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Optimization Of Scat Detection Methods For A Social Ungulate, The Wild Pig, And Experimental Evaluation Of Factors Affecting Detection Of Scat, David A. Keiter, Fred L. Cunningham, Olin E. Rhodes Jr., Brian J. Irwin, James C. Beasley

United States Department of Agriculture Wildlife Services: Staff Publications

Collection of scat samples is common in wildlife research, particularly for genetic capturemark- recapture applications. Due to high degradation rates of genetic material in scat, large numbers of samples must be collected to generate robust estimates. Optimization of sampling approaches to account for taxa-specific patterns of scat deposition is, therefore, necessary to ensure sufficient sample collection. While scat collection methods have been widely studied in carnivores, research to maximize scat collection and noninvasive sampling efficiency for social ungulates is lacking. Further, environmental factors or scat morphology may influence detection of scat by observers. We contrasted performance of novel radial search …

Effects Of Simulated Removal Activities On Movements And Space Use Of Feral Swine, Justin W. Fischer, Dan Mcmurty, Chad R. Blass, W. David Walter, Jeff Beringer, Kurt C. Vercauteren

United States Department of Agriculture Wildlife Services: Staff Publications

Abundance and distribution of feral swine (Sus scrofa) in the USA have increased dramatically during the last 30 years. Effective measures are needed to control and eradicate feral swine populations without displacing animals over wider areas. Our objective was to investigate effects of repeated simulated removal activities on feral swine movements and space use.We analyzed location data from 21 feral swine that we fitted with Global Positioning System harnesses in southern MO, USA. Various removal activities were applied over time to eight feral swine before lethal removal, including trapped-and-released, chased with dogs, chased with hunter, and chased with …

Widespread Detection Of Highly Pathogenic H5 Influenza Viruses In Wild Birds From The Pacific Flyway Of The United States, S. N. Bevins, R. J. Dusek, C. L. White, T. Gidlewski, B. Bodenstein, K. G, Mansfield, P. Debruyn, D. Kraege, E. Rowan, C. Gillin, B. Thomas, S. Chandler, J. A. Baroch, B. S. Schmit, M. J. Grady, R. S. Miller, M. L. Drew, S. Stopak, B. Zscheile, J. Bennett, J. Sengl, Caroline Brady, H. S. Ip, E. Spackman, M. L. Killian, M. K. Torchetti, J. M. Sleeman, T. J. Deliberto

United States Department of Agriculture Wildlife Services: Staff Publications

A novel highly pathogenic avian influenza virus belonging to the H5 clade 2.3.4.4 variant viruses was detected in North America in late 2014. Motivated by the identification of these viruses in domestic poultry in Canada, an intensive study was initiated to conduct highly pathogenic avian influenza surveillance in wild birds in the Pacific Flyway of the United States. A total of 4,729 hunter-harvested wild birds were sampled and highly pathogenic avian influenza virus was detected in 1.3% (n = 63). Three H5 clade 2.3.4.4 subtypes were isolated from wild birds, H5N2, H5N8, and H5N1, representing the wholly Eurasian lineage H5N8 …

Vertebrate Host Susceptibility To Heartland Virus, Angela M. Bosco-Lauth, Amanda E. Calvert, J. Jeffrey Root, Tom Gidlewski, Brian H. Bird, Richard A. Bowen, Atis Muehlenbachs, Sherif R. Zaki, Aaron C. Brault

United States Department of Agriculture Wildlife Services: Staff Publications

Heartland virus (HRTV) is a recently described phlebovirus initially isolated in 2009 from 2 humans who had leukopenia and thrombocytopenia. Serologic assessment of domestic and wild animal populations near the residence of 1 of these persons showed high exposure rates to raccoons, white-tailed deer, and horses. To our knowledge, no laboratory-based assessments of viremic potential of animals infected with HRTV have been performed. We experimentally inoculated several vertebrates (raccoons, goats, chickens, rabbits, hamsters, C57BL/6 mice, and interferon-α/β/γ receptor–deficient [Ag129]) mice with this virus. All animals showed immune responses against HRTV after primary or secondary exposure. However, neutralizing antibody responses were …

Tracking A Deadly Virus Highly Pathogenic Avian Influenza In Wild Birds, Tom Deliberto, Gail Keirn

United States Department of Agriculture Wildlife Services: Staff Publications

For Dennis Kohler, the call was somewhat unexpected. On a cold, windy morning in December 2014, Kohler sat in his Colorado office reviewing research plans for upcoming disease studies. A few hours later, Koehler's colleagues at the National Wildlife Health Center in Madison, Wis., called to notify him and other members of the U.S. Department of Agriculture's National Wildlife Disease Program (NWDP) that recent samples collected from wild birds in Washington State had tested positive for highly pathogenic avian influenza (HPAI).

