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Cardinalidae

Cardinalidae is a family of New World passerine birds that includes cardinals, grosbeaks, buntings, and several tanager-like genera such as ant-tanagers and saltators. Comprising 56 species across 13 genera, the family is characterized by medium-sized songbirds with robust, cone-shaped bills adapted for cracking seeds and fruits, and plumage featuring bold patterns in primary colors like red, yellow, blue, and black.^[1] These birds exhibit significant sexual dimorphism, with males typically displaying more vivid coloration to attract mates, while females are often duller for camouflage during nesting. Their diet primarily consists of seeds, insects, berries, and nectar, supplemented by foraging in pairs or small flocks. Cardinalidae species are renowned for their rich, whistled songs, which play a key role in territorial defense and mate attraction, contributing to their popularity among birdwatchers.^[2]^[3] The family is distributed across the Americas, ranging from southern Canada and the United States through Central America to northern Argentina and Chile, occupying diverse habitats such as open woodlands, shrublands, grasslands, forest edges, and even suburban areas, though they generally avoid dense rainforests and arid deserts. While most species are resident, some northern populations migrate southward seasonally. Conservation status varies, with a few species like the black-cheeked ant-tanager facing threats from habitat loss, but many remain common due to adaptability to human-modified landscapes.^[1]^[2]

Taxonomy and systematics

Classification history

In the 18th and 19th centuries, birds now recognized as belonging to Cardinalidae were classified within the family Fringillidae, primarily due to their seed-eating habits and possession of stout, conical bills adapted for cracking seeds.^[4] Carl Linnaeus, for instance, placed the northern cardinal (Cardinalis cardinalis) in the genus Loxia (now restricted to crossbills) under Fringillidae in his Systema Naturae (1758). This grouping reflected early taxonomic emphasis on dietary and morphological similarities with true finches, though it encompassed a heterogeneous assemblage of New World passerines.^[4] During the 20th century, classifications shifted as ornithologists recognized distinctions in skeletal morphology, vocalizations, and biogeography, leading to the reassignment of cardinal-like birds to the family Emberizidae (American sparrows and buntings).^[4] Debates persisted regarding close affinities with Thraupidae (tanagers), particularly for genera like Piranga and Habia, which some studies suggested belonged there based on plumage patterns and habitat preferences.00002-3) By the late 20th century, the American Ornithologists' Union (AOU, now American Ornithological Society) treated the group as the tribe Cardinalini within Emberizidae, acknowledging its distinctiveness but retaining a broader familial context.^[4] A landmark molecular study by Klicka, Burns, and Spellman in 2007 utilized mitochondrial DNA sequences from all recognized genera and 34 of 42 species to delineate a monophyletic Cardinalini, firmly establishing Cardinalidae as a distinct family within the superfamily Emberizoidea. This analysis resolved longstanding ambiguities by excluding tanager-like genera from Thraupidae and confirming cardinalids as a cohesive clade sister to Thraupidae, supported by shared morphological traits like robust skulls and carotenoid-based plumage. The work prompted the AOU to elevate Cardinalidae to family status in its 2009 supplement to the Check-list of North American Birds. Recent molecular advancements have further refined the family's internal structure. In 2024, a phylogenomic study incorporating whole-genome data across Passeriformes, including Cardinalidae, highlighted subclade relationships and supported ongoing taxonomic adjustments within Emberizoidea. Complementing this, Scott et al. (2024) introduced the genus Driophlox for four ant-tanager species previously in Habia, based on integrated genetic (mitochondrial and nuclear loci) and vocalization analyses that revealed paraphyly in the old arrangement.^[5] The family name Cardinalidae derives from the genus Cardinalis, itself from the Latin cardinalis (meaning "principal" or "chief"), alluding to the bright red plumage of species like the northern cardinal, which evoked the scarlet robes and caps worn by Catholic cardinals. This etymology, first applied by Linnaeus in 1758, underscores the visual prominence of these birds in early European descriptions of New World avifauna.

Phylogenetic relationships

The family Cardinalidae occupies a position within the order Passeriformes, specifically in the suborder Passeri (oscine passerines) and the superfamily Emberizoidea, which encompasses several New World songbird lineages including emberizids and icterids.^[1]^[6] Phylogenetic debates persist regarding the closest relatives of Cardinalidae, with molecular evidence supporting it as sister to Thraupidae (tanagers) in some analyses, while others position Mitrospingidae (mitrospingid tanagers) as the nearest kin, based on shared ultraconserved DNA elements and mitochondrial markers.^[7]^[8] A 2024 multilocus DNA study reinforced these ties by identifying common genetic signatures in nuclear and mitochondrial sequences across these groups, highlighting convergent adaptations in vocalization and plumage.^[9] Within Cardinalidae, molecular phylogenies divide the family into six main subclades: (1) the basal Pheucticus grosbeaks, (2) Piranga tanagers (previously classified in Thraupidae), (3) Cardinalis cardinals, (4) Granatellus chats, (5) Saltator grosbeaks, and (6) Emberizoides and buntings (including Passerina and allies).^[8] This structure stems from a 2022 phylogeny that rearranged Pheucticus as the earliest diverging lineage, attributing prior misplacements to convergences in vocal repertoires and morphological traits like bill shape, resolved via ultraconserved elements analysis.^[10] Molecular clock estimates date the origin of Cardinalidae to the Miocene approximately 12 million years ago, with major diversification occurring in the Holocene across the New World, driven by climatic shifts and habitat fragmentation.^[11] The 2025 eBird/Clements taxonomy update added a net 22 species globally through splits and lumps elsewhere but introduced no major changes to the core Cardinalidae phylogeny, such as splits or lumps, following the 2024 integrations.^[12]

Genera and species

The family Cardinalidae encompasses 14 genera and 53 species, all endemic to the New World, ranging from southern Canada and the United States through Central America to Argentina and Tierra del Fuego.^[13] In 2024, the genus Driophlox was newly recognized with four species split from Habia based on genetic analyses revealing distinct evolutionary lineages.^[5] The genera and their species are listed below, with representative examples and native ranges (as of IOC World Bird List v14.1, July 2024; consistent with Clements v2025).

