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Describing Willow Flycatcher habitats: scale perspectives and gender differences

We compared habitat characteristics of nest sites (female-selected sites) and song perch sites (male-selected sites) with those of sites unused by Willow Flycatchers ( Empidonax traillii ) at three different scales of vegetation measurement: (1) microplot (central willow [ Salix spp.] bush and four adjacent bushes); (2) mesoplot (0.07 ha); and, (3) macroplot (flycatcher territory size). Willow Flycatchers exhibited vegetation preferences at all three scales. Nest sites were distinguished by high willow density and low variability in willow patch size and bush height. Song perch sites were characterized by large central shrubs, low central shrub vigor, and high variability in shrub size. Unused sites were characterized by greater distances between willows and willow patches, less willow coverage, and a smaller riparian zone width than either nest or song perch sites. At all scales, nest sites were situated farther from unused sites in multivariate habitat space than were song perch sites, suggesting (1) a correspondence among scales in their ability to describe Willow Flycatcher habitat, and (2) females are more discriminating in habitat selection than males. Microhabitat differences between male-selected (song perch) and female-selected (nest) sites were evident at the two smaller scales; at the finest scale, the segregation in habitat space between male-selected and female-selected sites was greater than that between male-selected and unused sites. Differences between song perch and nest sites were not apparent at the scale of flycatcher territory size, possibly due to inclusion of (1) both nest and song perch sites, (2) defended, but unused habitat, and/or (3) habitat outside of the territory, in larger scale analyses. The differences between nest and song perch sites at the finer scales reflect their different functions (e.g., nest concealment and microclimatic requirements vs. advertising and territorial defense, respectively), and suggest that the exclusive use of either nest or song perch sites in vegetation analyses can result in misleading, or at least incomplete, descriptions of a species' habitat. Habitat interpretations for Willow Flycatchers (and perhaps for many passerines) are a function of the gender-specific behavior of the birds observed and the scale of vegetation measurement.

The Condor

Does nonrandom nest placement imply nonrandom nest predation?: A reply

In response to the critique by Schmidt and Whelan (Condor 101(4):916-920, 1999), we find that the relationship between nest success and tree selectivity is dependent upon inclusion or exclusion of particular tree species, whether or not years are pooled, and the selectivity index used. We question their use of point estimates of nest success with extremely high variances, defend our index, question the application of the Chesson (1983) index to our data, and explain the need to analyze years separately. Bottomland hardwood forest systems are extremely variable; hydroperiods alter the suitability of nesting substrates, availability of alternative food, and behavior of predators and their prey. Given these features, actively searching for Acadian Flycatcher (Empidonax virescens) nests is seldom an efficient predator foraging strategy. Therefore, these predation events are best described as random; nests are principally encountered opportunistically by generalist predators while searching for other prey.

Condor

Behavioral interactions of penned red and arctic foxes

Expansion of the geographical distribution of red foxes ( Vulpes vulpes ) into the far north tundra region may lead to competition between arctic ( Alopex lagopus ) and red foxes for space and resources. Behavioral interactions between red and arctic foxes were evaluated during 9 trials conducted in a 4.05-ha enclosure near Woodworth, North Dakota. Each trial consisted of introducing a male-female pair of arctic foxes into the enclosure and allowing them to acclimate for approximately a week before releasing a female red fox into the enclosure, followed by her mate a few days later. In 8 of 9 trials, red foxes were dominant over arctic foxes during encounters. Activity of the arctic foxes decreased upon addition of red foxes. Arctic foxes tried unsuccessfully to defend preferred den, resting, and feeding areas. Even though the outcome of competition between red and arctic foxes in the Arctic is uncertain, the more aggressive red fox can dominate arctic foxes in direct competition for den sites and other limited resources.

North Dakota

Social organization in a nesting population of eastern willets ( Catoptrophorus semipalmatus )

The breeding ecology of eastern Willets ( Catoptrophorus semipalmatus ) was studied over a 3-yr period in a salt marsh on the Atlantic coast of Virginia. During the study, 171 adults were color-marked. Willets are strictly monogamous and show strong site and mate fidelity from year to year. Normally a pair bond dissolves only after one member dies or fails to return in the spring. With an estimated annual adult mortality of 15%, the average life of a pair bond is 3 yr. The sexes share in most aspects of breeding, including brood-rearing, but females abandon both mate and brood about 2 weeks after the chicks hatch. Males apparently incubate at night and sometimes relieve the female at the nest during the middle of the day. In this area Willets defend separate nesting and feeding territories, though nest-site defense decreases through the incubation period. When mates are lost, males retain their original nesting and feeding territories, while females pair with available mates near their original nesting territory. Females compete for mates, and males select the most aggressive females. Nesting was particularly synchronous in lowland marsh sites, where the onset of egg laying was slightly earlier than in upland sites. Nest density was the highest reported for any scolopacid, and the highest densities were in sites closest to the marsh. Nest spacing is achieved through agonistic behavior, but there is evidence of a counter-tendency leading to nest clumping. Several features of the Willet social system resemble characteristics typical of colonial species of birds. These are discussed along with other aspects of social organization and the selective forces operating on it.

The Auk

Precocious breeding by yearling Giant Canada Geese

Many species of waterfowl are capable of breeding as yearlings. In the subfamily Anserinae, however, reproduction normally does not commence until individuals reach two to four years of age (Rohwer 1992). Most published accounts indicate that Canada Geese ( Branta canadensis ) conform to the pattern typically found in other geese, deferring reproduction as yearlings and initiating breeding when birds are two years of age or older (Hanson 1962, Brakhage 1965, Bellrose 1980, Moser and Rusch 1989). The few documented exceptions (Brakhage 1965, Cooper 1978, MacInnes and Dunn 1988) suggest that the likelihood of a successful breeding attempt is higher for yearling males than females and that when early breeding occurs, one member of the pair is often at least two years old. Only two records of nesting attempts by yearling female Canada Geese have been published (Hall and McGilvrey 1971, Mickelson, 1975). In each case, the female produced some fertile eggs but deserted the nest before the eggs hatched. The observations noted above demonstrate that reproduction by yearling Canada Geese is physiolog ically possible for both sexes, but they also raise a number of interesting questions regarding why most individuals defer breeding during their first year. Energetic or nutritional constraints, and the inability of yearlings to secure and successfully defend a territory, are likely physiological (Elder 1946) and social (Brakhage 1965) impediments to early breeding From an evolutionary perspective, reduced reproductive success of yearlings relative to adults, and increased mortality associated with early breeding. also may contribute to deferred sexual maturity in waterfowl (Rohwer 1992).

