Post-Breeding Aggressive Territorial Behaviour in Eurasian Tree Sparrows

Home , European pied flycatcher, Tit (bird)

Tichodroma 27: 2–10 (2015) ISSN 1337-026X

Post-breeding aggressive territorial behaviour in Eurasian Tree Sparrows (Passer montanus): territory or mate defence? Pohniezdne agresívne teritoriálne správanie u vrabca poľného (Passer montanus): hájenie teritória alebo partnera?

Jan Pinowski¹, Barbara Pinowska¹, Jerzy Romanowski2 & Radovan Václav3

1 Daniłowskiego 1/33, 01 833 Warsaw, Poland ² Faculty of Biology and Environmental Studies UKSW, Wóycickiego 1/3, 01-938, Warsaw, Poland 3 Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 84506 Bratislava, Slovakia; e-mail: [email protected]

Abstract. The genetic interests of males and females usually differ. Whenever female fitness depends on an access to limited resources such as male parental care, competitive interactions among females are expected to evolve. There are numerous studies reporting female-female aggression during the breeding season, but this subject has not been examined for the post-breeding period. Here we examine female-female aggression in the Eurasian Tree Sparrow Passer montanus occurring during the autumn courtship period. We show that aggressive behaviour between nest owners and intruders is common and often fierce during the post-breeding courtship period. Importantly, females more frequently perform this behaviour than male nest owners and it is sex- but not age-specific (i.e. birds interfere with individuals of the same sex, but regardless of their age). This study, focusing on the post-breeding period, corroborates the idea that female territorial aggression can be involved in mate monopolization rather than simply serving as a territory defence strategy. Moreover, aggressive territorial behaviour of nest owners towards intruders can also serve as a signal of mate quality during the pair-formation period. We propose that examining female territorial behaviours during the post-breeding period can provide additional insights into the causes of diversity in avian mating systems.

Key words: antagonistic behaviour, sexual conflict, post-breeding period, mating systems

Introduction perspective, revealing that females choose mates with superior genotypes, high paternal The genetic interests of males and females investment and skills enhancing fitness of their usually differ, which leads to sexual conflict re- progeny (e.g. Wasser & Waterhouse 1983, garding the level of parental care or the number Rosenqvist & Berglund 1992, Gowaty 1996, of simultaneous social partners (Trivers 1972, 1997, Shuster & Wade 2003). Whenever the Wittenberger & Tilson 1980, Clutton-Brock female’s fecundity or the ability to rear young 1991, see Chapman et al. 2003). depends on an access to limited resources such Traditionally, the literature on mating sys- as male parental care, competitive interactions tems is centred on the role of male behaviour. among females are expected to evolve (Hardy It was only since the 1970s when the subject & Williams 1983). Indeed, it has been proposed started to be examined from the female’s that the diversity of mating systems can partly

