J OURNAL OF C RUSTACEAN B IOLOGY, 34(1), 54-60, 2014 A COMPARISON OF TWO GEARS FOR QUANTIFYING ABUNDANCE OF LOTIC-DWELLING CRAYFISH Kristi Williams 1,∗ , Shannon K. Brewer 2,∗∗ , and Mark R. Ellersieck 3 1 302 Anheuser Busch Natural Resources Building, Department of Fisheries and Wildlife Sciences, University of Missouri, Columbia, MO 65201, USA 2 U.S. Geological Survey, Oklahoma Cooperative Fish and Wildlife Research Unit, 007 Agriculture Hall, Oklahoma State University, Stillwater, OK 74078, USA 3 Agriculture Experiment Station, Department of Statistics, University of Missouri, Columbia, MO 65211, USA ABSTRACT Crayfish (saddlebacked crayfish, Orconectes medius) catch was compared using a kick seine applied two different ways with a 1-m2 quadrat sampler (with known efficiency and bias in riffles) from three small streams in the Missouri Ozarks. Triplicate samples (one of each technique) were taken from two creeks and one headwater stream (n = 69 sites) over a two-year period. General linear mixed models showed the number of crayfish collected using the quadrat sampler was greater than the number collected using either of the two seine techniques. However, there was no significant interaction with gear suggesting year, stream size, and channel unit type did not relate to different catches of crayfish by gear type. Variation in catch among gears was similar, as was the proportion of young-of-year individuals across samples taken with different gears or techniques. Negative binomial linear regression provided the appropriate relation between the gears which allows correction factors to be applied, if necessary, to relate catches by the kick seine to those of the quadrat sampler. The kick seine appears to be a reasonable substitute to the quadrat sampler in these shallow streams, with the advantage of ease of use and shorter time required per sample. K EY W ORDS: capture efficiency, Orconectes medius, quadrat sampler, saddlebacked crayfish, seine DOI: 10.1163/1937240X-00002212 I NTRODUCTION The increasing interest in researching the ecology and monitoring the status of freshwater crayfishes is likely due to an evolving appreciation of their key ecological role in many lotic systems (Momot, 1995), their vulnerability to extirpation from habitat destruction (Taylor et al., 2007), and the damage invasive crayfish species can cause (Lodge et al., 2000). Studies involving crayfish ecology often require quantitative estimates of population numbers, and although numerous methods for sampling populations are commonly used, most have serious biases and unknown efficiencies. It follows that confidence in results and conclusions requires confidence in the sampling approach. Various active and passive sampling methods to study crayfish have been employed for several decades. Methods include baited traps, hand netting or hand collecting, visual assessments using SCUBA, snorkeling or bankside above stream observation, electrofishing, and enclosure nets and seines (see Haag et al., 2013 for a complete assessment). The usefulness of results from these methods is often questionable because of suspected, but unquantified biases, very low efficiencies that increase variation, and utility in only special situations. ∗ Present An enclosed quadrat sampler has been extensively used, and its characteristics examined, for collecting both crayfish (Rabeni, 1985; DiStefano et al., 2003) and benthic fishes (Fisher, 1987; Peterson and Rabeni, 2001). This sampler has been evaluated for its sampling variance and sampling precision for crayfish under a variety of conditions found in Ozark streams (DiStefano et al., 2003). Further work by Larson et al. (2008) determined actual efficiencies (% of individuals present that are collected) in riffle channel units of an Ozark stream. Because of these efforts, we believe this technique to be the best way currently available to estimate crayfish densities and abundances in small to medium-sized streams. Whereas the quadrat sampler has proven effective in many situations, it is not without problems. The quadrat is heavy, cumbersome, and labor intensive. Most importantly it is time consuming. Use in Ozark streams has shown two operators can reasonably collect 16 quadrat samples per day (two samples per h) depending on the number of crayfish collected for processing and if additional abiotic measurements are taken at each site (DiStefano, 2000). This rate of sampling discourages its use in situations requiring substantial numbers of samples to be taken over a short period of time. address; Arkansas Department of Environmental Quality, 5201 Northshore Drive, North Little Rock, AR 72118, USA. author; e-mail: [email protected] ∗∗ Corresponding © The Crustacean Society, 2014. Published by Brill NV, Leiden DOI:10.1163/1937240X-00002212 55 WILLIAMS ET AL.: GEARS FOR LOTIC-DWELLING CRAYFISH STUDIES Recently, a small kick seine has gained popularity in the Ozark region of Missouri and Arkansas (Flinders and Magoulick, 2003), and in other areas of the U.S. (Mather and Stein, 1993) for determining crayfish population characteristics. The kick seine is lightweight, easily constructed and used, and inexpensive. Most