Animals, including siblings, compete for resources such as food, territory, and potential mating partners. In animal sibling rivalry, individuals compete for parental care or limited resources, which can sometimes result in siblicide, the killing of siblings. [1] Sibling rivalry occurs in many different forms. Siblings may compete for resources in a prenatal and/or post-birth environment. The degree of rivalry varies, ranging from a low level of violence in non-aggressive to the killing of kin in siblicide.
When there are multiple offspring in a single brood, the potential for sibling rivalry arises due to competition for food and parental attention. Natural selection may favor behaviors that allow an individual offspring to gain more resources, even if the behavior decreases a sibling's fitness. Competition for food and resources can be seen in many bird species. For example, blue-footed booby (Sula nebouxii) siblings often exhibit aggression towards each other, with older chicks pecking at younger chicks. This behavior increases when there are food shortages, indicating more intense competition. [2] In other bird species, siblings compete for food through manipulation of parental behavior rather than direct aggressive acts. Increased parental attention may mean more food for the offspring, favoring the development of begging behavior in nestlings. American robin (Turdus migratorius) chicks compete for food provided by their parents through louder and more prominent cheeps or other vocalizations, with the most food given to chicks exhibiting the most intense begging behavior. [3]
Sibling rivalry may not seem to align with the kin selection theory, which predicts that altruistic behaviors may evolve if inclusive fitness benefits (including those of relatives) from such behaviors outweigh the costs. [4] Theoretically, helping relatives would allow individuals to spread genes related to their own. However, some species may show sibling rivalry when the fitness costs outweigh the benefits of helping relatives. Sibling relatedness can influence degree of rivalry. Canary nestlings are more selfish and competitive if other nestlings are less related. [5] When offspring beg for more food from their parents, they also are “competing” with their future siblings by decreasing the fitness of parents, reducing their ability to invest in future offspring. This is known as interbrood rivalry, which can lead to parent–offspring conflict.
Siblicide is a consequence of sibling rivalry, and its occurrence may be due to a variety of factors, such as food shortages and limited parental care. Killing a sibling could be advantageous for an animal because it monopolizes more resources through the elimination of a competitor. Moreover, there are different types of siblicide. For example, obligate siblicide is the unconditional killing of a sibling. In species of birds that exhibit this behavior, the larger chick commits the act of siblicide. On the other hand, facultative siblicide refers to situations in which the death of a sibling does not always occur, but is usually motivated by environmental factors such as limited resources. [6] Facultative siblicide is exhibited by the previously discussed case of the blue-footed booby (Sula nebouxii). In this species, senior chicks may sometimes eliminate siblings when there are food shortages. During these shortages, chicks exhibit higher levels of pecking, but this aggression decreases when food levels are brought back to sufficient levels. [7] In some species, parents have evolved behaviors to take advantage of siblicide to increase their own fitness. For example, the laughing gull exhibits asynchronous hatching patterns in order to cut parental losses. In this species, birds that lay their eggs at different times produce, on average, a larger number of fledglings per nest in comparison to birds with synchronous hatches. [8] Staggered hatching creates chicks in different stages of growth. Older, and thus larger, chicks kill their younger siblings, reducing brood size and allowing the parent to concentrate efforts when food is scarce. Once the brood is reduced, sibling rivalry decreases because there are fewer competitors, benefitting the surviving offspring. This also benefits the parents by minimizing unproductive parental investment in offspring that are unlikely to be successful. This is especially advantageous during food shortages when parents are unable to adequately feed all offspring.
Siblicide has also been noted in mammalian species. For instance, spotted hyenas (Crocuta crocuta) have been known to exhibit facultative siblicidal behavior. Intense sibling aggression begins only a few minutes after birth and then continues for a few days. [9] The function of sibling aggression is to establish and maintain rank relationship between litter mates, but this aggression does not always lead to sibling death. [10] However, during periods of intense feeding competition, aggression can escalate to siblicide. [11] Spotted hyena aggression demonstrates how facultative siblicide can be triggered by environmental factors.
