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Alan D. Afton

Publications and source records attributed to Alan D. Afton.

At least 19 recordsLinked to original sources

Evaluating the waterfowl breeding population and habitat survey for scaup

Potential bias in breeding population estimates of certain duck species from the Waterfowl Breeding Population and Habitat Survey (WBPHS) has been a concern for decades. The WBPHS does not differentiate between lesser ( Aythya affinis ) and greater ( A . marila ) scaup, but lesser scaup comprise 89% of the combined scaup population and their population estimates are suspected to be biased. We marked female lesser scaup (i.e., marked scaup) in the Mississippi and Atlantic Flyways, Canada and United States, with implantable satellite transmitters to track their spring migration through the traditional and eastern survey areas of the WBPHS, 2005–2010. Our goal was to use data independent of the WBPHS to evaluate whether breeding population estimates for scaup were biased and identify variables that might be used in the future to refine population estimates. We found that the WBPHS estimates of breeding scaup are biased because, across years, only 30% of our marked scaup had settled for the breeding period when the strata in which they settled were surveyed, 43% were available to be counted in multiple survey strata as their migration continued during the WBPHS, 32% settled outside the WBPHS area, the number of times a marked scaup was available to be counted by survey crews varied positively with the latitude that a marked scaup settled on breeding areas, the probability of a marked scaup being in a stratum while it was surveyed varied among years, and these probabilities were positively correlated with the traditional and eastern breeding population estimates for scaup. Annual population estimates derived from banding data provide a less biased and preferable method of monitoring scaup population status and trend. Development of models that include metrics such as survey stratum latitude and annual spring environmental conditions might potentially be used to improve scaup breeding population estimates derived from the WBPHS, but independent estimates from banding data would be important to evaluate such models.

Journal of Wildlife Management↗

Temporal variation in survival and recovery rates of lesser scaup: A response

We recently analyzed long-term (1951–2011) continental band-recovery data from lesser scaup ( Aythya affinis ) and demonstrated that harvest rates declined through time, but annual survival rates exhibited no such trends; moreover, annual harvest and survival rates were uncorrelated for all age-sex classes. We therefore concluded that declining fecundity was most likely responsible for recent population declines, rather than changes in harvest or survival. Lindberg et al. ( 2017 ) critiqued our conclusions, arguing that we did little more than fail to reject a null hypothesis of compensatory mortality, postulated ecologically unrealistic changes in fecundity, and failed to give sufficient consideration to additive harvest mortality. Herein, we re-summarize our original evidence indicating that harvest has been compensatory, or at most weakly additive, and demonstrate that our analysis had sufficient power to detect strongly additive mortality if it occurred. We further demonstrate that our conclusions were not confounded by population size, band loss, or individual heterogeneity, as suggested by Lindberg et al. ( 2017 ), and we provide additional support for our conjecture that low fecundity played a major role in declining scaup populations during 1983–2006. We therefore reiterate our original management recommendations: given low harvest rates and lack of demonstrable effect on scaup survival, harvest regulations could return to more liberal frameworks, and waterfowl biologists should work together to continue banding lesser scaup and use these data to explore alternative hypotheses to identify the true ecological causes of population change, given that it is unlikely to be excessive harvest.

Journal of Wildlife Management↗

Environmental factors influence lesser scaup migration chronology and population monitoring

Identifying environmental metrics specific to lesser scaup ( Aythya affinis ; scaup) spring migration chronology may help inform development of conservation, management and population monitoring. Our objective was to determine how environmental conditions influence spring migration of lesser scaup to assess the effectiveness of the Waterfowl Breeding Population and Habitat Survey in accurately estimating scaup populations. We first compared peak timing of mallard ( Anas platyrhynchos ) and scaup migration from weekly ground surveys in North Dakota, USA because the Waterfowl Breeding Population and Habitat Survey is designed to capture annual mallard migration. As predicted, we detected that peak timing of scaup and mallard migrations differed in 25 of 36 years investigated (1980–2010). We marked scaup with satellite transmitters ( n = 78; 7,403 locations) at Long Point, Lake Erie, Ontario, Canada; Pool 19 of the Mississippi River, Iowa and Illinois, USA; and Presque Isle Bay, Lake Erie, Pennsylvania, USA. We tested the assumption that our marked scaup were representative of the continental population using the traditional survey area by comparing timing of migration of marked birds and scaup counted in the North Dakota Game and Fish Department survey. We detected a strong positive correlation between marked scaup and the survey data, which indicated that marked scaup were representative of the population. We subsequently used our validated sample of marked scaup to investigate the effects of annual variation in temperature, precipitation, and ice cover on spring migration chronology in the traditional and eastern survey areas of the Waterfowl Breeding Population and Habitat Survey, 2005–2010. We evaluated competing environmental models to explain variation in timing and rate of scaup migration at large-scale and local levels. Spring migration of scaup occurred earlier and faster during springs with warmer temperatures and greater precipitation, variables known to influence energy budgets and wetland availability. Our results suggest that surveys designed to index abundance of breeding mallards is imprecise for estimating scaup abundance, and inaccurate at estimating breeding population size by survey stratum.

