Amphibians have evolved a fascinating variety of ways to produce and look after their young. Parental care occurs in many families, with either the mother or the father, or occasionally both, spending time and effort to give their young a better chance in life. Recent Croaking Science articles have described the glass frogs (July), where fathers guard incubating eggs on leaves above streams; Phyllomedusa tree frogs (August) where the parents enclose the eggs in folded leaves above water; Mannophryne stream frogs (September), where fathers guard their eggs on land, then transport the hatched tadpoles on their backs to safe water. This month, we look at the marsupial frogs, where mothers carry their developing eggs on their backs. As well as considering their reproductive arrangements, we look at their conservation status: overall, an alarming picture.
The Hemiphractidae are a family of New World frogs, composed of 118 species in six genera. They are distributed across Central and South America, as well as some Caribbean islands. They are very small arboreal frogs, with a primarily nocturnal lifestyle. The main distinguishing feature of the family is the storage and development of eggs on the backs of females, either externally or within a pouch (which has earned some the nickname of marsupial frogs). They also have unusually large embryonic external gills compared to other frogs.
Males of several species have been seen to actively place the eggs on the backs of females during amplexus, during which time they are assumed to contribute their half of the required gametes. They have mostly been seen to breed during wet seasons in tropical areas, as most frogs do. In northern parts of South America there are two wet seasons, a long one lasting from April until August and a shorter one lasting from November until January. Hemiphractid frogs have been seen carrying eggs during both of these, suggesting that they do not possess a strict reproductive timetable but carry out opportunistic breeding when conditions suit.
You can see a range of specialisations between hemiphractid genera, none more apparent than the egg-carrying strategy (to pouch or not to pouch). The genera that utilise enclosed dorsal pouches include Gastrotheca, Flectonotus and Fritziana. Cryptobatrachus, Stefania and Hemiphractus don’t have pouches, instead using mucosal secretions to keep the eggs in place.
Within the pouchless genera, Stefania stand out due to the development of young. In this genus there is no free-swimming tadpole stage, resulting in the hatching of froglets directly from the eggs carried on the female’s back. Female Stefania individuals have been found carrying up to 25 eggs on their back! With a gestation period of around 3 months, those kids are one heck of a burden. Within the pouched or “marsupial” species, the two belonging to the genus Flectonotus stand out due to their mysterious nature, with little research being conducted on either. It is known, however, that hatching occurs within the pouch and that individuals are then deposited into small bodies of water, such as in the ‘tanks’ enclosed by bromeliad leaves; they are at the stage of being well developed tadpoles, which soon metamorphose, without feeding. It is not clear why Flectonotus retains this requirement for a brief aquatic phase, rather than progressing to the froglet stage in the pouch.
Conservation status
Analysis of IUCN’s Red List status for the family Hemiphractidae shows 32% of species in the four threatened categories (critically endangered to near threatened), similar to the figure for amphibians as a whole. However, only 25% of species are in the least concern category, which provides a much more alarming picture. This is because 24% of species are listed as data deficient, and 20% more have yet to be assessed. This is the reality behind some of the global figures for the state of nature. In some parts of the world, especially the tropics, the state of our knowledge of many groups and individual species is very poor. Because of the continuing loss and alteration of natural habitats around the world, the expectation is that species whose status is poorly known are unlikely to be doing well. However, this is not always the case, as we show below for one species of hemiphractid frog.
The dwarf marsupial frog (Flectonotus fitgeraldi) was brought into the spotlight by a recent piece of collaborative research between researchers in Scotland, Trinidad, the USA, Portugal and Venezuela. This study aimed to clarify the conservation status of the species, by establishing an accurate picture of their distribution. Currently, IUCN lists the species as Endangered, but this status appeared to be based on no published original research. Finding these frogs is easier said than done, since they are sized between 2 and 3cm (roughly the size of a 2 pence coin). They have been reported to exist across the islands of Trinidad and Tobago, as well as northern parts of Venezuela. However, there was uncertainty as to whether these three populations fit within a single species, since there had been several cases previously when close examination of widely distributed amphibian species revealed the different populations to be divergent enough to merit description as separate species.
F. fitzgeraldi are most reliably found by identification of the plant species they use as homes and tadpole deposition sites. These are commonly bromeliads, heliconias and aroids. The larval young of F. fitzgeraldi are deposited in phytotelmata, small pools of water contained within plant structures. The hatching tadpoles are at a late non-feeding stage, having been nourished by yolk content of the egg. In the days following deposition in a bromeliad tank, tadpoles develop limbs and emerge from the water as froglets. Metamorphosis is normally complete within 5 days, during which the tail is reabsorbed and froglets graduate to become fully-fledged frogs.
In order to review the Endangered classification of F. fitzgeraldi, the study made use of molecular techniques as well as surveys over the course of several years. Genomic DNA samples were compared for sequence similarity to determine whether the populations in Trinidad, Tobago and Venezuela all belonged to the same species. Surveys were initially conducted to determine the hours of peak activity in F. fitzgeraldi and found that they exclusively call between 6pm and 8pm, roughly an hour either side of sunset. Based on this information, presence/absence surveys were carried out using the call to determine presence (seeing the frogs is difficult, and moving vegetation to attempt to see them disturbs the frogs) at locations across Trinidad, Tobago and Venezuela. These took the form of transects through forested areas, either on foot or in vehicles, while listening for the raspy calls of F. fitzgeraldi, which sound like running your finger along a comb. This presence/absence data was used to build up a map of their distribution.
Results from the study showed the three populations belong to the same species, with no taxonomic changes needed. It was also shown that F. fitzgeraldi was distributed more widely across Trinidad, Tobago and Venezuela than previously thought. This, combined with the lack of evidence showing deforestation on these two islands, was sufficient to demonstrate that ‘Endangered’ is too extreme a label for the species. However, deforestation in Venezuela, mainly for cattle farming, was found to threaten the habitat quality of F. fitzgeraldi, so the study concluded that a categorisation of the species as ‘Vulnerable’ was appropriate.
Although F. fitzgeraldi is sheltered from mainland deforestation by the island refuges provided by Trinidad and Tobago, many other hemiphractid frogs do not share the same luxury. This means that habitat loss caused by deforestation poses a threat to species whose distribution is restricted to northern areas of the South American continent, such as Venezuela and Guyana. For this reason, species of the hemiphractid family must be monitored closely in these areas over the following decade.
The work on F. fitzgeraldi that we summarise here shows that detailed fieldwork can sometimes reveal that the status of a species is better than previously thought. It also shows the value of student expeditions, which can deploy significant numbers of enthusiastic young people to carry out detailed field research which would be prohibitively expensive otherwise.
Acknowledgements
We thank Paul Hoskisson and Joanna Smith for permission to use their photographs.
Further reading
Our account of the conservation status of F. fitzgeraldi is based on a research paper currently under review.
Cameron Boyle and Roger Downie
University of Glasgow