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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Cladus: Avemetatarsalia
Cladus: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauriformes
Cladus: Dracohors
Cladus: Dinosauria
Ordo: Saurischia
Cladus: Eusaurischia
Subordo: Theropoda
Cladus: Neotheropoda
Cladus: Averostra
Cladus: Tetanurae
Cladus: Avetheropoda
Cladus: Coelurosauria
Cladus: Tyrannoraptora
Cladus: Maniraptoromorpha
Cladus: Maniraptoriformes
Cladus: Maniraptora
Cladus: Pennaraptora
Cladus: Paraves
Cladus: Eumaniraptora
Cladus: Avialae
Infraclassis: Aves
Cladus: Euavialae
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Ornithuromorpha
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Neognathae
Cladus: Neoaves
Cladus: Telluraves
Cladus: Australaves
Ordo: Passeriformes
Subordo: Passeri
Infraordo: Passerida
Superfamilia: Passeroidea

Familia: Cardinalidae
Genera: AmaurospizaCardinalisCaryothraustesChlorothraupisCyanocompsaCyanoloxiaGranatellusHabiaPasserinaPeriporphyrusPheucticusPirangaRhodothraupisSpiza

Name

Cardinalidae Ridgway, 1901

Typus: Cardinalis Bonaparte, 1838

References
Additional references

Klicka, J., K. Burns, & G. M. Spellman. 2007. Defining a monophyletic Cardinalini: A molecular perspective. Molecular Phylogenetics and Evolution 45: 1014–1032. DOI: 10.1016/j.ympev.2007.07.006 Full article (PDF)Reference page.

Vernacular names
azərbaycanca: Kardinal
català: Cardinàlid
čeština: Kardinálovití
Deutsch: Kardinäle
English: Cardinals
Esperanto: Kardinaledoj
español: cardinálidos
suomi: Kardinaalit
français: Cardinaux
magyar: Kardinálispintyfélék
日本語: ショウジョウコウカンチョウ科
lietuvių: Kardinoliniai
Nederlands: Kardinaalachtigen
norsk nynorsk: Kardinalar
norsk: Kardinaler
Diné bizaad: Tááʼyaʼniiltʼóshii
polski: Kardynały
русский: Кардиналовые
svenska: Kardinaler
Türkçe: Kardinal kuşugiller

Cardinalidae (often referred to as the "cardinal-grosbeaks" or simply the "cardinals") is a family of New World-endemic passerine birds that consists of cardinals, grosbeaks, and buntings. It also includes several birds such as the tanager-like Piranga and the warbler-like Granatellus. As such, membership of this group is not easily defined by a single or even a set of physical characteristics, but instead by molecular work. In general they are medium to large songbirds with stout features, some with large heavy bills.

Members of this group are beloved for their brilliant red, yellow, or blue plumages seen in many of the breeding males in this family. Most species are monogamous breeders that nest in open-cup nests, with many taking turn incubating the nest and taking care of their young. Most are arboreal species though the dickcissel is a ground-dwelling prairie bird.

Conservation-wise most members of this family are considered least concern by the IUCN Red List though a few like the Carrizal seedeater and black-cheeked ant tanager are listed as critically endangered and endangered respectively. Studies on the effects of climate change on species has suggested many more might be threatened with extinction in the near future. However, some species such as the northern cardinal have been expanding their range within the last century.

Field characteristics
The male of the blue grosbeak showing the brilliant blue coloration with brown shoulders
The female of the blue grosbeak is predominantly brown

The grosbeaks, seedeaters, and cardinals have large bills, while Granatellus and buntings have small bills. The cardinalid tanagers have stout, near pointed bills, with some species of Piranga having serrations along the edge of their upper bills.[1] This bill shape is not always an indicator of relationships, as the various species of blue cardinalid species, like the blue grosbeak and Cyanoloxia grosbeaks are related to the buntings. Similarly the cardinalid tanagers are closer to the cardinals and masked grosbeaks (see more in the systematics section). The head is medium to large in size, with a medium neck length. The body of cardinalids ranges from small to medium with lengths of 4.5 to 11 in (11 to 28 cm). Legs are also short to medium in length. The wings are medium and pointed. Cardinalids have nine visible primary feathers with the tenth primary feather being short in comparison.[2][3][4] The plumages in cardinalids are sexually dichromatic as many males of various species display bright reds, oranges, blues or blacks. In most temperate species males will undergo molting between seasons so that non-breeding males will either resemble the females of their species or in-between the two. These species such as the indigo bunting will exhibit a complex molt cycle where they go through four different stages of plumage coverage within their first year of life. In the spring to summer birds start with juvenile plumage to supplemental plumage, then to a first basic (nonbreeding) plumage in the fall to winter and finally the first alternate (breeding) plumage. Adults will typically have the basic two molt cycle from alternate to basic or partial (late summer or fall) to back again in the spring. Males of tropical species will have the same coloration year-round. Females of all species are either drabber in coloration by comparison, either a lighter coloration of the males, with oranges, greens or yellows. The molting pattern in most cardinalids exhibits delayed plumage maturation, so that often the first- year male birds would be in non-breeding plumage or at an intermediate stage.[2] The molting pattern in cardinalids is divided into two types. A preformative molt is a partial molt where only the body feathers get replaced, but not the wing and tail feathers, which is seen in a lot of temperate and neotropical species. The second type is an eccentric preformative molt when only the outer primary and inner secondaries are replaced. This molt is seen in some species of Cyanoloxia and Passerina.[5][6]
Systematics

