John C. Wingfield

University of California, Davis, Davis, California, United States

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Publications (423)1264.69 Total impact

  • Source
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    ABSTRACT: The physical and biological responses to rapid arctic warming are proving acute, and as such, there is a need to monitor, understand, and predict ecological responses over large spatial and temporal scales. The use of the normalized difference vegetation index (NDVI) acquired from airborne and satellite sensors addresses this need, as it is widely used as a tool for detecting and quantifying spatial and temporal dynamics of tundra vegetation cover, productivity, and phenology. Such extensive use of the NDVI to quantify vegetation characteristics suggests that it may be similarly applied to characterizing primary and secondary consumer communities. Here, we develop empirical models to predict canopy arthropod biomass with canopy-level measurements of the NDVI both across and within distinct tundra vegetation communities over four growing seasons in the Arctic Foothills region of the Brooks Range, Alaska, USA. When canopy arthropod biomass is predicted with the NDVI across all four growing seasons, our overall model that includes all four vegetation communities explains 63% of the variance in canopy arthropod biomass, whereas our models specific to each of the four vegetation communities explain 74% (moist tussock tundra), 82% (erect shrub tundra), 84% (riparian shrub tundra), and 87% (dwarf shrub tundra) of the observed variation in canopy arthropod biomass. Our field-based study suggests that measurements of the NDVI made from air- and spaceborne sensors may be able to quantify spatial and temporal variation in canopy arthropod biomass at landscape to regional scales.
    Ecological Applications 04/2015; 25(3):779-790. · 4.13 Impact Factor
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    ABSTRACT: Arctic weather in spring is unpredictable and can also be extreme, so Arctic-breeding birds must be flexible in their breeding to deal with such variability. Unpredictability in weather conditions will only intensify with climate change and this in turn could affect reproductive capability of migratory birds. Adjustments to coping strategies are therefore crucial, so here we examined the plasticity of the adrenocorticotropic stress response in two Arctic songbird species-the snow bunting (Plectrophenax nivalis) and Lapland longspur (Calcarius lapponicus)-breeding in northwest Greenland. Across the breeding season, the stress response was strongest at arrival and least robust during molt in male snow buntings. Snow bunting females had higher baseline but similar stress-induced corticosterone levels compared to males. Modification of the stress response was not due to adrenal insensitivity, but likely regulated at the anterior pituitary gland. Compared to independent nestlings and adult snow buntings, parental-dependent chicks had a more robust stress response. For Lapland longspurs, baseline corticosterone was highest at arrival in both male and females, and arriving males displayed a higher stress response compared to arriving females. Comparison of male corticosterone profiles collected at arrival in Greenland (76°N) and Alaska (67-71°N;) reveal that both species have higher stress responses at the more northern location. Flexibility in the stress response may be typical for birds nesting at the leading edges of their range and this ability will become more relevant as global climate change results in major shifts of breeding habitat and phenology for migratory birds. J. Exp. Zool. 9999A: 1-10, 2015. Copyright © 2015 Wiley Periodicals, Inc. © 2015 Wiley Periodicals, Inc.
    Journal of Experimental Zoology Part A Ecological Genetics and Physiology 03/2015; 323A:266-275. DOI:10.1002/jez.1923 · 1.35 Impact Factor
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    ABSTRACT: The role of polychlorinated biphenyls (PCBs) on exposure-related endocrine effects has been poorly investigated in wild birds. This is the case for stress hormones including corticosterone (CORT). Some studies have suggested that environmental exposure to PCBs and altered CORT secretion might be associated. Here we investigated the relationships between blood PCB concentrations and circulating CORT levels in seven free-ranging polar seabird species occupying different trophic positions, and hence covering a wide range of PCB exposure. Blood ∑7PCB concentrations (range: 61–115,632 ng/g lw) were positively associated to baseline or stress-induced CORT levels in three species and negatively associated to stress-induced CORT levels in one species. Global analysis suggests that in males, baseline CORT levels generally increase with increasing blood ∑7PCB concentrations, whereas stress-induced CORT levels decrease when reaching high blood ∑7PCB concentrations. This study suggests that the nature of the PCB-CORT relationships may depend on the level of PCB exposure.