The Auk

The influence of food abundance, food dispersion and habitat structure on territory selection and size of an Afrotropical terrestrial insectivore

Most tropical insectivorous birds, unlike their temperate counterparts, hold and defend a feeding and breeding territory year-around. However, our understanding of ecological factors influencing territory selection and size in tropical insectivores is limited. Here we examine three prominent hypotheses relating food abundance, food dispersion (spatial arrangement of food items), and habitat structure to territoriality in the Usambara Thrush Turdus roehli . We first compared leaf-litter macro-invertebrate abundance and dispersion, and habitat structure between territories and random sites. We then examined the relation between these same ecological factors and territory size. Invertebrate abundance and dispersion were sparsely and evenly distributed across our study system and did not vary between territories and random sites. In contrast, habitat structure did vary between territories and random sites indicating the Usambara Thrush selects territories with open understorey and closed overstorey habitat. Invertebrate abundance and dispersion within territories of the Usambara Thrush were not associated with habitat structure. We believe the most likely explanation for the Usambara Thrush’s preference for open understorey and closed overstorey habitat relates to foraging behavior. Using information-theoretic model selection we found that invertebrate abundance was the highest-ranked predictor of territory size and was inversely related, consistent with food value theory of territoriality.

Ostrich

Processes influencing the transport and fate of contaminated sediments in the coastal ocean– Boston Harbor and Massachusetts Bay

Most of the major urban centers of the United States including Boston, New York, Washington, Chicago, New Orleans, Miami, Los Angeles, San Francisco, and Seattle—are on a coast (fig. 1.1). All of these cities discharge treated sewage effluent into adjacent waters. In 2000, 74 percent of the U.S. population lived within 200 kilometers (km) of the coast. Between 1980 and 2002, the population density in coastal communities increased approximately 4.5 times faster than in noncoastal areas of the U.S. (Perkins, 2004). More people generate larger volumes of wastes, increase the demands on wastewater treatment, expand the area of impervious land surfaces, and use more vehicles that contribute contaminants to street runoff. According to the National Coastal Condition Report II (U.S. Environmental Protection Agency, 2005a), on the basis of coastal habitat, water and sediment quality, benthic index, and fish tissue, the overall national coastal condition is only poor to fair and the overall coastal condition in the highly populated Northeast is poor. Scientific information helps managers to prioritize and regulate coastal-ocean uses that include recreation, commercial fishing, transportation, waste disposal, and critical habitat for marine organisms. These uses are often in conflict with each other and with environmental concerns. Developing a strategy for managing competing uses while maintaining sustainability of coastal resources requires scientific understanding of how the coastal ocean system behaves and how it responds to anthropogenic influences. This report provides a summary of a multidisciplinary research program designed to improve our understanding of the transport and fate of contaminants in Massachusetts coastal waters. Massachusetts Bay and Boston Harbor have been a focus of U.S. Geological Survey (USGS) research because they provide a diverse geographic setting for developing a scientific understanding of the geology, geochemistry, and oceanography of coastal systems in general. Scientific data from this region can also be used to inform decisions about important economic, environmental, and political issues. From the economic viewpoint, the annual value of tourism and shipping in Massachusetts and Cape Cod Bays is about $1.5 billion and $1.9 billion, respectively. Commercial and recreational fishing generates about $240 million per year in the same region (U.S. Environmental Protection Agency, 2005b). The environmental issue is the 300-year history of waste discharge from the Boston metropolitan area into the harbor. This history is punctuated by cycles of environmental degradation, public outcry, and improvements in the sewage treatment system. With each improvement, however, the continuous growth of population in greater Boston (fig. 1.2) and the resulting increase in the volume of waste exceeded the capacity of the treatment system, thereby setting the stage for a new contamination crisis. By the 1980s, the levels of contaminants in sediments of Boston Harbor were among the highest in the nation (National Oceanic and Atmospheric Administration, 1987). Fish were diseased, shellfish beds were closed, and swimming beaches were unsafe after heavy rains; in general, water quality and aesthetics were below acceptable standards. Legal and political issues have always been part of Boston Harbor’s history. The environmental conditions in the 1980s were highlighted in a 1983 legal suit brought by the city of Quincy against the Metropolitan District Commission (MDC, the state agency responsible for sewage treatment) and heads of three state agencies for discharging untreated or poorly treated sewage into the harbor (Dolin, 2004). The suit never went to trial, but through the actions of a Massachusetts Superior Court, the issue of Boston Harbor contamination remained on the political and public agenda. The judge called the harbor “unsafe, unsanitary, indecent, in violation of the law (Clean Water Act), and a danger to the health and welfare of the people” (Forman, 1984). To force the state legislature to implement a plan to improve harbor conditions, the judge threatened to place the MDC in receivership and curtail new sewage hookups for industry. Under intense lobbying by business, the legislature created the Massachusetts Water Resources Authority (MWRA) in December 1984. The independent MWRA was established to manage Boston’s waste treatment system and was given the authority to float bonds to pay for major improvements in the treatment system. In 1985, a Federal court began hearings on a suit brought by the Conservation Law Foundation, the Environmental Protection Agency (USEPA), and towns of Quincy and Winthrop against the MDC and MWRA (as heir to responsibilities of the MDC) for years of violation of the Clean Water Act. The judge ruled against the defendants and required all the parties to submit a construction plan and schedule for a new sewage treatment system. From these submissions, he developed a schedule for treatment system upgrades that would give the “citizens of this commonwealth a public assurance that Boston Harbor will be cleaned up within a defined period of time” (Dolin, 2004). The MWRA’s Boston Harbor cleanup program (Levy and Connor, 1992) has transformed the Boston sewage system. Key improvements were to (1) reduce contaminants at the industrial source; (2) remediate leaks in the sewage-collection system; (3) eliminate sewage sludge discharge to the harbor; (4) upgrade sewage treatment from primary to secondary; (5) construct a new ocean outfall 15.2 km offshore in Massachusetts Bay for discharge of treated effluent (fig. 1.3); and (6) implement improvements in the combined-sewer-overflow system. As part of the harbor cleanup program, the MWRA developed a comprehensive monitoring program (summarized in MWRA, 2004) to assess changes in the harbor and bays that specifically related to the new sewage system. Additional information about conditions and processes in the coastal system on a regional scale and over a long time period was and continues to be important in predicting and interpreting local change. Implementation of the MWRA’s program and the mission of the USGS to understand the geology of the nation’s offshore waters provided an opportunity to conduct a cooperative multidisciplinary research program. This USGS program addresses basic scientific questions as well as concerns raised by management regarding the design, implementation, and assessment of the new sewage treatment system. Already active in Boston Harbor during the late 1970s, the USGS expanded research into Massachusetts Bay with a multidisciplinary program in 1989.

