Wisconsin Amphipod Atlas · 2017-07-12 · based on collections made in Fond du Lac County in 1986 and 1994. Interestingly, the SWIMS database includes a record of S putealis being

Wisconsin Amphipod Atlas

Wisconsin Department of Natural Resources P.O. Box 7921 Madison, WI 53707-7921 Miscellaneous Publication PUB-SS-1174 2017

_______________________________________________________________ Summary: This report documents the distribution of 15 species of amphipod (Crustacea: Amphipoda: Crangonyctidae, Gammaridae, Hyalellidae, and Pontoporeiidae) occurring in Wisconsin lakes and rivers. Technical literature, museum specimens, and Wisconsin DNR field staff investigations provide the basis for county records/maps for each. One additional nonnative species (Apocorophium lacustre, Corophiidae) has colonized the Upper Mississippi River and may eventually be found in the state, and a second, Dikerogammarus villosus (‘killer shrimp,’ Gammaridae), is regulated as a ‘prohibited species’ under the state’s invasive species laws in an effort to prevent its introduction and spread. This atlas can serve as a baseline for conservation planning purposes. Contents Introduction 1 Species Accounts 2 Conclusions 9 Acknowledgements 9 Literature Cited 9 Author Contact Information

Dreux J. Watermolen Wisconsin Department of Natural Resources P.O. Box 7921 Madison, Wisconsin 53707-7921

(608) 266-8931 [email protected]

Cover Illustrations: An amphipod, from J.D. Dana’s New Text-book of Geology (Ivison, Blakeman & Company, NY, 1883) provided by the Educational Technology Clearinghouse, https://etc.usf.edu/.

The Wisconsin Department of Natural Resources provides equal opportunity in its employment,

programs, services, and functions under an Affirmative Action Plan. If you have any questions regarding this plan, please write to Equal Opportunity Office, Department of Interior, Washington, DC 20240.

This publication is available in alternative format (large print, Braille, audio tape, etc.) upon request. Please call (608) 266-0531 for more information. _________________________________________________________________

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Introduction About 240 species of amphipods occur in North American freshwater habitats where they are primarily bottom-dwelling, omnivorous scavengers. These little crustaceans provide a significant food source for predatory fishes (Wagner 1911, Jackson 1912, Pearse and Achtenberg 1920, Wells 1980) and birds (Munro 1941, Bartonek and Hickey 1969, Peterson and Ellarson 1977, Ross et al. 2005), and some species serve as the intermediate hosts for parasites of these vertebrates (Swales 1936, Weller 1938, Crompton and Nickol 1985, McGurdy et al. 1999). Their presence, absence, and community composition can be used to gauge water quality and some species are used routinely in toxicity testing (Hilsenhoff 1982, Watton and Hawkes 1984, Gerhardt et al. 2004, Major et al. 2013, MacNeil 2014). In spite of their ecological significance and our state’s rich limnological history, we know relatively little about the distribution of these crustaceans in Wisconsin waters. Holsinger (1976) presented preliminary distribution maps for freshwater amphipods occurring in the United States, but predicted that additional collecting would extend the plotted ranges of many species. Building on Holsinger’s work, Jass and Klausmeier (1995, 2003) described the geographic distribution, community associations, life histories, and habitat characteristics of eight species based on their field work in 21 southeastern counties, specimens in the Milwaukee Public Museum (MPM) collection, and the published literature. Since then, at least two introduced species have become established in the state and a third has been found in surrounding areas. Over the past decade or so, Wisconsin Department of Natural Resources (DNR) biologists have collected macroinvertebrates, including amphipods, at numerous sites throughout the state and submitted samples to the macroinvertebrate laboratory at the University of Wisconsin-Stevens Point for identification and enumeration. Records from this work are maintained in the Wisconsin DNR’s Surface Water Integrated Monitoring System (SWIMS) database1 and are used for various water quality assessment purposes. To date, only limited efforts have used SWIMS data for species conservation planning purposes. Recently, I downloaded and analyzed the 3,411 amphipod records included in SWIMS, covering the period 2005 through 2016, and updated Jass and Klausmeier’s (1995) distribution maps for species found in Wisconsin2. I also identified additional literature reports and specimen records from MPM, the United States National Museum of Natural History (Smithsonian Institution, USNM), Harvard University’s Museum of Comparative Zoology (MCZ), and the Illinois Natural History Survey (INHS). I supplemented the records with information from the U.S. Geological Survey’s Nonindigenous Aquatic Species (NAS) database3 and unpublished data from the Wisconsin DNR’s Bureau of Natural Heritage Conservation (NHC). Finally, I searched the World Amphipoda Database4 for additional records and to verify current taxonomic nomenclature.