Earlier that month, a die-off of mallards (Anas platyrhynchos), American wigeon (A. Americana) and northern …

Pathogen Emergence In The Age Of Pandemics, Sarah N. Bevins

United States Department of Agriculture Wildlife Services: Staff Publications

The emergence of novel or previously rare pathogens is not new. Throughout recorded history, catastrophic mortality has followed the arrival of new infectious diseases to areas that had no previous experience with the offending agent. The second plague pandemic, more commonly known as the Black Death, was caused by a bacterium that originated in China but then spread throughout Asia, Europe, and Africa via the Silk Road and other routes of commerce. The emergence of rinderpest in Africa, associated with Asian cattle imported by European colonists, killed millions of domestic livestock and wild animal species in the 1890s. The result …

Is There A Future For Genome-Editing Technologies In Conservation?, J. A. Johnson, R. Altwegg, D. M. Evans, J. G. Ewen, I. J. Gordon, N. Pettorelli, J. K. Young

United States Department of Agriculture Wildlife Services: Staff Publications

In a recent review, Pimm et al. (2015) highlight emerging technologies in protecting biodiversity. While their list is noteworthy, the authors’ exclusion of innovations in genomic research, with the exception of single-species DNA barcoding methods, was surprising given recent advances in genome-editing technology and its potential application to conservation. Taylor & Gemmell (2016) address that deficiency in a subsequent commentary identifying three avenues where emerging genomic technologies have great potential for increasing our ability to conserve biodiversity. Those areas include the use of next-generation sequencing technologies and methods such as RADseq for monitoring genetic diversity, effective population size, and …

Composition And Diversity Of Avian Communities Using A New Urban Habitat: Green Roofs, Brian E. Washburn, Ryan M. Swearingin, Craig K. Pullins, Matthew E. Rice

United States Department of Agriculture Wildlife Services: Staff Publications

Green roofs on buildings are becoming popular and represent a new component of the urban landscape. Public benefits of green roof projects include reduced stormwater runoff, improved air quality, reduced urban heat island effects, and aesthetic values. As part of a city-wide plan, several green roofs have been constructed at Chicago’s O’Hare International Airport (ORD). Like some other landscaping features, green roofs on or near an airport might attract wildlife and thus increase the risk of bird–aircraft collisions. During 2007–2011, we conducted a series of studies to evaluate wildlife use of newly constructed green roofs and traditional (gravel) roofs on …

Integrating Early Detection With Dna Barcoding: Species Identification Of A Nonnative Monitor Lizard (Squamata: Varanidae) Carcass In Mississippi, U.S.A., Robert N. Reed, Matthew W. Hopken, David Steensen, Bryan G. Falk, Antoinette J. Piaggio

United States Department of Agriculture Wildlife Services: Staff Publications

Early detection of invasive species is critical to increasing the probability of successful management. At the primary stage of an invasion, invasive species are easier to control as the population is likely represented by just a few individuals. Detection of these first few individuals can be challenging, particularly if they are cryptic or otherwise characterized by low detectability. The engagement of members of the public may be critical to early detection as there are far more citizens on the landscape than trained biologists. However, it can be difficult to assess the credibility of public reporting, especially when a diagnostic digital …

No Filters, No Fridges: A Method For Preservation Of Water Samples For Edna Analysis, Kelly E. Williams, Kathryn P. Huyvaert, Antoinette J. Piaggio

United States Department of Agriculture Wildlife Services: Staff Publications

Background: Advancements in the detection of environmental DNA (eDNA) for detecting species of interest will likely allow for expanded use of these techniques in the field. One obstacle that continues to hinder applications in the field is the requirement of a cold chain of storage for water samples containing eDNA. While eDNA has been successfully preserved using Longmire’s lysis buffer applied to filters, it has yet to be tried with freshwater samples collected for eDNA detection of an invasive species. We tested the utility of Longmire’s solution (100 mM Tris, 100 mM EDTA, 10 mM NaCl, 0.5 % SDS, 0.2 …

