Publications - Dr. Stu Ludsin

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Professor, Director ODNR Partnership | ludsin.1@osu.edu | u.osu.edu/LudsinLab | Curriculum vitate [pdf]

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Recent Publications

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Testicular collections as a technique to increase milt availability in sauger (Sander canadensis)

Abstract
Sauger
Sauger (Sander canadensis)

This study was conducted to compare quality and quantity of sperm collected from sauger (S. canadensis) using two collection methods: stripping alone and testicular tissue collection combined with stripping. Sperm were collected from sauger broodstock (n = 20) during the breeding season. Fish were randomly assigned to two sperm collection groups: (1) stripping once or (2) stripping twice before testicular tissue collection for obtaining additional sperm. Sperm motility variables, morphology, total number produced, and fertilization (%) were compared using the two collection methods. Testicular sperm had greater total motility (70.1 ± 2.1% compared with 44.3 ± 5.7%) but there were fewer morphologically normal cells (76.4 ± 1.3% compared with 92.8 ± 1.0%) compared to sperm collected using the stripping procedure. Sperm collection regimen utilizing testicular collections and sperm extractions in combination with stripping resulted in a ∼ten fold increase in total number of motile and morphologically normal sperm (39.5 ± 4.1 × 10 9) compared with the currently utilized two sequential sperm stripping collection procedures alone (3.6 ± 4.1 × 10 9 sperm). In large-scale studies (150,000 eggs), fertilization, using sperm collected from testicular tissues (1.0 × 105 motile sperm/egg), was similar to sperm collected with only the stripping procedure (71.2 ± 5.5 %, 81.2 ± 5.5 %, P = 0.265). The results of this study indicate testicular collection combined with sperm extractions allows for collection of sperm of a quantity and quality to maximize fry production and reduce the problems with lack of broodstock availability for sperm collection.

Citation

Blawut, B., B. Wolfe, C.R. Moraes, D. Sweet, S.A. Ludsin, M.A.C. da Silva. 2020. Testicular collections as a technique to increase milt availability in sauger (Sander canadensis). Animal Reproduction Science 212, 106240. doi.org/10.1016/j.anireprosci.2019.106240

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Use of hypertonic media to cryopreserve sauger spermatozoa

Abstract

The objective of this study was to determine the effects of extender osmolality on postthaw sperm quality and fertility in Sauger Sander canadensis. Fresh milt from 10 male Saugers was diluted by using base extenders with osmolalities of 350, 500, or 750 mOsm/kg (E350, E500, and E750, respectively) containing 10% dimethyl sulfoxide, frozen in LN2 vapor, and stored. Sperm parameters (total motility, progressive motility, velocity, and viability) were assessed at different steps of the cryopreservation process (extended, equilibrated, and postthaw). Fertilization rates were compared between fresh and frozen sperm and at two sperm‐to‐egg ratios. All of the parameters that were measured, except for progressive motility, were reduced by cryopreservation. Extender 500 yielded the highest postthaw progressive motility (32.20 ± 3.86% [mean ± SD]) and velocity (84.97 ± 16.82 μm/s), whereas both E350 and E500 displayed the highest total motility (65.30 ± 4.24 and 68.70 ± 6.46%) and viability (80.60 ± 4.84 and 78.80 ± 3.91%), respectively. By contrast, E750 yielded the lowest postthaw velocity, viability, and total and progressive motility. Despite the increase in the motility parameters, fertilization in E350 (13.93%) was approximately double that in E500 (6.58%), although not statistically different. In conclusion, traditional isosmotic base extenders (E350) were found to be superior to hypertonic base extenders in the preservation of Sauger milt. These results serve as a starting point for future investigations of cryopreservation potential for Sauger spermatozoa that work toward developing a freezing protocol that is more suitable for large‐scale application.

Citation

Blawut, B., B. Wolfe, C.R. Moraes, S.A. Ludsin, M.A. Coutinho da Silva. 2020. Use of Hypertonic Media to Cryopreserve Sauger Spermatozoa. North American Journal of Aquaculture 82(1):84-91. doi.org/10.1002/naaq.10125

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Cyanobacterial blooms modify food web structure and interactions in western Lake Erie

Abstract
HAB season zooplankton tow being rinsed in a sieve cup
Zooplankton tow condensation in sieve cup; collected during 2014 harmful algal bloom in Lake Erie

With anthropogenic eutrophication and climate change causing an increase in cyanobacterial blooms worldwide, the need to understand the consequences of these blooms on aquatic ecosystems is paramount. Key questions remain unanswered with respect to how cyanobacteria blooms affect the structure of aquatic food webs, the foraging abilities of higher consumers, and the potential for cyanotoxins (e.g., microcystins [MCs]) to accumulate in fish. Toward addressing these uncertainties, physicochemical attributes, water (for MCs), phytoplankton, zooplankton, and epipelagic and benthic age-0 fish were sampled at 75 sites (44 sites for fish) of varying cyanobacteria concentration (0.1–44 μg/L) in western Lake Erie during the cyanobacteria bloom season, 2013–2014. Sites with high cyanobacteria biomass were characterized by Microcystis spp. (84–100% of biomass), detectible levels of MCs (maximum = 10.8 μg/L), and low water transparency (minimum = 0.25 m). Counter to expectations, strong positive relationships were found between cyanobacteria concentration and the biomass of several herbivorous zooplankton taxa (e.g., DaphniaDiaphanosoma spp., Bosmina (formerly Eubosminacoregoni, and Calanoida spp.). Expectations regarding fish were partly supported (e.g., diet selectivity varied across a cyanobacteria gradient) and partly not (e.g., consumption of zooplankton did not differ between bloom and non-bloom sites). These findings show that cyanobacterial blooms can strongly affect the distribution, composition, and interactions of zooplankton and fish, sometimes in surprising ways, highlighting the need to further explore their impact on aquatic food webs.

Citation

Briland, R.D., J.P. Stone, M. Manubolu, J. Lee, S.A. Ludsin. 2020. Cyanobacterial blooms modify food web structure and interactions in western Lake Erie. Harmful algae 92, 101586. doi.org/10.1016/j.hal.2019.03.004

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RAD-Seq refines previous estimates of genetic structure in Lake Erie walleye

Abstract
Lake Erie map with sampling sites noted
Lake Erie and locations of the walleye local spawning populations sampled for this study (from Figure 1 of this article).