1 SWIMS holds chemical, physical, and biological data and serves as the Wisconsin DNR’s data repository for Clean Water Act-related work. For more information regarding SWIMS, see http://dnr.wi.gov/topic/surfacewater/swims/. 2 Nine hundred forty-seven of the SWIMS records include identification to the genus level only and are therefore omitted from the maps included here. 3 The NAS information resource is an established central repository for spatially referenced biogeographic accounts of introduced aquatic species. For more information regarding NAS, see https://nas.er.usgs.gov/default.aspx. 4 The World Amphipoda Database is a component of the World Register of Marine Species (WoRMS), an authoritative and comprehensive list of names of marine organisms, including information on synonymy. To access the database, see http://www.marinespecies.org/amphipoda/.

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Species Accounts Fifteen amphipod species, representing four families, are now documented from Wisconsin. These include an endemic, groundwater-dwelling species and at least two introduced species. An additional nonnative species has colonized the Upper Mississippi River and may eventually be found in the state. Chapter NR 40, Wis. Admin. Code, regulates yet another nonnative species as a means of preventing its introduction and spread. County records for each species are summarized in the accompanying maps. Shaded counties reflect SWIMS data and may represent a single collection record, records from multiple collections, or collections from multiple waterbodies within the county. Dots centered in counties depict literature records; squares represent additional museum specimens (i.e. specimens not cited by Jass and Klausmeier [1995]). Records from the NAS database are depicted with triangles. Open circles denote unpublished NHC records. Literature records are from Jass and Klausmeier (1995) unless otherwise indicated. Museum collection information, complete SWIMS data, and additional literature citations for all species are available upon request. Family Crangonyctidae Zhang and Holsinger (2003) provide the most recent treatment of this family, including re- descriptions of Crangonyx species found in Wisconsin and identification keys. Crangonyx gracilis – Jass and Klausmeier (1995) reported this species from 11 counties in southeastern Wisconsin. The SWIMS database includes records from three additional counties: Door, Washington, and Wood. Additional MPM and USNM specimens are available for Calumet, Fond du Lac, Portage, and Waushara counties (Zhang and Holsinger 2003). The literature record for Columbia County comes from Weckel (1907). The Brown County record is based on a specimen collected in Green Bay that was tentatively identified as C. gracilis (Mozley and Howmiller 1977). Smith and Verrill (1872) and Bundy (1882) reported this species from Lake Superior. C. gracilis is considered a Species with Information Needs in the Wisconsin Wildlife Action Plan (Wisconsin DNR 2015).

Common name: northern lake crangonyctid Crangonyx minor – Jass and Klausmeier’s (1995, 2003) specimens from an unnamed stream in Kenosha County comprise the only available Wisconsin records for this species. This species is considered a Species with Information Needs in the Wisconsin Wildlife Action Plan (Wisconsin DNR 2015).

Common name: small stream crangonyctid

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Crangonyx pseudogracilis – This is the most widespread crangonyctid in Wisconsin. Jass and Klausmeier (1995) reported C. pseudogracilis from three counties. The SWIMS database includes 63 records from 25 additional counties. The literature record for Ashland County comes from Bousfield (1958). The records for Calumet, Columbia, Fond du Lac, and Walworth counties come from Zhang and Holsinger (2003) based on MPM specimens.

