Parental care

Paternal care Care of offspring by males may evolve when natural selection favouring parental care is stronger than sexual selection against paternal care. In approximately 1% of bird species, males exclusively provide care after eggs are laid. The Maternal Relief hypothesis proposes that males provide care to reduce the burdens associated with reproduction for the female, which ultimately generates shorter inter-birth intervals and produces more successful offspring. In polygynous species, where a single male mates with more than one female, the male's role as a caregiver therefore tends to be reduced. Conversely, males may be exclusively responsible for caring for their offspring in polyandrous species, where a single female mates with more than one male. One hypothesis regarding the evolution of male parental care in non-monogamous species suggests that parental behaviour is correlated with increased siring of offspring. Females of some species of reptiles may remain with their clutch to provide care, by curling around their eggs for the duration of the incubation period. The most intricate example of maternal care in this group can be seen in crocodilian species, as mothers may stay with their young for multiple months. The general mammalian tendency for female parents to invest more in offspring was focused on in the development of early hypotheses to describe sex differences in paternal care. It was initially suggested that different levels of investment by each sex in terms of gamete size and number may have led to the evolution of female-only care. This early hypothesis suggested that because females invest more in the production of fewer and larger gametes, compared with males who produce many, smaller gametes, maternal care would be favoured. This is because females have initially invested more, and would thus stand more to lose if they did not continue to invest in the offspring. Biparental care Biparental care tends to be favoured when sexual selection is not intense, and when the adult sex ratio of males to females is not strongly skewed. For two parents to cooperate in caring for young, the mates must be coordinated with each other as well as with the requirements of the developing young, and the demands of the environment. The selection of biparental care as a behavioural strategy is considered to be an important factor driving the evolution of monogamy, if the value of exclusive cooperation in care for mutual offspring by two parents outweighs the potential benefits of polygamy for either sex. Biparental care may increase offspring survival as well as allow parents to gain further mating opportunities with the pair mate. There is conflicting evidence for whether offspring fare equally, better or worse when receiving care by two parents rather a single parent. On one hand, it has been suggested that due to sexual conflict, parents should withhold the amount of care they provide and shift as much of the workload as possible to their partner. In this case, offspring may be worse off. Other experimental evidence contrasts this, and suggests that when both parents care for their mutual offspring, their individual contributions may have synergistic effects on the fitness of their young. In this case, offspring would benefit from biparental care. Biparental care is particularly prevalent in mammals and birds. Alloparental care Alloparental care, caring for non-descendant offspring, is a seemingly altruistic and reproductively costly behaviour; it has both adaptive benefits and evident costs. It has been observed in over 120 mammal and 150 bird species. It is a defining feature of eusociality, which is found in insects, including various ants, bees, and termites. For mammalian mothers, alloparenting may be beneficial in promoting earlier weaning of infants (as long as earlier weaning does not compromise infant survival). This strategy results in shorter inter-birth intervals and increased reproductive success. Frequent alloparenting may provide mothers more opportunities to feed without their young, which may ultimately increase their net energy gains and permits them to invest more energy in milk synthesis. However, potential costs of alloparenting may include the expenditure of time and resources in caring for non-descendant offspring with no apparent direct benefits to alloparents. The offspring that experience alloparental care may benefit from increased protection from predators and the learning of group dynamics through social interactions. In the eusocial insects, the evolution of a caste system has driven workers to sacrifice their own personal reproductive fitness to assist in the reproductive success of the colony. Indirect fitness benefits are gained instead through assisting related members of the colony. Similarly, worker ants tend to raise their sisters rather than their daughters, due to their greater relatedness. The survival of the colony is believed to be the main reward that drives the altruism of the workers. ==In groups of animals==

Invertebrates building mud nest for her offspring. Each nest is provisioned with food caught by the mother; one or more eggs are laid inside, and the nest is then sealed. Parental care is not frequently observed in invertebrate species. In Dipterans, oviposition is instead commonly observed. Adults lay their eggs before leaving them to hatch and develop into larva, then pupa, then adults. For example, Phormia regina adults lay their eggs preferentially on carrion and corpses. Though biparental and male-only care are rarely observed, female-only care does exist in some invertebrates. Some insects, including the Hymenoptera (ants, bees and wasps), invest substantial effort in caring for their young. The type and amount of care invested varies widely. Solitary wasps such as the potter wasps (Eumeninae) build nests for their young, provisioning them with food, often caterpillars, caught by the mother. The nests are then sealed, and the young live on the food until they leave the nest as adults. In contrast, social wasps and honeybees raise young in substantial colonies, with eggs laid mainly by queens (mothers), and the young cared for mainly by workers (sisters of the young). '' with eggs on his back Outside the Hymenoptera, parental care is found among the burying beetles and the magnificent salt beetle. Many species of Hemiptera take care of their young, for instance in the Belostomatidae genus Abedus. Among arachnids, several groups exhibit parental care. Wolf