Veterinary Preventive Medicine, Epidemiology, and Public Health Commons^™

The Economic Impacts Of Blackbird (Icteridae) Damage To Sunflower In The Usa, Karina Ernst, Julie Elser, George Linz, Hans Kandel, Jason Holderieath, Samantha Degroot, Steven Shwiff, Stephanie Shwiff

USDA Wildlife Services: Staff Publications

BACKGROUND: Blackbird (Icteridae) damage to ripening sunflower (Helianthus annuus L.) has been a persistent economic issue in the USA for the last five decades. To quantify losses, we surveyed blackbird damage from 2001 to 2013 (excluding 2004) to physiologically mature sunflower in eight states: North Dakota, South Dakota, Texas, Nebraska, Minnesota, Colorado, Kansas, and Vermont.

RESULTS:We pooled data gathered during the most recent 5 years (2009 to 2013) of the survey and found losses averaged $US2.5 million and $US11.3 million for confectionery and oilseed hybrids, respectively. Three states, North Dakota, South Dakota, and Nebraska, had sufficient acreage and bird damage …

Habitat Selection By American Beaverat Multiple Spatial Scales, Guiming Wang, Lance F. Mcclintic, Jimmy D. Taylor

USDA Wildlife Services: Staff Publications

Background: Semiaquatic mammals require both aquatic and terrestrial habitats, particularly interfaces between the two habitats. As ecosystem engineers, American beaver (Castor canadensis) consume and fell a great amount of deciduous trees. We tested the prediction that open water and amounts of food resources, including hardwood forests (i.e., deciduous trees as the dominant form of vegetation), herbaceous and woody wetlands, and shrubs, would influence the second-order habitat selection (i.e., placing home ranges on the landscape) by American beaver, whereas the third-order habitat selection of American beaver would be associated with woody wetland and shrub edges. We investigated hierarchical habitat selection by …

Epidemic Growth Rates And Host Movement Patterns Shape Management Performance For Pathogen Spillover At The Wildlife–Livestock Interface, Kezia R. Manlove, Laura M. Sampson, Benny Borremans, E. Frances Cassirer, Ryan S. Miller, Kim M. Pepin, Thomas E. Besser, Paul C. Cross

USDA Wildlife Services: Staff Publications

Managing pathogen spillover at the wildlife–livestock interface is a key step towards improving global animal health, food security and wildlife conservation. However, predicting the effectiveness of management actions across host–pathogen systems with different life histories is an on-going challenge since data on intervention effectiveness are expensive to collect and results are system-specific.We developed a simulation model to explore how the efficacies of different management strategies vary according to host movement patterns and epidemic growth rates. The model suggested that fast-growing, fast-moving epidemics like avian influenza were best-managed with actions like biosecurity or containment, which limited and localized overall spillover risk. …

Board Invited Review: Prospects For Improving Management Of Animal Disease Introductions Using Disease-Dynamic Models, Ryan S. Miller, Kim M. Pepin

USDA Wildlife Services: Staff Publications

Management and policy decisions are continually made to mitigate disease introductions in animal populations despite often limited surveillance data or knowledge of disease transmission processes. Science-based management is broadly recognized as leading to more effective decisions yet application of models to actively guide disease surveillance and mitigate risks remains limited. Disease-dynamic models are an efficient method of providing information for management decisions because of their ability to integrate and evaluate multiple, complex processes simultaneously while accounting for uncertainty common in animal diseases. Here we review disease introduction pathways and transmission processes crucial for informing disease management and models at the …

Improving Risk Assessment Of The Emergence Of Novel Influenza A Viruses By Incorporating Environmental Surveillance, Kim M. Pepin, Matthew W. Hopken, Susan A. Shriner, Erica Spackman, Zaid Abdo, Colin Parrish, Steven Steven, James O. Lloyd-Smith, Antoinette J. Piaggio

USDA Wildlife Services: Staff Publications

Reassortment is an evolutionary mechanism by which influenza A viruses (IAV) generate genetic novelty. Reassortment is an important driver of host jumps and is widespread according to retrospective surveillance studies.However, predicting the epidemiological risk of reassortant emergence in novel hosts from surveillance data remains challenging. IAV strains persist and cooccur in the environment, promoting co-infection during environmental transmission. These conditions offer opportunity to understand reassortant emergence in reservoir and spillover hosts. Specifically, environmental RNA could provide rich information for understanding the evolutionary ecology of segmented viruses, and transform our ability to quantify epidemiological risk to spillover hosts. However, significant challenges …

