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Andrew Huber EEOB 3310 Dr. Herbers Wikipedia Project- Annotated Bibliography 9/14/14

Proposed question: How did different parental care methods arise in poisonous species of frogs?

Townsend, D. S., Stewart M. M., Pough, F. H. (1984). Male Parental Care and Its Adaptive Significance in a Neotropical Frog. Animal Behaviour. Vol. 32, No. 2. pp. 421-431.

This article explores how certain animals invest in taking care of their offspring after they are born. This characteristic has evolved and is passed on from generation to generation, so it must have some kind of benefit. The study conducted follows the poison dart frog Eleutherodactylus coqui. It looks at how the males take care of their offspring by observing the brooding and defending of their nest. The article demonstrates that there was a much higher survival rate for eggs that were guarded by the male father frog than when left alone. This was the result of the male keeping the eggs moist, protecting against predators, and removing eggs with fungal infections. The male watch contributed to the eggs survival rate so natural selection favored this process.

Townsend, D. S., Moger, W. H. (1987). Plasma androgen levels during male parental care in a tropical frog (Eleutherodactylus). Hormones and Behavior. Vol. 21, No. 1. pp. 39-99.

This paper by Townsend and Moger looks into a physiological difference in a male poison dart frog while it is acting as a parent. They analyzed the hormones present in the frog Eleutherodactylus coqui for androgens. Levels of the hormone were compared among male frogs that were parenting, calling, mating, and not mating. The results show that a noticeable decline in androgens happened in males that were transitioning from mating to parenting. With less androgens, the frogs called for mates less and practiced less egg cannibalism. This study demonstrates that there is a genetic background for the male parenting approach and an evolutionary support for the increased safety of the developing offspring.

Brust, D. G. (1993). Maternal Brood Care by Dendrobates pumilio: A Frog that Feeds its Young. Journal of Herpetology. Vol. 27, No. 1. pp. 96-98.

This journal observes the parental care patterns of the strawberry poison dart frog known as Dendrobates pumilio. The male serves as the primary caretaker for the first week after the eggs are laid. He is responsible for keeping the eggs protected and moist by urinating on them. Once they became tadpoles, the female would make many trips to the place in the water where they were raised. Eggs would be released by the female, which would then be fed on by the tadpoles. There is evolutionary support for the males taking care of the eggs while the female gathers food and energy for producing eggs that can be eaten. This system of both parents allows the eggs and tadpoles to be safe and fed instead of choosing one or the other.

Brown, J. L., Morales, V., Summers K. (2010). A Key Ecological Trait Drove the Evolution of Biparental Care and Monogamy in an Amphibian. The American Naturalist. Vol. 175, No. 4. pp. 436-446.

This article observes the connections of the size of the breeding pool and trophic egg feeding to the development of various types of parental raising. For small pools, it is necessary that the eggs be taken care of by both parents because of the dramatic increase in survival of offspring. The large pools allowed one parent to take care of the offspring, but there was much greater predation happening in those areas. The male typically does the basic parenting necessities, and biparental care has developed from the evolution of females going back to take care of their offspring. Overall, the article shows a very logical and reasonable development of the parental care in the poison dart frogs in evolutionary time.

Caldwell, J. P., Veronica, R. L. (1999). Determinants of Biparental Care in the Spotted Poison Frog, Dendrobates vanzolinii. Copeia. Vol. 1999, No. 3. pp. 565-575.

This study looks into the complex systems that poison frogs have for raising their offspring. The article focuses in particular on the species Dendrobates vanzolinii, and it dives into questions about the roles of males and females. The frog was discovered to exhibit biparental rearing of the offspring. Two of the main causes of this pattern were the difficulties of finding places for laying eggs and the need for food at the eggs sites. The males were necessary because they provided the protection that allowed the females to feed the young with their eggs. The evolution of this biparental care was critical for the frogs to live on both land and water and to ensure the survival of their young.