Traditionally members of this group were classified as a tribe of the finch family Fringillidae (Cardinalini) characterized by heavy, conical, seed-crushing bills. The group consisted of the genera Pheucticus, Parkerthraustes, Saltator, Spiza, Cyanocompsa, Cyanoloxia, Porphyrospiza, Passerina, Caryothraustes, Periporphyrus, Rhodothraupis, and Cardinalis. The issue that taxonomists had faced, however, was there was no unifying morphological traits that were congruent in various studies.[7] In 2007 a mitochondrial DNA study by Klicka, Burns and Spellman sampling all of the aforementioned genera and 34 of the 42 species found that the genera Parkerthraustes, Saltator, and Porphyrospiza were not members of the cardinal-lineage, but instead are found throughout in the tanager-lineage (Thraupidae). Interestingly enough the genera classified as thraupids at the time, Piranga, Habia, Chlorothraupis, and Amaurospiza, are found to be part of cardinalid radiation. In addition the genus Granatellus, originally classified as a parulid warbler, are also found to be part of Cardinalidae.[8] The authors have found that with this new relationship Cardinalidae can be classified into five subgroups, which have been supported by subsequent studies. The five subclades consists of the Habia lineage (Habia and Chlorothraupis), the “masked” lineage (Piranga, Caryothraustes, Periporphyrus, Rhodothraupis, and Cardinalis), the Granatellus lineage, the “blue” lineage (Cyanocompsa, Amaurospiza, Cyanoloxia, Passerina, and Spiza), and the Pheucticus lineage.[8] These subclades and membership of these genera have been widely supported in subsequent studies.[9][10] In a 2021 paper by Guallar et al. found based on the preformative molting pattern of cardinalids suggested the ancestor of this group was a forest-dwelling bird that dispersed into open habitats on numerous occasions.[6]

The cardinalids are part of a larger grouping of American endemic songbirds, Emberizoidea, which also includes the aforementioned thraupids and parulids, as well as icterids (New World blackbirds), passerellids (New World sparrows), and several families that contain one or a couple of genera. Several studies have placed cardinalids as either the sister group to Thraupidae,[9] Mitrospingidae (a small family whose genera were formerly classified as thraupids),[11] or the sister to a clade containing thraupids and mitrospingids.[10] At least one study suggested that cardinalids could treated as a subfamily of Thraupidae.[12]
Phylogeny

Phylogeny of the cardinalid genera after Klicka et al. (2007).[8]

Cardinalidae
Habia‑lineage

Chlorothraupis

Habia

"Masked cardinalid"‑lineage

Piranga

Cardinalis

Caryothraustes

Rhodothraupis

Periporphyrus

Pheucticus‑lineage

Pheucticus

Granatellus‑lineage

Granatellus

"Blue cardinalid"‑lineage

Spiza

Amaurospiza

Cyanoloxia

Cyanocompsa

Passerina




Species list

These 53 species and this sequence are recognized by the IOC as of July 2021:[13]