Individual-Level Antibody Dynamics Reveal Potential Drivers Of Influenza A Seasonality In Wild Pig Populations, Kim M. Pepin, Kerri Pedersen, Xiu-Feng Wan, Fred L. Cunningham, Colleen T. Webb, Mark Q. Wilber

USDA Wildlife Services: Staff Publications

Swine are important in the ecology of influenza A virus (IAV) globally. Understanding the ecological role of wild pigs in IAV ecology has been limited because surveillance in wild pigs is often for antibodies (serosurveillance) rather than IAVs, as in humans and domestic swine. As IAV antibodies can persist long after an infection, serosurveillance data are not necessarily indicative of current infection risk. However, antibody responses to IAV infections cause a predictable antibody response, thus time of infection can be inferred from antibody levels in serological samples, enabling identification of risk factors of infection at estimated times of infection. Recent …

Biology And Impacts Of Pacific Island Invasive Species. 15. Psittacula Krameri, The Rose-Ringed Parakeet (Psittaciformes: Psittacidae)1, Aaron B. Shiels, Nicholas P. Kalodimos

USDA Wildlife Services: Staff Publications

The rose-ringed parakeet (RRP), Psittacula krameri, has become established in at least four Pacific Island countries (Hong Kong China, Japan, New Zealand, U.S.A.), including the Hawaiian islands of Kaua‘i, O‘ahu, and Hawai‘i. Most Pacific islands are at risk of RRP colonization. This species was first introduced to Hong Kong in 1903 and Hawai‘i in the 1930s–1960s, established since 1969 in Japan, and in New Zealand since 2005 where it has repeatedly established after organized removals. The founding birds were imported cage-birds from the pet trade. In native India, RRP are generally found associated with human habitation and are considered a …

Taking The Bait: Species Taking Oral Rabies Vaccine Baits Intended For Raccoons, Betsy S. Haley, Are R. Berentsen, Richard M. Engeman

USDA Wildlife Services: Staff Publications

Raccoon rabies in eastern USA is managed by strategically distributing oral rabies vaccine (ORV) baits. The attractiveness, palativity, density, and non-target species bait take affect ORV effectiveness. We examined raccoon and non-target species differences in investigating/removing fish-meal polymer and coated sachet baits applied to simulate two aerial bait distribution densities. Bait densities of 150 baits/km2 and 75 baits/km2 were evaluated, respectively, in zones expected to have high and low

Racc oon densities. Three primary non-target species visited baits: coyotes, white-tailed deer, and feral swine. The proportion of bait stations visited by raccoons during 1 week observation periods ranged from 50 …

Cause‐Specific Mortality Of The World’S Terrestrial Vertebrates, Jacob E. Hill, Travis L. Devault, Jerrold L. Belant

USDA Wildlife Services: Staff Publications

Aim: Vertebrates are declining worldwide, yet a comprehensive examination of the sources of mortality is lacking. We conducted a global synthesis of terrestrial vertebrate cause‐specific mortality to compare the sources of mortality across taxa and determine predictors of susceptibility to these sources of mortality.

Location: Worldwide.

Time period: 1970–2018.

Major taxa studied: Mammals, birds, reptiles and amphibians.

Methods: We searched for studies that used telemetry to determine the cause of death of terrestrial vertebrates. We determined whether each mortality was caused by anthropogenic or natural sources and further classified mortalities within these two categories (e.g. harvest, vehicle collision and predation). …

Movement Responses Inform Effectiveness And Consequences Of Baiting Wild Pigs For Population Control, Nathan P. Snow, Kurt C. Vercauteren

USDA Wildlife Services: Staff Publications

Wild pigs (Sus scrofa) damage agricultural and natural resources throughout their nearly global distribution. Subsequently, population control activities (e.g., trapping, shooting, or toxic baiting) frequently involve the deployment of bait to attract wild pigs. A better understanding of how wild pigs respond to bait sites can help maximize efficiency of baiting programs and identify any potential pitfalls. We examined the movement behaviors of 68 wild pigs during three stages of intensive baiting programs (i.e., 15 days each: prior, during, and post baiting) spread across two distinct study areas in southern and northern Texas, USA. We found that bait sites needed …

