Abstract
Abstract Ecological processes driving female-skewed adult sex ratios (ASRs; males:females) in populations with polygynous mating systems have been addressed theoretically, but empirical support is scarce. The theoretical framework of the female substitution hypothesis (FSH) asserts that a female-skewed ASR at carrying capacity reflects an overall fitness benefit for females and for males competitive in acquiring access to reproductive females. The FSH predicts that as population abundance increases females should acquire forage more efficiently than males, thereby leading to passive displacement of males. The result is declining ASR associated with differential habitat use by both sexes as food resources are depleted by female scramble competition. We characterized the temporal variation in ASR in a Roosevelt elk population inhabiting the Redwood National and State Parks, California, across 24 years, and determined which of two possible ecological mechanisms was the driver of a declining ASR. The first mechanism explored was that increasing female abundance associated with declining forage in the study area led to the passive displacement of males into the study periphery over time. The second mechanism explored was that a declining ASR was precipitated by a lack of males within the study area and the study periphery. Systematic population surveys from a vehicle were done to estimate abundance and ASR as well as assess male abundance in the study periphery. Forage biomass was estimated in quarter-m2 plots randomly placed in meadows inhabited by female elk. Our multiple regression analysis revealed an inverse relationship between abundance and ASR indicating density dependence. We found numerous males in the study periphery when females were abundant. Our least squares models indicated declining food resources across years when female abundance increased. Our results showed that the first, and not the second, ecological mechanism examined was responsible for a female-skewed ASR. Our findings provide empirical support for the theoretical framework of the FSH in a nonmigratory population protected from hunting.
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