Population dynamics of an expanding passerine at the distribution margin

This is the peer reviewed version of the following article: Karvonen, J.; Orell, M.; Rytkönen, S.; Broggi, J.; Belda Perez, EJ. (2012). Population dynamics of an expanding passerine at the distribution margin. Journal of Avian Biology. 43(2):102-108. doi:10.1111/j.1600-048X.2011.05376.x., which has been published in final form at http://dx.doi.org/10.1111/j.1600-048X.2011.05376.x. This article may be used for non-commercial purposes in accordance With Wiley Terms and Conditions for self-archiving [http://olabout.wiley.com/WileyCDA/Section/id-817011.html ]Individuals may be maladapted to novel environments at the species' distribution margin. We investigated population dynamics in a marginal habitat where reproduction has been proven poor. Survival, population growth rate (¿) and its components, breeding and natal dispersal were studied in great tits Parus major breeding at the northern margin of its distribution in northern Finland. We used long term capture-mark-recapture data sets. Study area size and population density were used to explain adult survival rates. The average annual estimates of adult survival rose from 0.371 to 0.388 between the periods of 1971-1984 and 1999-2009. The estimates are slightly lower than estimates of small passerines in Europe. Low local survival rate of fledglings (0.050-0.055) probably reflects intensified emigration from this low quality area. Temporal variation in ¿ was large (0.498-1.856). Despite of low adult survival and recruitment rates, the mean estimates of ¿ (1.008 and 1.033) indicate an overall stability in the population size. Indeed, our results suggest that the immigration has an important role in the population dynamics of northern great tits. Thus the population is demographically and genetically dependent on core habitats which may cause adaptive problems due to intensive gene flow. Given those limitations, options for evolution of local adaptations in northern distribution margins are discussed.Satu Lampila, Mikko Ojanen, Suvi Ponnikas, Kari Koivula and numerous other field workers helped with data collection over the years. Veli-Matti Pakanen and Emma Vatka helped with the manuscript. Financial support for this study was provided by the Research Council for Biosciences and Environment of the Academy of Finland.Karvonen, J.; Orell, M.; Rytkönen, S.; Broggi, J.; Belda Pérez, EJ. (2012). Population dynamics of an expanding passerine at the distribution margin. Journal of Avian Biology. 43(2):102-108. https://doi.org/10.1111/j.1600-048X.2011.05376.xS102108432Bauchau, V., & Van Noordwijk, A. J. (1995). Comparison of survival estimates obtained from three different methods of recapture in the same population of the great tit. Journal of Applied Statistics, 22(5-6), 1031-1038. doi:10.1080/02664769524775Broggi, J., Hohtola, E., Orell, M., & Nilsson, J.-Å. (2005). 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Immune Response to Newcastle Disease Virus Vaccination in a Wild Passerine

We studied the immune response of wild House Sparrows (Passer domesticus) experimentally challenged with different doses of inactivated Newcastle disease virus (NDV) vaccine. We evaluated within-individual cell-mediated and humoral responses in birds kept in outdoor aviaries, over a 6-wk period. Nonbreeding adult House Sparrows developed a significant humoral response to NDV experimental vaccination within 1 wk postchallenge, as measured by hemagglutination inhibition assay; values increased until week 4 and persisted until week 6. Differences among treatments appeared by week 1, with individuals challenged with the highest dose (0.2 mL) eliciting a higher humoral response than the rest (n = 18). By week 4, all individuals vaccinated displayed an increased humoral response, with individuals challenged with the highest dose remaining significantly above responses of individuals vaccinated with the middle dose (0.1 mL, n = 14), but not the lowest dose (0.05 mL, n = 15). The middle and lowest dose responded similarly and significantly different from controls (n = 23). Differences persisted through week 6 postchallenge. Cell-mediated responses were independent of the experimental treatment. All individuals experienced a rise in granulocyte concentration, whereas lymphocyte and monocyte concentrations decreased, most likely as a result of captivity. Adult wild House Sparrows immunochallenged with inactivated NDV vaccine developed a specific humoral response, highlighting the utility of this technique in immunologic and evolutionary ecology studies in wild birds.Peer reviewe

Interpopulation Variation in Contour Feather Structure Is Environmentally Determined in Great Tits

