A complete multilocus species phylogeny of the tits and chickadees (Aves: Paridae).

The avian family Paridae (tits and chickadees) contains c. 55 species distributed in the Palearctic, Nearctic, Afrotropics and Indomalaya. The group includes some of the most well-known and extensively studied avian species, and the evolutionary history, in particular the post-glacial colonization of the northern latitudes, has been comparably well-studied for several species. Yet a comprehensive phylogeny of the whole clade is lacking. Here, we present the first complete species phylogeny for the group based on sequence data from two nuclear introns and one mitochondrial gene for 67 taxa of parids. Our results strongly support the inclusion of the Fire-capped Tit (Cephalopyrus flammiceps), currently placed in the Remizidae, as the most basal member of the Paridae. The Yellow-browed Tit (Sylviparus modestus) and the Sultan Tit (Melanochlora sultanea) constitute the next two sequential branches whereas the remaining tits fall into two large clades, one of which contains the seed hoarding and nest excavating species. The indicated clades within these two groups are largely congruent with recent classifications, but with several unforeseen relationships, such as non-monophyly of the Sombre Tit (Poecile lugubris) and the Marsh Tit (Poecile palustris), as well as non-monophyly of both the African gray and the African black tits. Further, our results support a close relationship between the White-fronted Tit (Parus semilarvatus) and the varied Tit (Poecile varius) as well as a close relationship between the White-naped Tit (Parus nuchalis) and the Yellow-cheeked and Black-lored tits (Parus spilonotus and P. xanthogenys). Finally, Hume's Ground-tit (Pseudopodoces humilis) is found to be closely related to the Green-backed Tit (Parus monticolus) and the Great Tit (Parus major). We propose a new classification that is in accordance with this phylogeny.

Identifying causal relationships of cancer treatment and long-term health effects among 5-year survivors of childhood cancer in Southern Sweden.

Background: Survivors of childhood cancer can develop adverse health events later in life. Infrequent occurrences and scarcity of structured information result in analytical and statistical challenges. Alternative statistical approaches are required to investigate the basis of late effects in smaller data sets. Methods: Here we describe sex-specific health care use, mortality and causal associations between primary diagnosis, treatment and outcomes in a small cohort (n = 2315) of 5-year survivors of childhood cancer (n = 2129) in southern Sweden and a control group (n = 11,882; age-, sex- and region-matched from the general population). We developed a constraint-based method for causal inference based on Bayesian estimation of distributions, and used it to investigate health care use and causal associations between diagnoses, treatments and outcomes. Mortality was analyzed by the Kaplan-Meier method. Results: Our results confirm a significantly higher health care usage and premature mortality among childhood cancer survivors as compared to controls. The developed method for causal inference identifies 98 significant associations (p < 0.0001) where most are well known (n = 73; 74.5%). Hitherto undescribed associations are identified (n = 5; 5.1%). These were between use of alkylating agents and eye conditions, topoisomerase inhibitors and viral infections; pituitary surgery and intestinal infections; and cervical cancer and endometritis. We discuss study-related biases (n = 20; 20.4%) and limitations. Conclusions: The findings contribute to a broader understanding of the consequences of cancer treatment. The study shows relevance for small data sets and causal inference, and presents the method as a complement to traditional statistical approaches.

Delayed dispersal: youth costs carry lifetime gains.

An analysis of reproductive success in the green woodhoopoe Phoeniculus purpures challenges the view that delayed dispersal is costly. Females delaying dispersal for longer had more reproductive events in life and higher lifetime production of offspring.

Habitat-specific demography and source-sink dynamics in a population of Siberian jays.

