Estuarine fish and tetrapod evolution: insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent

The Waterloo Farm lagerstatte in South Africa provides a uniquely well-preserved record of a Latest Devonian estuarine ecosystem. Ecological evidence from it is reviewed, contextualised, and compared with that available from the analogous Swartvlei estuarine lake, with a particular emphasis on their piscean inhabitants. Although the taxonomic affinities of the estuarine species are temporally very different, the overall patterns of utilisation prove to be remarkably congruent, with similar trophic structures. Significantly, both systems show evidence of widespread use of estuaries as fish nurseries by both resident and marine migrant taxa. Holocene estuaries are almost exclusively utilised by actinopterygians which are overwhelmingly dominated by oviparous species. Complex strategies are utilised by estuarine resident species to avoid exposure of eggs to environmental stresses that characterize these systems. By contrast, many of the groups utilising Devonian estuaries were likely live bearers, potentially allowing them to avoid the challenges faced by oviparous taxa. This may have contributed to dominance of these systems by non-actinoptergians prior to the End Devonian Mass Extinction. The association of early aquatic tetrapods at Waterloo Farm with a fish nursery environment is consistent with findings from North America, Belgium and Russia, and may be implied by the estuarine settings of a number of other Devonian tetrapods. Tetrapods apparently replace their sister group, the elpistostegids, in estuaries with both groups having been postulated to be adaptated to shallow water habitats where they could access small piscean prey. Correlation of tetrapods (and elpistostegids) with fish nursery areas in the Late Devonian lends strong support to this hypothesis, suggesting that adaptations permitting improved access to the abundant juvenile fish within the littoral zone of estuarine lakes and continental water bodies may have been pivotal in the evolution of tetrapods.