Prairie rivers of the Great Plains, USA, provide important habitat for the federally threatened piping plover (“plover”, Charadrius melodus). Plovers nest on open to sparsely vegetated river sandbars, and their demographic rates are closely linked to habitat availability and quality, as well as river flow. The Niobrara River in northern Nebraska has supported 22–41% of the state’s plovers since species listing in 1986, but the population and habitat are relatively understudied, and both have declined since 2010. The objectives of this study were to understand plover demography, habitat, and the role of the Niobrara in the regional plover population. Periods of high river flow promote creation and maintenance of suitable sandbar nesting habitat, but increased river flow during the plover breeding season can decrease nest and chick survival. We estimated the effect of daily peak river flow on survival rates of 115 nests and 66 chicks on the Niobrara River, 2010–2016, using logistic exposure and Cormack-Jolly-Seber models, respectively. We monitored 1,874 banded hatch-year and adult birds across the regional population (Niobrara River, Lewis and Clark Lake, Gavins Point Reach segment of the Missouri River), and used multi-state mark-recapture models to estimate survival and inter-annual dispersal probabilities among sites relative to habitat availability. We developed land cover datasets from high-resolution aerial imagery to quantify suitable habitat and compare the relative effects of habitat characteristics on nest-site selection and nest success for a subset of years (2010, 2012, 2014, and 2016) using logistic regression models. We included data from a sympatric nester with similar nesting habitat needs, the interior least tern (“tern”, Sternula antillarum athalassos), to improve precision of our models. We compared 63 plover and 92 tern nests to 292 random unused points, and 73 successful (hatched ≥1 egg) to 79 failed nests. Low nest and chick survival and high emigration from the Niobrara appear to be important factors contributing to population decline. Daily nest and chick survival were negatively related to river flow. Nest-site selection was based primarily on distance to the river bank (i.e., the nearest potential source of predators), yet flooding (eggs submerged or washed out of the nest bowl during increased river flow) caused at least as many nest failures as predation. Nests predominantly were surrounded by dry sand habitat, indicating some degree of flood avoidance, but were no farther from water than random, and drier nest sites were no less likely to fail. Dispersal occurred throughout the regional population, but plovers were more likely to leave the Niobrara than to enter it. Expansive flood-created sandbars on the Missouri River, concurrent with a trend towards more vegetated and saturated habitat on the Niobrara, may have drawn birds from the Niobrara population, especially those that dispersed to the Niobrara during sustained Missouri River flooding 2010–2011. The outsized negative effect of flooding on nest success, the lack of protection afforded by dry sand nest sites, and selection for nesting habitat based more strongly on predator avoidance than flood avoidance suggest that plovers may have face more frequent and intense levels of breeding season flooding than is typical. Identifying and promoting the processes that contribute to creation and maintenance of high-elevation sandbars on the Niobrara is an important next step towards effective management of nesting birds. / Master of Science

Habitat loss and predation are threatening many shorebird populations worldwide. While habitat preservation often is preferable, sometimes habitat needs to be restored or created in order to stave off immediate declines. The Great Plains population of piping plovers (Charadrius melodus) was listed as threatened in 1986, and habitat loss and predation appear to be limiting the growth of this population. On the Missouri River, piping plovers nest on sandbars, but the damming of the mainstem of the Missouri in the mid-twentieth century reduced the natural capacity of the Missouri River to create sandbar habitat. In 2004, the United States Army Corps of Engineers (USACE) implemented a habitat creation project on the Gavins Point Reach of the Missouri River (stretch of river immediately downriver from the Gavins Point Dam) in an effort to promote recovery of piping plovers and the endangered least tern (Sternula antillarum). The USACE built 3 sandbars in 2004 – 2005 and built another sandbar on Lewis and Clark Lake in 2007. We studied the population dynamics of piping plovers in relationship to this newly engineered habitat. We monitored 623 nests on 16 sandbar complexes, to evaluate habitat selection, determine the factors affecting nesting success, and compare nesting