Genus	Species Count	Example Species	Native Range
Amaurospiza	4	Blue Seedeater (A. moesta)	Neotropical
Cardinalis	3	Northern Cardinal (C. cardinalis)	Nearctic to Neotropical
Caryothraustes	2	Yellow-shouldered Grosbeak (C. humeralis)	Neotropical
Chlorothraupis	4	Buff-throated Ant-tanager (C. rufiventer)	Neotropical
Cyanocompsa	1	Blue Bunting (C. parellina)	Neotropical
Cyanoloxia	4	Blue-black Grosbeak (C. cyanoides)	Neotropical
Driophlox	4	Crested Ant-tanager (D. cristata)	Neotropical
Emberizoides	2	Campo Troupial (E. herbicola)	Neotropical
Granatellus	3	Red-breasted Chat (G. venustus)	Neotropical
Habia	1	Red-throated Ant-tanager (H. fuscicauda)	Neotropical
Periporphyrus	2	Finsch's Tanager (P. erythromelas)	Neotropical
Pheucticus	5	Evening Grosbeak (P. melanocephalus)	Nearctic
Piranga	11	Scarlet Tanager (P. olivacea)	Nearctic to Neotropical
Saltator	16	Buff-throated Saltator (S. maximus)	Neotropical
Spiza	1	Dickcissel (S. americana)	Nearctic
Volatinia	1	Blue-black Grassquit (V. jacarina)	Neotropical

Physical characteristics

Morphology and size

Members of the Cardinalidae family display considerable variation in size, ranging from 11 to 24 cm in total length and 10 to 60 g in weight, reflecting adaptations to diverse ecological niches within the New World. The smallest species include those in the genus Granatellus, such as the rose-breasted chat (Granatellus pelzelni), which measures 12–12.5 cm in length and weighs 10–12.5 g, while the largest are found in the genus Saltator, exemplified by the cinnamon-bellied saltator (Saltator cinnamomeus) at 22–24 cm in length and up to 53 g.^[14]^[15]^[16] The overall body form in Cardinalidae is robust and compact, suited to an arboreal lifestyle, with a stocky build, short neck, and strong, medium-length legs that facilitate perching on branches and foraging in foliage. Wings are short and rounded, typically featuring 10 primaries, which enable agile maneuvers and short-distance flights rather than long migrations in most species. The tail is medium in length, often squared or slightly rounded, aiding in balance during perching and brief aerial pursuits.^[1]^[17]^[18] Bill morphology is a defining feature, generally stout and conical for cracking seeds, but showing adaptive variations across genera that correlate with diet. In seed specialists like grosbeaks (e.g., Pheucticus) and cardinals (Cardinalis), the bill is large, deep, and powerful, with a convex culmen and strong jaw muscles for handling hard seeds. In contrast, chats (Granatellus) and some buntings (Passerina) have slenderer, less deep bills suited for gleaning insects or probing for nectar, while tanagers in Piranga have stouter bills adapted for consuming fruit and insects.^[19]^[1]^[17] Sexual size dimorphism is minimal throughout the family, with males averaging only slightly larger than females in select genera such as Cardinalis, where males measure 22.2–23.5 cm compared to females at 20.9–21.6 cm.^[20]

Plumage and coloration

The family Cardinalidae exhibits pronounced sexual dichromatism in most genera, with males displaying vibrant plumage colors such as brilliant red, yellow, or blue to attract mates and defend territories, while females are typically duller with olive, brown, or gray tones that provide camouflage.^[21] For example, in the genus Cardinalis, the male Northern Cardinal (C. cardinalis) features vivid red body plumage, a black face mask, and a prominent crest, contrasting with the female's buffy brown overall coloration accented by subtle red tinges on the wings, crest, and tail.^[21] Similar patterns occur in Piranga tanagers, where males like the Scarlet Tanager (P. olivacea) are uniformly bright red, and females are yellowish-olive with faint streaking on the underparts. In Passerina buntings, males attain striking blue or purple hues, such as the Indigo Bunting (P. cyanea), while females are streaked brown. Grosbeaks in Pheucticus show bold black-and-white patterns with yellow accents in males, opposed to the streaked brown females. These bright male colors are driven by sexual selection, enhancing mate attraction and territorial displays, whereas female plumage evolves under natural selection pressures like predation risk.^[22] Plumage types vary across genera, reflecting diverse evolutionary adaptations. Species in Cardinalis and some grosbeaks possess distinctive crests of elongated feathers on the head, which can be raised during displays.^[21] Buntings in Passerina and Spiza typically feature streaked juvenile and female plumages for concealment, while Piranga species exhibit tanager-like patterns with spotted or streaked underparts in females. An exception is the genus Saltator, where species are largely monochromatic, with both sexes sharing grayish or olive plumage lacking strong dimorphism.^[23] Many Cardinalidae species also reflect ultraviolet (UV) light in their feathers, which is perceptible to avian vision and may contribute to intraspecific signaling, though this varies by species and patch. Molting patterns in Cardinalidae involve seasonal changes that maintain plumage condition and coloration. Adults undergo a complete prebasic molt annually, replacing all body feathers and flight feathers after breeding to achieve definitive bright or cryptic appearances. Juveniles enter a partial preformative molt shortly after fledging, typically replacing body feathers, wing coverts, and tertials but retaining juvenile remiges; this is the ancestral condition, with extensions to include primaries in species adapted to open habitats. Juvenile plumage often mimics that of adult females in dimorphic species, delaying full maturation and reducing aggression from adults. Some species, like the Indigo Bunting, exhibit eccentric molting with delayed replacement of certain wing feathers, contributing to gradual color transitions.^[24] These molts ensure functional plumage, with habitat openness influencing molt extent—non-forest species invest more in complete preformative molts.

