SPECIES

Northern Cardinal Cardinalis cardinalis

Sylvia L. Halkin, Daniel P. Shustack, M. Susan DeVries, Jodie M. Jawor, and Susan U. Linville
Version: 2.0 — Published February 12, 2021

Behavior

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Locomotion

Walking, Hopping, Climbing, etc.

Hops on ground and in vegetation.

Flight

Undulating flight in open areas; short flights between adjacent branches when foraging. During courtship, male flight display includes short, rapid wingbeats as he slowly descends (Song-Flight Display; see Sexual Behavior).

Self-Maintenance

Preening, Head-Scratching, Stretching, Bathing, Anting, etc.

Bathes throughout year , including winter. Commonly preens while perched in trees and known to engage in "anting" behavior (62). Cleans and likely sharpens bill (as in other species; e.g., 263) by wiping it on hard surfaces (SUL).

Sleeping, Roosting, Sunbathing

Roosts late in evening (time not quantified); pair may roost in close proximity ( ); gives loud call notes (probably chips) before approaching roost site; male roosts away from nest tree or shrub (distance not quantified; 177). Sunbathes frequently in early morning in cooler months, when individuals perch on exposed limbs with feathers fluffed (SUL, JMJ).

Daily Time Budget

In the non-breeding season, time budgets of males and females in a captive flock with unlimited food were the same: 74% perching, 13% feeding, and 5% flying; changes in flock size did not result in a distinct continuum of changes in time allocation (264). No data available on time budgets in wild birds. See Breeding: Parental Care.

Agonistic Behavior

Communicative and Physical Interactions

Agonistic interactions seen in multiple contexts: maintaining distance between individuals (usually during feeding in winter flocks), defense of territory (breeding and non-breeding seasons), and defense of nest during intraspecific brood parasitism attempts (females). Interactions include displays, chases, vocal behavior (calls, song), displacements, and physical attacks (JMJ, MSD).

Except where noted, the following is from Lemon (4, which includes additional descriptions and illustrations). Uses several displays during aggressive encounters. Demonstrates aggression with lowered crest and pee-too or chuck call notes. Head-Forward Display (crest and body low, mouth gaping, wings vibrating) indicates higher level of aggression and may be followed by chips, pee-toos, and chucks before bird lunges. In extreme circumstances, before lunging, bird crouches with crest down, feathers smooth, vision fixed on opponent. Responses to aggressive displays may involve departure (moving away by hopping or flying) or submission (crest raised, upper breast fluffed, bill pointing upward, avoiding direct eye contact). Individuals engage in displacement-feeding and bill-wiping during aggressive encounters. When individuals fight, they grapple with feet and strike with beaks, either in air or on ground; these encounters are rare and usually last just a few seconds (67). Continual chases between males, females, or pairs lasting 10–30 minutes or more, with little rest, have also been observed (67, SUL, JMJ).

Song commonly a component of aggressive interactions, with song noted frequently as part of naturally occurring and simulated territory intrusions (99, 265). Females in new pairs sing more frequently than females in established pairs, females sing at higher rates during the early breeding season (17), and singing by females during nest defense has been noted (15, 19). See additional information in Sounds and Vocal Behavior: Vocalizations: Social Context and Presumed Functions of Vocalizations.

Territorial Behavior

Defended territory sizes during breeding season range from 0.21 to 2.60 ha (mean 0.93–1.50) across data from southern Indiana (n = 47 territories; 67), southwestern Ohio (n = 122 territories; 266), eastern Tennessee (n = 20 territories; 211), and southern Arizona (n = 4 territories; 135). Territory is established by song, postures, pursuit, and combat. Males and females engage in intrasexual (male–male or female–female) and intersexual aggressive interactions during territory defense (15, 33, 265). Intruding individuals are frequently alone, although pairs have been observed intruding on territories together (MSD). Sexes may behave differently during intrusions; intruding males often sing during territory intrusions and move conspicuously around territories, while intruding females typically remain silent and their movements are less obvious (JMJ, MSD).