2 Tichodroma 27 (2015) be due to female intra-sexual competition e.g. by cal nest-box colony, established in 1960 and female aggression towards the potential mates comprising 33 nest boxes, was located along of their male partners (Veiga 1990). the forest edge (22 nest boxes) and interior (11 Wittenberger & Tilson (1980) suggested five nest boxes) of a 30-year-old pine forest near a conditions for monogamy to evolve in birds with former field station of the Institute of Ecology, the aggression of mated females towards other PAS. Nest boxes were arranged in five rows females representing one of the conditions. and were located 5–8 m apart. No natural tree There are numerous studies reporting female- holes were present in the focal colony. Nest female aggression during the breeding season boxes were used for nesting dominantly by (Breiehagen & Slagsvold 1988, Davies 1989, Tree Sparrows (1961: 18, 1962: 15 and 1963: 4 Møller 1986, Veiga 1990, 1992, Slagsvold et pairs). In addition, 6, 5 and 3 pairs of Great Tits al. 1992, Slagsvold 1993, Slagsvold & Lilfjeld Parus major and 1, 4 and 1 pairs of European 1994, Gowaty 1996, Liker & Székely 1997, Pied Flycatcher Ficedula hypoleuca nested in Rosvall 2011). Moreover, female aggression the focal colony in 1961, 1962, and 1963, re- can indeed inhibit multiple females from set- spectively. One pair of Eurasian Wryneck Jynx tling at the male’s territory (Struchbury & torquilla nested in the colony in 1961. Robertson 1987, Gowaty & Wagner 1988, The Tree Sparrow is a small, sedentary, Hobson & Sealay 1989, Sandell & Smith 1996, semi-colonial, cavity-nesting passerine (Sasvári 1997, Sandell 1998). Though the occurrence & Hegyi 1994, 2000, Summers-Smith 1995), of female-female aggression during the post- defending a territory within 2 m of its nest site breeding period would corroborate the role of (Sano 1974, 1988). At the end of the breed- aggression in preserving social monogamy, this ing season, adult and fully-grown juvenile subject has not been examined for the post- Tree Sparrows leave the breeding colony to breeding pair-formation period. moult. During this period flocks are formed Here we examine aggression behaviour by in feeding and roosting sites outside nesting nest owners towards nest intruders in Eurasian colonies (Pinowski 1965, 1967). Timing of Tree Sparrow Passer montanus (hereafter only moult completion of juvenile and adult Tree Tree Sparrow). The behaviour is studied during Sparrows is gradual: 15, 30 and 99% of all Tree the post-breeding period when pair formation Sparrows finish their moult by mid-September, takes place in sexually mature birds follow- end-September, and mid-October, respectively ing the breeding season and post-breeding/ (Pinowski et al. 2009). After the moult period post-juvenile moult. Based on ringing data and and a gradual return to nesting colonies, from behavioural observations, we aim to establish the beginning of September until the end of the basic qualitative and quantitative character- October, younger immature birds normally istics of aggressive territorial behaviour by nest prospect nest sites while adult and older imma- owners against intruders. We address if territo- ture birds defend a nest site and perform autumn rial aggression by pair members can represent courtship (Pinowski et al. 2014). a territory defence strategy or it can rather be In 1961, Tree Sparrow nest owners were viewed as a strategy by nest owners, particularly observed at 31 nest-boxes during the autumn females, to monopolize their social mates. courtship period, but nest owners were identified unambiguously only at 18 nest-boxes. At Materials and methods least one nest owner per nest box was identified at 23 of 32 and 8 of 23 nest-boxes in 1962 and 1963, respectively. Study area and species The study was carried out between the Kampinos Behavioural observations National Park and the Vistula River, NW of Antagonistic behaviour of Tree Sparrows, which Warsaw, Poland (52º20’ N, 20º50’ E). The fo- was the primary focus of this study, was collect-