Background: The plumage of birds is important for flying, insulation and social communication. Contour feathers cover most of the avian body and among other functions they provide a critical insulation layer against heat loss. Feather structure and composition are known to vary among individuals, which in turn determines variation in the insulation properties of the feather. However, the extent and the proximate mechanisms underlying this variation remain unexplored. Methodology/Principal Findings: We analyzed contour feather structure from two different great tit populations adapted to different winter regimes, one northern population in Oulu (Finland) and one southern population in Lund (Sweden). Great tits from the two populations differed significantly in feather structure. Birds from the northern population had a denser plumage but consisting of shorter feathers with a smaller proportion containing plumulaceous barbs, compared with conspecifics from the southern population. However, differences disappeared when birds originating from the two populations were raised and moulted in identical conditions in a common-garden experiment located in Oulu, under ad libitum nutritional conditions. All birds raised in the aviaries, including adult foster parents moulting in the same captive conditions, developed a similar feather structure. These feathers were different from that of wild birds in Oulu but similar to wild birds in Lund, the latter moulting in more benign conditions than those of Oulu. Conclusions/Significance: Wild populations exposed to different conditions develop contour feather differences either due to plastic responses or constraints. Environmental conditions, such as nutrient availability during feather growth play a crucial role in determining such differences in plumage structure among populations

Specific Appetite for Carotenoids in a Colorful Bird

Background: Since carotenoids have physiological functions necessary for maintaining health, individuals should be selected to actively seek and develop a specific appetite for these compounds. Methodology/Principal Findings: Great tits Parus major in a diet choice experiment, both in captivity and the field, preferred carotenoid-enriched diets to control diets. The food items did not differ in any other aspects measured besides carotenoid content. Conclusions/Significance: Specific appetite for carotenoids is here demonstrated for the first time, placing these compounds on a par with essential nutrients as sodium or calcium

Patterns of variation in energy management in wintering tits (<em>Paridae</em>)

Abstract Winter energy management in small passerines living year-round in boreal or alpine areas presumably results in strong selective pressure since they need to find food, at a time when natural resources diminish and become less available, and energy requirements increase dramatically. In this thesis energy management during the non-breeding season was studied in three species of tits (Parus spp.), from three different populations: Coll de Pal (Spanish Pyrenees), Lund (Southern Sweden) and Oulu (Northern Finland). Energy management strategies vary significantly between species and among populations and individuals of the same species. Such differences may depend on several environmental factors, food predictability and individual characteristics. Birds from the studied populations appear to react to energetic challenges on a short-term basis and in a highly flexible way. The coal tit (Parus ater) in Coll de Pal and the willow tit (Parus montanus) in Oulu, both hoarding species, relied mostly on short-term management of energy for winter survival. Social and residence status appeared to be the most important factors in determining the level of energy reserves, underlining the importance of food predictability for energy management in wintering tits. Further studies were carried out on two distinct populations of great tit (Parus major) exposed to different winter hardiness. Birds from both populations increased their resting metabolic rate (MR) with experimentally decreasing ambient temperatures. Birds from Oulu maintained higher expenditures than birds from Lund in all cases, but also experienced higher energetic cost of thermoregulation at the lowest temperatures. The differences probably did not arise from a differential insulation capacity between populations, despite the differences in plumage structure found, but from a differential metabolic acclimatization. Birds from Lund probably became hypothermic at the lowest temperatures, which may have exceeded the levels they were acclimatized for. The observed differences in basal MR in laboratory conditions were consistent in wild birds throughout the non-breeding season. Birds from both populations experienced similar patterns of variation in basal MR, with expenditures increasing with mass but decreasing with day length, size and age. Great tits modulate their energy expenditure in a flexible way as a means for surviving the non-breeding season. Further, despite such flexibility, populations appear to be locally adapted for such metabolic acclimatization. These results may have important implications on their life-history and distribution. Winter acclimatization appears to be a complex set of entangled strategies that are based on a metabolic adjustment to cope with changing energy requirements. Other mechanisms that apparently play a secondary role, for example the long term management of reserves through fattening or hoarding, or conserving heat through hypothermia and by developing a better insulative plumage, are certainly important emergency strategies that in natural conditions may explain how some populations can endure winter conditions