Assignment 2
https://en.wikipedia.org/wiki/Dendrobatinae 3 suggestions: My suggestions would be to include evolutionary background to the phenomenon, such as the statement that the biparental care system was an adaptation of frogs to breed on land due to the risk of predation in ponds. While the female gathers energy to feed the eggs, the male broods over the eggs, which increases their survival rate and is favored by natural selection. Finally, show how the breeding on land returns to water by stating that the female can deliver the tadpoles to the pond once the eggs are hatched. 1 sentence + citation: The males are responsible for protecting the eggs from predation and keeping the eggs from drying out by urinating on them. Brust, D. G. (1993). "Maternal Brood Care by Dendrobates pumilio: A Frog that Feeds its Young", Journal of Herpetology. Vol. 27, No. 1. pp. 96-98.

Post was checked Tuesday, 10/14/14 and no responses were present.

Evolution of Biparental Care in Tropical Frogs
Parental care is seen in many different types of forms in animals. When reproducing sexually, one or both parents may take care of their offspring to ensure that they have a better chance of survival. Forms of parental care include protecting, feeding, and guiding the offspring until they are self-sustainable. Sometimes, care given by the parents is necessary for the offspring to survive. Some frogs partake in parental care in various ways. The mother, father, or a combination of both sexes carry out this parental investment. The evolution of biparental care, which is the joint effort of both parents, is a topic that is still being investigated. Biparentalism arose in some species of tropical frogs as a result of the ecological conditions, the differences between the sexes, and the natural tendencies of the amphibians.

Tropical frogs show parental care in a number of ways despite the costs associated with the services. Dendrobates, a genus of poisonous frogs, carry out parental care by staying near the clutch, which is the group of eggs laid by the female (Brust 1993). In this location, the poisonous frogs can guard the eggs against predators, keep the eggs from drying out, and eat eggs infested with fungi (Brust 1993). Eleutherodactylus coqui, a tropical frog found in Puerto Rico, also does the same parental functions as well as brooding over the eggs (Townsend, Steward, and Pough 1984). However, E. coqui and other parental care species suffer some fitness costs as a result of its responsibilities. The frogs typically lose body weight or abdominal fat and miss out on all mating chances during this period of brooding and protecting (Townsend 1986). Additionally, the E. coqui frogs undertake the great risk of dying or losing their ability to mate successfully while defending their offspring. These costs must be outweighed by the benefits of parental care, or else the trait would be weeded out by natural selection. Taking part in parental care improves the parents’ direct fitness, which is measured by the number of offspring that survive to reproducing age. As shown in one study, the success of clutches drops from 77.5 percent to 23.1 percent when the E. coqui parents are taken away from their eggs (Townsend et al. 1984). The parents could elect to mate more often or take better care of themselves. However, they choose to improve the state of their offspring, who are reliant on their contributions.

An important aspect of parental care is who accepts the duties of parenting. In the Dendrobates genus, there are biparental care species, such as D. ventrimaculatus and D. fantasticus, as well as male and female parental care species (Summers and Earn 1999). Phylogenetic evidence shows that male parental care is the ancestral strategy in these Dendrobates (Summers and Earn 1999). The trend of using males to guard or brood eggs, even in the biparental care, can be understood from the perspective of the female. After oviposition, or laying of the eggs, the females need to replenish their bodies that have been dedicated to nurturing the eggs before they can mate again (Weygoldt 1987). Dedicating herself to watching over the eggs would delay the next opportunity for the mother to mate by about two to four weeks (Weygoldt 1987). Since this outcome would cause many males to compete for a few females able, the males are favored for the brooding (Weygoldt 1987). Females can also take full responsibility of the brood care, but males are favored for the task from the view of available mates and the gender ratio.

