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Overview

Comprehensive Description

Description

Size: 17-24 mm. The skin is relatively smooth, and the "usual" coloration includes a bright red back, some small black spots, black to dark blue hind legs, and a red belly, which occasionally is red and blue, and can vary toward tan and white in some Panamanian localities.

Unusual color variations occur on the small islands off the coast of Panama (in the Bocas del Toro archipelago), including these color combinations: blue above and below, without spots; green above and below with small spots; green above and white below, with small spots; red above and white below, with small spots; and olive green above and yellow below, with black flecks. In each population there is generally only a single color morph, but on the Island of Bastimentos there are different colors in one population. Males have a tan-grayish vocal pouch under the throat, visible when they call to defend their territory. When removed from their territory, they lose the vocal pouch coloration fairly rapidly (Summers et al. 1997).

A Spanish-language species account can be found at the website of Instituto Nacional de Biodiversidad (INBio).In 2011, the genus Dendrobates was subdivided into seven genera, including the new genus Oophaga by Brown et al (2011).

  • Savage, J. M. (2002). The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago and London.
  • Duellman, W. E., and Trueb, L. (1986). Biology of Amphibians. McGraw-Hill, New York.
  • Walls, J. G. (1994). Jewels of the Rainforest: Poison Frogs of the Family Dendrobatidae. J.F.H. Publications, Neptune City, New Jersey.
  • Crothers, L., Gering, E., and Cummings, M. (2011). ''Aposematic signal variation predicts male-male interactions in a polymorphic poison frog.'' Evolution, 65, 599-605.
  • Donnelly, M. A. (1989). ''Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica.'' Journal of Herpetology, 23, 362-367.
  • Donnelly, M. A. (1991). ''Feeding patterns of the Strawberry Poison Frog Dendrobates pumilio (Anura: Dendrobatidae).'' Copeia, 23, 723-730.
  • Graves, B. M. (1999). ''Diel activity patterns of the sympatric poison dart frogs, Dendrobates auratus and D. pumilio, in Costa Rica.'' Journal of Herpetology, 33(3), 375-381.
  • McVey, M. E., Robert, Z. G., Perry, D., and MacDougal, J. (1981). ''Territoriality and homing behavior in the poison-dart frog (Dendrobates pumilio).'' Copeia, 1981(1), 1-8.
  • Summers, K., Bermingham, E., Weigt, L., McCafferty, S. and Dahlstrom, L. (1997). ''Phenotypic and genetic divergence in three species of dart-poison frogs with contrasting parental behaviour.'' Journal of Heredity, 88, 8-13.
  • Summers, K., Cronin, T. W., and Kennedy, T. (2003). ''Variation in spectral reflectance among populations of Dendrobates pumilio, the strawberry poison frog, in the Bocas del Toro Archipelago, Panama.'' Journal of Biogeography, 30, 35-53.
  • Zimmermann, E. (1990). ''Behavioral signals and reproduction modes in the neotropical frog family Dendrobatidae.'' Biology and Physiology of Amphibians. W. Hanke, eds., Fischer, Stuttgart.
  • Brown J.L., Twomey E., Amézquita A., De Souza M.B., Caldwell J.P., Lötters S., Von May R., Melo-Sampaio P.R., Mejía-Vargas D., Perez-Peña P., Pepper M., Poelman E.H., Sanchez-Rodriguez M., and Summers K. (2011). ''A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae).'' Zootaxa, 3083, 1-120.
  • Liebermann, S. and Dock, C. F. (1982). ''Analysis of the leaf litter arthropod fauna of a lowland tropical evergreen forest site.'' Revista de Biología Tropical, 30, 27-34.
  • Pröhl, G. (1995). Territorial- und Paarungsverhalten von Dendrobates pumilio. Diplomarbeit (Master's thesis). Tierärztliche Hochschule Hannover, Germany.
  • Solís, F., Ibáñez, R., Jaramillo, C., Chaves, G., Savage, J., Köhler, G., and Cox, N. 2008. Oophaga pumilio. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. www.iucnredlist.org. Downloaded on 25 June 2011.
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Distribution

Range Description

This species' geographic range is the Atlantic versant, humid lowlands, and premontane slopes in eastern central Nicaragua (0-940m asl) south through the lowlands of Costa Rica and northwestern Panama (including many islands in Bocas del Toro), from 1-495m asl (Savage, 2002).
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Distribution and Habitat

Oophaga pumilio is found in the rainforests of the Caribbean coast of Central America, from Nicaragua to Panama, between sea level and 960 m (Walls 1994).

  • Savage, J. M. (2002). The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago and London.
  • Duellman, W. E., and Trueb, L. (1986). Biology of Amphibians. McGraw-Hill, New York.
  • Walls, J. G. (1994). Jewels of the Rainforest: Poison Frogs of the Family Dendrobatidae. J.F.H. Publications, Neptune City, New Jersey.
  • Crothers, L., Gering, E., and Cummings, M. (2011). ''Aposematic signal variation predicts male-male interactions in a polymorphic poison frog.'' Evolution, 65, 599-605.
  • Donnelly, M. A. (1989). ''Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica.'' Journal of Herpetology, 23, 362-367.
  • Donnelly, M. A. (1991). ''Feeding patterns of the Strawberry Poison Frog Dendrobates pumilio (Anura: Dendrobatidae).'' Copeia, 23, 723-730.
  • Graves, B. M. (1999). ''Diel activity patterns of the sympatric poison dart frogs, Dendrobates auratus and D. pumilio, in Costa Rica.'' Journal of Herpetology, 33(3), 375-381.
  • McVey, M. E., Robert, Z. G., Perry, D., and MacDougal, J. (1981). ''Territoriality and homing behavior in the poison-dart frog (Dendrobates pumilio).'' Copeia, 1981(1), 1-8.
  • Summers, K., Bermingham, E., Weigt, L., McCafferty, S. and Dahlstrom, L. (1997). ''Phenotypic and genetic divergence in three species of dart-poison frogs with contrasting parental behaviour.'' Journal of Heredity, 88, 8-13.
  • Summers, K., Cronin, T. W., and Kennedy, T. (2003). ''Variation in spectral reflectance among populations of Dendrobates pumilio, the strawberry poison frog, in the Bocas del Toro Archipelago, Panama.'' Journal of Biogeography, 30, 35-53.
  • Zimmermann, E. (1990). ''Behavioral signals and reproduction modes in the neotropical frog family Dendrobatidae.'' Biology and Physiology of Amphibians. W. Hanke, eds., Fischer, Stuttgart.
  • Brown J.L., Twomey E., Amézquita A., De Souza M.B., Caldwell J.P., Lötters S., Von May R., Melo-Sampaio P.R., Mejía-Vargas D., Perez-Peña P., Pepper M., Poelman E.H., Sanchez-Rodriguez M., and Summers K. (2011). ''A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae).'' Zootaxa, 3083, 1-120.
  • Liebermann, S. and Dock, C. F. (1982). ''Analysis of the leaf litter arthropod fauna of a lowland tropical evergreen forest site.'' Revista de Biología Tropical, 30, 27-34.
  • Pröhl, G. (1995). Territorial- und Paarungsverhalten von Dendrobates pumilio. Diplomarbeit (Master's thesis). Tierärztliche Hochschule Hannover, Germany.
  • Solís, F., Ibáñez, R., Jaramillo, C., Chaves, G., Savage, J., Köhler, G., and Cox, N. 2008. Oophaga pumilio. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. www.iucnredlist.org. Downloaded on 25 June 2011.
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Geographic Range

