Zimkus et al. (2012) found that most Phrynobatrachus species breed in small, lotic bodies of water and have aquatic eggs with free-living, feeding tadpoles. However, reproductive modes that provide autonomy from permanent water bodies evolved independently at least seven times. These shifts towards alternate reproductive modes are not linked to a common temporal event, clades that exhibit alternate reproductive modes have lower diversification rates than those that deposit eggs aquatically. In addition, adult habitat, pedal webbing and body size have no effect on diversification rates. Although these traits are not associated with increased speciation rates, they may still provide opportunities to extend into new niches, thus increasing overall diversity.
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