    Environmental Pollution 02/2015; 197:173-180. DOI:10.1016/j.envpol.2014.12.007 · 3.90 Impact Factor
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    ABSTRACT: Rufous-collared sparrows (Zonotrichia capensis peruviensis) from valleys in the Atacama Desert of Chile, live in an extremely stable environment, and exhibit overlap in molt and reproduction, with valley-specific differences in the proportion of birds engaged in both. To better understand the mechanistic pathways underlying the timing of life-history transitions, we examined the relationships among baseline and stress-induced levels of corticosterone (CORT), testosterone, and bacteria-killing ability of the blood plasma (BKA), as well as haemosporidian parasite infections and the genetic structure of two groups of sparrows from separate valleys over the course of a year. Birds neither molting nor breeding had the lowest BKA, but there were no differences among the other three categories of molt-reproductive stage. BKA varied over the year, with birds in May/June exhibiting significantly lower levels of BKA than the rest of the year. We also documented differences in the direction of the relationship between CORT and BKA at different times during the year. The direction of these relationships coincides with some trends in molt and reproductive stage, but differs enough to indicate that these birds exhibit individual-level plasticity, or population-level variability, in coordinating hypothalamo-pituitary-adrenal axis activity with life-history stage. We found weak preliminary evidence for genetic differentiation between the two populations, but not enough to indicate genetic isolation. No birds were infected with haemosporidia, which may be indicative of reduced parasite pressure in deserts. The data suggest that these birds may not trade off among different life-history components, but rather are able to invest in multiple life-history components based on their condition. Copyright © 2015. Published by Elsevier Inc.
    General and Comparative Endocrinology 02/2015; 213. DOI:10.1016/j.ygcen.2015.02.010 · 2.67 Impact Factor
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    ABSTRACT: Seabirds often have high loads of contaminants. These contaminants have endocrine disrupting properties but their relationships with some endocrine mechanisms are still poorly investigated in free-living organisms. This is the case for the stress response which shifts energy investment away from reproduction and redirects it towards survival. In birds, this stress response is achieved through a release of corticosterone and is also accompanied by a decrease in circulating prolactin, an anterior pituitary hormone widely involved in regulating parental cares. We measured blood concentrations of some legacy persistent organic pollutants (POPs) and mercury (Hg) and examined their relationships with the corticosterone and prolactin responses of known-age (9–46 years old) incubating snow petrels (Pagodroma nivea) to a standardized capture/handling stress protocol. In this Antarctic seabird, we also investigated whether high contaminant burden correlates with a higher occurrence of egg neglect, a frequently observed behavior in snow petrels. POPs and Hg were unrelated to age. Stress-induced corticosterone concentrations were positively related to POPs in both sexes, and stress-induced prolactin concentrations were negatively related to Hg in males. Egg-neglect behavior was not related to POPs burden, but males with higher Hg concentrations were more likely to neglect their egg. This suggests that in birds, relationships between age and contaminants are complex and that even low to moderate concentrations of POPs and Hg are significantly related to hormonal secretion. In this Antarctic species, exposure to legacy POPs and Hg could make individuals more susceptible to environmental stressors such as ongoing disturbances in Polar Regions.