Massachusetts

Water as an urban resource and nuisance

Generally, when people speak of water as a resource, they are considering its good aspects and recognizing that it is essential for life and living. Sometimes or at some places or to some people, the same water may be annoying or unpleasant and thus a nuisance-for example, rain at a picnic, snow at any time except Christmas Eve, ground water in a basement, floodwater inundating personal property, and any water after it has been polluted by somebody else. For purposes of this circular, water as a resource will be defined more broadly and as a nuisance more narrowly. Water is part of the natural resource base including all aspects of the land, the air, and the water that must be considered in planning if an environment suitable for the well-being of all life is to be maintained. A nuisance, private or public, is a cause of annoyance, inconvenience, or injury that is an invasion or disturbance of the rights of some particular person or of members of a community, and it is punishable under the laws or customs accepted by the community. An activity harmful to mankind but widely practiced by the general public may be condoned by human laws and thus not be defined legally as a nuisance; the punishment for this activity is deterioration of the natural environment until it becomes unsuitable for living. The water resource, which is widely and irregularly distributed on earth, is available to man for such enjoyment and development and use as he sees fit, some use being essential to his existence. Natural variations in the quantity and quality of water are inevitable and, if they cause annoyance or injury to someone, are accepted as one of the hardships that this planet imposes upon its inhabitants; such variations are recognized as "acts of God." However, if any man or society is partly responsible for these variations, which may cause such annoyance or injury, and may become a nuisance (an invasion or disturbance of the rights of others) such a man or society may perhaps be subject to injunctions and damage suits. Legal disputes over water as a nuisance are generally deeply involved with problems of the respective rights of plaintiff and defendant. These respective rights vary among the States.

Circular

Walden Pond, Massachusetts: Environmental setting and current investigations

Introduction Walden Pond, in Concord, Massachusetts, is famous among lakes because of its unique social history. Walden was the setting for American naturalist Henry David Thoreau's well-known essay 'Walden; or, Life in the Woods,' first published in 1854. Thoreau lived and wrote at Walden Pond from July 1845 to September 1847. In 'Walden,' Thoreau combined highly admired writing on Transcendental philosophy with pioneering observations of aquatic ecology and physical aspects of limnology, the study of lakes. Because Thoreau also defended so effectively the value of living close to nature in the Walden woods, the pond is considered by many to be the birthplace of the American conservation movement. Visitors come from all over the world to the pond, which has been designated a National Historic Landmark, and its fame has resulted in a major fund drive to preserve the surrounding woods. Walden Pond has no surfacewater inflow or outflow, and much of its ground-water contributing area likely is preserved within the Walden Pond Reservation area (fig. 1). Only 15 miles from Boston, the pond is unusually clear and pristine for an urban-area lake. However, point sources of nutrients near the pond, and a large annual visitor attendance, concentrated during the summer when the swimming beach (fig. 2) is open, may contribute a nutrient load sufficient to change the pond environment. The occurrence of nuisance algal species, a recent beach closing, and an awareness of water-quality problems suffered by other ponds in the region raise concerns about the risk of ecological change at Walden Pond. Despite the role of Walden Pond as a cultural and environmental icon, little is known about the pond's ecological features, such as its internal nutrient cycling or the structure of its food web, nor have consistent measurements been made to determine whether these features are changing or are stable. Production rates of aquatic plants in lakes and ponds naturally undergo a slow increase as plant nutrients, organic matter, and soil are leached and eroded from the surrounding watershed. This process is known as eutrophication. The term 'cultural eutrophication' refers to an accelerated form of the natural process in which extra soil and nutrients are derived from people's use of fertilizer, rerouting of surface drainage, and disposal of domestic and industrial waste. Cultural eutrophication can lead to excessive growth of aquatic plants, pond filling by decayed plants and eroded soils, reduced water clarity, and depletion of dissolved oxygen in deep water with subsequent loss of cold-water fish habitat. In order to document the longterm ecological health of Walden Pond, the U.S. Geological Survey (USGS), working in cooperation with the Massachusetts Department of Environmental Management (MDEM), is investigating factors that could contribute to cultural eutrophication of Walden. Through measurements of mass balance of nutrients and oxygen in the pond's deep water, the investigation will establish a baseline data set on Walden's trophic state, which is a measure of the pond's ability to support plant growth. The baseline data will be used to detect trends and give early warning of trophic changes or trophic response to pond remediation projects. This Fact Sheet provides background information on the environmental setting, limnological features, and cultural eutrophication of Walden Pond, and describes the joint USGS/MDEM study.

Massachusetts

Cold War legacy not a post-dismantlement environmental concern

Standing ready to defend our country or to assure mutual destruction, the mid-west’s contribution to nuclear proliferation of the 60s and 70s resided innocuously beneath concrete slabs in Missouri’s cornfields. In June 1961, the U.S. Department of Defense (DOD) implemented a plan to place 150 intercontinental ballistic missiles in west-central Missouri. The missile was the solid propellant Minuteman series rocket capable of accurately delivering three nuclear warheads to targets halfway around the world. Cold war tensions at the time necessitated quick deployment of this national defense system. Construction of the missile launch facilities in Missouri began in April 1962 and within 26 months the entire system was completed and on full-alert status.

