Michael Griesser

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.

Life history and the evolution of family living in birds

The reason why some bird species live in family groups is an important question of evolutionary biology that remains unanswered. Families arise when young delay the onset of independent reproduction and remain with their parents beyond independence. Explanations for why individuals forgo independent reproduction have hitherto focused on dispersal constraints, such as the absence of high-quality breeding openings. However, while constraints successfully explain within-population dispersal decisions, they fail as an ultimate explanation for variation in family formation across species. Most family-living species are long-lived and recent life-history studies demonstrated that a delayed onset of reproduction can be adaptive in long-lived species. Hence, delayed dispersal and reproduction might be an adaptive life-history decision rather than ‘the best of a bad job’. Here, we attempt to provide a predictive framework for the evolution of families by integrating life-history theory into family formation theory. We suggest that longevity favours a delayed onset of reproduction and gives parents the opportunity of a prolonged investment in offspring, an option which is not available for short-lived species. Yet, parents should only prolong their investment in offspring if this increases offspring survival and outweighs the fitness cost that parents incur, which is only possible under ecological conditions, such as a predictable access to resources. We therefore propose that both life-history and ecological factors play a role in determining the evolution of family living across species, yet we suggest different mechanisms than those proposed by previous models.

Referential Calls Signal Predator Behavior in a Group-Living Bird Species

Predation is a powerful agent of natural selection, driving the evolution of antipredator calls [1]. These calls have been shown to communicate predator category [2-4] and/or predator distance to conspecifics [5-7]. However, the risk posed by predators depends also on predator behavior [8], and the ability of prey to communicate predator behavior to conspecifics would be a selective advantage reducing their predation risk. I tested this idea in Siberian jays (Perisoreus infaustus), a group-living bird species. Predation by hawks, and to a lesser extent by owls, is substantial and the sole cause of mortality in adult jays [9]. By using field data and predator-exposure experiments, I show here that jays used antipredator calls for hawks depending on predator behavior. A playback experiment demonstrated that these prey-to-prey calls were specific to hawk behavior (perch, prey search, attack) and elicited distinct, situation-specific escape responses. This is the first study to demonstrate that prey signals convey information about predator behavior to conspecifics. Given that antipredator calls in jays aim at protecting kin group members [10, 11], consequently lowering their mortality [9], kin-selected benefits could be an important factor for the evolution of predator-behavior-specific antipredator calls in such systems.

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.

Fighting to stay: the role of sibling rivalry for delayed dispersal

In many bird species with delayed dispersal, siblings differ in how long they postpone independence. Some offspring remain with their parents for a year or more and generally forego personal reproduction, while other siblings disperse in the first summer of life. We studied the basis for this variation by following dispersal among newly fledged Siberian jays, Perisoreus infaustus, carrying radiotransmitters. Postponed dispersal was the preferred option. Sibling rivalry preceded dispersal, and the larger and socially dominant siblings within broods were more likely to stay. No sex effect was found: males and females were just as likely to delay dispersal. This role of sibling rivalry shows that models explaining delayed dispersal based on ecological constraints outside the natal territory are too simplistic. The process of within-brood competition in determining which individuals disperse and which ones delay dispersal indicates that there are benefits to be gained from remaining in the natal territory.