Tichodroma 27 (2015) 3 ed each year during the autumn courtship period 1 m of the nest box and identified by ring com- from the beginning of September to the begin- bination, 2) individuals on the roof of the nest ning of November in 1961–1963. The species’ box, looking at the entrance hole from the roof, autumn courtship activity normally commenced perching at the entrance hole and inspecting nest about an hour after sunrise and continues until content in the absence of identified owners, and noon, but the duration depends on weather. 3) individuals chased by identified owners and Courtship activity was shorter on rainy and cool engaged in fights with them, mostly with female days when birds spent more time away from the owners. In this study, the term “chasing” is used colony feeding in fields (Pinowski 1966, 1967, when the owners gave warning calls tre-tre Pinowski et. al. 2009). Birds were observed and actively drove the bird away from the nest nearly every day from early morning until noon. box area (ca. 1m radius). The term “fighting” In total, birds were observed during 128 days is used when the act of chasing was combined (38, 50 and 40 days in 1961, 1962 and 1963, with physical attack represented by aerial fights respectively). Observations were conducted eventually followed by combats on the ground. from two positions within the colony, using a 10 × 50 binocular. The observer rotated between Data analysis the two positions in hourly intervals during the The difference in the frequency of aggressive whole time of bird presence in the colony. behaviour occurrence between males and fe- The ring combination (see Pinowski et al. males within pairs was tested using the Sign 2014 for details on ringing) and behaviour of test because the data used represents matched individual birds and time of observation were samples. The difference in the proportion of recorded for all birds sighted at or near a nest aggressive males and females of different age box. In total, birds were observed for 760 hours was tested with the Binomial test. The deviation during the three years. In addition to the period between the observed and expected frequencies of autumn courtship, sexual and antagonistic of aggressive behaviour with respect to age was behaviour of Tree Sparrows was observed dur- tested with the Chi-square test. The tests were ing the breeding period (involving first broods calculated with Statistica 7 (StatSoft, Tulsa, only) to detect eventual cases of polygyny. OK, USA) and R software (R Core Team 2014). During the autumn courtship period multiple Tree Sparrows can be observed at or near single Results nests. The same birds observed at the nest at least three times during three different days or during two days each day spending at the nest Focal colony mating system structure at least three hours were considered to be pair The structure of a focal colony during the breed- members and nest owners. Most important ing and autumn courtship periods is quantita- evidence for the association of specific birds tively described in Table 1. No case of polygyny with a given nest box during the autumn court- was detected during the breeding period in ship period were observations of nest building 1961−1963. In 1961, one of the pairs (1/18, 6%), behaviour, copulation attempts, pair perching on for which pair member identity was determined, the nest box and night roosting in the nest box was polygynous during the post-breeding pe- in winter, and the occupation of the nest box by riod. An immature male, an immature female, the birds in the immediately preceding or fol- and a female of unknown age formed the pair. lowing breeding seasons (Pinowski & Noskov Each of the two females occupied a different 1981, Pinowski et al. 2006, 2008, 2009). In ad- nest box. In 1962, one polygynous pair (1/23, dition, nest owners were inferred based on their 4%) was recorded during the autumn courtship antagonistic responses to intruders. In contrast, period. In this case, it was an adult male first intruding/prospecting birds not holding a nest paired with a female of unknown age and an site refer to 1) strange individuals present within adult female. Later on during autumn, two im-

4 Tichodroma 27 (2015) Table 1. Breeding and post-breeding parameters of a Tree Sparrow colony studied between 1961 and 1963. Tab. 1. Hniezdne a pohniezdne ukazovatele sledovanej kolónie vrabca poľného v r. 1961 až 1963.

Year / Rok Colony parameters / Charakteristiky kolónie 1961 1962 1963 No. of all nest boxes / 33 33 33 Počet všetkých búdok No. of nest boxes used during the breeding season / 18 15 4 Počet búdok obsadených počas hniezdnej sezóny No. of nest boxes used during the autumn courtship period / 31 32 23 Počet búdok obsadených počas obdobia jesenného toku No. of nest boxes with autumn nests / 28 27 12 Počet búdok s vybudovanými jesennými hniezdami mature females replaced these females. In 1963, nest owners towards nest visitors. The single two monogamous pairs, in both cases comprised exception involved a case when a male visitor of adult birds, each occupied two nest boxes, chased a male nest owner, but tolerated the nest but no case of polygyny (0/8, 0%) was detected owner’s female partner. during the autumn courtship period. The fights between nest owners and intruders tended to occur more often at nest boxes Description of post-breeding situated 20 m from the forest edge compared to antagonistic behaviour nest boxes situated at the forest edge – of 12 nest Post-breeding antagonistic behaviour between boxes where we detected fights, 7 nest boxes nest box owners and nest box visitors was ob- were located in the forest interior. Furthermore, served from September 16 till October 21, from all (7/7) fights occurring during 1962 and 1963 September 8 till October 13, and from October 2 at nest boxes situated 20 m from the forest edge till 11 in 1961, 1962, and 1963, respectively. The occurred at those nest boxes, which had not been frequency of antagonistic interactions peaked at occupied in the previous breeding season and days when prospectors most intensively visited which were rarely prospected during the autumn nest boxes. The following stimuli elicited an courtship period. antagonistic response of nest owners in the post- breeding period: 1) nest visitors entering the nest Sex and age determinants of post- box and 2) nest visitors perching at the entrance breeding antagonistic behaviour hole and the roof of the nest box. When a pair Female nest box owners were more likely than held two nest boxes during the autumn courtship male owners to be involved in chases with period, female owners chased intruders away intruders. Namely, when single nest owners from the secondary nest box, then returning to momentarily defended nest boxes, 2 male and the primary nest box. In the polygynous trios, 15 female nest box owners were observed to each of the females occupied and defended their chase intruders (Sign binomial test, Z = 2.91, own nest box. p = 0.004). Similarly, 7 male and 16 female The antagonistic behaviour normally in- nest box owners chased intruders alone when volved only chasing. In some cases, female both partners were momentarily present at the owners entered their nest box immediately after nest box (Z = 1.67, p = 0.095). A joint response chasing. If nest owners failed to chase intruders by male and female nest owners was observed away, aggression escalated into an aerial foot- on 37 occasions with 54% (20/37) of the joint gripping fight with birds usually falling, locked chases occurring when multiple intruders visited together, to the ground where their combat the nest box. continued. It was possible to approach such With respect to fighting behaviour, female locked birds within the reach of the hand. The owners were generally more likely than male combatants flew up and down to the ground up owners to be involved in physical interference to six times during their fight. With one excep- with intruders. Five female and no male owners tion, we only recorded physical aggression of were observed to attack intruders in cases when

