Causes of sex‐biased adult survival in ungulates: sexual size dimorphism, mating tactic or environment harshness?

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Cohort effects and deer population dynamics (2003, background): “In most deer populations, males grow faster and longer than females (Jarman, 1983) and are more sensitive to environmental harshness (CluttonBrock, Guinness & Albon, 1982), leading females to outlive males (Toïgo & Gaillard, 2003).”
Empirical Evidence of Density- Dependence in Populations of Large Herbivores (2009, background): “Depending on mating system, males may be unable to replenish the energy expended during the rut and hencemight survive less than females, especially at high density (Byers, 1997; Hogg and Forbes, 1997; Toı̈go and Gaillard, 2003).”
Potential causes and life-history consequences of sexual size dimorphism in mammals (2005, background): “While males did appear to suffer costs of maintaining a larger size during times of environmental stress, under favourable environmental conditions and during times of good food supply, sexual size dimorphism appeared to be advantageous to males in terms of survival ( Toigo & Gaillard, 2003 ).”