Distribution and habitat

Geographic range

The Cardinalidae family is exclusively native to the New World, with a broad geographic distribution extending from the northern limits in southern Canada and Alaska—where species such as the Evening Grosbeak (Coccothraustes vespertinus) breed in coniferous forests—to the southern extremes in northern Argentina, exemplified by the Grayish Saltator (Saltator coerulescens), which inhabits semi-open woodlands and scrub across much of South America.^[25]^[26] This latitudinal span covers diverse ecosystems across the Nearctic and Neotropical realms, reflecting the family's adaptation to a variety of temperate and tropical environments, though no species occur in the Old World. Of the 53 species in the family, about 10 are primarily associated with the Nearctic region, including mainly migratory grosbeaks and cardinals such as the Northern Cardinal (Cardinalis cardinalis), Rose-breasted Grosbeak (Pheucticus ludovicianus), and Black-headed Grosbeak (Pheucticus melanocephalus), which breed across North America and often winter in the Neotropics.^[27]^[18] In contrast, the Neotropical region supports the vast majority, with 43 species exhibiting the highest diversity in Central and South America, particularly concentrated in the Amazon Basin and along the Andean slopes, where genera like Chlorothraupis (ant-tanagers) and Saltator (saltators) thrive in forest edges, second growth, and humid lowlands.^[28]^[29] While the family has few strict endemics to the Caribbean islands—such as the formerly debated placement of the Puerto Rican Spindalis (Spindalis portoricensis), now classified elsewhere—several species like the Summer Tanager (Piranga rubra) occur on these islands as residents or vagrants.^[30] Historically, range shifts have occurred within the family; notably, the Northern Cardinal has expanded northward across the northeastern United States and into southern Canada during the 20th century, facilitated by the proliferation of bird feeders, milder winters, and habitat modifications from human activity.^[31]

Habitat preferences

Members of the Cardinalidae family inhabit a diverse array of environments across the Americas, ranging from dense tropical forests to open grasslands and arid scrublands. The ancestral habitat of the family is inferred to have been forested, with subsequent evolutionary radiations leading to occupations of more open and semi-open areas. For instance, tanagers in the genus Piranga prefer the canopies of tropical dry and wet forests, where they forage in the upper layers of vegetation. In contrast, many buntings (Passerina spp.) favor shrublands, scrub, and grasslands, such as the Dickcissel (Spiza americana), which thrives in open prairies and agricultural fields. Cardinals (Cardinalis spp.) commonly occupy forest edges, secondary growth, and suburban thickets, demonstrating a tolerance for human-modified landscapes.^[1] Elevational preferences within Cardinalidae span from sea level to high montane zones, with some species occurring up to 1,500 m in the Andes, often in montane forests and shrubby slopes. Microhabitat selections vary by foraging and nesting behaviors; arboreal species such as certain grosbeaks (Pheucticus spp.) utilize mid- to upper forest strata, while ground-foraging seedeaters prefer open fields with scattered shrubs. Nesting often occurs in dense understory vegetation for many taxa, providing cover from predators, whereas some, like saltators (Saltator spp.), select thickets along forest edges. A few species exhibit nectar-feeding tendencies in flowering savannas, supplementing their diet during periods of abundant blooms.^[1]^[32] Adaptations to disturbed habitats are evident in several genera, including cardinals and buntings, which readily exploit agricultural edges, second-growth woodlands, and urban gardens, contributing to their range expansions. However, old-growth forest specialists, such as some Piranga tanagers and grosbeaks, show sensitivity to habitat fragmentation, preferring intact canopies with minimal disturbance. Climate influences favor warm-temperate to tropical zones for most species, though arid-adapted forms like the Varied Bunting (Passerina versicolor) endure desert washes and xeric scrub through behavioral adjustments to water scarcity and intense solar exposure. These preferences reflect phylogenetic shifts from closed to open environments, correlating with morphological and behavioral traits.^[33]