From Laskey (100), Kinser (67), and Merritt (211), and additional authors/observers as cited: Active defense of territory boundaries in Tennessee and southern Indiana ceases in late summer or early fall and resumes in late winter to early spring (January–March). Color-banded males and pairs have been observed on breeding season territories during the non-breeding season in Kentucky, New York, Ohio, Indiana, Wisconsin, and Mississippi populations (267, 212, JMJ, SLH). Gentry (212) reported in a Mississippi population that single males and pairs occupy and defend breeding season territories in the non-breeding season (October–February). Territory boundaries of established individuals are approximately the same from year to year (SUL, JMJ, SLH), although one male was observed to expand his territory to include that of a neighboring pair when he became the bigamous mate of the neighboring female after her mate's summer disappearance (SLH); it is not known whether that expansion lasted beyond one breeding season. Territory boundary locations in a given area may be similar from year to year even when different individuals occupy some of the territories (SLH, JMJ, MSD). Both males and females leave their territory, usually quietly and near the ground, to feed and to bathe during the breeding and non-breeding seasons. During the breeding season, banded females reported up to 75 m from the territory boundary, and banded males up to 100 m; JMJ observed banded adults of both sexes from a Mississippi population up to one km from their breeding territories during the early breeding season (March–April).

Males and females experiencing simulated territory or nest intrusions respond to experimental intrusions as if simulated intruders are real: behavior observed during experimental trials is similar to that observed opportunistically from naturally interacting individuals. DeVries et al. (33) assessed male response to simulated intrusions (song and taxidermy male presented) and found males react to intruders by returning song (song matching and over-singing, in which the resident starts to sing during the playback song), intense chipping, flights over the taxidermy male, and physical attacks involving contact with the taxidermy male. Comparison of aggressive behaviors with circulating testosterone levels found no co-variation between testosterone levels and aggression in males, although testosterone can be increased during aggressive interactions (31, 265). Gentry (212) found similar types of behavioral responses (song, flights over the speaker, etc.) when males responded to simulated male intrusions (song recordings only) during the breeding and non-breeding seasons in a Mississippi population although responses were generally weaker in the non-breeding season. Females may accompany mates during territory defense against intruding males. DeVries et al. (265) found that pairs matched the strength of their behavioral responses during simulated intrusions by males.

Female cardinals have been assessed for their willingness to defend their nest site from intraspecific brood parasites (15, 34, 19). Natural observations of females approaching nests that are not theirs have been made (JMJ; R. Breitwisch, personal communication) and genetic analyses have found broods that include young unrelated to the females that are caring for them (75). However, intraspecific nest parasitism is rare and may vary by population (75, 19; see also Breeding: Eggs). In Ohio and Mississippi populations, females will aggressively defend nests during simulated nest intrusions (taxidermy female cardinal placed at nest) and behaviors range from focal females returning to nests to incubate eggs without interacting with simulated intruders, to focal females singing at and diving at simulated intruders, and physically attacking simulated intruders (15, 34, 19). These behaviors match naturally occurring observations of females defending their nests from intraspecific intruders. In an Ohio population where intraspecific brood parasitism was documented (75), female face mask ornamentation co-varied with level of aggression shown during nest defense (15); females with larger and darker (dark gray or black) face masks showed more aggression during nest defense. In a Mississippi population, where no behavioral evidence of brood parasitism was found (268), female ornamentation did not co-vary with the level of aggression shown during nest defense (19). Last, DeVries et al. (34) did not find that circulating testosterone levels influenced female aggressive behavior shown during nest defense.

Sexual Behavior

Mating System and Operational Sex Ratio

Socially monogamous, but polygyny observed, perhaps because of persistence of an unmated female staying on a mated male's territory (4, MSD) or because of the male assuming territory where a neighboring male has disappeared (269, SLH). Polygyny may (SUL) or may not (4) result in successful rearing of offspring.

