Walter H. Piper
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Research Subjects
Research Interests
Walter H. Piper has a total of 43 co-authors. Here are the top co-authors by number of shared publications:
- Brian A. Hoover
Farallon Institute, Petaluma, USA
3 shared publications Michael W. Meyer
NOVA Ecological Services Arbor Vitae WI USA
3 shared publicationsGabriella Jukkala
Department of Natural Resources University of Illinois Champaign IL USA
2 shared publicationsKeren B. Tischler
Biology Program, Northland College, Ashland, Wisconsin, USA
2 shared publicationsKristin M. Brunk
Northland College
2 shared publicationsNathan Banfield
Rainbow Circle Montgomery City MO USA
2 shared publications
Latest Publications
- Open Access
- 22 September 2026
- Published by Oxford University Press (OUP) in Behavioral Ecology
- Vol. 37 (6)
Shoreline development influences common loon territory selection but not extent of nest defense
Shoreline development of temperate lakes has increased steadily in recent decades, putting pressure on animal populations that live in lacustrine habitats. Common loons (Gavia immer) nest along lakeshores, and shoreline development has been identified as a likely cause of population decline in this species. Taking advantage of a marked loon population from a long-term study in northern Wisconsin, USA between 1992 and 2023, we investigated impacts of shoreline development and human disturbance on territory choice and nest defense behavior of breeding loons. Loons settled on territories that resembled their natal lakes in terms of shoreline development, an indication of natal habitat imprinting. Shoreline development did not affect the distance at which loons flushed from their nest at the approach of humans. However, nesting loons were less likely to flush from the nest late in the incubation period, suggesting a tendency for loons to defend nests according to likely fitness benefits. Loons' tendency to breed on lakes similar in human activity to their natal lake might reflect their effort to seek out lakes of familiar fish species composition rather than human activity. Natal habitat imprinting of this kind illustrates the potential of anthropogenic factors to affect behavior and, in turn, population dynamics.
- 22 September 2026
- Published by Oxford University Press (OUP) in Behavioral Ecology
- Vol. 37 (6)
Open AccessShoreline development influences common loon territory selection but not extent of nest defense
Shoreline development of temperate lakes has increased steadily in recent decades, putting pressure on animal populations that live in lacustrine habitats. Common loons (Gavia immer) nest along lakeshores, and shoreline development has been identified as a likely cause of population decline in this species. Taking advantage of a marked loon population from a long-term study in northern Wisconsin, USA between 1992 and 2023, we investigated impacts of shoreline development and human disturbance on territory choice and nest defense behavior of breeding loons. Loons settled on territories that resembled their natal lakes in terms of shoreline development, an indication of natal habitat imprinting. Shoreline development did not affect the distance at which loons flushed from their nest at the approach of humans. However, nesting loons were less likely to flush from the nest late in the incubation period, suggesting a tendency for loons to defend nests according to likely fitness benefits. Loons' tendency to breed on lakes similar in human activity to their natal lake might reflect their effort to seek out lakes of familiar fish species composition rather than human activity. Natal habitat imprinting of this kind illustrates the potential of anthropogenic factors to affect behavior and, in turn, population dynamics.
- Open Access
- 24 November 2025
- Published by Springer Nature in Oecologia
- Vol. 207 (12) , 196
A silver spoon effect reduces lifetime fitness in a declining loon population
A complete understanding of factors that influence animal fitness requires that we measure not only those occurring day to day in the life of an animal but also those that operate on longer time scales. Here, we investigated silver spoon effects (fitness impacts resulting from conditions faced early in life) and carryover effects (fitness impacts caused by environmental factors in a previous season) in a northern Wisconsin population of the common loon ( Gavia immer ). The mass of a loon chick divided by its age, an indication of food it received from its parents in its first 4 to 6 weeks of life (“chick condition”), affected both the likelihood of survival to adulthood and, among territory settlers, the number of chicks it fledged as an adult. Only one carryover effect was evident: increased ocean pH on the wintering ground had a modest positive effect on territory settlement rate. However, cohorts of loons that faced unfavorable ocean conditions in their first year yielded adults that fledged many chicks, which suggests that selection resulting from poor ocean conditions removed weaker phenotypes. The robust silver spoon effect in this species helps us understand a current and alarming pattern in the Wisconsin loon population: the sharp decline in the survival of chicks to breeding age.
