Strawberry poison-dart frog

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Strawberry poison-dart frog
Oophaga pumilio (Strawberry poision frog) (2532163201).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Anura
Family: Dendrobatidae
Genus: Oophaga
Species:
O. pumilio
Binomial name
Oophaga pumilio
(Schmidt, 1857)
Synonyms

Dendrobates pumilioSchmidt, 1857

The strawberry poison frog, strawberry poison-dart frog or blue jeans poison frog (Oophaga pumilio, formerly Dendrobates pumilio) is a species of small poison dart frog found in Central America. [2] It is common throughout its range, which extends from eastern central Nicaragua through Costa Rica and northwestern Panama. The species is often found in humid lowlands and premontane forest, but large populations are also found in disturbed areas such as plantations. [3] The strawberry poison frog is perhaps most famous for its widespread variation in coloration, comprising approximately 15–30 color morphs, most of which are presumed to be true-breeding. [4] O. pumilio, while not the most poisonous of the dendrobatids, is the most toxic member of its genus.[ citation needed ]

Contents

Diet

The diet of O. pumilio causes the skin of the amphibian to become toxic in nature when certain subspecies of mites and ants are ingested very similar to many other poison dart frogs. [5] [6] Alkaloid toxins are organic in nature and contain nitrogenous bases that react with carbon and hydrogen groups. [7] Pumiliotoxin 251D is the specialized toxin that is sequestered by this species of frog. This toxin has a negative stimulating effect on cardiac function and is a severe disruptor of the sodium potassium ion channels within cells. Upon ingestion of Pumiliotoxin 251D, organisms preying on O. pumilio experience convulsions, paralysis, and death. [7]

It has been found that once O.pumilio reaches sexual maturity, their granular glands significantly increase in size and their diet shifts. In females, it is common to find about 53% more alkaloids than adult males. [8]

Oribatida mites belonging to the glandulate suborder Brachypylina are an important origin of pumiliotoxins in O. pumilio. Hexane-extraction techniques indicate presence of alkaloid toxins in Brachypylina. [9] Toxins appear to be biosynthesized in adult mites, as nymph and larval stages of the arachnid do not carry the toxins. Experimental analysis of this species of mite show alkaloid toxins are found almost exclusively in the opisthonotal glands of mites of the Scheloribatidae. [10] Oil glands of the mite contain the toxins and are then released internally as the amphibian digests the arthropod.

O. pumilio can also attribute its cutaneous toxicity to its rich diet of formicinae ants. [6] Species of the formicine genus Brachymyrmex contain pumiliotoxins which the frogs incorporate and accumulate poison from. [11] There is a variability of alkaloid profiles among populations and individuals of O. pumilio, which is indicative of varying levels of available prey within their infraspecific habitats. [12] Research and physical analysis reveal that maternally derived alkaloids exist in young tadpoles. [13] The increase in alkaloids in tadpoles suggests that the females are providing more chemical defenses to their more vulnerable young. This being one of the first found examples of provisioning that occurs after hatching. [8] During tadpole-rearing, mother frogs feed their young an unfertilized egg from their ovaries after dropping each individual tadpole into a repository of water usually found in a bromeliad. [14] Tadpoles lacking the obligate nutritive egg diet do not contain the alkaloid. [13] This step is crucial for the tadpoles to sequester the alkaloid from their mother; without such, young tadpoles become susceptible to predation by arthropods and other frogs.

Behavior

Oophaga pumilio is diurnal and primarily terrestrial, and can often be found in leaf litter in both forested and disturbed areas. Studies have shown that the optimal habitat is determined by the male, considering the resource benefits and defense costs. Males tend to expend more energy defending smaller but higher quality areas. [15] There has also been evidence that the better competitors and fighters are the males guarding smaller sites with higher female density. In most Anura the louder the vocalization when competing usually means they are larger in size and in better health. However, in the O. pumilio species researchers have determined that these frogs call out at a lower rate to limit their energetic expenditures. [15] Females, on the other hand, simply distribute themselves according to tadpole rearing sites. [15]

Though brightly colored and toxic, these frogs are relatively small, growing to approximately 17.5–22 mm (0.69–0.87 in) in standard length. [3]

Gnome-mime-sound-openclipart.svg
Listen to a pumilio call
Male advertisement call
Problems playing this file? See media help.