    Science of The Total Environment 02/2015; 505:180-188. DOI:10.1016/j.scitotenv.2014.10.008 · 3.16 Impact Factor
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    ABSTRACT: With the physical and biological responses to arctic warming proving acute, there is a need to monitor, understand, and predict ecological responses over large spatial and temporal scales. The normalized difference vegetation index (NDVI) has been used to addresses this need, as it is a widely adopted tool for detecting and quantifying spatial and temporal dynamics of tundra vegetation cover, productivity, and phenology. Such extensive use of the NDVI to quantify vegetation characteristics suggests that it may be similarly applied to characterizing consumer communities. We developed empirical models to predict canopy arthropod biomass based on canopy-level measurements of the NDVI. We did this both across and within distinct tundra vegetation communities in the arctic tundra of Alaska. This research is a first step towards assessing the potential for datasets acquired by air and spaceborne sensors to quantify spatial and temporal dynamics in canopy arthropod biomass at landscape and regional scales. When canopy arthropod biomass was predicted with the NDVI across four growing seasons (2010 through 2013), our overall model, which included four distinct vegetation communities, explained 63% of the variance in canopy arthropod biomass. Each of our four vegetation community-specific models explained 74% (moist tussock tundra), 82% (erect shrub tundra), 84% (riparian shrub tundra), and 87% (dwarf shrub tundra) of the observed variation in canopy arthropod biomass.
    Arctic Long-Term Ecological Research (ARC LTER), Woods Hole, MA; 02/2015
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    ABSTRACT: The hormonal stress response is flexible and can be modulated by individuals according to its costs and benefits. Therefore, it is predicted that parents in poor body condition should modify their hormonal stress response, and thus, redirect energy allocation processes from parental care to self-maintenance when stressors occur. To test this prediction, most studies on free-living vertebrates have only focused on the stress response while the stress recovery – how quickly hormonal levels return to baseline values - has been neglected. Moreover, most studies have only focused on corticosterone –the primary mediator of allostasis - without paying attention to prolactin despite its major role in mediating parental behaviours. Here, we examined the effect of a short-term fasting event on the corticosterone and prolactin stress responses and recoveries, and we subsequently explored their relationships with parental decision in the snow petrel (Pagadroma nivea). By comparing the hormonal profiles of fasting and non-fasting snow petrels, we showed that parents modulate their corticosterone (but not prolactin) stress response according to their energetic status. We also described for the first time the hormonal stress recoveries in wild birds and found that they did not differ between fasting and non-fasting birds. Importantly, egg neglect was negatively correlated with circulating prolactin but not corticosterone levels in this species, demonstrating therefore a complex link between body condition, parental behavior and circulating corticosterone and prolactin levels. We suggest that both corticosterone and prolactin play a major role in the way parents adjust to stressors. This multiple signaling may allow parents to fine-tune their response to stressor, and especially, to activate specific allostasis-related mechanisms in a timely manner.
    Hormones and Behavior 01/2015; 67:28-37. DOI:10.1016/j.yhbeh.2014.11.009 · 4.51 Impact Factor
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    ABSTRACT: Abstract Some sexually selected signals are thought to convey information about the current condition and genetic/epigenetic quality of the individual signaling, including the ability to resist parasites. However, it is unclear whether semistatic sexual signals that develop periodically and remain stable over protracted periods, such as avian breeding plumage, can relate to measures of current condition and health. We examined a semistatic signal (wing epaulet size) in male red-winged blackbirds (Agelaius phoeniceus) during the breeding season and looked for relationships between this trait and circulating testosterone (T), hematocrit, bacteria-killing ability (BKA) of the blood, and the infection status, richness, and abundance of four functional categories of parasite. We found that epaulet size was positively related to circulating levels of T and ectoparasite infections. We found no relationships between T and parasite infections. In adult males there was a negative relationship between T and BKA, whereas in yearling males there was no relationship. We found no evidence for a general reduction in immunocompetence in males with larger epaulets but rather an increase in susceptibility to specific types of parasites. Our results suggest that semistatic signals can be linked to measures of current condition, and we postulate that these relationships are modulated via activity levels related to breeding-season activities.