- 24 November 2025
- Published by Springer Nature in Oecologia
- Vol. 207 (12) , 196
Open AccessA silver spoon effect reduces lifetime fitness in a declining loon population
A complete understanding of factors that influence animal fitness requires that we measure not only those occurring day to day in the life of an animal but also those that operate on longer time scales. Here, we investigated silver spoon effects (fitness impacts resulting from conditions faced early in life) and carryover effects (fitness impacts caused by environmental factors in a previous season) in a northern Wisconsin population of the common loon ( Gavia immer ). The mass of a loon chick divided by its age, an indication of food it received from its parents in its first 4 to 6 weeks of life (“chick condition”), affected both the likelihood of survival to adulthood and, among territory settlers, the number of chicks it fledged as an adult. Only one carryover effect was evident: increased ocean pH on the wintering ground had a modest positive effect on territory settlement rate. However, cohorts of loons that faced unfavorable ocean conditions in their first year yielded adults that fledged many chicks, which suggests that selection resulting from poor ocean conditions removed weaker phenotypes. The robust silver spoon effect in this species helps us understand a current and alarming pattern in the Wisconsin loon population: the sharp decline in the survival of chicks to breeding age.
- Open Access
- 6 May 2024
- Published by Wiley in The Bulletin of the Ecological Society of America
- Vol. 105 (3)
Climate‐Related Loss of Water Clarity Threatens a Northern Icon
- 6 May 2024
- Published by Wiley in The Bulletin of the Ecological Society of America
- Vol. 105 (3)
Open AccessClimate‐Related Loss of Water Clarity Threatens a Northern Icon
- Open Access
Climate change‐associated declines in water clarity impair feeding by common loons
Climate change has myriad impacts on ecosystems, but the mechanisms by which it affects individual species can be difficult to pinpoint. One strategy to discover such mechanisms is to identify a specific ecological factor related to survival or reproduction and determine how that factor is affected by climate. Here we used Landsat imagery to calculate water clarity for 127 lakes in northern Wisconsin from 1995 to 2021 and thus investigate the effect of clarity on the body condition of an aquatic visual predator, the common loon (Gavia immer). In addition, we examined rainfall and temperature as potential predictors of water clarity. Body mass tracked July water clarity strongly in loon chicks, which grow chiefly in that month, but weakly in adult males and females. Long‐term mean water clarity was negatively related to chick mass but positively related to adult male mass, suggesting that loons foraging in generally clear lakes enjoy good foraging conditions in the long run but might be sensitive to perturbations in clarity during chick‐rearing. Finally, chick mass was positively related to the density of docks, perhaps because angling removes large fishes and thus boosts the abundance of the small fishes on which chicks depend. Water clarity itself declined strongly from 1995 to 2021, was negatively related to July rainfall, and was positively related to July air temperature. Our findings identified both long‐term and short‐term water clarity as strong predictors of loon foraging efficiency, and suggest that climate change, through water clarity, impacts freshwater ecosystems profoundly. Moreover, our results identified the recent decrease in water clarity as a likely cause of population decline in common loons.