Delineating population structure helps fishery managers to maintain a diverse “portfolio” of local spawning populations (stocks), as well as facilitate stock‐specific management. In Lake Erie, commercial and recreational fisheries for Walleye Sander vitreus exploit numerous local spawning populations, which cannot be easily differentiated using traditional genetic data (e.g., microsatellites). Here, we used genomic information (12,264 polymorphic loci) generated using restriction site‐associated DNA sequencing to investigate stock structure in Lake Erie Walleye. We found low genetic divergence (genetic differentiation index FST = 0.0006–0.0019) among the four Lake Erie western basin stocks examined, which resulted in low classification accuracies for individual samples (40–60%). However, more structure existed between western and eastern Lake Erie basin stocks (FST = 0.0042–0.0064), resulting in greater than 95% classification accuracy of samples to a lake basin. Thus, our success in using genomics to identify stock structure varied with spatial scale. Based on our results, we offer suggestions to improve the efficacy of this new genetic tool for refining stock structure and eventually determining relative stock contributions in Lake Erie Walleye and other Great Lakes populations.

Citation

Chen, K.Y., P.T. Euclide, S.A. Ludsin, W.A. Larson, M.G. Sovic, H.L. Gibbs, E.A. Marschall. 2020. RAD‐Seq Refines Previous Estimates of Genetic Structure in Lake Erie Walleye. Transactions of the American Fisheries Society 149(2):159-173. doi.org/10.1002/tafs.10215

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Otolith microchemistry shows natal philopatry of walleye in western Lake Erie

Abstract
Otolith with annuli visible
Walleye otolith with annuli visible

Natal philopatry is important to the structure of fish populations because it can lead to local adaptations among component stocks of a mixed population, reducing the risk of recruitment failure. By contrast, straying between component stocks may bolster declining populations or allow for colonization of new habitat. To examine rates of natal philopatry and straying among western Lake Erie walleye (Sander vitreus) stocks, we used the concentration of strontium [Sr] in otolith cores to determine the natal origin of adults captured at three major spawning sites: the Sandusky (n = 62) and Maumee (n = 55) rivers and the Ohio reef complex (n = 50) during the 2012–2013 spawning seasons. Mean otolith core [Sr] was consistently and significantly higher for individuals captured in the Sandusky River than for those captured in the Maumee River or Ohio reef complex. Although logistic regression indicates that no individuals with a Maumee River or Ohio reef complex origin were captured in the Sandusky River, quadratic discriminant analysis suggests low rates of straying of fish between the Maumee and Sandusky rivers. Our results suggest little straying and high rates of natal philopatry in the Sandusky River walleye stock. Similar rates of natal philopatry may also exist across western Lake Erie walleye stocks, demonstrating a need for stock-specific management.

Citation

Chen, K.Y., S.A. Ludsin, B.J. Marcek, J.W. Olesik, E.A. Marschall. 2020. Otolith microchemistry shows natal philopatry of walleye in western Lake Erie. Journal of Great Lakes Research, in pressdoi.org/10.1016/j.jglr.2020.06.006

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Towards more robust hydroacoustic estimates of fish abundance in presence of pelagic macroinvertebrates

Abstract
Chaoborus larva
Chaoborus larva

The inclusion of unwanted targets in hydroacoustic surveys biases estimates of fish abundance. Thus, knowledge of frequency-dependent responses of unwanted targets (e.g., pelagic macroinvertebrates) can help ensure that transducer frequencies are used that minimize this bias. We determined how fish density estimates varied across multiple frequencies when the larval stage of a midge, Chaoborus, was present in the water column. We hypothesized that fish density estimates would increase with increasing transducer frequency, owing to greater backscattering by Chaoborus at higher frequencies than lower ones, which allows it to be included with the backscattering caused by fish. We found that fish density estimates were always greater at higher frequencies (e.g., 120 and 200 kHz) compared to a lower one (70 kHz) in several productive north-temperate reservoirs. Furthermore, pairwise comparisons of total (i.e., fish plus Chaoborus) backscattering showed that significantly more backscattering occurred at higher rather than lower frequencies. We also found that fish density estimates varied between spring and summer, partially owing to inter-seasonal size variation in Chaoborus that influenced its backscattering. Beyond demonstrating why the presence of pelagic macroinvertebrates needs to be considered when estimating fish abundance with hydroacoustics, we provide methods to identify and reduce this bias.

Citation

Dillon, R.A., J.D. Conroy, L.G. Rudstam, P.F. Craigmile, D.M. Mason, S.A. Ludsin. 2020. Towards more robust hydroacoustic estimates of fish abundance in the presence of pelagic macroinvertebrates. Fisheries Research 230, 105667. doi.org/10.1016/j.fishres.2020.105667

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Spatial patterning of walleye recreational harvest in Lake Erie: Role of demographic and environmental factors

Abstract

Demographic and environmental factors can influence the spatial distribution of fish populations, potentially affecting the timing, location, and magnitude of harvest. Quantifying these relationships can be complicated, if their effects vary spatially over a population’s range or are non-additive (i.e., interactive), where one factor mediates the effect of another. Toward understanding the relative influence of demographic and environmental factors on fishery harvest in large freshwater lakes, we used varying-coefficient generalized additive models to explore the existence of non-additive, spatially-dependent effects of adult population size and thermal conditions on recreational harvest patterns of Lake Erie walleye (Sander vitreus) during 2006-2015. We identified nonlinear, additive, and generally positive effects of thermal conditions and adult population size on harvest rates. Their effects were, however, spatially-dependent, the accounting of which can help explain inter-annual and intra-annual variation in lake-wide harvest rates. Specifically, harvest rates increased more with increasing cumulative degree days in the eastern portion of the central basin, especially offshore, relative to the rest of the study area. Harvest rates also increased more with increasing walleye population size in the southwest portion of the west basin and the middle of the central basin compared to other study areas. As in marine ecosystems, our findings demonstrate the benefit of using modeling approaches that consider the spatial dependency of harvest rate on demographic and environmental factors to understanding broader harvest dynamics in large lakes. Their use could help managers and policy-makers ensure the sustained use of valued freshwater fish populations amidst demographic and environmental change.