Banding data show a slight female bias (1.03 females: 1 male, n = 532,785 banded birds of all ages with sex identified; for adults only with sex identified, the ratio is 1.1 females: 1 male, n = 293,492; U.S. Geological Survey Bird Banding Laboratory data as of November 2019). Sex ratios of birds banded between late summer or fall and spring were 1.37 females: 1 male in central Kentucky (n = 382 birds, 2-year study; 267) and 1.08 females: 1 male (n = 100 birds) one year and 1.03 females: 1 male (n = 152 birds) the next in north-central West Virginia (214).

Courtship, Copulation, and Pair Bond

Pairs may remain together throughout winter either on breeding territory or in flocks, but divorces occurred between seasons (2 of 10 banded pairs divorced) and within seasons (2 of 21 banded pairs divorced) in an Ohio population (28). Except where noted, the following is from Lemon (4, which includes additional descriptions and illustrations of displays of both sexes). Pair bond is initiated with various male courtship displays. During Lopsided Display, male twists and rotates his body to display breast, and holds crest low, wings spread, and feathers flat. Song-Dance Display is similar to Lopsided Display, but more upright, with crest erect. Male sings, swaying back and forth. During Song-Flight Display, male fluffs breast feathers, raises crest, sings, and descends slowly toward female, beating wings in short, rapid strokes. Females also perform displays to males during courtship and early pair formation, including an open wing display where the body is held upright and wings half-opened, displaying underwing coverts (see Breeding: Phenolology: Pair Formation) (270), and swaying displays similar to those described for males, during which song may or may not be sung (243, 271).

Male feeds female during courtship and incubation; female may assume begging posture to obtain food, quivering wings like a juvenile (4). Jawor and Breitwisch (272) did not find that male plumage ornamentation predicted mate provisioning rates during incubation; however, females that were provisioned at greater rates during incubation later fed their nestlings at lower rates, compared to females that had been provisioned at lower rates during incubation. When ready to mate, female gives Precopulatory Display: points beak and tail upward, lowers and quivers wings, fluffs breast feathers, and separates ventral feathers to expose cloacal opening (4). Viewing of copulations is rare in the field, so it is difficult to determine if any copulations are forced. Females have been observed soliciting extra-pair copulations on neighboring territories (SUL).

Ornamentation

Studies assessing the use of ornamentation in both sexes have focused on bright red plumage (upper breast in males, underwings in females), crest length and bill color in both sexes, face mask brightness (e.g., amount of reflected light) and area in males, and face mask score (color, light gray to black, and area combined; used by JMJ in Ohio and Mississippi populations) or face mask brightness (amount of reflected light; used by JMJ in Mississippi population) in females (113, 15, 16, 272, 19). Note, studies have used various color measurement techniques from Munsell color chips to color reflectance spectrometers and terminology for coloration (e.g., redder, brighter, darker, etc.) varies across studies, making comparisons across findings difficult; here we use the terminology that the authors of various studies employed in their work.

Rural males with redder plumage hue have been reported to hold territories with denser vegetation (78, 273), but this relationship breaks down in urban systems that provide many territories with dense vegetation and abundant food (273). Males of redder hue had greater reproductive success in a rural New York population (78), but not in rural or urban Ohio populations (273). Female preference for more colorful mates has not been found in captive mate-choice trials (76) and male dominance status in captive males (male only groups) was not impacted by plumage color (77). Male feeds nestlings more than female (28, 16, 272), and carotenoid plumage brightness can be positively correlated with feeding effort (see Breeding: Parental Care), although not always (16, 272, 15).

Male and female ornamentation has been compared with body condition (scoring based on residuals following regression of mass on tarsus length), feather growth rates, body size, parental care behavior, nest defense (against predators and intraspecific brood parasitism), and mate provisioning. Multiple ornaments on the same individual (plumage color, bill color, crest length, face mask expression) tend not to co-vary in expression, although carotenoid-based ornaments (plumage color and bill color) are often similar in expression (16, 272, 15).