Open AccessClimate change‐associated declines in water clarity impair feeding by common loons
Climate change has myriad impacts on ecosystems, but the mechanisms by which it affects individual species can be difficult to pinpoint. One strategy to discover such mechanisms is to identify a specific ecological factor related to survival or reproduction and determine how that factor is affected by climate. Here we used Landsat imagery to calculate water clarity for 127 lakes in northern Wisconsin from 1995 to 2021 and thus investigate the effect of clarity on the body condition of an aquatic visual predator, the common loon (Gavia immer). In addition, we examined rainfall and temperature as potential predictors of water clarity. Body mass tracked July water clarity strongly in loon chicks, which grow chiefly in that month, but weakly in adult males and females. Long‐term mean water clarity was negatively related to chick mass but positively related to adult male mass, suggesting that loons foraging in generally clear lakes enjoy good foraging conditions in the long run but might be sensitive to perturbations in clarity during chick‐rearing. Finally, chick mass was positively related to the density of docks, perhaps because angling removes large fishes and thus boosts the abundance of the small fishes on which chicks depend. Water clarity itself declined strongly from 1995 to 2021, was negatively related to July rainfall, and was positively related to July air temperature. Our findings identified both long‐term and short‐term water clarity as strong predictors of loon foraging efficiency, and suggest that climate change, through water clarity, impacts freshwater ecosystems profoundly. Moreover, our results identified the recent decrease in water clarity as a likely cause of population decline in common loons.
- 2 February 2024
- Published by Elsevier in Animal Behaviour
- Vol. 209, 169-177
Honest signalling of age in a territorial breeder
- 2 February 2024
- Published by Elsevier in Animal Behaviour
- Vol. 209, 169-177
Honest signalling of age in a territorial breeder
- 23 September 2023
- Published by Elsevier in Animal Behaviour
- Vol. 205, 79-87
Site familiarity makes an enduring impact on breeding success
- 23 September 2023
- Published by Elsevier in Animal Behaviour
- Vol. 205, 79-87
Site familiarity makes an enduring impact on breeding success
- 21 January 2022
- Published by Springer Nature in Behavioral Ecology and Sociobiology
- Vol. 76 (2) , 1-11
Territory holders are more aggressive towards older, more dangerous floaters
- 21 January 2022
- Published by Springer Nature in Behavioral Ecology and Sociobiology
- Vol. 76 (2) , 1-11
Territory holders are more aggressive towards older, more dangerous floaters
- Open Access
- 22 March 2021
- Published by Wiley in Journal of Animal Ecology
- Vol. 90 (5) , 1165-1176
Interrelated impacts of climate and land‐use change on a widespread waterbird
Together climate and land‐use change play a crucial role in determining species distribution and abundance, but measuring the simultaneous impacts of these processes on current and future population trajectories is challenging due to time lags, interactive effects and data limitations. Most approaches that relate multiple global change drivers to population changes have been based on occurrence or count data alone. We leveraged three long‐term (1995–2019) datasets to develop a coupled integrated population model‐Bayesian population viability analysis (IPM‐BPVA) to project future survival and reproductive success for common loons Gavia immer in northern Wisconsin, USA, by explicitly linking vital rates to changes in climate and land use. The winter North Atlantic Oscillation (NAO), a broad‐scale climate index, immediately preceding the breeding season and annual changes in developed land cover within breeding areas both had strongly negative influences on adult survival. Local summer rainfall was negatively related to fecundity, though this relationship was mediated by a lagged interaction with the winter NAO, suggesting a compensatory population‐level response to climate variability. We compared population viability under 12 future scenarios of annual land‐use change, precipitation and NAO conditions. Under all scenarios, the loon population was expected to decline, yet the steepest declines were projected under positive NAO trends, as anticipated with ongoing climate change. Thus, loons breeding in the northern United States are likely to remain affected by climatic processes occurring thousands of miles away in the North Atlantic during the non‐breeding period of the annual cycle. Our results reveal that climate and land‐use changes are differentially contributing to loon population declines along the southern edge of their breeding range and will continue to do so despite natural compensatory responses. We also demonstrate that concurrent analysis of multiple data types facilitates deeper understanding of the ecological implications of anthropogenic‐induced change occurring at multiple spatial scales. Our modelling approach can be used to project demographic responses of populations to varying environmental conditions while accounting for multiple sources of uncertainty, an increasingly pressing need in the face of unprecedented global change.