    Physiological and Biochemical Zoology 01/2015; 88(1):11-21. DOI:10.1086/679475 · 2.05 Impact Factor
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    ABSTRACT: Seabirds often have high loads of contaminants. These contaminants have endocrine disrupting properties but their relationships with some endocrine mechanisms are still poorly investigated in free-living organisms. This is the case for the stress response which shifts energy investment away from reproduction and redirects it towards survival. In birds, this stress response is achieved through a release of corticosterone and is also accompanied by a decrease in circulating prolactin, an anterior pituitary hormone widely involved in regulating parental cares. We measured blood concentrations of some legacy persistent organic pollutants (POPs) and mercury (Hg) and examined their relationships with the corticosterone and prolactin responses of known-age (9–46 years old) incubating snow petrels (Pagodroma nivea) to a standardized capture/handling stress protocol. In this Antarctic seabird, we also investigated whether high contaminant burden correlates with a higher occurrence of egg neglect, a frequently observed behavior in snow petrels. POPs and Hg were unrelated to age. Stress-induced corticosterone concentrations were positively related to POPs in both sexes, and stress-induced prolactin concentrations were negatively related to Hg in males. Egg-neglect behavior was not related to POPs burden, but males with higher Hg concentrations were more likely to neglect their egg. This suggests that in birds, relationships between age and contaminants are complex and that even low to moderate concentrations of POPs and Hg are significantly related to hormonal secretion. In this Antarctic species, exposure to legacy POPs and Hg could make individuals more susceptible to environmental stressors such as ongoing disturbances in Polar Regions.
    Science of The Total Environment 01/2015; 505:180–188. · 3.16 Impact Factor
  • John C Wingfield
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    ABSTRACT: The Earth has always been a changeable place but now warming trends shift seasons and storms occur with greater frequency, intensity and duration. This has prompted reference to the modern era as the Anthropocene caused by human activity. This era poses great challenges for all life on earth and important questions include why and how some organisms can cope and others cannot? It is of heuristic value to consider a framework for types of environmental signals and how they might act. This is especially important as predictable changes of the environment (seasonality) are shifting rapidly as well as unpredictable changes (perturbations) in novel ways. What we need to know is how organisms perceive their environment, transduce that information into neuroendocrine signals that orchestrate morphological, physiological and behavioral responses. Given these goals we can begin to address the questions: do neuroendocrine systems have sufficient flexibility to acclimate to significant change in phenology, are genetic changes leading to adaptation necessary, or both? Copyright © 2014 Elsevier Inc. All rights reserved.
    Frontiers in Neuroendocrinology 12/2014; DOI:10.1016/j.yfrne.2014.11.005 · 7.58 Impact Factor
  • Brian G. Walker, P. Dee Boersma, John C. Wingfield
    Canadian Journal of Zoology 12/2014; DOI:10.1139/cjz-2014-0216 · 1.35 Impact Factor
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    Jesse S. Krause, David Dorsa, John C. Wingfield
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    ABSTRACT: The aim of this study was to determine circulating patterns of the three major adrenal steroids in blood in response to stress during acute restraint handling in two subspecies of white-crowned sparrow (Zonotrichia leucophrys). Gambel’s white-crowned sparrows (Z.l. gambelii) are long distance migrants that breed at high latitudes and Nuttall’s white-crowned sparrows (Z.l. nuttalli) are residents of coastal California. Column partition chromatography was developed to separate progesterone, dehydroepiandrosterone (DHEA), and corticosterone from a small plasma sample. Each of these steroids has the capability to modulate the stress response through various mechanisms. For example, progesterone is bound to corticosterone binding globulin (CBG) with a higher affinity than corticosterone. If plasma levels of progesterone rise during acute stress, then this could displace corticosterone from CBG and increase the amount of biologically active, free, corticosterone in blood. Dehydroepiandrosterone (DHEA) has been implicated to have many anti-stress properties with the potential to mitigate some of the actions of corticosterone. Results indicate that progesterone levels in both subspecies are elevated in response to acute stress handling. DHEA levels declined in Gambel’s but did not change in Nuttall’s. Thus DHEA does not follow the same secretory pattern as in mammals. Corticosterone levels were elevated in response to acute stress handling in both subspecies. This study provides new insight into an integrated stress response among three steroids.