Reproduction and parental care

Oophaga pumilio is an external breeder, and other species of the genus Oophaga are notable in the amphibian world for exhibiting a high degree of parental care. [16] The strawberry poison frog has dual parental care. [17] The males defend and water the nests, and the females feed the oophagous tadpoles their unfertilized eggs. Although both male and female contribute to parental care, females invest more heavily in terms of energy expenditure, time investment, and loss of potential reproduction. [17] When choosing a partner for mating, females will choose the closest calling male rather than the highest quality male. [18] Females provide energetically costly eggs to the tadpoles for 6–8 weeks (until metamorphosis), remain sexually inactive during tadpole rearing, and care for only one clutch of four to six tadpoles at a time. [17] The males contribute via the relatively "cheap" (in terms of energy) act of watering and protecting the eggs for a relatively short period (10–12 days), and can care for multiple nests at one time. [17] The extreme maternal investment in their offspring is believed to be the result of high egg mortality. Only 5–12% of the clutch develops into tadpoles, so the female's fitness may be best increased by making sure those few eggs that form tadpoles survive. [19]

The la gruta morph from Colon Province, Panama OophagaPumilio colon lagruta to source dendrobase.hu.jpg
The la gruta morph from Colón Province, Panama

After mating, the female lays three to five eggs on a leaf or bromeliad axil. The male then ensures the eggs are kept hydrated by transporting water in his cloaca. After about 10 days, the eggs hatch and the female transports the tadpoles on her back to some water-filled location. [20] In captivity, on rare occasions, the male is observed transporting the tadpoles, though whether this is intentional, or the tadpoles simply hitch a ride, is unknown. Bromeliad axils are frequently used tadpole deposition sites, but anything suitable can be used, such as knots in trees, small puddles, or human trash such as aluminum cans.

Tadpoles are deposited singly at each location. Once this has been done, the female will come to each tadpole every few days and deposit several unfertilized food eggs. [3] In captivity, tadpoles have been raised on a variety of diets, ranging from algae to the eggs of other dart frogs, but with minimal success. O. pumilio tadpoles are considered obligate egg feeders, as they are unable to accept any other form of nutrition.

After about a month, the tadpole will metamorphose into a small froglet. Generally, it stays near its water source for a few days for protection as it absorbs the rest of its tail.

Taxonomy

Oophaga pumilio belongs to the genus Oophaga , [16] although the name Dendrobates pumilio is still sometimes used. There is evidence that the species of Oophaga (previously classified as the "female parental care group" of Dendrobates [21] ) are a monophyletic evolutionary group. Due to the low level of genetic divergence between the species analyzed in this genus, it is estimated that they speciated relatively recently, after the formation of the current Panamanian land bridge in the Pliocene (3–5 million years ago). [22] Oophaga pumilio is believed to be most closely related to Oophaga arborea [23] and Oophaga sylvatica. [16]