Tichodroma 27 (2015) 5 single nest owners were momentarily present owners, after accounting for the proportion of at nest boxes (Z = 1.79, p = 0.074), and eight adult to immature nest owners, the frequency of female and no male owners were observed to antagonistic encounters was not related to the fight intruders in cases when both owners were owner’s age for neither sex. Finally, antagonistic present at nest boxes (Z = 2.47, p = 0.013). Only encounters, particularly in terms of fighting a single case of a joint fight of nest owners with behaviour among females, occurred dominantly an intruder was observed during three years – in a relatively poorer breeding habitat. a male joined his female partner who initiated Intra-sexual antagonistic behaviour among the attack with an intruder. territory owners and intruders is not exclu- Chases by nest box owners of known age sive to Tree Sparrows (c.f. Sano 1974, 1988). were performed mostly by adult birds, but A similar pattern was described for Common this was significant only for male owners Magpie Pica pica (Birkhead 1979), Eurasian (male owners: 19 adult vs. 6 immature birds, Nuthatch Sitta europea (Mattyhsen 1986), Tree binomial test, p = 0.015; female owners: 13 Swallow Tachycineta bicolor (Stutchbury & adult vs. 11 immature birds, binomial test, p Robertson 1987), Common Blue Tit Cyanistes = 0.839). Yet, after considering the proportion caeruleaus (Kempaeners 1994), or Common of adult to immature nest box owners of both Starling Sturnus vulgaris (Sandell 1998). sexes (81.3% and 53.3% of males and females, Based on ringing data, we show for a sexually respectively, were adults), the frequency of monomorphic species that the frequency of chases performed by adult birds did not differ antagonistic behaviour is higher in female than from expected frequencies (males: χ2 = 0.45, male nest owners, but it is not related to age in p = 0.502; females: χ2 = 0.01, p = 0.935). Five neither sex. Thus, antagonistic territorial behav- adult and three immature female nest owners iour in Tree Sparrows during the post-breeding and a single immature male nest owner were courtship period is sex- but not age-specific. observed to fight intruders. Superficially, these results indicate that in Tree Chased or fought intruders predominantly Sparrows individuals of each sex select and comprised immature birds (2 adult vs. 24 defend a specific nest site and leave mate selec- immature birds, binomial test, p < 0.001), ir- tion up to intra-sexual competition. Moreover, respective of the sex of the birds chasing them our results suggest sexual conflict between nest (female chasers: 1 adult vs. 16 immature birds, owners whereby females attempt to monopolize binomial test, p < 0.001; male chasers: 1 adult the reproductive effort of their social partners. vs. 8 immature birds, binomial test, p = 0.039). Though a wealth of literature exists on The observed proportion of chased/attacked female mate and nest selection in birds, for immature to adult intruders did not differ from which males initially acquire nest sites, the the expected proportion (in 84.4% cases, intrud- subject is not clear. For example, Leffelaar & ers were immature birds; χ2 = 1.24, p = 0.266). Robertson (1985) suggested that female Tree Swallows compete over males with nest-sites, Discussion rather than for a male or a nest-site alone. For European Pied Flycatcher, however, which has This study reports that antagonistic behaviour similar nest site requirements as Tree Sparrows, between nest owners and intruders is common a series of studies indicated that females choose and often fierce during the post-breeding court- the quality of the nest site and its surroundings, ship period in Tree Sparrows, suggesting it can rather than the quality of male nest owners substantially affect the birds’ time and energy (Alatalo et al. 1984, 1986, Slagsvold 1986). budgets. Antagonistic behaviour was more fre- Yet, more recent studies have pointed out that quently performed by females and took place females routinely use multiple cues when select- within sexes. Though adult male nest owners ing a mate and his territory (Burley 1981, Dale initiated more chases than immature male nest & Slagsvold 1996, see Candolin 2003).