In females in an Ohio population, multiple ornaments co-varied positively with body size and body condition (underwing plumage color, bill color), date of first nest and reproductive success (underwing plumage color), nestling feeding rates (face mask score [size and color combined]), and intrasexual aggression (face mask score) (15). Ornament expressions did not co-vary with feather growth rates. In a Mississippi population, Winters and Jawor (19) did not find the face mask ornament (either feather brightness or a combined color/area score) indicative of behavioral aggression. This variation could be due to population variance in intraspecific brood parasitism behavior. Maney et al. (274) found that female carotenoid plumage saturation was positively correlated with higher white blood cell counts (typically a signal of infection).

In an Ohio population, male ornamentation was found to positively co-vary with body size and condition (breast plumage color, bill color respectively) and to negatively co-vary with reproductive success (face mask area) (16). Male ornamentation in this study did not co-vary with nest defense behavior or with feather growth rates. The association between male ornamentation and parental care behavior is complex. In an Ohio population, Linville et al. (29) found male breast plumage coloration to be positively associated with feeding rates, yet, Jawor and Breitwisch (16) found a negative association between nestling feeding and breast coloration, whereas Jawor and Brietwisch (272) found no association between ornamentation and nestling feeding rates. This variation has not been fully examined; its sources are unclear and could be linked to variation in food availability or environmental conditions; similar methodologies were used to determine nestling feeding rate and ornament coloration in all three of these studies. Jones et al. (53) found that males in better body condition had higher plumage brightness. Maney et al. (274) found that males with more saturated carotenoid-based plumage had higher heterophil to lymphocyte ratios (elevated heterophil to lymphocyte ratios are considered a sign of elevated physiological stress).

Carotenoid-based coloration in both sexes has been shown to be impacted by fruit availability during the molt (229). Captive male cardinals fed only white millet, red millet, and sunflower seeds during the molt, with no sources of carotenoids other than those found in the seeds, grew red plumage that was less saturated than that of wild males (74). Males, but not females, in urban habitat had decreased plumage brightness compared to rural individuals (53). McGraw et al. (72) assessed feather carotenoid content from a study skin of a male cardinal with yellow instead of red plumage, and suggested that yellow individuals may lack enzymes responsible for carotenoid conversion in feather follicles. The exact impacts and interactions of food, physiology, habitat and landscape context on coloration need further study.

Extra-Pair Mating Behavior/Paternity

DNA fingerprinting in two populations of cardinals (Ohio and Kentucky) has revealed extra-pair fertilizations (EPFs), with rates ranging from 9 to 35% of nestlings (45, 75). The male guards his mate during her fertile periods, maintaining contact with the mate on average 79% of observation time during initial nesting attempts in a Kentucky population (21 focal pairs, 20 April–31 August, n = 256 observation periods of 65 minutes each; 45); EPFs may be associated with less guarding time. Females that were mated with brighter males obtained more EPFs in an Ohio population, but relationships between plumage brightness and mate-guarding is unknown (75). Radio-tracked females on food-supplemented territories were found to make more movements off of their territories during fertile periods than did unsupplemented females (275).

Brood Parasitism of Conspecifics

Rare, and may vary between populations; see Breeding: Eggs: Egg Laying, Agonistic Behavior, and Mating System and Sex Ratio (above).

Social and Interspecific Behavior

Degree of Sociality

Highly territorial during breeding season and in some populations outside of the breeding season (212, JMJ). Winter flocks not strongly cohesive (see Behavior: Spacing and Territorial Behavior). No cooperative foraging. Pairs will mob predators near the nest, sometimes joining other mobbing passerines (44, SLH, JMJ), but no mobbing by larger groups of cardinals reported. Individuals using feeders in winter tended to base use on feeder occupancy; females tended to avoid feeders with other cardinals, males displaced females and avoided feeders with other males (276). Males will displace their mates at feeders during the non-breeding season, but become more tolerant of mates at feeders as the breeding season approaches (100). At a Kentucky feeding station, in observations of marked cardinals between 15 December and 5 April, males dominated females and adults dominated juveniles (267).