Biogeographic Regions: neotropical (Native )

  • Savage, J. 2002. THE AMPHIBIANS AND REPTILES OF COSTA RICA. Chicago: University of Chicago Press.
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Countries

Countries

Costa Rica, Nicaragua, Panama

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Physical Description

Morphology

Physical Description

Oophaga pumilio are slender frogs with bilateral symmetry. They are small frogs measuring 17 to 24 mm in length at adulthood. They feature four, un-webbed digits on each hand and foot, and the body is overall quite compact. These frogs have fairly large, dark eyes set on the sides of the head. The skin of a poison dart frog is very moist which gives them a somewhat glossy appearance in bright light. This species is sexually monomorphic.

They are typically bright red with blue legs although they vary greatly in coloration, and are known as being one of the most polymorphic, aposematic species. However, populations of O. pumilio tend to be the same color. Though typically strawberry red, the dorsal coloration can vary in color from red to blue, yellow, white, green, black or orange. The dorsal surface may also feature dark spots or mottling. Legs are typically darker and have some degree of blackish mottling. Their aposematic coloration has convergently evolved between some separate populations.

Tadpoles are dark brown above with lighter brown undersides and dark spots scattered throughout. They have small, ventral, oral discs with large, serrated beaks. They can reach 16 mm in length.

Range length: 17 to 24 mm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry ; poisonous

Sexual Dimorphism: sexes alike

  • Sandmeier, F. 2001. "Oophaga Pumilio" (On-line). Amphibiaweb. Accessed September 11, 2009 at http://amphibiaweb.org/.
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Diagnostic Description

Identification

Adult

Species description based on Savage (2002).  A small poison frog (males to 24 mm). Throats of females are usually red; throats of males have a pale spot in the center.

Dorsal

The general coloration of Oophaga pumilio consists of a red body (occassionally blue) with red, blue, purple, or black hind legs. The lower forearms sometimes also have these colors. In some individuals, the dorsum and ventral surface are flecked with black. The skin of the dorsum is smooth. On the islands of Bocas del Toro in Panama, this species exhibits an incredible amount of color and pattern polymorphism (see Evolution below).

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
A diurnal and mostly terrestrial frog of humid lowland and premontane forest, cacao plantations, and abandoned forest clearings. Males appear to be fiercely territorial; individual territories have been estimated at 2.5 m2 (Donnelly, 1983). Observations concerning mating behaviour suggest that some O. pumilio are at times polygynous (McVey et al., 1981; Donnelly, 1989; Zimmermann and Zimmermann, 1994). Females lay a clutch of three to nine eggs in moist leaf-litter; clutch sizes in captive specimens of six to 16 eggs have been recorded (Limerick, 1980; Silverstone, 1975). There appears to be no information on the number of clutches laid annually. O. pumilio eggs hatch approximately seven days after oviposition, adults then carry the developed tadpoles from the forest floor to water filled bromeliads (Limerick, 1980). O. pumilio tadpoles have a very specialized oophagous diet, feeding solely on food eggs supplied by the female (Heselhaus, 1992; McVey et al.; 1981, Zimmermann and Zimmermann, 1994). There is little available information on wild larval development; Heselhaus (1992) reports that captive tadpoles fed an artificial diet ‘grow slowly, taking four to six months, a third longer than with natural feeding, to reach metamorphosis’. Sexual maturity is reached at a minimum size of 19mm (approximately 10 months). There are few data on longevity; Donnelly (1983) concluded that the population at Finca La Selva, Costa Rica was mostly comprised of ‘long-lived’ adults; Zimmermann and Zimmermann (1994) gave longevity of 4 years for captive O. pumilio.

Systems
  • Terrestrial
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Oophaga pumilio typically live in rainforest habitats and also live in cacao and banana groves, but not banana plantations. Unlike some other Dendrobates, O. pumilio tend to live near the forest floor in leaf litter but they frequently climb trees and vines. Females oviposit on land, but will transport each tadpole to its own water-filled bromeliad to complete metamorphosis. Thus, these frogs require moist, terrestrial habitats with abundant water-filled plants for successful reproduction.

In Nicaragua, Oophaga pumilio live between 0 to 940 m above sea level and in Panama between 0 to 495 m. Costa Rican populations inhabit similar elevations.

Range elevation: 0 to 940 m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: rainforest

  • 2003. Poison frogs. Pp. 197-200 in M Hutchins, W Duellman, N Schlager, eds. Grzimek's Animal Life Encyclopedia, Vol. 6, 2nd Edition. Farmington Hills, MI: Gale Group.
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Habitat

Lowland and premotane forest to 940 m.

