Sex differences in preening behaviour in the Ciconia ciconia

Katarzyna Maria ZOLNIEROWICZ 1*, Marketa ONDROVA-NYKLOWA 2, Marcin TOBOLKA 1

1 Institute of Zoology, Poznań University of Life Sciences, Wojska Polskiego 71C, 60-625 Poznań, Poland, *e-mail: [email protected] (corresponding author) 2  Department of Zoology and Laboratory of Ornithology, Faculty of Science, Palacky University, 17. Listopadu 50, CZ-771 46 Olomouc, Czech Republic

ARTICLE INFO ABSTRACT Regular research paper Preening is a type of maintenance behaviour in , published in that fulfils an important role in grooming. Preening may also Pol. J. Ecol. (2016) 64: 406–410 be important for signalling quality of mates. Therefore we hypothesized that the frequency of preening may be related to sex received after revision and population density of White Stork Ciconia ciconia. We observed preening activity of 25 pairs in Western Poland at the beginning XXXX 2016 of incubation, when preening frequency is the highest. Birds were observed on the nest, because most preening behaviour occurs doi there. We found that being on the nest males spent proportionally 10.3161/15052249PJE2016.64.3.xxx more time on preening than females (on average 30% vs. 16%). Females spent more time preening when their mate was present at the nest. There was no significant relationship between preening key words frequency and indirect quality indicators (arrival date, laying date, preening in birds hatching date, clutch size, brood size) nor between the distance distance to neighbour, to nearest neighbours and the time males and females spent on sex-specific behaviour preening.

Preening is an important grooming behav- ver, preening behaviour can increase in fre- iour in birds which can help maintain feather quency when size increases due to social quality by removing dirt and oiling (Zampi- facilitation (Palestis and Burger 1998) or ga et al. 2004, Griggio et al. 2010) or reduc- as a displacement activity resulting from high ing number of ectoparasites (Møller 1991, group density (Mills and Faure 1989, Keel- Rózsa 1993, Waite et al. 2012). Preening ing 1994). also can play an important role in birds’ mat- Preening is a type of behaviour described ing. Males of budgerigars Melopsittacus un- and studied in many species, mainly in dulatus which preened more often were more captivity, like e.g. budgerigars (Zampiga attractive for females due to better et al. 2004, Griggio and Hoi 2006, Grig- reflectance which is called “attractive preen- gio et al. 2010), domestic canaries Serinus ca- ing” hypothesis (Griggio et al. 2010). On naria (Lenouvel et al. 2009), Anas the other hand it is found that self-preening platyrhynchos (Delogu et al. 2010), feral pi- (grooming of own feathers) and allo-preen- geons Columba livia (Rózsa 1993, Waite et ing (grooming of feathers of other individual, al. 2012), zebra finches Taeniopygia guttata e.g. a partner) may promote horizontal trans- (Kulkarni and Heeb 2007), with only few fer of bacteria (Kulkarni and Heeb 2007) studies in the wild: Hirundo rustica and viruses (Delogu et al. 2010). Therefore (Møller 1991) and Sterna spp. (Van an alternative hypothesis has been devel- Iersel and Bol 1958, Palestis and Burger oped that bird females may avoid too fre- 1998). All of these birds are known as hosts quent preening males to constraint parasites for mites and lice, which consume feather spread. It is called “preening avoidance” hy- keratin (Loye and Zuk 1991, Møller 1991). pothesis (Griggio and Hoi 2006). Moreo- Therefore some authors suggested that preen- Preening in the White Stork Ciconia ciconia 407 ing frequency may be a proxy for individual Stork, and to assess if preening in this species fitness which is a predictor for arrival date, may be an indicator of fitness and related to clutch size and breeding success (Møller breeding parameters. We tested if preening 1991). However the exact role of preening is is sex related (may play a role in mating or still being discussed. pair display) and also if preening frequency The European White Stork Ciconia cico- may be modified by the distance to neigh- nia is an example of long-lived monomorphic bours, which is an indicator of population bird species. It nests solitary on the top of hu- density and intraspecific interactions. There- man made structures like electricity poles, fore we put forward following hypotheses: 1) chimneys, roofs or trees, however can form Frequency of preening differs between pair aggregations of several pairs or even colonies members; 2) Preening behaviour is related to (Tryjanowski et al. 2006). Also in White distance to nearest neighbours; and finally 3) Stork assemblages of lice (Fryderyk and Preening (as a proxy for fitness) is related to Izdebska 2009) and their removal were well several breeding parameters like arrival dates, described (Clayton and Cotgreave 1994, time of breeding and number of egg. Bocheński and Jerzak 2006) but preen- The study was conducted in Western ing was not quantified or studied in details. Poland near the town of Leszno (51°51′N, Preening in White Stork occurs mostly on 16°34′E). This is an area of arable fields -in the nest, during incubation and after juve- terspersed with meadows, pastures, human niles have fledged Bocheński( and Jerzak settlements, small river valleys and woods. 2006). In contrast to many dimorphic spe- In this location White Storks generally build cies, there is no suggestion that of isolated nests on electricity poles, chimneys stork as a monomorphic bird is important and roofs of buildings. Rarely White Storks for mate choice (Bocheński and Jerzak also nest on trees far from human settlements 2006). However, in the light of fact that birds (Tobolka et al. 2013). In the studied popu- can see in Ultraviolet (Benett and Cuthill lation the mean distance to three nearest ac- 1994) preening can play an important role in tive nests was 2.36 km (range: 0.14–5.9) so it feather light reflectance and therefore in mate can influence time budget of several pairs, i.e. choice. Hence, we hypothesized that male preening behaviour (Bocheński and Jer- storks preen more frequent than females to zak 2006). attract females and show better reflectance of Fieldwork was carried out from March to feathers as an effect of good fitness. Moreo- May 2011. Each of 25 pairs was observed once ver, if the preening frequency is an effect of in the beginning of incubation period. Ob- ectoparasites occurrence it may diminish fit- servation lasted for two hours and was con- ness and finally arrival dates on the breed- ducted only in good weather conditions (no ing ground, clutch size (in females) and final rain or strong wind). During the incubation breeding success. there is always at least one member of the pair White Stork is also known for its very good on the nest. For each observed bird we noted eyesight. It can see predator or neighbour time spent on nest and on preening. Because from even 2–3 km. Occurrence of other stork both birds in each pair did not spend the en- can modify its time budget (Bocheński and tire two hours observation period on the nest, Jerzak 2006). Feather maintenance is time the percentage of time spent on preening dur- and energy consuming (Croll and McLar- ing their presence on the nest was calculated. en 1993). Therefore, we hypothesized that in In addition, we used mean distance to the higher densities White Storks have to spend three nearest neighbours (Clark and Evans more time on food foraging and defending 1954), as an index of the density of stork nests their nests, so they have less time and energy around each focal nest. We used QUANTU- for preening, especially in the beginning of MGIS with geoportal.gov.pl wms label. breeding season when interactions and ag- In our study we could not catch adult gressive behaviour are frequently observed birds and directly measure individuals’ body (Bocheński and Jerzak 2006). size or assess the number of ectoparasites. We Here we attempt to describe the preening decided to use indirect indicators of individ- behaviour (i.e. self-preening) of the White ual condition such as arrival date and brood 408 Katarzyna Maria Zolnierowicz et al.