PRED & STAR_ data

Los datos corresponden a un estudio experimental sobre el compromiso entre del riesgo de depredacion e inhanición en el Carbonero común (Parus major) y su efecto sobre el peso y el metabolismo basal. Encontramos que aves cautivas en aviarios al exterior, si disponen del tiempo necesario, promueven cambios en el coste energetico de mantenimiento (tasa metabolica basal) antes que el peso, que indicaria el nivel de reservas interno. Estos resultados tienen gran relevancia dado que cuestionan el enfoque tradicional los estudios de energética en vertebrados endotermos, que primava la regulación de las reservas como practicamente el unico mecanismo adaptativo de regulacion.Peer reviewe

Individual response in body mass and basal metabolism to the risks of predation and starvation in passerines

Wintering energy management in small passerines has focused on the adaptive regulation of the daily acquisition of energy reserves within a starvation-predation trade-off framework. However, the possibility that the energetic cost of living, i.e. basal metabolic rate (BMR), is being modulated as part of the management energy strategy has been largely neglected. Here, we addressed this possibility by experimentally exposing captive great tits (Parus major) during winter to two consecutive treatments of increased starvation and predation risk for each individual bird. Body mass and BMR were measured prior to and after each week-long treatment. We predicted that birds should be lighter but with a higher metabolic capacity (higher BMR) as a response to increased predation risk, and that birds should increase internal reserves while reducing their cost of living (lower BMR) when exposed to increased starvation risk. Wintering great tits kept a constant body mass independently of a week-long predation or starvation treatment. However, great tits reduced the cost of living (lower BMR) when exposed to the starvation treatment, while BMR remained unaffected by the predation treatment. Energy management in wintering small birds partly relies on BMR regulation, which challenges the current theoretical framework based on body mass regulation

DATA Fed_Non_Fed

Los datos corresponden a un estudio experimental sobre el efecto de los comederos en dos especies de paridos boreales. Encontramos que el acceso a comederos afectaba al peso, el metabolismo y su interaccion con las condiciones ambientales, de modo distinto en cada especie.Peer reviewe

Population differences in the structure and coloration of great tit contour feathers

Contour feathers cover most of the avian body and play critical roles in insulation, social communication, aerodynamics, and water repellency. Feather production is costly and the development of the optimum characteristics for each function may be constrained by limited resources or time, and possibly also lead to trade-offs among the different characteristics. Populations exposed to different environmental conditions may face different selective pressures, resulting in differences in feather structure and coloration, particularly in species with large geographical distributions. Three resident populations of great tit Parus major L. from different latitudes differed in feather structure and coloration. Individuals from the central population exhibited less dense and longer contour feathers, with a higher proportion of plumulaceous barbs than either northern or southern birds, which did not differ in their feather structure. Ultraviolet reflectance and brightness of the yellow of the contour feathers of the breast was higher for the southern than for the northern population. Birds with greener plumage (higher hue) had less dense but longer feathers, independently of the population of origin. Differences in feather structure across populations appear to be unrelated to the contour feather colour characteristics except for hue. Nutritional and time constraints during molt might explain the pattern of feather structure, whereas varying sexual selection pressure might underlie the coloration patterns observed. Our results suggest that different selective pressures or constraints shape contour feather traits in populations exposed to varying environmental conditions.Peer reviewe

Transgenerational effects enhance specific immune response in a wild passerine

Vertebrate mothers transfer diverse compounds to developing embryos that can affect their development and final phenotype (i.e., maternal effects). However, the way such effects modulate offspring phenotype, in particular their immunity, remains unclear. To test the impact of maternal effects on offspring development, we treated wild breeding house sparrows (Passer domesticus) in Sevilla, SE Spain with Newcastle disease virus (NDV) vaccine. Female parents were vaccinated when caring for first broods, eliciting a specific immune response to NDV. The immune response to the same vaccine, and to the PHA inflammatory test were measured in 11-day-old chicks from their following brood. Vaccinated chicks from vaccinated mothers developed a stronger specific response that was related to maternal NDV antibody concentration while rearing their chicks. The chicks’ carotenoid concentration and total antioxidant capacity in blood were negatively related to NDV antibody concentration, whereas no relation with PHA response was found. Specific NDV antibodies could not be detected in 11-day-old control chicks from vaccinated mothers, implying that maternally transmitted antibodies are not directly involved but may promote offspring specific immunity through a priming effect, while other immunity components remain unaffected. Maternally transmitted antibodies in the house sparrow are short-lived, depend on maternal circulation levels and enhance pre-fledging chick specific immunity when exposed to the same pathogens as the mothers