Behavior

Walking, Running, Hopping, Climbing, etc.

Movements on foot include strong hops, jumping, clambering, sidling, and creeping (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ).

Flight

Flies in short, direct bursts, with loose wing beats; longer flights are more undulating (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). Tends to avoid longer flights over open spaces (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close ).

Take-off speed (in alarmed/escape flights) is 3.84 m/s in dominant adult males (n = 11) and 3.72 m/s in subordinate juvenile females (n = 11), and declines over the course of a day in the latter (93 Krams, I. (2002). Mass-dependent take-off ability in wintering great tits (Parus major): comparison of top-ranked adult males and subordinate juvenile females. Behavioral Ecology and Sociobiology 51:345–349. Close ).

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

Information needed.

Sleeping, Roosting

Summer roosts are generally in foliage, except for nesting females which roost in the nest cavity until nestlings are ca. 14 days old, or sometimes until fledging (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 94 Berndt, R., and W. Winkel (1972). Über das Nächtigen weiblicher Meisen (Parus) während der Jungenaufzucht [On nocturnal brooding by female tits (Parus) while rearing young]. Journal für Ornithologie 113:357–365. Close , 19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). Winter roosts include tree cavities, nest boxes, holes in man-made structures (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ), dense conifers (95 Lehtonen, L. (1947). Zur Winterbiologie der Kohlmeise, Parus m. major L. Ornis Fennica 24:32–47. Close ), holes in snow (96 Gogel, R. (1962). Gîte sous la neige d'une Mésange charbonnière. Nos Oiseaux 26:216. Close , 97 Zonov, G. B. (1967). On the winter roosting of Paridae in Cisbaikal. Ornithologija 8:351–4. Close ). Roost holes are usually occupied singly, sometimes in pairs (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ).

Individuals in northern populations with longer days go to roost proportionately earlier in the day with respect to twilight (i.e. at higher light levels) than birds from more southern latitudes (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 91 Mace, R. (1989). A comparison of Great Tits' (Parus major) use of time in different daylengths at three European sites. Journal of Animal Ecology 58(1):143–151. Close )

Daily Time Budget

Males during the breeding season are active ~83–93% of daylight hours, with lower percentages of daylight active at higher latitudes (91 Mace, R. (1989). A comparison of Great Tits' (Parus major) use of time in different daylengths at three European sites. Journal of Animal Ecology 58(1):143–151. Close ). Males spend proportionately more time than expected displaying and less time foraging at the start and ends of days, while time spent at rest is distributed evenly across a day (91 Mace, R. (1989). A comparison of Great Tits' (Parus major) use of time in different daylengths at three European sites. Journal of Animal Ecology 58(1):143–151. Close ).

Descriptions based on Hinde (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close ) and Cramp and Perrins (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ), except where noted.

Physical and Communicative Interactions

Integrated flock movements are initiated by a bird stopping feeding, giving tink calls, flying higher and moving gradually away, until other individuals respond and follow.

Individuals within foraging flocks fight over food and establish dominance hierarchies. Such interactions include scolding and other calls, which might attract more birds to a skirmish. The visual aspects of displays emphasize the white cheeks and black bib and ventral line. The width of the ventral line is a badge of social status, with more aggressive and dominant individuals having wider bands (98 Järvi, T., and M. Bakken (1984). The function of the variation in the breast stripe of the Great Tit (Parus major). Animal Behaviour 32:590–596. Close ).

A feeding bird that is approached too closely by another will respond with Wings-Raised Posture or Head-Forward Posture. In the Wings-Raised Posture, wings are held half-open and raised level with or over the back, the tail is spread and sometimes raised or lowered, and the body feathers are sleeked. Birds sometimes continue feeding in this posture. Higher intensity threats are met by stopping feeding and adopting the Head-Forward Posture, where the beak is pointed at the threat, the head is held down and forward, swaying sometimes with the beak open. At highest intensities, the Head-Forward Posture is accompanied by Wings-Raised Posture, and the bird moves towards the threat, which usually retreats. Displacement behaviors during interactions include pecking at twigs, bill-wiping, and leaf-turning.

Supplanting Attacks involve an aggressor diving down from above, sometimes with wings raised, at another individual, which usually retreats and abandons the food it had. Some fight back against Supplanting Attacks by jumping to meet aggressors in the air, resulting in combat. Fighting birds hold their bodies upright and facing each other, attacking with beak and claws, and falling to the ground; such fights rarely last for more than a few seconds. Supplanting Attacks can be directed by males at females during the establishment of pairs and territories. In these, males hold the body vertically, and make a series of short upward sweeps during the attack.