Fact Sheet

Public acceptance of management actions and judgments of responsibility for the wolves of the southern Greater Yellowstone Area: Report to Grand Teton National Park

Introduction Wolves of Grand Teton National Park and the Greater Yellowstone Area Gray wolves (Canis lupus) appeared in Grand Teton National Park (GRTE) in October of 1998, two years after being reintroduced to Yellowstone National Park (YNP). Since that time, five packs have been within the GRTE borders - Gros Ventre Pack, Nez Perce Pack, Yellowstone Delta Pack, Teton Pack, and Green River Pack (Table 1). Wolves in the Greater Yellowstone Area are increasing and spreading out geographically (USFWS and others, 2004). This dispersion was demonstrated recently by the death of a 2-year-old female wolf from the Swan Lake pack on I-70 in Colorado (June 7, 2004; http://mountain-prairie.USFWS.gov/pressrel /04-43.htm). The organization of wolf packs in the GYA is dynamic and highly structured. In 2003, for example, a wolf from the Teton Pack joined with the Green River Pack, and several young wolves left the Teton Pack and moved south (USFWS and others, 2004). Pack size (averaging five to ten members) is dependent on hunting efficiency, which depends on prey size, type, and density. Each pack defends home ranges of several hundred square miles. The social structure of the pack is based on a breeding pair (an alpha male and female). Other wolves in the pack can be categorized as betas (males and/or females second in rank to the alphas), subordinates, pups, and occasional omegas (outcasts). Because generally only the alpha pair breeds, subordinate wolves of reproductive age must disperse from their packs and form new associations in order to breed. (http://www.nps.gov/grte/wolf/biolo.htm). The reintroduced wolves are classified by the U.S. Fish and Wildlife Service (USFWS) as "nonessential experimental" under section 10(j) of the Endangered Species Act. The recovery criteria for the GYA wolves were met in 2002 for removing the wolves from the Endangered Species List (30 or more breeding pairs). Currently, the USFWS manages wolf populations in the GYA until delisting occurs. After delisting, state Fish and Wildlife Services in Montana, Idaho, and Wyoming will be responsible for managing wolves. Each state must submit a wolf management plan to the USFWS which then must be approved before management shifts occur. As of this writing, the process of delisting the wolves in the state of Wyoming is ongoing. However, the reclassification of wolves nationwide was completed on April 1, 2003. Wolves outside of YNP changed in status from endangered to threatened. The wolves classified in the experimental nonessential population did not change in status (USFWS and others, 2004). This classification of experimental nonessential population allows for flexibility in management decisions concerning the wolves (Smith and others, 2004). For example, control actions in the GYA included trapping and radio-collaring four wolves; intensive monitoring; increasing riders on grazing allotments; harassing wolves with rubber bullets, cracker shells, and lights; moving livestock to different pastures; and issuing four shoot on-sight permits. When non-lethal control methods were not effective, wolves were killed in an attempt to prevent further livestock depredations (USFWS and others, 2004; Table 1). At the same time that wolf numbers are rising, human population statistics in the GRTE area are also rising. The population of Teton County, Wyoming in 1990 was just over 11,000 people; today that number has increased to approximately 19,000 (U.S. Census Bureau, 2005). park visitation for GRTE has been substantial over the last several years with an average visitation of 2.5 million visitors (NPS, 2004a). Furthermore, land ownership surrounding GRTE and the establishment of grazing rights within park boundaries are problem areas for wolf-human interactions due to livestock depredation. With increasing numbers of visitors, residents, and livestock it is reasonable to assume that conflicts are going to increase also. In 1950, GRTE was expanded to in

Open-File Report

Southwestern Willow Flycatcher Breeding Site and Territory Summary - 2007

The Southwestern willow flycatcher (Empidonax traillii extimus; hereafter references to willow flycatcher and flycatcher refer to E.t. extimus, except where specifically noted) is an endangered bird that breeds only in dense riparian habitats in parts of six Southwestern states (Arizona, New Mexico, southern California, extreme southern Nevada, southern Utah, and southwestern Colorado). Since 1993, hundreds of Southwestern willow flycatcher surveys have been conducted each year, and many new flycatcher breeding sites located. This document synthesizes the most current information available on all known Southwestern willow flycatcher breeding sites. This rangewide data synthesis was designed to meet two objectives: (1) identify all known Southwestern willow flycatcher breeding sites and (2) assemble data to estimate population size, location, habitat, and other information for all breeding sites, for as many years as possible, from 1993 through 2007. This report provides data summaries in terms of the number of flycatcher sites and the number of territories. When interpreting and using this information, it must be kept in mind that a 'site' is a geographic location where one or more willow flycatchers establishes a territory. Sites with unpaired territorial males are considered breeding sites, even if no nesting attempts were documented. A site is often a discrete patch of riparian habitat but may also be a cluster of riparian patches; there is no standardized definition for site, and its use varies within and among states. For example, five occupied habitat patches along a 10-km stretch of river might be considered five different sites in one state but only a single site in another state. This lack of standardization makes comparisons based on site numbers problematic. Researchers for this report generally deferred to statewide summary documents or to local managers and researchers when delineating a site for inclusion in the database. However, to avoid inflating the number of sites and to establish more consistent definitions of the term 'site', adjacent and nearby sites from statewide reports were sometimes considered as a single site for the purposes of the rangewide data summary. Any combining or splitting of sites at the rangewide level was done on a case-by-case basis. Because of differences in site definitions, one should not evaluate the relative importance of a geographic region (such as drainage, watershed, or state) simply according to the number of flycatcher sites. A 'territory' is an exclusive defended area within a breeding site. Although detailed monitoring studies have identified unpaired territorial males and polygynous males at some flycatcher breeding sites, for the purposes of this report a territory is equivalent to the exclusive breeding area of a pair of flycatchers. In general, the concept of territory is more similar among states and different investigators than site; thus, it is a more robust unit to use for summaries and comparisons. However, note that the definition of a polygynous territory is not consistent among states; a male polygynously paired with two females would be considered one territory in some states and two territories in other states. For each site, we referred to reports or spoke directly with researchers and managers to gather information such as management entity/agency, location (state, drainage, elevation), gross habitat type (native, exotic, or mixed; dominant tree species), and number of flycatcher territories. Synthesizing the information on more than 200 breeding sites is challenging because annual data-collection and survey-reporting requirements are not standardized rangewide, and the nature and degree of readily available information varies widely from state to state. This is particularly true for areas such as California, where there are many flycatcher sites but surveyors are not required to submit standardized flycatcher survey forms. The lack of c

Open-File Report

Quality assurance and quality control of geochemical data—A primer for the research scientist

