Wetland Use and Feeding by Lesser Scaup During Spring Migration Across the Upper Midwest, Usa
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WETLANDS, Vol. 29, No. 2, June 2009, pp. 704–712 ’ 2009, The Society of Wetland Scientists WETLAND USE AND FEEDING BY LESSER SCAUP DURING SPRING MIGRATION ACROSS THE UPPER MIDWEST, USA Michael J. Anteau1,2 and Alan D. Afton3 1School of Renewable Natural Resources, Louisiana State University Baton Rouge, Louisiana, USA 70803 2Present address: U.S. Geological Survey, Northern Prairie Wildlife Research Center, Jamestown North Dakota, USA 58401, USA E-mail: [email protected] 3U.S. Geological Survey Louisiana Cooperative Fish and Wildlife Research Unit, Louisiana State University Baton Rouge, Louisiana, USA 70803 Abstract: Low food availability and forage quality and concomitant decreased lipid reserves of lesser scaup (Aythya affinis; hereafter scaup) during spring migration in the upper Midwest may partially explain reductions in the continental population of scaup. In springs 2004–2005, we examined wetland use and feeding activity of scaup on 356 randomly-selected wetlands within 6 regions in Iowa, Minnesota, and North Dakota. We examined wetland characteristics that favor high scaup use in 286 of these wetlands. We found that probabilities of wetland use and feeding by scaup increased with turbidity up to 45 and 30 NTU, respectively, but then declined at higher turbidity levels. Wetland use was positively correlated with size of open-water zone and amphipod densities, but was not correlated with chironomid densities. Feeding increased with amphipod density up to 26 m23 and then declined at higher amphipod densities; scaup seemingly forage most efficiently at amphipod densities above 26 m23. Wetland use was higher in North Dakota than in southern Minnesota and Iowa. Our results indicate that effective wetland restoration efforts to benefit scaup require maintaining abundant populations of amphipods (generally near 26 m23 landscape geometric mean) in wetlands with large (. 500 m diameter) open-water zones throughout the upper Midwest, but especially within Iowa and southern Minnesota. Key Words: behavioral cue, patch occupancy, optimal foraging, resource selection, staging, stopover areas, waterfowl INTRODUCTION 2006). Accordingly, restoration, and management of wetland habitats used by scaup during spring The continental scaup population (lesser [Aythya migration is a major goal of recent conservation affinis] and greater scaup [A. marila] combined) has efforts in the upper Midwest (Ducks Unlimited declined markedly since 1978 (Austin et al. 1998, 2004; S. Stephens, Ducks Unlimited Inc., personal Afton and Anderson 2001, Wilkins et al. 2007). The communication; G. Zenner, Iowa Department of largest declines are associated with a major segment Natural Resources, personal communication). of the lesser scaup population that winters in states Three important questions must be addressed to bordering the Gulf of Mexico and migrates along target types of wetlands that scaup use and regions the Mississippi River valley and through the upper of greatest conservation need. First, what are the Midwest (Afton and Anderson 2001). preferred foods of scaup during spring migration? Food availability and forage quality for female Amphipods and chironomids are principal foods lesser scaup (hereafter scaup) currently are low and consumed by scaup during spring migration (Afton apparently decreased from that historically during et al. 1991, Anteau and Afton 2006, 2008b). spring migration in the upper Midwest (Anteau and Amphipod densities are positively correlated with Afton 2006, 2008a, 2008b, Strand et al. 2008). scaup use on breeding areas (Lindeman and Clark Moreover, scaup currently have low lipid-reserve 1999, Fast et al. 2004), but it is unclear if this is a levels and catabolize lipid reserves during stopover direct link or only because amphipod densities might periods throughout the upper Midwest (Anteau be correlated with densities of other scaup foods. 2006, Anteau and Afton 2004, Badzinski and Petrie Therefore, we simultaneously examined influences 704 Anteau & Afton, WETLAND USE BY LESSER SCAUP 705 of amphipod and chironomid densities on wetland converged basin (Johnson and Higgins 1997) or use and feeding activity of scaup to determine the comparable National Wetland Inventory data (see relative importance of these food items to migrating Anteau 2006). In Iowa, we broadened the constraint to females. Second, what characteristics of wetlands include townships with 200 ha of either semiperma- might scaup use to select foraging wetlands? The size nent or permanent wetlands. Constraining township and class of wetlands used by migrating scaup during selection helped ensure that there were enough migration must be identified so that conservation wetlands to