    Comparative Biochemistry and Physiology - Part A Molecular & Integrative Physiology 11/2014; DOI:10.1016/j.cbpa.2014.07.019 · 2.37 Impact Factor
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    ABSTRACT: The acute stress response in vertebrates is a highly adaptive suite of physiological and behavioural mechanisms that promote survival in the face of deleterious stimuli from the environment. Facultative changes of physiology and behaviour are mediated through changes in circulating levels of glucocorticoids (corticosterone, cortisol) and their subsequent binding to the high affinity mineralocorticoid receptor (MR) or the low affinity glucocorticoid receptor (GR). Free-living wild Gambel's White-crowned sparrows (Zonotrichia leucophrys gambelii) display annual fluctuations in the stress response with marked attenuation during the transition from the pre-parental to the parental stage. We investigated whether this rapid reduction in the stress response is mediated through changes in MR and GR mRNA expression in the brain using in situ hybridisation. MR mRNA expression was significantly lower in the hippocampus as the birds became parental. No changes were observed in GR mRNA expression in the paraventricular nucleus (PVN) or preoptic area (POA) at this time. No significant correlations were found between initial capture levels of corticosterone and GR or MR mRNA expression. No differences were found in basal levels of corticosterone between pre-parental and parental in birds collected for in situ hybridisation. Stress response data revealed no difference at baseline but reductions in peak levels of corticosterone as birds became parental. These data suggest that changes in MR expression may be important for the regulation of the stress response or behavioural stress sensitivity in order to promote parental care and investment.This article is protected by copyright. All rights reserved.
    Journal of Neuroendocrinology 11/2014; DOI:10.1111/jne.12237 · 3.51 Impact Factor
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    ABSTRACT: The physical and biological responses to rapid arctic warming are proving acute, and as such, there is a need to monitor, understand, and predict ecological responses over large spatial and temporal scales. The use of the Normalized Difference Vegetation Index (NDVI) acquired from airborne and satellite sensors addresses this need as it is widely used as a tool for detecting and quantifying spatial and temporal dynamics of tundra vegetation cover, productivity, and phenology. Such extensive use of the NDVI to quantify vegetation characteristics suggests that it may be similarly applied to characterizing primary and secondary consumer communities. Here we develop empirical models to predict canopy arthropod biomass with canopy-level measurements of the NDVI both across and within distinct tundra vegetation communities over four growing seasons in the arctic foothills region of the Brooks Range, Alaska. When canopy arthropod biomass is predicted with the NDVI across all four growing seasons, our overall model that includes all four vegetation communities explains 63% of the variance in canopy arthropod biomass. Whereas each of our four vegetation community-specific models explain 74% (moist tussock tundra), 82% (erect shrub tundra), 84% (riparian shrub tundra), and 87% (dwarf shrub tundra) of the observed variation in canopy arthropod biomass. Our field-based study suggests that measurements of the NDVI made from air and spaceborne sensors may be able to quantify spatial and temporal variation in canopy arthropod biomass at landscape to regional scales.
    Ecological Applications 10/2014; DOI:10.1890/14-0632.1 · 4.13 Impact Factor
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    ABSTRACT: Hummingbirds present a unique combination between extremely high life costs and a number of efficient adaptations to fuel these demands. In addition to cognitive abilities, territorial hummingbirds display aggressive behaviors that allow for access to better food resources. In year-round territorial species, male-male territorial aggression is similar between breeding and non-breeding seasons; however, the endocrine mechanisms underlying control of territoriality during these distinct seasonal periods may differ. In many species, testosterone (T) triggers increased aggression during the breeding season whereas territoriality in the non-breeding season can be regulated by circulating the biologically inert sex steroid precursor dehydroepiandrosterone (DHEA) and converting it to T in target tissues. The seasonal hormonal regulation of hummingbird territorial behavior has heretofore been unknown. Our goal was to assess seasonal changes in sex steroids, territorial aggression levels, and body condition during reproductive and non-reproductive seasons in hummingbirds. To validate the use of cloacal fluid (CF) for the study of sex steroids, steroid levels in plasma and CF were correlated in Sephanoides sephaniodes. During the reproductive season, Calypte. anna, Archilochus alexandri, and Selasphorus rufus males showed high levels of T that were positively correlated with aggression, but the relationship between T and body condition was not consistent across species. As expected, T levels in females were significantly lower than in males in all seasons, however still detectable. During the non-reproductive season, CF DHEA of Calypte anna was high and positively correlated with aggressive behaviors and body condition. Our results suggest that hummingbirds display aggressive behaviors that could be linked to different hormones during the breeding and non-breeding seasons.