Geochemistry is a constantly expanding science. More and more, scientists are employing geochemical tools to help answer questions about the Earth and earth system processes. Scientists may assume that the responsibility of examining and assessing the quality of the geochemical data they generate is not theirs but rather that of the analytical laboratories to which their samples have been submitted. This assumption may be partially based on knowledge about internal and external quality assurance and quality control (QA/QC) programs in which analytical laboratories typically participate. Or there may be a perceived lack of time or resources to adequately examine data quality. Regardless of the reason, the lack of QA/QC protocols can lead to the generation and publication of erroneous data. Because the interpretations drawn from the data are primary products to U.S. Geological Survey (USGS) stakeholders, the consequences of publishing erroneous results can be significant. The principal investigator of a scientific study ultimately is responsible for the quality and interpretation of the project's findings, and thus must also play a role in the understanding, implementation, and presentation of QA/QC information about the data. Although occasionally ignored, QA/QC protocols apply not only to procedures in the laboratory but also in the initial planning of a research study and throughout the life of the project. Many of the tenets of developing a sound QA/QC program or protocols also parallel the core concepts of developing a good study: What is the main objective of the study? Will the methods selected provide data of enough resolution to answer the hypothesis? How should samples be collected? Are there known or unknown artifacts or contamination sources in the sampling and analysis methods? Assessing data quality requires communication between the scientists responsible for designing the study and those collecting samples, analyzing samples, treating data, and interpreting results. This primer has been developed to provide basic information and guidance about developing QA/QC protocols for geochemical studies. It is not intended to be a comprehensive guide but rather an introduction to key concepts tied to a list of relevant references for further reading. The guidelines are presented in stepwise order beginning with presampling considerations and continuing through final data interpretation. The goal of this primer is to outline basic QA/QC practices that scientists can use before, during, and after chemical analysis to ensure the validity of the data they collect with the goal of providing defendable results and conclusions.

Open-File Report

Least Bell's Vireos and Southwestern Willow Flycatchers at the San Luis Rey flood risk management project area in San Diego County, California—Breeding activities and habitat use—2020 annual report

Executive Summary Surveys and monitoring for the endangered Least Bell’s Vireo ( Vireo bellii pusillus ; vireo) were done at the San Luis Rey Flood Risk Management Project Area (Project Area) in the city of Oceanside, San Diego County, California, between March 31 and July 20, 2020. We completed four protocol surveys during the breeding season, supplemented by weekly territory monitoring visits. We identified a total of 161 territorial male vireos; 145 were confirmed as paired and 4 were confirmed as single males. For the remaining 12 territories, we were unable to confirm pair status. Three transient vireos were detected in 2020. The vireo population in the Project Area increased by 26 percent from 2019 to 2020. Vireo populations increased across San Diego County, with a 39-percent increase documented at Marine Corps Base Camp Pendleton (MCBCP); a 58-percent increase at Marine Corps Air Station; a 78-percent increase on the Otay River; and a 7-percent increase in the population on the middle San Luis Rey River. We used an index of treatment (Treatment Index) to evaluate the impact of on-going vegetation clearing on the Project Area vireo population. The Treatment Index measures the cumulative effect of vegetation treatment within a territory (since 2005) by using the percent area treated weighted by the number of years since treatment. We found that the Treatment Index for unoccupied habitat was more than five times that of occupied habitat, indicating that vireos selected less disturbed habitat in which to settle. We monitored vireo nests at three general site types: (1) within the flood channel where exotic and native vegetation removal has occurred regularly (Channel), (2) three sites next to the flood channel where limited exotic and native vegetation removal has occurred (Off-channel), and (3) three sites that have been actively restored by planting native vegetation (Restoration). Nesting activity was monitored in 100 territories, 4 of which were occupied by single males. Hatching success was higher in the Channel relative to the Off-channel. We found no other differences between Channel, Off-channel, and Restoration nests in terms of clutch size or fledging success. There also was no difference in measures of productivity per pair between Channel, Off-channel, Restoration, and Mixed territories (territories that were classified as one site type but nesting occurred in another site type, or where multiple site types were used for nesting). Overall, breeding success and productivity were lower in 2020 than in 2019, with 69 percent of pairs fledgling at least one young and pairs fledging an average of 2.1±1.7 young. To investigate whether the cumulative years of treatment had an impact on vireo reproductive effort, we looked at the effects of the Treatment Index on reproductive parameters. Results from generalized linear models indicated that treatment did not have an effect on vireo nesting effort or the number of vireo fledglings per pair produced in 2020. Similarly, our analysis of nest survival for 2020 revealed no effect of Treatment Index on daily survival rate. Analysis of vegetation data collected at vireo nests from 2006 to 2020 revealed that vegetation at 1–2 meters (m) from the ground was the most important predictor of daily survival rate. There were differences in nest-placement characteristics among site types and successful/unsuccessful nests. Channel nests were placed higher in the vegetation than Off-channel or Restoration nests. Host plant height, distance to edge of host plant, and distance to edge of vegetation clump were greater at Channel sites compared with Off-channel sites, but were not different from Restoration sites. Within sites, we found only one difference between successful and unsuccessful nests. At Off-channel sites, successful nests were placed higher in the vegetation than unsuccessful nests. Red/arroyo willow ( Salix laevigata or Salix lasiolepis ) and mule fat ( Baccharis salicifolia ) were the species most commonly selected for nesting by vireos in all 3 site types. Vireos used a wider variety of species for nesting in Channel and Off-channel sites (7 and 10 species, respectively) compared to Restoration sites (3 species). Ninety-three vireos banded before the 2020 breeding season were resighted and identified at the Project Area in 2020, all of which were originally banded at the Project Area. Adult birds of known age ranged from 1 to 9 years old. A total of 171 vireos were newly banded in 2020. Twenty-eight adult vireos were banded with a unique color combination, and 143 nestlings were banded with a single dark blue numbered federal band on the left leg. Between 2006 and 2020, survivorship of males (67±10 percent) was consistently higher than females (59±11 percent). First-year birds from 2006 to 2020 had an average over-winter survivorship of 17±5 percent. First-year dispersal in 2020 averaged 2.9±2.9 kilometers (km), with the longest dispersal (13.5 km) by a female that was recaptured at Las Flores Creek, MCBCP. From 2007 to 2012, most returning first-year vireos returned to the Project Area, whereas from 2013 to 2017, the majority of returning birds dispersed to areas outside of the Project Area. In 2018, the trend shifted, and most first-year vireos returned to the Project area. This trend continued in 2020 with most first-year vireos returning to the Project Area; 77 percent of all re-encountered first-year birds returned to the Project Area and 23 percent dispersed to areas outside of the Project Area (upstream to the middle San Luis Rey River and to drainages on MCBCP). Most of the returning adult male vireos showed strong between-year site fidelity to their previous territories. Eighty percent of males (45/56) occupied a territory in 2020 that they had defended in 2019 (within 100 m). Thirty-three percent of females (2/6) detected in 2020 returned to a territory that they occupied in 2019. The average between-year movement for returning adult vireos was 0.1±0.5 km. We completed four protocol surveys for the endangered Southwestern Willow Fycatcher ( Empidonax traillii extimus ; flycatcher) at the Project Area between May 20 and July 20, 2020. No Willow Flycatchers were detected in the Project Area in 2020. A total of 46 vegetation transects (526 points) were sampled at the San Luis Rey Flood Risk Management Project Area in 2020. Seventy-one percent (376/526) of points were in the Channel and 22 percent (115/526) were at Upper Pond. The remaining 7 percent (35/526) were at the Whelan Restoration site. Foliage cover below 1 m was higher at the Channel points compared to Upper Pond and Whelan Restoration. Higher foliage cover in the Channel was attributed to the higher herbaceous component. However, foliage cover from 1 to 3 m was higher at the Whelan Restoration site compared to both Upper Pond and the Channel. Average canopy height was similar at all three site types and was 4.4 m or less. From 2006 to 2020, total foliage cover declined above 1 m in the Channel, from 4 to 5 m at Upper Pond, and above 8 m at Whelan Restoration. Within the Channel, the steepest declines occurred between 2009 and 2013 and between 2014 and 2016. Since 2016, we observed an increase in percent foliage between 0 and 2 m within the Channel, but for other height classes, percent cover remained below levels detected before 2009. Changes in cover at Upper Pond and Whelan Restoration appeared to be driven by the loss of tall tree cover. The vegetation mowing and treatment activities, in combination with lack of precipitation (especially between 2012 and 2016), may have contributed to the decline in foliage cover observed from 2006 to 2020. We sampled vegetation at 49 vireo nests and 49 random plots (“territory” plots) within territories in the Channel and Upper Pond following the 2020 breeding season. Vireos in the Channel selected territories with significantly more foliage cover above 2 m but less cover below 1 m relative to the available habitat. In contrast, Channel vireos selected nest sites within their territories with lower foliage cover above 3 m and were non-selective with regard to cover below 2 m. Vireos at Upper Pond generally were less selective with regard to territory and nest sites but tended to select territories with more foliage cover from 1 to 2 m and above 8 m, and they selected nest sites within their territories with greater foliage cover from 0 to 1 m.

