Christopher M. Tonra

A call for full annual cycle research in animal ecology

For vertebrates, annual cycles are organized into a series of breeding and non-breeding periods that vary in duration and location but are inextricably linked biologically. Here, we show that our understanding of the fundamental ecology of four vertebrate classes has been limited by a severe breeding season research bias and that studies of individual and population-level responses to natural and anthropogenic change would benefit from a full annual cycle perspective. Recent emergence of new analytical and technological tools for studying individual and population-level animal movement could help reverse this bias. To improve understanding of species biology and reverse the population declines of many vertebrate species, a concerted effort to move beyond single season research is vital.

Early elevation of testosterone advances migratory preparation in a songbird.

SUMMARY The timing of events associated with spring migration can be an important determinant of fitness for migratory birds. The need to prepare and maintain physical condition for migration is one demand that must be met for early arrival on breeding areas, and this demand is compounded by the energetic demands of the physiological transition to breeding. Here, we examined whether migratory birds, specifically males, can adequately meet both of these demands by elevating the sex steroid testosterone early during migratory preparation. To test this, we used a captive experiment in which we photostimulated male dark-eyed juncos (Junco hyemalis) and manipulated circulating testosterone. We predicted males with elevated testosterone would prepare to migrate and breed faster than control males or males where the ability of testosterone to bind with receptors, or be converted to estradiol, was inhibited (testosterone inhibited). We measured migratory preparation using mass, food intake, fat deposits and nocturnal activity (Zugunruhe). To estimate breeding preparation, we measured the diameter of the cloacal protuberance (CP). We found that males in the testosterone treatment group began migratory preparation approximately 10 days before controls. There was no difference in the magnitude of peak migratory condition between testosterone-implanted birds and controls. Males implanted with testosterone also reached the onset of breeding preparation faster and had larger peak CP diameter compared with controls. Testosterone-inhibited birds showed no signs of migratory preparation and only a weak increase in CP diameter. These results demonstrate that early elevation of testosterone during migratory preparation could incur benefits for males in terms of the ability to depart earlier from non-breeding areas and arrive in a more advanced breeding condition. This experiment demonstrates that there may be important physiological underpinnings to known winter to breeding season carry-over effects in migratory birds.

How to capture wild passerine species to study baseline corticosterone levels

Baseline corticosterone levels have been suggested to be an effective tool to assess and monitor the health status of individuals and populations of wild birds. However, measuring baseline corticosterone levels in the field is difficult because the capture protocol may affect rates of corticosterone secretion. In passerine birds, two methods of capture are widely used: (1) passive netting consisting of monitoring mist-nets frequently to check if a bird has been caught; and (2) target netting consisting of installing a conspecific decoy and/or playing a tape-recorded conspecific song in the vicinity of a mist-net until the territorial bird is captured. Our objective here was to determine whether these methods of capture are effective at achieving non-disturbed baseline corticosterone levels in American Redstarts (Setophaga ruticilla). We found that passive netting was associated with elevated corticosterone levels, suggesting that hanging in a net for several minutes activates the HPA axis and, thus, elicits an adreno-cortical stress response. In contrast, target netting was not associated with an increase in corticosterone levels, regardless of the length of time between when the target bird first approached the net and when it was captured. Therefore, we suggest that researchers interested in estimated baseline corticosterone in this species use target netting and not passive netting.

Experimental and observational studies of seasonal interactions between overlapping life history stages in a migratory bird

Prior to reproduction, migratory animals are at the juxtaposition of three life history stages in which they must finish the non-breeding stage, initiate and complete migration, and prepare for the onset of breeding. However, how these stages interact with one another is not fully understood. We provide evidence that, for migratory birds that begin breeding development prior to departure from non-breeding sites, the level of breeding preparation can drive migration phenology, a critical behavioral determinant of reproductive success. Specifically, male American redstart (Setophaga ruticilla) plasma androgen levels, which increase in males during the period leading into migration, were positively correlated with energetic condition. We empirically tested the hypothesis that elevated androgen simultaneously supports migratory and breeding preparation in a hormone manipulation field experiment. Males with testosterone implants showed advanced preparation for migration and breeding, and ultimately departed on migration earlier than controls. It is assumed that early departure leads to early arrival at breeding areas, which increases breeding success. Collectively, our observational and experimental results demonstrate how overlapping life history stages can interact to influence important components of an individuals fitness. This highlights the critical need for understanding population processes across the full life cycle of an organism to better understand the ecological and evolutionary origins of complex life history events.