    10/2014; 155(4):1017-1025. DOI:10.1007/s10336-014-1088-y
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    ABSTRACT: Climate warming is affecting the Arctic in multiple ways, including via increased dominance of deciduous shrubs. Although many studies have focused on how this vegetation shift is altering nutrient cycling and energy balance, few have explicitly considered effects on tundra fauna, such as the millions of migratory songbirds that breed in northern regions every year. To understand how increasing deciduous shrub dominance may alter breeding songbird habitat, we quantified vegetation and arthropod community characteristics in both graminoid and shrub dominated tundra. We combined measurements of preferred nest site characteristics for Lapland longspurs (Calcarius lapponicus) and Gambel's White-crowned sparrows (Zonotrichia leucophrys gambelii) with modeled predictions for the distribution of plant community types in the Alaskan arctic foothills region for the year 2050. Lapland longspur nests were found in sedge-dominated tussock tundra where shrub height does not exceed 20 cm, whereas White-crowned sparrows nested only under shrubs between 20 cm and 1 m in height, with no preference for shrub species. Shrub canopies had higher canopy dwelling arthropod availability (i.e. small flies and spiders) but lower ground dwelling arthropod availability (i.e. large spiders and beetles). Since flies are the birds’ preferred prey, increasing shrubs may result in a net enhancement in preferred prey availability. Acknowledging the coarse resolution of existing tundra vegetation models, we predict that by 2050 there will be a northward shift in current White-crowned sparrow habitat range and a 20-60% increase in their preferred habitat extent, while Lapland longspur habitat extent will be equivalently reduced. Our findings can be used to make first approximations of future habitat change for species with similar nesting requirements. However, we contend that as exemplified by the current study's findings, existing tundra modeling tools cannot yet simulate the fine-scale habitat characteristics that are critical to accurately predicting future habitat extent for many wildlife species.This article is protected by copyright. All rights reserved.
    Global Change Biology 10/2014; DOI:10.1111/gcb.12761 · 8.22 Impact Factor
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    ABSTRACT: Emberizid sparrows (emberizidae) have played a prominent role in the study of avian vocal communication and social behavior. We present here brain transcriptomes for three emberizid model systems, song sparrow Melospiza melodia, white-throated sparrow Zonotrichia albicollis, and Gambel's white-crowned sparrow Zonotrichia leucophrys gambelii. Each of the assemblies covered fully or in part, over 89% of the previously annotated protein coding genes in the zebra finch Taeniopygia guttata, with 16,846, 15,805, and 16,646 unique BLAST hits in song, white-throated and white-crowned sparrows, respectively. As in previous studies, we find tissue of origin (auditory forebrain versus hypothalamus and whole brain) as an important determinant of overall expression profile. We also demonstrate the successful isolation of RNA and RNA-sequencing from post-mortem samples from building strikes and suggest that such an approach could be useful when traditional sampling opportunities are limited. These transcriptomes will be an important resource for the study of social behavior in birds and for data driven annotation of forthcoming whole genome sequences for these and other bird species.