importantly, we have found sampling time is generally 10-15 min per sample (approximately 4 samples per h) depending on habitat conditions and the number of crayfish that need to be processed. This means use of the kick seine could result in 32 samples taken in one day and that estimate generally agrees with the findings of others (Allert et al., 2008). However, no analyses of the characteristics of the kick seine for collecting crayfish have been conducted. The objective of this study was to test the performance of the kick seine as compared to the quadrat sampler whose biases and efficiencies are known under some common stream situations. We assumed that the efficiency of the quadrat sampler would be similar in runs and pools of small, shallow streams but realize that the ability to seal the sampler to the bottom (likely better in these habitats than riffles) will affect efficiency. If the kick seine captures comparable numbers of crayfish as the quadrat sampler under varying conditions, or if measurable biases exist and can be accounted for, the seine may be a less labor intensive substitute for the quadrat sampler. M ATERIALS AND M ETHODS Study Area Three streams were chosen for crayfish sampling, each located in the Ozark Highlands, Missouri, USA. The Ozark Highlands ecoregion consists largely of karst topography with relatively clear streams that carry low suspended sediment loads, have well-developed riffle-pool complexes, and have significant spring influences (Nigh and Schroeder, 2002). One stream was a small (mean channel width 2.3 m), shallow (mean depth 0.12 m) unnamed headwater stream (1st order, Strahler, 1957) with an average velocity of 0.20 m · s−1 . The other two streams sampled were Lost and Courtois creeks, both fourth-order streams. Courtois Creek had similar available velocity conditions (mean velocity 0.20 m · s−1 ) but as expected, was wider (mean width 13.2 m) and deeper (mean depth 0.33 m) than the headwater stream. Depth and velocity profiles were not taken at Lost Creek but the channel width was similar to Courtois Creek (mean width 11.2 m). Particle sizes indicate dominant and subdominant substrate types were primarily pebble and cobble (modified Wentworth scale, Gordon et al., 1993) in all of the streams sampled (Williams, 2006). The maximum depth sampled was 0.40 m in the headwater stream and 0.55 m in Courtois Creek. In each stream, riffle, run, and pool channel units (discrete morphological habitat features formed by high flows) were delineated based on gradient, velocity, and depth. Riffles had higher gradients than the surrounding areas, increased velocities, and shallow depths. Runs were transition areas with intermediate depths and velocities whereas, pools were depositional areas with limited velocity and were generally deeper than surrounding habitat (Rabeni and Jacobson, 1993 for a review of channel unit classification in Ozark streams). Sampling Methods Sample sites were chosen in each channel unit to minimize variation due to microhabitat differences among samples. Criteria for sample sites were: a channel unit area that would allow three 1-m2 samples to be taken while leaving at least a 1 m2 undisturbed between each sample, a similar dominant substrate type, and little variation in depth and velocity. Samples were taken in triplicate, one quadrat and two seines, from the same channel units with the underlying assumption being each sample was representative of the crayfish densities within that channel unit. During the first year of sampling, the order of gear used was not randomized and the seine samples were always taken first. During the second year of sampling, gear order was randomized at each site in case more crayfish were sampled via the first gear. Samples were taken beginning downstream and working in an upstream direction. Crayfish were sampled in autumn 2002 and summer 2003 from riffles, runs, and pools of three streams using two gear types (quadrat sampler and seine) and one variation of technique (seine variation). The 1-m2 quadrat sampler has a known efficiency for sampling lotic crayfish in riffle habitats (Larson et al., 2008). The sampler is 0.5 m high and is covered on each side by 3-mm netting and has a 1.2 m bag seine attached to the downstream end (see diagram in Larson et al., 2008). Sampling procedures for using the quadrat sampler were described by DiStefano et al. (2003). Briefly, the sampler was placed in the substrate and the side nets were sealed to the streambed to prevent crayfish escape. The substrate within the sampler was disturbed for approximately 3-5 minutes, and large substrate was removed from the interior portion of the sampler to a depth of 15 cm. Water and debris were swept into the downstream seine and examined for crayfish at the end of the sample. The kick seine was compared to the quadrat sampler using two different techniques from qualified field personnel. Both techniques required two people for operation and both seines had 3-mm netting and a 22.7-kg lead line attached to the bottom. With the first technique (hereafter referred to as seine 1), one individual held the seine handles about 1 m apart while the second person secured