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Trophic Strategy

Food Habits

Strawberry poison frogs feed by "wide foraging" in which frogs use their tongues to catch large numbers of small prey. All of their diet consists of small arthropods, some of which (particularly formicine ants) provide toxins which the frogs can excrete through their skin. Oophaga pumilio consume mostly ants but mites also make up a significant portion of their diet. The tadpoles are oophages, so called because they eat unfertilized eggs either by cutting a hole and sucking the contents out or in the case of larger tadpoles, consume the egg whole. Strawberry poison frogs will typically eat from 7 prey per hour (for juveniles) to 14 prey per hour (adults).

Animal Foods: eggs; insects; terrestrial non-insect arthropods

Primary Diet: carnivore (Insectivore )

  • Donnelly, M. 1991. Feeding Patterns of the Strawberry Poison Frog, Dendrobates pumilio (Anura: Dendrobatidae). Copeia, 3: 723-730.
  • Grant, T., D. Frost, J. Caldwell, R. Gagliardo, C. Haddad, P. Kok, D. Means, B. Noonan, W. Schargel, W. Wheeler. 2006. Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia, Athesphatanura, Dendrobatidae). Bulletin of the American Museum of Natural History, 299: 1-266. Accessed November 18, 2009 at http://hdl.handle.net.login.ezproxy.library.ualberta.ca/2246/5803.
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Diet

Diet

The diet consists of a variety of small arthropods, including ants (Savage 2002).

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Associations

Ecosystem Roles

Dendrobates pumilo fills a niche of ant and mite population control. They play a significant role in pest control for local plant life. Though the poisonous adults are rarely preyed upon, defenseless tadpoles are likely a food source for predators.

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Predation

Strawberry poison dart frogs have few major predators because their aposematic coloration warns predators that it is very poisonous. However, night ground snakes are immune to the toxins of Oophaga pumilio. Tadpoles are often consumed because their poison glands are underdeveloped.

Known Predators:

Anti-predator Adaptations: aposematic

  • Myers, C., J. Daly, B. Malkin. 1978. A dangerously toxic new frog (Phyllobates) used by the Emberá Indians of western Colombia, with discussion of blowgun fabrication and dart poisoning.. Bulletin of the American Museum of natural history, 161 (2): 307–365.
  • Wang, I., H. Shaffer. 2008. Rapid Color Evolution in an aposematic Species: A phylogentic Analysis of Color Variation in the Strikingly Polymorphic Stawberry Poison-dart Frog. Evolution, 62/11: 2742-2759.
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General Ecology

Ecology

Ecology

Oophaga pumilio is commonly encountered hopping along in leaf litter (Savage 2002). Their coloration is aposematic, warning predators of the toxic alkaloids exuded from their skin (Saporito et al 2007).

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Life History and Behavior

Behavior

Communication and Perception

The calls of Oophaga pumilio consist of a series of short chirps lasting 5 to 32 seconds with 5 to 9 notes per second most often used for territorial announcement by males and for males and females to announce availability for mating. During mating, males and females will engage in mutual tactile stimulation, but unlike many amphibians they do not engage in amplexus.

It has been recently discovered that Oophaga pumilio feature eyes with unique rods and cones that enable them to differentiate between the many conspecific color morphs. Females rely heavily on this ability to select mates of the same color morph.

Communication Channels: visual ; tactile ; acoustic

Perception Channels: visual ; tactile ; acoustic

  • Forester, D., A. Wisnieski. 1991. The Significance of airbone Olfactory Cues to the Recognition of Home area by the Poison Dart Frog. Journal fo Herpetology, 25/4: 502-504.
  • Siddiqi, A., T. Cronin, E. Loew, M. Vorobyev, K. Summers. 2004. Interspecific and intraspecific views of color signals in the strawberry poison frog Dendrobates pumilio. The Journal of Experimental Biology, 207: 2471-2485.
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Behaviour

Call

A series of harsh "chirps" Myers et al 1995).

Behavior and communication

Males are territorial (Savage 2002). They may engage in extensive wrestling bouts with intruding males if necessary (Savage 2002). After eggs are deposited in a male's territory, he guards them and keeps them hydrated (Weygoldt 1980, Brust 1990, Brust 1993). When the eggs hatch, the female returns and carries her offspring to water-filled phytotelmata. She feeds her larvae unfertilized eggs on a daily basis (Weygoldt 1980, Brust 1990, Brust 1993). When she arrives, she places her rear-end in the water, and the tadpoles touch her and vibrate to signal their presence (Weygoldt 1980, Brust 1990, Brust 1993). The tadpoles do not survive witout the eggs supplied by their mothers (Brust 1990, Brust 1993).

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Life Cycle

Development

Strawberry dart frog zygotes are formed when a male fertilizes eggs from a female. The zygotes divide until they become tadpoles at which point they swim onto their mother’s back and are individually taken to axils of bromeliads. These plants have small pools of water which the tadpoles complete metamorphosis in. The tadpoles require a food source within 3 days of being re-located or they will starve. Females provide unfertilized eggs for the young to consume. Tadpoles begin metamorphosis after growing to 11 mm in length, and the process takes 6 to 8 weeks to complete.

  • Duellman, W., L. Trueb. 1986. Biology of Amphibians. New York, NY, USA: McGraw-Hill Book Company.
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Life History

Breeding season

Breeding occurs year-round (Savage 2002). If a female approaches a male, they engage in a series of courtship behaviors prior to amplexus (Savage 2002). Amplexus is unusual--the male and female face away from one another and place their vents together (Savage 2002).

Egg

Females typically lay between 3 and 5 eggs, and may lay another clutch as often as every week (Savage 2002). Eggs hatch in one week (Limerick 1980).

Tadpole

The tadpole is small and very dark brown, with a moderate tail with almost no tail fins (Savage 2002). The tail is lighter than the body, with some darker pigmentation (Savage 2002). Metamorphosis occurs after 43-52 days (Savage 2002).

Metamorph juvenile

Metamorphs are deep red (Savage 2002).

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Life Expectancy

Lifespan/Longevity

Information regarding the life span of strawberry poison frogs is rare, but closely related Dendrobates auratus are known to live up to 17 years in captivity.

Typical lifespan

Status: captivity:
17 (high) years.