Reducing Prairie Dog Populations And Damage By Castration Of Dominant Males, Gary W. Witmer

USDA Wildlife Services: Staff Publications

Black-tailed prairie dogs (Cynomys ludovicianus) occur widely across the prairie states of North America. They compete with livestock for forage, transmit plague, and damage lawns, landscaping, and property. Interest in non-lethal methods, such as immunocontraception, is growing; however, reductions in the population due to contraception may be offset by increases in survival because adults and yearlings are not subject to the energetic demands of reproduction, and lower densities may increase the amount of resources available to growing offspring. Surgical sterilization provides a means for modeling these effects. Thus, we castrated males prior to the 1998 breeding season to simulate the …

The Role Of Scavenging In Disease Dynamics, Joaquín Vicente, Kurt C. Vercauteren

USDA Wildlife Services: Staff Publications

Contents

Introduction................ 161

The Use of Animal Remains and the Exposure of Scavengers to Disease........ 163

The Relevance of Scavenging for Pathogens to Spread and Persist.......... 166

Human Related Factors Resulting in Increased Risk for Disease Transmission Through Scavenging.............. 170

Management of Scavenging to Reduce Disease Risks.............. 173

Restoration of Large Predators.................. 174

Elimination of Hunting of Scavengers............ 174

Destruction of Big Game and Domestic Animal Carcasses........... 174

Restoration of the Effects of Overabundance............. 175

Excluding Mammalian and Avian Scavengers from Natural Carrions.......... 176

Excluding Mammalian and Avian Scavengers from Vulture Restaurants........... 176

Conclusions and Future Perspectives........... 178

References............... 178

Home Range And Habitat Use Of West Virginia Canis Latrans (Coyote), Lauren M. Mastro, Dana J. Morin, Eric M. Gese

USDA Wildlife Services: Staff Publications

Canis latrans (Coyote) has undergone a range expansion in the United States over the last century. As a highly opportunistic species, its home range and habitat use changes with ecological context. Coyotes were first reported in West Virginia in 1950 but were not commonly observed until the 1990s, and there is scant information on Coyotes in the region. We used telemetry data from 8 radiocollared Coyotes in West Virginia to estimate home-range size and third-order habitat selection. Home-range areas (95% utilization distributions; UDs) varied from 5.22 to 27.79 km2 (mean = 12.48 ± 2.61 km2), with highly concentrated use of …

Carrion Availability In Space And Time, Marcos Moleón, Nuria Selva, David M. Bailey, David M. Bailey, Ainara Cortés-Avizanda, Travis L. Devault

USDA Wildlife Services: Staff Publications

Introduction

Availability of carrion to scavengers is a central issue in carrion ecology and management, and is crucial for understanding the evolution of scavenging behaviour. Compared to live animals, their carcasses are relatively unpredictable in space and time in natural conditions, with a few exceptions (see below, especially Sect. “Carrion Exchange at the Terrestrial-Aquatic Interface”). Carrion is also an ephemeral food resource due to the action of a plethora of consumers, from microorganisms to large vertebrates, as well as to desiccation (i.e., loss of water content; DeVault et al. 2003; Beasley et al. 2012; Barton et al. 2013; Moleón et …

Locally Fixed Alleles: A Method To Localize Gene Drive To Island Populations, Jaye Sudweeks, Brandon Hollingsworth, Dimitri V. Blondel, Karl J. Campbell, Sumit Dhole, John D. Eisemann, Owain Edwards, John Godwin, Gregg R. Howald, Kevin P. Oh, Antoinette J. Piaggio, Thomas A.A. Prowse, Joshua V. Ross, J. Royden Saah, Aaron B. Shiels, Paul Q. Thomas, David W. Threadgill, Michael R. Vella, Fred Gould, Alun L. Lloyd

USDA Wildlife Services: Staff Publications

Invasive species pose a major threat to biodiversity on islands. While successes have been achieved using traditional removal methods, such as toxicants aimed at rodents, these approaches have limitations and various off-target effects on island ecosystems. Gene drive technologies designed to eliminate a population provide an alternative approach, but the potential for drive-bearing individuals to escape from the target release area and impact populations elsewhere is a major concern. Here we propose the “Locally Fixed Alleles” approach as a novel means for localizing elimination by a drive to an island population that exhibits significant genetic isolation from neighboring populations. Our …