- 22 March 2021
- Published by Wiley in Journal of Animal Ecology
- Vol. 90 (5) , 1165-1176
Open AccessInterrelated impacts of climate and land‐use change on a widespread waterbird
Together climate and land‐use change play a crucial role in determining species distribution and abundance, but measuring the simultaneous impacts of these processes on current and future population trajectories is challenging due to time lags, interactive effects and data limitations. Most approaches that relate multiple global change drivers to population changes have been based on occurrence or count data alone. We leveraged three long‐term (1995–2019) datasets to develop a coupled integrated population model‐Bayesian population viability analysis (IPM‐BPVA) to project future survival and reproductive success for common loons Gavia immer in northern Wisconsin, USA, by explicitly linking vital rates to changes in climate and land use. The winter North Atlantic Oscillation (NAO), a broad‐scale climate index, immediately preceding the breeding season and annual changes in developed land cover within breeding areas both had strongly negative influences on adult survival. Local summer rainfall was negatively related to fecundity, though this relationship was mediated by a lagged interaction with the winter NAO, suggesting a compensatory population‐level response to climate variability. We compared population viability under 12 future scenarios of annual land‐use change, precipitation and NAO conditions. Under all scenarios, the loon population was expected to decline, yet the steepest declines were projected under positive NAO trends, as anticipated with ongoing climate change. Thus, loons breeding in the northern United States are likely to remain affected by climatic processes occurring thousands of miles away in the North Atlantic during the non‐breeding period of the annual cycle. Our results reveal that climate and land‐use changes are differentially contributing to loon population declines along the southern edge of their breeding range and will continue to do so despite natural compensatory responses. We also demonstrate that concurrent analysis of multiple data types facilitates deeper understanding of the ecological implications of anthropogenic‐induced change occurring at multiple spatial scales. Our modelling approach can be used to project demographic responses of populations to varying environmental conditions while accounting for multiple sources of uncertainty, an increasingly pressing need in the face of unprecedented global change.
- Open Access
- 28 December 2020
- Published by Wiley in Ecology and Evolution
- Vol. 11 (3) , 1310-1319
Early evidence of natal‐habitat preference: Juvenile loons feed on natal‐like lakes after fledging
Many species show natal habitat preference induction (NHPI), a behavior in which young adults select habitats similar to those in which they were raised. However, we know little about how NHPI develops in natural systems. Here, we tested for NHPI in juvenile common loons (Gavia immer) that foraged on lakes in the vicinity of their natal lake after fledging. Juveniles visited lakes similar in pH to their natal lakes, and this significant effect persisted after controlling for spatial autocorrelation. On the other hand, juveniles showed no preference for foraging lakes of similar size to their natal one. When lakes were assigned to discrete classes based on size, depth, visibility, and trophic complexity, both juveniles from large lakes and small lakes preferred to visit large, trophically diverse lakes, which contained abundant food. Our results contrast with earlier findings, which show strict preference for lakes similar in size to the natal lake among young adults seeking to settle on a breeding lake. We suggest that NHPI is relaxed for juveniles, presumably because they select lakes that optimize short‐term survival and growth. By characterizing NHPI during a poorly studied life stage, this study illustrates that NHPI can take different forms at different life stages.
- 28 December 2020
- Published by Wiley in Ecology and Evolution
- Vol. 11 (3) , 1310-1319
Open AccessEarly evidence of natal‐habitat preference: Juvenile loons feed on natal‐like lakes after fledging
Many species show natal habitat preference induction (NHPI), a behavior in which young adults select habitats similar to those in which they were raised. However, we know little about how NHPI develops in natural systems. Here, we tested for NHPI in juvenile common loons (Gavia immer) that foraged on lakes in the vicinity of their natal lake after fledging. Juveniles visited lakes similar in pH to their natal lakes, and this significant effect persisted after controlling for spatial autocorrelation. On the other hand, juveniles showed no preference for foraging lakes of similar size to their natal one. When lakes were assigned to discrete classes based on size, depth, visibility, and trophic complexity, both juveniles from large lakes and small lakes preferred to visit large, trophically diverse lakes, which contained abundant food. Our results contrast with earlier findings, which show strict preference for lakes similar in size to the natal lake among young adults seeking to settle on a breeding lake. We suggest that NHPI is relaxed for juveniles, presumably because they select lakes that optimize short‐term survival and growth. By characterizing NHPI during a poorly studied life stage, this study illustrates that NHPI can take different forms at different life stages.