Journal of Wildlife Management↗

Temporal variation in survival and recovery rates of lesser scaup

Management of lesser scaup (Aythya affinis) has been hindered by access to reliable data on population trajectories and vital rates. We conducted a Bayesian analysis of historical (1951–2011) band-recovery data throughout North America to estimate annual survival and recovery rates for juvenile and adult male and female lesser scaup to determine if increasing harvest or declining survival rates have contributed to population changes and to determine if harvest has been primarily additive or compensatory. Annual recovery rates were low, ranging from 1% to 4% for adults and 2% to 10% for juveniles during most years, with trend models indicating that recovery rates have declined through time for all age–sex classes. Annual survival (mid-Aug to mid-Aug) averaged 0.402 (σ ̂ 0.043) for juvenile males, 0.416 (σ ̂ 0.067) for juvenile females, 0.689 (σ ̂ 0.109) for adult males, and 0.602 (σ ̂ 0.115) for adult females, where σ ̂ represents an estimate of annual process variation in each survival rate. Annual survival rates exhibited no evidence of long-term declines or negative correlations with annual recovery rates (i.e., an index of harvest intensity) for any age–sex class, suggesting that declining fecundity was the most likely explanation for population declines during 1975–2005. We conclude that hunting mortality played a minor role in affecting population dynamics of lesser scaup and waterfowl managers could take a less cautious approach in managing harvest, especially if recruiting or maintaining waterfowl hunters are viewed as important management objectives.

Journal of Wildlife Management↗

Time constraints in temperate-breeding species: Influence of growing season length on reproductive strategies

Organisms that reproduce in temperate regions have limited time to produce offspring successfully, and this constraint is expected to be more pronounced in areas with short growing seasons. Information concerning how reproductive ecology of endotherms might be influenced by growing season length (GSL) is rare, and species that breed over a broad geographic range provide an opportunity to study the effects of time constraints on reproductive strategies. We analyzed data from a temperate‐breeding bird, the lesser scaup Aythya affinis ; hereafter scaup, collected at eight sites across a broad gradient of GSL to evaluate three hypotheses related to reproductive compensation in response to varying time constraints. Clutch initiation date in scaup was unaffected by GSL and was unrelated to latitude; spring thaw dates had a marginal impact on timing of breeding. Clutch size declined during the nesting season, as is reported frequently in bird species, but was also unaffected by GSL. Scaup do not appear to compensate for shorter growing seasons by more rapidly reducing clutch size. This study demonstrates that this species is remarkably consistent in terms of timing of breeding and clutch size, regardless of growing season characteristics. Such inflexibility could make this species particularly sensitive to environmental changes that affect resource availabilities.

Alaska, Manitoba, Montana, North Dakota, Northwest↗

Lipid catabolism of invertebrate predator indicates widespread wetland ecosystem degradation