    05/2014; 2:e396. DOI:10.7717/peerj.396
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  • Wolfgang Goymann, John C. Wingfield
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    ABSTRACT: Testosterone is a key hormone for the development of secondary sexual characters and dimorphisms in behavior and morphology of male vertebrates. Because females often express detectable levels of testosterone, testosterone has been suggested to also play a role in the modulation of secondary sexual traits in females. Previous comparative analyses in birds and fish demonstrated a relationship between male-to-female testosterone ratios and the degree of sexual dimorphism. Furthermore, female maximum testosterone was related to mating system and coloniality. Here, we reevaluate these previous ideas using phylogenetic analyses and effect size measures for the relationship between birds’ male-to-female maximum testosterone levels. Further, we investigate the seasonal androgen response of female birds (the difference from baseline to maximum testosterone), which in males is strongly related to mating system. We could not confirm a relationship between male-to-female testosterone, maximum female testosterone, or the seasonal androgen response of females with any life-history parameter. We conclude that the expectation that testosterone regulates traits in females in a similar manner as in males should be reconsidered. This expectation may be partially due to hormone manipulation studies using pharmacological doses of testosterone that had similar effects in females than in males but may be of limited importance for the physiological range of testosterone concentrations occurring within ecological and evolutionary contexts. Thus, the assumption that circulating testosterone should covary with ecologically relevant secondary sexual traits in females may be misleading: selection pressures on females differ from those on males and females may regulate behavior differently.
    Behavioral Ecology 04/2014; 25(4):685-699. DOI:10.1093/beheco/aru019 · 3.16 Impact Factor
  • Wolfgang Goymann, John C. Wingfield
    Behavioral Ecology 04/2014; 25(4):704-705. DOI:10.1093/beheco/aru106 · 3.16 Impact Factor

Publication Stats

24k Citations
1,264.69 Total Impact Points

Institutions

  • 2008–2015
    • University of California, Davis
      • Department of Neurobiology, Physiology and Behavior
      Davis, California, United States
    • University of Antwerp
      • Departement Biologie
      Antwerpen, VLG, Belgium
  • 1975–2015
    • University of Washington Seattle
      • • Department of Biology
      • • Department of Psychology
      Seattle, Washington, United States
  • 2013
    • Oklahoma State University - Stillwater
      • Department of Zoology
      SWO, Oklahoma, United States
  • 2010
    • University of Oregon
      • Institute of Neuroscience
      Eugene, Oregon, United States
  • 2008–2010
    • Queen's University
      • Department of Biology
      Kingston, Ontario, Canada
  • 2009
    • Northern Arizona University
      • Department of Biological Sciences
      Flagstaff, Arizona, United States
  • 2006
    • University of Texas at Austin
      • Department of Integrative Biology
      Austin, TX, United States
    • Purdue University
      • Department of Biological Sciences
      West Lafayette, Indiana, United States
    • New York State
      New York City, New York, United States
  • 2004–2006
    • Virginia Polytechnic Institute and State University
      • Department of Biological Sciences
      Blacksburg, VA, United States
    • Max Planck Institute for Ornithology
      • Max Planck Institute for Ornithology (Radolfzell)
      Pöcking, Bavaria, Germany
    • University of Groningen
      • Centre for Ecological and Evolutionary Studies (CEES)
      Groningen, Groningen, Netherlands
  • 2005
    • The Richard Stockton College of New Jersey
      Эгг Харбор Сити, New Jersey, United States
    • University of Massachusetts Boston
      • Department of Biology
      Boston, MA, United States
  • 2002
    • Koninklijk Nederlands Instituut voor Onderzoek der Zee - NIOZ
      Burg, North Holland, Netherlands
  • 2000
    • Arizona State University
      • School of Life Sciences
      Phoenix, Arizona, United States
  • 1997
    • Tokyo Medical and Dental University
      Edo, Tōkyō, Japan
  • 1991–1995
    • University of North Carolina at Greensboro
      • Department of Psychology
      Greensboro, NC, United States
  • 1994
    • University of Wollongong
      • Department of Biological Sciences
      City of Greater Wollongong, New South Wales, Australia
    • Waseda University
      Edo, Tōkyō, Japan
  • 1992–1994
    • Indiana University Bloomington
      • Department of Biology
      Bloomington, Indiana, United States
  • 1988
    • University of Victoria
      • Department of Biology
      Victoria, British Columbia, Canada
  • 1985–1988
    • The Rockefeller University
      • Center for Field Research in Ethology and Ecology
      New York, New York, United States
  • 1981
    • University of Bristol
      Bristol, England, United Kingdom