- Open Access
- 26 August 2020
- Published by Oxford University Press (OUP) in Ornithological Applications
- Vol. 122 (4)
Plunging floater survival causes cryptic population decline in the Common Loon
Populations of many vertebrates are declining and geographic ranges contracting, largely as a consequence of anthropogenic threats. Many reports of such decline, however, lack the breadth and detail to narrow down its causes. Here we describe population decline in the Common Loon (Gavia immer), a charismatic aquatic bird, based on systematic resighting and measurement of a marked population. During our 27-year investigation, age-adjusted chick mass has fallen by 11%, mortality among young and old chicks has increased by 31% and 82%, respectively, and fledging success has declined by 26%. Meanwhile, the return rate of marked nonbreeders (“floaters”) has plunged by 53%, and the adult population overall has declined by 22%. Consistent with the thinning ranks of floaters, the rate of territory eviction has decreased by 52% during the study. Despite the decline in floaters, territory occupancy remains unchanged. However, a matrix model, updated with recent estimates for breeding success, juvenile survival, and senescence, yields a recalculated deterministic population growth rate (λ) of 0.94 for our study population, which suggests that declines in vital rates could lead to a loss of 52% of the current population and a decline of 37% in territory occupancy by 2031. Lack of data on floaters in other upper Midwest and New England loon populations leaves their status in doubt. • The adult loon population in northern Wisconsin has declined by 22% over the past 27 years. • Numbers of chicks and chick mass have declined significantly. • The population of young nonbreeders (floaters) has plummeted by 46%. • Despite these steep declines, there has been no measurable drop in the number of territorial pairs. • The hidden nature of the Wisconsin decline points out that floater populations can mask major drops in breeding populations and throws into question population trends in other loon populations, whose floater populations are largely unknown.
- 26 August 2020
- Published by Oxford University Press (OUP) in Ornithological Applications
- Vol. 122 (4)
Open AccessPlunging floater survival causes cryptic population decline in the Common Loon
Populations of many vertebrates are declining and geographic ranges contracting, largely as a consequence of anthropogenic threats. Many reports of such decline, however, lack the breadth and detail to narrow down its causes. Here we describe population decline in the Common Loon (Gavia immer), a charismatic aquatic bird, based on systematic resighting and measurement of a marked population. During our 27-year investigation, age-adjusted chick mass has fallen by 11%, mortality among young and old chicks has increased by 31% and 82%, respectively, and fledging success has declined by 26%. Meanwhile, the return rate of marked nonbreeders (“floaters”) has plunged by 53%, and the adult population overall has declined by 22%. Consistent with the thinning ranks of floaters, the rate of territory eviction has decreased by 52% during the study. Despite the decline in floaters, territory occupancy remains unchanged. However, a matrix model, updated with recent estimates for breeding success, juvenile survival, and senescence, yields a recalculated deterministic population growth rate (λ) of 0.94 for our study population, which suggests that declines in vital rates could lead to a loss of 52% of the current population and a decline of 37% in territory occupancy by 2031. Lack of data on floaters in other upper Midwest and New England loon populations leaves their status in doubt. • The adult loon population in northern Wisconsin has declined by 22% over the past 27 years. • Numbers of chicks and chick mass have declined significantly. • The population of young nonbreeders (floaters) has plummeted by 46%. • Despite these steep declines, there has been no measurable drop in the number of territorial pairs. • The hidden nature of the Wisconsin decline points out that floater populations can mask major drops in breeding populations and throws into question population trends in other loon populations, whose floater populations are largely unknown.