Thesis (M. Phil.)--University of Hong Kong, 1974. / Offset from typescript.

Thesis (Ph. D.)--University of Hong Kong, 1981.

Predators are a threat to many ground-nesting shorebirds, although it remains largely unknown how they interact with passive predator management techniques such as nest exclosures. I examined the effects of nest exclosures on incubating Piping Plovers (Charadrius melodus melodus) and their predators on nesting beaches in Kouchibouguac and PEI National Parks. A combination of behavioural observations, video monitoring of nests and an artificial nest experiment was used to examine the effects of nest exclosures in this study system. The behaviour of Piping Plovers did not differ between exclosed and unexclosed nests, although different types of predators seemed to have an effect on plover nest attentiveness. Predators visited exclosed nests more often than unexclosed nests and spent more time in the vicinity of exclosed nests than unexclosed nests. Since increased adult mortality and nest abandonment have been documented at exclosed Piping Plover nests, as well as nests of other shorebirds, the results of this study provide evidence of a link between predator harassment and these negative effects.

Thesis (Ph.D.)--University of Hong Kong, 1981. / Also available in print.

Red foxes (Vulpes vulpes) have been identified as a key predator of the threatened piping plover (Charadrius melodus) along the U.S. Atlantic coast. However, little is known about coastal red fox ecology, making it difficult to create effective red fox management strategies in these settings. Here, we quantify aspects of red fox population, spatial, and dietary ecology and interactions with threatened piping plovers on Fire Island, New York. We conducted remote camera surveys, scat and sign surveys, den monitoring, and GPS tracking of red foxes on the island in 2015–2018. We used these data to estimate red fox occupancy, reproduction, survival, and population density. We used GPS data to investigate red fox space use, habitat selection, and responses to piping plover nest exclosure setup, pre-hatch pipping, and hatching. We used fecal dietary analyses and data from den prey item surveys to quantify the frequency of piping plover predation and to identify major prey items of red foxes on the island. Red fox occupancy remained high even after substantial decreases in abundance, population density, annual reproduction, and seasonal survival following 2 sarcoptic mange outbreaks. Within their home ranges, red foxes selected areas that were closer to vegetation during the daytime and twilight hours, but farther from vegetation at night. We did not find clear evidence that red foxes in our study area keyed in on piping plover nest exclosure setup, pipping, or hatching at the spatial scales considered in our comparisons, although fox penetration of and digging at exclosures was an issue in 2015 at Smith Point County Park. Items from Orders Rodentia (rodents, 43% of scats), Coleoptera (beetles, 38%), and Decapoda (crabs and other crustaceans, 29%) were most frequently found in 293 red fox scats examined. Skates (Family Rajidae, 89% of dens with food items) and Atlantic surf clams (Spisula solidissima, 67%) were found most frequently outside of dens. We did not find any identifiable piping plover remains in red fox scats or outside of dens. Our results suggest that direct interactions between red foxes and piping plovers during our study period and in our study area were less frequent than expected, but concurrent work by collaborators documented that the trap success of red foxes was negatively related to piping plover reproductive output during our study period. Lethal removal of red foxes is unlikely to eliminate them from shorebird nesting areas unless complete eradication of foxes from the island can be achieved. We recommend strategic vegetation management in and around piping plover nesting areas to reduce daytime resting areas and hunting cover for red foxes, and continued use of nest exclosures. We also recommend further investigation into indirect impacts of red foxes on piping plover populations, and into the possibility that anthropogenic food resources could be subsidizing the island's red fox population. / Doctor of Philosophy / Red foxes have been identified as a key predator of the piping plover, a small migratory shorebird that breeds along the U.S. Atlantic coast and is considered 'threatened' (at risk of becoming endangered and eventually disappearing) within the United States. The lack of information about red fox ecology in coastal settings has been a challenge for wildlife biologists tasked with reducing predation on piping plovers. We investigated red fox ecology, behavior, and interactions with piping plovers on Fire Island, New York. We used trail cameras, collected scat (feces), monitored dens, and tracked red foxes on the island with global positioning system (GPS) collars in 2015–2018. We used these data to estimate red fox distribution, litter sizes, survival rates, and population sizes. We used GPS data to estimate red fox territory sizes, describe habitat selection, and investigate responses to piping plover nest exclosure setup, pipping (a period before hatching during which chicks vocalize inside the eggs), and hatching. We dissected red fox scats and recorded prey items found outside of dens to determine what red foxes on the island were eating. The proportion of each study area used by red foxes remained high even after substantial decreases in abundance, population density, annual reproduction, and seasonal survival following 2 parasitic disease (sarcoptic mange) outbreaks. Within their territories, red foxes selected areas that were closer to vegetation during the daytime and twilight hours but farther from vegetation at night. We did not find clear evidence that red foxes in our study area keyed in on piping plover nest exclosure setup, pipping, or hatching, although fox penetration of and digging at exclosures was an issue in some years at a site not included in those comparisons. Rodents, beetles, and crustacean remains were found most frequently in red fox scats. Skates and surf clams were found most frequently outside of dens. We did not find any identifiable piping plover remains in red fox scats or outside of dens. Our results suggest that that direct interactions between red foxes and piping plovers may be less frequent than previously believed, but concurrent work by collaborators documented that the trap success of red foxes was negatively related to piping plover reproductive output during our study period. Lethal removal of red foxes is unlikely to eliminate red foxes from shorebird nesting areas unless all foxes on the island are removed. We recommend strategic vegetation removal in and around piping plover nesting areas to reduce daytime resting spots and hunting cover for red foxes, and continued use of nest exclosures. We also recommend further investigation into indirect impacts of red foxes on piping plover populations, and into the possibility that anthropogenic food resources could be subsidizing the island's red fox population.

There is little information on demographic trends of, or threats to the Wilson's Plover (Charadrius wilsonia), despite concerns that habitat degradation may be causing the species to decline. I studied Wilson's Plover demography, nest site selection, and physiological and behavioral responses to overflights at Cape Lookout National Seashore, North Carolina, where the National Park Service and U.S. Marine Corps recently lowered the altitude for overflights at tactical speeds. I monitored the responses of incubating Wilson's Plovers to overflights and other human activities and compared heart rate, incubation rate, and vigilance behaviors during time periods with and without these stimuli. I compared habitat use vs. availability at the island-scale, and nest placement relative to geomorphic features and vegetative cover, both on a local and island-wide scale. Wilson's Plovers increased vigilance during military rotary-wing and civilian fixed-wing overflights, but not during military fixed-wing overflights. Plovers were vigilant more and incubated less when researchers were present. Wilson's Plovers selected for interdune areas, flats and isolated dunelets on flats, and against beach and dunes. At the local scale, nests were more likely to be near dense vegetation than random points. Mayfield nest survival was 25%, and predation caused most of the nest failure. Daily nest survival was negatively correlated with nest age and initiation date. Nests with cameras and heart rate monitors had lower survival, and nests with predator exclosures had higher survival. The mean number of chicks fledged per pair was 0.78. / Master of Science

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

Thesis (M. Sc.)--University of Hong Kong, 1995. / Includes bibliographical references (leaves 78-80).

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.