Review Of Anthraquinone Applications For Pest Management And Agricultural Crop Protection, Shelagh T. Deliberto, Scott J. Werner

United States Department of Agriculture Wildlife Services: Staff Publications

We have reviewed published anthraquinone applications for international pest management and agricultural crop protection from 1943 to 2016. Anthraquinone (AQ) is commonly found in dyes, pigments and many plants and organisms. Avian repellent research with AQ began in the 1940s. In the context of pest management, AQ is currently used as a chemical repellent, perch deterrent, insecticide and feeding deterrent in many wild birds, and in some mammals, insects and fishes. Criteria for evaluation of effective chemical repellents include efficacy, potential for wildlife hazards, phytotoxicity and environmental persistence. As a biopesticide, AQ often meets these criteria of efficacy for the …

Effects Of Renewable Energy Production And Infrastructure On Wildlife, José Antonio Sánchez-Zapata, Miguel Clavero, Martina Carrete, Travis L. Devault, Virgilio Hermoso, Miguel Angel Losada, María José Polo, Sonia Sánchez-Navarro, Juan Manuel Pérez-García, Francisco Botella, Carlos Ibáñez, José Antonio Donázar

United States Department of Agriculture Wildlife Services: Staff Publications

The high levels of human demands of resources—from food to space and energy—are one of the main drivers of global change and are causing large negative impacts on ecosystems functioning worldwide (Vitousek et al. 1997 ). Global change components range from climate change to habitat destruction, species invasions, pollution and eutrophication. Although these factors can produce ecosystem changes independently, the final descent is often driven by synergistic processes. The resulting amplifying feedbacks can be disconnected from the original driver of change, leading to a state shift in the biosphere with unexpected consequences (Barnosky et al. 2012 ). Climate change seems …

Novel And Current Rodenticides For Pocket Gopher Thomomys Spp. Management In Vineyards: What Works?, Roger A. Baldwin, Ryan Meinerz, Gary W. Witmer

United States Department of Agriculture Wildlife Services: Staff Publications

BACKGROUND: Rodenticides are often included as part of an integrated pest management approach for managing pocket gophers (Thomomys spp.) given that they are relatively quick and inexpensive to apply. Strychnine has historically been the most effective toxicant for pocket gophers, but its use is currently limited in the United States; alternative registered toxicants have not proven effective. Recent research with baits containing cholecalciferol plus anticoagulant toxicants proved effective against pocket gophers in a lab setting. Therefore, we established a field study to compare cholecalciferol plus anticoagulant combinations [0.03% cholecalciferol plus 0.005% diphacinone (C+D), 0.015% cholecalciferol plus 0.0025% brodifacoum (C+B1), …

A Quantitative Rt-Pcr Assay For Rapid Detection Of Eurasianlineage H10 Subtype Influenza A Virus, Hailiang Sun, Jian-Li Xue, Elizabeth Bailey, Yifei Xu, Guoliang Hu, John A. Baroch, Yi Zhang, Lanny Pace, Thomas J. Deliberto, Xiu-Feng Wan

United States Department of Agriculture Wildlife Services: Staff Publications

Influenza A viruses (IAVs) are single-stranded, negative sense RNA viruses. IAV subtype is determined on the basis of the viral surface glycoproteins, hemagglutinin (HA), and neuraminidase (NA). To date, 18 HA and 11 NA subtypes have been reported (Tong et al., 2012). IAVs can cause sporadic infections, local epidemics, and global pandemics among humans. In addition to humans, IAVs can naturally infect avian, swine, equines, canines, and sea mammals (Webster et al., 1992). Migratory waterfowl are the natural reservoir for IAVs, and the avianorigin IAVs play an important role in influenza ecology and have been involved in generation of the …

Cougar Predation Rates And Prey Composition In The Pryor Mountains Of Wyoming And Montana, Linsey W. Blake, Eric M. Gese