Common name: northern river crangonyctid Crangonyx richmondensis – Jass and Klausmeier (1995) reported this species from four counties in southeastern Wisconsin. The SWIMS database includes a record from one additional county in the north-eastern part of the state. The records for Juneau County come from Zhang and Holsinger (2003) and an MPM specimen. Unpublished NHC records extend the range in the northwest.

Common name: bog crangonyctid Stygobromus putealis – Holmes (1910) originally described this endemic groundwater-dwelling species based on specimens collected from a well at Waupun (Dodge County). The type specimens were deposited in the USNM collection. Hubricht (1943) also collected it from a well near Waupun, and it was later recorded from four additional sites in Fond du Lac and Green counties (Holsinger 1976). Jass and Klausmeier (2011) recently re-documented the presence of this species based on collections made in Fond du Lac County in 1986 and 1994. Interestingly, the SWIMS database includes a record of S putealis being collected from the Waupaca River (Waupaca County) in September 2010. The 20 specimens collected are no longer available for examination, but this record almost certainly repre-sents a misidentification of one of the lotic species. This species is considered a Species with Information Needs in the Wisconsin Wildlife Action Plan (Wisconsin DNR 2015).

Common name: Wisconsin well stygobromid

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Family Gammaridae Hou and Sket (2016) recently verified the monophyly of this large and diverse family. As discussed by Pinkster (1983), Jass and Klausmeier (1995), and Hou and Sket (2016), several factors make the taxonomic identification of gammarids difficult and uncertain. Nonetheless, I have treated published identifications as accurate. As such, the accounts that follow must be viewed in light of such uncertainties. Holzinger (1976) addresses species occurring in the United States and includes keys for their identification and notes on distribution and ecology. Gammarus fasciatus – Holzinger’s (1976) range map for this species covers the central and eastern portions of the state (stippled area on map). Jass and Klausmeier (1995) reported this species from 20 southeastern counties and Crawford County in the southwest based entirely upon earlier literature records, but noted that these earlier records may be unreliable because of confusion in separating G. fasciatus and the more common G. pseudolimnaeus. The SWIMS database includes G. fasciatus records from five additional counties and extends the distribution of this species to the western edge of the state. Additional MPM specimens are available for Bayfield, Milwaukee, and Waupaca counties. The U.S. Geological Survey’s NAS database includes records from Lake Michigan and Lake Superior. The Brown County record is from Howmiller (1971) and Howmiller and Beeton (1971) who reported this species at several sites in Green Bay. The literature record for Douglas County comes from Grigorovich et al. (2003, 2005) and represents specimens collected in Lake Superior at Superior. Specimens collected from Lake Superior at Ashland are available in the USNM collection. Pearse and Achtenberg (1920) reported this species in the diet of yellow perch (Perca flavescens) caught in Dane County lakes.

Common name: none available Gammarus lacustris – Holzinger’s (1976) range map for this species covers the northern half of the state (stippled area on map), but his work provided no specific locality records. The SWIMS database includes records from Ashland and Iron counties; the MPM collection includes specimens from Washburn County. Smith and Verrill (1872) reported G. lacustris in Lake Superior.