Evolution

Strawberry poison frog, O. pumilio, shows extreme variation in color and pattern between populations that have been geographically isolated for more than 10,000 years. [24] When populations are separated by geographic distances and landscape barriers, they frequently experience restricted gene flow, which can enable phenotypic divergence between populations through selection or drift. [25] Their variety in warning coloration is used for their visibility, toxicity and resistance to predators. When divergent phenotypes are mostly restricted to separate islands, the biogeography of color polymorphism suggests a major role for neutral process. However, Summers et al. (1997) [26] provide evidence that neutral divergence alone is unlikely to have caused the variation in color patterns. As shown by Lande, rapid evolution in sexually selecting species is led by the interaction of random genetic drift with natural and sexual selection such as random genetic drift in female mating preferences. [27] Color is known to play a role in male–female signaling, mate attraction, and male–male signaling in anurans. Based on Tazzyman and Iwasa's study that involved collections of samples from main islands in the Bocas del Toro archipelago, its results proved that female preference on male calls led to call divergence and therefore divergence was driven by sexual selection. Mate choice plays a critical role in generating and maintaining biodiversity. [28] Furthermore, spatial variation in predators or habitat features could exert divergent natural selection on coloration in response to its subjection to predator selection. [24] It is still unclear to what extent sexual selection has driven the evolution of color morphs rather than reinforcing the reproductive isolation of morphs. [29] In an aposematic organism such as O. pumilio, phylogenetic signal of selection cannot be attributed to female mate choice alone but is quite possible that genetic drift would interact with female color preferences to trigger divergence [24] Researchers Maan and Cummings had also found that in some cases female O. pumilio preferred male mates that had very different coloration than their own phenotype. [30] In nature, the equality of color through evolution is very unlikely considering the various sensory biases of predators and the different background colors of the environments these frogs inhabit. Due to this variability in color evolution, it is unlikely to say there is superiority of aposematic purpose of color selection in Oophaga. Species such as O. pumilio have been known to thrive and compete very well on disturbed and converted land. With temperatures rising in many different biomes, the success of many species is going to be determined by its ability to acclimate and adapt. In a study done by Rivera and Nowakowski, they discovered that in many cases O. pumilio is experiencing greater temperature stress in converted habitats than forests. [30]

Habitat niche

This frog species utilizes scattered structures throughout disturbed lands to relieve some thermal stress. However, O. pumilio is still warmer than any other species in the forested areas, being exposed to temperatures up to 27 degrees Celsius. These findings suggest that this species of dart frog acts as an ecological buffer and will be more successful than other species as land use changes and temperatures rise. [31]

Captivity

Oophaga pumilio is a popular frog in captivity, due to its striking colors and unique life cycle. They have been imported in vast quantities to the United States and Europe since the early 1990s, when they would typically be available for around US$75 each. However, these shipments have since stopped, and O. pumilio is much less common and available in reduced diversity. A select number of hobbyists and breeders are successfully reproducing these frogs in captivity, and healthy, captive-bred individuals have become much easier to find (in the 21st century).

In Europe, O. pumilio is much more diverse due to an increased frequency of smuggling and the resulting offspring of smuggled animals. Smuggling of dart frogs is less common elsewhere, but is still a large problem as it kills large numbers of animals and often degrades or destroys viable habitat.[ citation needed ]

"Blue jeans" color morph Dendrobates pumilio.jpg
"Blue jeans" color morph

Recently, O. pumilio has been exported from Central America again in small numbers from frog farms. Because of this, they have seen a huge increase in numbers in the dart frog community and are regularly available.

Common color morphs in captivity

One example of a color morph is the blue jeans morph. It is most common throughout the species range, but is relatively rare in the United States pet trade. Most of these animals came from imports during the 1990s, or are their descendants. [32] As of 2003, it was observed that this morph could be found throughout Costa Rica, as well as mainland Panama. [4] [24] [25] [26] [27] [28] [29]

Related Research Articles

<span class="mw-page-title-main">Tadpole</span> Larval stage in the life cycle of an amphibian

A tadpole or polliwog is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found in adult amphibians such as a lateral line, gills and swimming tails. As they undergo metamorphosis, they start to develop functional lungs for breathing air, and the diet of tadpoles changes drastically.

<span class="mw-page-title-main">Poison dart frog</span> Family of amphibians

Poison dart frog is the common name of a group of frogs in the family Dendrobatidae which are native to tropical Central and South America. These species are diurnal and often have brightly colored bodies. This bright coloration is correlated with the toxicity of the species, making them aposematic. Some species of the family Dendrobatidae exhibit extremely bright coloration along with high toxicity — a feature derived from their diet of ants, mites and termites— while species which eat a much larger variety of prey have cryptic coloration with minimal to no amount of observed toxicity. Many species of this family are threatened due to human infrastructure encroaching on their habitats.