Using Unmanned Aerial Vehicles For Bird Harassment On Fish Ponds, Ciera A. Rhoades, Peter J. Allen, D. Tommy King

USDA Wildlife Services: Staff Publications

The effects of aquaculture decline on piscivorous birds in the Mississippi Delta concern catfish farmers, with possible increases in fish loss and disease transmission. Piscivorous birds quickly habituate to most current methods of harassment (loud noises and visual disturbances) leading to increased depredation and disease. Our study was designed to test the efficacy of using unmanned aerial vehicles (UAVs) to effectively control piscivorous birds at fish farms. We hypothesized that a UAV would be more efficient at reducing the number of fish-eating birds on fish ponds than current forms of harassment. We conducted pre-treatment bird surveys, harassment observations, and post-treatment …

Comparing Live-Capture Methods For Nutria: Single- Versus Multiple-Capture Cage Traps, Trevor R. Sheffels, Mark D. Sytsma, Jacoby Carter, Jimmy D. Taylor

USDA Wildlife Services: Staff Publications

Herbivory and burrowing by nutria (Myocastor coypus) cause substantial ecological and economic damage. Trapping is a common, effective practice for reducing nutria damage; however, trapping approaches must continually be adapted to keep pace with evolving animal welfare and ethical issues and to more effectively target pest species of interest. Our objective was to evaluate the efficacy of 2 nonlethal trap types for nutria: single-capture (SCT) and multi-capture (MCT) cage traps. We established 3 MCTs and 3 SCTs at each of 7 sites on a 10,500-ha mixed-use island located 15 km northwest of Portland, Oregon, USA. We pre-baited using carrots, apples, …

Ecological Functions Of Vertebrate Scavenging, James C. Beasley, Zachary H. Olson, Nuria Selva, Travis L. Devault

USDA Wildlife Services: Staff Publications

Introduction

The role of vertebrate scavenging in food web dynamics has historically been minimalized and portrayed as the activity of a select group of obligate scavengers (e.g., vultures), with a simplistic linkage between carrion and detritivores in food webs. Research in the last few decades, however, has revealed that the role of carrion in food web dynamics is severely underestimated, highly complex, and pervasive among ecosystems across the globe (DeVault et al. 2003; Selva and Fortuna 2007; Wilson and Wolkovich 2011). Such observations have led to a surge in research interest in scavenging ecology that continues to reveal new information …

In Situ Evaluation Of An Automated Aerial Bait Delivery System For Landscape-Scale Control Of Invasive Brown Treesnakes On Guam, Shane Siers, Will Pitt, John Eisemann, Larry Clark, Aaron B. Shiels, C. S. Clark, R. J. Gosnell, M. C. Messaros

USDA Wildlife Services: Staff Publications

After decades of biodiversity loss and economic burden caused by the brown treesnake invasion on the Pacific island of Guam, relief hovers on the horizon. Previous work by USDA Wildlife Services (WS) and its National Wildlife Research Center (NWRC) demonstrated that brown treesnake numbers in forested habitats can be dramatically suppressed by aerial delivery of dead newborn mouse (DNM) baits treated with 80 mg of acetaminophen. However, manual bait preparation and application is impractical for landscape-scale treatment. WS, NWRC, and the US Department of the Interior have collaborated with Applied Design Corporation to engineer an automated bait manufacturing and delivery …

Isolation Of Rabies Virus From The Salivary Glands Of Wild And Domestic Carnivores During A Skunk Rabies Epizootic, Cornell University Jimenez, Terry R. Spraker, Jessica Anderson, Richard Bowen, Amy Gilbert

USDA Wildlife Services: Staff Publications

Rabies is a fatal zoonotic disease of global importance. Rabies virus is shed in the saliva of infected hosts and is primarily transmitted through bite contact. Canine rabies has been eliminated from the US, but wildlife constitutes more than 90% of the reported cases of animal rabies in the US each year. In the US, several wild carnivore species are reservoirs of distinct variants of rabies virus (RV). After decades of apparent absence, the south-central skunk (SCSK) RV variant was detected in Colorado in 2007 and resulted in a large-scale epizootic in striped skunk (Mephitis mephitis) populations in northern Colorado …