Migration patterns

Approximately 20% (about 11 of 53 species) in the Cardinalidae family are fully migratory, with the remainder exhibiting partial migration or remaining sedentary throughout their ranges.^[34] For example, the Scarlet Tanager (Piranga olivacea) breeds across much of North America and winters in northern South America, while the Summer Tanager (Piranga rubra) migrates from breeding grounds in the southern United States and Mexico to wintering areas in Central and South America.^[35]^[36] In contrast, species like the Northern Cardinal (Cardinalis cardinalis) and Pyrrhuloxia (Cardinalis sinuatus) are year-round residents in their temperate and subtropical habitats, showing only limited dispersal movements averaging 100-150 km in some individuals.^[37]^[38] Migratory routes vary by species and population, with many North American breeders following trans-Gulf paths across the Gulf of Mexico or overland Central American flyways to reach Neotropical wintering grounds. The Rose-breasted Grosbeak (Pheucticus ludovicianus), for instance, typically migrates southeast from breeding areas in eastern North America, crossing the Gulf in non-stop flights of up to 800 km, though some individuals travel overland around its western edge.^[39] Similarly, the Painted Bunting (Passerina ciris) shows population-specific patterns, with western birds staging in Arizona and northwestern Mexico before continuing south, while eastern populations migrate directly across the Caribbean to winter in southern Florida, the Bahamas, and Cuba.^[40] Temperate species like the Blue Grosbeak (Passerina caerulea) often undertake shorter-distance migrations within the continent, moving southward to Mexico and Central America.^[41] Spring northbound migration generally occurs from March to May, with fall southbound movements peaking between August and October; many species, including tanagers and buntings, travel nocturnally to reduce predation risk and conserve energy.^[42]^[39] Photoperiod changes serve as primary triggers, prompting physiological preparations such as fat accumulation to fuel extended flights, with migrants like grosbeaks building reserves for non-stop journeys exceeding 1,000 km.^[43] In mountainous regions, some species engage in altitudinal migration, shifting downslope during the non-breeding season; the Hepatic Tanager (Piranga flava), for example, exhibits vertical movements in its resident subspecies across elevational gradients in the southwestern United States and Mexico.^[44] Most tropical members of the family, including buntings and saltators such as the Blue-black Grosbeak (Cyanocompsa cyanoides), remain sedentary within their local ranges year-round.^[1]

Natural history

Diet and foraging

Members of the Cardinalidae family are generally omnivorous, with diets varying across genera but commonly consisting of seeds, fruits, insects, and occasionally nectar. In cardinals (Cardinalis), vegetable matter dominates, comprising about 70% of annual intake, including weed seeds, grains, and berries like sunflower seeds and dogwood fruits, whereas in grosbeaks (Pheucticus), the diet is more balanced, with vegetable matter comprising around 40-50% annually.^[45]^[46] Insect consumption, such as beetles, larvae, and spiders, increases during the breeding season to around 50% in some species, providing essential protein. Tanager-like species in genera such as Piranga and Granatellus emphasize insects (e.g., Lepidoptera larvae and bees) year-round, supplemented by fruits like mulberries and serviceberries, while some cardinals opportunistically feed on nectar from flowers such as cardinal flower.^[47]^[48]^[49] Foraging techniques in Cardinalidae include gleaning from foliage, ground probing, and hawking insects, adapted to arboreal or terrestrial habitats. Cardinals and grosbeaks often crack hard seeds and nuts using their robust conical bills, foraging solitarily or in pairs within territories, while tanagers hover briefly or sally to capture flying insects in the canopy.^[45]^[50]^[47] Species like the Dickcissel (Spiza americana) employ ground gleaning for seeds and insects, typically in flocks during non-breeding periods to exploit communal food patches in grasslands.^[51] Pyrrhuloxia (Cardinalis sinuatus) and similar taxa show opportunistic behaviors, shifting between arboreal fruit-picking and ground-level insect hunting based on availability.^[50] Dietary composition shifts seasonally, with insects forming a larger proportion in summer across the family to meet higher energy demands, while seeds and fruits predominate in winter for sustenance.^[45] Social foraging varies; non-migratory cardinals maintain solitary or paired routines year-round, whereas species like the Dickcissel form large winter flocks for efficient resource location.^[51]^[45] These patterns reflect adaptations such as the family's characteristic thick bills, which facilitate processing tough seeds and fruits, as detailed in morphological descriptions.^[1]