Fig. 1. Differences in time spent on preening between females and males of White Storks Ciconia cico- nia. Bold line – median, box – 25 and 75 quartile, whiskers – min and max, ₒ – outlayer. size (Kosicki et al. 2004). Arrival date of 2006). To explain differences in time spent on the first and the second bird from pair, lay- preening in comparison to distance to neigh- ing date, hatching date, clutch size and final bours or breeding parameters we used Pearson breeding success were obtained by direct in- correlation. Results are presented as means ± spections in the nests and special question- SD. All statistical analyses were prepared us- naire forms delivered to farmers living nearby ing IBM SPSS Statistics 20 for Windows. the nests. We defined arrival date as the day Females spent significantly more time on when the particular bird occurred on its nest nests than males (respectively, on average. 97 (day 1 = 1 January) (details in Ptaszyk et al. and 66 min during 120 min. of observation, 2003 and Tobolka et al. 2015). We include in t = -2.84, P = 0.009, n = 25). The percent- the present study only pairs with detailed and age of time spent on preening was greater in certain arrival dates collected by experienced males (30.1 ± 24.8%) than in females (16.2 ± observers because sometimes, some nest are 15.1%) (t =2.10, P = 0.046, n = 25). We found visited not only by particular breeding pair, that an individual’s preening activity was un- also by nonbreeders (Wuczyński 2005) We related to the preening activity of their mate defined date of laying as the day when the first (r= -0.33, P = 0.11, n = 25). Females’ relative egg was laid in the nest. This was estimated on time spent on preening was positively corre- the basis of direct inspection in nest (details lated with the presence of males (r = 0.49, P = in Tobolka et al 2015). Clutch size is a num- 0.014, n = 25). ber of eggs in the nest during the first inspec- We did not find any relationship between tion when the clutch was complete. Breeding time spent on preening by males and females success was a number of fledglings able to fly during the 2h observation and the mean dis- (Tryjanowski et al. 2006). During the ob- tance to three closest nests of neighbours servation the sex of pair members was easily (respectively r = 0.19 P = 0.36, and r = 0.11, determined by observing the position during P = 0.59, n = 25). copulation, which is a reliable method for this We did not find any statistically signifi- bird species (Chernetsov et al. 2006). cant relationships (P> 0.20 in all cases) be- We used t paired-test because female be- tween breeding parameters like arrival date, haviour is potentially related to male behav- time of egg laying or clutch size and preening iour and vice versa (Bocheński and Jerzak of both adults. Preening in the White Stork Ciconia ciconia 409