6 Tichodroma 27 (2015) Tree Sparrows rarely survive four years and lize the reproductive effort of their mates and both sexes participate in parenting (Summers- keep them socially monogamous (c.f. Veiga Smith 1995). Even though males do not develop 1990). Such a female adaptation would not a brood patch and their incubation is less ef- be superfluous because paternal investment in ficient (Pinowski et al 1973, Anderson 1978), Tree Sparrows is substantial during the breeding both sexes take an approximately equal share in season with males contributing to nest build- feeding young (Summers-Smith 1995, Samchuk ing, incubation and chick rearing (Pinowski et al. 1981). Consequently, in line with the lat- 1966, 1967, Pinowski et al. 1973, Pinowski ter studies, it is unlikely that female choice in & Pinowska 2009). Moreover, though such Tree Sparrows operates passively and solely information is not available for Tree Sparrow, through territory acquisition because male pa- polygynous male House Sparrows were found rental contribution is an important determinant to invest in the nests of secondary females of female nesting success in this biparental and significantly less than in their primary nests short-lived species. (North 1980, Veiga 1990). Finally, similarly as One of our important findings is a high reported for House Sparrow, we observed male proportion of joint chases conducted by male Tree Sparrows to tolerate strange females pros- and female nest owners against intruders. pecting their territory and occasionally create Moreover, male chases against intruders oc- polygynous bonds during the autumn courtship curred several times more often when their period. Nevertheless, polygyny has only rarely mates were momentarily with them at nest been detected during the breeding season in boxes. Therefore, even though antagonistic Tree Sparrow (Creutz 1949, Berck 1961–1962, behaviour in Tree Sparrows is sex-specific, both Deckert 1962, Sano 1974, 1988, Weise 1992) mates often conduct it simultaneously and its with some of the aforementioned studies attain- occurrence appears to depend on mate presence. ing polygamy only by manipulating nest-site Consequently, antagonistic behaviour in Tree availability using nest-boxes. Therefore, while Sparrows during the autumn period may not female aggressive behaviour can be viewed as a only serve to defend a territory from intruders, female monopolization attempt of male parental but can also be a component of courtship display care, one would expect polygyny to occur more with the intensity of this behaviour reflecting regularly and at higher rates in Tree Sparrow. individual quality. This suggestion is similar to It is intriguing to contrast the results of this that on female infanticidal behaviour in House study with those reported by Veiga (1990, 1992, Sparrows Passer domesticus, stating that female 2004) for a closely related House Sparrow. aggressive behaviour can serve as a signal of First, while female-female territorial interfer- quality and facilitate female acceptance in the ence is common in both species during the pair male’s territory (Veiga 2004). formation period, social polygamy during the Our results revealed that the frequency breeding period appears to show higher rates in of antagonistic behaviour was disproportion- House Sparrows (see Anderson 2006). Second, ately higher in female than male nest owners. in contrast to social polygamy rates, genetic Moreover, with a single exception, physical polygamy rates seem comparable between the interference among nest owners and intruders two species (Cordero et al. 2002, Seress et al. occurred only among females. Finally, female- 2007), even though the range of extra-pair female physical interference dominantly oc- paternity levels in House Sparrow is known curred in a poorer nesting habitat of woodland to vary considerably across populations and interior where male parental assistance is likely years (see Anderson 2006). We suggest that it of high importance. These results conclusively would be worthwhile to investigate the within- point to the existence of sexual conflict in Tree population relationships between the intensity Sparrows with female aggressive behaviour of female-female competition and the degree possibly representing an adaptation to monopo- of social and genetic polygamy in the species

