Mimic poison frog

Ranitomeya imitator (formerly Dendrobates 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,[1][2] and it is one of the best known dart frogs.[3] 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.[3]

Mimic poison frog
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Anura
Family: Dendrobatidae
Genus: Ranitomeya
Species:
R. imitator
Binomial name
Ranitomeya imitator
(Schulte, 1986)
Synonyms

Dendrobates imitator Schulte, 1986

Color morphology
Captive bred R. imitator 'Chazuta'

Among different populations of R. imitator, different color morphs are present to mimic other poison frog species. There are four color morphs among R. imitator populations that mimic closely related sympatric (existing in the same area) species: varadero (R. fantastica), striped (lowland R. variabilis), spotted (highland R. variabilis), and banded (R. summersi).[4] The geographic distribution of these morphs primarily includes geographically isolated populations in different areas of Peru, however there are a number of "transition zones" between populations where different color morphs interact.[5] The striped morph is the most widely spread, mimicking the striped Ranitomeya variabilis and can be found throughout the lower Huallaga River drainage in Peru.[6] The spotted morph mimics the highland spotted frog Ranitomeya variabilis with mainly blue-green coloration, but can be found in other forms, sometimes in yellow.[6] Although R. imitator closely resembles R. variabilis in coloration with its striped morph, the two species differ in many key aspects. R. imitator provides biparental care and is a monogamous species, while R. variabilis provides solely paternal care and is polygamous.[7] The aradero morph is a lowland form that lives nearby another but does not resemble it.[6] Last, the banded morph, a mimic of Ranitomeya summersi, lives in much drier climates than the average R. imitator and is most often found in Dieffenbachia and Heliconia plants.[6]

Interactions between different morphs

While populations of different color morphs are concentrated in different areas of Peru, there are transition zones between populations where different color morphs may interact.[5] These transition zones have been particular areas of interest to study reproductive isolation and incipient speciation, and recent research suggests that color pattern is likely a driving factor in reproductive isolation, as some transition zones show evidence of assortative mating, where individuals of one morph prefer to mate with others of the same morph.[4][5] Mating calls by males are an important factor in mating, and often need to be distinct in order to attract females of the same species. In R. imitator, male calls vary little among different color morphs, however it has been suggested that color pattern is the principal component that drives speciation.[8][9]

Toxicity

Like most other Ranitomeya species, R. imitator has a mild toxicity compared to other poison dart frogs. It produces the potent pumiliotoxin B, but its small size limits the amount of poison it can secrete. Like other poison dart frogs, it does not produce toxin in captivity. It probably gains its poison from consuming toxic insects or other invertebrates in the wild. Frogs of the related genus Phyllobates may derive their toxins from local melyrid beetles of genus Choresine.[10]

Reproduction and parental care
Ranitomeya imitator's developmental life stages.

The reproductive and parental care behaviors exhibited by this species have been of great interest to a variety of researchers, namely because R. imitator is the first amphibian to show evidence of monogamy, and because it independently evolved biparental care.[11] Further, the occurrence of both of these factors supports the hypothesis that biparental care favors the evolution of monogamy, for which evidence has previously been restricted to other groups of vertebrates.[12] The parental care provided to R. imitator offspring has been demonstrated to be vital for offspring growth and success.[12]

Courtship and oviposition

To attract mates, males will initiate calls while approaching a female. If the female is receptive to the male's courtship, she will follow him to an oviposition site where they will mate. She will then deposit fertilized eggs on a plant to allow them to develop.[5][11] The number of tadpoles that successfully hatch and make it to their breeding pool is relatively small, where one study observed a range of 1-4 tadpoles per mated pair.[12]

Male parental care

After mating occurs, the male will guard the fertilized eggs. Upon hatching, the male will transport individual tadpoles to their own respective phytotelma, which are small pools of water within plants.[13] Additionally, the male will strategically place tadpoles in certain rearing sites, specifically avoiding predators.[14] After tadpoles are in their rearing sites, the male will call while next to tadpoles which signals to the mother to feed an individual.[11] Tadpole begging, a behavior in which tadpoles vibrate their body, also encourages the mother to provide trophic (eggs specifically for nutrition) eggs.[15]

Female parental care

Female parental care is provided in the form of trophic egg feeding to offspring. After mating, the female will have a surplus of unfertilized eggs, which she provisions to offspring as food that aids in offspring growth and development.[12] This form of biparental care is not uncommon in poison frogs. Researchers have found that poison frogs that use phytotelma to rear offspring are more likely to exhibit egg-feeding parental care, and that this combination favors the evolution of biparental care in these species.[11] The hormonal regulation of parental care in this species remains somewhat unknown.[16]

Ownership

Compared to many other dart frog species, Ranitomeya imitator has relatively large and stable wild populations.[17] However, they are often illegally collected and exported.[18] They have also been imported legally, and multiple captive-bred varieties exist in the pet trade.