Using Dna To Identify The Source Of Invasive Mongooses, Herpestes Auropunctatus (Carnivora: Herpestidae) Captured On Kaua‘I, Hawaiian Islands, Darren J. Wostenberg, Matthew W. Hopken, Aaron B. Shiels, Antoinette J. Piaggio

USDA Wildlife Services: Staff Publications

Two small Indian mongooses (Herpestes auropunctatus) were live-captured in 2012 at separate locations on the Hawaiian Island of Kaua'i, which was previously considered to be free of this invasive species. Genotypes from these two individuals were compared to genotypes of H. auropunctatus from the islands of Hawai'i (n =39), O'ahu (n =91), Maui (n = 39), and Moloka'i (n = 19) to determine the island of origin of the Kaua'i individuals. Genotypes were generated from each individual using five microsatellite loci. Genetic clustering was estimated by Bayesian inference of spatial clustering of individuals and …

Raccoon (Procyon Lotor) Response To Ontario Rabies Vaccine Baits (Onrab) In St. Lawrence County, New York, Usa, Kerri Pedersen, Amy T. Gilbert, Kathleen M. Nelson, Daniel P. Morgan, Amy J. Davis, Kurt C. Vercauteren, Dennis Slate, Richard B. Chipman

USDA Wildlife Services: Staff Publications

Oral rabies vaccination (ORV) campaigns have been conducted annually in the US over the past two decades to prevent raccoon (Procyon lotor) rabies, which is enzootic along the eastern region of the country from southeastern Canada to Alabama. Because raccoon rabies has been eliminated from neighboring Canadian provinces, continued detection of the variant in the US is of concern due to the potential for infected raccoons to cross the border via the St. Lawrence River. Ontario Rabies Vaccine Baits (ONRAB) containing a live, recombinant human adenovirus expressing the rabies virus glycoprotein have been under experimental use in the …

Trail Cameras Are A Key Monitoring Tool For Determining Target And Non-Target Bait-Take During Rodent Removal Operations: Evidence From Desecheo Island Rat Eradication, Aaron B. Shiels, D. Will, C. Figuerola-Hernández, K.J. Swinnerton, S. Silander, C. Samra, G.W. Witmer

USDA Wildlife Services: Staff Publications

Efforts to remove invasive rodents (e.g. Rattus spp. and Mus musculus) from islands often use toxicant-laced baits containing the anticoagulants brodifacoum or diphacinone. Rodenticide baits are generally delivered through aerialor hand-broadcast, or in bait stations. These baits are not rodent-specific and are subject to non-target consumption or secondary exposure (e.g. an individual preying upon another individual that has consumed bait). During rodenticide applications, it is generally unknown which animals are visiting and consuming bait; and to quantify this, we recommend using trail cameras (e.g. Reconyx^tm motion-activated infra-red) positioned to monitor individual bait pellets. To demonstrate the importance and effectiveness …

What Are The Transmission Mechanisms Of Influenza A Viruses In Wild Mammals?, J. Jeffrey Root

USDA Wildlife Services: Staff Publications

Some influenza A viruses (IAVs) represent serious potential threats to public and agricultural health, with 3 notable examples from the past decade. During 2009, a novel H1N1 IAV (A[H1N1]pdm09), which was first detected in the United States, spread rapidly throughout many regions of the world. In the United States alone, the Centers for Disease Control and Prevention estimated that >60 million human cases were associated with this emergent and pandemic virus. During 2013, a novel H7N9 avian-origin IAV (Asian lineage avian influenza A[H7N9] virus) was first detected in China. This virus not only cost the poultry industry more than $1 …

Mind The Gap: Experimental Tests To Improve Efficacy Of Fladry For Nonlethal Management Of Coyotes, Julie K. Young, John Draper, Stewart Breck

USDA Wildlife Services: Staff Publications

Coyotes (Canis latrans) are the top predator of livestock in the contiguous United States. Developing more effective nonlethal tools to prevent coyote depredation will facilitate coexistence between livestock producers and coyotes. Fladry is a nonlethal deterrent designed to defend livestock by creating a visual barrier to wolves (C. lupus). Fladry may also be effective with coyotes, but large gap spacing between flags may reduce its efficacy. To address this issue, we performed 2 experiments on captive coyotes using fladry modified to reduce gap spacing at the U.S. Department of Agriculture, Predator Research Facility in Millville, Utah, USA, during 2015–2016 and …