success between natural and engineered habitat. From these 623 nests, we banded 357 adults and 685 chicks to investigate the factors affecting adult and juvenile survival. We used a logistic-exposure model to calculate nest survival. Adult and juvenile survival was calculated using Cormack-Jolly-Seber based models in Program MARK. We used the estimates from these studies to create a matrix population model for piping plovers nesting on the Gavins Point Reach. We used this model to predict the effects of engineered habitat on the population growth rate. Piping plovers selected for engineered sandbars and against natural and natural/modified habitats. Daily survival rate (DSR) on engineered habitats was significantly higher than on natural or natural modified habitats (log odds: 2.71, 95% CI: 1.20 – 6.08). Predator exclosures around nests did not affect DSR after controlling for the effects of date, nest age, and clutch size. Piping plover juvenile survival to recruitment was negatively related to nesting density on the relatively densely populated engineered sandbars. On the less dense natural sandbars, survival to recruitment was positively correlated with density. Adult survival did not appear to be related to density within our study. Movement within the study area was related also to density. Juveniles from densely populated engineered sandbars were more likely to leave engineered habitat to nest on natural sandbars than were juveniles hatched on less densely populated engineered sandbars. Movements among sandbars by breeding adults suggested that adults preferred engineered habitat. It is possible that juveniles moved to natural habitats because they were unable to compete with adults for the more desirable engineered habitats. Adults and juveniles emigrated from the study area at a higher rate after the 2006 breeding season, a year when water discharge was higher, nesting densities were higher, and reproductive success was lower (as a result of predation) than in the other years. Deterministic modeling suggested that engineered habitat significantly increased population growth. Decreased productivity over time and associated predicted negative population growth suggest that the amount of engineered habitat created was inadequate to sustain population growth, and/or that relatively high water discharge and nesting densities coupled with low reproductive rates and high emigration rates could lead to rapid declines in the plover population. Continued research is needed to determine the effects of these factors on long-term population growth. Our results suggest that habitat creation could be a viable short-term solution to population declines in shorebird populations limited by habitat loss, but high densities and increased predation associated with habitat creation indicate that other, long-term solutions may be required. / Ph. D.

Globally, riparian ecosystems are in decline due to anthropogenic modifications including damming, channelization and the conversion of the floodplain for human use. These changes can profoundly affect riparian species as many have adapted to the historical dynamism of these ecosystems. On the managed Missouri River, an imperiled shorebird, the piping plover (Charadrius melodus) uses riverine sandbars to breed. From 2004 to 2009, due to limited breeding habitat and low population numbers, the U.S. Army Corps of Engineers constructed 255 ha of sandbar habitat to benefit piping plovers and least terns (Sternula antillarum). During the breeding seasons of 2010 and 2011, historically high flows resulted in the creation of 1,887 ha of suitable sandbar habitat. Our study compared the demographic response and the condition of piping plovers to these anthropogenic and natural habitat creation events. From 2005–2014 we monitored 1,071 nests, and from those nests we uniquely banded 968 adults and 2,021 piping plover chicks. We obtained 405 egg (clutch) mass measurements, 1,285 mass measurements from 633 adults, and 7,093 mass measurements from 1,996 plover chicks resulting in 3,175 mass measurements from 654 broods of chicks. We also collected 3,347 invertebrate prey samples. We used a random effects logistic exposure model to estimate nest success, a random effects Cormack-Jolly-Seber model in RMARK to estimate pre-fledge chick survival and the Barker model in RMARK to estimate hatch-year (HY) and after hatch-year (AHY) survival and fidelity to our study area. We then used estimates from these analyses to calculate reproductive output, reproductive output necessary for a stationary population, and population growth (λ). For adult condition and egg (clutch) mass we used generalized linear mixed regression, and for pre-fledge chick growth rates we used a modified Richard's model to estimate the effects of habitat type (pre- vs. post-flood). We also tested for