Common name: none available

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Gammarus pseudolimnaeus – Holzinger’s (1976) range map for G. pseudolimnaeus shows this species to be widespread in the southern and eastern portions of the state (stippled area on map) and absent from the northwest. Jass and Klausmeier (1995) reported this species from 21 southeastern counties. Its presence in several of these counties was confirmed by Stepenuck (1999) and Stepenuck et al. (2008). The SWIMS database includes 2,378 records, from every county except Menominee County, significantly expanding the known range of this species in the state. The literature records for Marathon, Portage, Shawano, and Waupaca counties are from Hooper (1993) and Scott (2007). Other new county records include Bayfield (Clamp 1987), Grant (Peterka 1969), Iowa (Hilsenhoff 1971, Engel 1985), Sauk (Miller 1982, 1984), and Washburn (Clamp 1987) and additional museum Common name: specimens from Barron (MPM), Bayfield (MPM), Dane none available (INHS, MPM), Dunn (MPM), Iowa (MPM), La Crosse (USNM), Oconto (MPM), and Vernon (MPM, USNM). Gammarus tigrinus – This species was introduced from the North American Atlantic Coast and is now present in all of the Great Lakes (Grigorovich et al. 2005, Trebitz et al. 2010, NAS database), as well as in the upper Mississippi River near the Wisconsin-Minnesota border (Grigorovich et al. 2008). G. tigrinus is likely more widespread in Wisconsin waters then the few available records in the NAS database suggest. The expansion of its range will likely have implications for native species and aquatic food webs (Grigorovich et al. 2005).

Common name: none available Chaetogammarus (=Echinogammarus) ischnus – This nonnative species was introduced from the Ponto-Caspian region and was first documented in Lake Michigan in 1998 (Nalepa et al. 2001), in Lake Superior in 2001 (Grigorovich et al. 2003), and in the Upper Mississippi River in 2004 (Grigorovich et al. 2008). The SWIMS database includes a collection record from Sheboygan County; the literature record for this county comes from Scudder Eikenberry et al. (2016). Wisconsin records of C. ischnus in the NAS database include: Brown County–Fox River above (south) of the De Pere dam, 2008; Fox River below (north) of the De Pere dam, 2008; Door County–Lake Michigan, in Green Bay south of Egg Harbor, 2008; Mink River, 2011; Marinette County–Peshtigo River near CR-BB at Keller Rd, 2011; Oconto County–Green Bay just south of Little Suamico River mouth, 2011. The literature record for Brown County comes from De Stasio (2013) and Kiehnau (2015). The Door, Manitowoc, Marinette, and Milwaukee county records are reported in Scudder Eikenberry et al. (2014, 2016). The range expansion of C. ischnus over wide areas will likely have implications for native species and aquatic food webs. Common name: none available

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Dikerogammarus villosus – This species has been introduced and spread readily across central Europe and the United Kingdom (Bij de Vaate et al. 2002, MacNeil et al. 2010, Rewicz et al. 2014). Known as the ‘killer shrimp,’ D. villosus is a predatory species that feeds on a wide range of macroinvertebrates and has physiological and behavioral characteristics that facilitate its spread (Bij de Vaate et al. 2002, Dick et al. 2002, Kley and Maier 2003). The experience in Europe demonstrates how D. villosus can quickly dominate macroinvertebrate communities (both in numbers and biomass), reduce biodiversity, and potentially alter energy cycling (Rewicz et al. 2014). D. villosus has not become established in North America. Nonetheless, Wisconsin regulates this species as a ‘prohibited species’ under the state’s invasive species laws5 in an effort to prevent its introduction and spread.

Common name: killer shrimp Family Corophiidae Apocorophium lacustre – Although not yet reported from Wisconsin, this nonnative species was documented in the Upper Mississippi River drainage in 2005 (Grigorovich et al. 2008) and will likely be found in the state with further field collecting. The effects it may have on native macroinvertebrate communities remain unknown.

Common name: none available Family Hyalellidae The taxonomy of this family is not currently well understood. Hyalella azteca was long considered North America’s most widely distributed amphipod, occurring in permanent and semi-permanent habitats across nearly the entire continent. In fact, most of the reports of Hyalella recorded in North and Central America have been assigned to Hyalella azteca. However, at least 11 other related species are known from the region, and although morphological studies suggested that it was a single species (Bousfield 1996), more recent evidence has demonstrated that H. azteca is actually a species complex of as many as seven or more related species (Hogg et al. 1998, Witt and Hebert 2000, Major et al. 2013, Soucek et al. 2015). Complicating matters further, Souceck et al. (2015) recently found co-occurring species in many habitats. The accounts that follow must be considered in light of the numerous records currently synonymized under the H. azteca species complex. As such, it may be best practice to refer historical records that cannot be verified with voucher specimens to ‘Hyalella spp.’ Soucek et al. (2015) contains the most recent key for the identification of North American Hyalella species.