the lead line to the streambed using cobble or larger substrates. The second person would then use a side-to-side kicking action over an estimated 1-m2 area upstream of the seine. The area was disturbed once, large substrates were removed, and then a second pass was made over the 1-m2 area. The second technique (hereafter referred to as seine 2) used a seine of the exact same dimensions as seine 1 but used slight variations in the technique. The seine was held with the top of the handles together and near the water’s surface to create more of a bag in the back of the seine and less effort was spent creating a seal with the leadline and the streambed. The area was disturbed as with the other technique, but instead used a front-to-back kicking motion and swept water into the seine while kicking, rather than making two passes. Personnel operating each of the techniques were alternated at each sampling event to negate any influence related to sampler and operator bias. Regardless of technique, adult and juvenile crayfish were enumerated on the stream bank following each sample, held in buckets and returned after to the stream following the completion of sampling in that channel unit. Lengthfrequency histograms were used to delineate a carapace length threshold for young-of-year versus adult crayfish (Rabeni, 1992). Analyses Analyses were restricted to captures of saddlebacked crayfish Orconectes medius (Faxon, 1884) because the species represented 95% and 97% of the total number of individuals captured in year 1 and year 2, respectively. Prior to analyses, some variables were combined into a single metric so multiple models could be created to account for missing data from one sample year (pools were not sampled from one stream during one year) and to include the influence of stream size even though only one headwater stream was sampled. Stream sampled and sampling year were combined to form a single variable called ‘environment.’ The first general linear model excluded pool channel units because they were not sampled from one stream during one sample year. By omitting these data, we could analyze for possible influences related to sample year and stream size. For the second model, a subset of data from the ‘environment’ variable (2002 data from Courtois Creek) was omitted because pools were not sampled during that period. Removing these data and then creating a second model allowed us to assess the influence of all three channel units sampled (riffle, run, and pool). We were primarily interested in the interactive effects to determine if stream size or channel unit sampled influenced densities of crayfish captured by each gear or technique. In addition, we were interested in examining possible interactive effects between environment and gear to assess whether lack of randomization of gear and technique during the first year of sampling had any relationship to the observed results. Two general linear mixed models were created to assess the influence of channel unit, stream size, year sampled, and gear on crayfish catch. Our primary interest was assessing the significance of gear and any interactions with gear (channel unit, stream size, year sampled). The results of these models guided the second portion of the analyses (regressions) by identifying environmental conditions that would interact with gear thereby creating a situation where regressions would need to be created for varying environmental conditions, e.g., in headwater streams versus creeks. Data were analyzed as two split plots in space using the ‘Proc 56 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 1, 2014 Glimmix’ procedure in SAS (SAS, 2000) to account for possible spatial autocorrelation. Both models used a log link and a Poisson distribution. Plots of each subset of data indicated either a Poisson or negative binomial distribution would be appropriate but Poisson seemed to fit these data subsets slightly better. One was added to all of the adult crayfish densities because the log of zero was missing. The first model contained the main effects of environment (year and stream) and the subplot contained the effects of channel unit (riffles and runs) and gear (two seining methods and the quadrat sampler) and all possible combinations with the main effect environment. The second model contained the main effects of environment, but this time with year by stream combinations by omitting 2002 data from sampled creeks because pools were not sampled from creeks during that year. The subplot contained the effects of channel unit (riffle, run, and pool), and gear (two seining methods and the quadrat sampler) and all possible combinations with the main effect environment. For both models, we used the Tukey-Kramer honestly significant difference post-hoc test to determine where significant differences occurred given the overall model was significant. Two negative binomial linear regressions were created to examine the relationship between the number of crayfish collected with the quadrat sampler compared to the number of crayfish collected with each seining technique (seine 1 and seine 2) to allow for adjustments to be made to seine samples taken under