Color expression in experimentally regrown feathers of an overwintering migratory bird: implications for signaling and seasonal interactions

Plumage coloration in birds plays a critical role in communication and can be under selection throughout the annual cycle as a sexual and social signal. However, for migratory birds, little is known about the acquisition and maintenance of colorful plumage during the nonbreeding period. Winter habitat could influence the quality of colorful plumage, ultimately carrying over to influence sexual selection and social interactions during the breeding period. In addition to the annual growth of colorful feathers, feather loss from agonistic interactions or predator avoidance could require birds to replace colorful feathers in winter or experience plumage degradation. We hypothesized that conditions on the wintering grounds of migratory birds influence the quality of colorful plumage. We predicted that the quality of American redstart (Setophaga ruticilla) tail feathers regrown after experimental removal in Jamaica, West Indies, would be positively associated with habitat quality, body condition, and testosterone. Both yearling (SY) and adult (ASY) males regrew feathers with lower red chroma, suggesting reduced carotenoid content. While we did not observe a change in hue in ASY males, SY males shifted from yellow to orange plumage resembling experimentally regrown ASY feathers. We did not observe any effects of habitat, testosterone, or mass change. Our results demonstrate that redstarts are limited in their ability to adequately replace colorful plumage, regardless of habitat, in winter. Thus, feather loss on the nonbreeding grounds can affect social signals, potentially negatively carrying over to the breeding period.

Variation in Stable Hydrogen Isotope Values in a Wetland-Associated Songbird

Abstract. Stable isotope analysis is often used to identify the geographic origins of migratory bird populations. While this method can accurately predict the provenance of migratory species, stable hydrogen isotope values measured in feathers (δ2Hf) can be variable within a site and may be influenced by differences among age class (second year vs. after second year), type of feather (primary vs. rectrix), year of sampling, species, and local hydrology. In this study, sources of variation in δ2Hf values were assessed in a wetland-associated Neotropical migratory bird, the Prothonotary Warbler (Protonotaria citrea), by comparing δ2Hf values among age classes, sexes, years and feather types in individuals breeding in eastern Virginia, USA. Age and year were found to influence δ2Hf values, with individuals in their second year having more depleted δ2Hf values (-62.43 ± 9.56‰) than individuals older than 2 years (-53.73 ± 9.04‰). Differences between primaries and rectrices were within the range of sampling error (-2.98‰), and there was no effect of sex. For wetland-associated songbirds, age-related differences in the dietary proportion of aquatic vs. terrestrial prey may provide an additional mechanism for differences observed in δ2Hf values between age classes. In studies that use stable isotopes to assign geographic feather origin, researchers should attempt to account for and propagate known variation in δ2Hf values in assignment models whenever possible.

Prealternate molt-migration in Rusty Blackbirds and its implications for stopover biology

ABSTRACT To achieve greater understanding of the full annual cycles of birds, it is critical to describe the spatial nature of little-understood phases. One of the least understood aspects of avian annual cycles is the ecology of molt: the periodic replacement of feathers. While work on the spatial nature of molt in migratory passerines has increasingly found incidences of species and populations completing molt during migration, this work has been limited entirely to prebasic flight feather molt. We examined the prevalence and progression of contour feather molt in a migratory songbird, the Rusty Blackbird (Euphagus carolinus), during spring stopover. We found that 98% of birds exhibited a partial prealternate molt during stopover, primarily in the head region. Furthermore, molt intensity peaked in the middle of the migration period and was negatively associated with fat score. This is the first evidence in the passerine literature of an obligate prealternate molt completed during migration, which is in many ways similar to the molt strategy of a variety of shorebirds (Families Charadriidae and Scolopacidae). These findings could prove crucial to understanding the constraints on spring migration in this declining species. Furthermore, we argue that molt schedules such as those of the Rusty Blackbird and shorebirds should be referred to as “prealternate molt-migration,” broadening the traditional definition of molt-migration beyond prebasic flight feather molt.

Expanding the traditional definition of molt-migration

ABSTRACT The occurrence of molt during migration, known as “molt-migration,” has increasingly received attention across many avian taxa since first being described in waterfowl in the 1960s. However, despite the many different types of molt stages and strategies, most, if not all, uses of the term “molt-migration” apply to the definitive prebasic molt of flight feathers in post-breeding adults, whereas fewer studies address migration for body-feather molts. Here, we argue that the current definition of molt-migration, as applied, is limited in focus relative to the diverse ways in which it can manifest in avian populations. We suggest a new, broader definition of molt-migration and highlight examples of molt-migration as traditionally defined, and the many examples that have not been defined as such. We propose a new, 2-tiered typology for defining different forms of molt-migration, based on (1) its progression relative to stationary portions of the annual cycle, and (2) the stage of molt involved. In order to advance our understanding of the ecology and evolution of this increasingly documented phenomenon and apply this knowledge to conservation and management, avian researchers must begin to utilize a common framework for describing molt-migration in its various forms.