<span class="mw-page-title-main">Blue poison dart frog</span> Species of amphibian

The blue poison dart frog or blue poison arrow frog is a poison dart frog found in the "forest islands" surrounded by the Sipaliwini Savanna in southern Suriname. Its indigenous Tiriyo name is okopipi. The name "azureus" comes from its azure blue color. While first described as a valid species and usually recognized as such in the past, recent authorities generally treat it as a morph of D. tinctorius, although a few treat it as a subspecies of D. tinctorius or continue to treat it as its own species. To what extent it differs from the blue D. tinctorius in southern Guyana, adjacent Pará (Brazil) and possibly far southwestern Suriname, is also a matter of dispute, and many herpetologists, as well as many people keeping poison dart frogs in captivity, have not distinguished these, with all commonly being identified as "azureus".

<span class="mw-page-title-main">Dyeing poison dart frog</span> Species of amphibian

The dyeing poison dart frog, also known as the cobalt poison frog, tinc, is a species of poison dart frog. It is among the most variably colored and largest species of poison dart frogs, typically reaching snout–vent lengths of about 50 mm (2.0 in). It is distributed in the eastern portion of the Guiana Shield, including parts of French Guiana, Guyana, Suriname, and Brazil. These types of frogs usually like to stay in the ground as it is easier to catch prey. They tend to also stay in bodies of water like rivers or climb trees. Because of this they mostly inhabit near the Amazon Rainforest. To mate they have special rituals and leave their tadpoles in freshwater.

<span class="mw-page-title-main">Harlequin poison frog</span> Species of amphibian

The harlequin poison frog, also known as harlequin poison-dart frog, is a species of poison dart frog endemic to the Chocó region of western Colombia. The frog is normally found on the ground of tropical rain forests, among fallen limbs or leaf litter. Some frogs traditionally classified as Oophaga histrionica were separated as new species in 2018. These are Oophaga anchicayensis, Oophaga andresi and Oophaga solanensis.

<span class="mw-page-title-main">Green and black poison dart frog</span> Species of amphibian

The green-and-black poison dart frog, also known as the green-and-black poison arrow frog and green poison frog, is a brightly-colored member of the order Anura native to southern Central America and Colombia. This species has also been introduced to Oahu, Hawaii in an effort to lower mosquito numbers. It is one of the most variably colored species of poison dart frogs, after D. tinctorius, Adelphobates galactonotus and some Oophaga species. From a conservation standpoint, it is considered to be of least concern by the International Union for Conservation of Nature. Within the reptile and amphibian hobby, D. auratus remains one of the most commonly kept, and easiest-to-breed, dart frogs in captivity, as well as in zoos.

<span class="mw-page-title-main">Golden poison frog</span> Species of amphibian

The golden poison frog, also known as the golden dart frog or golden poison arrow frog, is a poison dart frog endemic from the rainforests of Colombia. The golden poison frog has become endangered due to habitat destruction within its naturally limited range. Despite its small size, this frog is considered to be the most poisonous extant animal species on the planet.

<i>Dendrobates</i> Genus of amphibians

Dendrobates is a genus of poison dart frogs native to Central and South America. It once contained numerous species, but most originally placed in this genus have been split off into other genera such as Adelphobates, Ameerega, Andinobates, Epipedobates, Excidobates, Oophaga, Phyllobates and Ranitomeya, leaving only five large to medium-sized species in the genus Dendrobates. All the other genera used to be grouped in with Dendrobates because it was previously thought that all brightly colored poison dart frogs came from the same ancestor but this has since been proven to be incorrect. Dendrobates and Phyllobates evolved conspicuous coloration from the same common ancestor but not the same as any of the other genera listed above.

<span class="mw-page-title-main">Mimic poison frog</span> Species of amphibian

Ranitomeya imitator, is a species of poison dart frog found in the north-central region of eastern Peru. Its common names include mimic poison frog and poison arrow frog, and it is one of the best known dart frogs. It was discovered in the late 1980s by Rainer Schulte who later split it up into more subspecies; describing each as a specific color morph, and sometimes having a separate behavioral pattern. The acoustics, morphs, and behavior of the species have been extensively researched.

The polkadot poison frog is a species of frogs in the family Dendrobatidae endemic to Panama, where it is known as rana venenosa in Spanish. Its natural habitats are humid lowland and montane forests. It is threatened by habitat loss and is listed by the IUCN as being "critically endangered".