similar environmental conditions (small headwater streams and creeks). Because there were no significant interactions between gear and other fixed effects in the general linear mixed models, data were combined to include all of the quadrat and seine samples into one dataset. Two samples were omitted from the analyses because no quadrat sample was taken with the two seine samples. We examined the data for outliers and checked the observations to ensure data were entered correctly. The dataset included 69 observations for each gear (quadrat, seine 1 and seine 2) because the gears were combined at each site (one of each seine and one quadrat sample were taken at each of 69 sites, n = 39 in 2002 and n = 30 in 2003). Descriptive statistics were calculated for each gear to determine if over-dispersion was indicated by these data (variance > mean; Osgood, 2000). In addition, a dispersion parameter was estimated for each model (Proc Genmod; SAS, 2000) to assess model fit. If dispersion is >0 (by examining 95% confidence intervals around the dispersion estimate), then a negative binomial distribution is appropriate rather than a Poisson distribution. The negative binomial distribution is a generalization of the Poisson distribution but includes an additional parameter allowing the variance to exceed the mean (Osgood, 2000). Given fit was appropriate, negative binomial regressions were fit to the data using a negative binomial distribution and log link (Proc Genmod; SAS, 2000). The count of crayfish collected using the quadrat sampler was the dependent variable in each model since some estimate of efficiency is known for the quadrat sampler (i.e., riffles). Models were considered significant at α 0.05. We assumed that correcting data from one gear to another would be most relevant if the precision among the gears or techniques (seine 1 and seine 2) were comparable. We calculated the coefficients of variation in crayfish catch for each gear and technique so comparisons of precisions could be made given the samples represent different methods. Adjustments for young-of-year crayfish could not be made because the efficiency of the quadrat sampler in collecting juveniles is unknown. However, we compared the ratio of young-of-year to total crayfish collected for individual streams in each of two years to determine if size selectivity was evident between gears or techniques. If differences exist, the choice of gear might then relate more to project objective rather than simply ease of collection. R ESULTS Influence of Year and Habitat In the first model, the number of adult crayfish sampled was significantly related to the main effects of environment (F4,156 = 13.69, P < 0.01), channel unit (F1,156 = 7.64, P < 0.01) and gear (F2,156 = 6.92, P < 0.01). In addition, the expected interactive effect of environment ∗ channel unit was also significant indicating the number of crayfish sampled in a particular stream and year depended on the channel unit sampled (riffle or run) (F4,156 = 7.11, P < 0.01). Densities of crayfish collected using the quadrat Fig. 1. Mean crayfish densities (±95% confidence limits) associated with two gears (quadrat and seine) and two seining techniques (seine 1 and seine 2). The top panel represents data included in Model 1 and the bottom panel represents data included in Model 2. sampler were greater than when using either of the two seining techniques (seine 1: t156 = −3.19, P < 0.01; seine 2: t156 = −2.96, P < 0.01; Fig. 1). Generally, the quadrat sampler collected approximately 20% more crayfish than either of the two seining techniques. No significant interactions were detected with gear indicating stream size (headwater or creek) and year did not alter the number of crayfish collected. These results suggest lack of randomizing gear type in year one had no significant effect on the outcome and there was no influence on catch by gear between the stream sizes we sampled. In the second model created, the number of adult crayfish sampled was significantly related to the main effects of environment (F3,113 = 30.97, P < 0.01) and gear (F2,113 = 6.92, P < 0.01). As we expected and similar to model 1, the interactive effect of environment ∗ channel unit (F6,113 = 7.11, P < 0.01) was also significant indicating the number of crayfish sampled in a particular stream and year depended on the channel unit sampled (riffle, run, or pool). Again, the quadrat sampler collected more crayfish than either of the two seining techniques (seine 1: t113 = −2.44, P = 0.04; 57 WILLIAMS ET AL.: GEARS FOR LOTIC-DWELLING CRAYFISH STUDIES Table 1. Maximum likelihood parameter estimates for two negative binomial distribution regression models and untransformed data for each seining technique (no significance is assumed with the models created using the actual data due to distributional assumptions). Parameter df Estimate SE Wald 95% confidence limits χ2 P > χ2 <0.01 <0.01 Model 1: negative binomial model (seine 1) Intercept 1 1.58 Seine 1 1 0.07 Dispersion 1 0.50 0.15 0.02 0.11 1.29 0.03 0.32 1.87 0.10 0.77 116.10 15.60 Model 1: actual data (seine 1) Intercept 1 Seine 1 1 4.64 0.63 1.12 0.12 2.45 0.40 6.83 0.86 17.24 28.49 Model 2: negative