1. There are a number of models describing population structure, many of which have the capacity to incorporate spatial habitat effects. One such model is the source-sink model, that describes a system where some habitats have a natality that is higher than mortality (source) and others have a mortality that exceeds natality (sink). A source can be maintained in the absence of migration, whereas a sink will go extinct. 2. However, the interaction between population dynamics and habitat quality is complex, and concerns have been raised about the validity of published empirical studies addressing source-sink dynamics. In particular, some of these studies fail to provide data on survival, a significant component in disentangling a sink from a low quality source. Moreover, failing to account for a density-dependent increase in mortality, or decrease in fecundity, can result in a territory being falsely assigned as a sink, when in fact, this density-dependent suppression only decreases the population size to a lower level, hence indicating a 'pseudo-sink'. 3. In this study, we investigate a long-term data set for key components of territory-specific demography (mortality and reproduction) and their relationship to habitat characteristics in the territorial, group-living Siberian jay (Perisoreus infaustus). We also assess territory-specific population growth rates (r), to test whether spatial population dynamics are consistent with the ideas of source-sink dynamics. 4. Although average mortality did not differ between sexes, habitat-specific mortality did. Female mortality was higher in older forests, a pattern not observed in males. Male mortality only increased with an increasing amount of open areas. Moreover, reproductive success was higher further away from human settlement, indicating a strong effect of human-associated nest predators. 5. Averaged over all years, 76% of the territories were sources. These territories generally consisted of less open areas, and were located further away from human settlement. 6. The source-sink model provides a tool for modelling demography in distinct habitat patches of different quality, which can aid in identifying key habitats within the landscape, and thus, reduce the risk of implementing unsound management decisions.

Reduced mortality selects for family cohesion in a social species.

Delayed dispersal is the key to family formation in most kin-societies. Previous explanations for the evolution of families have focused on dispersal constraints. Recently, an alternative explanation was proposed, emphasizing the benefits gained through philopatry. Empirical data have confirmed that parents provide their philopatric offspring with preferential treatment through enhanced access to food and predator protection. Yet it remains unclear to what extent such benefits translate into fitness benefits such as reduced mortality, which ultimately can select for the evolution of families. Here, we demonstrate that philopatric Siberian jay (Perisoreus infaustus) offspring have an odds ratio of being killed by predators 62% lower than offspring that dispersed promptly after independence to join groups of unrelated individuals (20.6% versus 33.3% winter mortality). Predation was the sole cause of mortality, killing 20 out of 73 juveniles fitted with radio tags. The higher survival rate among philopatric offspring was associated with parents providing nepotistic predator protection that was withheld from unrelated group members. Natal philopatry usually involves the suppression of personal reproduction. However, a lower mortality of philopatric offspring can overcome this cost and may thus select for the formation of families and set the scene for cooperative kin-societies.

Out of Gondwanaland; the evolutionary history of cooperative breeding and social behaviour among crows, magpies, jays and allies.

Cooperative breeding is comparatively rare among birds in the mainly temperate and boreal Northern Hemisphere. Here we test if the distribution of breeding systems reflects a response to latitude by means of a phylogenetic analysis using correlates with geographical range among the corvids (crows, jays, magpies and allied groups). The corvids trace their ancestry to the predominantly cooperative 'Corvida' branch of oscine passerines from the Australo-Papuan region on the ancient Gondwanaland supercontinent, but we could not confirm the ancestral state of the breeding system within the family, while family cohesion may be ancestral. Initial diversification among pair-breeding taxa that are basal in the corvid phylogeny, represented by genera such as Pyrrhocorax and Dendrocitta, indicates that the corvid family in its current form could have evolved from pair-breeding ancestors only after they had escaped the Australo-Papuan shield. Within the family, cooperative breeding (alloparental care/family cohesion) is strongly correlated to latitude and its predominance in species maintaining a southerly distribution indicates a secondary evolution of cooperative breeding in the lineage leading away from the basal corvids. Multiple transitions show plasticity in the breeding system, indicating a response to latitude rather than evolutionary inertia. The evolutionary background to the loss of cooperative breeding among species with a northerly distribution is complex and differs between species, indicating a response to a variety of selection forces. Family cohesion where the offspring provide alloparental care is a main route to cooperatively breeding groups among corvids. Some corvid species lost only alloparental care, while maintaining coherent family groups. Other species lost family cohesion and, as a corollary, they also lost the behaviour where retained offspring provide alloparental care.

Experimentally increased food resources in the natal territory promote offspring philopatry and helping in cooperatively breeding carrion crows.

