Emilio Barba

The design of artificial nestboxes for the study of secondary hole-nesting birds: a review of methodological inconsistencies and potential biases

Abstract. The widespread use of artificial nestboxes has led to significant advances in our knowledge of the ecology, behaviour and physiology of cavity nesting birds, especially small passerines. Nestboxes have made it easier to perform routine monitoring and experimental manipulation of eggs or nestlings, and also repeatedly to capture, identify and manipulate the parents. However, when comparing results across study sites the use of nestboxes may also introduce a potentially significant confounding variable in the form of differences in nestbox design amongst studies, such as their physical dimensions, placement height, and the way in which they are constructed and maintained. However, the use of nestboxes may also introduce an unconsidered and potentially significant confounding variable due to differences in nestbox design amongst studies, such as their physical dimensions, placement height, and the way in which they are constructed and maintained. Here we review to what extent the characteristics of artificial nestboxes (e.g. size, shape, construction material, colour) are documented in the ‘methods’ sections of publications involving hole-nesting passerine birds using natural or excavated cavities or artificial nestboxes for reproduction and roosting. Despite explicit previous recommendations that authors describe in detail the characteristics of the nestboxes used, we found that the description of nestbox characteristics in most recent publications remains poor and insufficient. We therefore list the types of descriptive data that should be included in the methods sections of relevant manuscripts and justify this by discussing how variation in nestbox characteristics can affect or confound conclusions from nestbox studies. We also propose several recommendations to improve the reliability and usefulness of research based on long-term studies of any secondary hole-nesting species using artificial nestboxes for breeding or roosting.

Brominated flame retardants and organochlorines in the European environment using great tit eggs as a biomonitoring tool

Large-scale studies are essential to assess the emission patterns and spatial distribution of organohalogenated pollutants (OHPs) in the environment. Bird eggs have several advantages compared to other environmental media which have previously been used to map the distribution of OHPs. In this study, large-scale geographical variation in the occurrence of OHPs, such as polychlorinated biphenyls (PCBs), polybrominated diphenyl ethers (PBDEs) and organochlorine pesticides (OCPs), was investigated throughout Europe using eggs of a terrestrial residential passerine species, the great tit (Parus major). Great tit eggs from 22 sampling sites, involving urban, rural and remote areas, in 14 European countries were collected and analysed (5-8 eggs per sampling site). The environmentally most important congeners/compounds of the analysed pollutants were detectable in all sampling locations. For PCBs, PBDEs and OCPs, no clear geographical contamination pattern was found. Sum PCB levels ranged from 143 ng/g lipid weight (lw) to 3660 ng/g lw. As expected, PCB concentrations were significantly higher in the sampled urban compared to the remote locations. However, the urban locations did not show significantly higher concentrations compared to the rural locations. Sum PBDEs ranged from 4.0 ng/g lw to 136 ng/g lw. PBDEs were significantly higher in the urbanized sampling locations compared to the other locations. The significant, positive correlation between PCB and PBDE concentrations suggests similar spatial exposure and/or mechanisms of accumulation. Significantly higher levels of OCPs (sum OCPs ranging from 191 ng/g lw to 7830 ng/g lw) were detected in rural sampling locations. Contamination profiles of PCBs, PBDEs and OCPs differed also among the sampling locations, which may be due to local usage and contamination sources. The higher variance among sampling locations for the PCBs and OCPs, suggests that local contamination sources are more important for the PCBs and OCPs compared to the PBDEs. To our knowledge, this is the first study in which bird eggs were used as a monitoring tool for OHPs on such a large geographical scale.

Feather mites and birds: an interaction mediated by uropygial gland size?

Feather mites (Arachnida: Acari: Astigmata) feed mainly on secretions of the uropygial gland of birds. Here, we use analyses corrected for phylogeny and body size to show that there is a positive correlation between the size of this gland and mite abundance in passerine birds at an interspecific level during the breeding season, suggesting that the gland mediates interactions between mites and birds. As predicted on the basis of hypothesized waterproofing and antibiotic functions of uropygial gland secretions, riparian/marsh bird species had larger glands and higher mite loads than birds living in less mesic terrestrial environments. An unexpected pattern was a steeper relationship between mite load and gland size in migratory birds than in residents. If moderate mite loads are beneficial to a host but high loads detrimental, this could create complex selection regimes in which gland size influences mite load and vice versa. Mites may exert selective pressures on gland size of their hosts that has resulted in smaller glands among migratory bird species, suggesting that smaller glands may have evolved in these birds to attenuate a possible detrimental effect of feather mites when present in large numbers.