<span class="mw-page-title-main">Lehmann's poison frog</span> Species of amphibian

Lehmann's poison frog or the red-banded poison frog is a species of frogs in the family Dendrobatidae endemic to a small part of western Colombia. Its natural habitats are submontane tropical rainforests. It is threatened by habitat loss and collection for the pet trade, and the IUCN lists it as being "critically endangered". It was named after Colombian conservation biologist Federico Carlos Lehmann.

<span class="mw-page-title-main">Blue-bellied poison frog</span> Species of amphibian

The blue-bellied poison frog or bluebelly poison frog is a species of frog in the family Dendrobatidae.

<i>Oophaga sylvatica</i> Species of amphibian

Oophaga sylvatica, sometimes known as its Spanish name diablito, is a species of frog in the family Dendrobatidae found in Southwestern Colombia and Northwestern Ecuador. Its natural habitat is lowland and submontane rainforest; it can, however, survive in moderately degraded areas, at least in the more humid parts of its range. It is a very common frog in Colombia, but has disappeared from much of its Ecuadorian range. It is threatened by habitat loss (deforestation) and agricultural pollution and sometimes seen in the international pet trade.

<span class="mw-page-title-main">Vicente's poison frog</span> Species of amphibian

Vicente's poison frog is a species of frog in the family Dendrobatidae that is endemic to the Veraguas and Coclé Provinces of central Panama.

Silverstone's poison frog is a species of frog in the family Dendrobatidae endemic to Peru. Its natural habitats are subtropical or tropical moist lowland forests and rivers.

<span class="mw-page-title-main">Golfodulcean poison frog</span> Species of amphibian

The Golfodulcean poison frog or Golfodulcean poison-arrow frog is a species of frog in the family Dendrobatidae endemic to Costa Rica.

<span class="mw-page-title-main">Pumiliotoxin 251D</span> Chemical compound

Pumiliotoxin 251D is a toxic organic compound. It is found in the skin of poison frogs from the genera Dendrobates, Epipedobates, Minyobates, and Phyllobates and toads from the genus Melanophryniscus. Its name comes from the pumiliotoxin family (PTXs) and its molecular mass of 251 daltons. When the toxin enters the bloodstream through cuts in the skin or by ingestion, it can cause hyperactivity, convulsions, cardiac arrest and ultimately death. It is especially toxic to arthropods, even at low concentrations.

<i>Oophaga</i> Genus of amphibians

Oophaga is a genus of poison-dart frogs containing twelve species, many of which were formerly placed in the genus Dendrobates. The frogs are distributed in Central and South America, from Nicaragua south through the El Chocó to northern Ecuador. Their habitats vary with some species being arboreal while other being terrestrial, but the common feature is that their tadpoles are obligate egg feeders. Most species in this genus are seriously threatened and O. speciosa is already extinct.

<i>Adelphobates</i> Genus of amphibians

Adelphobates is a small genus of poison dart frogs. They are found in the central and lower Amazon basin of Peru and Brazil, possibly Bolivia. It was originally erected as a sister group to the Dendrobates and Oophaga genera. The validity of the genus is still being discussed, with the alternative being "Dendrobates galactonotus group" within Dendrobates. One species originally placed in this genus as Adelphobates captivus has since been moved to the genus Excidobates erected in 2008.

<span class="mw-page-title-main">Dendrobatinae</span> Subfamily of amphibians

Dendrobatinae is the main subfamily of frogs in the family Dendrobatidae, the poison dart frogs of Central and South America, found from Nicaragua to the Amazon basin in Brazil.