Animals frequently undergo periods when they accumulate lipid reserves for subsequent energetically expensive activities, such as migration or breeding. During such periods, daily lipid-reserve dynamics (DLD) of sentinel species can quantify how landscape modifications affect function, health, and resilience of ecosystems. Aythya affinis (Eyton 1838; lesser scaup; diving duck) are macroinvertebrate predators; they migrate through an agriculturally dominated landscape in spring where they select wetlands with the greatest food density to refuel and accumulate lipid reserves for subsequent reproduction. We index DLD by measuring plasma-lipid metabolites of female scaup (n = 459) that were refueling at 75 spring migration stopover areas distributed across the upper Midwest, USA. We also indexed DLD for females (n = 44) refueling on a riverine site (Pool 19) south of our upper Midwest study area. We found that mean DLD estimates were significantly (P<0.05) less than zero in all ecophysiographic regions of the upper Midwest, and the greatest negative value was in the Iowa Prairie Pothole region (-31.6). Mean DLD was 16.8 at Pool 19 and was markedly greater than in any region of the upper Midwest. Our results indicate that females catabolized rather than stored lipid reserves throughout the upper Midwest. Moreover, levels of lipid catabolism are alarming, because scaup use the best quality wetlands available within a given stopover area. Accordingly, these results provide evidence of wetland ecosystem degradation across this large agricultural landscape and document affects that are carried-up through several trophic levels. Interestingly, storing of lipids by scaup at Pool 19 likely reflects similar ecosystem perturbations as observed in the upper Midwest because wetland drainage and agricultural runoff nutrifies the riverine habitat that scaup use at Pool 19. Finally, our results underscore how using this novel technique to monitor DLD, of a carefully selected sentinel species, can index ecosystem health at a landscape scale.

PLoS ONE↗

Fish and land use influence Gammarus lacustris and Hyalella azteca (Amphipoda) densities in large wetlands across the upper Midwest

Gammarus lacustris and Hyalella azteca (hereafter G. lacustris and H. azteca , respectively) are important components of secondary production in wetlands and shallow lakes of the upper Midwest, USA. Within the past 50 years, amphipod densities have decreased while occurrences of fish and intensity of agricultural land use have increased markedly across this landscape. We investigated influences of fish, sedimentation, and submerged aquatic vegetation (SAV) on densities of G. lacustris and H. azteca in semipermanent and permanent wetlands and shallow lakes ( n = 283) throughout seven eco-physiographic regions of Iowa, Minnesota, and North Dakota during 2004–2005. G. lacustris and H. azteca densities were positively correlated with densities of SAV ( P < 0.001 and P < 0.001, respectively). Both species were negatively correlated with densities of large fish (non-Cyprinidae; P = 0.01 and P = 0.013, respectively) and with high densities of fathead minnows ( Pimephales promelas ; P < 0.001 and P = 0.033, respectively). H. azteca densities also were negatively correlated with densities of small fish (e.g., other minnows [Cyprinidae] and sticklebacks [Gasterosteidae]; P = 0.048) and common carp ( Cyprinus spp.; P = 0.022). G. lacustris densities were negatively correlated with high levels of suspended solids (an index for sedimentation; P < 0.001). H. azteca densities were positively correlated with the width of upland-vegetation buffers ( P = 0.004). Our results indicate that sedimentation and fish reduce amphipod densities and may contribute to the current low densities of amphipods in the upper Midwest. Thus, removing/excluding fish, and providing a thick buffer of upland vegetation around wetlands may help restore amphipod densities and wetland and water quality within this landscape.

Iowa, Minnesota, North Dakota↗

Testing competing hypotheses for chronology and intensity of lesser scaup molt during winter and spring migration

We examined chronology and intensity of molt and their relationships to nutrient reserves (lipid and protein) of Lesser Scaup ( Aythya affinisK/i>) to test predictions of two competing hypotheses. The "staggered cost" hypothesis states that contour-feather molt is nutritionally costly and should not occur during nutritionally costly periods of the annual cycle unless adequate nutrients are available. The "breeding plumage" hypothesis states that prealternate molt must be complete prior to nesting, regardless of nutrient availability. Males and females were completing prebasic molt during winter (Louisiana) and had similar molt intensities. Females underwent prealternate molt during spring migration (Illinois and Minnesota) and prebreeding (Manitoba) periods; 53% and 93% of females were in moderate to heavy molt in Minnesota and Manitoba, respectively, despite experiencing other substantial nutritional costs. Intensity of prealternate molt was not correlated with lipid reserves even though females, on average, were nutritionally stressed. Molt intensity was not negatively correlated with protein reserves at any location. Chronology and intensity of prealternate molt varied little and were not temporally staggered from other nutritionally costly events. Prealternate molt did not influence nutrient reserves, and nutrient reserves likely were not the ultimate factor influencing chronology or intensity of prealternate molt of females. We surmise that nutrients required for prealternate molt come from exogenous sources and that the "staggered cost" hypothesis does not explain chronology of prealternate molt in female Lesser Scaup; rather, it appears that molt must be complete prior to nesting, consistent with the "breeding plumage" hypothesis.