Image Genus Living species
Piranga leucoptera - Waraira Repano National Park, Venezuela -male-8 (cropped).jpg Piranga Vieillot, 1808
  • Flame-colored tanager, Piranga bidentata
  • Tooth-billed tanager, Piranga lutea
  • Red tanager, Piranga flava
  • Hepatic tanager, Piranga flava
  • Summer tanager, Piranga rubra
  • Rose-throated tanager, Piranga roseogularis
  • Scarlet tanager, Piranga olivacea
  • Western tanager, Piranga ludoviciana
  • White-winged tanager, Piranga leucoptera
  • Red-headed tanager, Piranga erythrocephala
  • Red-hooded tanager, Piranga rubriceps
Habia rubica - Red-crowned Ant-Tanager (male).JPG Habia Blyth, 1840
  • Red-crowned ant tanager, Habia rubica
  • Red-throated ant tanager, Habia fuscicauda
  • Black-cheeked ant tanager, Habia atrimaxillaris
  • Sooty ant tanager, Habia gutturalis
  • Crested ant tanager, Habia cristata
Chlorothraupis carmioli -Costa Rica-8.jpg Chlorothraupis Salvin & Godman, 1883
  • Carmiol's tanager, Chlorothraupis carmioli
  • Olive tanager, Chlorothraupis frenata
  • Lemon-spectacled tanager, Chlorothraupis olivacea
  • Ochre-breasted tanager, Chlorothraupis stolzmanni
Yellow Grosbeak RWD.jpg Pheucticus L. Reichenbach, 1850
  • Yellow grosbeak, Pheucticus chrysopeplus
  • Black-thighed grosbeak, Pheucticus tibialis
  • Golden grosbeak, Pheucticus chrysogaster
  • Black-backed grosbeak, Pheucticus aureoventris
  • Rose-breasted grosbeak, Pheucticus ludovicianus
  • Black-headed grosbeak, Pheucticus melanocephalus
Red-breasted Chat - Oaxaca - Mexico S4E8587 (23040934886).jpg Granatellus Bonaparte, 1850
  • Red-breasted chat, Granatellus venustus
  • Grey-throated chat, Granatellus sallaei
  • Rose-breasted chat, Granatellus pelzelni
Northern Cardinal (34121550154).jpg Cardinalis Bonaparte, 1838
  • Northern cardinal, Cardinalis cardinalis
  • Vermilion cardinal, Cardinalis phoeniceus
  • Pyrrhuloxia, Cardinalis sinuatus
Black-faced Grosbeak - Panama H8O0281 (23184904673).jpg Caryothraustes L. Reichenbach, 1850
  • Black-faced grosbeak, Caryothraustes poliogaster
  • Yellow-green grosbeak, Caryothraustes canadensis
Crimson-colored Grosbeak From The Crossley ID Guide Eastern Birds.jpg Rhodothraupis Ridgway, 1898
  • Crimson-collared grosbeak, Rhodothraupis celaeno
Periporphyrus erythromelas - Red-and-black Grosbeak.jpg Periporphyrus L. Reichenbach, 1850
  • Red-and-black grosbeak, Periporphyrus erythromelas
Amaurospiza moesta - Blackish-blue seedeater (male).JPG Amaurospiza Cabanis, 1861
  • Cabanis's seedeater, Amaurospiza concolor
  • Ecuadorian seedeater, Amaurospiza aequatorialis
  • Blackish-blue seedeater, Amaurospiza moesta
  • Carrizal seedeater, Amaurospiza carrizalensis
DickcisselA.jpg Spiza Bonaparte, 1824
  • Dickcissel, Spiza americana
Cyanocompsa brissonii -Vale do Ribeira, Registro, Sao Paulo, Brasil -male-8.jpg Cyanoloxia Bonaparte, 1850
  • Glaucous-blue grosbeak, Cyanoloxia glaucocaerulea
  • Blue-black grosbeak, Cyanoloxia cyanoides
  • Amazonian grosbeak, Cyanoloxia rothschildii
  • Ultramarine grosbeak, Cyanoloxia brissonii
Blue Bunting From The Crossley ID Guide Eastern Birds.jpg Cyanocompsa Cabanas, 1861
  • Blue bunting, Cyanocompsa parellina
Passerina ciris-20090208.jpg Passerina Vieillot, 1816 North American buntings
  • Blue grosbeak, Passerina caerulea
  • Indigo bunting, Passerina cyanea
  • Lazuli bunting, Passerina amoena
  • Varied bunting, Passerina versicolor
  • Painted bunting, Passerina ciris
  • Rose-bellied bunting, Passerina rositae
  • Orange-breasted bunting, Passerina leclancherii

Natural history

The cardinalids can be found from Canada to northern Argentina and Uruguay, with Central America having the most concentrated amount of species. Species are found year-around in the Central United States and the Eastern United States down to the neotropics. Cardinalids found in the West Indies are non-breeding migrants and those in the Western United States and Canada are breeding migrants.[3] The western tanager is the northernmost species in the family, with their breeding ranges occurring in the southern portions of the Northwest Territories. The northern cardinal has been introduced in Hawaii and Bermuda. They occupy a variety of habitats from forests to grassland and arid scrubland. Most North American cardinalid species migrate south for the winter, whether further south in the continent or extending into the neotropics, except the northern cardinal and pyrrhuloxia which stay year-round. The neotropical species are residential year-round in their range.[2]

Feeding ecology
A scarlet tanager foraging in a flowering dogwood tree.