Patterns of nestling provisioning by a single-prey loader bird, Great Tit Parus major

Capsule Nestling provisioning rates depend on nestling age and number, and on time of season, but not on time of day. Aims To determine patterns of nestling provisioning, the effort made by the parents, and the factors which affect them. Methods Mechanical counters to determine food provisioning patterns in 229 Great Tit Parus major nests over 4 years. Results Feeding frequency per chick showed a linear increase with nestling age and total feeding frequency stabilized towards the end of the nestling period. The number of visits per nest increased linearly, while those per nestling decreased linearly with brood‐size. Feeding rates per nest declined throughout the breeding season parallel to the seasonal brood‐size decline; feeding frequencies per nestling did not vary seasonally. Adult effort was almost constant throughout the day, independently of variations in brood‐size, ambient temperature and date. Conclusion Previous studies of nestling provisioning patterns were either contradictory or poorly supported by low sample sizes. The present study involved a much larger sample and clarifies nestling provisioning patterns and the factors with which they are associated.

Laying date and clutch size of Great Tits(Parus major) in the Mediterranean region: a comparison of four habitat types

Laying data and clutch size of Great Tits were studied in four different habitats in eastern Spain: two holm oak(Quercus ilex) forests, at 500 and 900–950 m a.s.l., a zeen oak(Quercus faginea) forest, at 900–1100 m a.s.l., a pine(Pinus sylvestris) forest, at 1000–1050 m a.s.l., and orange(Citrus aurantium) plantations, at 30 m a.s.l. All sites were placed at about the same latitude (39–41°N), and all were studied during the same years (1992–95). Our results show that (1) laying date did not differ between the natural habitats at the same altitude (range of the means of yearly means 4–8 May); (2) within the same habitat type (holm oak forest) laying date was earlier at low altitude (30 Aprilvs. 8 May); (3) laying date was earlier in the orange plantations (21 April) than in natural habitats; (4) among natural habitats at the same altitude, clutch size decreased from zeen oak (mean of yearly means 7.3 eggs) to holm oak (7.0 eggs) to pine forests (6.4 eggs), though only the difference between zeen oak and pine forests was significant; (5) within the same habitat type (holm oak forest), the clutch size tended to be larger at high altitude (7.0vs. 5.9 eggs); and (6) clutch size in orange plantations (7.7 eggs) did not differ significantly from that of the zeen oak forest, but was larger than in the holm oak and pine forests. We discuss the effect of the habitat type on laying date and clutch size of Great Tits. Legedatum und Gelegegröße der Kohlmeise wurden in vier unterschiedlichen Biotopen in Ostspanien untersucht: zwei Steineichenwälder(Quercus ilex) in 500 m und 900–950 mNN, ein Bergeichenwald(Quercus faginea) in 900–1100 mNN, ein Kiefernwald(Pinus sylvestris) in 1000–1050 mNN und eine Orangenpflanzung(Citrus aurantium) in 30 mNN. Alle fünf Gebiete lagen auf etwa demselben Breitengrad (39–41°N) und wurden 1992–1995 parallel untersucht. (1) Auf gleicher Meereshöhe unterscheidet sich der Legebeginn nicht zwischen den verschiedenen Waldbiotopen (im Mittel 4.–8. Mai). (2) Innerhalb desselben Biotoptyps (Steineichenwald) war der Legebeginn auf niedrigerer Meereshöhe früher als in höheren Lagen (30. April vs. 8. Mai). (3) Im Orangenhain wurde früher mit der Eiablage begonnen (21. April) als in den Waldbiotopen. (4) Auf gleicher Meereshöhe nahm die Gelegegröße vom Bergeichenwald (Mittelwert 7.3 Eier) über die Steineichenwälder (7,0 Eier) zum Kiefernwald hin ab (6.4 Eier), jedoch ist nur der Unterschied zwischen Bergeichenwald und Kiefernwald signifikant. (5) Innerhalb der Steineichenwälder besteht die Tendenz zu größerer Gelegegröße in den höheren Lagen (7.0 vs. 5.9 Eier). (6) Im Orangenhain war die Gelegegröße mit durchschnittlich 7.7 Eier ähnlich der im Bergeichenwald, aber größer als in den Steineichen- und Kiefernwälder.

