Tawny Owl Females of Intermediate Colour Morph Reproduce Better and Increase in a Local Polymorphic Population
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Ornis Fennica 94: 33–44. 2017 Tawny Owl females of intermediate colour morph reproduce better and increase in a local polymorphic population Gintarë Graytë, Saulius Rumbutis, Mindaugas Dagys, Daiva Vaitkuvienë & Rimgaudas Treinys* Graytë, G., Dagys, M., Vaitkuvienë, D., Treinys, R., Nature Research Centre, Akademi- jos str. 2, 08412 Vilnius, Lithuania. * Corresponding authors e-mail: rimga.t@gmail. com Rumbutis, S., T. Ivanauskas Zoological Museum, Laisvës ave. 106, 44253 Kaunas, Lithu- ania Received 31 August 2016, accepted 15 January 2017 Individuals of the same population with differing melanin plumage colouration can re- spond differently through fitness to environmental stressors. In a local population of Tawny Owls (Strix aluco), we collected individual-based data on nesting attempts of this well-known colour polymorphic species between 1985 and 2014. We studied whether body size, condition, demographic parameters and habitat around occupied nest sites dif- fer among grey, intermediate and brown coloured females. The proportions of females belonging to each of the three colour morphs in the local population changed during the study period, withintermediate individua ls becoming more frequent in the last decade. Females were similar in body size and achieved similar body condition during breeding irrespective of plumage colouration. Although most of the analysed demographic param- eters were not statistically significantly related to the colour morph, the intermediate owls raised more nestlings in a lifetime and tended to have higher recapture probability. We found little support for habitat and morph associations. In summary, we propose that the increase in intermediate Tawny Owl colour morphs in this local population is associated with their better reproductive performance compared to the other morphs. 1. Introduction in birds (Fowlie & Krüger 2003, Galeotti et al. 2003). However, regardless of the mechanism that Colour polymorphism is the coexistence of two or results in colour polymorphism in any particular more distinct and genetically determined colour species, this phenomenon is usually regarded as forms in an interbreeding population (Roulin providing adaptive functions to local environmen- 2004). This is a phenomenon observed in many tal conditions (Tate et al. 2016 and references vertebrate taxa including birds, especially birds of therein). The ratio of different colour morphs prey and owls (Fowlie & Krüger 2003, Roulin within a local population of raptor species tends to 2004). Apostatic, disruptive and sexual selection vary spatially along geographical or environmen- are three major hypotheses proposed to explain the tal gradients (Galeotti et al. 2003, Amar et al. origin and maintenance of colour polymorphism 2014, Roulin & Randin 2015, Tate et al. 2016). 34 ORNIS FENNICA Vol. 94, 2017 The relationship between melanin-based plu- Tawny Owl is under strong genetic control, highly mage colouration and fitness in birds is species- heritable and is not sensitive to environmental dependent (Meunier et al. 2011). At temperate lat- conditions (Brommer et al. 2005, Karell et al. itudes in Germany, individuals of the dominant in- 2011b, Emaresi et al. 2014). Tawny Owl individu- termediate morphof Common Buzzard ( Buteo als differing in plumage colouration show some buteo) produced two times more nestlings than the behavioural (Da Silva et al. 2013), physiological individuals of the rare light and dark morph during (Galeotti & Sacchi 2003, Gasparini et al. 2009a, their lifetime (Krüger & Lindström 2001). In the 2009b, Karell et al. 2013, Emaresi et al. 2016), case of Tawny Owls (Strix aluco) breeding at morphological (Roulin et al. 2003) and fitness dif- northern latitudes in Finland, individuals of the ferences (Roulin et al. 2003, 2004, 2008, Brom- grey colour morphsurvived better and produced mer et al. 2005, Piault et al. 2009, Karell et al. more nestlings during their lifetime than brown in- 2011a, 2011b, Da Silva et al. 2013, Emaresi et al. dividuals (Brommer et al. 2005). In a long-term 2014). Differently coloured Tawny Owls tend to study of Swainsons Hawk (Buteoswainsoni )in be associated withspecifi c regional environmental California, however, Briggs et al. (2011a) did not conditions (Galeotti & Cesaris 1996). This expec- find any significant differences in survival and tation, however, may be valid in a local population various reproductive parameters among individu- as well (Roulin et al. 2003) because environmental als of three colour morphs. Similarly, brood size heterogeneity can promote the evolution of was independent of colour morph in the dimorphic phenotypic plasticity with individuals matching Booted