Another aggressive display is the Head-up Posture. A bird perches with the body held vertical, body feathers sleeked, tail spread, the neck extended and bill pointing up, and the head swaying side to side, which emphasizes the white cheeks and black bib and ventral line. Two birds adopt this posture facing each other for up to 10 seconds.

Territorial Behavior

In Western Europe, it is territorial only during the breeding season. Territories are formed within several weeks of the breakup of winter foraging flocks. Older birds reestablish the same territories. Territory boundaries are strongest during nest building, and decline during incubation. In Israel, the Great Tit is territorial year-round (99 Yavin, S. (1987). Nest site selection of the Great Tit Parus major terrae-sanctae. Ph.D. dissertation, Tel-Aviv University, Tel-Aviv, Israel. Close ).

Mating System and Operational Sex Ratio

Monogamous (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ).

Courtship, Copulation, and Pair Bond

Behaviors summarized from Hinde (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close ) and Cramp and Perrins (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ) except where noted.

Early in the season, males follow females around the territory for almost the entire day, rising to sing before the female emerges from the nest hole, and going to roost after the female enters the hole in the evening. Females typically perch lower than accompanying males, and males call to relocate female when no longer visible.

Pair bonding behaviors include flights by both sexes with shallow wingbeats and brief glides (often associated with nest inspection displays, see Breeding: Nest Site), the male perching or hopping with the body held horizontal, the wings raised, and the tail raised and spread, and males giving a warbling call along with wing shivering and a slight pumping of the tail. The nest inspection display might have a role in pair bonding, especially when performed outside the breeding season. Courtship feeding is solicited by the female crouching with shivering wings, a slightly raised tail, and giving a zeedling call. The male responds by feeding either without display or with slight wing shivering. Female begging is most frequent during egg laying and early incubation.

Copulation typically takes place in thick cover, in the early morning, most often right after the female emerges from the nest hole, and rarely at other times of day. The male initiates by crouching with the head held back, the wings shivering, the tail slightly raised and spread, and calling. Females initiate or signal receptivity by crouching with shivering wings and calling. Copulation is sometimes be followed by bill wiping, wing shivering, or courtship feeding.

Pairs are maintained year-round in Israel (99 Yavin, S. (1987). Nest site selection of the Great Tit Parus major terrae-sanctae. Ph.D. dissertation, Tel-Aviv University, Tel-Aviv, Israel. Close ). In western Europe, pairs break up after breeding, and re-form, if both survive, by January and the start of territoriality (58–77% of cases; 19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). In a five-year study in Germany, divorce occurred in 5 out of 29 pairs [= 17.2%] where both members of the pair were recaptured in multiple years (100 Lubjuhn, T., Strohbach, S., Brun, J., Gerken, T. and Epplen, J.T. (1999). Extra-pair paternity in Great Tits (Parus major) – a long term study. Behaviour. 136(9): 1157-1172. Close ).