sample within each township. efforts can be efficiently targeted. Moreover, infor- We allocated numbers of sampling clusters among mation on the importance of vegetation structure or the 6 regions (3 to 6 clusters per region; Table 1) water clarity of wetlands as visual cues used by scaup based on region size and number of candidate to select foraging wetlands would be useful for townships available (Table 1, Figure 1). Each region management. Consequently, we examined how var- was divided into sub-regions based on latitude, such ious wetland characteristics (e.g., turbidity, depth, that there was one sampling cluster per sub-region size of open-water zone, water regime, amount and each year (except in North Dakota Glaciated Plains species of emergent vegetation, and densities of two [NDGP]). In NDGP, we assigned two sampling principle foods) influenced wetland use and feeding clusters for each sub-region because of the larger by scaup. Third, what areas or regions in the upper width of this region (east to west) relative to other Midwest are in most need of conservation efforts; are regions (Figure 1). there regions that scaup use heavily, but where they Each sampling cluster was comprised of three are unable to acquire nutrient reserves? Accordingly, adjacent 36-square-mile townships (27,972 ha total). we examined inter-regional variation in wetland use The centroids of the second and third townships and feeding by scaup in relation to nutritional and selected were constrained within 50 km of the dietary data from these same areas (Anteau 2006, centroid of the first randomly selected township. Anteau and Afton 2008b). We randomly selected 3 semipermanent or perma- nent wetlands (each . 4 ha) in each township for wetland sampling. We randomly selected new METHODS townships and wetlands annually within each sub- Study Area region from the list of candidate townships to maximize representation of spatial variability in We sampled wetlands and observed scaup behav- wetland characteristics. ior within the Prairie Pothole Regions of Iowa, In springs 2004 and 2005, we observed use and Minnesota, and North Dakota. This area encompasses behavior of scaup on 356 wetlands and conducted the most important spring migration stopover areas complete wetland sampling on a random subset of for scaup in the upper Midwest, based on observations 286 wetlands (Table 1). In 2005, we supplemented of wildlife biologists and managers, band-recovery our sample in Minnesota with 9 additional high data, sightings of color-marked scaup, aerial surveys in scaup use wetlands because preliminary analyses spring, and scaup marked with satellite transmitters indicated that random sampling alone did not (Anteau and Afton 2008b, Afton 2008). provide an optimal balance of high-use wetlands We stratified the three-state area into six eco- for examining factors that influenced wetland use physiographic regions (hereafter regions) based on and feeding by scaup. watershed and groundwater hydrology, geology, and Sampling coincided with estimated middle of the plant communities (Figure 1; Kantrud et al. 1989; scaup migration period, when relatively large num- Minnesota Department of Natural Resources, unpub- bers of migrating scaup were present on our study lished data). The Minnesota Glaciated Plains (MNGP) area (see Anteau 2006). We started sampling in the and Minnesota Morainal (MOR) eco-physiographic southern portion of the study area and worked north. regions included areas outside the traditional Prairie This approach ensured an equal probability of scaup Pothole Region (Figure 1); however, we included these being present within each sub-region, and that regions so that results would be relevant to state- sampling occurred at similar times relative to spring specific management and conservation plans. phenology. Sampling was conducted during 3 April We used a constrained-random clustered sampling to16May2004and30Marchto3May2005. approach to select wetlands; this approach minimized travel time between wetlands. We first estimated Determination of Wetland Use and Feeding numbers of townships within each region that had at least 200 ha of semipermanent wetlands (candidate We assessed wetland use and feeding by scaup on townships for random selection; Table 1), based on each selected wetland with four visits (twice a day 706 WETLANDS, Volume 29, No. 2, 2009 Figure 1. Study area depicting 6 eco-physiographic regions for observations of lesser scaup wetland use and feeding during spring migration (2004 and 2005) in the upper Midwest. Areas in white were not sampled. Table 1. Numbers of sampling clusters (C), randomly selected wetlands observed for lesser scaup use and feeding (NRandom), wetlands with compete wetland surveys (NComplete) by region in springs 2004 and 2005, and candidate townships (T) available for random selection