In the Eleutherodactylus coqui frogs, the males assume the parental responsibilities once the eggs are laid (Townsend 1986). The E. coqui females lay eggs on a plant so that the male can brood, or sit over, the eggs (Townsend et al. 1984). This act prevents moisture from leaving the egg and serves as protection for the developing eggs (Townsend et al. 1984). Males show distinctly different behavior when they are guarding eggs. Those E. coqui males that are defending eggs have a significant decline in the amount of androgens in their blood compared to mating and non-mating males (Townsend and Moger 1987). The lowered amount of the hormone causes them to stop sending out calls for mates and to lessen their cannibalistic tendencies for the eggs (Townsend and Moger 1987). This species does not have a tadpole stage outside of the eggs, which allows the hatchlings to come out and survive in the land environment (Townsend et al. 1984). Because offspring develop conveniently in one place, the male can raise E. coqui offspring by himself.

The environment can have a substantial impact on the uses of parental care. Not all tropical frogs have the ability to lay their eggs just on land or plants. Unlike Eleutherodactylus coqui, the eggs many species of frogs hatch into tadpoles that need water in order to survive (Summers and Earn 1999). Tropical frogs can choose from a variety of water sources, such as lakes, streams, and small puddles. There is greater risk involved with reproducing in bigger bodies of water because of the higher likelihood of fish and other aquatic predators being there (Brown, Morales, and Summers 2010). Instead, frogs can choose to place eggs in phytotelmata, which are small pools of water that collect at the bases of leaves or natural holes in trees (Brown et al. 2010; Caldwell and de Oliveira 1999). However, there is a trade-off that comes with electing a smaller water source. Not much sunlight reaches these locations, so algae and other food sources cannot grow to feed the inhabitants (Caldwell and de Oliveira 1999). Tropical frogs must use alternative methods of feeing their tadpole offspring. In the case of using phytotelmata, it is very difficult for one parent to guard and feed his or her offspring in possibly several different places. Roles performed by both parents provide a great advantage to the offspring (Caldwell and de Oliveira 1999). As seen in the Amazon Rain Forrest, the different size of the bodies of water chosen for breeding correlates with the amount of biparental care in two very similar species of the Ranitomeya genus (Brown et al. 2010). Ranitomeya imitator favors smaller pools and used biparental care (Brown et al. 2010). Conversely, Ranitomeya variabilis utilizes larger bodies of water for breeding and only males took part in parental care (Brown et al. 2010). The ecologic aspects of a species habitat can have significant impacts on the development of certain types of parental care.

The process that caused biparental care to come about in the environment is still up for debate. There are many ideas and theories about how it arose in certain species. With the environmental pressures of choosing phytotelmata over bigger bodies of water, natural selection could have shaped this process. Trophic egg feeding plays a key role in the ability of frogs to breed in smaller bodies of water that lack food sources (Brown et al. 2010). This is the process of females laying eggs for tadpoles to eat. Many species of tropical frogs have an inherent nature of cannibalism, such as Dendrobates vanzolinii, that allow their tadpoles to utilize the eggs for nourishment (Caldwell and de Oliveira 1999). With a male guarding the eggs, an intermediate step to developing biparental care may have been using the eggs from a mating with another female to feed existing tadpoles (Summers and Earn 1999). Males could direct where the eggs should be positioned, and then he could move them into the water that holds his tadpole offspring (Summers and Earn 1999). This polygynous relationship puts a cost on the female because she loses eggs to benefit offspring that are not her own (Summers and Earn 1999). It is possible that the female could counter this effect by participating in biparental care with her mate (Summers and Earn 1999). Also, since there may be a lack of males, females could benefit from attacking other clutches that her mate might have fertilized (Wells 1978). Eliminating competition of a female’s offspring might result in higher survival of those she is trying to protect. Females have been seen eating other females’ eggs in captivity by Dendrobates auratus (Wells 1978). This intrasexual competition among the females might have been another important driving force in bringing about biparental care. Many factors contributed to the evolution of biparental care in some species of tropical frogs.