We found that male White Storks spent is establishing (Bocheński and Jerzak 2006). relatively more time on preening at the nest Probably during the , even in than females. We also found that the time conditions of high breeding pairs density ag- spent on preening by females was positively gressive interactions are less frequent. There- correlated with male presence at the nest, fore more detailed studies on White Stork which may suggest the role of preening in behaviour overlapping entire breeding season communication between mates. In the light (each breeding stages) are needed to assess of Griggio and Hoi (2006) who tested if the role of preening in mating and pair dis- female budgerigars use male preening time play having regard external factors like popu- (“attractive preening” hypothesis) as a qual- lation density or even natural predators (e.g. ity signal of males, our results suggest that White-tailed Eagle) occurrence. preening also for White Storks can play a role in pair display. Female budgerigars spent sig- ACKNOWLEDGEMENTS: The study was support- nificantly more time near the preened males ed by a grant from the National Science Centre N/ than unpreened (Zampiga et al. 2004). NZ8/01186, a scholarship NSC T/NZ8/01001 and However our results did not allow us to con- a scholarship from the European Social Fund in firm nor to reject the “preening avoidance” 2010/2011 (MT). We would like to thank to Tomáš hypothesis saying that females should avoid Grim, Daniel Hanley and Piotr Tryjanowski for their males which spend a lot of time on preening, helpful comments to manuscript. because of probable high number of ectopar- asites on them (Griggio and Hoi 2006. To test this hypothesis appropriately we should REFERENCES take into account more variables and collect more numerous data. In our study there was Bennett A.T.D., Cuthill I.C 1994 – Ultraviolet vi- no relationship between condition indicators sion in birds: What is its function? – Vision we chose (arrival date, laying date, breeding Res. 34: 1471–1478. success, clutch size) and time spent on preen- Bocheński M., Jerzak J. 2006 – Behaviour of the ing. But this issue needs more detailed stud- White Stork Ciconia ciconia: a review (In: The ies including adults’ catching, body condition White Stork in Poland: studies in biology, ecol- measurements and detailed ectoparasites ogy and conservation, Eds: P. Tryjanowski, analyses. T.H. Sparks, L. Jerzak) – Bogucki Wydawnict�- We did not find a positive correlation be- wo Naukowe, Poznan, pp. 295–324. tween the distance to the nearest neighbours, Chernetsov N., Chromik W., Dolata P.T., Profus which may be an indicator of higher den- P., Tryjanowski P. 2006 – Sex-related natal sity in the local population (Janiszewski dispersal of White Storks (Ciconia ciconia) in et al. 2013), and time spent on preening. In Poland: how far and where to? – The Auk, 123: contrast, the results of a study conducted on 1103–1109. Common terns show that males that nested Clark P.J., Evans F.C. 1954 – Distance to nearest close to other males spent more time preen- neighbor as a measure of spatial relationships ing (Palestis and Burger 1998). However, in populations – Ecology, 35: 445–453. distances between nests in Terns colonies Clayton D.H., Cotgreave P. 1994 – Relationship are much shorter then between White Stork of bill morphology to grooming behaviour in nests, especially in Western Poland (Try- birds – Anim. Behav. 47: 195–201. janowski et al. 2006). Intraspecific interac- Croll D.A., McLaren E. 1993 – Diving metabo- tions connected with the distance to neigh- lism and in common and bours in studied White Stork population thick-billed murres – J. Comp. Physiol. B. 163: could modify the time budget to spend more 160–166. time on nest defending, screening or forag- Delogu M., De Marco M.A., Di Trani L., Raffini E., ing than preening, especially in the initial pe- Cotti C., Puzelli S., Ostanello F., Webster R.G., riod of breeding season. However, aggressive Cassone A., Donatelli I. 2010 – Can preening behaviour is more often in the early begin- contribute to influenza A virus infection in ning of the breeding season when hierarchy wild waterbirds? – PLoS ONE, 5: e11315. 410 Katarzyna Maria Zolnierowicz et al.

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