Citation

Dippold, D.A., G.D. Adams, S.A. Ludsin. 2020. Spatial patterning of walleye recreational harvest in Lake Erie: Role of demographic and environmental factors. Fisheries Research 230, 105676. doi.org/10.1016/j.fishres.2020.105676

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Forecasting the combined effects of anticipated climate change and agricultural conservation practices on fish recruitment dynamics in Lake Erie

Abstract
  1. Many aquatic ecosystems are experiencing multiple anthropogenic stressors that threaten their ability to support ecologically and economically important fish species. Two of the most ubiquitous stressors are climate change and non‐point source nutrient pollution.
  2. Agricultural conservation practices (ACPs, i.e. farming practices that reduce runoff, prevent erosion, and curb excessive nutrient loading) offer a potential means to mitigate the negative effects of non‐point source pollution on fish populations. However, our understanding of how ACP implementation amidst a changing climate will affect fish production in large ecosystems that receive substantial upstream sediment and nutrient inputs remains incomplete.
  3. Towards this end, we explored how anticipated climate change and the implementation of realistic ACPs might alter the recruitment dynamics of three fish populations (native walleye Sander vitreus and yellow perch Perca flavescens and invasive white perch Morone americana) in the highly productive, dynamic west basin of Lake Erie. We projected future (2020–2065) recruitment under different combinations of anticipated climate change (n = 2 levels) and ACP implementation (n = 4 levels) in the western Lake Erie catchment using predictive biological models driven by forecasted winter severity, spring warming rate, and Maumee River total phosphorus loads that were generated from linked climate, catchment‐hydrology, and agricultural‐practice‐simulation models.
  4. In general, our models projected reduced walleye and yellow perch recruitment whereas invasive white perch recruitment was projected to remain stable or increase relative to the recent past. Our modelling also suggests the potential for trade‐offs, as ACP implementation was projected to reduce yellow perch recruitment with anticipated climate change.
  5. Overall, our study presents a useful modelling framework to forecast fish recruitment in Lake Erie and elsewhere, as well as offering projections and new avenues of research that could help resource management agencies and policy‐makers develop adaptive and resilient management strategies in the face of anticipated climate and land‐management change.
Citation

Dippold, D.A., N.R. Aloysius, S.C. Keitzer, H. Yen, J.G. Arnold, P. Daggupati, M.E. Fraker, J.F. Martin, D.M. Robertson, S.P. Sowa, M.V. Johnson, M.J. White, S.A. Ludsin. 2020. Forecasting the combined effects of anticipated climate change and agricultural conservation practices on fish recruitment dynamics in Lake Erie. Freshwater Biology 65(9): 1487-1508. doi.org/10.1111/fwb.13515