differences in invertebrate prey abundance between habitat types using negative binomial regression. Our results indicated that AHY survival varied throughout our study and was lowest during the flood (2010 and 2011). We found that nest success, pre-fledge chick survival, reproductive output, and HY survival and fidelity were consistently higher on the flood-created habitat than engineered habitat, leading to sustained population growth after the flooding, as compared to just one year of population growth prior to the flood. Unlike pre-flood engineered habitat, the demographic parameters we measured did not decrease as the post-flood habitat aged. These differences were related to increased sandbar habitat, low nesting densities, and decreased nest and chick predation on the post-flood habitat. Although we hypothesized that increased demographic rates would be reflected by increased piping plover condition following the flood, we found that our measured condition variables (adult mass, clutch mass, and pre-fledge chick growth rates) remained unchanged following the flood. We also found evidence that clutch mass, chick growth rates and invertebrate prey abundance decreased as the post-flood sandbar habitat aged. As the condition of individuals did not appear to contribute directly to the increased demographic rates following the flood, we suggest that the change in density-dependent predation pressure may explain the discrepancy. As many ecosystems have previously been altered, it's rare that ecologists have the opportunity to compare management practices with natural ecosystem processes. Results from this study suggest that management intervention may not be an equivalent substitute for natural ecosystem processes and provide insight on future management of riparian ecosystem. / Master of Science

The piping plover (Charadrius melodus) is a federally-listed shorebird that nests on barrier islands along the U.S. Atlantic Coast and is highly vulnerable to habitat change and predation. We have addressed these two threats by 1) developing and implementing a linked model system that predicts future change to piping plover habitat resulting from sea-level rise and beach management efforts by joining dynamic models of sea-level rise, shoreline change, island geomorphology and piping plover nest habitat suitability, and 2) quantifying occupancy and movement of the red fox (Vulpes vulpes), a key shorebird predator at Assateague Island, Maryland and Virginia. We constructed and tested a model that links changes in geomorphological characteristics to piping plover nesting habitat suitability. We then linked this model to larger scale shoreline change resulting from sea level rise and storms. Using this linked model to forecast future sea-level rise and beach management efforts, we found that modest sea-level rise rates (3 mm and 4.1 mm/yr; similar to current rates) may increase suitable piping plover nesting habitat area in 50-100 years and some beach management strategies (beach nourishment and artificial dune modifications) also influence habitat availability. Our development and implementation of this tool to predict change in piping plover habitat suitability provides a vital starting point for predicting how plover nesting habitat will change in a context of planned human modifications intended to address climate change-related threats. Our findings regarding red fox occupancy and movement complement the use of this model for planning future management actions by providing vital information on the effects of certain predator management activities and habitat use of a key mammalian predator, the red fox, for shorebirds along the U.S. Atlantic Coast. Overall, we found that 1) red fox occupancy was strongly tied to eastern cottontail (Sylvilagus floridanus) trap success, increasing sharply with increased eastern cottontail trap success, 2) red fox occupancy did not change in response to an intensive eradication program, and 3) red foxes in our study area generally moved little between camera stations spaced 300 m from each other, but may move large distances (> 6km) at times, likely to occupy new territory available after lethal control efforts. Our findings have important ramifications for the sustainability of long-term predator removal programs and our understanding of future habitat change on the red fox. For example how vegetation changes affect eastern cottontails, how resulting fluctuations in eastern cottontails affect red fox occupancy, and how consequential changes in red fox occupancy affect plover breeding productivity. Our predictive model combined with these predator findings will allow wildlife managers to better plan and implement effective management actions for piping plovers in response to the multiple stressors of SLR-induced habitat change and predation. / Ph. D.