binomial model (seine 2) Intercept 1 1.56 Seine 2 1 0.07 Dispersion 1 0.45 0.14 0.02 0.11 1.29 0.04 0.29 1.83 0.11 0.71 129.78 20.25 Model 2: actual data (seine 2) Intercept 1 Seine 2 1 1.03 0.11 2.21 0.51 6.27 0.95 16.79 41.04 4.24 0.73 seine 2: t113 = −3.47, P < 0.01; Fig. 1). As with model 1, no interactive effects occurred with gear suggesting channel unit (including sampled pools) did not relate to different catches of crayfish by gear. Correction Factors The variance associated with number of crayfish collected with each gear type was greater than the means suggesting over-dispersion was present and that a negative binomial model would be appropriate. The 95% confidence intervals of the dispersion parameter for each model were greater than zero providing further support for use of a negative binomial distribution (Table 1). The first model indicated a significant linear relationship between the number of crayfish collected via the quadrat sampler compared to the number of crayfish collected via seine method 1 (Table 1, Fig. 2). The intercept of the first model represents the negative binomial regression <0.01 <0.01 estimate when the count of crayfish collected using seine 1 is zero. In this case, when the seine count = 0, the log of the expected quadrat count = 1.58 (Table 1). Seine method 1 was significant to the model and the maximum likelihood parameter estimate indicates that if we increase our seine 1 sample by one crayfish, the difference in the logs of expected counts would increase by 0.06, while holding the intercept constant. The actual data values (non-transformed data with a normal distribution) are reported in Table 1 and should approximate the mean (but should not be used to interpret significance). As we would expect given that there were no differences between crayfish catches between the two seine techniques, the second model results were similar to the first model. The second model indicated a significant linear relationship between crayfish collected via the quadrat sampler and crayfish collected using seine 2 (Table 1). The parameter estimates for the intercept and seine 2 (1.56 and 0.07, respectively) are very similar to those presented above for seine 1. Precision A comparison of variation among the three techniques indicates a greater sampling precision (as measured by the coefficient of variation) for the quadrat compared to the seines (Table 2), but the 9% or 13% difference between gears is likely not biologically meaningful. Size Selectivity Graphs comparing gears and techniques (Fig. 3) reveal no consistent differences in the catch of different ages, i.e., sizes, of crayfish. Although year 2 results show slightly Table 2. A comparison of sampling precision of collections of adult crayfish between the seine techniques and quadrat sampler. Fig. 2. Scatterplot of crayfish densities obtained using the quadrat sampler versus densities estimated using the seines (one of two techniques). Black circles indicate approximate densities using seine one whereas open circles represent approximate densities obtained using seine two. Data presented are the 69 observations presented in the regression analyses. Method n Mean SD CV Seine 1 Seine 2 Quadrat 69 69 69 7.0 6.6 9.0 6.4 6.2 9.0 91% 95% 82% 58 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 1, 2014 Fig. 3. Percent of young-of-year crayfish that occurred in catch by sampling method from two streams that were sampled in 2002 and 2003. Gears and techniques are represented by: S1: seine technique one, Q: quadrat sampler, and S2: seine technique two. higher ratios of young-of-year, year 1 results are very similar with a range of results of only 2% in the creek and 5% in the headwater stream. D ISCUSSION Accurate estimates of crayfish population densities and abundances are required to increase understanding and management of this important faunal group. The ideal crayfish sampler would be efficient, have quantifiable biases, and be time resourceful. We believe the 1-m2 quadrat sampler to be the only stream crayfish sampler with known efficiencies under some conditions (riffles) and with minimal size selection (see catch curve in Rabeni, 1992), except for very small (young) individuals that may pass through the net mesh, and very large individuals that occupy specialized habitats. Large crayfish often use deeper pools with large boulders and limestone bluffs making their capture difficult (DiStefano et al., 2003). However, the larger individuals in these streams contribute little to overall secondary production (Brewer et al., 2009). Although the quadrat sampler can be used to estimate densities over a known area, it is relatively time consuming and often logistically cumbersome to use. Field experience (DiStefano et al., 2003) has shown a 2-person team can complete about two quadrat samples per h. Larson et al. (2008) indicate with power analysis that even at a power of 0.8, approximately 20 quadrat samples would be necessary for confidence intervals (half width) to be within 40% of the mean. A day and a half effort to produce a meaningful density estimate for a crayfish population in a reach of stream may not be realistic in many