Variation in clutch size in relation to nest size in birds

Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nests of 21 species of hole and non-hole-nesting birds. There was a significant, positive relationship between clutch size and the base area of the nest box or the nest, and this relationship did not differ significantly between open nesting and hole-nesting species. The slope of the relationship showed significant intraspecific and interspecific heterogeneity among four species of secondary hole-nesting species, but also among all 116 slope estimates. The estimated relationship between clutch size and nest box base area in study sites with more than a single size of nest box was not significantly different from the relationship using studies with only a single size of nest box. The slope of the relationship between clutch size and nest base area in different species of birds was significantly negatively related to minimum base area, and less so to maximum base area in a given study. These findings are consistent with the hypothesis that bird species have a general reaction norm reflecting the relationship between nest size and clutch size. Further, they suggest that scientists may influence the clutch size decisions of hole-nesting birds through the provisioning of nest boxes of varying sizes.

Clutch-size variation in Western Palaearctic secondary hole-nesting passerine birds in relation to nest box design

Secondary hole-nesting birds that do not construct nest holes themselves and hence regularly breed in nest boxes constitute important model systems for field studies in many biological disciplines with hundreds of scientists and amateurs involved. Those research groups are spread over wide geographic areas that experience considerable variation in environmental conditions, and researchers provide nest boxes of varying designs that may inadvertently introduce spatial and temporal variation in reproductive parameters. We quantified the relationship between mean clutch size and nest box size and material after controlling for a range of environmental variables in four of the most widely used model species in the Western Palaearctic: great tit Parus major, blue tit Cyanistes caeruleus, pied flycatcher Ficedula hypoleuca and collared flycatcher F.albicollis from 365 populations and 79610 clutches. Nest floor area and nest box material varied non-randomly across latitudes and longitudes, showing that scientists did not adopt a random box design. Clutch size increased with nest floor area in great tits, but not in blue tits and flycatchers. Clutch size of blue tits was larger in wooden than in concrete nest boxes. These findings demonstrate that the size of nest boxes and material used to construct nest boxes can differentially affect clutch size in different species. The findings also suggest that the nest box design may affect not only focal species, but also indirectly other species through the effects of nest box design on productivity and therefore potentially population density and hence interspecific competition.

Fuel management and stopover duration of Blackcaps Sylvia atricapilla stopping over in northern Spain during autumn migration period

Capsule Fuel load is correlated with fuel deposition rate; stopover duration is affected by arrival fuel load. Aims To determine the stopover duration, fuel management and flight ranges at departure of Blackcaps stopping over in northern Spain. Methods Systematic mist-netting and ringing allowed the use of mark–recapture Cormack–Jolly–Seber models for the estimation of stopover duration. Trapped birds were measured and weighed in order to estimate mass gain. FLIGHT software was used to estimate flight ranges. Results Stopover duration ranged from 3.6 to 13.6 days, and was negatively correlated with arrival body mass (assessed by body mass at the first capture event). On average, arrival body mass was 18.4 g, whilst body mass at departure was 19.8 g. No significant differences in arrival body mass and departure body mass were observed between age or sex classes. Mass deposition rate did not differ between age or sex classes (mean = 0.20 g/day). Birds recaptured one day after the first capture event lost mass, whilst recaptures from the second day onwards had a mean gain of mass; mass was observed to increase linearly with the stopover duration. Mass deposition rate was positively correlated with departure body mass. Finally, with a mean departure body mass of 19.8 g, a Blackcap stopping over in northern Spain should be able to fly up to 1100 km. Conclusions Stopover duration assessed by Cormack–Jolly–Seber models was longer than that observed in birds recaptured more than once (‘minimum stopover duration’). Stopover was longer for birds arriving with less fuel. The positive relationship between departure body mass and mass deposition rate suggests a time-minimizing strategy. The lack of difference in fuel deposition rate between age and sex classes suggests a relatively abundant food supply at the study site, but other explanations might also account for the lack of age and sex differences, for example if competition for food was not determined by social hierarchies but by scramble competition. Departing fuel load would allow these birds to arrive at their wintering areas in southern Spain under still-air conditions, without needing to refuel.

