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I am broadly interested in growing organisms, especially within-species (intraspecific) variation in the growth rates of wild songbirds. I studied the impact of environmental variation on grassland songbird growth for my MSc and currently study the impact of maternally-invested resources (especially hormones!) on the growth of American Robins and commercially-raised Coturnix quail. 

Quickly navigate to the following sections:

-Ongoing Research

-Published Research

-ground-nesting robins (2022)

-blackbird mite infections (2022)

-growth-tuning cowbirds (2021)

-sparrow territory aggregation (2020)

-Published Collaborations (side-projects)

-Undergraduate Research


Ongoing Research:

The impact of maternal hormones on the growth and development of birds 

(fondly known on Twitter as project #GrowBirdGrow)

We are investigating the ways in which the hormones passed from mom to egg ultimately influence the growth and behavior of bird embryos, chicks, and juveniles. We are also exploring sibling competition, egg-laying order, egg resources, and nestling microbiome development in our study species, American Robins (Turdus migratorius) and commercially-raised Japanese Quail (Coturnix japonica). Keep checking in using the @GrowBirdGrow Twitter  as I update everyone on our progress!

The role of brood parasitism in shaping nestling growth and development strategies

(fondly known on Twitter as project #PrairieBabies)

In my master’s research, I seek to identify the effect(s) of Brown-headed Cowbird (Molothrus ater) brood parasitism on the growth and development of nestlings of three grassland-obligate host species: Grasshopper Sparrow (Ammodramus savannarum), Dickcissel (Spiza americana), and Eastern Meadowlark (Sturnella magna). Previous work on nestling development has illustrated the importance of perceived nest predation risk (e.g. Cheng and Martin 2012) and food availability (e.g. Ricklefs 1993)—in order to identify the variation explained by cowbird parasitism, we need to quantify the effect(s) of and interactions between food availability and predation risk as well. We will locate nests of each of the host species and identify the perceived predation risk, the food available to the nestlings, and the presence and risk of cowbirds at subsets of the total nests. We will then relate these factors to the skeletal growth, development (eyes opening, movement capacity), feather growth, and fat/muscle gain of the host nestlings, which we will calculate from nestling measurements taken every other day. At the conclusion of this study we hope to advance our knowledge of the impact of cowbirds, identify the development strategies that produce highest nest and post-fledge success in cowbird-dominated systems, and provide insight on the past and future evolution of cowbird hosts.

pink ugly newborn bird in a hand

Published Research:

Ground-nesting by arboreal American robins (Turdus migratorius)

Ecology and Evolution 2022, coauthors: M.E. Hauber, T.J. Benson, and M. Abolins-Abols, check it out here! 

Animals with dependent and vulnerable young need to decide where to raise their offspring to minimize ill effects of weather, competition, parasitism, and predation. These decisions have critical fitness consequences through impacting the survival of both adults and progeny. Birds routinely place their nest in specific sites, allowing species to be broadly classified based on nest location (e.g., ground- or tree-nesting). However, from 2018 to 2020, we observed 24 American robin (Turdus migratorius) nests placed not on their species-typical arboreal substrates or human-made structures but on the ground at a predator-rich commercial tree-farm in Illinois, USA. This behavior does not appear to be in response to competition and did not affect nest daily survival rate but was restricted to the early half of the breeding season. We hypothesize that ground nesting may be an adaptive response to avoid exposure and colder temperatures at sites above the ground early in the breeding season or a nonadaptive consequence of latent robin nest-placement flexibility.

Knemidokoptic mite infestation of island populations of Red-winged Blackbirds (Agelaius phoeniceus)

Wilson Journal of Ornithology 2022, J.S. Marshall and S.K. Winnicki, check it out here!

Knemidokoptic mites infest a number of bird species around the world and may lead to decreased survival, although relatively few studies have documented long-term effects on populations. We present 9 years of data monitoring the survival of Red-winged Blackbirds (Agelaius phoeniceus) in the Lake Erie, Ohio, USA islands. Blackbirds in our study first appeared with mite infestation in 2012. Percentage of the total number of blackbirds infested with mites peaked at 7.3% in 2013 and persisted at between 5 and 6% through the summer of 2018. We saw no differences in infestation rates based on blackbird sex or age. Unlike most other studies, we did recapture birds that had previously had infestations, and we detected no decline in recapture rate due to mite infestation. While that does not mean that blackbirds do not experience negative consequences of infestation, our evidence does suggest that low level infestation may be persistent in a population without dramatic declines in survivorship.

Male Red-winged Blackbird that we banded

Developmental asynchrony and host species identity predict variability in nestling growth of an obligate brood parasite: a test of the “growth-tuning” hypothesis

Canadian Journal of Zoology 2021, coauthors B.M. Strausberger, N.D. Antonson, D.E. Burhans, J. Lock, A.M. Kilpatrick, and M.E. Hauber, check it out here!