Experimental Infections Of Norway Rats With Avian‑Derived Low‑Pathogenic Influenza A Viruses, Kaci K. Vandalen, Nicole M. Nemeth, Nicholas O. Thomas, Nicole L. Barrett, Jeremy W. Ellis, Heather J. Sullivan, Alan B. Franklin, Susan A. Shriner

USDA Wildlife Services: Staff Publications

Influenza A viruses (IAVs) are a public-health, veterinary, and agricultural concern. Although wild birds are considered the primary reservoir hosts for most IAVs, wild-bird IAV strains are known to spill over into poultry, domestic or wild mammals, and humans. Occasionally, spillover events may result in adaptation or reassortment with other strains. Moreover, some IAV strains found in wild waterfowl mutate into highly pathogenic forms in poultry, causing tremendous economic losses. When domestic animals, wildlife, and humans dwell in close proximity to each other, such as may be the case with agricultural operations, wildlife may represent a potential risk for interspecies …

Taking The Bait: Species Taking Oral Rabies Vaccine Baits Intended For Raccoons, Betsy S. Haley, Are R. Berentsen, Richard M. Engeman

USDA Wildlife Services: Staff Publications

Raccoon rabies in eastern USA is managed by strategically distributing oral rabies vaccine (ORV) baits. The attractiveness, palativity, density, and non-target species bait take affect ORV effectiveness. We examined raccoon and non-target species differences in investigating/removing fish-meal polymer and coated sachet baits applied to simulate two aerial bait distribution densities. Bait densities of 150 baits/km² and 75 baits/km² were evaluated, respectively, in zones expected to have high and low raccoon densities. Three primary non-target species visited baits: coyotes, white-tailed deer, and feral swine. The proportion of bait stations visited by raccoons during 1 week observation periods ranged from …

Feral Swine Harming Insular Sea Turtle Reproduction: The Origin, Impacts, Behavior And Elimination Of An Invasive Species, Richard M. Engeman, Robert W. Byrd, Jamie Dozier, Mark A. Mcalister, James O. Edens, Elizabeth M. Kierepka, Timothy J. Smyser, Noel Myers

USDA Wildlife Services: Staff Publications

Feral swine are among the world's most destructive invasive species wherever they are found, with translocations figuring prominently in their range expansions. In contrast, sea turtles are beloved species that are listed as threatened or endangered throughout the world and are the focus of intense conservation efforts. Nest predation by feral swine severely harms sea turtle reproduction in many locations around the world. Here we quantify and economically assess feral swine nest predation at North Island, South Carolina, an important loggerhead sea turtle nesting beach. Feral swine depredation of North Island sea turtle nests was first detected in 2005, with …

Predation And Disease-Related Economic Impacts Of Wild Pigs On Livestock Producers In 13 States, Aaron M. Anderson, Chris Slootmaker, Erin Harper, Ryan S. Miller, Stephanie A. Shwiff

USDA Wildlife Services: Staff Publications

We report the results of a survey on wild pigs (Sus scrofa) damage to livestock producers in 13 US states (Alabama, Arkansas, California, Florida, Georgia, Louisiana, Mississippi, Missouri, North Carolina, Oklahoma, South Carolina, Tennessee, Texas). The survey was distributed by the USDA National Agricultural Statistical Service in the summer of 2017 to a sample of livestock producers in the 13-state region. Findings indicate that predation and disease-related damage can be substantial in certain states and for certain types of livestock. In particular, damage to cattle operations in Texas and Arkansas was substantially higher than damage in other states …

Persistence And Conspecific Observations Improve Problem-Solving Abilities Of Coyotes, Julie K. Young, Laura Touzot, Stacey P. Brummer

USDA Wildlife Services: Staff Publications

Social learning has important ecological and evolutionary consequences but the role of certain factors, such as social rank, neophobia (i.e., avoidance of novel stimuli), persistence, and task-reward association, remain less understood. We examined the role of these factors in social learning by captive coyotes (Canis latrans) via three studies. Study 1 involved individual animals and eliminated object neophobia by familiarizing the subjects to the testing apparatus prior to testing. Studies 2 and 3 used mated pairs to assess social rank, and included object neophobia, but differed in that study 3 decoupled the food reward from the testing apparatus (i.e., altered …