California

Least Bell's Vireos and Southwestern Willow Flycatchers at the San Luis Rey Flood Risk Management Project Area in San Diego County, California—Breeding activities and habitat use—2022 annual report

Executive Summary We completed four protocol surveys for Least Bell’s Vireos ( Vireo bellii pusillus ; vireo) during the breeding season, supplemented by weekly territory monitoring visits. We identified a total of 133 territorial male vireos; 114 were confirmed as paired, and 3 were confirmed as single males. For the remaining 16 territories, we were unable to confirm breeding status. Two transient vireos were detected in 2022. The vireo population in the Project Area increased by 9 percent from 2021 to 2022. The vireo population at Marine Corps Base Camp Pendleton also increased (4 percent), whereas the population at Marine Corps Air Station remained relatively stable (decreased from 10 pairs to 9) and the Otay River population decreased by 10 percent (2 territories). We used an index of treatment (Treatment Index) to evaluate the effect of on-going vegetation clearing on the Project Area vireo population. The Treatment Index measures the cumulative effect of vegetation treatment within a territory (since 2005) by using the percentage area treated weighted by the number of years since treatment. We determined that the Treatment Index for unoccupied habitat was more than four times that of occupied habitat, indicating that vireos selected habitat that was less treated in which to settle. We monitored vireo nests at three general site types: (1) within the flood channel where exotic and native vegetation removal has occurred regularly (Channel), (2) three sites near the flood channel where limited exotic and native vegetation removal has occurred (Off-channel), and (3) three sites that have been actively restored by planting native vegetation (Restoration). Nesting activity was monitored in 80 territories, 3 of which were occupied by single males and 1 by a male whose breeding status could not be confirmed. Overall, 38 percent of completed nests were successful and nest success did not differ among the three sites. In 2022, there were no differences with regard to clutch size, hatching, or fledging success among Channel, Off-channel and Restoration sites. Overall breeding success and productivity were slightly higher in 2022 than in 2021, with 72 percent of pairs fledgling at least one young and pairs fledging an average of 2.2±1.7 young. To investigate if the cumulative years of treatment had an effect on vireo reproductive effort, we looked at the effects of the Treatment Index on reproductive parameters. Results from generalized linear models indicated that treatment did not have an effect on vireo nesting effort or the number of vireo fledglings per pair produced in 2022. Similarly, we did not detect an effect of Treatment Index on daily survival rate (DSR) of nests. Analysis of vegetation data collected at vireo nests from 2006 to 2022 did not reveal an effect of vegetation cover at the nest on DSR. We did find, however, that Channel nests were placed higher in the host plant than Off-channel nests. In the Channel and Off-channel sites, successful nests were placed closer to the edge of the host plants than unsuccessful nests. Additionally, successful Off-channel nests were placed lower in the vegetation, in shorter host plants, and closer to the edge of the vegetation clump than unsuccessful nests. Red/arroyo willow ( Salix laevigata or Salix lasiolepis ) were the species most commonly selected for nesting by vireos in all three site types. Black willow ( Salix gooddingii ) and mule fat ( Baccharis salicifolia ) also were commonly used. Vireos used a wider variety of species for nesting in Channel and Off-channel sites (eight and six species, respectively) compared to Restoration sites (two species), although there was limited nesting in Restoration sites in 2022. There were 43 vireos banded before the 2022 breeding season that were resighted and identified at the Project Area in 2022, all of which were originally banded in the Project Area. Adult birds of known age ranged from 1 to 7 years old. A total of 146 vireos were newly banded in 2022. There were 8 adult vireos banded with a unique color combination, and 138 nestlings were banded with a single dark blue numbered federal band on the left leg. Between 2006 and 2022, survivorship of males (66±11 percent) was consistently higher than that of females (59±12 percent). First-year birds from 2006 to 2022 had an average annual survivorship of 15±6 percent. First-year dispersal in 2022 averaged 6.7±7.4 kilometers (km), with the longest dispersal (15.3 km) by a male that was recaptured at Fallbrook Creek, Fallbrook Naval Weapons Station (FNWS). From 2007 to 2011, most returning first-year vireos returned to the Project Area, whereas from 2014 to 2016, the majority of returning birds dispersed to areas outside of the Project Area. From 2018 to 2021, the trend shifted, and more first-year vireos returned to the Project area. In 2022, only one first-year vireo returned to the project area and two dispersed to sites outside the Project Area (upstream to the middle San Luis Rey River and to Fallbrook Creek, FNWS). However, the total number of identified first-year vireos was low and the trend in 2022 will likely shift as additional returning first-year vireos are identified in subsequent years. Most of the returning adult male vireos showed strong between-year site fidelity to their previous territories. Seventy-three percent of males (27/37) occupied a territory in 2022 that they had defended in 2021 (within 100 meters [m]). There were no females (0/4) detected in 2022 that returned to a territory they occupied in 2021; however, 50 percent of females (2/4) detected in 2022 returned to areas adjacent to their previous territories (within 300 m). The average between-year movement for returning adult vireos was 0.3±0.7 km. The amount of treatment at adults’ 2021 territories did not affect the distance adults moved to their 2022 territories. We completed four protocol surveys for the endangered Southwestern Willow Flycatcher ( Empidonax traillii extimus ; flycatcher) at the Project Area between May 16 and July 25, 2022. Four transient Willow Flycatchers were detected in the Project Area in 2022. Two transients were detected in Reach 1, one in Reach 3a, and one in Pilgrim Pond. There were not any resident flycatchers documented in the Project Area in 2022. A total of 46 vegetation transects (528 points) were sampled at the Project Area in 2022. Seventy-one percent (378/528) of points were located in the Channel, and 22 percent (115/528) were in Upper Pond. The remaining 7 percent (35/528) of points were at the Whelan Restoration site. Foliage cover below 2 m was higher at the Channel points compared to Upper Pond and Whelan Restoration, which can be attributed to the dense herbaceous vegetation that grows after mowing. Above 2 m, foliage cover was similar at the Channel and Whelan Restoration sites and was higher than at Upper Pond. Average canopy height was higher in the Channel (5.6±3.4 m) compared to Upper Pond (4.7±2.9 m) and Whelan Restoration (4.6±1.9 m). From 2006 to 2022, total foliage cover declined above 2 m in the Channel, in contrast to Upper Pond and Whelan Restoration, where little directional change in vegetation cover has occurred and where vegetation cover has largely recovered to 2006 levels. Within the Channel, the steepest declines occurred between 2009 and 2013 and between 2014 and 2016. Since 2016, we observed an increase in foliage cover, largely herbaceous, between 0 and 2 m within the Channel. The percent cover remained below levels detected before 2009 for other height classes. We sampled vegetation at 44 vireo nests and 44 random plots (“territory” plots) within territories in the Channel and Upper Pond after the 2022 breeding season. Vireos in the Channel established territories in areas with significantly more cover from 3 to 6 m but less cover below 2 m relative to the available habitat. Within territories, Channel vireos selected nest sites with significantly more foliage cover from 2 to 3 m. Vireos at Upper Pond established territories in areas with significantly more foliage cover from 5 to 6 m and below 1 m relative to available habitat. However, within territories, Upper Pond vireos selected nest sites with significantly less foliage cover from 5 to 6 m and below 1 m. Data either are not available or have limited availability owing to restrictions of the funding entity (U.S. Army Corps of Engineers). Please contact Christopher Chabot, Planning Division, Los Angeles District, U.S. Army Corps of Engineers, for more information.