Extra-pair Mating Behavior/Paternity

The frequency and degree of extra-pair paternity is variable by location and by year (101 Strohbach, S., E. Curio, A. Bathen, J. Epplen, and T. Lubjuhn (1998). Extrapair paternity in the great tit (Parus major): a test of the “good genes” hypothesis. Behavioral Ecology 9(4):388–396. Close , 100 Lubjuhn, T., Strohbach, S., Brun, J., Gerken, T. and Epplen, J.T. (1999). Extra-pair paternity in Great Tits (Parus major) – a long term study. Behaviour. 136(9): 1157-1172. Close , see review in 102 Lubjuhn, T., T. Gerken, J. Brün, and T. Schmoll (2007). Yearling male great tits, Parus major, suffer more strongly from cuckoldry than older males. Zoology 110(5):387–397. Close ), but is often high in the Great Tit. Examples include: in England, 31% of broods (n = 160 broods) contained at least one extra-pair young (103 Roth, A. M., J. A. Firth, S. C. Patrick, E. F. Cole, and B. C. Sheldon (2019). Partner’s age, not social environment, predicts extrapair paternity in wild great tits (Parus major). Behavioral Ecology 30(6):1782–1793. Close ), and an estimated 14% of nestlings were sired by an extra-pair male (n = 831 nestlings; 104 Blakey, J. K. (1994). Genetic evidence for extra-pair fertilizations in a monogamous passerine, the Great Tit Parus major. Ibis 136:457–462. Close ). In the Netherlands, 8.5% of broods (n = 82) contained extra-pair young, and 3.5% of nestlings (n = 516) were extra-pair (105 Verboven, N., and A. C. Mateman (1997). Low frequency of extra-pair fertilizations in the Great Tit Parus major revealed by DNA fingerprinting. Journal of Avian Biology 28:231–239. Close ), or 25.3% of broods (n = 99) and 6.5% of nestlings (n = 667) (106 van Oers, K., P. J. Drent, N. J. Dingemanse, and B. Kempenaers (2008). Personality is associated with extrapair paternity in great tits, Parus major. Animal Behaviour 76(3):555–563. Close ). In Denmark, 39.1% of broods (n = 23) and 9.9% of nestlings (n = 192) (107 Otter, K. A., I. R. K. Stewart, P. K. McGregor, A. M. R. Terry, T. Dabelsteen, and T. Burke (2001). Extra-pair paternity among Great Tits Parus major following manipulation of male signals. Journal of Avian Biology 32:338–344. Close ), and in Germany, 40% of broods (n = 78) and 8.5% of nestlings (n = 681) were extra-pair (101 Strohbach, S., E. Curio, A. Bathen, J. Epplen, and T. Lubjuhn (1998). Extrapair paternity in the great tit (Parus major): a test of the “good genes” hypothesis. Behavioral Ecology 9(4):388–396. Close ). Showing the interannual variability, in a five-year study in Germany, the proportion of broods containing at least one extra-pair young varied by year from 27.8–44.2% (n = 265), and the total percentage of extra-pair young in the population varied from 5.4–8.6% of nestlings (n = 2,013) (100 Lubjuhn, T., Strohbach, S., Brun, J., Gerken, T. and Epplen, J.T. (1999). Extra-pair paternity in Great Tits (Parus major) – a long term study. Behaviour. 136(9): 1157-1172. Close ).

Significant research has gone into examining the factors that influence the rate of extra-pair paternity. Some of the factors can include: 1) habitat: significantly more broods contain extra-pair young in urban habitats (48.9% of broods, n = 45; 10.5% of nestlings, n = 393) than in forest habitats (24.4% of broods, n = 41, 4.15% of nestlings, n = 458) in Hungary (108 Pipoly, I., K. Szabó, V. Bókony, B. Preiszner, G. Seress, E. Vincze, J. Schroeder, and A. Liker (2019). Higher frequency of extra-pair offspring in urban than forest broods of Great Tits (Parus major). Frontiers in Ecology and Evolution 7:229. Close ); 2) age: males paired with older females sire more extra-pair young (103 Roth, A. M., J. A. Firth, S. C. Patrick, E. F. Cole, and B. C. Sheldon (2019). Partner’s age, not social environment, predicts extrapair paternity in wild great tits (Parus major). Behavioral Ecology 30(6):1782–1793. Close ), and males in their first year are more likely to be cuckolded, and have a higher proportion of young in their nest sired by extra-pair males (102 Lubjuhn, T., T. Gerken, J. Brün, and T. Schmoll (2007). Yearling male great tits, Parus major, suffer more strongly from cuckoldry than older males. Zoology 110(5):387–397. Close ); 3) behavior/personality: pairs with similar behavioral syndromes have higher extra-pair paternity than pairs with dissimilar personalities (106 van Oers, K., P. J. Drent, N. J. Dingemanse, and B. Kempenaers (2008). Personality is associated with extrapair paternity in great tits, Parus major. Animal Behaviour 76(3):555–563. Close ), and females that are more innovative and less neophobic have more extra-pair young (109 Bókony, V., I. Pipoly, K. Szabó, B. Preiszner, E. Vincze, S. Papp, G. Seress, T. Hammer, and A. Liker (2017). Innovative females are more promiscuous in great tits (Parus major). Behavioral Ecology 28(2):579–588. Close ); and 4) individual quality: cuckolded males have less genetic variation than those who are not cuckolded (107 Otter, K. A., I. R. K. Stewart, P. K. McGregor, A. M. R. Terry, T. Dabelsteen, and T. Burke (2001). Extra-pair paternity among Great Tits Parus major following manipulation of male signals. Journal of Avian Biology 32:338–344. Close ). Males that sire extra-pair young have more fledglings than cuckolded males, but do not recruit more offspring in to the population (101 Strohbach, S., E. Curio, A. Bathen, J. Epplen, and T. Lubjuhn (1998). Extrapair paternity in the great tit (Parus major): a test of the “good genes” hypothesis. Behavioral Ecology 9(4):388–396. Close ).

