The Causations and Applications of Alloparental Care Across Animal Taxa

By Chloe Bender
Introduction
Parenting is when an animal provides effort and investment into raising its young.
Parental care has evolved in many species because the benefits of offspring care has outweighed
the costs. Such benefits include a better survival chance for the offspring and increases the
fitness of the parent(s). Parental care encompasses several systems such as mating, sexual
selection, and reproductive biology (Lee et. al. 2016, Wisenden et. al. 2014, Ramos et. al. 2012)
Yet some animals will provide care to nonfilial offspring which is called alloparental care. This
behavior could be in collaboration with the biological parent or full adoption of the offspring.
This behavior has been seen in all types of animals ranging from mammals and birds to insects
and reptiles (Wysocki et. al. 2017, Riedman 1982). Depending on the species, which sex
alloparents can change. In mammals, females tend to show more alloparent behaviors while in
birds it is the males. Insects have equal amounts of alloparent behavior in both sexes (Reeve and
Reeve 1997).
Alloparental care is most common in social animals when animals focus on the survival
and fitness of the group rather than each other. Typically this leads to the parents and older
offspring co-parenting the current young offspring (Samuk and Aviles 2013, Biedremann et. al.
2011, Riedman 1982). There is two types of co-parenting: the first involves that both parents are
somehow related to the offspring they are taking care of. The second is that one parent is related
to the offspring while the helper is a non-breeding adult (Junghanns et. al. 2017). Co-parenting
can have the helping adult preform tasks such as predator detection, food gathering, cleaning,
shelter creation, and antipredator defense (Josi et. al. 2018).
There has been several hypotheses that were created to try to explain why animals would
display alloparental behavior. One of these hypotheses is kin selection (Hamilton’s rule). Kin
selection is when an animal adopts or co-parents offspring that are closely related to them. The
helpers/adoptees gained indirect fitness benefits from raising the offspring. Kin selection tends to
apply more to social groups in which animals who are closely related are in close proximity to
one another (Dias et. al. 2017, Junghanns et. al. 2017, Kalmbach 2006). Kin selection also
explains that offspring who are more closely related will show less aggression to one another
while those who are less related will show more aggression (Boos et. al. 2015).
Another hypothesis is reproductive error hypothesis. Reproductive error is when an
animal fails to discriminate between its own offspring and the offspring of another animal. This
leads to the animal adopting and raising offspring that are non-related. However, the animal
believes that it is their biological offspring. Some instances of this is if an animal recently lost its
young, they may mistake another’s offspring as their young (Wysocki et. al. 2017, Riedman
1982).
This review will try to explain what causes animals to preform alloparental behavior. The
animals that will be analyzed are mammals, birds, fish, insects, and reptiles/amphibians. The
hypotheses explained above along with other potential causations will be used to help explain the
causes of alloparental behavior. After looking at alloparental behavior in each classification of
animal, the applications of this knowledge will be analyzed. Knowing what causes an animal to