Breeding biology

Members of the Cardinalidae family exhibit predominantly socially monogamous mating systems, forming pair bonds for the duration of a breeding season, though some species display genetic polygyny through extra-pair copulations or occasional polygynous arrangements where one male mates with multiple females.^[1] In the Northern Cardinal (Cardinalis cardinalis), pairs often remain together for multiple seasons or even lifelong, providing a notable example of pair fidelity within the family.^[20] However, in species like the Indigo Bunting (Passerina cyanea), social monogamy coexists with extra-pair fertilizations, affecting up to 15-20% of offspring in some populations.^[52] Courtship behaviors are relatively simple, with males relying on vivid plumage displays and persistent singing to attract females, often accompanied by mate-feeding where the male presents food to the female as a precopulatory gesture.^[53] Unlike some avian families, cardinalids lack elaborate dances or aerial displays, emphasizing vocal and visual signals instead.^[54] Breeding seasons vary by latitude: in temperate regions, it occurs from spring through summer (typically April to July), influenced by increasing day length and food availability, while tropical species may breed year-round with peaks tied to resource abundance.^[55] Pairs typically produce 1-3 broods per year, with nesting commencing shortly after pair formation. Nests are open cups constructed primarily by the female, using twigs, grasses, bark strips, and rootlets, often lined with finer materials such as hair or moss; they are sited in dense shrubs, vines, or low to mid-canopy branches of trees, 1-4 meters above ground.^[54] Clutches consist of 1-6 eggs, averaging 3-4, which are pale blue to white or greenish with brownish spots or blotches for camouflage.^[56] Incubation lasts 11-14 days, primarily by the female but with some male assistance in many species, ensuring the altricial young hatch in a helpless state.^[57] Parental care is biparental, with both sexes feeding the nestlings a diet rich in insects to support rapid growth; the male often takes a larger role in provisioning fledglings after they leave the nest at 9-12 days post-hatching.^[58] This shared investment enhances fledgling survival, though variations exist, such as reduced male care in polygynous contexts among some buntings.^[59] Nestlings briefly shift to a higher-protein diet compared to adult foraging, emphasizing arthropods during this vulnerable period. Members of the Cardinalidae family produce a diverse vocal repertoire that includes songs and calls adapted for various communicative purposes. Songs are typically melodious whistles, as exemplified by the Northern Cardinal's (Cardinalis cardinalis) clear, repeated phrases such as "cheer-cheer" or "birdie-birdie," delivered in series that can last several seconds and span frequencies from 1 to 8 kHz.^[60]^[61] Calls often consist of sharp chip notes used for alarm or contact, with the Northern Cardinal exhibiting at least 16 distinct call types, including metallic "chip" sounds to signal potential threats.^[60] In grosbeaks, such as the Black-headed Grosbeak (Pheucticus melanocephalus), songs are more complex, featuring repertoires of 14–37 syllable types per individual (mean 24.5), organized into variable phrases that enhance territorial signaling.^[62] Similarly, Rose-breasted Grosbeaks (Pheucticus ludovicianus) possess repertoires of 15–23 syllables, producing rich, robin-like warbles that are sweeter and more varied.^[63]^[64] These vocalizations serve multiple functions, including territory defense through persistent dawn singing by males, which advertises ownership and deters rivals during the breeding season.^[65]^[53] Mate attraction is facilitated by elaborate songs, often performed from prominent perches, while flock coordination occurs via short calls in non-breeding aggregations. Duetting, where paired individuals alternate songs, is observed in species like the Northern Cardinal, strengthening pair bonds and potentially aiding in joint territory maintenance; females contribute softer responses to the male's louder phrases.^[66]^[61] Songs also play a role in courtship displays, where they accompany visual behaviors to solicit mating responses.^[67] Socially, Cardinalidae species are generally solitary or paired during the breeding season, with males vigorously defending territories through vocal and physical confrontations against intruders.^[20] In winter, they form flocks typically numbering 10–20 individuals, though groups can reach up to 100 in some areas, allowing for communal roosting and resource sharing without intense competition.^[68]^[20] Aggression is expressed through displays such as wing-flashing and bill-wiping, which intimidate rivals during territorial disputes.^[69] Non-vocal communication supplements these sounds, including tail-pumping or flicking to signal alarm and crest-raising to convey excitement or dominance in interactions.^[20] Paired birds engage in allopreening, mutual grooming that reinforces bonds and maintains plumage hygiene.^[70] Variations in vocalizations include rare instances of mimicry, as Northern Cardinals generally do not imitate other species' sounds, unlike more versatile mimics in related families.^[71] Dialects occur in some populations, such as the Dickcissel (Spiza americana), where song structure shows geographic patterns of similarity, with local "dick" phrases varying in syllable count and buzz quality across neighborhoods.^[72]^[73]

Conservation

Population status

The Cardinalidae family comprises 52 species (as of 2025 taxonomic lists), of which approximately 88% (46 species) are assessed as Least Concern on the IUCN Red List, reflecting their generally wide distributions and stable or increasing populations for many common taxa.^[74] Approximately 12% (6 species) are classified as Near Threatened, 4% (2 species) as Endangered, with one species considered Critically Endangered by experts though not yet formally assessed by the IUCN, highlighting localized vulnerabilities despite the family's overall resilience. The species considered Critically Endangered is the Carrizal Seedeater (Amaurospiza carrizalensis), restricted to a small area in southeastern Venezuela where ongoing threats limit its viability.^[75] Populations of widespread species remain stable or are expanding, particularly in North America; for instance, the Northern Cardinal (Cardinalis cardinalis) has shown an increasing trend there, contributing to a global family estimate exceeding 100 million individuals, driven largely by abundant common species.^[76] However, about 15% of species exhibit population reductions, such as the Vermilion Cardinal (Cardinalis phoeniceus), which has declined due to habitat loss and trapping for the pet trade. The 2025 IUCN Red List update, via BirdLife International assessments, indicates no major category shifts from 2021 levels, though continued monitoring is essential given fluctuating trends in some taxa.^[77] Regionally, North American Cardinalidae populations are largely stable and abundant, supported by diverse habitats and conservation efforts, whereas Neotropical hotspots show greater vulnerability, with approximately 20% of bunting species (Passerina spp.) experiencing declines linked to habitat pressures in breeding and wintering areas. These trends are informed by eBird abundance data and BirdLife International monitoring, emphasizing the need for targeted surveys in understudied tropical regions.^[78]

Threats and measures

Habitat loss and degradation due to deforestation represent the primary threat to many species in the Cardinalidae family, particularly in tropical regions where clearance for agriculture and urbanization fragments forests essential for tanagers and grosbeaks. For instance, ongoing Amazon deforestation has contributed to population declines in species like the Yellow Cardinal (Gubernatrix cristata), which requires grassland habitats.^[79] Agricultural expansion and associated pesticide use further exacerbate risks by reducing insect and seed availability, key components of the diet for seedeaters and buntings.^[1] Climate change poses additional challenges, including altered migration patterns and increased drought frequency that affect breeding success in species such as the Scarlet Tanager (Piranga olivacea), which shows vulnerability to prolonged dry periods impacting insect prey abundance. Collisions with windows and predation by domestic cats are significant for migratory cardinalids, with estimates indicating that cats alone kill billions of birds annually in North America, including grosbeaks and orioles during stopovers. Trapping for the international pet trade threatens several species, notably the Vermilion Cardinal (Cardinalis phoeniceus) and Yellow Cardinal (Gubernatrix cristata), where illegal capture has led to rapid population declines in South America.^[80]^[77]^[79] Conservation efforts include the establishment of protected areas in the Neotropics, such as national parks in Venezuela and Brazil that safeguard habitats for threatened species like the Carrizal Seedeater (Amaurospiza carrizalensis), whose range is limited to a small area threatened by hydroelectric development. Reforestation initiatives and sustainable agriculture programs aim to restore fragmented landscapes, while monitoring through platforms like eBird and Partners in Flight surveys track population trends across the Americas. The Northern Cardinal (Cardinalis cardinalis) has maintained stable populations partly due to supplemental feeding in urban and suburban areas, which provides consistent seed resources amid habitat changes.^[75]^[81] International measures encompass CITES Appendix II listings for traded species like the Yellow Cardinal to regulate capture and export, alongside partnerships led by BirdLife International that support on-the-ground protection in key ranges. Despite these actions, gaps persist in studying tropical populations, where data scarcity hinders threat assessment, and enhanced climate modeling beyond 2025 is needed to predict range shifts for vulnerable taxa.^[79]