Snowy plovers (Charadrius nivosus) are small, ground-nesting shorebirds that are a species of conservation concern throughout North America. Despite increased efforts to understand factors contributing to the decline of snowy plover, little is known about habitat selection and breeding ecology of snowy plover for the large population found in the Great Basin. We tested hypotheses concerning the occupancy and nesting success of snowy plover. First, we identified factors influencing snowy plover nest survival at Great Salt Lake, Utah. We hypothesized that snowy plover would demonstrate differences in nest survival rates across years due to differences in habitat characteristics, predator abundance, human influence, resource availability, and fluctuating water levels. We conducted nest surveys at five sites along the Great Salt Lake to locate new nests or monitor known nests until nest fate was determined. We found 608 nests between 2003, 2005-2010, and 2012. The most common cause of nest failure was predation, followed by weather, abandonment, and trampling. Nest survival estimates ranged from 4.6 -- 46.4% with considerable yearly variation. There was no correlation between researcher activity (visits to nests and trapping of adults) and nest survival. Nests in close proximity to roads had lower survival than nests far from roads. Nests located on barren mudflats also had lower survival than nests in vegetated areas or near debris. We found that nests had a higher probability of survival as they increased in incubation stage. Because nesting areas around the Great Salt Lake host some of the largest concentrations of breeding snowy plover in North America, we suggest that managers consider measures to maintain suitable nesting habitat for snowy plover. Second, we determined factors affecting snowy plover occupancy and detection probabilities in western Utah between 2011 and 2012. We hypothesized that snowy plover would be associated with spring water flows and sparsely vegetated salt flats. We made repeated visits to randomly selected survey plots recording the number of snowy plover adults and habitat characteristics within each plot. We modeled the relationship between snowy plover detection probability and habitat and environmental characteristics. The detection probability was 77% (95% CI = 64 -- 86%) and did not vary by year. There was a positive relationship between ambient temperature and detection probability. Next, we modeled the relationship between snowy plover occupancy and individual habitat characteristics including distance to water, distance to roads, land cover types, and vegetative characteristics. Snowy plover occupancy did not vary by year and was estimated at 12% (95% CI = 7 -- 21%). Occupancy was best predicted by close proximity to water, playa land cover, and minimal shrub cover. We used habitat characteristics that best predicted snowy plover occupancy to generate a predictive habitat model that can help prioritize future snowy plover surveys and guide conservation efforts.

The Wilson’s Plover (Charadrius wilsonia) is a species of concern in most southeastern U.S. coastal states, where it breeds and winters. The U.S. Shorebird Conservation Plan listed this species as a Species of High Concern (Prioritization Category 4), and the U.S. Fish and Wildlife Service has designated it as a Bird of Conservation Concern (BCC). Despite its conservation status, Wilson’s Plover population trends are poorly understood and little research has been conducted examining habitat factors affecting this species’ breeding and foraging ecology. I collected Wilson’s Plover demographic data and explored which habitat characteristics influenced breeding success and foraging site selection among three coastal habitat types (i.e. fiddler crab (Uca spp.) mud flats, beach front, and interdune sand flats) at Onslow Beach, Marine Corps Base Camp Lejeune, North Carolina, 2008-2009. I observed little difference between years in nest success (≥ 1 egg hatched), failure, and overall nest survival. The majority of nest failures were caused by mammalian predators. For those nests that hatched successfully, greater proportions were located in clumped vegetation than on bare ground or sparsely vegetated areas. In-season chick survival for both years was higher for nests that hatched earlier in the season, and for nests farthest from the broods’ final foraging territory. Productivity estimates (chicks fledged per breeding pair) were not significantly different between years (0.88 ± 0.26 fledged/pair in 2008, 1.00 ± 0.25 fledged/pair in 2009) despite a shift in foraging behavior, possibly related to habitat alterations and availability in 2009. My findings indicate that Wilson’s Plover adults and broods were flexible in establishing final foraging territories; in 2008 all final brood foraging territories were on fiddler flats while in 2009, final foraging territories were sometimes split between fiddler flats, beach front, and interdune sand flats. For those Wilson’s Plovers establishing territories on fiddler flats, area of the flat was the most important feature explaining use versus non-use of a particular flat; area ≥ 1250 m² was preferred. Close proximity to water and vegetative cover were also important habitat features in foraging site selection on fiddler crab mud flats, and in all habitat types combined. My findings will directly contribute to population and habitat research goals outlined in the U.S. Shorebird Plan and will supplement limited data about foraging and habitat use related to Wilson’s Plover breeding ecology. / Master of Science