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care for non-related offspring could help with conservation efforts and the preservation of
endangered species.
Mammals
Mammals may adopt non-offspring due to hormones and chemical imbalances that are
occurring in the brain. In a postpartum state, mothers have estrogens and progestins that are used
to create a mother-offspring bond and facilitate parental care (Nunez et. al. 2013). Even if the
mother recently lost her offspring, these hormones are still activated within her brain. Therefore
if mother sees an offspring who is abandon or orphaned, she is likely to confuse the non-related
offspring as her own. When a mother mistakes a nonfilial offspring as her own, it is called
reproductive error. This behavior has been seen in feral horses (Equus caballus), bottlenose
dolphins (Tursiops truncates), and polar bears (Ursus maritimus) (Nunez et. al. 2013, Howells et.
al. 2009, Atkinson 1996). One of the main chemicals responsible for reproductive error is
oxytocin. When higher levels of oxytocin are found in certain areas of the brain, nucleus
accumbens and caudate putamen (both are located in coronal section of the frontal lobe), the
mother is more likely to preform alloparental behavior and adopt the offspring (Figure 1)
(Olazabal and Young 2006).
Another reason for adoption of non-offspring is to increase fitness of both the mother and
the child. Polar bears, cougars (Puma concolor), California sea lions (Zalophus californianus),
and red squirrels (Tamiasciurus hudsonicus) have all demonstrated instances in which the mother
would adopt nonfilial offspring in order to improve upon her own fitness (Bartnick et. al. 2014,
Gorrell et. al. 2010, Flatz and Gerber 2010, Atkinson 1996). If the adopting mother is a young
adult, she might adopt other orphaned young in order to gain experience in parenting. This would
lead to an increase in her fitness as she is preparing herself on raising her own young (Flatz and
Gerber 2010, Atkinson 1996). Other mothers might adopt older nonfilial offspring, even if they
currently have their own offspring, in order to increase the current offspring’s fitness. A cougar
adopted two non-offspring males even though she had her own litter. She did this so the adopted
males could protect her younger ones while she was hunting. It allowed her to be out longer and
increase her chancing of catching prey to feed her offspring. If a predator were to attack the litter
while the mother was gone, the predator would have attacked the older nonfilial males increasing
her related offspring’s fitness (Bartnick et. al. 2014).
Hamilton’s rule can also be used to explain alloparental behavior in mammals.
Hamilton’s rule states that a mother has a higher chance of adopting non-offspring if the benefit
of adoption is less than the cost and if the non-offspring are closely related (cousins or
nieces/nephews) to the mother (Figure 2). (Vitikainen et. al. 2017, Gorrell et. al. 2010, Atkinson
1996). Usually Hamilton’s rule applies to social species such as red squirrels. Since the mothers
are adopting their close relatives, this increases the fitness of their overall family. Offspring who
do not have close relatives are not adopted. The degree of adoption decreases as the relatedness
to the orphaned offspring decreases (Gorrell et. al. 2010).
Birds
Birds show alloparental behavior by adopting non-related young into their broods and
treating them as if they were biological offspring. Like mammals, birds also experience
reproductive error. If the orphaned fledgling and the related offspring are young and about the
same age, they would make very similar calls. This causes the parents to mistake the orphaned

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fledgling for one of their own and they will adopt the fledgling into their group (Wysocki et. al.
2017, Lengyel 2002). Offspring recognition in parents is to increase with the age of the chick.
Therefore, very young chicks can be mistaken as part of the brood. Hormones in the parents from
having recently hatched chicks also cause the parents to adopt non-related offspring as they are
reacting to the stimulus that the orphan chick creates. The hormones is the cause to reproductive
error and adoption as the hormones make the parent believe the orphaned chick is one of their
biological offspring who is in need (Kalmbach 2006).
Fledglings will leave their natal nest and seek refuge in other nests in order to increase
their fitness. Pied avocets (Recurvirostra avoset) and white stork (Ciconia ciconia) fledglings
will leave their natal nest if the amount of food they are receiving from their related parents is
not enough to sustain them (Lengyel 2002, Redondo et. al. 1995). In pied avocets, this usually
happens very early on, within 24 hours of hatching (Lengyel 2002). Since the chicks are so
young when they go looking for a new nest, the foster parents are influenced by hormones and
reproductive error causing them to adopt the chicks into their broods (Wysocki et. al. 2017,
Kalmbach 2006, Lengyel 2002). However, the main drive for this adoption is that the chicks left
voluntarily in order to increase their chances of survival and ultimately their own fitness. In
white storks this behavior occurs when the fledglings are older and are learning to fly. As the
fledglings age, the parent storks start feeding them less often and aggression between siblings
increase. When aggression between siblings and lack of food cross a threshold (Figure 3), the
fledgling will fly to a different nest that has a younger chick in hopes of being adopted and fed
more regularly, thus increasing its fitness (Redondo et. al. 1995). The chicks are making an
active decision to leave their natal nest since it is causing a decrease in fitness and find a new
nest that leads to an increase in fitness.
Brood size effects the costs and benefits of non-related offspring adoption. Having larger
broods increases predator detection and can increase the ability for the brood to defend
themselves from a predator (Lengyel 2002). However, increased predator detection also means
less time for foraging. Food also has to be divided between more chicks meaning the biological
offspring are being feed less than before the adoption (Kalmbach 2006). Parents of larger broods
tend to have a higher dominance over the other birds in the area. This leads to the larger broods
having more territory and consequently, more food/higher quality feeding patches. Lastly, young
in larger broods had a higher survival rate than young in smaller broods. Thus, parents may adopt
non-related young in order in increase their own young’s chance of survival (Kalmbach 2006,
Lengyel 2002). While there is a cost to adopting the offspring, most adult breeding pairs will
adopt the nonfilial offspring as the benefits outweigh the costs.
Instead of a mother or a pair of parents caring for young, some birds use co-parenting
techniques. This means that some birds who currently have no offspring will aid in taking care of
others young. Typically this behavior is in more social birds such as El Oro parakeet (Pyrrhura
orcesi) (Kramer et. al. 2016) and campo flickers (Colaptes campestris) (Dias et. al. 2017). In
both species, the offspring were taken care of by the biological parents and other closely related
adult birds. Thus, the likelihood that an adult bird will help care for offspring increases as the
relatedness to that offspring increases. Co-parenting occurs due to a fitness benefit to both the
offspring and the non-biological adults. Since they are social birds if they make sure their close
relatives survive, it increases their fitness in return (Dias et. al. 2017, Kramer et. al. 2016).
Fish