Tichodroma 27 (2015) 7 with variable mating systems such as House dá sa očakávať evolúcia zložitých kompetetív- and Tree Sparrows. That is to say, in addition nych vzťahov medzi samicami. Hoci existuje to male paternity assurance and female extra- viacero štúdií o sexuálnej agresivite medzi pair copulation behaviours, female-female samicami, pre mimohniezdne obdobie nebola aggressive behaviour could be an important táto problematika doposiaľ študovaná. V tejto determinant not only of social but also genetic práci sme sledovali teritoriálnu agresivitu medzi mating systems. samicami vrabca poľného (Passer montanus) In conclusion, we provide correlational v pohniezdnom období počas jesenného toku. evidence that female territorial aggression can Zistili sme, že agresívne správanie medzi maji- be involved in mate monopolization rather than teľmi hniezdnych búdok a cudzími vtákmi je aj simply serving as a territory defence strategy. v pohniezdnom období bežné a často veľmi in- It is not surprising that this female behaviour is tenzívne. Samice vykazovali všeobecne vyššiu conspicuous during the post-breeding courtship frekvenciu agresívneho správania ako samci, period because, due to elevated winter mortality, pričom agresívne správanie bolo pohlavne the number of sexually active birds is markedly ale nie vekovo špecifické (t.j. vtáky napádali higher in the autumn compared to spring court- jedincov rovnakého pohlavia ale nezávisle ship periods (Pinowski 1968). Consequently, od veku). Táto práca podporuje hypotézu, že we propose that examining female territorial samičie teritoriálne agresívne správanie môže behaviours during the post-breeding period u niektorých vtáčích druhov predstavovať stra- can provide important insights into the causes tégiu na monopolizáciu partnerov a nemusí byť of diversity in avian mating systems. Finally, prostým prejavom hájenia teritória. Agresívne our study lends tentative support to the idea teritoriálne správanie u vrabca poľného tiež that aggressive territorial behaviour of nest môže slúžiť ako signál individuálnej kvality, owners towards intruders during the pair- ktorý je určený partnerom počas obdobia formation period can also serve as a signal of vytvárania a utužovania párových zväzkov. female quality, facilitating mate acceptance at Výsledky tejto práce naznačujú, že teritoriálne the territory and strengthening the pair bond in agresívne správanie v pohniezdnom období biparental species such as Tree Sparrow. Further môže objasniť určitú variabilitu v rozmanitosti studies should experimentally assess if male vtáčích rozmnožovacích systémov. quality or operational sex ratio affect the level of female intra-sexual aggression during the References pair-formation period and how the latter relates to variation in mating systems. Alatalo R. V., Lundberg A. & Glynn C. 1986: Female pied flycatchers choose territory quality and not male Acknowledgments characteristics. — Nature 323: 152–153. We are indebted to M. Borowiec, A. Dyrcz, T. Svagsvold, K. Alatalo R. V., Lundberg A. & Ståhlbrandt K. 1984: Vincent and two anonymous referees for important critical Female mate choice in the pied flycatcher Ficedula comments on early drafts of the manusxcript. We thank to all hypoleuca. — Behav. Ecol. Sociobiol. 14: 253–261. colleagues who helped with field work. The methods applied Anderson T. R. 1978: Population studies of European in this study conform to the animal welfare regulations valid sparrows in North America. — Occ. Papers Mus. Nat. in the 1960s when the study was conducted. Hist. Kansas. 70: 1–58. Anderson T. R. 2006: Biology of the ubiquitous house spar- Súhrn row: from genes to populations. — Oxford University Press, Oxford. Genetické záujmy samcov a samíc sú zvyčajne Berck K. H. 1961–1962: Beiträge zur Ethologie des Feld- odlišné. Kedykoľvek reprodukčný úspech samíc sperlings (Passer montanus) und dessen Beziehung závisí od prístupu k limitujúcim zdrojom, akými zum Haussperling (Passer domesticus). — Die Vo- sú napríklad samčia rodičovská starostlivosť, gelwet 82/83: 129–173, 8–16.

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