Brood Parasitism of Conspecifics

No or little apparent intraspecific brood parasitism (8 putative instances in n = 616 nests, 110 Rohwer, F. C., and S. Freeman (1989). The distribution of conspecific nest parasitism in birds. Canadian Journal of Zoology 67:239–253. Close ; 1 instance in n = 82 broods, 105 Verboven, N., and A. C. Mateman (1997). Low frequency of extra-pair fertilizations in the Great Tit Parus major revealed by DNA fingerprinting. Journal of Avian Biology 28:231–239. Close ). Experimental egg additions are generally accepted and not removed (111 Kempenaers, B., R. Pinxten, and M. Eens (1995). Intraspecific brood parasitism in two tit Parus species: occurrence and responses to experimental parasitism. Journal of Avian Biology 26(2):114–120. Close ).

Brood Parasitism of Other Species

The Great Tit is not an obligate brood parasite, but intense competition for cavity nest sites leads to some facultative brood parasitism. The Great Tit and the Eurasian Blue Tit (Cyanistes caeruleus) occasionally form mixed clutches in each others' nests (112 Petrassi, F., A. Sorace, F. Tanda, and C. Consiglio (1998). Mixed clutches of Blue Tits Parus caeruleus and Great Tits Parus major in nest boxes in Central Italy. Ornis Svecica 8:49–52. Close , 113 Barrientos, R., J. Bueno-Enciso, E. Serrano-Davies, and J. J. Sanz (2015). Facultative interspecific brood parasitism in tits: a last resort to coping with nest-hole shortage. Behavioral Ecology and Sociobiology 69:1603–1615. Close ). In Spain, mixed-species clutches had a prevalence of 3.0% (n = 1,285 nests), with higher prevalence in smaller woodland patches compared to large (113 Barrientos, R., J. Bueno-Enciso, E. Serrano-Davies, and J. J. Sanz (2015). Facultative interspecific brood parasitism in tits: a last resort to coping with nest-hole shortage. Behavioral Ecology and Sociobiology 69:1603–1615. Close ). The Eurasian Blue Tit tends to dump eggs in Great Tit nests as a brood parasite, whereas Great Tit tends to usurp Eurasian Blue Tit nests and take over the clutch, adding its own eggs (113 Barrientos, R., J. Bueno-Enciso, E. Serrano-Davies, and J. J. Sanz (2015). Facultative interspecific brood parasitism in tits: a last resort to coping with nest-hole shortage. Behavioral Ecology and Sociobiology 69:1603–1615. Close ). Mixed-species clutches are more frequent at higher population densities (113 Barrientos, R., J. Bueno-Enciso, E. Serrano-Davies, and J. J. Sanz (2015). Facultative interspecific brood parasitism in tits: a last resort to coping with nest-hole shortage. Behavioral Ecology and Sociobiology 69:1603–1615. Close ). Nestlings of the Great Tit that are raised by Eurasian Blue Tit show misimprinting, which results in low recruitment and reproductive success (114 Slagsvold, T. and B. T. Hansen (2001). Sexual imprinting and the origin of obligate brood parasitism in birds. The American Naturalist 158(4):354–367. Close ).

The Great Tit also competes with Ficedula flycatchers for nest sites (see Social and Interspecific Behavior), and in rare instances, the Great Tit has raised nestlings of European Pied Flycatcher (Ficedula hypoleuca) in mixed broods, and vice-versa (115 Slagsvold, T. (1975). Competition between the Great Tit Parus major and the Pied Flycatcher Ficedula hypoleuca in the breeding season. Ornis Scandinavica 6(2):179–190. Close ).

Degree of Sociality

Occurs singly or in pairs during the breeding season. More gregarious from late summer to early spring, forming large groups. Large groups may consist solely of juveniles, and in winter flock sizes reach up to 50 individuals (including first-years and adults) as part of a mixed-species foraging flock (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). Summer flocks are more loosely organized and less integrated than winter flocks (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). When the Great Tit forms single-species flocks, those average 5.4 individuals per flock, with the range of mean flock sizes across studies 2.0–12.0 (n = 5 studies) (116 Beauchamp, G., and I. Krams (2023). Flock size increases with the diversity and abundance of local predators in an avian family. Oecologia 202:629–637. Close ).