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 Adoption of non-related offspring in fish are caused by hormones and scents that
brooding mothers produce. Prolactin is a hormone found in fish that regulates parental care. In
daffodil cichlid species (Neolamprologus pulcher) lower levels of prolactin are in brood-caring
females rather than females who currently have no broods. However, females still have more
prolactin then males, meaning, they are more likely to display parenting behavior (Bender et. al.
2008). Foster parents and biological parents have the same levels of hormones and provided the
same level of care to the young regardless of degree of relatedness (Ramos et. al. 2012, Bender
et. al. 2008). In other cichlid species, such as convict cichlids (Amatitlania siquia), young use
hormones in order to track brood-caring females. Smaller, younger offspring follow the scent
produced by their mother in order to follow her and stay within her care. However, the older the
offspring do not show as much of a preference to their biological mothers scent and will follow
the scent of any brood-caring mother (Wisenden et. al. 2014). Since the cost of adopting
offspring is low and the more non-related offspring in the brood, the chances of a predator
attacking the offspring related to the mother decreases (Lee et. al. 2016, Wisenden et. al. 2014).
Adoptions of unrelated offspring can occur unintentionally through the act of cuckoldry
or intentionally and act as helpers to the brooding parent. While some fish adopt later in the
spawning season, primarily those in small and medium size territories, cuckoldry occurs early in
spawning season and causes the main care-giver, mainly males, to be stuck caring for unrelated
offspring. Either eggs are deposited into the brood and left to be taken care for by the foster
parent or the eggs are fertilized by a different male. Typically, the cuckoldry is preformed by
smaller fish who could not get a mate (Stiver et. al. 2012). Larger parents have a smaller
proportion of their brood containing non-related offspring meaning the larger fish are better at
defending against cuckoldry (Lee et. al. 2016). However, it is rare for a fish to have a brood that
is exclusively its own offspring (Stiver et. al. 2012). Like birds, some fish species exhibit co-
parenting. Male helpers will assist the females with caring for the young and will actually tend to
the young more often then the biological females. The females spend more time defending the
brood area while the helps clean the brood area and eggs. Helpers are normally related to one of
the two parents and have their natal brood site nearby. Helpers help females since they are
related to the offspring and increasing their survival increases the both the offspring, helper, and
mother’s fitness (Josi et. al. 2018).
Insects
When it comes to alloparental behavior and adopting nonfilial offspring, insects tend to
primarily use co-parenting techniques. A variety of insect species show co-parenting behavior. In
carpenter bees (Ceratina calcarata) and allodapine bees (Exoneurella eremophila) the first
emerging female bees will take over caring for their siblings if the mother dies shortly after
emergence. The smallest, oldest daughters are the ones that take on the new role as the mother.
Typically the mother focuses on rearing the daughter first in order to have someone to care for
her offspring in case something happens to the mother and she cannot care for her young. This
act of raising the females first so they can take over offspring care is called Bull’s “insurance by
protogyny” model (Lewis and Richards 2017, Hogendoorn et. al. 2001). The frequency of
sibling-reared nest sites increased as the year went on (Figure 4) meaning the adult females were
no longer surviving long enough to fully take care of their offspring and their oldest females took
over as the caregiver (Hogendoorn et. al. 2001).