Califoria

Least Bell's Vireos and Southwestern Willow Flycatchers—Breeding activities and habitat use—2023 annual report

Executive Summary We completed four protocol surveys for Least Bell’s Vireos ( Vireo bellii pusillus ; hereinafter vireo) during the breeding season, supplemented by weekly territory monitoring visits between April 6 and July 20 at the San Luis Rey Flood Risk Management Project Area (hereinafter Project Area). We identified a total of 136 territorial male vireos; 121 were confirmed as paired, and 4 were confirmed as single males. For the remaining 11 territories, we were unable to confirm breeding status. In 2023, two transient vireos were detected. The vireo population in the Project Area increased by 2 percent from 2022 to 2023. Populations in southern San Diego County also increased (by 6 percent on the Otay River) or were stable (Salt Creek/Wolf Canyon). In contrast, the vireo population at Marine Corps Base Camp Pendleton (MCBCP) and at Marine Corps Air Station decreased by 2 and 10 percent, respectively. We used an index of treatment (hereinafter Treatment Index) to evaluate the effect of ongoing vegetation clearing on the Project Area vireo population. The Treatment Index measures the cumulative effect of vegetation treatment within a territory by using the percentage area treated weighted by the number of years since treatment. We determined that the Treatment Index for an unoccupied habitat was more than four times higher than that of an occupied habitat, indicating that vireos selected habitats that were less treated in which to settle. We monitored vireo nests at three general site types: (1) within the flood channel where non-native and native vegetation removal has occurred regularly (hereinafter Channel), (2) three sites near the flood channel where limited non-native and native vegetation removal has occurred (hereinafter Off-channel), and (3) three sites that have been actively restored by planting native vegetation (hereinafter Restoration). Nesting activity was monitored in 84 territories, 4 of which were occupied by single males. Overall, 46 percent of completed nests were successful, and nest success did not differ among the three sites. In 2023, we found that territories in the Channel had greater hatching success per egg compared to Off-channel, but there were no other differences with regard to clutch size, hatching, or fledging success among Channel, Off-channel, and Restoration sites. Overall breeding success and productivity were slightly higher in 2023 than in 2022, with pairs fledging an average±standard deviation of 3.1±2.1 young and 79 percent of pairs fledging at least 1 young. To investigate if the cumulative years of treatment had an effect on vireo reproductive effort, we looked at the effects of the Treatment Index on reproductive parameters. Results from generalized linear models indicated that treatment did not have an effect on vireo nesting effort (the number of nest attempts) or the number of vireo fledglings per pair produced in 2023. Similarly, we did not detect an effect of Treatment Index on the daily survival rate (DSR) of nests. Analysis of vegetation data collected at vireo nests from 2006 to 2023 did not reveal an effect of vegetation cover at the nest on DSR. We did find, however, that Channel nests were placed higher in and farther from the edge of the host plant than Off-channel nests. Within sites, we did not detect any differences in vegetation cover between successful and unsuccessful nests. Red/arroyo willow ( Salix laevigata or Salix lasiolepis ) and mule fat ( Baccharis salicifolia ) were the species most commonly selected for nesting by vireos in all three site types. Black willow ( Salix gooddingii ) and sandbar willow ( Salix exigua ) also were commonly used. Vireos used a wider variety of species for nesting in Channel and Off-channel sites (10 and 13 species, respectively) compared to Restoration sites (2 species), although there was limited nesting in Restoration sites in 2023. There were 51 vireos banded before the 2023 breeding season that were resighted and identified at the Project Area in 2023. Two of these vireos were originally banded outside of the Project Area, at the Santa Margarita River on MCBCP. Adult birds of known age ranged from 1 to 7 years old. Between 2006 and 2023, survival of males (66±11 percent) was consistently higher than that of females (60±12 percent). First-year birds from 2006 to 2022 had an average annual survival of 15±5 percent. First-year dispersal in 2023 averaged 20.2±31.3 kilometers (km), with the longest dispersal (76.3 km) by a female that was recaptured at Wolf Canyon, a tributary to Otay River. From 2007 to 2012, most returning first-year vireos returned to the Project Area, whereas from 2014 to 2016, a greater proportion of returning birds dispersed to areas outside of the Project Area. From 2018 to 2022, the trend shifted, and more first-year vireos returned to the Project Area, except for 2022 when only one out of five first-year vireos returned to the Project Area. This trend continued in 2023: 71 percent of all first-year vireos returned to the Project Area, and 29 percent dispersed to areas outside of the Project Area (San Diego River and Wolf Canyon). Most of the returning adult male vireos showed strong between-year fidelity to their previous territories. In 2023, 94 percent of males (34/36) occupied a territory that they had defended in 2022 (within 100 meters [m]). In 2023, 33 percent of females (1/3) detected returned to a territory they occupied in 