Single-species flocks form dominance hierarchies, with males dominant over females and adults dominant over first-years (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). Dominance in winter flock is correlated with higher winter survival and more surviving offspring (117 Lambrechts, M. M., and A. A. Dhondt (1986). Male quality, reproduction, and survival in the Great Tit (Parus major). Behavioral Ecology and Sociobiology 19:57–64. Close ). Birds in suburban areas show less flocking behavior and establish dominance hierarchies around bird feeders (85 Gompertz, T. (1961). The vocabulary of the Great Tit. British Birds 54:369–394. Close ).

Nonpredatory Interspecific Interactions

The Great Tit frequently joins, and is a major component of, mixed-species foraging flocks, particularly in the nonbreeding season. In Europe, the most common species that join the Great Tit in mixed-species foraging flocks are other species of tit, including Eurasian Blue Tit (Cyanistes caeruleus), Coal Tit (Periparus ater), Marsh Tit (Poecile palustris), Willow Tit (Poecile montanus), Gray-headed Chickadee (Poecile cinctus), Sombre Tit (Poecile lugubris), Crested Tit (Lophophanes cristatus), and Long-tailed Tit (Aegithalos caudatus); other core species of mixed flocks include Eurasian Nuthatch (Sitta europaea), Eurasian Treecreeper (Certhia familiaris), and Goldcrest (Regulus regulus) (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 118 Alerstam, T., S. G. Nilsson, and S. Ulfstrand (1974). Niche differentiation during winter in woodland birds in southern Sweden and the Island of Gotland. Oikos 25(3):321–330. Close , 119 Morse, D. H. (1978). Structure and foraging patterns of flocks of tits and associated species in an English woodland during the winter. Ibis 120(3):298–312. Close , 120 Herrera, C. M. (1979). Ecological aspects of heterospecific flocks formation in a Mediterranean passerine bird community. Oikos 33:85–96. Close , 121 Alatalo, R. V. (1981). Interspecific competition in tits Parus spp. and the Goldcrest Regulus regulus: foraging shifts in multispecific flocks. Oikos 37(3):335–344. Close , 122 Székely, T., and T. Juhász (1993). Flocking behaviour of tits (Parus spp.) and associated species: the effect of habitat. Ornis Hungarica 3:1–6. Close , 123 Panayotopoulou, M., J.-A. Nilsson, and H. Kallander (2005). The winter social system of sombre tits Parus lugubris, in Greece. Lund University, Sweden. Close ). Other species or less frequent species that join mixed-flocks include Lesser Spotted Woodpecker (Dryobates minor), Great Spotted Woodpecker (Dendrocopos major), Short-toed Treecreeper (Certhia brachydactyla), Common Chaffinch (Fringilla coelebs), Common Firecrest (Regulus ignicapilla), Common Chiffchaff (Phylloscopus collybita), Willow Warbler (Phylloscopus trochilus), Eurasian Blackcap (Sylvia atricapilla), and Spotted Flycatcher (Muscicapa striata) (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 119 Morse, D. H. (1978). Structure and foraging patterns of flocks of tits and associated species in an English woodland during the winter. Ibis 120(3):298–312. Close , 120 Herrera, C. M. (1979). Ecological aspects of heterospecific flocks formation in a Mediterranean passerine bird community. Oikos 33:85–96. Close , 122 Székely, T., and T. Juhász (1993). Flocking behaviour of tits (Parus spp.) and associated species: the effect of habitat. Ornis Hungarica 3:1–6. Close ).

Beauchamp and Krams (116 Beauchamp, G., and I. Krams (2023). Flock size increases with the diversity and abundance of local predators in an avian family. Oecologia 202:629–637. Close ) summarized mixed-flock sizes across studies, and found a mean size of 11.2 birds, with a range of means across studies of 3.3–37.0 (n = 20 studies). Mixed flocks establish dominance hierarchies among species, with Great Tit being most dominant, likely on account of its size relative to other tit species (119 Morse, D. H. (1978). Structure and foraging patterns of flocks of tits and associated species in an English woodland during the winter. Ibis 120(3):298–312. Close ). Mixed flocks show some partitioning of foraging strata, with the Great Tit tending to forage more on the inner and lower parts of trees (119 Morse, D. H. (1978). Structure and foraging patterns of flocks of tits and associated species in an English woodland during the winter. Ibis 120(3):298–312. Close , 121 Alatalo, R. V. (1981). Interspecific competition in tits Parus spp. and the Goldcrest Regulus regulus: foraging shifts in multispecific flocks. Oikos 37(3):335–344. Close ). Mixed flocks are larger and more diverse in areas with higher predation risk (124 Székely, T., T. Szép and T. Juhász (1989). Mixed species flocking of tits (Parus spp.): a field experiment. Oecologia 78:490–495. Close , 116 Beauchamp, G., and I. Krams (2023). Flock size increases with the diversity and abundance of local predators in an avian family. Oecologia 202:629–637. Close ).