Intrauterine cannibalism (see also: cannibalism) occurs when siblings eat each other in the embryonic phase. This can take the form of embryophagy when siblings eat embryos, and oophagy, when siblings eat eggs. [12] Intrauterine cannibalism can benefit embryos by providing increased nutrition. Fire salamander (Salamandra salamandra) populations exhibiting intrauterine cannibalism have embryos that develop more quickly into larvae due to nutritional supplementation from feeding on siblings. [13] Lamnoid shark embryos have adaptations to facilitate intrauterine cannibalism in the form of precocious teeth, which they use to feed on intrauterine eggs and siblings after their yolk supply is used up. [14] In sand tiger sharks (Carcharias taurus), the first embryo to reach a certain size, referred to as "the hatchling," will always consume the smaller, less-developed siblings in the womb. [15] Because the first embryo may have a different father from the eaten embryos, this form of siblicide early during development may be indirectly involved in male competition. This phenomenon of embryonic cannibalism may play a role in sexual selection, as males compete post-fertilization for paternity. [16] Thus, intrauterine cannibalism in sharks may reflect not only sibling rivalry, but also male competition for successful mating with females, which is an example of sexual selection.
Not all forms of sibling rivalry in animals involve direct aggression or death of a sibling. This is not an extremely aggressive form of rivalry; however, it still results in reduced sibling fitness.
Fetal programming (see also: Barker's Hypothesis) refers to the persisting effects in adult life caused by the fetal environment. In sheep, competition for resources within the uterus may lead to impaired reproductive abilities and different body composition. Sheep that were born 600 grams lighter than their twins have an impaired reproductive ability - the lighter the sheep weighed in comparison to their twin, the more impaired. Although the sheep are not competing physically like in intrauterine cannibalism, the disparity in birth weight suggests an overall fitness difference. [17]
Is the teat that he fattens his flesh on.
He fights for his teat with tenacity
Against any sibling's audacity.
The piglet, to arm for this mission,
Is born with a warlike dentition
Of eight tiny tusks, sharp as sabres,
Which help in impressing the neighbors;
But to render these weapons less harrowing,
Most farmers remove them at farrowing.
We studied pig sisters and brothers
When some had their teeth, but not others.
We found that when siblings aren't many,
The weapons help little if any,
But when there are many per litter,
The teeth help their owners grow fitter.
But how did selection begin
To make weapons to use against kin? [18]
(Abstract from the paper “Armed Sibling Rivalry
among Suckling Piglets” by Fraser and Thompson)
Domestic piglets (Sus scrofa) have been shown to exhibit different forms of non-lethal competition such as uterine competition. The relative development of the embryo in the uterus can affect their chance of survival. Pig embryos follow different developmental paths because during estrus, sows will ovulate the majority of their follicles during a short period of time and then a few during a longer period of time. [19] This pattern causes a difference in development, so the less developed embryos are less likely to survive. Competition also exists over space in the uterus of the sow. The central portion of the uterus is the most crowded and the site of the greatest competition. This competition prevents some embryos from fully growing, often resulting in a low birthweight that may put piglets at a disadvantage once out of the womb. [20] Neonatal competition also exists among piglets as they directly compete against their siblings for their mother's teats only hours after their birth. [21] Competition is responsible for 43% of piglet neonatal death due to starvation. Under normal conditions (i.e. stable environment, average litter size), larger piglets appear to have an advantage in survival partially due to their ability to win more fights against smaller piglets over access to teats. [22] It is believed that teeth in piglets evolved as a product of an evolutionary arms race caused by sibling competition, resulting in armed sibling rivalry. Teeth become more important when the litter size is larger than normal, causing increased competition. In these situations, the teeth can help the individual piglet compete against siblings. [23]
Sibling rivalry can be mediated and/or encouraged by parents, especially in bird species. Sibling rivalry can also have a negative impact on the parent and the fitness of future offspring due to interbrood conflict which may lead to parent–offspring conflict. This conflict may force parents to exert extra energy at the expense of future broods.