References

  1. ↑ IUCN SSC Amphibian Specialist Group (2015). "Oophaga pumilio". IUCN Red List of Threatened Species . 2015: e.T55196A3025630. doi: 10.2305/IUCN.UK.2015-4.RLTS.T55196A3025630.en . Retrieved 19 November 2021.
  2. ↑ Frost, Darrel R. (2014). "Oophaga pumilio (Schmidt, 1857)". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 12 September 2014.
  3. 1 2 3 Savage, J. M. 2002. The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago and London.
  4. 1 2 Summers, K.; Cronin, T. W.; Kennedy, T. (2003). "Variation in spectral reflectance among population of Dendrobates pumilio, the strawberry poison frog, in the Bocas del Toro Archipelago, Panama". Journal of Biogeography. 30: 35–53. doi:10.1046/j.1365-2699.2003.00795.x. hdl: 11603/13558 . S2CID   33860218.
  5. ↑ Saporito, Ralph; Donnelly, Maureen; Norton, Roy; Garraffo, Martin; Spande, Thomas; Daly, John (2007). "Oribatid Mites as a Major Dietary Source for Alkaloids in Poison Frogs". Proceedings of the National Academy of Sciences of the United States of America. 104 (21): 8885–8890. Bibcode:2007PNAS..104.8885S. doi: 10.1073/pnas.0702851104 . PMC   1885597 . PMID   17502597.
  6. 1 2 Saporito, Ralph; Garraffo, Martin; Donnelly, Maureen; Edwards, Adam; Longino, John; Daly, John (2004). "Formicine ants: an arthropod aource for the pumiliotoxin alkaloids of dendrobatid poison frogs". Proceedings of the National Academy of Sciences of the United States of America. 101 (21): 8045–8050. Bibcode:2004PNAS..101.8045S. doi: 10.1073/pnas.0402365101 . PMC   419554 . PMID   15128938.
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  8. 1 2 Stynoski, Jennifer L.; Torres-Mendoza, Yaritbel; Sasa-Marin, Mahmood; Saporito, Ralph A. (March 2014). "Evidence of maternal provisioning of alkaloid-based chemical defenses in the strawberry poison frogOophaga pumilio". Ecology. 95 (3): 587–593. doi:10.1890/13-0927.1. hdl: 10669/76946 . ISSN   0012-9658. PMID   24804437. S2CID   4094179.
  9. ↑ Takada, W.; Sakata, T.; Shimano, S.; Enami, Y.; Mori, N.; Nishida, R. & Kuwahara, Y. (2005). "Scheloribatid mites as the source of pumiliotoxins in dendrobatid frogs". Journal of Chemical Ecology. 31 (10): 2403–2415. doi:10.1007/s10886-005-7109-9. PMID   16195851. S2CID   535250.
  10. ↑ Raspotnig, G.; Norton, R. A.; Heethoff, M. (2011). "Oribatid mites and skin alkaloids in poison frogs". Biology Letters. 7 (4): 555–556. doi:10.1098/rsbl.2010.1113. PMC   3130211 . PMID   21345855.
  11. ↑ Staudt, K.; Meneses, O.; S., Mebs; D., Proehl, H. (2010). "Foraging behaviour and territoriality of the strawberry poison frog (Oophaga pumilio) in dependence of the presence of ants". Amphibia-Reptilia. 31 (2): 217–227. doi: 10.1163/156853810791069100 .{{cite journal}}: CS1 maint: multiple names: authors list (link)
  12. ↑ Mebs, D.; Pogoda, W.; Batista, A.; Ponce, M.; Koelher, G.; Kauert, G. (2008). "Variability of alkaloid profiles in Oophaga pumilio (Amphibia: Anura: Dendrobatidae) from Western Panama and Southern Nicaragua". Salamandra. 44 (4): 241–247.
  13. 1 2 Stynoski, J. L.; Torres-Mendoza, Y.; Sasa-Marin, M.; Saporito, R. A. (2014). "Evidence of maternal provisioning of alkaloid-based chemical defenses in the strawberry poison frog Oophaga pumilio" (PDF). Ecology. 95 (3): 587–593. doi: 10.1890/13-0927.1 . PMID   24804437.
  14. ↑ Forsyth, A., Fogden, M., Fogden, P., Wilson, E. O., 2008. Nature of the Rainforest: Costa Rica and Beyond. Cornell University Press. Ithaca, New York, USA.