The Great Tit competes with other species over cavities suitable as nest sites, which are limited. Competition with Ficedula flycatchers can be fatal, with the Great Tit killing intruder European Pied Flycatcher (Ficedula hypoleuca) or Collared Flycatcher (Ficedula albicollis) during attempted nest takeovers by the flycatcher (115 Slagsvold, T. (1975). Competition between the Great Tit Parus major and the Pied Flycatcher Ficedula hypoleuca in the breeding season. Ornis Scandinavica 6(2):179–190. Close , 125 Merilä, J., and D. A. Wiggins (1995). Interspecific competition for nest holes causes adult mortality in the Collared Flycatcher. The Condor 97(2):445–450. Close , 126 Ahola, M. P., T. Laaksonen, T. Eeva, and E. Lehikoinen (2007). Climate change can alter competitive relationships between resident and migratory birds. Journal of Animal Ecology. 76: 1045-1052. Close , 127 Samplonius, J. M., and C. Both (2019). Climate change may affect fatal competition between two bird species. Current Biology 29:327–331. Close ). European Pied Flycatcher attempt to take over Great Tit nests by harassing the adult flying in/out of the nest until it deserts, or by attempting to rapidly build a new nest over the Great Tit nest or eggs while the cavity is unoccupied (115 Slagsvold, T. (1975). Competition between the Great Tit Parus major and the Pied Flycatcher Ficedula hypoleuca in the breeding season. Ornis Scandinavica 6(2):179–190. Close ). When the Great Tit encounters a flycatcher inside its nest cavity, the flycatcher is most often killed (115 Slagsvold, T. (1975). Competition between the Great Tit Parus major and the Pied Flycatcher Ficedula hypoleuca in the breeding season. Ornis Scandinavica 6(2):179–190. Close ). Despite this, the direction of competition is strongly biased, with European Pied Flycatcher evicting and destroying Great Tit nests but Great Tit only rarely evicting and destroying European Pied Flycatcher nests (128 Busse, P., and J. Gotzman (1962). Nesting competition and mixed clutches among some birds inhabiting the nest-boxes. Acta Ornithologica 7:1–32. Close , 115 Slagsvold, T. (1975). Competition between the Great Tit Parus major and the Pied Flycatcher Ficedula hypoleuca in the breeding season. Ornis Scandinavica 6(2):179–190. Close ). In Gotland, Sweden, a mean total of 4.2% of adult Collared Flycatcher were killed by tits annually in nest box study populations (range 0–17%), and most were killed during the Great Tit egg laying period (125 Merilä, J., and D. A. Wiggins (1995). Interspecific competition for nest holes causes adult mortality in the Collared Flycatcher. The Condor 97(2):445–450. Close ). In Groningen, Netherlands, between 0.4–8.9% of adult male European Pied Flycatcher in study populations are killed annual by Great Tit (127 Samplonius, J. M., and C. Both (2019). Climate change may affect fatal competition between two bird species. Current Biology 29:327–331. Close ). Mortality of the European Pied Flycatcher is highest for late-arriving and unpaired males, and there is no apparent negative population effect of the male mortality caused by interactions with Great Tit (127 Samplonius, J. M., and C. Both (2019). Climate change may affect fatal competition between two bird species. Current Biology 29:327–331. Close ). Fatal competition between these species increases at higher densities of both species (125 Merilä, J., and D. A. Wiggins (1995). Interspecific competition for nest holes causes adult mortality in the Collared Flycatcher. The Condor 97(2):445–450. Close , 126 Ahola, M. P., T. Laaksonen, T. Eeva, and E. Lehikoinen (2007). Climate change can alter competitive relationships between resident and migratory birds. Journal of Animal Ecology. 76: 1045-1052. Close , 127 Samplonius, J. M., and C. Both (2019). Climate change may affect fatal competition between two bird species. Current Biology 29:327–331. Close ). Competition between European Pied Flycatcher and Great Tit in the Netherlands may be declining over time as climate change drives more asynchrony in the timing of breeding between these species, with Great Tit shifting to earlier laying dates at a faster rate (127 Samplonius, J. M., and C. Both (2019). Climate change may affect fatal competition between two bird species. Current Biology 29:327–331. Close ).

