Mannophryne is a genus of 20 frog species found in the forests of Venezuela and Trinidad and Tobago. They belong to the family Aromobatidae, the sister family to the well-known Dendrobatidae, or poison dart frogs. Unlike poison dart frogs, that are brightly coloured to advertise their toxicity, aromobatids use cryptic colouration to blend into their surroundings as they do not have toxic compounds in their skins. In addition, they have a very effective escape response and can identify and reach cover quickly. This has led to one of their common names of rocket frogs, as well as being known as stream frogs, because of their normal habitat of stream margins.
In Trinidad and Tobago, we studied the endemic Mannophryne trinitatis (Trinidad) and M. olmonae (Tobago). Previous assessments of both species for IUCN indicated that they are threatened with extinction on account of habitat loss or degradation. An additional worry was the finding that both species were infected with chytrid fungus, the cause of numerous amphibian population declines across the world, although Mannophryne seemed not to suffer from the infection. We conducted population surveys of M. trinitatis in the Northern Range of Trinidad and found them to be numerous, no longer infected with chytrid, and with no sign of significant habitat loss. In Tobago, M. olmonae are very restricted in their distribution and are found at much lower numbers along streams in the northeast of the island. However, they also showed a lack of chytrid infection and no serious loss of habitat. We contributed to a review of IUCN’s frog assessments and recommend that M. trinitatis now be classed as Least Concern, and M. olmonae as Vulnerable.
Mannophryne are small mottled brown frogs, only a few centimetres in length, which are active during the day close to small mountain streams in primary and secondary forests. They feed on small arthropods such as beetles, flies and spiders present on the abundant rotting fruit. During the rainy season (June to December), calling males can be heard morning and afternoon trying to attract a mate with their quick “weep, weep, weep” whistling calls. If you follow the calls and look carefully, they can be seen perched on rocks, roots, plants or at the mouths of small caves but will jump quickly away to safety if approached. Meanwhile, females hold a territory, which they defend from other females. When a female has chosen a mate, she follows the male into his cave and lays up to 14 eggs on the damp leaf litter. The males provide parental care, defending and cleaning the eggs until they hatch. This takes around 21 days and then each tadpole flips onto its father’s back and he begins the journey to look for a safe pool of water for his tadpoles to develop in. When a pool free from predatory fish or crustaceans and with a good supply of food is located, the father deposits some of the tadpoles and then searches for another. Tadpoles from a single clutch may be deposited in several bodies of water and will take around 2 months to develop to the stage of metamorphosis.
One of the most interesting and recently studied aspects of this genus is their colouration. Sexual dichromatism is the difference in colouration between the males and females of a species. This is often used in communication as a visual signal to warn off competitors or highlight quality to potential mates. Sexual dichromatism can be split into two categories. Dynamic dichromatism only occurs in males and involves the development of temporary colouration, often as part of a courtship display, whilst ontogenetic dichromatism can occur in males or females and involves the development of a permanent colour difference as they mature. Mannophryne are a very rare example among frogs in showing both, where males turn a jet-black colour when they call, and females develop a yellow patch on their throats as they mature.
This is noteworthy, because both forms of dichromatism are energetically expensive, stressing their importance. As male Mannophryne are active and calling during the day, their colour change seems counter- productive as it makes them more conspicuous. Additionally, it appears that open areas are a preferred calling site, and in turn attract a mate, further increasing their predation risk. Therefore, both signals (visual and audio) must be essential in attracting a mate. The role of the yellow throat in females is likely a signal of quality. We found that throat colour is individually variable in the extent of the colour patch and in its hue, ranging from pale yellow to bright orange. Larger and heavier females have more orange throats. Animals often use bright or bold colours to signal their quality to others and to avoid physical confrontation. In M. trinitatis, females can be see raising their heads to show their pulsing throat patch when confronted by another female. This signal shows that the defending female has the additional energy to dedicate to the “expensive” orange colouration and often provides enough incentive to dissuade the challenger from further conflict. Yellow and orange colours are based on carotenoid pigments, and there is evidence from many kinds of animals that carotenoids signal health and quality, as they are related to immune system functions. As a test of this idea that colour differences in the females are a sign of quality, we tested their escape responses in a semi-natural experiment: frogs with the brightest yellow throats showed more effective escape responses than the others.
An extra facet to throat colour use by Mannophryne females may be signalling their quality to males. As males take such exhaustive care of their clutch through to deposition of tadpoles, it is unlikely that they could manage more than one at a time. Therefore, it is important that they choose a high-quality female. Mate selection on both sides may be a more significant and understudied aspect to sexual selection in animals where males take all or the majority of parental care responsibilities.
Another fascinating piece of the puzzle is the timing of these colour changes. We observed the colour change on a male M. trinitatis, with the whole process taking around 20 minutes. The male was located by erratic calling and found to still be in its mottled brown colouration. As the calling became more consistent and frequent, the colouration changed to become darker until its change was complete. This is interesting because colour change in frogs is generally quite slow, mediated by hormones, and it will be interesting to discover if there is a special mechanism in Mannophryne. We found through rearing juveniles for several months after metamorphosis, that the yellow colouration develops very gradually in females, well before they reach sexual maturity.
Even though they do not have the bright colours or toxicity of their dart frog cousins, these little frogs are a lot more interesting and special than you might think at first glance.
Further Reading
Downie, J. R., Livingstone, S. R., & Cormack, J. R. (2001). Selection of tadpole deposition sites by male Trinidadian stream frogs, Mannophryne trinitatis (Dendrobatidae): an example of anti-predator behaviour. Herpetological Journal 11, 91-100.
Downie, J. R., Robinson, E., Linklater‐McLennan, R. J., Somerville, E., & Kamenos, N. (2005). Are there costs to extended larval transport in the Trinidadian stream frog, Mannophryne trinitatis (Dendrobatidae)?. Journal of Natural History 39, 2023-2034.
Greener, M. S., Hutton, E., Pollock, C. J., Wilson, A., Lam, C. Y., Nokhbatolfoghahai, M., … & Downie, J. R. (2020). Sexual dichromatism in the neotropical genus Mannophryne (Anura: Aromobatidae). PloS ONE 15(7), e0223080.
Greener, M. S., Shepherd, R., Hoskisson, P. A., Asmath, H., & Downie, J. R. (2017). How many Trinidad stream frogs (Mannophryne trinitatis) are there, and should they be regarded as vulnerable to extinction?. Herpetological Journal 27, 5-11.
Jowers, M. J., & Downie, J. R. (2005). Tadpole deposition behaviour in male stream frogs Mannophryne trinitatis (Anura: Dendrobatidae). Journal of Natural History 39, 3013-3027.
Lehtinen, R. M., Mannette, R. P., Naranjit, K. T., & Roach, A. C. (2007). Ecological observations on the critically endangered Tobago endemic frog Mannophryne olmonae. Applied Herpetology 4, 377-386.
Manzanilla, J., Jowers, M. J., La Marca, E., & García-París, M. (2007). Taxonomic reassessment of Mannophryne trinitatis (Anura: Dendrobatidae) with a description of a new species from Venezuela. Herpetological Journal 17, 31-42.
Contributing authors : Mark S. Greener and Roger Downie