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Rose-breasted Chat - Granatellus pelzelni - Birds of the World
Oct 28, 2025 · Identification. 12–12.5 cm; 10–12.5 g. Long tail frequently cocked and fanned. Male has black head, white supercilium behind eye ...Identification · Systematics History · VocalizationsMissing: size | Show results with:size
[16]
Cinnamon-bellied Saltator | Mexican Birds.org
The Cinnamon-bellied Saltator has an average length of 22 cm (8.7 inches) to 24 cm (9.4 inches), and sexes do not differ in size. They are grey above and ...
[17]
Slate-colored Grosbeak Saltator grossus - Birds of the World
Identification. 19–20·5 cm; 41–53 g (Venezuela and Suriname). Male nominate race is mostly uniformly slaty blue above and ...Missing: size | Show results with:size
[18]
Bird Cardinalidae - Cardinals, Grosbeaks & Allies - Fat Birder
Cardinalidae is a family of New World-endemic passerine birds that consists of cardinals, grosbeaks, and buntings. Species Links. Given the number of species in ...
[19]
Northern Cardinal Identification - All About Birds
Relative Size · Measurements. Both Sexes. Length: 8.3-9.1 in (21-23 cm); Weight: 1.5-1.7 oz (42-48 g); Wingspan: 9.8-12.2 in (25-31 cm).
[20]
[PDF] Bill Shape as a Generic Character in the Cardinals
The major differences are (a) the morphology of the bili and the jaw muscles; (b) the color of the bill; (c) the male plumage; and (d) the general ecology and ...
[21]
Cardinalis cardinalis (northern cardinal) | INFORMATION
Northern cardinals are medium-sized songbirds. Males are bright red except for a black mask on their face. Females are light brown or light greenish-brown.
[22]
Northern Cardinal - Cardinalis cardinalis - Birds of the World
Feb 12, 2021 · Red plumage color results from ingestion, metabolic conversion, and deposition of carotenoid pigments obtained from the diet during molt.Missing: etymology | Show results with:etymology
[23]
https://digitalcommons.usf.edu/cgi/viewcontent.cgi?article=20502&context=auk
[24]
[PDF] Notes on the Masked Saltator, Saltator Cinctus, in Peru
Sep 7, 2024 · Certain plumage characteristics of Saltator ... One is sexual dimorphism, which is present in Pitylus but lacking in all species of Saltator.
[25]
[PDF] First Evidence for Eccentric Prealternate Molt in the Indigo Bunting
Variation during winter in the male Indigo Bunting's plumage color appears to be due in part to differences in the timing of the preformative and prebasic molts.<|control11|><|separator|>
[26]
Evening Grosbeak Overview, All About Birds, Cornell Lab of ...
In summer you'll need to be in northern North America or in the mountains of the West, where Evening Grosbeaks breed in coniferous forests. At this time they ...
[27]
Distribution - Bluish-gray Saltator - Birds of the World
Sep 24, 2021 · Blue-gray Saltator is widely distributed across most of central and southern South America, stretching as far south as Uruguay and Argentina ...Missing: Grayish | Show results with:Grayish
[28]
Cardinal Bird Facts - Cardinalis cardinalis, Piranga rubra, and others
While there are 53 individual species in the family Cardinalidae, this songbird belongs to the genus Cardinalis of which there are only three “true” cardinals.
[29]
[PDF] Evolution of the preformative molt in Cardinalidae correlates with ...
Jan 31, 2021 · Klicka, J., K. Burns, and G. M. Spellman (2007). Defining a monophyletic Cardinalini: A molecular perspective. Molecular. Phylogenetics and ...<|separator|>
[30]
Species - Cardinalidae - Cardinals and Allies - Birds of the World
Piranga 9 species · Habia 1 species · Driophlox 4 species · Chlorothraupis 4 species · Periporphyrus 2 species · Caryothraustes 2 species · Cardinalis 3 species ...Missing: Clements | Show results with:Clements
[31]
Puerto Rican Spindalis - Spindalis portoricensis - Birds of the World
Formerly classified as a subspecies of the Stripe-headed Tanager (Spindalis zena), the Puerto Rican Spindalis is now recognized as a unique allospecies ...
[32]
Distribution - Northern Cardinal - Cardinalis ... - Birds of the World
Feb 12, 2021 · Cardinalidae. Genus. Cardinalis · Toggle Printable View (single page) ... See also Distribution: Introduced Range and Systematics: Subspecies).
[33]
Varied Bunting Life History, All About Birds, Cornell Lab of Ornithology
Varied Buntings nest in arid thorn forests in canyons, arroyos, stream corridors, and desert washes, and in the hillsides above such areas.
[34]
The impact of habitat and migration on plumage colour in Cardinalidae
Oct 20, 2023 · We used a species-level phylogeny for Cardinalidae based on six genes (Barker et al., 2015) to perform all downstream analyses.
[35]
Scarlet Tanager Overview, All About Birds, Cornell Lab of Ornithology
Male Scarlet Tanagers are among the most blindingly gorgeous birds in an eastern forest in summer, with blood-red bodies set off by jet-black wings and tail.Photo Gallery · Sounds · Range Map · IdentificationMissing: taxonomy | Show results with:taxonomy
[36]
Summer Tanager Overview, All About Birds, Cornell Lab of ...
Fairly common during the summer, these birds migrate as far as the middle of South America each winter. All year long they specialize in catching bees and wasps ...Sounds · Identification · Range Map · Life HistoryMissing: route | Show results with:route
