Year-round territorial aggression is independent of plasma DHEA in the European nuthatch Sittaeuropaea



Abstract
Plasma testosterone can play an important role in promoting aggressive behaviors relating to territory defense in breeding male birds. Some birds defend territories also during the non-breeding phase, when testosterone circulates at basal levels. In such species, plasma levels of the pro-hormone dehydroepiandrosterone (DHEA) may support non-breeding territoriality by acting as a local substrate for sex steroids. To test this possible role of plasma DHEA, we examined the seasonal DHEA profile of male (and female) European nuthatches Sitta europaea: a male and female nuthatch pair will defend an all-purpose territory throughout the year. We hypothesized that plasma DHEA would be detectable in wintering nuthatches with a territory. However, only ca. half of the territorial wintering males (and females) displayed detectable DHEA levels, suggesting that plasma DHEA is not a major sex steroid precursor during non-breeding. Further, among hatching-year birds, plasma DHEA was significantly lower in territorial birds than in “floaters”, i.e., subordinate birds without a territory. To experimentally examine the role of DHEA in non-breeding territoriality, we treated adult wintering males with DHEA and measured effects on aggressive responses to conspecific challenge. DHEA treatment elevated plasma levels of DHEA (and testosterone), but did not enhance territorial behaviors or their persistence. Taken together, our data suggest that DHEA (and, indeed, sex steroids per se) do not regulate non-breeding territoriality in the nuthatch. Given that territorial aggression in nuthatches is expressed year-round, a hormone for its activation may be redundant.

Highlights
► We examine the seasonal DHEA profile in a year-round territorial songbird. ► We test effects of DHEA treatment on non-breeding territoriality in autumn. ► Plasma DHEA shows no seasonal variation and is often undetectable. ► DHEA treatment does not enhance non-breeding aggression or its persistence. ► We report that DHEA does not relate to non-breeding territoriality in this species.

Keywords
Nuthatch; Seasonal DHEA profile; Implant; Territorial aggression; Testosterone; Non-breeding; Wintering