United States Department of Agriculture Wildlife Services: Staff Publications

Understanding predator-prey dynamics is a fundamental principle of ecology and an ideal component for management decisions. Across North America, the impact of cougars (Puma concolor) on their prey varies regionally. To document the relationships between cougars, bighorn sheep (Ovis canadensis canadensis), mule deer (Odocoileus hemionus), and feral horses (Equus caballus) on the Bighorn Canyon National Recreation Area and the Pryor Mountains, we deployed GPS collars on 6 cougars (the total number residing on the study area), and visited their clusters to determine predation rates and foraging patterns. We examined the composition of …

Inferring Invasive Species Abundance Using Removal Data From Management Actions, Amy J. Davis, Mevin B. Hooten, Ryan S. Miller, Matthew L. Farnsworth, Jesse Lewis, Michael Moxcey, Kim M. Pepin

United States Department of Agriculture Wildlife Services: Staff Publications

Evaluation of the progress of management programs for invasive species is crucial for demonstrating impacts to stakeholders and strategic planning of resource allocation. Estimates of abundance before and after management activities can serve as a useful metric of population management programs. However, many methods of estimating population size are too labor intensive and costly to implement, posing restrictive levels of burden on operational programs. Removal models are a reliable method for estimating abundance before and after management using data from the removal activities exclusively, thus requiring no work in addition to management. We developed a Bayesian hierarchical model to estimate …

Plant Community Shifts Caused By Feral Swine Rooting Devalue Florida Rangeland, Brittany Bankovich, Elizabeth Boughton, Raoul Boughton, Michael L. Avery, Samantha M. Wisely

United States Department of Agriculture Wildlife Services: Staff Publications

Invasive species threaten agriculture by changing agroecosystem structure and function, reducing habitat value, decreasing biodiversity and ecosystem services and increasing management costs. Grazing lands in south central Florida are a mosaic of sown pastures, native grasslands, wetlands and woodlands that provide a variety of ecosystem services. Disturbance of these pastures and native grasslands by invasive feral swine (Sus scrofa) can have negative consequences for both economic productivity and biodiversity. In this study, we show that the effect of rooting on plant diversity depends on ecosystem type and initial levels of plant species diversity. For example, in native grassland pasture, rooting …

Chapter 8 Keeping Wildlife Out Of Your Food: Mitigation And Control Strategies To Reduce The Transmission Risk Of Food-Borne Pathogens, Alan B. Franklin, Kurt C. Vercauteren

United States Department of Agriculture Wildlife Services: Staff Publications

In this chapter, we provide a general framework for developing strategies to mitigate the contamination of agricultural operations with pathogens carried by wildlife. As part of this framework, we present adaptive management as a viable approach to developing these strategies to reduce the uncertainty over time as to whether management methods are being effective. We provide the general steps to developing an adaptive management strategies as well as generic mitigation methods that can be applied to agricultural operations as part of an adaptive management strategy.

Nutritional Effects On Reproductive Performance Of Captiveadult Female Coyotes (Canis Latrans), Eric M. Gese, Beth M. Roberts, Frederick F. Knowlton

United States Department of Agriculture Wildlife Services: Staff Publications

Interactions between animals and their environment are fundamental to ecological research. Field studies of coyote (Canis latrans) reproductive performance suggest mean litter size changes in response to prey abundance. However, this relationship has been assessed primarily by using carcasses collected from trappers. The objective of this study was to assess whether nutritional manipulation prior to mating affected reproduction in adult female coyotes. We examined the effects of caloric restriction during the 7 months prior to estrus on the reproductive rates of 11 captive female coyotes and the subsequent initial survival of pups through two reproductive cycles. This was …

Molecular Forensics In Avian Conservation: A Dna‑Based Approach For Identifying Mammalian Predators Of Ground‑Nesting Birds And Eggs, Matthew W. Hopken, Elizabeth K. Orning, Julie K. Young, Antoinette J. Piaggio

United States Department of Agriculture Wildlife Services: Staff Publications

Background: The greater sage-grouse (Centrocercus urophasianus) is a ground-nesting bird from the Northern Rocky Mountains and a species at risk of extinction in in multiple U.S. states and Canada. Herein we report results from a proof of concept that mitochondrial and nuclear DNAs from mammalian predator saliva could be non-invasively collected from depredated greater sage-grouse eggshells and carcasses and used for predator species identification. Molecular forensic approaches have been applied to identify predators from depredated remains as one strategy to better understand predator–prey dynamics and guide management strategies. This can aid conservation efforts by correctly identifying predators most …