Cardinals, the dickcissel, seedeaters, buntings, and grosbeaks have the thicker, seed-crushing bills that enabled them to feed heavily on fruits and seeds outside of the breeding season (especially in the winter for northern species like the aforementioned dickcissel and northern cardinal). Once their breeding season begins, members of this group will supplement themselves with invertebrate prey, vital when raising their young and refueling their energetic costs of reproduction and other daily activities. The genera Chlorothraupis, Habia, Piranga, and Granatellus have slightly longer and less deep bills, which their diet mostly consists of insects, fruit, nectar and sap, less so on seeds.[4] Cardinalids typically forage alone low level or on the ground, though some like Piranga and grosbeaks will forage high in the tree canopy. Many will come to birdfeeders especially during the winter.[2]

Breeding and reproduction
A northern cardinal nest showing the nest structure and key features of their egg in Cardinalidae.

Nearly all cardinalids are monogamous breeders and are highly territorial. Despite being monogamous this is only during the breeding season, and each year the birds might partner up with a different bird. The only exception is the dickcissel which is a polygynous species which nest in dense grasses and sedges. Other non-monogamous species include the lazuli and painted buntings which perform extra-copulation with multiple partners. The family is known for their intense brilliant songs. In some species like the lazuli bunting and indigo bunting the bird learn singing by match-based, meaning that first year breeding males will learn by copying the songs of nearby males, as opposed of learning it while they are in the nest. Even more unusual is the females of a few species, such as the scarlet tanager,[1] northern cardinal, pyrrhuloxia, and black-headed grosbeak, which sing as well. In temperate species the breeding season is occurs annual while in tropical species it is year-around. The breeding seasons is in sync with the abundance of insects. Most species build open-cup nests made of grasses and twigs depending on the species. These nests would be in the trees, often high up in the crown. The nest building is done by both partners or by the female alone. The male and female take turns incubating the nest, often the male would feed the female.[2] In a clutch on average there are 1 to 6 six eggs, with tropical species laying the fewest.[14] Cardinalids produce one to three broods per season. As with other passerines, the young are born altricial and fledged between one and two weeks.[14]
Conservation
The black-cheeked ant-tanager is one of the few threatened species of Cardinalidae. Endemic to Osa Peninsula, Costa Rica, this species faces habitat loss.

As of 2021, the IUCN Red List has nearly 82 percent of cardinalids to be least concern. However, there are a handful of species that are of conservation concern. The rose-bellied bunting is an endemic near-threatened species as they are found in a small area of Oaxaca and Chiapas, Mexico; the black-cheeked ant-tanager is another endemic species found in Osa Peninsula in Costa Rica and the carrizal seedeater a critically endangered species found in the spiny bamboo thickets in the understory of deciduous forest in a remote southeastern corner of Venezuela. All of these species are threatened with habitat loss and the confinement within their much smaller range. The IUCN has not yet reevaluate the other species of seedeaters in the genus Amaurospiza.[4]

Despite the vast majority of species being classified as least concern, there has been a growing concern in how the ongoing climate crisis will affect the distribution and migration of many species across the globe. One study led by Dr. Brooke L. Bateman published in July 2020 focused on the risk North American birds will face from climate change and the measures needed to protect them. The first study assessed 604 species from the United States found that if the planet warmed by 3.0 degrees Celsius many species, especially arctic birds, waterbirds, and boreal and western forest birds, will be highly vulnerable to climate change and future conservation efforts will need to be in place.[15] Among the species sampled, the North American species of Piranga and Pheucticus are found to be most climate vulnerable of the cardinalids.[16] These species will either lose some substantial amount of their range or they will migrate up north to escape the sudden change in their habitat.