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Full Publications List

  1. Blawut, B., B. Wolfe, C.R. Moraes, D. Sweet, S.A. Ludsin, M.A.C. da Silva. 2020. Testicular collections as a technique to increase milt availability in sauger (Sander canadensis). Animal Reproduction Science 212, 106240. doi.org/10.1016/j.anireprosci.2019.106240
  2. Blawut, B., B. Wolfe, C.R. Moraes, S.A. Ludsin, M.A. Coutinho da Silva. 2020. Use of Hypertonic Media to Cryopreserve Sauger Spermatozoa. North American Journal of Aquaculture 82(1):84-91. doi.org/10.1002/naaq.10125
  3. Briland, R.D., J.P. Stone, M. Manubolu, J. Lee, S.A. Ludsin. 2020. Cyanobacterial blooms modify food web structure and interactions in western Lake Erie. Harmful algae 92, 101586. doi.org/10.1016/j.hal.2019.03.004
  4. Chen, K.Y., P.T. Euclide, S.A. Ludsin, W.A. Larson, M.G. Sovic, H.L. Gibbs, E.A. Marschall. 2020. RAD‐Seq Refines Previous Estimates of Genetic Structure in Lake Erie Walleye. Transactions of the American Fisheries Society 149(2):159-173. doi.org/10.1002/tafs.10215
  5. Chen, K.Y., S.A. Ludsin, B.J. Marcek, J.W. Olesik, E.A. Marschall. 2020. Otolith microchemistry shows natal philopatry of walleye in western Lake Erie. Journal of Great Lakes Research, in pressdoi.org/10.1016/j.jglr.2020.06.006
  6. Dillon, R.A., J.D. Conroy, L.G. Rudstam, P.F. Craigmile, D.M. Mason, S.A. Ludsin. 2020. Towards more robust hydroacoustic estimates of fish abundance in the presence of pelagic macroinvertebrates. Fisheries Research 230, 105667. doi.org/10.1016/j.fishres.2020.105667
  7. Dippold, D.A., G.D. Adams, S.A. Ludsin. 2020. Spatial patterning of walleye recreational harvest in Lake Erie: Role of demographic and environmental factors. Fisheries Research 230, 105676. doi.org/10.1016/j.fishres.2020.105676
  8. Dippold, D.A., N.R. Aloysius, S.C. Keitzer, H. Yen, J.G. Arnold, P. Daggupati, M.E. Fraker, J.F. Martin, D.M. Robertson, S.P. Sowa, M.V. Johnson, M.J. White, S.A. Ludsin. 2020. Forecasting the combined effects of anticipated climate change and agricultural conservation practices on fish recruitment dynamics in Lake Erie. Freshwater Biology 65(9): 1487-1508. doi.org/10.1111/fwb.13515
  9. Fraker, M.E., S.C. Keitzer, J.S. Sinclair, N.R. Aloysius, D.A. Dippold, H. Yen, J.G. Arnold, P.Daggupati, M.V. Johnson, J.F. Martin, D.M. Robertson, S.P. Sowa, M.J. White, S.A. Ludsin. 2020. Projecting the effects of agricultural conservation practices on stream fish communities in a changing climate. Science of the Total Environment, 747, 141112. doi.org/10.1016/j.scitotenv.2020.141112
  10. Marcek, B.J., E.A. Burbacher, K. Dabrowski, K.P. Winslow, S.A. Ludsin. 2020. Interactive effects of hypoxia and temperature on consumption, growth, and condition of juvenile hybrid striped bass. Transactions of the American Fisheries Society 149(1):71-83. doi.org/10.1002/tafs.10210
  11. May, C.J., S.A. Ludsin, D.C. Glover, E.A. Marschall. 2020. The influence of larval growth rate on juvenile recruitment in Lake Erie walleye (Sander vitreus). Canadian Journal of Fisheries and Aquatic Sciences 77(3):548-555. doi.org/10.1139/cjfas-2019-0059
  12. Stone, J.P., K.L. Pangle, S.A. Pothoven, H.A. Vanderploeg, S.B. Brandt, T.O. Höök, T.H. Johengen, S.A. Ludsin. 2020. Hypoxia’s impact on pelagic fish populations in Lake Erie: A tale of two planktivores. Canadian Journal of Fisheries and Aquatic Sciences,  77(7): 1131-1148. doi.org/10.1139/cjfas-2019-0265
  13. Bade, A.P., T.R. Binder, M.D. Faust, C.S. Vandergoot, T.J. Hartman, R.T. Kraus, C.C. Krueger, and S.A. Ludsin. 2019. Sex-based differences in spawning behavior account for male-biased harvest in Lake Erie walleye (Sander vitreus). Canadian Journal of Fisheries and Aquatic Sciences 76(11): 2003-2012. doi.org/10.1139/cjfas-2018-0339
  14. Brown, T., M.E. Fraker, and S.A. Ludsin. 2019. Space use of predatory larval dragonflies and tadpole prey in response to chemical cues of predation. American Midland Naturalist 181:53-62. doi.org/10.1674/0003-0031-181.1.53
  15. Dillon, R.A., J.D. Conroy, S.A. Ludsin. 2019. Hydroacoustic data‐analysis recommendations to quantify prey‐fish abundance in shallow, target‐rich ecosystems. North American Journal of Fisheries Management 39(2):270-288. doi.org/10.1002/nafm.10266
  16. Glaspie, C.N, M. Clouse, K.B. Huebert, S.A. Ludsin, D.M. Mason, J.J. Pierson, M.R. Roman, and S.B. Brandt. 2019. Fish diet shifts associated with the Northern Gulf of Mexico hypoxic zone. Estuaries and Coasts 42(8):2170-2183. doi.org/10.1007/s12237-019-00626-x
  17. Becher, C. M.G. Strahan, and S.A. Ludsin. 2018. Coded wire tag use with juvenile channel catfish: evaluation of mortality, retention, and growth. North American Journal of Fisheries Management 38:1367-1374. doi.org/10.1002/nafm.10238
  18. Blawut, B., B. Wolfe, C.R. Moraes, S.A. Ludsin, and M.A. Coutinho da Silva. 2018. Increasing saugeye (S. vitreus × S. canadensis) production efficiency in a hatchery setting using assisted reproduction technologies. Aquaculture 495:21-26. doi.org/10.1016/j.aquaculture.2018.05.027
  19. Chen, K.Y., E.A. Marschall, M.G. Sovic, A.C. Fries, H.L. Gibbs, S.A. Ludsin. 2018. assignPOP: An R package for population assignment using genetic, non‐genetic, or integrated data in a machine‐learning framework. Methods in Ecology and Evolution 9(2):439-446. doi.org/10.1111/2041-210X.12897
  20. Glaspie, C.N, M. Clouse, A.T. Adamack, Y. Cha, S.A. Ludsin, D.M. Mason, M.R. Roman, C.A. Stow, and S.B. Brandt. 2018. Effect of hypoxia on diet of Atlantic bumpers in the northern Gulf of Mexico. Transactions of the American Fisheries Society 147:740-748. doi.org/10.1002/tafs.10063