Flocks observed during or after fall molt (end of September in southern Indiana and Iowa; 277, 67). May start as family groups consisting of adults with their last brood, and enlarge when birds from adjacent breeding territories join together (67), and/or when joined by juveniles lacking mates and territories (e.g., Alabama flock reported in Breeding: Immature Stage). In some areas, pairs that remain on breeding territories year-round only join flocks as they pass through their territories (central Tennessee [100]; southwestern Kentucky [243]; central Kentucky [267]; southern Illinois and southern New York [278]; southwestern Ohio [99]; southeastern Mississippi [MSD, JMJ]).

Winter flocks reported in subspecies cardinalis (62 and other references below) and superbus (135). Flock size may vary geographically within cardinalis; flocks of > 100 reported only from West Virginia (279), southern Indiana (67) and probably South Carolina (T. Nuttall in 62). In Ohio, winter flocks of 4–60 (M. B. Trautman in 214); in Tennessee, 6–25 (280, 100). Mean winter flock size 13.8 in southern Indiana (n = 430 flocks; 67), but 4.6 in central Kentucky (n = 922 observations of flocks that included at least 1 of 8 radio-tagged cardinals; 267); these two studies used different methods and possibly different criteria for flock membership, but both found variation in flock sizes over course of winter and day, and with temperature; Kinser (67) also reported different flock sizes in different habitats.

Flock movement described as "rolling" or "tank-tread like," with front birds settling on the ground to feed until most of the flock has passed over them, and then flying up to settle at the front again (281, 67). Flock membership is fluid; not only do birds spending winter on breeding territories temporarily join a flock as it passes over them, but within a six hour period, marked birds were observed to move 500–900 m, changing the flocks with which they were associating (67). Flocks may move "through several square miles," but in this case individual birds rarely remained with the flock for distances > 1.6 km, and small groups joined the flock as it "rolled" over them, while similar numbers of different individuals left flocks to remain behind feeding (281). Members of winter flocks roost in aggregations in thickets or conifer groves (214, 135, 211). Flocks gradually disband January–March as birds leave to establish territories (67, 211).

Flocks are usually made up of approximately equal numbers of males and females and contain a mix of hatch-year young and older adults (243, 279, 100, 67). Banded hatch-year young have been seen in winter flocks 0.8–4.0 km away from where they hatched (100). Cardinal flocks may be associated with other species (see Nonpredatory Interspecific Interactions, below).

Play

No reports.

Nonpredatory Interspecific Interactions

Competes with Gray Catbird (Dumetella carolinensis) and American Robin (Turdus migratorius) for nest sites and feeding grounds, giving way to robins for nesting position (282). Drives House Sparrow (Passer domesticus), Field Sparrow (Spizella pusilla), and Harris's Sparrow (Zonotrichia querula) away from feeding stations (244) and supplants White-throated Sparrow (Zonotrichia albicollis; 283). Parasitized by Brown-headed and Bronzed Cowbird within their ranges (see Breeding: Brood Parasitism). In southwestern United States, where sympatric with Pyrrhuloxia, breeding cycles are identical, territories overlap, and songs can be similar; interspecific aggression was not noted (135), but male Pyrrhuloxia have been observed responding aggressively to recorded male Northern Cardinal song (JMJ). Govoni et al. (284) reported nest sharing between a female Northern Cardinal and a female American Robin.

Male Northern Cardinal has occasionally been recorded feeding nestlings or fledglings of other species (e.g., 285); one male fed a group of 7 large goldfish in an outdoor pool (286). Other bird species have assisted Northern Cardinal in incubating eggs and feeding young, including Song Sparrow (Melospiza melodia) in a nest shared with a Northern Cardinal pair (287).

Cardinal flocks may be closely or loosely associated with other species, including Dark-eyed Junco (Junco hyemalis), White-throated Sparrow, Tufted Titmouse (Baeolophus bicolor), Song Sparrow, American Tree Sparrow (Spizelloides arborea), American Goldfinch (Spinus tristis), Eastern Towhee (Pipilo erythrophthalmus), and Pyrrhuloxia (Cardinalis sinuatus) (281, 135, 67).