Kin-based societies, where families represent the basic social unit, occur in a relatively small number of vertebrate species. In the majority of avian kin societies, families form when offspring prolong their association with the parents on the natal territory. Therefore, the key to understanding the evolution of families in birds is to understand natal philopatry (i.e. the tendency to remain on the natal territory). It has been shown that, within populations, the strength of the association between parents and offspring (i.e. family stability) increases when offspring dispersal is constrained by external environmental factors, but it is unclear whether and how family wealth influences juvenile dispersal decisions. Here, we show that young carrion crows (Corvus corone corone) from territories that were food-supplemented year-round were more philopatric and more likely to help at their family's nest than the unfed ones. The results suggest that offspring philopatry and helping behaviour are influenced by the quality of 'home' and that the availability of food resources positively affects the cohesion of the family.

Predation risk induces changes in nest-site selection and clutch size in the Siberian jay.

Life-history theory predicts that an individual should reduce its reproductive efforts by laying a smaller clutch size when high risk of nest predation reduces the value of current reproduction. Evidence in favour of this 'nest predation hypothesis', however, is scarce and based largely on correlative analyses. Here, we manipulated perceived risk of nest predation in the Siberian jay Perisoreus infaustus using playback involving a mixture of calls by corvid nest predators in the vicinity of nest sites. In response to being exposed to this acoustic cue simulating increased risk of nest predation, the jays chose a nest site offering more protective covering and reduced clutch size. This is the first experimental demonstration of clutch size adjustment and nest site selection as a result of phenotypic plasticity in an open nesting passerine reflecting a facultative response to the perceived risk of nest predation.

Delayed dispersal as a route to breeding: territorial inheritance, safe havens, and ecological constraints.

The relative roles of ecological constraints, the benefits of philopatry, and the role of life history continue to be debated in the evolution of natal philopatry and cooperative breeding. We compare three routes to breeding: departing to search for territories as a floater, staying and queuing to inherit the natal territory, or queuing and eventually shifting to a neighboring vacancy. Our model assumed a dominance-structured population. It quantifies the benefits of philopatry for varying-rank subordinates and contrasts it against the benefit of dispersal. We apply the model to data on Siberian jay Perisoreus infaustus, a species in which retained offspring do not help at the nest. The results indicate that territorial inheritance plays a small role in this species (presumably due to inbreeding avoidance), and territory acquisition is less constrained for dispersing than philopatric offspring. Nevertheless, small family groups-one or, at the most, two same-sex queuers-are predicted to form because philopatric offspring gain nepotistic benefits that improve their survival. This fits with data on group sizes and supports the idea of the natal territory as a safe haven for waiting for breeding opportunities. We also discuss our predictions in the light of ecological constraints and clarify recent confusingly different predictions on the role of habitat saturation as an explanation for delayed dispersal and cooperative breeding. We argue that "ecological constraint" is too wide a term to yield useful predictive power and that it is more appropriate to examine the consequences of specific life-history traits on the success of dispersers.

Why offspring delay dispersal: experimental evidence for a role of parental tolerance.

Approximately 3% of all bird species live in families based on a prolonged parent-offspring association. Formation of family groups often appears to be associated with various constraints on the possibilities of independent reproduction for the offspring. However, delayed dispersal is not the only alternative for offspring that defer reproduction. To account for the formation of a family group it is also necessary to explain why non-dispersing offspring forego the alternative options to join other groups as 'extra birds' or to become 'floaters' and roam between territories. We removed fathers from Siberian jay Perisoreus infaustus family groups to test the proposal that nepotistic parental tolerance gives the natal territory a special value for the offspring, which they cannot find elsewhere and that makes them stay. In this species, parents are more tolerant of their retained offspring than towards immigrant extra birds. In response to the removal of fathers, retained offspring dispersed, whereas there was no similar response among the unrelated extra birds. Retained offspring, however, left only after despotic immigrant alpha-males replaced removed fathers, indicating that the presence of fathers is an essential motive for offspring to delay their dispersal. By blocking immigrant and unrelated males from becoming alpha-males and by being tolerant themselves, fathers provide a 'safe haven' in the natal territory where retained offspring can avail themselves of available resources without any, or only mild, competitive interference.

History, environment and social behaviour: experimentally induced cooperative breeding in the carrion crow.