  21. Hu, C., S.A. Ludsin, J.F. Martin, E. Dittmann, and J. Lee. 2018. Mycosporine-like amino acids (MAAs)—producing Microcystis in Lake Erie: Development of a qPCR assay and insight into its ecology. Harmful algae 77:1-10. doi.org/10.1016/j.hal.2018.05.010
  22. Manubolu, M., J. Lee, K.M. Riedl, Z.X. Kua, L.P. Collart, and S.A. Ludsin. 2018. Optimization of extraction methods for quantification of microcystin-LR and microcystin-RR in fish, vegetable, and soil matrices using UPLC-MS/MS. Harmful algae 76:47-57. doi.org/10.1016/j.hal.2018.04.009
  23. Marin Jarrin, J.R., T.B. Johnson, S.A. Ludsin, J.M. Reichert, and K.L. Pangle. 2018. Do models parameterized with observations from the system predict larval yellow perch (Perca flavescens) growth performance better in Lake Erie? Canadian Journal of Fisheries and Aquatic Sciences 75:82-94. doi.org/10.1139/cjfas-2016-0392
  24. Niu, Q., M. Xia, S.A. Ludsin, P.Y. Chu, D.M. Mason, and E.S. Rutherford. 2018. High‐turbidity events in Western Lake Erie during ice‐free cycles: Contributions of river‐loaded vs. resuspended sediments. Limnology and Oceanography 63:2545-2562. doi.org/10.1002/lno.10959
  25. Chen, K.Y., S.A. Ludsin, M.M. Corey, P.D. Collingsworth, M.K. Nims, J.W. Olesik, K. Dabrowski, J.J. van Tassell, E.A. Marschall. 2017. Experimental and field evaluation of otolith strontium as a marker to discriminate between river-spawning populations of walleye in Lake Erie. Canadian Journal of Fisheries and Aquatic Sciences 74:693-701. doi.org/10.1139/cjfas-2015-0565
  26. Collingsworth, P.D., D.B. Bunnell, M.W. Murray, Y.C. Kao, Z.S. Feiner, R.M. Claramunt, B.M. Lofgren, T.O. Höök, and S.A. Ludsin. 2017. Climate change as a long-term stressor for the fisheries of the Laurentian Great Lakes of North America. Reviews in Fish Biology and Fisheries. 27:363–391. doi.org/10.1007/s11160-017-9480-3
  27. Goto, D., J.J. Roberts, S.A. Pothoven, S.A. Ludsin, H.A. Vanderploeg, S.B. Brandt, and T.O. Höök. 2017. Size-mediated control of perch-midge coupling in Lake Erie transient dead zones. Environmental Biology of Fishes 100:1587-1600. doi.org/10.1007/s10641-017-0667-1
  28. Lee, S., X. Jiang, M. Manubolu, K. Riedl, S.A. Ludsin, J.F. Martin, and J. Lee. 2017. Fresh produce and their soils accumulate cyanotoxins from irrigation water: implications for public health and food security. Food Research International 102:234-245. doi.org/10.1016/j.foodres.2017.09.079
  29. Wituszynski, D.M., C. Hu, F. Zhang, J.D. Chaffin, J. Lee, S.A. Ludsin, and J.F. Martin. 2017. Microcystin in Lake Erie fish: risk to human health and relationship to cyanobacterial blooms. Journal of Great Lakes Research 43:1084-1090. doi.org/10.1016/j.jglr.2017.08.006
  30. Brodnik, R.L., M.E. Fraker, E.J. Anderson, L. Carreon-Martinez, K.M. DeVanna, D.D. Heath, J.M. Reichert, E.F. Roseman, and S.A. Ludsin. 2016. Larval dispersal underlies demographically important intersystem connectivity in a Great Lakes yellow perch (Perca flavescens) population. Canadian Journal of Fisheries and Aquatic Sciences 73:416-426. doi.org/10.1139/cjfas-2015-0161
  31. Culbertson, A., J.F. Martin, N. Aloysius, and S.A. Ludsin. 2016. Anticipated impacts of climate change on 21st century Maumee River discharge and nutrient loads. Journal of Great Lakes Research 42:1332-1342. doi.org/10.1016/j.jglr.2016.08.008
  32. DeVanna-Fussell, K.M., R.E.H. Smith, M.E. Fraker, L. Boegman, K.T. Frank, T.J. Miller, J.T. Tyson, K.K. Arend, D. Boisclair, S.J. Guildford, R.E. Hecky, T.O. Hӧӧk, O.P. Jensen, J.K. Llopiz, C.J. May, R.G. Najjar, L.G. Rudstam, C.T. Taggart, Y.R. Rao, and S.A. Ludsin. 2016. A perspective on needed research, modeling, and management approaches that can enhance Great Lakes fisheries management under changing ecosystem conditions. Journal of Great Lakes Research 42:743-752. doi.org/10.1016/j.jglr.2016.04.007
  33. Keitzer, S.C., S.A. Ludsin, S.P. Sowa, G. Annis, J.G. Arnold, P. Daggupati, A.M. Froehlich, M.E. Herbert, M.V. Johnson, A.M. Sasson, H. Yen, M.J. White, and C.A. Rewa. 2016. Thinking outside of the lake: Can controls on nutrient inputs into Lake Erie benefit stream conservation in its watershed? Journal of Great Lakes Research 42:1322-1331. doi.org/10.1016/j.jglr.2016.05.012
  34. Lauber, T.B., R.C. Stedman, N.A. Connelly, L.G. Rudstam, R.C. Ready, G.L. Poe, D.B. Bunnell, T.O. Höök, M.A. Koops, S.A. Ludsin, and E.S. Rutherford. 2016. Using scenarios to assess possible future impacts of invasive species in the Laurentian Great Lakes. North American Journal of Fisheries Management 36:1292-1307. doi.org/10.1080/02755947.2016.1214647
  35. Watson, S.B., C. Miller, G. Arhonditsis , G.L. Boyer, W. Carmichael, M.N. Charlton, R. Confesor, D.C. Depew, T.O. Höök, S.A. Ludsin, G. Matisoff, S.P. McElmurry, M.W. Murray, P. Richards, Y.R. Rao, M.M. Steffen, and S.W. Wilhelm. 2016. The re-eutrophication of Lake Erie: harmful algal blooms and hypoxia. Harmful algae 56:44–66. doi.org/10.1016/j.hal.2016.04.010
  36. Yen, H., M.J. White, J.G. Arnold, S.C. Keitzer, M.V.V. Johnson, J.D. Atwood, P. Daggupati, M.E. Herbert, S.P. Sowa, S.A. Ludsin, D.M. Robertson, R. Srinivasan, and C.A. Rewa. 2016. Western Lake Erie Basin: Soft-data-constrained, NHDPlus resolution watershed modeling and exploration of applicable conservation scenarios. Science of the Total Environment 569:1265-1281. doi.org/10.1016/j.scitotenv.2016.06.202
  37. Carreon-Martinez, L.B., R.P. Walther, T.B. Johnson, S.A. Ludsin, and D.D. Heath. 2015. Benefits of turbid river plume habitat for Lake Erie yellow perch (Perca flavescens) recruitment determined by juvenile to larval genotype assignment. Plos One 10(5):e0125234. doi.org/10.1371/journal.pone.0125234