Estimates Of Small Indian Mongoose Densities: Implications For Rabies Management, Shylo R. Johnson, Are R. Berentsen, Christine Ellis, Amy Davis, Kurt C. Vercauteren

United States Department of Agriculture Wildlife Services: Staff Publications

The small Indian mongoose (Herpestes auropunctatus) is an invasive species and rabies reservoir in Puerto Rico. In the continental United States, terrestrial wildlife rabies is primarily managed by the National Rabies Management Program (NRMP) of the United States Department of Agriculture through oral rabies vaccination (ORV); the distribution of the vaccine baits is influenced by the population density of the target species. TheNRMPuses a density index for estimating raccoon (Procyon lotor) population density to guide bait distribution. In Puerto Rico, a wildlife rabies vaccination program does not exist and vaccination of domestic animals is limited and not compulsory. …

Bait Preference Of Free-Ranging Feral Swine For Delivery Of A Novel Toxicant, Nathan P. Snow, Joseph M. Halseth, Michael J. Lavelle, Thomas E. Hanson, Chad R. Blass, Justin A. Foster, Simon T. Humphrys, Linton D. Staples, David G. Hewitt, Kurt C. Vercauteren

United States Department of Agriculture Wildlife Services: Staff Publications

Invasive feral swine (Sus scrofa) cause extensive damage to agricultural and wildlife resources throughout the United States. Development of sodium nitrite as a new, orally delivered toxicant is underway to provide an additional tool to curtail growth and expansion of feral swine populations. A micro-encapsulation coating around sodium nitrite is used to minimize detection by feral swine and maximize stability for the reactive molecule. To maximize uptake of this toxicant by feral swine, development a bait matrix is needed to 1) protect the micro-encapsulation coating so that sodium nitrite remains undetectable to feral swine, 2) achieve a high …

Evaluation Of Harvest Indices For Monitoring Cougar Survival And Abundance, Michael L. Wolfe, Eric M. Gese, Pat Terletzky, David C. Stoner, Lise M. Aubry

United States Department of Agriculture Wildlife Services: Staff Publications

Harvest indices are used by state wildlife management agencies to monitor population trends and set harvest quotas for furbearer species. Although harvest indices may be readily collected from hunters, the reliability of harvest indices for monitoring demography and abundance of the harvested species is rarely examined, particularly amongst large carnivores. The overall objective of this study was to assess whether cougar (Puma concolor) harvest statistics collected by wildlife managers were correlated with changes in cougar demography, mainly survival rates and abundance. We estimated key demographic parameters for 2 cougar populations in Utah over 17 years during which we monitored 235 …

Cholecalciferol Plus Diphacinone Baits For Vole Control: A Novel Approach To A Historic Problem, Roger A. Baldwin, Ryan Meinerz, Gary W. Witmer

United States Department of Agriculture Wildlife Services: Staff Publications

Combination baits containing cholecalciferol plus an anticoagulant are effective against commensal rodents resistant to anticoagulants, and they likely pose less risk than anticoagulant-only rodenticides due to lower concentrations of active ingredients and shorter time to death. However, these combination baits have not been tested for agricultural rodent pests. Therefore, we established a study to test the efficacy of cholecalciferol plus diphacinone artichoke bract and pellet baits to determine their ability to manage California voles Microtus californicus in artichokes, where resistance to anticoagulants is known to occur. Field tests using radiocollared voles indicated that bract baits were highly efficacious (85 %), …

Defending Against Disparate Marine Turtle Nest Predators: Nesting Success Benefits From Eradicating Invasive Feral Swine And Caging Nests From Raccoons, Richard M. Engeman, David Addison, J. C. Griffin

United States Department of Agriculture Wildlife Services: Staff Publications

Nest predation can threaten marine turtle nesting success, and having to address dissimilar predator species complicates nest protection efforts. On Florida’s Keewaydin Island predation by raccoons Procyon lotor and invasive feral swine Sus scrofa are disparate, significant threats tomarine turtle nests. Using 6 years of nesting data (mostly for loggerhead marine turtles Caretta caretta) we examined the impacts of swine predation on nests and the benefits of swine eradication, caging nests to protect them from raccoon predation, and the effects of nest caging on swine predation. Nest predation by swine began in mid nesting season 2007, after which swine …