The males perform a certain set of functions in the biparental care system. The males typically take care of the basic parental duties involved with ensuring the eggs hatch successfully (Brown et al. 2010). The males have the important role of guarding the eggs against predators, and these threats can even be from the same species because of cannibalism (Brust 1993). Also, cannibalism in frogs puts a limit on the number of eggs that can be placed in each phytotelmata because of the likelihood that only one tadpole will consume the other eggs once it hatches (Caldwell and de Oliveira 1999). Thus, frogs may use several water sources for laying their eggs. Males can be very territorial and lead females to certain sites where eggs can be placed, as in Dendrobates vanzolinii (Caldwell and de Oliveira 1999). Ensuring the phytotelmata are located in a good area increases the probability that the eggs will develop into healthy tadpoles. If phytotelmata are close to the frogs’ natural living area, then the two sexes can easily cooperate for biparental care (Summers and Earn 1999). Increasing the distance between the living area and the breeding area makes it more difficult to maintain this bond between the male and female (Summers and Earn 1999). Finally, in Dendrobates vanzolinii and some other species, males can deliver tadpoles to a new body of water once they are mature enough (Caldwell and de Oliveira 1999). Males perform a number of roles in biparental care to ensure the success of their offspring.

The tropical frog females also have certain responsibilities in the parenting effort. An obvious contribution is that the females carry out the trophic egg feeding. Females put fertilized or unfertilized eggs into the tadpole’s water in order to feed the growing offspring (Ryan 2010). To ensure that the phytotelmata contain their offspring, the females look into the water source (Brust 1993). The tadpoles can poke their head near their mothers and wiggle their bodies to ensure they are seen (Brust 1993). Then the mother can be sure that she is giving eggs to the right tadpole. As in Dendrobates vanzolinii, the mothers usually stay around where the eggs are and refrain from coming into contact with other frogs (Caldwell and de Oliveira 1999). There they can replenish themselves and take in energy for producing more eggs, which is a significant investment. Females are sometimes the sex that transports the tadpoles from the clutch to certain ponds or bodies of water, such as females in Dendrobates pumilio (Brust 1993). Females contribute to the biparental care through their various roles.

Because of their life histories, the male and female functions, and the environmental conditions, some tropical frog species have evolved biparental care systems. Many frogs have a tadpole stage in their development immediately after the hatching. The frogs avoid some predators by breeding in smaller bodies of water. Moving away from large bodies of water brings other challenges for the tropical frogs, which can be dealt with through cooperation and division of labor in a mating couple. For future research, the method that caused biparental care to arise should be investigated in a variety of tropical frog species. Also, one should look further into the relationship between mating systems, such as polygyny and monogamy, and biparental care to see if there is any correlation or causation involved. Biparentalism in tropical frogs is difficult topic, and many aspects of this parental system still need to be researched.

Posted Webpage
https://en.wikipedia.org/wiki/Evolution_of_biparental_care_in_tropical_frogs

Many tropical frogs have developed a parental care system where both the mother and father partake in raising their offspring. The evolution of biparental care, which is the joint effort of both parents, is a topic that is still under investigation. Biparentalism arose in some species of tropical frogs as a result of the ecological conditions, the differences between the sexes, and their natural tendencies.

Male parental care could have served as the basis for the development of biparental care. Phylogenetic evidence shows that male parental care is the ancestral strategy in Dendrobates. Currently there are Dendrobates species, such as D. ventrimaculatus and D. fantasticus, that exhibit biparental care. The trend of using males to guard or brood eggs for biparental care or paternal care can be understood from the perspective of the female. After oviposition, or when the eggs are laid, the females need to replenish their bodies that have been dedicated to nurturing the eggs before they can mate again. Brooding by the females would delay the opportunity to mate by about two to four weeks. Since this outcome would cause many males to compete for a few females that are able to mate, the males are favored for the brooding.