  38. DuFour, M.R., C.J. May, E.F. Roseman, S.A. Ludsin, C.S. Vandergoot, J.J. Pritt, M.E. Fraker, J.J. Davis, J.T. Tyson, J.G. Miner, E.A. Marschall, and C. Mayer. 2015. Portfolio theory as a management tool to guide conservation and restoration of multi‐stock fish populations. Ecosphere, 6(12), pp.1-21. doi.org/10.1890/ES15-00237.1
  39. Farmer, T.M., E.A. Marschall, K. Dabrowski, and S.A. Ludsin. 2015. Short winters threaten temperate fish populations. Nature Communications 6:7724. doi.org/10.1038/ncomms8724
  40. Fraker, M.E., E.J. Anderson, C.J. May, K.Y. Chen, J.J. Davis, K.M. DeVanna, M.R. DuFour, E.A. Marschall, C.M. Mayer, J.G. Miner, K.L. Pangle, J.J. Pritt, E.F. Roseman, J.T. Tyson, Y. Zhao, S.A. Ludsin. 2015. Stock-specific advection of larval walleye (Sander vitreus) in western Lake Erie: Implications for larval growth, mixing, and stock discrimination. Journal of Great Lakes Research 41(3):830–845. doi.org/10.1016/j.jglr.2015.04.008
  41. Fraker, M.E., E.J. Anderson, R. Brodnik, L. Carreon-Martinez, K.M. DeVanna, B.J. Fryer, D.D. Heath, J.M. Reichert, and S.A. Ludsin. 2015. Particle backtracking improves breeding subpopulation discrimination and natal-source identification in mixed populations. Plos One 10(3):e0120752. doi.org/10.1371/journal.pone.0120752
  42. Jiang, L., M. Xia, S.A. Ludsin, E.S. Rutherford, D.M. Mason, J. Marin Jarrin, and K.L. Pangle. 2015. Biophysical modeling assessment of the drivers for plankton dynamics in dreissenid-colonized western Lake Erie. Ecological Modelling 308:18-33. doi.org/10.1016/j.ecolmodel.2015.04.004
  43. Kane, D.D., S.A. Ludsin, R.D. Briland, and D.A. Culver. 2015. Ten+ years gone: continued degradation of offshore planktonic communities in U.S. waters of Lake Erie's western and central basins (2003–2013). Journal of Great Lakes Research 41(3):930–933. doi.org/10.1016/j.jglr.2015.06.002
  44. Marin Jarrin, J.R., K.L. Pangle, J.M. Reichert, T.B. Johnson, J. Tyson, and S.A. Ludsin. 2015. Influence of habitat heterogeneity on the foraging ecology of first feeding yellow perch larvae, Perca flavescens, in western Lake Erie. Journal of Great Lakes Research 41(1):208–214. doi.org/10.1016/j.jglr.2014.12.024
  45. Bunnell, D.B., R.P. Barbiero, S.A. Ludsin, C.P. Madenjian, G. Warren, D. Dolan, T. Brenden, R. Briland, O.T. Gorman, J.X. He, T.H. Johengen, B.F. Lantry, T.F. Nalepa, S.C. Riley, C.M. Riseng, T.J. Treska, I. Tsehaye, D.M. Warner, M.G. Walsh, and B.C. Weidel. 2014. Changing ecosystem dynamics in the Laurentian Great Lakes: bottom-up and top-down regulation. BioScience 64(1):26-39. Cover of BioScience Issue. doi.org/10.1093/biosci/bit001
  46. Carreon-Martinez, L.B., K.W. Wellband, T.B. Johnson, S.A. Ludsin, and D.D. Heath. 2014. Novel molecular approach demonstrates turbid river plumes reduce predation mortality on larval fish. Molecular Ecology 23(21):5366–5377. doi.org/10.1111/mec.12927
  47. Gebremariam, S., J.F. Martin, C. DeMarchi, N.S. Bosch, R. Confesor., and S.A. Ludsin. 2014. A comprehensive approach to evaluating watershed models for predicting river flow regimes critical to downstream ecosystem services. Environmental Modelling & Software 61:121-134. doi.org/10.1016/j.envsoft.2014.07.004
  48. Gover, T.R., M.K. Nims, J.J. Van Tassell, P.D. Collingsworth, J.W. Olesik, S.A. Ludsin, and E.A. Marschall. 2014. How much cleaning is needed when processing otoliths from fish larvae for microchemical analysis? Transactions of the American Fisheries Society 143(3):779-783. doi.org/10.1080/00028487.2014.889749
  49. Hosack, G.R., G.W. Peters, and S.A. Ludsin. 2014. Interspecific relationships and environmentally driven catchabilities estimated from fisheries data. Canadian Journal of Fisheries and Aquatic Sciences 71(3):447-463. doi.org/10.1139/cjfas-2013-0236
  50. Lochet, A., B.J. Fryer, S.A. Ludsin, and J.E. Marsden. 2014. Identifying natal origins of spawning adult sea lamprey (Petromyzon marinus): re-evaluation of the statolith microchemistry approach. Journal of Great Lakes Research 40(3):763-770. doi.org/10.1016/j.jglr.2014.04.014
  51. Ludsin, S.A., K.M. DeVanna, and R.E.H. Smith. 2014. Physical-biological coupling and the challenge of understanding fish recruitment in large lakes. Canadian Journal of Fisheries and Aquatic Sciences 71(5):775-794. doi.org/10.1139/cjfas-2013-0512
  52. Scavia, D., J.D. Allan, K.K. Arend, S. Bartell, D. Beletsky, N.S. Bosch, S.B. Brandt, R.D. Briland, I. Daloğlu, J.V. DePinto, D.M. Dolan, M.A. Evans, T.M. Farmer, D. Goto, H. Han, T.O. Höök, R. Knight, S.A. Ludsin, D. Mason, A.M. Michalak, R.P. Richards, J.J. Roberts, D.K. Rucinski, E. Rutherford, D.J. Schwab, T. Sesterhenn, H. Zhang, and Y. Zhou. 2014. Assessing and addressing the re-eutrophication of Lake Erie: central basin hypoxia. Journal of Great Lakes Research 40(2):226–246. doi.org/10.1016/j.jglr.2014.02.004
  53. Zhang, H., D.M. Mason, C.A. Stow, A.T. Adamack, S.B. Brandt, X. Zhang, D.G. Kimmel, M.R. Roman, and W.C. Boicourt, and S.A. Ludsin. 2014. Effects of hypoxia on habitat quality of pelagic planktivorous fishes in the northern Gulf of Mexico. Marine Ecology Progress Series 505:209-226. doi.org/10.3354/MEPS10768
  54. Dabrowski, K., M. Korzeniowska, T.M. Farmer, S.A. Ludsin, and E.A. Marschall. 2013. The function of wax esters in larval fish transition from endogenous to exogenous nutrition - are freshwater fish the exception or the rule? Communications in Agricultural and Applied Biological Sciences 78(4):102-103. PMID: 25141637