Kin-based cooperative breeding, where grown offspring delay natal dispersal and help their parents to rear new young, has a long history in some avian lineages. Family formation and helping behaviour in extant populations may therefore simply represent the retention of ancestral features, tolerated under current conditions, rather than a current adaptive process driven by environmental factors. Separating these two possibilities challenges evolutionary biologists because of the tight coupling that normally exists between phylogeny and the environmental distribution of species and populations. The carrion crow Corvus corone corone, which exhibits extreme interpopulational variation in the extent of cooperative breeding, with populations showing no delayed dispersal and helping at all, provides a unique opportunity for an experimental approach. Here we show that offspring of non-cooperative carrion crows from Switzerland will remain on the natal territory and express helping behaviour when raised in a cooperative population in Spain. When we transferred carrion crow eggs from Switzerland to Spain, five out of six transplanted juveniles delayed dispersal, and two of those became helpers in the following breeding season. Our results provide compelling experimental evidence of the causal relationship between current environmental conditions and expression of cooperative behaviour.

Impact of forestry practices on fitness correlates and population productivity in an open-nesting bird species.

In the boreal forests of Fennoscandia, over 99% of forest area has been altered by forestry practices, which has created forest with age structures and stand characteristics that differ from primary forest stands. Although many researchers have investigated how forestry affects species abundance, few have assessed how forestry practices affect fitness correlates of species living in altered habitats, and this has negatively affected management efforts. We experimentally addressed the effect of standard forestry practices on fitness correlates of an open-nesting, long-lived bird species typical to boreal forests of Eurasia, the Siberian Jay (Perisoreus infaustus). We used a before-after comparison of reproductive data on the level of territories and found that standard forestry practices had a strong negative effect on the breeding success of jays. Both partial thinning of territories and partial clearcutting of territories reduced future breeding success by a factor of 0.35. Forestry practices reduced territory occupancy. Thus, over the 15 years of the study, productivity of the affected population declined over 50% as a result of territory abandonment and reduced breeding success. Results of previous studies on Siberian Jays suggest that the strong effect of forest thinning on fitness is explained by the fact that most common predators of nests and adults are visually oriented and thus thinning makes prey and nests more visible to predators. The consequences of thinning we observed are likely to apply to a wide range of species that rely on understory to provide visual protection from predators. Thus, our results are important for the development of effective conservation management protocols and for the refinement of thinning practices.

REPRODUCTIVE COST, AGE-SPECIFIC SURVIVAL AND A COMPARISON OF THE REPRODUCTIVE STRATEGY IN TWO EUROPEAN TITS (GENUS PARUS).

Theoretical analyses of optimal reproductive rates usually assume a trade-off between offspring production and parental survival. This study verified a survival cost for willow tit males; nonbreeding males survived better than males attending a brood. Theory also predicts a smaller clutch size in birds that are less successful in transforming reproductive investments into mature offspring. As predicted, we found that crested tits, suffering a higher nest predation rate, laid smaller clutches than willow tits. The generally lower survival rate of willow tit adults may largely be attributed to their higher reproductive commitment (larger willow tit clutch size), because no significant interspecific survival difference remained between nonbreeding males. Finally, in willow tits we found a positive correlation between average clutch size and juvenile survival rate (density-dependent) the ensuing year, suggesting that willow tits may adjust clutch size in response to changing survival prospects for their young by using the breeding density as a cue.

Kin selection in cooperative alliances of carrion crows.

In most cooperative vertebrates, delayed natal dispersal is the mechanism that leads to the formation of kin societies. Under this condition, the possibility that kin-based cooperative breeding is an unselected consequence of dispersal patterns can never be ruled out because helpers can only help their relatives. Here we show that a population of carrion crows (Corvus corone corone) fully fits the central prediction of kin selection theory that cooperative breeding should arise among relatives. On their territory, resident breeders are aided not only by nonbreeding retained offspring but also by immigrants (mainly males), with whom they share matings. Philopatry cannot account, however, for the high degree of genetic relatedness found between breeders and immigrants of the same sex that cooperate at a nest, indicating that crows actively choose to breed cooperatively with their relatives.