The environment can have a substantial impact on the uses of parental care. Not all tropical frogs have the ability to lay their eggs plainly on land or plants. Tropical frogs can choose from a variety of water sources, such as lakes, streams, and small puddles. There is greater risk involved with reproducing in bigger bodies of water because of the higher likelihood of fish and other aquatic predators being there. Instead, frogs can choose to place eggs in phytotelmata. However, there is a trade-off that comes with electing a smaller water source. Not much sunlight reaches these locations, so algae and other food sources cannot grow to feed the inhabitants. Tropical frogs must use alternative methods of feeding their tadpole offspring. In the case of using phytotelmata, it is very difficult for one parent to guard and feed his or her offspring in possibly several different places. Roles performed by both parents provide a great advantage to the offspring. As seen in the Amazon Rain Forrest, the different size of the bodies of water chosen for breeding correlates with the amount of biparental care in two very similar species of the Ranitomeya genus. Ranitomeya imitator favors smaller pools and uses biparental care. Conversely, Ranitomeya variabilis utilizes larger bodies of water for breeding and only males take part in parental care. The ecologic aspects of a species habitat can have significant impacts on the type of parental care exhibited.

Trophic egg feeding plays a key role in the ability of frogs to breed in smaller bodies of water that lack food sources. Many species of tropical frogs have an inherent nature of cannibalism, such as Dendrobates vanzolinii, that allow their tadpoles to utilize the eggs for nourishment. With a male guarding the eggs, an intermediate step to developing biparental care may have been using the eggs from a mating with another female to feed existing tadpoles. Males could direct where the eggs should be positioned, and then he could move them into the water that holds his tadpole offspring. This polygynous relationship puts a cost on the female because she loses eggs to benefit offspring that are not her own. It is possible that the female could counter this effect by participating in biparental care with her mate. Also, since there may be a lack of males, females could benefit from attacking other clutches that her mate might have fertilized. Eliminating competition of a female’s offspring might result in higher survival of those she is trying to protect. Females have been seen eating other females’ eggs in captivity by certain species, such as Dendrobates auratus. This intrasexual competition among the females might have been another important driving force for bringing about biparental care. Many factors contributed to the evolution of biparental care in some species of tropical frogs.

Final Draft
Evolution of Biparental Care in Tropical Frogs Andrew Huber; Recitation Thursday 3 p.m.

Parental care is seen in many different forms in animals. When reproducing sexually, one or both parents may take care of their offspring to ensure that they have a better chance of survival. Methods of parental care include protecting, feeding, and guiding the offspring until they are self-sustainable. Sometimes care given by the parents is necessary for the offspring to survive. Certain frogs partake in parental care in various ways. The mother, father, or a combination of both sexes carry out this parental investment. The evolution of biparental care, which is the joint effort of both parents, is a topic that is still under investigation. Biparentalism arose in some species of tropical frogs as a result of their natural tendencies, the ecological conditions, and the differences between the sexes.

Tropical frogs show parental care in a number of ways despite the costs associated with the services. Dendrobates, a genus of poisonous frogs, carry out parental care by staying near the clutch, which is the group of eggs (Brust 1993). In this location, the poisonous frogs can guard the eggs against predators, keep the eggs from drying out, and eat eggs infested with fungi (Brust 1993). Eleutherodactylus coqui, a tropical frog found in Puerto Rico, also does the same parental functions as well as brooding, or sitting, over the eggs (Townsend, Steward, and Pough 1984). However, E. coqui and other parental care species suffer individual fitness costs as a result of their responsibilities. The frogs typically lose body weight or abdominal fat and miss out on all mating chances during this period of brooding and protecting (Townsend 1986). Additionally, the E. coqui frogs undertake the great risk of dying or losing their ability to mate successfully while defending their offspring. These costs must be outweighed by the benefits of parental care, or else the trait would be weeded out by natural selection. Taking part in parental care improves the parents’ direct fitness, which is measured by the number of offspring that survive to reproducing age. As shown in one study, the success of clutches drops from 77.5 percent to 23.1 percent when the E. coqui parents are taken away from their eggs (Townsend et al. 1984). The parents could elect to take better care of themselves or mate more often. However, the frogs choose to improve the state of their offspring, who are reliant on their contributions.