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 Beetles display co-parenting behavior through the act of late dispersal. Female ambrosia
beetles (Xyleborus affinis) will delay dispersing from their natal nest in order to help their
mother, or siblings, raise young. Some females stay at their natal site in order to breed and raise
their own young. This has fitness benefits to both the females and their offspring since they have
their siblings to help raise the young (Biedremann et. al. 2011). Certain beetle species such as
the burying beetle (Nicrophorus vespilloides) will have reproductive error. When challenging
another beetle for territory, if the beetles are the same sex, the intruding beetle can experience
reproductive error and think the eggs/larvae in that territory is theirs. The beetle will then begin
to raise the offspring as if they biologically related. This only occurs if the beetles are the same
sex however, suggesting that some beetle species do not yet have ways to correctly identify their
own offspring (Shippi et. al. 2018).
Lastly, many spider species display alloparental behavior in the form of egg
switching or co-parenting. Social spider species tend to display egg sac switching more
frequently than solidarity spider species, however, both display the behavior. This is when
female spiders will switch webs/egg sacs and care for another spiders offspring. The spiders will
raise the offspring the exact same way regardless of the relatedness to the offspring. This
provides support for the idea that spiders lack the ability to discriminate which egg sacs are theirs
and which is another females. The female spiders just recognize that there is an egg sac to take
care of and will start providing the proper care for the young (Samuk and Aviles 2013). Female
spiders will also aid one another in taking care of the offspring. This behavior is found most
commonly in social spiders, rather than solitary ones. Adult females, including those who have
never had offspring before, will aid mothers in care of their young. This care includes the tending
of the egg sac, food regurgitation, and matriphagy. Female helpers who never had offspring
before specialize more in prey capture while the biological mother specialized more in egg sac
care (Figure 5). Females act as helpers in order to increase their fitness. Since this is most
common in social spiders, the helpers are usually closely related to the mother spider. Thus, the
helper spiders are trying to increase the fitness of their close relatives, and by association, their
own fitness (Junghanns et. al. 2017).

Reptiles/Amphibians
Amphibians and some reptile species demonstrate alloparental care. Most amphibians
provide care for their eggs until the eggs hatch. Reptiles, however, very rarely show any form of
parental care. Over 95% of reptiles show no parental care following egg laying or egg hatching
(Zilkha et. al. 2017). Some reptiles do provide care such as several species of skink: the ground
skink (Scincella latera1e) and the prairie skink (Eumeces septentrionalis). In each species, a
adult female who has laid eggs was found lying on another female’s eggs. Since parental care is
not common in reptiles, it can be assumed that in the few reptiles that due show parental care that
they have yet to adopt a method of detecting their own offspring. Therefore, the reptiles are
undergoing the reproductive error hypothesis in which they cannot determine which set of eggs
is their own and which is another females. This causes nest switching to occur in both species
(Somma 1987).
Amphibians are more likely to care for their eggs by tending to them and defending the
nest. However most leave before or slightly after the eggs hatch. Several frog species
demonstrate parental behavior by caring for their eggs then also tending to the tadpoles after they
5
hatch: glass frogs (Ikakogi tayrona), butter frogs (Leptodactylus aff. Latrans), pointed belly frogs
(Leptodactylus podicipinus) and ditch frogs (Leptodactylus aff. Leptodactyloides). In the related
species, butter frogs, pointed belly frogs, and ditch frogs, all three tend to mix broods and take
care of the others. Glass frogs have been found mixing their eggs/tadpoles with other glass frogs.
For all four species, the mixing of offspring is caused by reproductive error as amphibians, like
reptiles, have yet to evolve the mechanisms needed to detect their own young from another’s.
(Valencia and Delia 2015, Rodrigues et. al. 2011).