situations, depending on the objective. This is especially true when gear need to be carried over long distances to sample sites as the quadrat sampler is heavy. A two-person team can collect an average of >30 kickseine samples in a day, although this value will be reduced under some habitat conditions or if large numbers of crayfish are collected. Although the speed and ease of taking a sample may be important in some situations, investigators must be sure not to have a significant loss of sample precision when using alternative gears. We investigated precision by examining coefficients of variation, using our side-by-side comparisons, and found the seines and quadrat sampler to have comparable precisions, although the quadrat was generally better. The quadrat sampler also collected more crayfish, and could be deployed in some habitats where the seine is likely to be very ineffective, e.g., vegetated habitats or deeper pools. Furthermore, use of the seine as reported in the current study has inherent operator bias because the 1-m2 area sampled was estimated, although recent modification have ameliorated this problem by attaching a 1-m2 frame at the bottom of the seine (Allert et al., 2012). Variation obtained from sampling benthic 59 WILLIAMS ET AL.: GEARS FOR LOTIC-DWELLING CRAYFISH STUDIES stream organisms is consistently high (Downes et al., 1993) and the fact that all of these samplers have SD < mean values is an indication of acceptable levels of precision. Size bias of many crayfish collection methods is well known (Momot, 1967; Haag et al., 2013). Our examination of size bias is not conclusive, but the catch ratio of young of year to adult is certainly comparable between all methods. Comparisons need to be made with caution because the efficiency of catching very small crayfish with the quadrat is unknown as all efficiency estimates have been made using adults in select habitats (Larson et al., 2008). We do, however, know that high crayfish mortalities occur during the first year of life and temporal changes should be expected (Brewer et al., 2009). Further, reduced sampling precision should be anticipated if sampling during periods of crayfish hatching (DiStefano et al., 2003). It is likely that no method accurately portrays young-of-year densities, but this subject is certainly in need of further investigation. We conclude that while the quadrat sampler catches significantly more crayfish than does the kick seine, the kick seine results are easily corrected to be comparable to the quadrat, with little loss of precision under the conditions tested in this study. Whereas correction factors likely do not improve variation of density estimates, they do provide a truer estimate of density and therefore useful if transitioning from a historical sampling approach to a new approach in a long-term study of the same system. The kick-seine technique appears to have appropriate characteristics to make it a suitable sampler for crayfishes under some conditions, e.g., relatively shallow, non-vegetated habitats. However, we suggest additional testing is warranted to assess the efficiency of either gear in pool or other slackwater habitats or areas of increased habitat complexity, e.g., vegetated areas. The kick seine comparison was conducted on small streams, with similar conditions of depth and substrate composition; however, results might differ if obtained under less homogeneous conditions. Projects involving crayfish populations have varying objectives – e.g., research, determining population status, or monitoring over space and time. If abundances or densities of crayfish are central to the collection effort, investigators or managers would be well served to further test characteristics of their sampler under conditions specific to the study area. A pilot project examining efficiencies and biases such as described by Peterson and Paukert (2009) would be relatively easy to accomplish and would give greater confidence in study results. Results from this study show a simple approach to adjust catch from one gear to another given samples are taken from small, relatively shallow, pebble-cobble streams with riffle-pool morphologies. The need to adjust data in this study appears minimal but the approach is useful where greater differences may be evident. This ‘pairedsamples approach’ may also be taken when monitoring using one gear and then switching to another so historic data may continue to be useful in conservation and management decisions. ACKNOWLEDGEMENTS This research is a contribution of the Missouri Cooperative Fish and Wildlife Research Unit (U.S. Geological Survey, Missouri Department of Conservation, University of Missouri, and Wildlife Management Institute cooperating) with collaboration from the Oklahoma Cooperative Fish and Wildlife Research Unit. Any use of trade, firm, or product names is for descriptive purposes and does not imply endorsement by the U.S. Government. We thank Robert DiStefano for input on study design and two anonymous reviewers for improving the quality of the manuscript. The Missouri Department of Conservation provided field equipment. 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