An important aspect of parental care is to whom the duties of parenting are given. In the Dendrobates genus, there are biparental care species, such as D. ventrimaculatus and D. fantasticus, as well as male and female parental care species (Summers and Earn 1999). Phylogenetic evidence shows that male parental care is the ancestral strategy in these Dendrobates (Summers and Earn 1999). The tendency of using males to guard or brood eggs, even in biparental care, can be understood from the perspective of the female. After laying the eggs, also known as oviposition, the females need to replenish their bodies that have been dedicated to nurturing the eggs before they can mate again (Weygoldt 1987). Brooding by the females would delay the opportunity to mate by about two to four weeks (Weygoldt 1987). Since this outcome would cause many males to compete for a few females that are able to mate, the males are favored for the brooding (Weygoldt 1987). Females can also take full responsibility of the brood care, but males are favored for the task from the view of available mates and the gender ratio.

In the Eleutherodactylus coqui frogs, the males assume the parental responsibilities once the eggs are laid (Townsend 1986). The E. coqui females lay eggs on a plant so that the male can brood over the eggs (Townsend et al. 1984). This act prevents moisture from leaving the egg and serves as protection for the developing eggs (Townsend et al. 1984). Males show distinctly different behavior when they are guarding eggs. E. coqui males that are defending eggs have a significant decline in the amount of androgens in their blood compared to mating and non-mating males (Townsend and Moger 1987). The lowered amount of the hormone causes them to stop sending out calls for mates and to lessen their cannibalistic tendencies for the eggs (Townsend and Moger 1987). This species does not have a tadpole stage outside of the eggs, which allows the hatchlings to come out and survive on land (Townsend et al. 1984). Because offspring develop conveniently in one place, the male can raise E. coqui offspring by himself.

The environment can have a substantial impact on the uses of parental care. Not all tropical frogs have the ability to lay their eggs solely on land or plants. Unlike Eleutherodactylus coqui, the eggs of many species of frogs hatch into tadpoles that need water in order to survive (Summers and Earn 1999). Tropical frogs can choose from a variety of water sources, such as lakes, streams, and small puddles. There is greater risk involved with reproducing in bigger bodies of water because of the higher likelihood of fish and other aquatic predators being there (Brown, Morales, and Summers 2010). Instead, frogs can choose to place eggs in phytotelmata, which are small pools of water that collect at the bases of leaves or natural holes in trees (Brown et al. 2010; Caldwell and de Oliveira 1999). However, there is a trade-off that comes with electing a smaller water source. Not much sunlight reaches these locations, so algae and other food sources cannot grow to feed the inhabitants (Caldwell and de Oliveira 1999). Tropical frogs must use alternative methods of feeding their tadpole offspring. In the case of using a few phytotelmata, it is very difficult for one parent to guard and feed offspring in several different places. Roles performed by both parents provide a great advantage to their offspring (Caldwell and de Oliveira 1999). As seen in the Amazon Rain Forrest, the different size of the bodies of water chosen for breeding correlates with the amount of biparental care in two very similar species of the Ranitomeya genus (Brown et al. 2010). Ranitomeya imitator favors smaller pools and uses biparental care (Brown et al. 2010). Conversely, Ranitomeya variabilis utilizes larger bodies of water for breeding and only males take part in parental care (Brown et al. 2010). The ecologic aspects of a species habitat can have significant impacts on the type of parental care exhibited.

