| Cardinalids | |
|---|---|
.jpeg) | |
| Male breedingdickcissel (Spiza americana). | |
 | |
| Malenorthern cardinal (Cardinalis cardinalis). | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Aves |
| Order: | Passeriformes |
| Superfamily: | Emberizoidea |
| Family: | Cardinalidae Ridgway, 1901 |
| Type species | |
| Loxia cardinalis | |
| Genera | |
Pheucticus | |
Cardinalidae (sometimes referred to as "cardinal-grosbeaks" or simply "cardinals") is a family ofNew World-endemicpasserinebirds that consists ofcardinals,grosbeaks, andbuntings. It also includes several other genera such as the tanager-likePiranga and the warbler-likeGranatellus. Membership of this family is not easily defined by a single or even a set of physical characteristics, but instead by molecular work. Among songbirds, they range from average-sized to relatively large and have stout features. Some species have 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 inopen-cup nests, with parents taking turns incubating the eggs and taking care of their young. Most arearboreal species, although thedickcissel (Spiza americana) is a ground-dwellingprairie bird.
In terms of conservation, most members of this family are consideredleast concern by theIUCN Red List. However, a few birds, such as theCarrizal seedeater (Amaurospiza carrizalensis), are consideredendangered.
The grosbeaks, seedeaters, and cardinals have large bills, whileGranatellus and buntings have small bills. The cardinalid tanagers have stout, near-pointed bills, with some species ofPiranga having serrations along the edge of their upper bills.[1] Bill shape is not always an indicator of species' relationships, however, as the various species of blue cardinalid species, like theblue grosbeak (Passerina caerulea) andCyanoloxia grosbeaks, are related to the buntings. Similarly, the cardinalid tanagers are closer to the cardinals and masked grosbeaks (see more in thesystematics section). The head is medium to large, with a medium neck length. Cardinalid species bodies range from small to medium, with lengths of 11 to 28 cm (4.5 to 11 in). 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 shorter than the others.[2][3][4]
The plumages in cardinalids aresexually dichromatic. In many species, the males are bright red, orange, blue, or black. In mosttemperate species, however, malesmolt between seasons, such that non-breeding males will somewhat resemble the females of their species. These species, such as theindigo bunting (Passerina cyanea), exhibit a complex molt cycle going through four different stages of plumage within their first year of life. From spring to summer, birds start with juvenile plumage to supplemental plumage, then change to a first basic (non-breeding) plumage from fall to winter, and finally reach the first alternate (breeding) plumage. Adults will typically have the basic two molt cycle changing to basic or partial in the late summer or fall, and then back to alternate again in the spring. Males of tropical species retain the same coloration year-round. Females of all species are drabber in coloration and are often a lighter color than the males.
The molting pattern in mostcardinalids exhibits delayed plumage maturation, so that first-year male birds are in non-breeding plumage or an intermediate state.[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 andneotropical species. The second type is an eccentric preformative molt, in which only the outer primary and inner secondary feathers are replaced. This molt is seen in some species ofCyanoloxia andPasserina.[5][6]
Traditionally, members of this group were classified as a tribe of thefinch familyFringillidae (Cardinalini), characterized by heavy, conical, seed-crushing bills. The group consisted of the generaPheucticus,Parkerthraustes,Saltator,Spiza,Cyanocompsa,Cyanoloxia,Porphyrospiza,Passerina,Caryothraustes,Periporphyrus, andCardinalis. The issue that taxonomists faced was that there were no unifying morphological traits that were in agreement across various studies.[7] In 2007, a mitochondrial DNA study by Klicka, Burns and Spellman sampling all of the genera above and 34 of the total 42 species, found that the generaParkerthraustes,Saltator, andPorphyrospiza were not members of the cardinal lineage, but instead are found throughout in the tanager lineage (Thraupidae). Several genera classified as thraupids at the time—Piranga,Habia,Chlorothraupis, andAmaurospiza—were found to be part of cardinalid radiation. In addition, the genusGranatellus, originally classified as aparulid warbler, was also found to be part of Cardinalidae.[8] The study found that with these new relationships, Cardinalidae can be classified into six subgroups, which subsequent studies have supported. The six subclades consists of thePheucticus lineage, theGranatellus lineage, the “blue” lineage (Spiza,Cyanoloxia,Amaurospiza,Cyanocompsa, andPasserina), theHabia lineage (Habia andChlorothraupis), the “masked” lineage (Caryothraustes,Periporphyrus, andCardinalis), and thePiranga lineage (Piranga andDriophlox).[8][9] These subclades and membership of these genera have been widely supported in subsequent studies.[10][11] A 2021 paper by Guallar et al. 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 asicterids (New World blackbirds),passerellids (New World sparrows), and several smaller families that contain one or a couple of genera. Several studies have placed cardinalids as either the sister group to Thraupidae,[10]Mitrospingidae (a small family whose genera were formerly classified as thraupids),[12] or as a sister to a clade containing thraupids and mitrospingids.[11] At least one study suggested that cardinalids could be treated as a subfamily of Thraupidae.[13]
The genus levelcladogram of the Cardinalidae shown below is based onmolecular phylogenetic study published in 2024 which analyzed DNA sequences flankingultraconserved elements (UCEs).[9] The number of species in each genus is taken from the list maintained byFrank Gill,Pamela C. Rasmussen and David Donsker on behalf of theInternational Ornithological Committee (IOC).[14]
| Cardinalidae |
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The following 53 species and 14 genera are recognized by the IOC as of July 2024:[14]
| Image | Genus | Living species |
|---|---|---|
 | PheucticusL. Reichenbach, 1850 |
