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Although the advantage of polyandry stemming from genetic bet-hedging has been questioned, especially when it incurs a cost to females (Yasui 1998), there is empirical evidence of reduced parasite load and higher fitness payoffs resulting from a genetic bet-hedging (Baer / Schmid-Hempel 1999). Further, a direct cost of multiple mating may be substantially lower in externally fertilizing species compared to specieswithinternal transferof sperm(Arnqvist & Rowe 2005), although there may be indirect costs to females associated with behaviour, such as aggression,refusal to spawn, or decreased paternal care (see below).