Breeding Ecology of the Blue Tit Parus caeruleus in Eastern Spain: A Comparison with Other Localities with Special Reference to Corsica

Aspects of the breeding ecology of the Blue Tit Parus caeruleus were studied in intermediate stages of succession in a holm oak forest at Monte Poblet (eastern Spain) from 1985 to 1991. In different years the density in two plots without nestboxes varied between 1.0 and 6.5 pairs per 10 ha. Nestbox introduction seems to have increased the density of breeding pairs. The mean laying date was earlier (6 vs. 13 May) in low altitude (500-750 m a.s.l.) plots than in a high altitude (900-1000 m a.s.l.) plot, but clutch size (6.9 vs. 6.5 eggs) was not significantly different (mean 6.6 eggs). No differences in mean laying dates were found between years. Significant differences in clutch size were found only between two years at the high altitude site. There was a slight, non-significant, seasonal decline in clutch size. No second clutches were laid. These results are discussed in the context of other Mediterranean studies with special focus on the differences and similarities between Corsican and mainland populations. It is shown that most of the characteristics of the Corsican population (i.e. density, habitat use, laying date, clutch size, number of broods per year) are shared by some populations on the mainland. On the other hand, some breeding traits of Blue Tit populations in southern France are closer to those of Central and North European populations than to the Iberian ones.

Effect of fuel load, date, rain and wind on departure decisions of a migratory passerine

Meteorological conditions, fuel load and date in the season can affect the departure decisions among migratory birds. However, it is poorly understood to what extent the departure decisions are more influenced by some parameters in relation to others, and how they interact with each other. We explored here how fuel load, date, rain and wind (measured on the ground and at high altitude, codified as a tailwind component) influenced the departure decisions of migratory Blackcaps (Sylvia atricapilla) from a stopover site. We used mark–recapture data of 947 Blackcaps collected during the autumn migration period 2005 at a stopover site in northern Iberia, estimating the emigration likelihood with Cormack-Jolly-Seber models, in which we tested for the effect of these four study variables. Best models fitting data showed an additive and positive effect of tailwind and fuel load on the emigration likelihood.ZusammenfassungDie meteorologischen Verhältnisse, der Energievorrat sowie das konkrete Datum können bei Zugvögeln die Entscheidung darüber beeinflussen, wann sie ihren Zug beginnen, bzw. fortsetzen. Dabei ist jedoch nur wenig dazu bekannt, in welchem Ausmaß welcher dieser Parameter relativ zu den anderen die Abflugentscheidung beeinflusst und inwieweit sie untereinander interagieren. In dieser Studie haben wir für ziehende Mönchsgrasmücken (Sylvia atricapilla) untersucht, wie ihr Energievorrat, das jeweilige Datum, Regen und Wind (gemessen am Boden und in größerer Höhe, angegeben als Rückenwindkomponente) die Entscheidung für den Weiterflug von einem Zwischenstop beeinflussten. Hierfür verwendeten wir die Daten von 947 markierten Mönchsgrasmücken, die während des Herbstzuges 2005 in einem Zwischenstop-Gebiet im Norden der iberischen Halbinsel wiedergefangen wurden. Die Weiterzug-Wahrscheinlichkeit wurde anhand von Cormack-Jolly-Seber-Modellen abgeschätzt, mit denen wir die möglichen Auswirkungen der vier untersuchten Variablen testeten. Das den Daten am besten angepasste Modell zeigte einen additiven, positiven Effekt von Rückenwind und Energievorrat auf die Weiterzugswahrscheinlichkeit.