References
  1. East Carolina University. March 12, 2010. Biologists find proof of first confirmed species of monogamous frog. ScienceDaily. Retrieved May 31, 2013.
  2. Sherratt, T. (2008). "The Evolution of Müllerian Mimicry". Die Naturwissenschaften. 95 (8): 681–695. Bibcode:2008NW.....95..681S. doi:10.1007/s00114-008-0403-y. PMC 2443389. PMID 18542902.
  3. J.L. Brown, E. Twomey (2011). “ZOOTAXA: A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae)” Magnolia Press pp. 68-72. In Schulte, Rainer (1989). “Eine Neue Dendrobates- Art aus Ostperu (Amphibia: Salentia: Dendrobatidae) pp. 11-21
  4. Twomey, Evan; Vestergaard, Jacob S.; Summers, Kyle (2014-08-27). "Reproductive isolation related to mimetic divergence in the poison frog Ranitomeya imitator". Nature Communications. 5 (1): 4749. doi:10.1038/ncomms5749. ISSN 2041-1723.
  5. Twomey, Evan; Vestergaard, Jacob S.; Venegas, Pablo J.; Summers, Kyle (February 2016). "Mimetic Divergence and the Speciation Continuum in the Mimic Poison Frog Ranitomeya imitator". The American Naturalist. 187 (2): 205–224. doi:10.1086/684439. ISSN 0003-0147.
  6. Ranitomeya imitator”. dendrobates.org. In Schulte, Rainer (1989). “Eine Neue Dendrobates- Art aus Ostperu (Amphibia: Salentia: Dendrobatidae) Sauria 8(3): 11-20.
  7. Brown, J. L.; et al. (2010). "A key ecological trait drove the evolution of biparental care and monogamy in an amphibian" (PDF). The American Naturalist. 175 (4): 436–46. doi:10.1086/650727. PMID 20180700. S2CID 20270737.
  8. Twomey, Evan; Mayer, Michael; Summers, Kyle (December 2015). "Intraspecific Call Variation in the Mimic Poison Frog Ranitomeya imitator". Herpetologica. 71 (4): 252–259. doi:10.1655/HERPETOLOGICA-D-15-00004. ISSN 0018-0831.
  9. Twomey, Evan; Kain, Morgan; Claeys, Myriam; Summers, Kyle; Castroviejo-Fisher, Santiago; Van Bocxlaer, Ines (May 2020). "Mechanisms for Color Convergence in a Mimetic Radiation of Poison Frogs". The American Naturalist. 195 (5): E132–E149. doi:10.1086/708157. ISSN 0003-0147.
  10. Dumbacher, J. P.; et al. (2004). "Melyrid beetles (Choresine): A putative source for the batrachotoxin alkaloids found in poison-dart frogs and toxic passerine birds". Proceedings of the National Academy of Sciences of the United States of America. 101 (45): 15857–60. Bibcode:2004PNAS..10115857D. doi:10.1073/pnas.0407197101. PMC 528779. PMID 15520388.
  11. Brown, Jason L.; Morales, Victor; Summers, Kyle (April 2010). "A Key Ecological Trait Drove the Evolution of Biparental Care and Monogamy in an Amphibian". The American Naturalist. 175 (4): 436–446. doi:10.1086/650727. ISSN 0003-0147.
  12. Tumulty, James; Morales, Victor; Summers, Kyle (2014). "The biparental care hypothesis for the evolution of monogamy: experimental evidence in an amphibian". Behavioral Ecology. 25 (2): 262–270. doi:10.1093/beheco/art116. ISSN 1465-7279.
  13. Summers, Kyle; Brown, Jason; Morales, Victor; Twomey, Evan (2008). "Phytotelm size in relation to parental care and mating strategies in two species of Peruvian poison frogs". Behaviour. 145 (9): 1139–1165. doi:10.1163/156853908785387647. ISSN 0005-7959.
  14. Brown, J. L.; Morales, V.; Summers, K. (November 2008). "Divergence in parental care, habitat selection and larval life history between two species of Peruvian poison frogs: an experimental analysis". Journal of Evolutionary Biology. 21 (6): 1534–1543. doi:10.1111/j.1420-9101.2008.01609.x.
  15. Yoshioka, M.; Meeks, C.; Summers, K. (March 2016). "Evidence for begging as an honest signal of offspring need in the biparental mimic poison frog". Animal Behaviour. 113: 1–11. doi:10.1016/j.anbehav.2015.12.024.
  16. Schulte, Lisa M.; Summers, Kyle (May 2017). "Searching for hormonal facilitators: Are vasotocin and mesotocin involved in parental care behaviors in poison frogs?". Physiology & Behavior. 174: 74–82. doi:10.1016/j.physbeh.2017.03.005.
  17. Javier Icochea, Ariadne Angulo, Karl-Heinz Jungfer 2004. Ranitomeya imitator. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.1.
  18. “Clare, John (July 30, 2011). “Imitating Dart Frog, Ranitomeya imitator/Dendrobates imitator (Schulte, 1986) - Care and Breeding”. frogforum.net
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