  • 1970. Arrow-poison Frogs. Pp. 94 in M Burton, R Burton, eds. The international wildlife encyclopedia, Vol. 1, Third Edition. Tarrytown NY: Marshall Cavendish Corporation.
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Lifespan, longevity, and ageing

Maximum longevity: 7.5 years (captivity)
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Reproduction

There is a certain amount of sexual selection for Oophaga pumilio. Females tend to choose males with similar colored dorsal sides which usually means that they are from the same population. Males tend to have a better chance of mating if their territory is larger, therefore they will compete by wrestling for large territories. Perch height and calling frequency also influence the likelihood of male mating. This may not necessarily be a matter of sexual selection as much as it is good advertising.

In general, O. pumilio are polygynandrous, with both males and females mating with different partners multiple times per breeding season. The female comes to the male by following his call and after a brief period of mutual tactile stimulation the couple breed by egg laying and fertilizing. They assume a vent to vent posture facing away from each other. The whole breeding process takes between 10 and 180 minutes.

There is evidence that these frogs may have some degree of color-vision. Their eyes contain structures required to see different wavelengths, and thus may be able to differentiate between color morphs of their species. This may play a role in sexual selection, but more research must be done.

Mating System: polygynandrous (promiscuous)

Oophaga pumilio may breed throughout the year, but only under favorable, moist conditions. Females generally do not ovulate during drier periods and males are less likely to call at this time. After mating, females lay a clutch of 3 to 5 fertilized eggs in moist leaf litter. Under the male's care, the eggs develop into tadpoles after 10 to 14 days, at which time the female take sole responsibility for the young. After being transported to individual, water-filled bromeliads, tadpoles metamporphosize into adults after 43 to 52 days. Both male and female Oophaga pumilio reach sexual maturity at 10 months of age.

Breeding interval: Under optimal, moist conditions, Oophaga pumilio may breed year round.

Breeding season: The breeding season for Oophaga pumilio may last 8 to 10 months.

Range number of offspring: 3 to 5.

Average number of offspring: 4.6.

Range time to hatching: 10 to 14 days.

Average age at sexual or reproductive maturity (female): 10 months.

Average age at sexual or reproductive maturity (male): 10 months.

Key Reproductive Features: iteroparous ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (External ); oviparous

Oophaga pumilio select terrestrial locations to lay eggs, which then require significant additional moisture to avoid dessication. To ensure the clutch is moist, a male will urinate on the eggs on a daily basis. The male also defends the egg clutch, removes fungus, and rotates the eggs before they become tadpoles.

When the eggs hatch after 10 to 14 days, the female transports tadpoles from one to four at a time to a watery hollow in the vegetation (often a water-filled bromeliad). One tadpole is deposited in each location, because they will consume the smaller of their siblings if they are left to grow together. The female strawberry poison frogs must provide food for each tadpole within 3 days of transport or they will starve. Afterwards, she will make morning, daily visits to feed each tadpole 1 to 5 unfertilized eggs. A female will back into the bromeliad and submerge her vent into the pool of water, where the tadpole will stiffen its body and vibrate to solicit eggs. Females will only feed their own tadpoles even if solicited by others.

Parental Investment: male parental care ; female parental care ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Male, Protecting: Male); pre-weaning/fledging (Provisioning: Female, Protecting: Male); pre-independence (Provisioning: Female, Protecting: Male)

  • Savage, J. 2002. THE AMPHIBIANS AND REPTILES OF COSTA RICA. Chicago: University of Chicago Press.
  • 2003. Poison frogs. Pp. 197-200 in M Hutchins, W Duellman, N Schlager, eds. Grzimek's Animal Life Encyclopedia, Vol. 6, 2nd Edition. Farmington Hills, MI: Gale Group.
  • Cohen, N., R. Stebbins. 1995. A natural History of Amphibians. Princeton, New Jersey: Princeton University Press.
  • Duellman, W., L. Trueb. 1986. Biology of Amphibians. New York, NY, USA: McGraw-Hill Book Company.
  • Maan, M., M. Cummings. 2008. Female Preferences for Aposematic Signal Components in a Polymorphic Poison Frog. Evolution, 62/9: 2334/2345.
  • Proehl, H., W. Hoedl. 1999. Parental Investment, Potential reproductive rates, adn mating system in the strawberry dart-poison frog Dendrobates Pumilio. Behav Ecol Sociobiol, 46: 215-220.
  • Sandmeier, F. 2001. "Oophaga Pumilio" (On-line). Amphibiaweb. Accessed September 11, 2009 at http://amphibiaweb.org/.
  • Weiskittle, J. 2008. "Dendrobates pumilio: strawberry poison dart frog" (On-line). Accessed October 10, 2009 at http://jrscience.wcp.muohio.edu/fieldcourses02/PapersCostaRicaArticles/Dendrobatespumilio.strawb.html.
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Evolution and Systematics

Evolution

Evolotion

Daly and Myers (1967) and Myers and Daly (1983) initially characterized the great variation in color and patterning in Oophaga pumilio in the Bocas del Toro archipelago. This has sparked an incredible amount of research on sexual selection, behavior, and genetics in this species (Prohl et al 2007, Prohl 2002, Rudh et al 2007, Maan and Cummings 2008, Siddiqi et al 2004, Summers et al 1997, Summers and Kennedy 2004, Tazzyman and Iwasa 2010, Wang and Summers 2009, Wang and Summers 2010).

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Physiology and Cell Biology

Physiology

Physiology

Oophaga pumilio has become a model system for understanding the relationship between toxicity and diet in poison frogs (Saporito et al 2006, Saporito 2007, Saporito et al 2010)

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Cell Biology

Karyotype

Karyotype

2N = 20 (Bogart 1991).