Eagle (Aquila pennata) population in their phenotypes with the environment (Roulin et south-eastern Spain (Martínez et al. 2016). Re- al. 2008). Galeotti and Sacchi (2003) showed that gardless of whether colour polymorphism influ- differently coloured individuals are associated ences fitness or not, the ratios of morphs usually withvarying habitatsat a fine scale. Moreover, in- remain stable over time within local populations of dividuals of similar plumage colouration in dis- various raptor species (Brommer et al. 2005, tinct populations under different environmental Briggs et al. 2011a, Martínez et al. 2016). At the conditions may even show opposite patterns of the distribution range scale, however, temporal same fitness trait (cf. Brommer et al. 2005, Ema- changes in plumage pigmentation may be region- resi et al. 2014) indicating that morphs are adapted specific (Roulin et al. 2011). The stable long-term to specific environmental conditions that vary ratio may be a result of different and temporally al- geographically. Further, the effect of global warm- ternating performance strategies of the colour ing may vary spatially and subsequently, distinct morphs at a fine scale (Roulin et al. 2003). Global colour morphs may respond differently, in terms warming, however, can selectively affect individ- of fitness, to the same environmental stress uals in different ways in a given population (Roulin 2014). Karell et al. (2011b) demonstrated (Roulin 2014). Karell et al. (2011b) demonstrated how environmental changes differently drive the that changes in the climatic conditions resulted in fitness of the colour morphs of Tawny Owls in the an improvement in the fitness of brown-coloured same geographical area. Thus, former findings for individuals in Tawny Owls, leading to an increase any particular species in a specific geographical in the number of individuals of this morph in the area, habitat and/or time period may not be a uni- population. versal reflection of the fitness of a colour morph In this study, we investigated fitness and habi- and its change over time within that species. tat in the well-known polymorphic species Tawny Using data from breeding Tawny Owl females, Owl. Individuals of this generalist nocturnal pre- captured over a period of 30 years in a temperate dator vary from pale grey to reddish-brown, but forest in Lithuania, Central Europe, we investi- two (i.e., grey and brown) or three (i.e., grey, inter- gated whether body size, reproductive perfor- mediate and brown) distinct morphs are most fre- mance and habitat around nest sites is colour quently described withtheir proportions spatially morphdependent. Additionally, we predict that: 1) distributed in Europe without a clear geographical proportions of morphs change over years; 2) re- cline (Galeotti & Cesaris 1996, Karell et al. production and survival are morphdependent; 3) 2011b). Melanin-based plumage colouration in the relationship between mean winter tempera- Graytë et al.: The importance of colour morph for fitness in Tawny Owl 35 Fig. 1. Location of the study site (forest area = black fill) in Këdainiai district (bold line), Li- thuania. tures, mean spring temperatures and reproductive using one of two methods, namely accounting for parameters differs among morphs; and 4) habitat a) the age of nestlings; b) egg laying intervals (ev- composition differs between nest sites occupied ery second day) in cases of incomplete clutches by females of the three colour morphs. were found during the early nest site checks. The age of the oldest nestling was estimated using a nestling age identification key compiled by S. 2. Material and methods Rumbutis (unpubl. data). The start of breeding then was determined by backdating calculation, 2.1. Data accounting for the incubation duration (30 days; Southern 1970). For further analyses, we ex- Data for this study were collected between 1985 pressed the start of breeding in Julian days and 2014 in the Dotnuva forest (area: 39 km2), sur- (wherein 1st January is day 1), also accounting for rounded by agricultural fields in Këdainiai district, leap years. Thelengthand maximum breadthof Central Lithuania (55°2327 N, 23°4625 E) the eggs in full clutches were measured using a (Fig. 1). The terrain is flat and deciduous tree spe- sliding calliper to the nearest 0.1 mm. Egg volume cies dominate in the forest. All available nest was calculated as p × egg length× breadth 2/6000 boxes (nest boxes were first erected in 1977; from (Tatums index; Hakkarainen & Korpimäki 1993). 30 to 55 nest boxes were available annually for oc- For further analyses, the mean volume of the eggs cupation between 1985 and 2014) were checked in theclutchand clutchsize were used as response by S. Rumbutis for occupation from February to variables. Brood size was measured as the number March, depending on the winter weather