Nepotistic alarm calling in the Siberian jay, Perisoreus infaustus

From a life history perspective, parents have an incentive to protect their reproductive investment, and so may provide care even after their offspring are independent. Such prolonged parental care could lead to postponed dispersal of the offspring and thereby facilitate the formation of kin groups. We tested whether alpha birds in Siberian jays protected their independent, retained offspring by giving alarm calls during simulated predator attacks. We compared the responses to predator attacks simulated by flying a hawk model over a dyad of birds on a feeder for dyads composed of an alpha bird and either a relative or a nonrelative. Alpha females were nepotistic in their alarm-calling behaviour, in that they called more frequently when accompanied by their retained offspring than by unrelated immigrants, but alpha males called indiscriminately. This difference in alarm calling could reflect dominance relationships in Siberian jay groups, because the presence of immigrants may be less costly to alpha males, but alpha females are more vulnerable to competition from immigrants. Alarm calls were usually given during escape, when both individuals in the dyad had left the feeding site. However, results of a playback experiment suggest that alarm calls conveyed information about danger and incited an immediate escape reaction. Our results indicate that alarm calling can be nepotistic, and that factors other than kinship influence alarm-calling behaviour. Nepotistic antipredator behaviours are benefits that offspring can gain only in their natal territory. Hence, in the absence of preferential treatment by their parents, offspring may be more likely to disperse and kin groups are prevented from forming.

Nepotistic mobbing behaviour in the Siberian jay (Perisoreus infaustus)

Antipredator theory has rarely been invoked to explain why birds associate in kin groups. Instead, it has been argued that offspring delay dispersal in response to poor dispersal opportunities and changes in the social environment as a consequence of dispersal decisions have been overlooked. To test whether parents provide their offspring with nepotistic antipredator protection, we conducted a field experiment to investigate predator mobbing in the Siberian jay. We used models of a mounted hawk and owl, which represent the two main predators as confirmed by recoveries of killed radiotagged individuals. Kin groups mobbed the predator models for longer and uttered both more calls in total and more predator-specific calls than nonkin groups. However, not all categories of individual called equally. In particular, alpha males gave most calls, especially in the presence of their retained offspring. Unrelated immigrants behaved least cautiously and approached the models closer and for longer than any other individual category. Furthermore, jays mobbed the hawk model, their most dangerous predator, for longer and took fewer risks than they did when mobbing the owl model. Our experiments show that effort and risk taking while mobbing in the Siberian jay not only depend on the danger posed by the predator, but are also influenced by the social environment. Parents invest more in the presence of their offspring. We suggested that such nepotistic antipredator behaviour gives offspring an incentive to delay dispersal and that nepotism could therefore have a pivotal role for the evolution of kin group living in animals.

Mobbing calls signal predator category in a kin group-living bird species

Many prey species gather together to approach and harass their predators despite the associated risks. While mobbing, prey usually utter calls and previous experiments have demonstrated that mobbing calls can convey information about risk to conspecifics. However, the risk posed by predators also differs between predator categories. The ability to communicate predator category would be adaptive because it would allow other mobbers to adjust their risk taking. I tested this idea in Siberian jays Perisoreus infaustus, a group-living bird species, by exposing jay groups to mounts of three hawk and three owl species of varying risks. Groups immediately approached to mob the mount and uttered up to 14 different call types. Jays gave more calls when mobbing a more dangerous predator and when in the presence of kin. Five call types were predator-category-specific and jays uttered two hawk-specific and three owl-specific call types. Thus, this is one of the first studies to demonstrate that mobbing calls can simultaneously encode information about both predator category and the risk posed by a predator. Since antipredator calls of Siberian jays are known to specifically aim at reducing the risk to relatives, kin-based sociality could be an important factor in facilitating the evolution of predator-category-specific mobbing calls.

Archiving primary data: solutions for long-term studies

The recent trend for journals to require open access to primary data included in publications has been embraced by many biologists, but has caused apprehension amongst researchers engaged in long-term ecological and evolutionary studies. A worldwide survey of 73 principal investigators (Pls) with long-term studies revealed positive attitudes towards sharing data with the agreement or involvement of the PI, and 93% of PIs have historically shared data. Only 8% were in favor of uncontrolled, open access to primary data while 63% expressed serious concern. We present here their viewpoint on an issue that can have non-trivial scientific consequences. We discuss potential costs of public data archiving and provide possible solutions to meet the needs of journals and researchers.

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 bird’ or to become‘floater’ 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 α–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 α–males and by being tolerant themselves, fathers provide a‘safe have’ in the natal territory where retained offspring can avail themselves of available resources without any, or only mild, competitive interference.