  55. Howe, E.A., C.P. Hand, J.E. Marsden, S.A. Ludsin, and B.J. Fryer. 2013. Discriminating natal origin of spawning adult sea lamprey (Petromyzon marinus) in Lake Champlain using statolith elemental signatures. Journal of Great Lakes Research 39(2):239-246. doi.org/10.1016/j.jglr.2013.02.006
  56. Kinter, B.T., and S.A. Ludsin. 2013. Nutrient inputs versus piscivore biomass as the primary driver of reservoir food webs. Canadian Journal of Fisheries and Aquatic Sciences 70(3):367-380. doi.org/10.1139/cjfas-2012-0369
  57. Lochet, A. J.E. Marsden, B.J. Fryer, and S.A. Ludsin. 2013. Instability of statolith elemental signatures revealed in newly metamorphosed sea lamprey (Petromyzon marinus). Canadian Journal of Fisheries and Aquatic Sciences 70(4):565-573. doi.org/10.1139/cjfas-2012-0410
  58. Mayer, C.M., L.E. Burlakova, P. Eklöv, D. Fitzgerald, A. Karatayev, S.A. Ludsin, S. Millard, E.L. Mills, A.P. Ostapenya, L.G. Rudstam, B. Zhu, T.V. Zhukova. 2013. Benthification of freshwater lakes: exotic mussels turning ecosystems upside down. Pages 575-585 in T.F. Nalepa and D.W. Schloesser, editors. Quagga and Zebra Mussels: Biology, Impacts, and Control, Second Edition. Taylor and Francis, New York. 816 pp. doi.org/10.1201/B15437-44
  59. Goto, D., K. Lindelof, D.L. Fanslow, S.A. Ludsin, J.J. Roberts, H.A. Vanderploeg, A.E. Wilson, and T.O. Höök. 2012. Indirect consequences of hypolimnetic hypoxia on zooplankton growth in a large eutrophic lake. Aquatic Biology 16:217-227. doi.org/10.3354/ab00442
  60. Lowe, M.R., S.A. Ludsin, B.J. Fryer, R.A. Wright, D.R. DeVries, and T.M. Farmer. 2012. Response to “Comment on ‘Otolith microchemistry reveals substantial use of freshwater by southern flounder in the northern Gulf of Mexico’”. Estuaries and Coasts 35:907–910. doi.org/10.1007/s12237-012-9493-z
  61. Mack, H.R., J.D. Conroy, K.A. Blocksom, R.A. Stein, and S.A. Ludsin. 2012. A comparative analysis of zooplankton field collection and sample enumeration methods. Limnology and Oceanography Methods 10(1):41-53. doi.org/10.4319/lom.2012.10.41
  62. Pangle, K.L, T.D. Malinich, D.R. DeVries, D.B. Bunnell, and S.A. Ludsin. 2012. Context-dependent planktivory: interacting effects of turbidity and predation risk on adaptive foraging. Ecosphere 3: article 114 (18 pp). doi.org/10.1890/ES12-00224.1
  63. Pothoven, S.A., H.A. Vanderploeg, D. M. Warner, J.S. Schaeffer, S.A. Ludsin, R.M. Claramunt, and T.F. Nalepa. 2012. Influences on Bythotrephes longimanus life-history characteristics in the Great Lakes. Journal of Great Lakes Research 38(1):134-141. doi.org/10.1016/j.jglr.2011.10.003
  64. Pothoven, S.A., H.A. Vanderploeg, T.O. Höök, and S.A. Ludsin. 2012. Hypoxia modifies planktivore–zooplankton interactions in Lake Erie. Canadian Journal of Fisheries and Aquatic Sciences 69(12):2018-2028. doi.org/10.1139/cjfas-2012-0144
  65. Roberts, J.J., S.A. Ludsin, S.A. Pothoven, H.A. Vanderploeg, and T.O. Höök. 2012. Evidence of hypoxic foraging forays by yellow perch (Perca flavescens) and potential consequences for prey consumption. Freshwater Biology 57(5):922–937. doi.org/10.1111/j.1365-2427.2012.02753.x
  66. Arend, K.K., D. Beletsky, J.V. DePinto, S.A. Ludsin, J.J. Roberts, D.K. Rucinski, D. Scavia, D.J. Schwab, and T.O. Höök. 2011. Seasonal and interannual effects of hypoxia on fish habitat quality in central Lake Erie. Freshwater Biology 56(2):366-383. doi.org/10.1111/j.1365-2427.2010.02504.x
  67. Brandt, S.B., M. Costantini, S. Kolesar, S.A. Ludsin, D.M. Mason, C.M. Rae, and H. Zhang. 2011. Does hypoxia reduce habitat quality for Lake Erie walleye? A bioenergetics perspective. Canadian Journal of Fisheries and Aquatic Sciences 68(5):857–879. doi.org/10.1139/f2011-018
  68. Carreon-Martinez, L.B., T.B. Johnson, S.A. Ludsin, and D.D. Heath. 2011. Utilization of stomach content DNA to determine diet diversity in piscivorous fish. Journal of Fish Biology 78(4):1170-1182. doi.org/10.1111/j.1095-8649.2011.02925.x
  69. Lowe, M.R., D.R. DeVries, R.A. Wright, S.A. Ludsin, and B.J. Fryer. 2011. Otolith microchemistry reveals substantial use of freshwater by southern flounder in the northern Gulf of Mexico. Estuaries and Coasts 34:630-639. doi.org/10.1007/s12237-010-9335-9
  70. Roberts, J.J., S.B. Brandt, D. Fanslow, S.A. Ludsin, S.A. Pothoven, D. Scavia, and T.O. Höök. 2011. Effects of hypoxia on consumption, growth, and RNA:DNA ratios of young yellow perch. Transactions of the American Fisheries Society 140(6):1574-1586. doi.org/10.1080/00028487.2011.638576
  71. Legler, N.D., T.B. Johnson, D.D. Heath, and S.A. Ludsin. 2010. Water temperature and prey size effects on the rate of digestion of larval and early juvenile fish. Transactions of the American Fisheries Society 139(3):868–875. doi.org/10.1577/T09-212.1
  72. Pangle, K.L., S.A. Ludsin, and B.J. Fryer. 2010. Otolith microchemistry as a stock identification tool for freshwater fishes: testing its limits in Lake Erie. Canadian Journal of Fisheries and Aquatic Sciences 67(9):1475–1489. doi.org/10.1139/F10-076
  73. Reichert, J.M., B.J. Fryer, K.L. Pangle, T.B. Johnson, J.T. Tyson, A.B. Drelich, and S.A. Ludsin. 2010. River-plume use during the pelagic larval stage benefits recruitment of a lentic fish. Canadian Journal of Fisheries and Aquatic Sciences 67(6):987-1004. doi.org/10.1139/F10-036
  74. Lowe, M.R., D.R. DeVries, R.A. Wright, S.A. Ludsin, and B.J. Fryer. 2009. Coastal largemouth bass (Micropterus salmoides) movement in response to changing salinity. Canadian Journal of Fisheries and Aquatic Sciences 66(12):2174-2188. doi.org/10.1139/F09-152
  75. Ludsin, S.A., X. Zhang, S.B. Brandt, M.R. Roman, W.C. Boicourt, D.M. Mason, and M. Costantini. 2009. Hypoxia-avoidance by planktivorous fish in Chesapeake Bay: implications for food web interactions and fish recruitment. Journal of Experimental Marine Biology and Ecology 381(Suppl. 1):S121-S131. doi.org/10.1016/j.jembe.2009.07.016