5 Section NR 40.04(2)(d)5m, Wisconsin Administrative Code, classifies Dikerogammarus villosus as a ‘prohibited species.’ Section NR 40.04(3)(a), Wis. Admin. Code, prohibits the possession, transport, transfer, or introduction of D. villosus.

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Hyalella azteca – Jass and Klausmeier (1995) reported this species from 21 counties in southeastern Wisconsin. Its presence in several of those counties was confirmed by Stepenuck (1999) and Stepenuck et al. (2008). The SWIMS database includes records from two additional counties. The literature record for La Crosse County comes from Elstad (1986) and Chilton (1990) and represents the occurrence of this species in Pool 7 of the Upper Mississippi River. Howmiller (1971) reported H. azteca from several sites in Green Bay. Other new county records are as follows: Brown (De Stasio 2013), Door (Metzler and Sager 1986, Scudder Eikenberry 2014), Dunn (Northern States Power 1977), Iowa (Hilsenhoff 1971, Engel 1985), Marinette (Scudder Eikenberry 2014), Oconto (Scudder Eikenberry 2014), Outagamie (DeStasio 2013, Kiehnau 2015), Portage (Hooper 1993), Vernon (Elstad 1986), Vilas (Patrick 2013, Soucek et al. 2015), and Waupaca (Scott 2007). Pearse and Achtenberg (1920) reported this species in the diets of several fish caught in Dane County lakes. Barton and Hynes (1976) recorded it in Lake Superior. Additional museum specimens are available from Bayfield (MPM), Iowa (MPM), Lafayette (MPM), Marinette (MPM), Portage (MPM), Sauk (MCZ), Shawano (MPM), Trempealeau (MCZ), and Waupaca (MPM) counties. Gonzalez and Watling (2002) recently provided a detailed morphological description of H. azteca based on the syntype series used when the species was originally described.

Common name: none available Hyalella inermis – Stebbing (1906) synonymized H. inermis under H. azteca, but Gonzalez and Watling (2002) contended that H. inermis is a valid species, and Witt and Hebert (2000) provided evidence of such based, in part, on analysis of specimens collected in Vilas County. These latter authors noted, however, that detailed genetic analyses and morphological appraisals of specimens from the type locality would be necessary to confirm assignment of their specimens to this species.

Common name: none available

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Hyalella knickerbockeri – Stebbing (1906) synonymized H. knickerbockeri under H. azteca, but Gonzalez and Watling (2002) considered it to be a valid species. Jackson (1912) reported H. knicker-bockeri as “excessively abundant” in southern Wisconsin and “plentiful in many of the shallower lakes” in the northern part of the state. He also reported finding it “in a little rivulet fed largely by springs” near Monroe (Green County) and in the diets of several fishes collected in Storrs Lake (Rock County) and Lake Mendota (Dane County). Weckel (1907) and Baker (1924) reported H. knickerbockeri in the Lake Winnebago region. Other records include Delavan and Geneva lakes (Walworth County), the Wisconsin River, and Lakes Michigan and Superior (Weckel 1907).

Common name: none available Hyalella spinicauda – Soucek et al. (2015) recently described this species based on individuals collected from lentic and lotic habitats in Illinois, Michigan, Wisconsin, and Ontario, Canada. These authors collected specimens from Lake Winnebago in Calumet County and the La Crosse River in La Crosse County.