The Condor↗

Daily movements of female mallards wintering in Southwestern Louisiana

Understanding daily movements of waterfowl is crucial to management of winter habitats, especially along the Gulf Coast where hunting pressure is high. Radio-telemetry was used to investigate movements of female Mallards ( Anas platyrchychos ) wintering in southwestern Louisiana. Movement distances were analyzed from 2,455 paired locations (diurnal and nocturnal) of 126 Mallards during winters 2004–2005 and 2005–2006 to assess effects of winter, female age, areas closed (Lacassine National Wildlife Refuge [LAC], Cameron Prairie National Wildlife Refuge [CAM], Amoco Pool [AMOCO] or open to hunting [OPEN]), and habitat type, including all interactions. Movement distances from the various land management categories were not consistent by age, date, or by winter. Flight distances from LAC increased with date, whereas those from CAM and OPEN did not vary significantly by date. Female Mallards moved short distances between diurnal and nocturnal sites (ranging from 3.1 to 15.0 km by land management category), suggesting that they are able to meet their daily energy requirements within a smaller area than Northern Pintails ( Anas acuta , hereafter Pintails), and thus minimize transit energy costs.

Louisiana↗

Habitat use by female mallards in the lower Mississippi alluvial valley

Mallard ( Anas platyrhynchos ) populations in the lower Mississippi Alluvial Valley (LMAV), USA, historically averaged 1.6 million and represented the largest concentrations of wintering mallards in North America. Effective management of this wintering population requires current information on use of habitats. Accordingly, we employed radiotelemetry techniques to assess proportional use of habitats by female mallards during winters 2004-2005 and 2005-2006. We divided winters into 4 time periods defined by hunting seasons (FIRST, SPLIT, SECOND, and POST) and recorded diurnal and nocturnal locations. We examined variations in proportional use of habitats and use of areas closed to hunting due to effects of age (immature or ad), winter (2004-2005 or 2005-2006), time period (SECOND or POST), individual female, and all potential interactions of these effects, using locations recorded during the latter 2 time periods. We found that diurnal and nocturnal proportional use of habitats varied inconsistently among time periods and winters. Mean proportional use of forested wetlands ranged from 0.475 to 0.816 and from 0.428 to 0.764 during diurnal and nocturnal sampling periods, respectively. Diurnal proportional use of areas closed to hunting varied inconsistently among time periods and winters. Mean proportional use of areas closed to hunting ranged from 0.183 to 0.423 during diurnal sampling periods. Nocturnal use of areas closed to hunting varied inconsistently among female ages and time periods and among female ages and winters. Mean proportional use of areas closed to hunting ranged from 0.211 to 0.445 during nocturnal sampling periods. Our research suggests that forested wetlands in the LMAV provide important wintering habitats for female mallards; continued restoration and establishment of these habitats should benefit female mallards.

Journal of Wildlife Management↗

Hepatic element concentrations of lesser scaup (Aythya affinis) during spring migration in the upper Midwest

High concentrations of some hepatic elements might be contributing to the decline of the continental lesser scaup ( Aythya affinis ) population. We evaluated hepatic element concentrations of male and female lesser scaup collected from the upper Midwest (Iowa, Minnesota, North Dakota, and South Dakota) during the 2003 and 2004 spring migrations. We measured concentrations of 24 elements in livers of 117 lesser scaup. We found that only selenium concentrations were at levels (>3.0 μg/g wet weight [ww)]) proposed to adversely affect reproduction. Approximately 49% of females ( n = 61) had individual hepatic concentrations >3.0 μg/g ww selenium (Se). Our observed hepatic concentration of Se was similar to that reported in lesser scaup collected from the mid-continental United States but less than Se concentrations reported from the Great Lakes region. We found that the liver cadmium (Cd) concentration for males was significantly higher than that for females. Gender differences in hepatic Cd concentrations have not been previously reported for lesser scaup, but Cd is known to have negative impacts on male reproduction. Our results indicate that lesser scaup migrating through the upper Midwest in spring have elevated Se levels and that males carry a significantly greater Cd burden than females. Moreover, elemental concentrations might be high enough to affect reproduction in both male and female lesser scaup, but controlled laboratory studies are needed to adequately assess the effects of Se and Cd on lesser scaup reproduction.

