, except for one cocooned N. aquilonius with a higher concentration of 394·pg·ml -1 . For both species, AVT showed no relationship with plasma osmolality over the lower range of plasma osmolalities but was appreciably increased at the highest osmolality recorded. This study provides the first evidence that cocoon formation following burrowing is not obligatory in species that are capable of doing so, but that cocoon formation occurs when soil water conditions are more desiccating than for non-cocooned frogs.
It is well established that forming a cocoon, for frog species capable of doing so, markedly reduces evaporative water loss; however, the capacity of cocooned frogs to maintain hydration during extended estivation is not well understood. The combined effects of long-term estivation and water loss were examined in the cocoon-forming species Cyclorana platycephala by assessing the hydration state of the frogs throughout a 15-mo estivation period. Frogs lost mass throughout the 15-mo period to a maximum of 36%+/-6.5% of their initial standard mass. Plasma osmolality reached maximal levels by the ninth month of estivation at 487 mOsm kg(-1) and then remained stable to the fifteenth month of estivation. Urine osmolality continued to increase to the fifteenth month of estivation, at which point plasma and urine concentrations were isosmotic. The use of bladder water to counter losses from circulation was indicated by the relatively slow rate of increase in plasma osmolality with mass loss and the progressive increase in urine osmolality. For estivating frogs, evidence was found for a possible threshold relationship between plasma osmolality and plasma arginine vasotocin (AVT) concentration. After estivation, plasma AVT concentrations decreased markedly after 15-mo estivators were placed in water for 2 h, suggesting that high levels of AVT may not be integral to rapid rehydration in this species.
Arenophryne rotunda is a small (2-8 g) terrestrial frog that inhabits the coastal sand dunes of central Western Australia. While sand burrowing is a strategy employed by many frog species inhabiting Australia's semi-arid and arid zones, A. rotunda is unique among burrowing species because it lives independently of free water and can be found nocturnally active on the dune surface for relatively extended periods. Consequently, we examined the physiological factors that enable this unique frog to maintain water balance. A. rotunda was not found to have any special adaptation to reduce EWL (being equivalent to a free water surface) or rehydrate from water (having the lowest rehydration rate measured for 15 Western Australian frog species), but it was able to maintain water balance in sand of very low moisture (1-2%). Frogs excavated in the field were in dune sand of 4.4% moisture content, as a consequence of recent rain, which was more than adequate for these frogs to maintain water balance as reflected by their low plasma and urine osmotic concentrations. We suggest that in dry periods of the year, A. rotunda can achieve positive water balance by cutaneous water uptake by burrowing deeper into the substrate to where the percent water content is greater than 1.5%.
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