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Oophaga pumilio

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 47 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

ACCCTGTATCTAGTATTCGGAGCATGAGCCGGGATGGTCGGCACAGCTCTA---AGCCTCCTTATCCGAGCAGAATTAAGTCAACCTGGGGCCCTTTTAGGCGAT---GACCAAATCTACAATGTGATCGTCACCGCCCATGCCTTCGTAATAATTTTTTTTATAGTTATACCTATTCTAATTGGAGGCTTTGGGAACTGACTCGTCCCCTTAATA---ATTGGTGCCCCTGATATAGCTTTCCCTCGAATGAATAATATAAGCTTCTGACTTCTTCCTCCATCTTTTCTTCTCCTACTGGCTTCTGCCGGAGTAGAAGCTGGGGCGGGGACAGGATGAACTGTATACCCCCCACTTGCAGGAAATCTAGCTCACGCTGGCCCTTCAGTAGACCTAATC---ATCTTTTCTCTTCACCTAGCTGGCGTTTCATCCATCCTTGGAGCAATTAACTTTATTACCACAACCCTAAACATGAAACCTCCTTCTCTTTCACAGTATCAAACCCCCCTGTTTGTGTGGTCGGTCTTAATTACAGCAGTCCTGCTTCTTCTATCACTACCAGTGTTAGCTGCA---GGCATCACTATACTACTAACTGATCGAAATCTTAATACAACCTTTTTTGACCCCTCCGGAGGAGGCGACCCA---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------GTC
-- end --

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Statistics of barcoding coverage: Oophaga pumilio

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 46
Specimens with Barcodes: 57
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2010

Assessor/s
Frank Solís, Roberto Ibáñez, César Jaramillo, Gerardo Chaves, Jay Savage, Gunther Köhler, Neil Cox

Reviewer/s
Global Amphibian Assessment Coordinating Team (Simon Stuart, Janice Chanson, Neil Cox and Bruce Young)

Contributor/s

Justification
Listed as Least Concern in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category.

History
  • 2004
    Least Concern
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Climate change as well as deforestation in the habitat of O. pumilio could have drastic affects as the tadpole rearing process is extremely habitat specific. Strawberry poison frogs are very popular in the pet trade and populations may be threatened by illegal capture. Despite these potential threats, population numbers are currently high and they are considered least concern by the IUCN Red List.

CITES: no special status

IUCN Red List of Threatened Species: least concern

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Population

Population
This is a common species throughout its range. The species exhibits significant colour and pattern polymorphism especially among populations in the Bocas del Toro archipelago of Panama (Summers et al., 1997). Populations in forest habitat at La Selva, Costa Rica, seem to have experienced a decline (Whitfield et al., 2007).

Population Trend
Decreasing
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Life History, Abundance, Activity, and Special Behaviors

More males call during the rainy season and female do not ovulate during the drier period. The male call is described as a low buzz or ticking note call, and is used to attract females and as a territorial advertisement call (Walls 1994). There are 4 different calls (Zimmermann 1990). The call most heard is used for territorial defense in the morning, between 8 and 10 am. Males defend territories, approximately 3 m apart from other males (Walls 1994).

The female approaches the male to initiate breeding. It is the role of the male to tend to the terrestrial clutch of eggs (3-17 eggs), and to keep them moist by periodically emptying their bladders on the eggs (hydric brooding) until they hatch (after 5-15 days). Males may protect more than one clutch at a time. The female then carries the individual tadpoles (1-2) to separate, water filled leaf axils of bromeliads or other plants. If two tadpoles are brought to one bromeliad only one will survive. The female lays unfertilized eggs in with the tadpoles to serve as a food source, and she is thus able to maintain as many as six tadpoles through metamorphosis (Duellman and Trueb 1986). Tadpoles are obligately oophagous and must receive an egg meal within 3 days of being placed in a bromeliad water pool in order to survive. They metamorphosize when they reach approximately 11 mm long (Walls 1994). Males have been observed eating the eggs or carrying the tadpoles of unattended clutches to bromeliad water pools where they will die since the female will not be able to feed them (Duellman and Trueb 1986). Metamorphosis is complete in 6-8 weeks.

Adults of this species primarily consume ants.

These frogs are brightly colored and toxic, and the bright coloration has multiple functions. First, it functions in defense: the coloration warns predators that these frogs are not palatable (aposematic coloration). Second, it functions in mate choice: female strawberry poison dart frogs have been shown to preferentially choose males based on brightness and color. Now a third role has been shown in a recent Evolution paper: male-male competition (Crothers et al. 2011). Brighter males defend their territory more strongly, preferentially approaching brighter intruders rather than dull-colored ones, initiating confrontation more quickly, and directing more calls to brighter rivals. This may be particularly relevant in highly polymorphic populations of O. pumilio such as those of the Bocas del Toro Archipelago. In species where males provide some parental care (hydric brooding of the egg clutch, in the case of O. pumilio), sexually selected traits such as male color brightness are predicted to function as indicators of individual condition/quality.

  • Savage, J. M. (2002). The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago and London.
  • Duellman, W. E., and Trueb, L. (1986). Biology of Amphibians. McGraw-Hill, New York.
  • Walls, J. G. (1994). Jewels of the Rainforest: Poison Frogs of the Family Dendrobatidae. J.F.H. Publications, Neptune City, New Jersey.
  • Crothers, L., Gering, E., and Cummings, M. (2011). ''Aposematic signal variation predicts male-male interactions in a polymorphic poison frog.'' Evolution, 65, 599-605.
  • Donnelly, M. A. (1989). ''Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica.'' Journal of Herpetology, 23, 362-367.
  • Donnelly, M. A. (1991). ''Feeding patterns of the Strawberry Poison Frog Dendrobates pumilio (Anura: Dendrobatidae).'' Copeia, 23, 723-730.
  • Graves, B. M. (1999). ''Diel activity patterns of the sympatric poison dart frogs, Dendrobates auratus and D. pumilio, in Costa Rica.'' Journal of Herpetology, 33(3), 375-381.
  • McVey, M. E., Robert, Z. G., Perry, D., and MacDougal, J. (1981). ''Territoriality and homing behavior in the poison-dart frog (Dendrobates pumilio).'' Copeia, 1981(1), 1-8.
  • Summers, K., Bermingham, E., Weigt, L., McCafferty, S. and Dahlstrom, L. (1997). ''Phenotypic and genetic divergence in three species of dart-poison frogs with contrasting parental behaviour.'' Journal of Heredity, 88, 8-13.
  • Summers, K., Cronin, T. W., and Kennedy, T. (2003). ''Variation in spectral reflectance among populations of Dendrobates pumilio, the strawberry poison frog, in the Bocas del Toro Archipelago, Panama.'' Journal of Biogeography, 30, 35-53.
  • Zimmermann, E. (1990). ''Behavioral signals and reproduction modes in the neotropical frog family Dendrobatidae.'' Biology and Physiology of Amphibians. W. Hanke, eds., Fischer, Stuttgart.
  • Brown J.L., Twomey E., Amézquita A., De Souza M.B., Caldwell J.P., Lötters S., Von May R., Melo-Sampaio P.R., Mejía-Vargas D., Perez-Peña P., Pepper M., Poelman E.H., Sanchez-Rodriguez M., and Summers K. (2011). ''A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae).'' Zootaxa, 3083, 1-120.
  • Liebermann, S. and Dock, C. F. (1982). ''Analysis of the leaf litter arthropod fauna of a lowland tropical evergreen forest site.'' Revista de Biología Tropical, 30, 27-34.
  • Pröhl, G. (1995). Territorial- und Paarungsverhalten von Dendrobates pumilio. Diplomarbeit (Master's thesis). Tierärztliche Hochschule Hannover, Germany.
  • Solís, F., Ibáñez, R., Jaramillo, C., Chaves, G., Savage, J., Köhler, G., and Cox, N. 2008. Oophaga pumilio. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. www.iucnredlist.org. Downloaded on 25 June 2011.
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Threats