2022. The average between-year movement for returning adult vireos was 0.2±0.9 km. The amount of treatment at adults’ 2022 territories did not affect the distance adults moved to their 2023 territories. We completed four protocol surveys for the endangered Southwestern Willow Flycatcher ( Empidonax traillii extimus ; hereinafter flycatcher) at the Project Area between May 15 and July 21, 2023. In 2023, four transient Willow Flycatchers were detected in the Project Area. Two transients were detected in Reach 1, one in Reach 3a, and one in Whelan Mitigation. No resident flycatchers were documented in the Project Area in 2023. A total of 46 vegetation transects (516 points) were sampled in the Project Area in 2023. There were 71 percent (368/516) of points located in the Channel, and 22 percent (113/516) were in Upper Pond. The remaining 7 percent (35/516) of points were at the Whelan Restoration site. Foliage cover below 1 m was higher at the Channel points and Upper Pond compared to Whelan Restoration. From 1 to 3 m, foliage cover was similar at all 3 sites; however, above 3 m foliage cover was higher in the Channel compared to the Upper Pond and Whelan Restoration sites. Average canopy height was higher in the Channel (5.6±3.8 m) compared to Upper Pond (4.7±2.7 m) and Whelan Restoration (4.0±2.0 m). From 2006 to 2023, total foliage cover declined from 2 to 3 m and above 6 m in the Channel, in contrast to Upper Pond and Whelan Restoration, where little directional change in vegetation cover has occurred and where vegetation cover has largely recovered to 2006 levels. Within the Channel, the steepest declines occurred between 2009 and 2013 and between 2014 and 2016. Since 2016, we observed an increase in foliage cover, largely herbaceous, between 0 and 2 m within the Channel. Although increases were observed at all height classes after 2016, percentage cover has remained below levels measured before 2009. We sampled vegetation at 45 vireo nests and 45 random plots (territory plots) within territories in the Channel and Upper Pond after the 2023 breeding season. Vireos in the Channel established territories in areas with significantly more cover from 3 to 7 m but less cover below 1 m relative to the available habitat. Within territories, Channel vireos selected nest sites largely at random, but with significantly less foliage cover from 4 to 5 m. Vireos at Upper Pond established territories in areas with significantly more foliage cover below 4 m and from 5 to 6 m relative to available habitat. Within territories, Upper Pond vireos also selected nest sites at random except for a preference for sites with significantly less foliage cover below 1 m.

Open-File Report

The effects of management practices on grassland birds—Rates of Brown-headed Cowbird (Molothrus ater) parasitism in nests of North American grassland birds

The Brown-headed Cowbird ( Molothrus ater ) is an obligate brood parasite that relies entirely on host species to incubate its eggs and rear its young. The Brown-headed Cowbird is a host generalist; eggs of the cowbird have been found in nests of more than 220 species of North American birds. Although cowbirds are not considered a serious threat to their primary avian hosts, cowbird brood parasitism has been recognized as a potential threat to the survival of threatened and endangered species and has been implicated in the declines of other bird species through reduced reproductive output. In this chapter, we summarized the rates of cowbird brood parasitism for 40 grassland bird species. The reported rates of brood parasitism by Brown-headed Cowbirds varied considerably among and within grassland host species, and the factors that influence parasitism rates were described. Defenses possessed by cowbird hosts in grasslands include aggressively defending their nests against female cowbirds, egg rejection, desertion of parasitized nests, and burying cowbird eggs in the nest lining.

Professional Paper

Distribution and abundance of fallow deer leks at Point Reyes National Seashore, California

Only two species of ungulates (hoofed mammals) are native to Marin County, tule elk (Cervis elaphus nannodes) and Columbian black-tailed deer (Odocoileus hemionus columbianus). In the 1940s, European fallow deer (Dama dama) obtained from the San Francisco Zoo were released at Point Reyes. When Point Reyes National Seashore was established in 1962, fallow deer were well established within the boundaries of the National Seashore. The fallow deer population was estimated to be 500 in 1973 (Wehausen, 1973) and that number increased to 860 by 2005 (National Park Service, unpubl. data). Fallow deer have an unusual mating system. During the fall mating season (or rut), male fallow deer establish areas known as leks where they display to potential mates (Hirth, 1997). This behavior is unique among deer and elk, but it is similar to breeding systems used by grouse and a few other birds and mammals. Formation of leks in ungulates decreases the number of aggressive encounters in which dominant males are involved when the local male density becomes too high, because the spatial stability of territories in leks reduces the number of aggressive encounters between males (Hovi et al., 1996; Pelabon et al., 1999). A fallow deer lek is typically an area of about 100-150 m2 and typically includes two to five males. Using their hooves and antlers, each male clears away most or all of the vegetation and digs a rutting pit that he defends throughout the breeding season.

Scientific Investigations Report