|
.jpg) | GranatellusBonaparte, 1850 |
|
 | SpizaBonaparte, 1824 |
|
 | CyanoloxiaBonaparte, 1850 |
|
.JPG) | AmaurospizaCabanis, 1861 |
|
 | CyanocompsaCabanas, 1861 |
|
 | PasserinaVieillot, 1816 North American buntings |
|
.JPG) | HabiaBlyth, 1840 |
|
 | ChlorothraupisSalvin & Godman, 1883 |
|
.jpg) | CardinalisBonaparte, 1838 |
|
.jpg) | CaryothraustesL. Reichenbach, 1850 |
|
 | PeriporphyrusL. Reichenbach, 1850 |
|
 | DriophloxScott, BF, Chesser, Unitt & Burns, KJ, 2024 |
|
.jpg) | PirangaVieillot, 1808 |
|
Thecardinalids can be found fromCanada to northernArgentina andUruguay, withCentral America having the largest concentration of species. Species are found year-round in theCentral United States and theEastern United States down to theneotropics. Cardinalids found in theWest Indies are non-breeding migrants, and those in theWestern United States and Canada are breeding migrants.[3] Thewestern tanager (Piranga ludoviciana) is the northernmost species in the family; their breeding ranges occur in southern portions of theNorthwest Territories. Thenorthern cardinal (Cardinalis cardinalis) has been introduced inHawaii andBermuda. They occupy a variety of habitats, including forests, grasslands, and arid scrublands. Most North Americancardinalid species migrate south for the winter, whether further south in the continent or extending into the neotropics, except the northern cardinal andpyrrhuloxia, which stay year-round. The neotropical species are resident year-round in their range.[2]
.jpg)
Cardinals, the dickcissel, seedeaters, buntings, and grosbeaks have the thicker, seed-crushing bills that enable them to feed heavily on fruits and seeds outside of the breeding season (especially in the winter for northern species like the dickcissel mentioned above and northern cardinal). Once their breeding season begins, members of this group will supplement themselves withinvertebrate prey, vital when raising their young and refueling the energetic costs of reproduction and other daily activities. The generaChlorothraupis,Habia,Piranga, andGranatellus have slightly longer and less deep bills, whose diet mostly consists ofinsects,fruit,nectar and sap, less so onseeds.[4]Cardinalids typically forage alone, low level, or on the ground, though some likePiranga and grosbeaks will forage high in the tree canopy. Many will come tobird feeders, especially during the winter.[2]
Nearly allcardinalids aremonogamous breeders, and are highly territorial. Most species are monogamous during the breeding season, and each year, birds may find different partners. The only exception is thedickcissel, which is apolygynous species that nests in dense grasses andsedges. Other non-monogamous species include thelazuli andpainted bunting, which perform extra-copulation with multiple partners. The family is known for their intense, brilliant songs. In some species like thelazuli bunting and indigo bunting, the birds learn singing by match-based learning, meaning that first-year breeding males will learn by copying the songs of nearby males, as opposed to learning it while they are in the nest. Even more unusual is the females of a few species, such as thescarlet tanager,[1] northern cardinal,pyrrhuloxia, andblack-headed grosbeak, which sing as well. In temperate species, the breeding season occurs annually, while in tropical species it is year-round. The breeding seasons are in sync with the abundance of insects. Most species build open-cup nests made ofgrasses andtwigs, 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 feeds the female.[2] A clutch averages one to six eggs, with tropical species laying the fewest.[15]Cardinalids produce one to three broods per season. As with otherpasserines, the young are bornaltricial and fledged between one and two weeks.[15]
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As of 2021, the IUCN Red List has nearly 82 percent of cardinalids to beleast concern. However, there are a handful of species that are ofconservation concern. Therose-bellied bunting is an endemicnear-threatened species as they are found in a small area ofOaxaca andChiapas,Mexico; the black-cheeked ant-tanager is another endemic species found inOsa Peninsula inCosta Rica and the carrizal seedeater a critically endangered species found in the spinybamboo thickets in the understory ofdeciduous forest in a remote southeastern corner ofVenezuela. All of these species are threatened with habitat loss and confinement within their much smaller range. The IUCN has not yet reevaluate the other species of seedeaters in the genusAmaurospiza.[4]
Despite the vast majority of species being classified asleast concern, there is growing recognition that theclimate crisis may impact the distribution and migration of many cardinalid species. One study led by Dr. Brooke L. Bateman, published in July 2020, focused on the risk North American birds will face fromclimate change, as well as the measures required to protect them. The first study assessed 604 species from theUnited States, and found that if the planet warmed by 3.0 degrees Celsius, many species, especially arctic birds, waterbirds, and boreal and western forest birds, would be highlyvulnerable to climate change. Future conservation efforts will need to be in place.[16] Among the species sampled, the North American species ofPiranga andPheucticus are found to be most climate vulnerable of thecardinalids.[17] These species will either lose a substantial amount of their range or they will migrate north to escape the sudden change in their habitat.
A possible extinct species is the controversialTownsend's bunting (Spiza townsendi), a supposed enigmatic species related to thedickcissel. The Townsend's bunting is only known from a single type specimen collected fromChester County,Pennsylvania byJohn Kirk Townsend and described byJohn James Audubon in 1834. The specimen is housed in theNational Museum of Natural History. Genetic analysis has not been done on this specimen, but a study of the plumage has been conducted. Researchers are unsure about the specimen's status as an extinct species, a rare color-variant of the dickcissel, or a hybrid (of a female dickcissel and maleblue grossbeak). If the bird is indeed a dickcissel, however, it lacks any of the known field characteristics seen in the species in all its life stages and sexes.[18]
