavior of chickadees and titmice: an integrated approach". Oxford University Press, Oxford. 310 pp

External links

Wikidata ☰ Q282687 entry



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