Evaluation Of An Avian Radar System In A Midwestern Landscape, Michael B. Gerringer, Steven L. Lima, Travis L. Devault

United States Department of Agriculture Wildlife Services: Staff Publications

Bird strikes in aviation are an increasing threat to both aircraft and human safety. Management efforts have focused largely on the immediate airport environment. Avian radar systems could potentially be useful in assessing bird strike threats at greater distances from the airport, at higher altitudes, and at night, but few studies have been conducted to assess the capabilities of avian radar systems. Thus, our goal was to assess the detection and tracking abilities of a commercially available avian radar system in an airport environment in Indiana, USA, during October 2011–March 2012. Transits by free-flying birds allowed us to assess radar …

Development Of Artificial Bait For Brown Treesnake Suppression, Bruce A. Kimball, Scott A. Sterling, Thomas W. Mcauliffe, Randal S. Stahl, Rafael A. Garcia, William C. Pitt

United States Department of Agriculture Wildlife Services: Staff Publications

The brown treesnake (Boiga irregularis) was accidentally introduced to Guam in the 1940s from the Admiralty Islands. A native of Australia, Papua New Guinea, and the Solomon Islands, the brown treesnake (BTS) continues to threaten the economy and ecology of Guam and is currently the subject of a cooperative program to control snake populations on the island and prevent its spread throughout the Pacific Rim. Delivery of toxic baits is a primary component of population suppression efforts. While many food items tested as baits for toxicant delivery provide relevant food prey cues leading to investigatory behaviors in BTS, …

Retention Time Of Chlorophacinone In Black-Tailed Prairie Dogs Informs Secondary Hazards From A Prairie Dog Rodenticide Bait, Gary W. Witmer, Nathan P. Snow, Rachael S. Moulton

United States Department of Agriculture Wildlife Services: Staff Publications

BACKGROUND: Secondary toxicity in mammals and birds that consume animals containing residues of anticoagulant rodenticides represents a persistent conflict between conservation, agriculture and environmental contamination. Chlorophacinone residues in black-tailed prairie dogs (Cynomys ludovicianus) represent a secondary exposure hazard to predatory and scavenging avian and mammalian species in the Central Plains of the United States, especially considering efforts to re-establish black-footed ferrets (Mustela nigripes). Rozol® Prairie Dog Bait (chlorophacinone 0.005%) is registered to control black-tailed prairie dogs in ten states throughout themidwestern and western United States.

RESULTS:We fed Rozol Prairie Dog Bait to captive black-tailed prairie dogs …

Susceptibility Of Rock Doves To Low-Pathogenic Avian Influenza A Viruses, Susan A. Shriner, J. Jeffrey Root, Nicole L. Mooers, Jeremy W. Ellis, Scott R. Stopak, Heather J. Sullivan, Kaci K. Vandalen, Alan B. Franklin

United States Department of Agriculture Wildlife Services: Staff Publications

Following a 2008 outbreak of North American low-pathogenic H5N8 influenza A virus at an upland gamebird farm, we sero-sampled rock doves (pigeons, Columba livia) at the outbreak site and conducted experimental inoculations of wild-caught pigeons using the H5N8 virus and another low-pathogenic virus (H4N6). While 13 % of pigeons at the outbreak site were seropositive, none were positive for exposure to H5, and one was positive for N8. Challenged pigeons exhibited low susceptibility and limited viral RNA excretion for both viruses tested, but at least one individual had RNA loads indicative of the potential for viral transmission to other …

Supplementary Feeding And Endangered Avian Scavengers: Benefits, Caveats, And Controversies, Ainara Cortés- Avizanda, Guillermo Blanco, Travis L. Devault, Anil Markandya, Munir Z Virani, Joseph Brandt, José A Donázar

United States Department of Agriculture Wildlife Services: Staff Publications

Large avian scavengers are among the most vulnerable vertebrates, and many of their populations have declined severely in recent decades. To help mitigate this marked reduction in abundance, supplementary feeding stations (SFS; colloquially termed “vulture restaurants”) have been created worldwide, often without consideration of the scientific evidence supporting the suitability of the practice. SFS have been effective and important tools for conservation and reintroduction of avian scavengers. However, negative consequences can result from large aggregations of individual birds, disrupting intraguild processes and promoting density-dependent decreases in productivity. At the community level, SFS favor the congregation of predators (ie facultative scavengers), …