spiders are known for carrying their young on their abdomens for several weeks after they hatch. Nursery web spiders and some jumping spiders guard their young in silk nests after they hatch. The jumping spider species Toxeus magnus is notable for nursing its young through a form of lactation. Some crustaceans also show parental care. Mothers of the crab species Metopaulias depressus raises their young in water-filled bromeliads, cleaning them of debris, defending them against predators and feeding them with captured prey. In the desert isopod Hemilepistus reaumuri, juveniles share their parents' burrow for the first 10–20 days of their life, and are supplied with food by their parents. Finally, some species of Synalpheus shrimps are eusocial, living in colonies with one or a few breeders of each sex together with non-breeders that defend the colony. Fish seahorse Several groups of fish have evolved parental care. The ratio of fish genera that exhibit male-only: biparental: female-only care is 9:3:1. Some fish such as pipefish, sea dragons and seahorses (Syngnathidae) have a form of male pregnancy, where the female takes no part in caring for the young once she has laid her eggs. Males in other species may take a role in guarding the eggs before they hatch. Mouthbrooding is the care given by some groups of fish (and a few other animals such as Darwin's frog) to their offspring by holding them in their mouth for extended periods of time. Mouthbrooding has evolved independently in several different families of fish including the cardinalfish, sea catfish, bagrid catfish, cichlids, snakeheads, jawfishes, gouramis, and arowanas. Amphibians There is an equal prevalence of female-only and male-only care in amphibians. However, biparental care is uncommon. Provisioning in this animal group tends to be rare, and offspring guarding is more prevalent. For example, in the frog species Bibron's Toadlet, male frogs are left to care for the nest. Parental care after the laying of eggs has been observed in 5% of caecilian species, 18% of salamander species and 6% of frog species, though this number is likely an underestimate due to taxonomic bias in research and the cryptic nature of many species. Six modes of parental care are recognized among the Amphibia, in different species: egg attendance, egg transport, tadpole attendance, tadpole transport, tadpole feeding, and internal gestation in the oviduct (viviparity and ovoviviparity). the gastric-brooding frog raised tadpoles (and potentially eggs) in their stomach and the common Suriname toad raises eggs embedded in the skin on its back. Reptiles ns care for their young even after they have hatched. Reptiles provide less parental care than other tetrapods. When it does occur, it is usually female-only or biparental care. Many species within this group produce offspring that are self-sufficient, and are able to regulate their body temperatures and forage for themselves immediately after birth, thereby eliminating the need for parental care. Maternal care exists in crocodilians, where the mother assists hatchlings by transporting them in her mouth from the nest to the water. She may stay with the young for up to several months. Parental behavior have also been observed in Cunningham's skink, a viviparous lizard that protects its offspring against predators. Birds Birds are distinctive in the way they care for their young. 90% of bird species display biparental care, including 9% of species with alloparental care, or helpers at the nest. Biparental care may have originated in the stem reptiles (archosaurs) that gave rise to the birds, before they developed flight. In the remainder of bird species, female-only care is prevalent, and male-only care is rare. Most birds, including passerines (perching birds), have their young born blind, naked and helpless (altricial), totally dependent for their survival on parental care. The young are typically raised in a nest; the parents catch food and regurgitate it for the young. Some birds such as pigeons create a "crop milk" which they similarly regurgitate. David Lack developed a hypothesis that clutch size has evolved in response to the costs of parental care known as Lack's principle. It has since seen modifications but is still used as a general model. Mammals mother suckling its young mother grooming her young. There is maternal care in all species of mammals, and while 95% of species exhibit female-only care, in only 5% biparental care is present. Thus, there are no known cases of male-only care in mammals. The major adaptation shared by all live-bearing mammals for care of their young after birth is lactation (the feeding of milk from the mammary glands). Further, many mammals exhibit other parental care behaviors to increase the fitness of their offspring, for example, building a den, feeding, guarding, carrying, huddling, grooming and teaching their young. Others, consider also as a type of care when males provision the pregnant females. Humans Parenting or child rearing in humans is the process of promoting and supporting the physical, emotional, social, financial, and intellectual development of a child from infancy to adulthood. This goes far beyond anything found in other animals, including not only the provision of food, shelter, and protection from threats such as predators, but a prolonged period of support during which the child learns whatever is needed to live successfully in human society. ==In evolutionary biology==

In evolutionary biology, parental investment is the expenditure of time and effort towards rearing offspring that benefits the offspring's evolutionary fitness at a cost to parents' ability to invest in other components of the species' fitness. Parental care requires resources from one or both parents that increases the fitness of their offspring and of themselves. For example, if the strategy of parental care involves parents choosing to give each of a relatively small number of offspring an increased chance of surviving to reproduce themselves, they may accordingly have evolved to produce a small number of zygotes at a time, possibly only one. The ideal amount of parental investment would guarantee the survival and quality of both broods. Types of parental care and the amount of resources invested by parents vary considerably across the animal kingdom. The evolution of male-only, female-only, biparental or alloparental care in different groups of animals may be driven by multiple factors. Firstly, different groups may have diverse