Biological invasions provide a unique opportunity to study the mechanisms that regulate community composition and ecosystem function. Invasive species that are also ecosystem engineers can substantially alter physical features in an environment, and this can lead to cascading effects on the biological community. Aquatic-terrestrial interface ecosystems are excellent systems to study the interactions among invasive ecosystem engineers, physical features, and biological communities, because interactions among vegetation, sediment, and fluids within biophysical feedbacks create and modify distinct physical features. Further, these systems provide important ecosystem services including coastal protection afforded by their natural features. In this dissertation, I investigate the interactions and feedbacks among sand-binding beach grass species (a native, Elymus mollis (Trin.), and two non-natives, Ammophila arenaria (L.) Link and A. breviligulata Fernald), sediment supply, and dune shape along the U.S. Pacific Northwest coast. Dunes dominated by A. arenaria tend to be taller and narrower compared to the shorter, wider dunes dominated by A. breviligulata. These patterns suggest an ecological control on dune shape, and thus, coastal vulnerability to overtopping waves. I investigate the causes and consequences of these patterns with experiments, field observations, and modeling. Specifically, I investigate the relative roles of vegetation and sediment supply in shaping coastal dunes over inter-annual and multi-decadal time scales (Chapter 2), characterize a biophysical feedback between beach grass species growth habit and sediment supply (Chapter 3), uncover the mechanisms leading to beach grass coexistence and whether A. breviligulata can invade and dominate new sections of coastline (Chapter 4), and examine the non-target effects resulting from management actions that remove Ammophila for the recovery of the threatened Western Snowy plover (Charadrius alexandrinus nivosus) (Chapter 5). I found that vegetation and sediment supply play important roles in dune shape changes across inter-annual and multi-decadal time scales (Chapter 2). I determined that a biophysical feedback between the beach grass growth habits and sediment supply results in species-specific differences in sand capture ability, and thus, is a likely explanation for differences in dune shape (Chapter 3). I found that all three beach grass species can coexist across different sediment deposition rates, and that this coexistence is largely mediated by positive direct and indirect species interactions. I further determined that A. breviligulata is capable of invading and dominating the beach grass community in regions where it is currently absent (Chapter 4). Combined, these findings indicate that A. breviligulata is an inferior dune building species as compared to A. arenaria, and suggest that in combination with sediment supply gradients, these species differences ultimately lead to differences in dune shape. Potential further invasions of A. breviligulata into southern regions of the Pacific Northwest may diminish the coastal protection ability of dunes currently dominated by A. arenaria, but this effect could be moderated by the predicted near co-dominance of A. arenaria in these lower sediment supply conditions. Finally, I found that the techniques used to remove Ammophila for plover recovery have unintended consequences for the native and endemic dune plant communities, and disrupt the natural disturbance regime of shifting sand. A whole-ecosystem restoration focus would be an improvement over the target-species approach, as it would promote the return of the natural disturbance regime, which in turn, would help recover the native biological community. The findings from this dissertation research provide a robust knowledge base that can guide further investigations of biological and physical changes to the coastal dunes, can help improve the management of dune ecosystem services and the restoration of native communities, and can help anticipate the impacts of future beach grass invasions and climate change induced changes to the coast. / Graduation date: 2012 / Access restricted to the OSU Community at author's request from Sept. 22, 2011 - March 22, 2012