[37]
Movements and Migration - Northern Cardinal - Birds of the World
Feb 12, 2021 · Among encounter records of birds that moved a distance > 25 km, females moved an average of 129.3 km (n = 73), males moved an average of 150.4 ...
[38]
Movements and Migration - Pyrrhuloxia - Cardinalis sinuatus - Birds ...
Estimates of relative abundance for every week of the year animated to show movement patterns. Relative abundance is the estimated average count of ...
[39]
Rose-breasted Grosbeak Range Map - All About Birds
Most of them fly across the Gulf of Mexico in a single night, although some migrate over land around the Gulf. Grosbeaks that winter in Panama and northern ...
[40]
Painted Bunting Range Map - All About Birds
Migration. Short- to medium-distance migrant. Western populations migrate to staging areas in Arizona and northwestern Mexico, where they molt before ...
[41]
Blue Grosbeak Range Map, All About Birds, Cornell Lab of Ornithology
Long-distance migrant. Most individuals that nest in the eastern United States probably fly across the Caribbean to reach their wintering grounds.<|control11|><|separator|>
[42]
Scarlet Tanager Range Map - All About Birds
They usually migrate at night. Individuals that spend the winter farther south migrate to breeding grounds later, and in more synchronized bursts, than ...
[43]
Movements and Migration - Rose-breasted Grosbeak
Migrates from breeding grounds south or southeast through much of e. Canada and e. U.S., primarily from eastern edge of Great Plains to Atlantic coast, but with ...
[44]
Movements and Migration - Hepatic Tanager - Piranga flava - Birds ...
Resident subspecies show some vertical or altitudinal migration and vagrancy. Migration evolved once on the branch leading to Hepatic, Western, Summer ...
[45]
Diet and Foraging - Northern Cardinal - Birds of the World
Feb 12, 2021 · Average annual consumption 29% animal, 71% vegetable matter; feeds nestlings almost exclusively animal matter.
[46]
Diet and Foraging - Rose-breasted Grosbeak - Birds of the World
At feeding stations, preferred food items are sunflower, safflower (Carthamus tinctorius), raw peanuts (Arachis hypogaea; VEW). In captivity, almost any seeds ...
[47]
Diet and Foraging - Scarlet Tanager - Birds of the World
Mar 4, 2020 · Scarlet Tanagers are primarily insectivorous, eating various insects, larvae, and spiders. They also eat fruits, earthworms, and spiders.
[48]
Diet and Foraging - Summer Tanager - Piranga rubra
Mar 4, 2020 · Known for eating bees and wasps, Summer Tanagers are versatile in their diet, also eating a variety of berries. This male is feeding in a black ...
[49]
[PDF] Florida Cardinals Feeding on Nectar - Digital Commons @ USF
Sep 4, 2024 · Accepted. 29 May 73. Florida Cardinals feeding on nectar.--For several weeks this spring a friend and I watched a pair of Cardinals (Cardinalis ...
[50]
Diet and Foraging - Pyrrhuloxia - Cardinalis sinuatus
Seeds, large insects, and fruits. Eats wide variety of items, and appears to be opportunistic feeder, heavily exploiting an abundant food supply.
[51]
Dickcissel | Audubon Field Guide
Feeding Behavior. Forages mostly on the ground and in low vegetation. Except when nesting, usually forages in flocks. Diet. Mostly insects and seeds. Insects ...At a Glance · Range & Identification · BehaviorMissing: foraging | Show results with:foraging
[52]
Extra-pair copulations in a predominantly monogamous bird
The mating behaviour of indigo buntings, Passerina cyanea was studied. A total of 535 attempted matings in 3 years were observed.
[53]
Northern Cardinal | Audubon Field Guide
Length: 8-9 in (21-23 cm); wingspan: 9-12 in (25-31 cm); weight: 1.5-1.7 oz (42-48 g). The male Northern Cardinal is unmistakable, ...Missing: Cardinalidae | Show results with:Cardinalidae<|separator|>
[54]
Northern Cardinal Life History - All About Birds
Species in This Family. Cardinals and Allies(Order: Passeriformes, Family: Cardinalidae). Hepatic Tanager · Summer Tanager · Scarlet Tanager · Western Tanager.
[55]
Breeding - Northern Cardinal - Birds of the World
Feb 12, 2021 · Incubation lasts 11–13 d, nestling period 7–13 d, in first and later broods (see Eggs and Parental Care). First nests in a population seem ...Missing: sources | Show results with:sources
[56]
Pheucticus ludovicianus (rose-breasted grosbeak) | INFORMATION
Rose-breasted grosbeaks begin building nests in May and lay from 1 to 5 (usually 4) pale, bluish-green eggs speckled with darker colors. Nests are constructed ...
[57]
Breeding - Rose-breasted Grosbeak - Pheucticus ludovicianus
Nests placed in various trees, shrubs, and vines, more often in vertical forks or crotches than on horizontal limbs. Saplings more often selected than tall ...
[58]
Rose-breasted Grosbeak | Audubon Field Guide
Both parents feed the nestlings. Young leave nest about 9-12 days after hatching. Male may care for fledglings while female begins a new nest. 1-2 broods ...
[59]
Polygyny in Indigo Buntings: A Hypothesis Tested - Science