Generalist obligate brood parasites are excellent models for studies of developmental plasticity, as they experience a range of social and environmental variation when raised by one of their many hosts. Parasitic Brown-headed Cowbirds (Molothrus ater (Boddaert, 1783)) exhibit host-specific growth rates, yet cowbird growth rates are not predicted by hosts’ incubation or brooding periods. We tested the novel “growth-tuning” hypothesis which predicts that total asynchrony between cowbirds’ and hosts’ nesting periods results in faster parasitic growth in nests where host young fledge earlier than cowbirds. We tested this prediction using previously-published and newly-added nestling mass data across diverse host species. Total nesting period asynchrony (summed across incubation and brooding stages) predicted cowbird growth; 8-day old cowbirds were heavier in host nests with relatively shorter nesting periods. We further explored the drivers of variation in growth using mass measurements of cowbirds in Song Sparrow (Melospiza melodia (Wilson, 1810)) and Red-winged Blackbird (Agelaius phoeniceus (Linnaeus, 1766)) nests. Our top models included host species (cowbirds grew faster in sparrow nests), numbers of nestmates (slowest when raised alone), and sex (males grew faster). These results confirm that multiple social and environmental factors predict directional patterns of developmental plasticity in avian generalist brood parasites. 


Social interactions do not drive territory aggregation in a grassland songbird

Ecology 2020coauthors: S.M. Munguía, E.J. Williams, and W.A. Boyle


Understanding the drivers of animal distributions is a fundamental goal of ecology and informs habitat management. The costs and benefits of colonial aggregations in animals are well established, but the factors leading to aggregation in territorial animals remain unclear. Territorial animals might aggregate to facilitate social behavior such as (1) group defense from predators and/or parasites, (2) cooperative care of offspring, (3) extra-pair mating, and/or (4) mitigation of extra-pair mating costs through kin selection. Using experimental and observational methods, we tested predictions of all four hypotheses in a tallgrass prairie in northeast Kansas, United States. Grasshopper Sparrow (Ammodramus savannarum) males formed clumps of territories in some parts of the site while leaving other apparently suitable areas unoccupied. Despite substantial sampling effort (653 territories and 223 nests), we found no support for any hypothesized social driver of aggregation, nor evidence that aggregation increases nest success. Our results run counter to previous evidence that conspecific interactions shape territory distributions. These results suggest one of the following alternatives: (1) the benefits of aggregation accrue to different life‐history stages, or (2) the benefits of territory aggregation may be too small to detect in short‐term studies and/or the consequences of aggregation are sufficiently temporally and spatially variable that they do not always appear to be locally adaptive, perhaps exacerbated by changing landscape contexts and declining population sizes. Check it out here!

aggregation comic.jpg

Published Collaborations (side projects):

Effects of patch-burn grazing on reproductive success of grassland songbirds

B.H.F. Verheijen, A.N. Erickson, W.A. Boyle, K.S. Leveritte, J.L. Sojka, L.A. Spahr, C.C. Tehn, E.J. Williams, S.K. Winnicki, and B.K. Sandercock, Ornithological Applications 2022, check it out here!


Referential alarm calling elicits future vigilance in a host of an avian brood parasite

S.L. Lawson, J.K. Enos, C.S. Wolf, K. Stenstrom, S.K. Winnicki, T.J. Benson, M.E. Hauber, and S.A. Gill, Biology Letters 2021, check it out here! Popular science article here!

The limits of egg recognition: testing acceptance thresholds of American robins in response to decreasingly egg-shaped objects in the nest

M.E. Hauber, S.K. Winnicki, J.P. Hoover, D. Hanley, and I.R. Hays,

Royal Society Open Science, 2021, check out the New York Times feature here and the paper here!


My undergraduate biology research with T.D. Schultz and W.A. Boyle became the sparrow aggregation manuscript ultimately published in Ecology (see above). My history research with A.J. Davis is available at the Denison University Library or below: 

The Ideals of Female Sanctity in Merovingian and Carolingian Vitae

My undergraduate history research explored the ways that "holiness" was expressed in texts from Merovingian and Carolingian Gaul (5th-8th century, modern-day France). These saints' lives, or vitae, were written to commemorate the lives of and campaign for the canonization of deceased individuals. As such, they do not provide us with stories of historical reality, but rather give us insight into the values of the writers at the time. Therefore, I use vitae of female saints (especially the pretty awesome St. Radegund and St. Genovefa) to explore the ways gender identity interacted with religion; what language and motifs were acceptable ways to portray women's holiness? How did that change over time? How did gender and religion interact with other sociopolitical realities at the time? Ultimately I argued that these vitae portray a shift from warrior-like, action-based valor stories to passive, introspective piety as time progressed. This change mirrored the rise of the Church in Gaul; in the early Merovingian dynasty the spreading Church valued miraculous visible saints of all genders, representing the power of Christianity to the masses as the Church expanded. By the end of the Carolingian dynasty Christianity was firmly established in Gaul--at this point the Church did not need saints who showcased the incredible power of the religion, but rather saints who exhibited quiet, pious deference to the Church hierarchy. Please note that I have not revised this in years-- there are likely problematic references to a gender binary throughout the text. Read it here by clicking on the purple box!


St. Radegund, public domain image found here

Undergraduate Research:

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