Parents may play a passive role in encouraging sibling rivalry. The most common case is in food distribution. When parent birds distribute food, they do not favor any individual offspring. Great tits (Parus major) preferentially distribute food from similar locations, so offspring compete for those prime spots closest to the feeding site where they will get more food, leading to unequal growth of offspring. This behavior exemplified in the young can be interpreted as an optimal foraging strategy driven by scramble competition. [24]
Alternatively, parents may take an active role in mediating the intensity of rivalry between siblings. Again, great tit parents can choose what location to distribute food to as a means of controlling competition between nestlings. When parents feed chicks at one or two feeding locations in the nest, the distance between the male and female parents influences the level of competition between nestlings. In other words, parents may decrease sibling competition by varying the locations within the nest that receive food.
Interbrood conflict occurs when the current brood demands more at the expense of future broods. This sibling rivalry can lead to parent–offspring conflict, in which there are different optimal levels of parental investments whether viewed from the parental or offspring perspective. Biological signalling theory suggests that young can communicate with parents to maximize the amount of food they can get. Therefore, young that need or want more food may solicit or beg at higher levels. Parents can respond to this by providing more food, but this is based on the assumption of honest begging, in which chicks beg only when they actually need more food. However, if all the young start soliciting at higher levels, this incurs a cost on the parents because they will need to expend more energy searching for food. The increased travel to and from the nest may attract predators as well. These factors reduce the energy and resources parents have for future offspring. This is an example of where nestling intrabrood competition can influence the parents' investment in future broods. [25]
The black-headed gull is a small gull that breeds in much of the Palearctic including Europe and also in coastal eastern Canada. Most of the population is migratory and winters further south, but some birds reside in the milder westernmost areas of Europe. The species also occurs in smaller numbers in northeastern North America, where it was formerly known as the common black-headed gull.
Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when studying animal behaviors: What are the proximate causes, ontogeny, survival value, and phylogeny of a behavior?
The tree swallow is a migratory bird of the family Hirundinidae. Found in the Americas, the tree swallow was first described in 1807 by French ornithologist Louis Vieillot as Hirundo bicolor. It has since been moved to its current genus, Tachycineta, within which its phylogenetic placement is debated. The tree swallow has glossy blue-green upperparts, with the exception of the blackish wings and tail, and white underparts. The bill is black, the eyes dark brown, and the legs and feet pale brown. The female is generally duller than the male, and the first-year female has mostly brown upperparts, with some blue feathers. Juveniles have brown upperparts, and grey-brown-washed breasts. The tree swallow breeds in the US and Canada. It winters along southern US coasts south, along the Gulf Coast, to Panama and the northwestern coast of South America, and in the West Indies.
A supernormal stimulus or superstimulus is an exaggerated version of a stimulus to which there is an existing response tendency, or any stimulus that elicits a response more strongly than the stimulus for which it evolved.
Helpers at the nest is a term used in behavioural ecology and evolutionary biology to describe a social structure in which juveniles and sexually mature adolescents of either one or both sexes remain in association with their parents and help them raise subsequent broods or litters, instead of dispersing and beginning to reproduce themselves. This phenomenon was first studied in birds where it occurs most frequently, but it is also known in animals from many different groups including mammals and insects. It is a simple form of co-operative breeding. The effects of helpers usually amount to a net benefit, however, benefits are not uniformly distributed by all helpers nor across all species that exhibit this behaviour. There are multiple proposed explanations for the behaviour, but its variability and broad taxonomic occurrences result in simultaneously plausible theories.
The blue-footed booby is a marine bird native to subtropical and tropical regions of the eastern Pacific Ocean. It is one of six species of the genus Sula – known as boobies. It is easily recognizable by its distinctive bright blue feet, which is a sexually selected trait and a product of their diet. Males display their feet in an elaborate mating ritual by lifting them up and down while strutting before the female. The female is slightly larger than the male and can measure up to 90 cm (35 in) long with a wingspan up to 1.5 m (5 ft).
Parent–offspring conflict (POC) is an expression coined in 1974 by Robert Trivers. It is used to describe the evolutionary conflict arising from differences in optimal parental investment (PI) in an offspring from the standpoint of the parent and the offspring. PI is any investment by the parent in an individual offspring that decreases the parent's ability to invest in other offspring, while the selected offspring's chance of surviving increases.