Applications

By understanding the different causations of alloparental behavior in animals (Table 1),

humans can use this information to manipulate animals into adopting non-related offspring.
Some insects use hydrocarbons in order to distinguish themselves from others. They use this
hydrocarbons on their young in order to tell the young that they are all related. However even if
offspring are not related to each other or to the parenting mother, placing the hydrocarbons on
the eggs/hatch offspring causes the mother to adopt the unrelated offspring. The mother will then
start to actively place her hydrocarbons on the new offspring and her own offspring in order to
minimize aggression (Boos et. al. 2015). Mammals use a technique similar to insects in offspring
identification. Through manipulation of offspring’s odor, the female may believe that an
unrelated young is actually her offspring and start to provide care for the offspring (Price et. al.
2003). While insects had a very high adoption rate of young (99%), mammals tend to have a
lower adoption rate (85%) after manipulation (Boos et. al. 2015, Price et. al. 2003). Some social
animals require the groups acceptance in order for a non-related offspring to be adopted.
Chimpanzee’s can take over four weeks before a mother actually adopted the offspring. While
the troop was interacting and engaging with the offspring, there was no parental engagement
until four weeks after the offspring was introduced to the troop (Thunstrom et. al. 2013).
Therefore, we can use these manipulations in order to illicit adoption. These efforts can be used
in conservation of endangered species, in which every individual’s survival is important to the
overall species survival. Manipulating a mother to provide care to unrelated offspring allows the
offspring to have a higher chance at survival, as well as, the ability to learn directly from a
member of the same species instead of with human-facilitated care.

Conclusion
Each group of animals have their own causations and motivations to perform alloparental
care. Mammals have hormones as a causation to alloparental care while increased fitness and kin
selection/Hamilton’s rule acted as a incentive. The hormones would make the mammals believe
the nonfilial offspring was actually their biological offspring and would proceed to provide care
for that offspring, otherwise known as reproductive error. However if the offspring and foster
parent were related to some degree, then kin selection can be used as a explanation. Since the
foster parent and offspring are related, the parent gets an increase in fitness through association
even though they are not related. Birds also demonstrated that hormones can affect alloparental
behavior by causing reproductive error. Especially if the biological offspring and the orphaned
offspring are very young in age, the parent(s) will react to the stimulus of the orphan in the same
manner as their own offspring. Fitness was also a driving factor in adoptions. However, it was
about the fitness of the fledglings as the fledglings actively left their nests in search of a nest that