Major Threats
Habitat loss and over-collection for the pet trade are the principal threats to this species. It is believed that the species is currently being unsustainably collected, and because of the apparently low fecundity of this species, the possibility exists that over-harvesting might lead to localized population declines. Distinct island forms are particularly susceptible to both over-collection, and the development of islands for tourism purposes. The great majority of reported trade over the period 1991 to 1996 was in live animals, presumably by the herpetological pet market. The largest overall exporter of O. pumilio between 1991 and 1996 was Nicaragua (>95% of exports); the USA consistently accounted for over 80% of recorded live O. pumilio imports from Nicaragua during this period. Current export levels from range states are not known. Museum specimens of this species have been found to have chytrid fungi, the current impact of this pathogen on O. pumilio is not known. At La Selva, declines seem to be driven by climate-driven reductions in quantity of standing leaf litter (Whitfield et al., 2007).
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Life History, Abundance, Activity, and Special Behaviors

It is generally common throughout its range. Habitat loss and overcollection for the pet trade are problems for some populaions. Tourism (e.g., www.redfrogbeachclub.com) also affects some populations. It is found within several protected areas, including the Finca La Selva Biological Reserve (Costa Rica) and Isla Bastimentos National Marine Park (Bocas del Toro Archipelago, Panama), and may occur within other protected areas, particularly in Costa Rica. Export data from 1991-1996 showed that the majority of specimens (>95%) came from Nicaragua, which has established a CITES 2001 export quota of 3,450 specimens for O. pumilio (Solís et al. 2008).

  • Savage, J. M. (2002). The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago and London.
  • Duellman, W. E., and Trueb, L. (1986). Biology of Amphibians. McGraw-Hill, New York.
  • Walls, J. G. (1994). Jewels of the Rainforest: Poison Frogs of the Family Dendrobatidae. J.F.H. Publications, Neptune City, New Jersey.
  • Crothers, L., Gering, E., and Cummings, M. (2011). ''Aposematic signal variation predicts male-male interactions in a polymorphic poison frog.'' Evolution, 65, 599-605.
  • Donnelly, M. A. (1989). ''Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica.'' Journal of Herpetology, 23, 362-367.
  • Donnelly, M. A. (1991). ''Feeding patterns of the Strawberry Poison Frog Dendrobates pumilio (Anura: Dendrobatidae).'' Copeia, 23, 723-730.
  • Graves, B. M. (1999). ''Diel activity patterns of the sympatric poison dart frogs, Dendrobates auratus and D. pumilio, in Costa Rica.'' Journal of Herpetology, 33(3), 375-381.
  • McVey, M. E., Robert, Z. G., Perry, D., and MacDougal, J. (1981). ''Territoriality and homing behavior in the poison-dart frog (Dendrobates pumilio).'' Copeia, 1981(1), 1-8.
  • Summers, K., Bermingham, E., Weigt, L., McCafferty, S. and Dahlstrom, L. (1997). ''Phenotypic and genetic divergence in three species of dart-poison frogs with contrasting parental behaviour.'' Journal of Heredity, 88, 8-13.
  • Summers, K., Cronin, T. W., and Kennedy, T. (2003). ''Variation in spectral reflectance among populations of Dendrobates pumilio, the strawberry poison frog, in the Bocas del Toro Archipelago, Panama.'' Journal of Biogeography, 30, 35-53.
  • Zimmermann, E. (1990). ''Behavioral signals and reproduction modes in the neotropical frog family Dendrobatidae.'' Biology and Physiology of Amphibians. W. Hanke, eds., Fischer, Stuttgart.
  • Brown J.L., Twomey E., Amézquita A., De Souza M.B., Caldwell J.P., Lötters S., Von May R., Melo-Sampaio P.R., Mejía-Vargas D., Perez-Peña P., Pepper M., Poelman E.H., Sanchez-Rodriguez M., and Summers K. (2011). ''A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae).'' Zootaxa, 3083, 1-120.
  • Liebermann, S. and Dock, C. F. (1982). ''Analysis of the leaf litter arthropod fauna of a lowland tropical evergreen forest site.'' Revista de Biología Tropical, 30, 27-34.
  • Pröhl, G. (1995). Territorial- und Paarungsverhalten von Dendrobates pumilio. Diplomarbeit (Master's thesis). Tierärztliche Hochschule Hannover, Germany.
  • Solís, F., Ibáñez, R., Jaramillo, C., Chaves, G., Savage, J., Köhler, G., and Cox, N. 2008. Oophaga pumilio. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. www.iucnredlist.org. Downloaded on 25 June 2011.
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Management

Conservation Actions

Conservation Actions
This species is listed on Appendix II of CITES and is found in several protected areas within its range. A well-studied population of O. pumilio is present in the Finca La Selva Biological Reserve, northeastern Costa Rica (Donnelly, 1989; Limerick, 1980; McVey, 1981; Pröhl, 1997); populations are suspected to occur in Braulio Carrillo National Park, Corcovado National Park, and Tortuguero National Park, Costa Rica. Within Panama the species is present in a number of protected areas including Isla Bastimentos National Marine Park within the Bocas del Toro archipelago (Summers et al., 1997). Nicaragua has established a CITES 2001 export quota of 3,450 specimens for this species.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

There is no negative impact to humans from Dendrbates pumilio.