  76. Pothoven, S.A., H.A. Vanderploeg, S.A. Ludsin, T.O. Höök, and S.B. Brandt. 2009. Feeding ecology of emerald shiners and rainbow smelt in central Lake Erie. Journal of Great Lakes Research 35(2):190-198. doi.org/10.1016/j.jglr.2008.11.011
  77. Roberts, J.J., T.O. Höök, S.A. Ludsin, S.A. Pothoven, H.A. Vanderploeg, and S.B. Brandt. 2009. Effects of hypolimnetic hypoxia on foraging and distributions of Lake Erie yellow perch. Journal of Experimental Marine Biology and Ecology 381(Suppl. 1):S132-S142. doi.org/10.1016/j.jembe.2009.07.017
  78. Vanderploeg, H.A., S.A. Ludsin, J.F. Cavaletto, T.O. Höök, S.A. Pothoven, S.B. Brandt, J. Liebig, and G.A. Lang. 2009. Hypoxic zones as habitat for zooplankton in Lake Erie: Refuges from predation or exclusion zones? Journal of Experimental Marine Biology and Ecology 381(Suppl. 1):S108-S120. doi.org/10.1016/j.jembe.2009.07.015
  79. Vanderploeg, H.A., S.A. Ludsin, S.A. Ruberg, T.O. Höök, S.A. Pothoven, S.B. Brandt, G.A. Lang, J. Liebig, and J.F. Cavaletto. 2009. Hypoxia affects spatial distributions and overlap of pelagic fish, zooplankton, and phytoplankton in Lake Erie. Journal of Experimental Marine Biology and Ecology 381(Suppl. 1):S92-S107. doi.org/10.1016/j.jembe.2009.07.027
  80. Zhang, H., S.A. Ludsin, M.R. Roman, W.C. Boicourt, X. Zhang, D.G. Kimmel, A.T. Adamack, D.M. Mason, S.B. Brandt, and J.J. Pierson. 2009. Hypoxia-driven changes in the behavior and spatial distribution of pelagic fish and mesozooplankton in the northern Gulf of Mexico. Journal of Experimental Marine Biology and Ecology 381(Suppl. 1):S80-S91. doi.org/10.1016/j.jembe.2009.07.014
  81. Costantini, M., S.A. Ludsin, D.M. Mason, X. Zhang, W.C. Boicourt, and S.B. Brandt. 2008. Effect of hypoxia on habitat quality of striped bass (Morone saxatilis) in Chesapeake Bay. Canadian Journal of Fisheries and Aquatic Sciences 65(5):989-1002. doi.org/10.1139/f08-021
  82. Hand, C.P, S.A. Ludsin, B.J. Fryer, and J.E. Marsden. 2008. Statolith microchemistry as a technique for discriminating among Great Lakes sea lamprey (Petromyzon marinus) spawning tributaries. Canadian Journal of Fisheries and Aquatic Sciences 65(6):1153-1164. doi.org/10.1139/F08-045
  83. Lu, Y., S.A. Ludsin, D.L. Fanslow, and S.A. Pothoven. 2008. Comparison of three micro-quantity techniques for measuring total lipids in fish. Canadian Journal of Fisheries and Aquatic Sciences 65(10):2233-2241. doi.org/10.1139/F08-135
  84. Melancon, S., B.J. Fryer, J.E. Gagnon, and S.A. Ludsin. 2008. Mineralogical approaches to the study of biomineralization in fish otoliths. Mineralogical Magazine 72(2):627-637. doi.org/10.1180/minmag.2008.072.2.627
  85. Pothoven, S.A., S.A. Ludsin, T.O. Höök, D.L. Fanslow, D.M. Mason, P.C. Collingsworth, and J.J. Van Tassell. 2008. An evaluation of Bioelectrical Impedance Analysis for estimating fish energy content. Transactions of the American Fisheries Society 137(5):1519-1529. doi.org/10.1577/T07-185.1
  86. Hawley, N., T.H. Johengen, Y.R. Rao, S.A. Ruberg, D. Beletsky, S.A. Ludsin, B.J. Eadie, D.J. Schwab, T.E. Croley, and S.B. Brandt. 2006. Lake Erie hypoxia prompts Canada-U.S. study. EOS Transactions 87(32):313-319. doi.org/10.1029/2006EO320001
  87. Ludsin, S.A., B.J. Fryer, and J.E. Gagnon. 2006. Comparison of solution-based versus laser-ablation ICPMS for analysis of larval fish otoliths. Transactions of the American Fisheries Society 135(1):218–231. doi.org/10.1577/T04-165.1
  88. Melancon, S. B.J. Fryer, S.A. Ludsin, J.E. Gagnon, and Z. Yang. 2005. Effects of crystal structure on the uptake of metals by lake trout (Salvelinus namaycush) otoliths. Canadian Journal of Fisheries and Aquatic Sciences 62(11):2609-2619. doi.org/10.1139/f05-161
  89. Hedges, K.J., S.A. Ludsin, and B.J. Fryer. 2004. Effects of ethanol preservation on otolith micro-chemistry. Journal of Fish Biology 64(4):923-947. doi.org/10.1111/j.1095-8649.2004.00353.x
  90. Mora, C., P.M. Chittaro, P.F. Sale, J.P. Kritzer, and S.A. Ludsin. 2003. Patterns and processes in reef fish diversity. Nature 421:933-936. doi.org/10.1038/nature01393
  91. Sale, P.F., and S.A. Ludsin. 2003. The extent and spatial scale of connectivity among reef fish populations: implications for marine protected areas designated for fisheries enhancement. Gulf and Caribbean Research 14(2):119-128. doi.org/10.18785/gcr.1402.09
  92. Hobbs, B.F., S.A. Ludsin, R.L. Knight, P.A. Ryan, J. Biberhofer, and J.H.H. Ciborowski. 2002. Fuzzy cognitive mapping as a tool to define management objectives for complex ecosystems. Ecological Applications 12(5):1548-1565. doi.org/10.1890/1051-0761(2002)...
  93. Ludsin, S.A., and A.D. Wolfe. 2001. Biological invasion theory: Darwin’s contributions from The Origin of Species. BioScience 51(9):780-789. doi.org/10.1641/0006-3568(2001)...
  94. Ludsin, S.A., M.W. Kershner, K.A. Blocksom, R.L. Knight, and R.A. Stein. 2001. Life after death in Lake Erie: nutrient controls drive fish species richness, rehabilitation. Ecological Applications 11(3):731-746. doi.org/10.1890/1051-0761(2001)...
  95. Pine, W.E., S.A. Ludsin, and D.R. DeVries. 2000. First-summer survival of largemouth bass cohorts: is early spawning really best? Transactions of the American Fisheries Society 129(2):504-513. doi.org/10.1577/1548-8659(2000)...
  96. Ludsin, S.A., and D.R. DeVries. 1997. First-year recruitment of largemouth bass: the interdependency of early life stages. Ecological Applications 7(3):1024-1038. doi.org/10.1890/1051-0761(1997)...