Kinds of Predators

The Eurasian Sparrowhawk (Accipiter nisus) and the Eurasian Pygmy-Owl (Glaucidium passerinum) are well-studied as major predators of the Great Tit. Other potential avian predators (reviewed in 116 Beauchamp, G., and I. Krams (2023). Flock size increases with the diversity and abundance of local predators in an avian family. Oecologia 202:629–637. Close ) include Northern Hawk Owl (Surnia ulula), Boreal Owl (Aegolius funereus) (129 Korpimäki, E. (1988). Diet of breeding Tengmalm's Owls Aegolius funereus: long-term changes and year-to-year variation under cyclic food conditions. Ornis Fennica 65(1):21–30. Close ), Tawny Owl (Strix aluco) (130 Sasvari, L., and Z. Hegyi (1998). Bird predation by tawny owls (Strix aluco L.) and its effect on the reproductive performance of tits. Acta Oecologica 19(6):483–490. Close ), Ural Owl (Strix uralensis) (131 Vrezec, A., P. Saurola, A. Avotins, S. Kocijančič, S. Sulkava (2018). A comparative study of Ural Owl Strix uralensis breeding season diet within its European breeding range, derived from nest box monitoring schemes. Bird Study 65, supplement 1:85–95. Close ), Little Owl (Athene noctua), Merlin (Falco columbarius), Eurasian Hobby (Falco subbuteo) (132 Zawadzka, D., and J. Zawadzki (2001). Breeding populations and diets of the Sparrowhawk Accipiter nisus and the Hobby Falco subbuteo in the Wigry National Park (NE Poland). Acta Ornithologica 36.1:25–31. Close ), Peregrine Falcon (Falco peregrinus), and Great Gray Shrike (Lanius excubitor) (133 Lorek, G., P. Tryjanowski and J. Lorek. (2000). Birds as prey of the Great Grey Shrike (Lanius excubitor). Ring 22 (1):37-44. Close ) .

Major nest predators include weasels, such as the least weasel (Mustela nivalis, the most significant nest predator in southern England; 134 Dunn, E. (1977). Predation by weasels (Mustela nivalis) on breeding tits (Parus spp.) in relation to the density of tits and rodents. Journal of Animal Ecology 46(2):633–652. Close ) and European pine marten (Martes martes) (135 Kaliński, A., J. Wawrzyniak, M. Bańbura, J. Skwarska, P. Zieliński, M. Glądalski, and J. Bańbura (2014). Does the threat of European pine marten (Martes martes) predation influence the height of nests built by Blue Tits (Cyanistes caeruleus) and Great Tits (Parus major)? Avian Biology Research 7(2):83–90. Close ), squirrels, including the red squirrel (Sciurus vulgaris) and the introduced eastern gray squirrel (Sciurus carolinensis) (136 Broughton, R. K. (2020). Current and future impacts of nest predation and nest‐site competition by invasive eastern grey squirrels Sciurus carolinensis on European birds. Mammal Review 50:38–51. Close ), other rodents, such as yellow-necked mouse (Apodemus flavicollis) or forest dormouse (Dryomys nitedula)(137 Maziarz, M., T. Wesolowski, G. Hebda, M. Cholewa, and R. K. Broughton (2016). Breeding success of the Great Tit Parus major in relation to attributes of natural nest cavities in a primeval forest. Journal of Ornithology 157:343–354. Close ), the Great Spotted Woodpecker (Dendrocopos major) (138 Skwarska, J. A., A. Kalinski, J. Wawrzyniak, and J. Banbura (2009). Opportunity makes a predator: Great Spotted Woodpecker predation on Tit broods depends on nest box design. Ornis Fennica 86(3):109–112. Close ), and snakes, for example the Montpellier snake (Malpolon monspessulanus) and ladder snake (Zamenis scalaris) in Spain (139 Navalpotro, H., D. Mazzoni, and J. C. Senar (2021). A plastic device fixed around trees can deter snakes from predating bird nest boxes. Animal Biodiversity and Conservation 44.1:103–108. Close ). Fledglings are predated by Eurasian Jay (Garrulus glandarius), Great Spotted Woodpecker, Eurasian Sparrowhawk, and martens (140 Naef-Daenzer, B., F. Widmer, and M. Nuber (2001). Differential post-fledging survival of great and coal tits in relation to their condition and fledging date. Journal of Animal Ecology 70(5):730–738. Close ).

Response to Predators

The Great Tit responds to potential predators spotted in flight by given a high-pitched trill alarm call and diving to cover (inducing flock members to do the same), where it stays silent for minutes (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ).