Alloparenting is a term used to classify any form of parental care provided by an individual towards young that are not its own direct offspring. These are often referred to as "non-descendant" young, even though grandchildren can be among them. Among humans, alloparenting is often performed by a child's grandparents and older siblings. Individuals providing this care are referred to using the neutral term of alloparent.
A trophic egg is an egg whose function is not reproduction but nutrition; in essence, the trophic egg serves as food for offspring hatched from viable eggs. In most species that produce them, a trophic egg is usually an unfertilised egg. The production of trophic eggs has been observed in a highly diverse range of species, including fish, amphibians, spiders and insects. The function is not limited to any particular level of parental care, but occurs in some sub-social species of insects, the spider A. ferox, and a few other species like the frogs Leptodactylus fallax and Oophaga, and the catfish Bagrus meridionalis.
Siblicide is the killing of an infant individual by its close relatives. It may occur directly between siblings or be mediated by the parents, and is driven by the direct fitness benefits to the perpetrator and sometimes its parents. Siblicide has mainly, but not only, been observed in birds.
Cross-fostering is a technique used in animal husbandry, animal science, genetic and nature versus nurture studies, and conservation, whereby offspring are removed from their biological parents at birth and raised by surrogates, typically of a different species, hence 'cross.' This can also occasionally occur in nature.
Cannibalism is the act of consuming another individual of the same species as food. Cannibalism is a common ecological interaction in the animal kingdom and has been recorded in more than 1,500 species. Human cannibalism is also well documented, both in ancient and in recent times.
The scissortail sergeant or striptailed damselfish is a large damselfish. It earns its name from the black-striped tail and sides, which are reminiscent of the insignia of a military Sergeant, being similar to those of the sergeant major damselfish. It grows to a length of about 16 centimetres (6.3 in).
In animals, infanticide involves the intentional killing of young offspring by a mature animal of the same species. Animal infanticide is studied in zoology, specifically in the field of ethology. Ovicide is the analogous destruction of eggs. The practice has been observed in many species throughout the animal kingdom, especially primates but including microscopic rotifers, insects, fish, amphibians, birds and mammals. Infanticide can be practiced by both males and females.
Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal kingdom. There is great variation in different animal groups in terms of how parents care for offspring, and the amount of resources invested by parents. For example, there may be considerable variation in the amount of care invested by each sex, where females may invest more in some species, males invest more in others, or investment may be shared equally. Numerous hypotheses have been proposed to describe this variation and patterns in parental care that exist between the sexes, as well as among species.
Filial cannibalism occurs when an adult individual of a species consumes all or part of the young of its own species or immediate offspring. Filial cannibalism occurs in many species ranging from mammals to insects, and is especially prevalent in various types of fish species with males that engage in egg guardianship. The exact evolutionary purpose of the practice in those species is unclear and debated among zoologists, though there is consensus that it may have, or may have had at some point in species' evolutionary history, certain evolutionary and ecological implications.
Begging in animals is when an animal solicits being given resources by another animal. This is usually a young animal soliciting food from their parents, brood hosts or other adults. However, the resource is sometimes non-food related or may be solicited by adult animals. Begging behavior is most widely studied in birds, however, mammals, amphibians, and invertebrates perform begging displays. Generally in food solicitation, begging behavior is instinctive, although in some instances it is learned.
Matriphagy is the consumption of the mother by her offspring. The behavior generally takes place within the first few weeks of life and has been documented in some species of insects, nematode worms, pseudoscorpions, and other arachnids as well as in caecilian amphibians.
Allofeeding is a type of food sharing behaviour observed in cooperatively breeding species of birds. Allofeeding refers to a parent, sibling or unrelated adult bird feeding altricial hatchlings, which are dependent on parental care for their survival. Allofeeding also refers to food sharing between adults of the same species. Allofeeding can occur between mates during mating rituals, courtship, egg laying or incubation, between peers of the same species, or as a form of parental care.
Brood reduction occurs when the number of nestlings in a birds brood is reduced, usually because there is a limited amount of resources available. It can occur directly via infanticide, or indirectly via competition over resources between siblings. Avian parents often produce more offspring than they can care for, resulting in the death of some of the nestlings. Brood reduction was originally described by David Lack in his brood-reduction hypothesis to explain the existence of hatching asynchrony in many bird species.