The process that caused biparental care to come about in the environment is still up for debate. There are many ideas and theories about how it arose in certain species. With the environmental pressures of choosing phytotelmata over bigger bodies of water, natural selection could have shaped this process. Trophic egg feeding plays a key role in the ability of frogs to breed in smaller bodies of water that lack food sources (Brown et al. 2010). This feeding involves the females laying eggs for tadpoles to eat. Many species of tropical frogs have an inherent nature of cannibalism, such as Dendrobates vanzolinii, that allow their tadpoles to utilize the eggs for nourishment (Caldwell and de Oliveira 1999). With a male guarding the eggs, an intermediate step to developing biparental care may have been using the eggs from a mating with another female to feed existing tadpoles (Summers and Earn 1999). A male frog could direct where the eggs should be positioned, and then he could move them into the water that holds his tadpole offspring (Summers and Earn 1999). This polygynous relationship puts a cost on the female because she wastes eggs to benefit offspring that are not her own (Summers and Earn 1999). It is possible that the female could counter this effect by participating in biparental care with her mate (Summers and Earn 1999). Also, since there may be a lack of males, females could benefit from attacking other clutches that her mate might have fertilized (Wells 1978). Eliminating competition of a female’s offspring might result in higher survival of those she is trying to protect. Females have been seen eating other females’ eggs in captivity by certain species, such as Dendrobates auratus (Wells 1978). This intrasexual competition among the females might have been another important driving force for bringing about biparental care. Many factors contributed to the evolution of biparental care in some species of tropical frogs. The males perform a certain set of functions in the biparental care system. The males typically take care of the basic parental duties involved with ensuring the eggs hatch successfully (Brown et al. 2010). The males have the important role of guarding the eggs against predators, and these threats can even be from the same species because of cannibalism (Brust 1993). Also, cannibalism in frogs puts a limit on the number of eggs that can be placed in each phytotelmata because of the likelihood that only one tadpole will consume the other eggs once it hatches (Caldwell and de Oliveira 1999). Thus, frogs may use several water sources for laying their eggs. Males can be very territorial and lead females to certain sites where eggs can be placed, as in Dendrobates vanzolinii (Caldwell and de Oliveira 1999). Ensuring the phytotelmata are located in a good area increases the probability that the eggs will develop into healthy tadpoles. If phytotelmata are close to the frogs’ natural living area, then the two sexes can easily cooperate for biparental care (Summers and Earn 1999). Increasing the distance between the living area and the breeding area makes it more difficult to maintain this bond between the male and female (Summers and Earn 1999). Finally, in Dendrobates vanzolinii and some other species, males can deliver tadpoles to a new body of water once they are mature enough (Caldwell and de Oliveira 1999). Males perform a number of roles in biparental care to ensure the success of their offspring.

The tropical frog females also have certain responsibilities in the parenting effort. An obvious contribution is that the females carry out the trophic egg feeding. Females put fertilized or unfertilized eggs into the tadpole’s water in order to feed the growing offspring (Ryan 2010). To ensure that the phytotelmata contain their offspring, the females look into the water (Brust 1993). The tadpoles can poke their head near their mothers and wiggle their bodies to ensure they are seen (Brust 1993). Then the mother can be sure that she is giving eggs to her own tadpole. As in Dendrobates vanzolinii, the mothers usually stay around where the eggs are and refrain from coming into contact with other frogs (Caldwell and de Oliveira 1999). There they can replenish themselves and take in energy for producing more eggs, which is a significant investment. Females are sometimes the sex that transports the tadpoles from the clutch to certain ponds or bodies of water, such as females in Dendrobates pumilio (Brust 1993). Females contribute to the biparental care through their various roles.

Because of their life histories, the male and female functions, and the environmental conditions, some tropical frog species have evolved biparental care systems. Many frogs have a tadpole stage in their development immediately after the hatching. The frogs avoid some predators by breeding in smaller bodies of water. Moving away from large bodies of water brings other challenges for the tropical frogs, which can be dealt with through cooperation and division of labor in a mating couple. For future research, the method that allowed biparental care to arise should be investigated in a variety of tropical frog species. Also, one should look further into the relationship between mating systems, such as polygyny and monogamy, and biparental care to see if there is any correlation or causation involved. Biparentalism in tropical frogs is difficult topic, and many aspects of this parental system still need to be researched.

Responses
Thanks for your new article, Evolution of biparental care in tropical frogs. It's a nice addition to Wikipedia, but it lacks any wikilinks. If you linked to other articles, that would help readers learn more about any unfamiliar terms they might encounter in the article. Ian (Wiki Ed) (talk) 15:08, 17 November 2014 (UTC)

I added 13 wikilinks to the article in response to this suggestion.

There were also contributions by two users that added the categories frogs, behavioral ecology, and evolutionary biology.