A possible extinct species is the controversial Townsend's bunting, a supposed enigmatic species related to the dickcissel. The Townsend's bunting is only known from a single type specimen collected from Chester County, Pennsylvania by John Kirk Townsend and described by John James Audubon in 1834. The specimen is housed in the National Museum of Natural History. Genetic work has not been done on this bird, but observation of the plumage has been done. The controversy stems from the uncertain from authors whether the bird is an extinct species, a rare color-variant of the dickcissel, or a hybrid female dickcissel and male blue grosbeak. If the bird is indeed simply a dickcissel it lacks any of the known field characteristics seen in the species in all life stages and sexes.[17]
References

Wells, A. C.; Wells, J. V. (2001). "Tanagers". In Sibley, D.; Elphick, C.; Dunning, J. B. Jr. (eds.). The Sibley Guide to Bird Life & Behavior. New York City: Alfred A. Knopf, Inc. pp. 512–515. ISBN 1400043867.
Thompson, C. W. (2001). "Cardinals and Allies". In Sibley, D.; Elphick, C.; Dunning, J. B. Jr. (eds.). The Sibley Guide to Bird Life & Behavior. New York City: Alfred A. Knopf, Inc. pp. 536–541. ISBN 1400043867.
Dunne, P.; Karlson, K. T. (2021). Bird Families of North America. Location: Mariner Books. pp. 1–288. ISBN 978-0358164074.
Winkler, D. W.; Billerman, S. M.; Lovette, I. J. (4 March 2020). "Cardinals and Allies (Cardinalidae), version 1.0". Birds of the World. doi:10.2173/bow.cardin1.01. S2CID 216193779. Retrieved 7 January 2022.
Guallar, S.; Rueda-Hernández, R.; Pyle, P. (2020). "Preformative molt in Neotropical Cardinalidae". Ornithology Research. 28 (4): 250–257. doi:10.1007/s43388-020-00024-z. hdl:2072/377740. S2CID 228930759.
Guallar, S.; Rueda-Hernández, R.; Pyle, P. (2021). "Evolution of the preformative molt in Cardinalidae correlates with transitions from forest to open habitats". The Auk. 138 (1). doi:10.1093/ornithology/ukaa070.
Hellack, J. J.; Schnell, G. D. (1977). "Phenetic analysis of the subfamily Cardinalinae using external and skeletal characters". The Wilson Bulletin. 89 (1): 130–148. JSTOR 4160878.
Klicka, J.; Burns, K.; Spellman, G. M. (2007). "Defining a monophyletic Cardinalini: a molecular perspective". Molecular Phylogenetics and Evolution. 45 (3): 1014–1032. doi:10.1016/j.ympev.2007.07.006. PMID 17920298.
Barker, F. K.; Burns, K. J.; Klicka, J.; Lanyon, S. M.; Lanyon, I. J. (2014). "New insights into New World biogeography: An integrated view from the phylogeny of blackbirds, cardinals, sparrows, tanagers, warblers, and allies". The Auk: Ornithological Advances. 132 (2): 333–348. doi:10.1642/AUK-14-110.1. S2CID 53058340.
Oliveros, C.H.; et al. (2019). "Earth history and the passerine superradiation". Proceedings of the National Academy of Sciences of the United States. 116 (16): 7916–7925. doi:10.1073/pnas.1813206116. PMC 6475423. PMID 30936315.
Barker, F. Keith; Burns, Kevin J.; Klicka, John; Lanyon, Scott M.; Lovette, Irby J. (2013). "Going to extremes: contrasting rates of diversification in a recent radiation of New World passerine birds". Systematic Biology. 62 (2): 298–320. doi:10.1093/sysbio/sys094. PMID 23229025.
Selvatti, A. P.; Gonzaga, L. P.; de Moraes Russo, C. A. (2015). "A Paleogene origin for crown passerines and the diversification of the Oscines in the New World". Molecular Phylogenetics and Evolution. 88: 1–15. doi:10.1016/j.ympev.2015.03.018. PMID 25837731.
International Ornithological Committee (15 July 2021). "Cardinals, grosbeaks and "tanager" allies". IOC World Bird Names: Version 11.2. WorldBirdNames.org. Retrieved 6 January 2022.
Dittman, D. L.; Cardiff, C. W. (2009). "Grosbeaks and Allies". In Harris, T. (ed.). National Geographic Complete Birds of the World. Washington, D.C.: National Geographic. pp. 362–363. ISBN 9781426204036.
Bateman, B. L.; Wilsey, C.; Taylor, L.; Wu, J.; LeBaron, G. S.; Langham, G. (2020). "North American birds require mitigation and adaptation to reduce vulnerability to climate change". Conservation Science and Practice. 2 (8). doi:10.1111/csp2.242. S2CID 225453243.
Waters, H. (10 October 2019). "The Future for Birds". Audubon. Retrieved 7 January 2022.
Hume, J. P. (2017). Extinct Birds. London: Bloomsbury Publishing. pp. 1–560. ISBN 9781472937469.

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