Predation

Kinds of Predators; Manner of Predation

Adults are known to be taken by domestic cats (Felis domesticus; 288) and dogs (Canis familiaris; 214), Cooper's Hawk (Accipiter cooperii; 177), Loggerhead Shrike (Lanius ludovicianus), Northern Shrike (L. borealis) (289, 290), and eastern gray squirrel (Sciurus carolinensis; 291). Northern Cardinal remains identified in owl pellets, including those of Long-eared Owl (Asio otus; K. Russell, personal communication) and Eastern Screech-Owl (Megascops asio; D. McGovern, personal communication).

In most cases of predation on eggs or nestlings, the nest is undamaged, suggesting predation by snakes, birds, or small mammals (67, 292, 43). Known predators of eggs include milk snake (Lampropeltis triangulum), black racer (Coluber constrictor), Blue Jay (Cyanocitta cristata), fox squirrel (Sciurus niger), red squirrel (Tamiasciurus hudsonicus), domestic cat, and eastern chipmunk (Tamias striatus); eggs punctured by House Wren (Troglodytes aedon; 293, 62, 67); and Brown-headed Cowbird removes eggs and eats or discards them (see Breeding: Brood Parasitism). Nestlings or newly fledged birds taken by eastern chipmunk, black racer, and gray ratsnake (Pantherophis spiloides) (67, 294), as well as eastern ratsnake (P. alleghaniensis) . Nest predators confirmed by video monitoring of 89 Northern Cardinal nests in urban and rural forests in central Ohio included Brown-headed Cowbird (n = 18), raccoon (Procyon lotor; n = 9), Common Grackle (Quiscalus quiscula; n = 6), Blue Jay (n = 5), Gray Catbird (n = 5), Virginia opossum (Didelphis virginiana; n = 5), Cooper’s Hawk (n = 4), eastern gray squirrel (n = 4), Barred Owl (Strix varia; n = 4), American Crow (Corvus brachyrhynchos; n = 4), domestic cat (n = 4), red squirrel (n = 3), gray ratsnake (n = 2), and one instance each of Northern Cardinal (adult), eastern garter snake (Thamnophis sirtalis), eastern chipmunk, and House Wren; for two additional nest predation events by hawks, and two by songbirds, the predators could not be identified to species (252; L. Kearns and A. Rodewald, unpublished data); the authors suspected that the cardinal was removing a cowbird egg, but this could not be confirmed.

Response to Predators

In controlled experiments with predator models (taxidermic mounts of eastern chipmunk and Blue Jay, rubber model of ratsnake), females responded more strongly at the egg stage than did males, giving chip calls and flying toward the predator model (44, 15). Similar responses were given by both parents at the nestling stage, although females gave more alarm calls. No evidence for predator-specific defense. Ornamentation has not been found to co-vary with nest defense behaviors when facing predators (15, 16).

Northern Cardinals have been assessed for their reaction to humans (e.g., flight initiation distances, FID) in urban and rural habitats (295); individuals in urban habitats showed greater tolerance and shorter FIDs than did rural individuals. Incubating females in forested parks with recreational trails tended to flush more often when their nests were lower and closer to trails (296, 297). Northern Cardinals will respond to interspecific predator alarm calls with anti-predator behavior (298); however, Grade and Sieving (299) found reduced anti-predator reactions to broadcasts of interspecific predator alarm calls in urban cardinals compared to individuals in rural/natural areas. Stanback and Powell (300) found that Northern Cardinals reduced their use of feeders in open habitat in the evidence of predators (Cooper’s Hawk calls) and Duncan and Bednekoff (258) found that male Northern Cardinals altered territorial singing behavior in the presence of Cooper’s Hawk nest sites (see Sounds and Vocal Behavior: Vocalizations: Vocal Array: Places of Vocalizing).

Recommended Citation

Halkin, S. L., D. P. Shustack, M. S. DeVries, J. M. Jawor, and S. U. Linville (2021). Northern Cardinal (Cardinalis cardinalis), version 2.0. In Birds of the World (P. G. Rodewald and B. K. Keeney, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.norcar.02