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Economic Importance for Humans: Positive

Oophaga pumilio are sometimes captured (illegally) and sold as pets. As with other species of the genus Dendrobates, O. pumilio secrete a very powerful alkaloid poison which may offer significant medical prospects, but no major breakthroughs have been made. Native human populations use the powerful skin toxins to lace arrow heads, which significantly aids in hunting. The extensive variety of color morphs displayed by Oophaga pumilio make them an ideal species for research to gain better understanding of aposematic coloration.

Positive Impacts: pet trade ; research and education

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Risks

Relation to Humans

This frog can be found in gardens. The species is often smuggled for the pet trade.

  • Savage, J. M. (2002). The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago and London.
  • Duellman, W. E., and Trueb, L. (1986). Biology of Amphibians. McGraw-Hill, New York.
  • Walls, J. G. (1994). Jewels of the Rainforest: Poison Frogs of the Family Dendrobatidae. J.F.H. Publications, Neptune City, New Jersey.
  • Crothers, L., Gering, E., and Cummings, M. (2011). ''Aposematic signal variation predicts male-male interactions in a polymorphic poison frog.'' Evolution, 65, 599-605.
  • Donnelly, M. A. (1989). ''Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica.'' Journal of Herpetology, 23, 362-367.
  • Donnelly, M. A. (1991). ''Feeding patterns of the Strawberry Poison Frog Dendrobates pumilio (Anura: Dendrobatidae).'' Copeia, 23, 723-730.
  • Graves, B. M. (1999). ''Diel activity patterns of the sympatric poison dart frogs, Dendrobates auratus and D. pumilio, in Costa Rica.'' Journal of Herpetology, 33(3), 375-381.
  • McVey, M. E., Robert, Z. G., Perry, D., and MacDougal, J. (1981). ''Territoriality and homing behavior in the poison-dart frog (Dendrobates pumilio).'' Copeia, 1981(1), 1-8.
  • Summers, K., Bermingham, E., Weigt, L., McCafferty, S. and Dahlstrom, L. (1997). ''Phenotypic and genetic divergence in three species of dart-poison frogs with contrasting parental behaviour.'' Journal of Heredity, 88, 8-13.
  • Summers, K., Cronin, T. W., and Kennedy, T. (2003). ''Variation in spectral reflectance among populations of Dendrobates pumilio, the strawberry poison frog, in the Bocas del Toro Archipelago, Panama.'' Journal of Biogeography, 30, 35-53.
  • Zimmermann, E. (1990). ''Behavioral signals and reproduction modes in the neotropical frog family Dendrobatidae.'' Biology and Physiology of Amphibians. W. Hanke, eds., Fischer, Stuttgart.
  • Brown J.L., Twomey E., Amézquita A., De Souza M.B., Caldwell J.P., Lötters S., Von May R., Melo-Sampaio P.R., Mejía-Vargas D., Perez-Peña P., Pepper M., Poelman E.H., Sanchez-Rodriguez M., and Summers K. (2011). ''A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae).'' Zootaxa, 3083, 1-120.
  • Liebermann, S. and Dock, C. F. (1982). ''Analysis of the leaf litter arthropod fauna of a lowland tropical evergreen forest site.'' Revista de Biología Tropical, 30, 27-34.
  • Pröhl, G. (1995). Territorial- und Paarungsverhalten von Dendrobates pumilio. Diplomarbeit (Master's thesis). Tierärztliche Hochschule Hannover, Germany.
  • Solís, F., Ibáñez, R., Jaramillo, C., Chaves, G., Savage, J., Köhler, G., and Cox, N. 2008. Oophaga pumilio. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. www.iucnredlist.org. Downloaded on 25 June 2011.
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Wikipedia

Strawberry poison-dart frog

The strawberry poison frog or strawberry poison-dart frog (Oophaga pumilio or Dendrobates pumilio) is a species of small amphibian poison dart frog found in Central America. 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.[1] 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.[2] O. pumilio, while not the most poisonous of the dendrobatids, is the most toxic member of its genus.[citation needed]

Diet[edit]

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. Alkaloid toxins are organic in nature and contain nitrogenous bases that react with carbon and hydrogen groups.[3] Pumiliotoxin 251D, is the specialized toxin that is sequestered by this species of frog.1 This toxin has a negative stimulating effect on cardiac function and is a severe disruptor of the sodium potassium ion channels within cells.1 Upon ingestion of Pumiliotoxin 251D, organisms preying on O. pumilio experience convulsions, paralysis, and death.1

Oribatida mites belonging to the glandulate suborder Brachypylina are an important origin of pumiliotoxins in O. pumilio.[4] Hexane-extraction techniques indicate presence of alkaloid toxins in Brachypylina.2 Toxins appear to be biosynthesized in adult mites, as nymph and larval stages of the insect do not carry the toxins.2 Experimental analysis of this species of mite show alkaloid toxins are found almost exclusively in the opisthonotal glands of the Scheloribatidae, mite.[5] Oil glands of the mite contain the toxins and are then released internally as the amphibian digests the insect.

O. pumilio

O. pumilio [6]




O. pumilio also attributes its cutaneous toxicity from its rich diet of the Formicine ant. Species of the Formicine genus Brachymyrmex, contain pumiliotoxins which the frogs incorporate and accumulate poison from.[7]

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 intraspecific habitats.[8] Research and physical analysis reveal that maternally derived alkaloids exist in young tadpoles.[9] 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.[10] Tadpoles lacking the obligate nutritive egg diet do not contain the alkaloid.6 This step is crucial for the tadpoles to sequester the alkaloid from their mother; without such, young tadpoles become susceptible to predation from nearby arthropods and other frogs.