physiological or evolutionary constraints that may predispose one sex to care more than the other. Secondly, the costs and benefits of care by each sex may be influenced by ecological conditions and mating opportunities. Thirdly, operational and adult sex ratios may influence which sex has more mating opportunities, and thus predisposes one sex to care more. Furthermore, parenting decisions may be influenced by the confidence of either sex in being the genetic parent of the offspring, or paternity certainty. Male care is most prevalent in species with external fertilisation, while female care is more common with internal fertilisation. Explanations include the suggestion by Trivers (1972) that this depends on paternity certainty, A second explanation is Richard Dawkins and T. R. Carlisle's (1976) theory that the order of gamete release, and therefore the opportunity for each parent to desert, may influence which sex provides care. Internal fertilisation may provide the male parent with an opportunity to desert first, as is seen in some bird and mammal species; the roles may be reversed with external fertilisation. In fish, males often wait until a female lays her eggs before he can fertilise them, to prevent his small gametes from floating away. This allows the female to desert first, and leave male parents to care for the eggs. Male territoriality is particularly common with external fertilisation. Therefore, the male is most closely associated with the embryos. Males may defend their territories and thereby incidentally defend their eggs and young. This may preadapt males to provide care. Male care consequently involves less opportunity costs in this case, since males can still attract mates while simultaneously guarding territory and eggs. Females may even be more attracted to, and preferentially select to mate with, males that already have eggs in their nest. Male territoriality with internal fertilisation exists in some bird species. Nest size and nest building behaviour are two sexually selected traits that may attract a female to a male's territory for mating. Since the female lays her eggs in the nest within the male's territory, paternal care may evolve, even though fertilisation is internal. Amount of care Increasing parental investment in any one young benefits that particular offspring, but decreases resources for other offspring, possibly decreasing parental fitness. Therefore, there is a theoretical optimal brood size that maximises productivity for each brood. This demonstrates that both parents have the capacity to work harder and provide greater levels of care. One parent may be tempted to cheat, relying on the other parent. In biparental care, the key theoretical prediction is that parents should respond to reduced partner effort with incomplete compensation. A parent who does not put in their fair share of work then suffers reduced fitness, because their offspring receive less resources from both parents. This has been experimentally demonstrated with birds. == Flexibility of parental care ==

Parental care may vary in relation to costs and benefits When one parent is not sufficient, both parents may need to care for offspring. Each parent would like to minimise the level of care they must invest at the expense of the other parent. If one parent were to die or cease providing care, then the remaining partner may be obliged to desert the eggs or young. The extent of parental care provided to a current brood may also be influenced by prospects of future reproduction. Field experiments on a passerine bird species indicated that in areas where broods were fed with extra carotenoids, their mouths became redder. This consequently enhanced their begging displays and led parents to increase their provisioning. This was likely because the redder mouths indicated that offspring were healthier, and thus worth investing in. In other territories, the adults were also provided carotenoid-rich sugar diets, which increased the likelihood of them having a second brood in that season. Since parents that had second broods did not respond to the increased begging signals of their current brood, this indicates that parents strategically vary their sensitivity to their current broods demands in relation to their future prospects of reproducing in that season. The act of eating one's own offspring, or filial cannibalism, may be an adaptive behaviour for a parent to use as an extra source of food. Parents may eat part of a brood to enhance the parental care of the current brood. Alternatively, parents may eat the whole brood to cut their losses and improve their future reproductive success. Parental care may vary in relation to mate attractiveness In theory, a parent should invest more when paired with a mate of a high phenotypic or genetic quality. This is explained by the differential allocation hypothesis. This was shown through experimentation on zebra finches. Males were made more attractive to females by experimentally giving them red leg bands. Females increased their provisioning and raised more young when paired with these attractive males compared to when they were paired with less attractive males that had blue or green leg bands. Further experimentation on mallard ducks has displayed that females lay larger eggs and increase their provisioning when paired with more attractive males. Female peacocks have also been shown to lay more eggs after mating with males that possess more elaborate tails. Furthermore, female birds are generally more likely to care for the offspring of males that spend more time nest building, and build more elaborate nests. As a consequence, the reproductive success of males tends to increase with nest size and building behaviour. Therefore, differential allocation is expected because the offspring of these pairings would likely inherit the quality of the attractive parent, if attractiveness signifies genetic quality. Differential allocation may also work the other way around, where parents may invest less in their offspring if paired with unattractive mates. By reducing the amount of care invested in these offspring, individuals may save resources for future reproductive attempts with a more attractive mate. Differential allocation is mostly expected from females, since in many animal groups females are more choosy when assessing potential mates. However, in many bird species, males are known to be involved in caring for young, which may lead to differential allocation by males as well as females. == See also ==