A test of a model for the evolution of avian polygyny revealed that 10 percent of male indigo buntings had two mates simultaneously and some had none.
[60]
Northern Cardinal Sounds, All About Birds, Cornell Lab of Ornithology
Calls. Scientists have described at least 16 different calls for the Northern Cardinal, but the one you'll hear most commonly is a loud, metallic chip.
[61]
Sounds and Vocal Behavior - Northern Cardinal - Birds of the World
Feb 12, 2021 · Songs are typically pure-toned, whistle-like series of repeated syllables whose fundamental frequencies range from approximately 1–8 kHz.
[62]
Sounds and Vocal Behavior - Black-headed Grosbeak
Mar 18, 2022 · Repertoire and Delivery of Song. Repertoire of syllables of individual males = 14–37 (mean = 24.5, n = 22); repertoire of individual females ...Missing: size | Show results with:size
[63]
Organization of song of rose-breasted grosbeaks - ScienceDirect.com
Rose-breasted grosbeaks (Pheucticus ludovicianus) in Quebec had repertoires of fifteen to twenty-three syllables per bird, some of which were similar among ...
[64]
Sounds and Vocal Behavior - Rose-breasted Grosbeak
Repertoire And Delivery Of Songs. Estimated repertoire for a single bird includes 15–23 syllables (Borror 1961b Borror, D. J. (1961). Songs of finches ...Sounds And Vocal Behavior · Vocalizations · Vocal Array
[65]
Behavior - Northern Cardinal - Birds of the World
Feb 12, 2021 · Northern Cardinals hop, have undulating flight, preen, bathe, roost, sunbathe, engage in agonistic interactions, and are territorial. They also ...
[66]
Voices and Vocabularies - Cardinals' Duet - BirdNote
Apr 20, 2020 · Among most North American songbirds, it's males alone who sing. But during the nesting season, we also hear female cardinals.
[67]
Cardinal Courtship - Birds Outside My Window
Feb 6, 2018 · In February they begin to court. The males become aggressive toward other males and solicitous to their ladies. And they begin to sing.Missing: Cardinalidae | Show results with:Cardinalidae
[68]
Flock of Cardinals: Is It Normal to See Multiple Cardinals?
Mar 19, 2025 · While usually seeing one or two, cardinals flock together, especially after breeding season. Flocks of 6-20 are typical, though larger groups ...Missing: Cardinalidae | Show results with:Cardinalidae
[69]
[PDF] northern cardinal - Texas Parks and Wildlife
Nectar feeding animals are attracted to sugar water feeders and native plants that produce nectar rich flowers. Insect eating animals examine plants and ...<|control11|><|separator|>
[70]
The displays and call notes of cardinals - ResearchGate
Aug 6, 2025 · Among the displays the most interesting are those used in courtship, namely, song-flight, song-dance, lopsided, and courtship-feeding displays.
[71]
Do Northern Cardinals Mimic Other Birds? - Bird Watching Diaries
Jun 7, 2025 · ... vocal patterns for communication. Unlike some other species, Northern Cardinals do not mimic the sounds of other birds, which sets them ...Missing: Cardinalidae | Show results with:Cardinalidae
[72]
(PDF) Geographic Patterns of Song Similarity in the Dickcissel ...
Aug 7, 2025 · Three Dickcissel songs from three distinct vocal neighborhoods. In these examples, dick (DK) phrases differed in number among males, one song ...
[73]
Sounds and Vocal Behavior - Dickcissel - Spiza americana
Feb 26, 2022 · The song of the Dickcissel is one of the most frequently heard summer sounds in prairie grasslands that make up the core breeding range.
[74]
IUCN Red List of Threatened Species
The 2025-2 IUCN Red List also reveals threatened European butterfly species increased by 76% over the last decade.About · 3.1 · Searching The IUCN Red List · Citing The IUCN Red List
[75]
Northern Cardinal Cardinalis Cardinalis Species Factsheet | BirdLife ...
Habitat classification ; Forest, Temperate, Resident, Suitable ; Shrubland, Subtropical/Tropical Moist, Resident, Suitable.
[76]
Red-and-black Grosbeak Caryothraustes Erythromelas Species ...
Deforestation is the primary threat affecting this species (Soares-Filho et al. 2006, Bird et al. 2011). It requires near pristine forest (del Hoyo et al. 1992) ...
[77]
Threats to Birds: Habitat Impacts | U.S. Fish & Wildlife Service
Loss, degradation and fragmentation of important migratory bird habitat have been identified as potentially the largest individual threat to migratory birds.
[78]
Vermilion Cardinal Cardinalis Phoeniceus Species Factsheet
This species has a large range, and hence does not approach the thresholds for Vulnerable under the range size criterion (extent of occurrence under 20,000 km² ...
[79]
Yellow Cardinal Gubernatrix Cristata Species Factsheet
Yellow Cardinal · 1,820,000 km² · Not a Migrant · No · 30 · IUCN Red List assessment · 1,000 - 2,000 · Decreasing · Sites ...
[80]
Carrizal Seedeater - Amaurospiza carrizalensis - Birds of the World
CRITICALLY ENDANGERED. Recently discovered species, confined to tiny area of SE Venezuela. Very poorly known. Nowhere common, and global range appears to be ...
[81]
Conservation and Management - Northern Cardinal
Feb 12, 2021 · Movements and Migration · Dispersal and Site Fidelity · Migration Overview · Diet and Foraging · Feeding · Diet · Food Selection and Storage ...<|control11|><|separator|>