Behavior[edit]

Oophaga pumilio is diurnal and primarily terrestrial, and can often be found in leaf litter in both forested and disturbed areas. Though brightly colored and toxic, these frogs are relatively small, growing to approximately 17.5–22 mm in standard length.[1] Males are extremely territorial, guarding small territories;[11] Females and juveniles are far more sociable.

Male advertisement call

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Reproduction and parental care[edit]

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.[12] The strawberry poison frog has dual parental care.[13] The males defend and water the nests, and the females feed the oophagous tadpoles their unfertilized eggs. Although both sexes contribute to parental care, females invest more heavily in terms of energy expenditure, time investment, and loss of potential reproduction.[13] 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.[13] The males, though, contribute 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.[13] 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.[14]

The la gruta morph from Colón Province, Panama

Unlike many frog species, amplexus is absent in O. pumilio, with mating individuals instead exhibiting a distinct vent-to-vent position in which the female lays eggs and the male fertilizes them.[15] 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.[16] 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. [17] 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[edit]

Oophaga pumilio belongs to the genus Oophaga,[12] although the name Dendrobates pumilio is still most commonly used. There is evidence that the species of the Oophaga genus (previously classified as the "female parental care group" of Dendrobates[18]) 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).[19] Oophaga pumilio is believed to be most closely related to Oophaga arborea.[20]

Captivity[edit]

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$30 each. However, these shipments have since stopped, and O. pumilio is much less common and available in reduced diversity. In Europe, O. pumilio is much more diverse and available due to an increased frequency of smuggling and the resulting offspring of smuggled animals. Smuggling of dart frogs is less common elsewhere, but still problematic as it kills large numbers of animals and frequently degrades or destroys viable habitat.

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[edit]

"Blue jeans" color morph

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.[citation needed] This morph can be found throughout Costa Rica, as well as mainland Panama.[2]

Bastimentos color morph

References[edit]

  1. ^ a b Savage, J. M. 2002. The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago and London.
  2. ^ a b Summers, K., Cronin, T.W. and 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.
  3. ^ Vandendriessche,T., Abdel-Mottaleb, Y., Maertens, C., Cuypers, E., Sudau, A., Nubbemeyer, U., Mebs, D., Tytgat, J. March 2008.Modulation of Voltage-gated Na+ and K+ Channels by Pumiliotoxin 251 D: A "Joint Venture" Alkaloid from Arthropods and Amphibians. Toxicon, Vol. 51, No. 3:334-344
  4. ^ Takada, W., Sakata, T., Shimano, S., Enami, Y., Mori, N., Nishida, R., and Kuwahara, Y. October 2005.scheloribatid Mites as the Source of Pumiliotoxins in Dendrobatids. Journal of Chemical Ecology, Vol. 31, No. 10:2403-2415.
  5. ^ Raspotnig, G., Norton, Ray, A., Heethoff, M. 23 August 2011. Oribatid Mites and Skin Alkaloids in Poison Frogs. Biology Letters. Vol. 7, No. 4:555-556.
  6. ^ "Oophaga pumilio (Strawberry poision frog) (2532163201)" by Marshal Hedin from San Diego - Oophaga pumilio (Strawberry poision frog) Uploaded by Jacopo Werther. Licensed under Creative Commons Attribution 2.0 via Wikimedia Commons - https://commons.wikimedia.org/wiki/File:Oophaga_pumilio_(Strawberry_poision_frog)_(2532163201).jpg#mediaviewer/File:Oophaga_pumilio_(Strawberry_poision_frog)_(2532163201).jpg
  7. ^ Staudt, K., Meneses, O., S., Mebs, D., Proehl, H. 2010. Foraging Behaviour and Territoriality of the Strawberry Poison Dart Frog (Oophaga pumilio) in Dependence of the Presence of Ants. Amphibia-Reptilia Vol. 31, No. 2:217-227.
  8. ^ Mebs, D., Pogoda, W., Batista, A., Ponce, M., Koelher, G., Kauert, G. 20 November 2008. Variability of Alkaloid profiles in Oophaga pumilio (Amphibia: Anura: Dendrobatidae) from Western Panama and Southern Nicaragua. Salamandra Vol. 44, No. 4:241-247.
  9. ^ Superscript textStynoski, J., L., Torres-Mendoza, Y., Sasa-Marin, M., Saporito, R., A. March 2014. Evidence of Maternal Provisioning of Alkaloid-based Chemical Defenses in the Strawberry Poison Frog Oophaga pumilio. Ecology (Washington DC), Vol. 95, No. 3: 587-593.
  10. ^ 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.
  11. ^ Donnelly, M. A. 1989. Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica. Journal of Herpetology, 23:362-367.
  12. ^ a b 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., and Wheeler, W. 2006. Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia, Athesphatanura, Dendrobatidae). Bulletin of the American Museum of Natural History 299: 1-262. PDF fulltext
  13. ^ a b c d Haase, A., and H. Prohl. 2002. Female activity patterns and aggressiveness in the strawberry poison frog Dendrobates pumilio (Anura: Dendrobatidae). Amphibia-Reptilia 23: 129-140.
  14. ^ Prohl, H., and W. Hodl. 1999. Parental investment, potential reproductive rates, and mating system in the strawberry dart-poison frog, Dendrobates pumilio. Behavioral Ecological Sociobiology 46: 215-220
  15. ^ Myers, C.W., Daly, J.W., and Martinez, V. 1984. An arboreal poison frog (Dendrobates) from western Panama. American Museum Novitates 2783:1-20.
  16. ^ Limerick, S. 1980. Courtship behavior and oviposition of the poison-arrow frog Dendrobates pumilio. Herpetologica 36:69-71.
  17. ^ (Savage, 2002).
  18. ^ Zimmermann, H. and Zimmermann, E. 1988. Etho-Taxonomie und zoogeographische Artengruppenbildung bei Pfeilgiftfröschen (Anura: Dendrobatidae). Salamandra 24:125-160.
  19. ^ Summers, K., Weigt, L.A., Boag, P., and 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.
  20. ^ Roberts, J.L., Brown, J.L., von May, R., Arizabal, W., Presar, A., Symula, R., Schulte, R., and 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.
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