  15. 1 2 3 Meuche, Ivonne; Linsenmair, K. Eduard; Pröhl, Heike (2012-01-04). "Intrasexual competition, territoriality and acoustic communication in male strawberry poison frogs (Oophaga pumilio)". Behavioral Ecology and Sociobiology. 66 (4): 613–621. doi:10.1007/s00265-011-1309-9. ISSN   0340-5443. S2CID   9380176.
  16. 1 2 3 Grant, T.; Frost, D. R.; Caldwell, J. P.; Gagliardo, R.; Haddad, C. F. B.; Kok, P. J. R.; Means, D. B.; Noonan, B. P.; Schargel, W. E. & Wheeler, W. C. (2006). "Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae)" (PDF). Bulletin of the American Museum of Natural History. 299: 1–262. CiteSeerX   10.1.1.693.8392 . doi:10.1206/0003-0090(2006)299[1:PSODFA]2.0.CO;2. hdl:2246/5803. S2CID   82263880.
  17. 1 2 3 4 Haase, A.; Prohl, H. (2002). "Female activity patterns and aggressiveness in the strawberry poison frog Dendrobates pumilio (Anura: Dendrobatidae)". Amphibia-Reptilia. 23 (2): 129–140. doi: 10.1163/156853802760061778 .
  18. ↑ Meuche, Ivonne; Brusa, Oscar; Linsenmair, K Eduard; Keller, Alexander; Pröhl, Heike (2013-05-20). "Only distance matters – non-choosy females in a poison frog population". Frontiers in Zoology. 10 (1): 29. doi: 10.1186/1742-9994-10-29 . ISSN   1742-9994. PMC   3665588 . PMID   23688371.
  19. ↑ Prohl, H. & Hodl, W. (1999). "Parental investment, potential reproductive rates, and mating system in the strawberry dart-poison frog, Dendrobates pumilio". Behavioral Ecology and Sociobiology. 46 (4): 215–220. doi:10.1007/s002650050612. S2CID   41934325.
  20. ↑ Limerick, S (1980). "Courtship behavior and oviposition of the poison-arrow frog Dendrobates pumilio". Herpetologica. 36: 69–71.
  21. ↑ Zimmermann, H. & Zimmermann, E. (1988). "Etho-Taxonomie und zoogeographische Artengruppenbildung bei Pfeilgiftfröschen (Anura: Dendrobatidae)". Salamandra. 24: 125–160.
  22. ↑ Summers, K.; Weigt, L. A.; Boag, P.; Bermingham, E. (1999). "The evolution of female parental care in poison frogs of the genus Dendrobates: Evidence from mitochondrial DNA sequences". Herpetologica. 55 (2): 254–270.
  23. ↑ Roberts, J. L.; Brown, J. L.; von May, R.; Arizabal, W.; Presar, A.; Symula, R.; Schulte, R. & Summers, K. (2006). "Phylogenetic relationships among poison frogs of the genus Dendrobates (Dendrobatidae): A molecular perspective from increased taxon sampling". Herpetological Journal. 16: 377–385.
  24. 1 2 3 4 Brown, J.; Maan, M.; Cummings, M.; Summers, K. (2010). "Evidence for selection on coloration in a Panamanian poison frog: a coalescent-based approach". Journal of Biogeography. 37 (5): 891–901. doi:10.1111/j.1365-2699.2009.02260.x. S2CID   49231830.
  25. 1 2 Wang, I.; Summers, K. (2010). "Genetic structure is correlated with phenotypic divergence rather than geographic isolation in the highly polymorphic strawberry poison-dart frog". Molecular Ecology. 19 (3): 447–458. doi:10.1111/j.1365-294x.2009.04465.x. PMID   20025652. S2CID   205362447.
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  31. ↑ Rivera-Ordonez, Juana M.; Justin Nowakowski, A.; Manansala, Adrian; Thompson, Michelle E.; Todd, Brian D. (2019-08-02). "Thermal niche variation among individuals of the poison frog, Oophaga pumilio, in forest and converted habitats". Biotropica. 51 (5): 747–756. doi: 10.1111/btp.12691 . ISSN   0006-3606.
  32. ↑ Sheppard, Lisa. "Study finds ethical and illicit sources of poison frogs in the U.S. pet trade". blogs.illinois.edu. Retrieved 2022-11-15.

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