Predators spotted while perched are mobbed. Mobbing birds raise their crests, flick their wings, give various scolding calls, and make diving attacks towards the predator (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). The Great Tit sometimes mobs the Common Cuckoo (Cuculus canorus), which mimics the Eurasian Sparrowhawk in appearance, and is not a brood parasite of the Great Tit (141 Davies, N. B., and J. A. Welbergen (2008). Cuckoo-hawk mimicry? An experimental test. Proceedings of the Royal Society B Biological Sciences 275:1817–1822. Close ). Dominance is inversely correlated with return to foraging after the appearance of a predator; dominant birds wait longer than subordinates (142 de Laet, J. F. (1985). Dominance and anti-predator behaviour of Great Tits Parus major: a field study. Ibis 127:372–377. Close ).

The Great Tit varies its D call (also called churring calls or mobbing calls, see Vocal Array) according to the predator and context (143 Carlson, N. V., S. D. Healy, and C. N. Templeton (2017). A comparative study of how British tits encode predator threat in their mobbing calls. Animal Behaviour 125:77–92. Close , 88 Kalb, N., and C. Randler (2019). Behavioral responses to conspecific mobbing calls are predator-specific in great tits (Parus major). Ecology and Evolution 9:9207–9213. Close , 144 Kalb, N., F. Anger, and C. Randler (2019). Subtle variations in mobbing calls are predator-specific in great tits (Parus major). Scientific Reports 9:6572. Close ). Kalb et al. (144 Kalb, N., F. Anger, and C. Randler (2019). Subtle variations in mobbing calls are predator-specific in great tits (Parus major). Scientific Reports 9:6572. Close ) found that avian predators perceived as a higher threat (e.g. Eurasian Sparrowhawk, which preys on small birds like Great Tit; 132 Zawadzka, D., and J. Zawadzki (2001). Breeding populations and diets of the Sparrowhawk Accipiter nisus and the Hobby Falco subbuteo in the Wigry National Park (NE Poland). Acta Ornithologica 36.1:25–31. Close ) are given longer D calls with more syllables, longer syllables, and longer intervals between syllables, compared to those D calls given in response to avian predators perceived as a lower threat (e.g. Tawny Owl, which mainly eats small mammals but occasionally eats passerines like Great Tit; 145 Galeotti, P., F. Morimando, and C. Violani (1991). Feeding ecology of the tawny owls (Strix aluco) in urban habitats (northern Italy). Italian Journal of Zoology 58:143–150. Close , 130 Sasvari, L., and Z. Hegyi (1998). Bird predation by tawny owls (Strix aluco L.) and its effect on the reproductive performance of tits. Acta Oecologica 19(6):483–490. Close , 146 Ýmihorski, M., and G. Osojca (2006). Diet of the tawny owl (Strix aluco) in the Romincka Forest (NE Poland). Acta Zoologica Lituanica 16:46–52. Close ). The Great Tit is attracted to a mob by conspecific’s mob calls, and it approachs faster and closer in response to lower-threat D calls (88 Kalb, N., and C. Randler (2019). Behavioral responses to conspecific mobbing calls are predator-specific in great tits (Parus major). Ecology and Evolution 9:9207–9213. Close ). Males tend to approach mobbing calls faster and closer than females (88 Kalb, N., and C. Randler (2019). Behavioral responses to conspecific mobbing calls are predator-specific in great tits (Parus major). Ecology and Evolution 9:9207–9213. Close ).

The response of incubating or brooding females against intruders in the nest cavity is a variable Hissing DIsplay, given with the head raised, white cheeks fluffed, the crown sleeked, wings drooped and spread (or raised), and the tail spread. The bird stands and hisses with head thrust forward, or waves the head side to side with the bill open. This display is accompanied by bill snaps and by clapping the wings to produce loud sounds (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). Males react to react to red squirrels (Sciurus vulgaris) outside the nest with hissing, flicking wings, tink calls, and aggressive flights at the squirrel (19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ). A rare distraction display in response to young distress calls includes "impeded, moth-like flight" (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ).

Nestlings (starting at 6–8 d old) and fledglings respond to parental alarm calls by going silent and cowering down; excessive disturbance in the nest may cause premature fledging (84 Hinde, R. A. (1952). The Behaviour of the Great Tit Parus major and Some Related Species. Behaviour Supplement 2. E. J. Brill, Leiden, Netherlands. Close , 19 Cramp, S., and C. M. Perrins, Editors (1993). The Birds of the Western Palearctic. Volume 7. Flycatchers to shrikes. Oxford University Press, Oxford, UK. Close ).