Iowa, Minnesota, North Dakota, South Dakota↗

Movement distances and habitat switching by female Mallards wintering in the Lower Mississippi Alluvial Valley

Mallard (Anus platyrhynchos) populations in the Lower Mississippi Alluvial Valley (hereafter, LMAV) comprise the largest concentration of wintering Mallards in North America. Radiotelemetry techniques were employed to assess movements by female Mallards during winters of 2004-2005 and 2005-2006; 467 paired (diurnal and nocturnal) observations on 80 radiomarked females were attained to assess effects of date, female age, hunting regime and habitat type on distances moved and potential habitat switching. Distance moved increased with date for females diurnally located in RICE but decreased for females in FOREST. Median movement distance was 1.5 km, suggesting suitable habitats for all activities were in close proximity. Habitat switching varied among diurnal habitat types used by females. Females diurnally located in FOREST and REFOR switched habitats in <= 22% of paired observations, whereas females diurnally located in RICE switched habitats in 55% of paired observations. The decreased movement distances and less frequent habitat switching by females using forested habitats should reduce their energy expenditure and exposure to predation risk. The results document the importance of forested wet-lands to Mallards wintering in the LMAV and provide support for programs that protect and restore such habitats. Received 18 November 2009, accepted 16 January 2010.

Lousiana↗

Intraspecific variation in nutrient reserve use during clutch formation by Lesser Scaup

We studied nutrient reserve dynamics of female Lesser Scaup ( Aythya affinis ) to identify sources of intraspecific variation in strategies of nutrient acquisition for meeting the high nutritional and energetic costs of egg formation. We collected data from interior Alaska and combined these with data for Lesser Scaup from midcontinent breeding areas ( Afton and Ankney 1991 ), allowing a rangewide analysis for the species. We found little evidence that nutrient reserve use differed between Alaskan and midcontinent Lesser Scaup, except that subarctic birds used a small amount of protein reserves when forming eggs, whereas midcontinent birds did not. Mineral reserves contributed relatively little to the clutch, but endogenous lipid accounted for approximately two-thirds of the lipid in the clutch. Levels of endogenous lipid and protein at initiation of clutch formation declined with date of initiation. Also, absolute amounts of lipid and protein reserves used declined through the season, corresponding to smaller clutch sizes. Our data are consistent with a seasonally variable threshold of lipid reserves for initiation of clutch formation and considerable reliance on lipid reserves, suggestive of lipid control of productivity via effects on clutch size and initiation dates. However, our data cannot refute the hypothesis that clutch size or initiation dates are set by other factors that in turn dictate the amount of lipid reserves that are stored and used. Despite uncertainty regarding the role of nutrient limitations on productivity, maintenance of adequate food resources on winter, migration, and breeding areas should be a management concern, given the high costs of clutch formation by Lesser Scaup, evidence of recent population declines, and potential links between nutrition and productivity.

Alaska↗

Effects of winter marsh burning on abundance and nesting activity of Louisiana seaside sparrows in the Gulf Coast Chenier Plain

Louisiana Seaside Sparrows ( Ammodramus maritimus fisheri ) breed and winter exclusively in brackish and saline marshes along the northern Gulf of Mexico. Many Gulf Coast marshes, particularly in the Chenier Plain of southwestern Louisiana and southeastern Texas, are burned intentionally in fall or winter as part of waterfowl management programs. Fire reportedly has negatively affected two Seaside Sparrow subspecies ( A. m. nigrescens and A. m. mirabilis ) in Florida, but there is no published information regarding effects of fire on A. m. fisheri. We compared abundance of territorial male Louisiana Seaside Sparrows, number of nesting activity indicators, and vegetation structure in paired burned and unburned plots in Chenier Plain marshes in southwestern Louisiana during the 1996 breeding season (April–July) before experimental winter burns (January 1997) and again during two breeding seasons post-burn (1997–1998). We found that abundance of male sparrows decreased in burned plots during the first breeding season post-burn, but was higher than that of unburned plots during the second breeding season post-burn. Indicators of nesting activity showed a similar but non-significant pattern in response to burning. Sparrow abundance and nesting activity seemingly are linked to dead vegetation cover, which was lower in burned plots during the first breeding season post-burn, but did not differ from that in unburned plots during the second breeding season post-burn. We recommend that marsh management plans in the Gulf Coast Chenier Plain integrate waterfowl and Seaside Sparrow management by maintaining a mosaic of burned and unburned marshes and allowing vegetation to recover for at least two growing seasons before re-burning a marsh.