6
would provide a higher fitness. Most of the time, the parent(s) would accept the orphan
chick/fledgling since the larger the brood meant during a predator attack, their offspring would
have a higher survival rate. Fish had hormonal effects on alloparental care as well, however, one
difference was that some offspring use scent in order to find care-giving mothers. Therefore, any
mother who was providing care was acceptable and the offspring would often end up in other
broods being adopted by other fish. Fish have a unique causation called cuckoldry. This lead to
unrelated offspring being placed in a fish’s brood. The fish would take care of the offspring as
they could not discriminate between their offspring and the unrelated offspring. Insects had an
emphasis on co-parenting since insects are more likely to be social species. The reproductive
error hypothesis can also be applied to insects as they often can’t distinguish between their
young and unrelated young. Lastly reptiles had very little evidence of any type of parental care.
In the ones that did show parental care, it was assumed that alloparental care only took place
since the mothers could not tell their nest/eggs apart from the others in the area. Meaning
females would nest swap and care for another’s eggs. Amphibians provided more care than
reptiles but there was still a lack of evidence supporting parental care in a variety of amphibian
species. Mainly it was frogs who provided care to their young and would provide care to other
young that would end up in their brood. Again this is due to reproductive error as the females
could not tell which eggs were theirs and which were foreign offspring.
Some of the groups of animals share the same causations/incentives of alloparental care.
This includes mammals, birds, and fish all having hormones affecting the adoption of nonfilial
offspring. However, mammals, birds, fish, and insects all have evidence of reproductive error.
Mammals and birds would adopt in order to increase their own fitness and co-parenting was
found in fish, birds, and insects. Using this information, humans can manipulate animals into
adopting nonfilial/unrelated young. This would help improve conservation efforts of endangered
species. However, due to the lack of parental care in most reptiles and amphibian species, others
ways of conserving those species would have to be further researched. Other future research can
be done on the frequency of alloparental care in the wild, specifically in reptiles and amphibians,
and other possible causations to the behavior. Since each species has different underlying causes
for alloparental behavior, looking at the commonalities between varying animal species can help
aid in understanding if this behavior was evolutionary selection and aid in conservation efforts.
More research can be done to analyze the effectiveness of forced adoptions and their role in
conservation.

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Figures and Tables

Figure 1: The amount of oxytocin (OTR) that is was found in the nucleus accumbens in several
rodent species: prairie voles (Microtus ochrogaster), rats (Rattus norvegicus domestica), mice
(Mus musculus), and meadow voles (Microtus pennsylvanicus). The oxytocin levels were
compared to the amount of time the rodents spent crouching over their offspring. The more
oxytocin in the brain, the more the rodents preformed maternal care to their young (Olazabal and
Young 2006).

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Figure 2: The cost/benefit of adoption compared to current litter size in red squirrels
(Tamiasciurus hudsonicus). The solid line represents the cost of adopting one nonfilial offspring.
The dashed lines are the degrees of relatedness between the mother and nonfilial offspring with
the most related being the top dashed line. The closed circles are adoption that increased fitness
for the mother and adopted offspring. The open circles are offspring who remained unadopted
since the adoption would have resulted in a loss in fitness (Gorrell et. al. 2010).

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Figure 3: The closed dots indicate the amount of trips biological parents made to the nest to
deliver food. Open dots are the amount of aggressive encounters that occurred between siblings
sharing a nest. Nest switching occurs when sibling aggression becomes higher than the amount
of trips the biological parents are making to the nest with food (Redondo et. al. 1995).

Figure 4: The percentage of three different types of nests found throughout the year. Black boxes
were allodapine nests that had young females raising their siblings, grey boxes are the biological
mother raising her own offspring, and white boxes are sub-social nest sites. As fall approached,
the amount of observations of sibling-reared nests increased while sub-social nests dramatically
decreased (Hogendoorn et. al. 2001).

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Figure 5: The African social spider (Stegodyphus dumicola) has female helpers aiding in the care
of other female’s young. This figure compares the female helper’s (virgin female) task in co-
parenting compared to the biological mother’s (Mother) task. In egg sac care, the biological
mother provided most of the care. However in prey attacks, the helper females preformed most
of the attacks (Junghanns et. al. 2017).

Table 1: This is a summary table comparing the different types of animals with the different
causations/behaviors of alloparenting. It is found that all animals have evidence to support the
reproductive error hypothesis. Hormones and co-parenting were the next most common among
each species. Increased fitness and cuckoldry had the least in common between the different
animal types.
Causes/Behaviors Increase Reproductive
Hormones Co-parenting Cuckoldry
of alloparenting fitness error
Mammal ✓ ✓ ✓
Bird ✓ ✓ ✓ ✓
Fish ✓ ✓ ✓ ✓
Insect ✓ ✓
Reptiles &
✓
Amphibians

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