Louisiana↗

Survival of female northern pintails wintering in southwestern Louisiana

The North American breeding population of northern pintails ( Anas acuta ) has reached previously unprecedented low numbers 4 times since 1983. Because pintails show high fidelity to wintering areas, regional survival estimates and identification of factors influencing survival are needed to guide management of wintering pintails. We used radiotelemetry to estimate survival rates of female pintails wintering in southwestern Louisiana. We tested for variation in survival and hunting mortality rates in relation to age (immature or adult), winter (1990-91, 1991-92, 1992-93), time period (prehunting season, first hunting season, time between split hunting seasons, second hunting season, posthunting season), body condition (body mass when released, adjusted for body size), and region (southwestern Louisiana or elsewhere on the Texas-Louisiana Gulf Coast or Mississippi Alluvial Valley). Within southwestern Louisiana, the 147-day (5 Oct-28 Feb) survival rate of adults (0.714 ± 0.045; x̅ ± SE) was greater ( P = 0.02) than that of immatures (0.550 ± 0.068), primarily because immatures had higher hunting mortality. Female survival was lower ( P s < 0.01) during hunting than during nonhunting seasons but did not differ between first and second hunting seasons ( P = 0.58) or among non-hunting seasons ( P s >0.25). Survival did not differ in relation to winter or condition ( P s > 0.12). Hunting mortality did not differ ( P s > 0.13) in relation to winter, condition, or region, but hunting mortality of immatures (0.287 ± 0.046) was greater ( P < 0.001) than that of adults (0.130 ± 0.025). Despite conservative hunting regulations (30-day seasons and 1 pintail daily), hunting mortality rates of pintails in southwestern Louisiana were high. Pintails in southwestern Louisiana rely on a small number of key refuges for protection but feed almost exclusively in privately owned agricultural fields. If increased winter survival of female pintails in southwestern Louisiana is desired, we recommend that managers increase food available.

Louisiana↗

Use of mini-refuges by female northern pintails wintering in southwestern Louisiana

The Gulf Coast Joint Venture of the North American Waterfowl Management Plan began contracting private agricultural lands (hereafter mini-refuges) in 1988 to expand existing sanctuaries for northern pintails ( Anas acuta ) in southwestern Louisiana. Previous research suggested that mini-refuges may prove more attractive to pintails than permanent, open-water pools (pools) on refuges because mini-refuges provide sanctuary and food during the day, whereas pools generally provide only sanctuary (Rave and Cordes 1993). We used radiotelemetry to compare diel use of mini-refuges and pools (Lacassine Pool and Amoco Pool) by female pintails in southwestern Louisiana during winters of 1991-1992 and 1992-1993. We examined variation in use of these areas in relation to female age (immature or adult), time period (prehunting season, first hunting season, time between split hunting seasons, second hunting season, and posthunting season), and winter (1991-1992 and 1992-1993). Diurnal use of min-refuges and pools differed among time periods, but differences were not consistent between winters. Mini-refuges accounted for <2% of diurnal use by pintails in 7 of 10 time-period and winter comparisons. Diurnal use of mini-refuges was lower than that of Lacassine Pool in 8 of 10 time-period and winter comparisons. Diurnal use of mini-refuges was lower than tha of Amoco pool during first hunting season in 1992-1993, but use of these areas did not differ within other time periods and winters. Nocturnal use of mini-refuges and pools did not differ in relation to female age, time period, winter, or individual bird. Nocturnal use of mini-refuges did not differ from that of Lacassine Pool. In contrast to predictions and findings by Rave and Cordes (1993), we found that: (1) female pintails did not use mini-refuges more than pools, and (2) female pintails used mini-refuges at night. We believe that use of mini-refuges by pintails could be increased if mini-refuges were (1) located in areas of traditionally high pintail use, (2) increased in size, (3) flooded immediately prior to hunting season, and (4) cleared of dense vegetation by rolling, disking, or burning.

Louisiana↗