Common name: none available Family Pontoporeiidae Jass and Klausmeier (1995) did not treat members of this family as these deepwater species had previously been “collected in this region only from Lake Michigan and Green Lake (Juday and Birge 1927).” I discuss them here for the sake of completeness. At least two different species of Diporeia, possibly as many as eight, occur in the Great Lakes (Bousfield 1989). Pilgrim et al. (2009) recently concluded that Lake Superior Diporeia represent a distinct lineage that diverged from populations of the other lakes, but stopped short of referring them to a new species. Aids for identification of species are limited and all species records from the Great Lakes are currently referred to as Diporeia spp. Diporeia spp. – Diporeia spp. have historically been the most abundant benthic invertebrates in the offshore waters of the Great Lakes, comprising 60% to 80% of the benthic biomass (Cook and Johnson 1974, Mozley and Howmiller 1977, Winnell and White 1984, Nalepa 1987, Dermott et al. 2005). They serve as an important food source for various fishes and water birds (Peterson and Ellarson 1977, Wells 1980, Hondorp et al. 2005). Howmiller (1971) noted the abundance of Diporeia sp. in upper Green Bay. By the mid-1990s, however, Diporeia spp. populations declined precipitously in the lower Great

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Lakes (Nalepa et al. 1998, Dermott 2001, Nalepa et al. 2008). These declines have adversely affected fisheries (Pothoven et al. 2001, Hondorp et al. 2005). The Diporeia spp. in Lake Superior appear to have not experienced similar declines (Auer and Kahn 2004, Auer et al. 2009, Scharold et al. 2004). Specimens collected from Green Lake in 1922 are available in the USNM collection. Wagner (1911) noted the importance of Diporeia sp. in the diets of Green Lake’s ciscoes. Common name: none available Conclusions Successful biological diversity conservation efforts rely on knowledge of what species occupy a given area, where the species occur within the area, and the factors that influence observed distributions. The above records and accompanying maps illustrate that amphipods are more diverse and widely distributed in Wisconsin than previously published records suggest. Additional fieldwork, evaluation of museum collections, and further analyses of SWIMS data will allow a better understanding of their basic biology and the measures that might be needed for conservation purposes, particularly in light of the introduction and spread of nonnative and invasive species. Publication of these updated maps will hopefully stimulate interest in such work. Acknowledgements The literature review for this atlas was initiated as a result of the “Wisconsin’s Species Diversity: The State of Scientific Knowledge” symposium sponsored by the Aldo Leopold Chapter of the Society for Conservation Biology in April 1997. I am indebted to the chapter’s Non-insect Invertebrates Working Group for providing the push needed to begin an assessment of our knowledge. This report is dedicated to Joan P. Jass, a colleague who devoted a considerable part of her professional life to increasing our knowledge of Wisconsin crustaceans. Her work provides a foundation for this atlas. Terrell Hyde shared unpublished records from the Bureau of Natural Heritage Conservation. Dougal Walker prepared the distribution maps. I appreciate their assistance and willingness to help. Literature Cited Auer, N.A., B.A. Cannon, and M.T. Auer. 2009. Life history, distribution, and production of Diporeia

near the Keweenaw Peninsula Lake Superior. Journal of Great Lakes Research 35(4):579-590. Auer, N.A., and J.E. Kahn. 2004. Abundance and distribution of benthic invertebrates, with emphasis

on Diporeia, along the Keweenaw Peninsula, Lake Superior. Journal of Great Lakes Research 30(Suppl. 1):340–359.

Baker, F.C. 1924. The fauna of the Lake Winnebago region. Transactions of the Wisconsin Academy of

Science, Arts, and Letters 21:109-146. Barton, D.R., and H.B.N. Hynes. 1976. The distribution of Amphipoda and Isopoda on the exposed

shores of the Great Lakes. Journal of Great Lakes Research 2(2):207-214. Bartonek, J., and J. Hickey, 1969. Food habits of canvasbacks, redheads, and lesser scaup in

Manitoba. Condor 71(3):280-290.

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Bij de Vaate, A., K. Jazdzewski, H.A.M. Ketelaars, S. Gollasch, and G. Van der Velde. 2002. Geographical patterns in range extension of Ponto-Caspian macroinvertebrate species in Europe. Canadian Journal of Fisheries and Aquatic Sciences 59(7):1159-1174.

Bousfield, E.L. 1958. Fresh-water amphipod crustaceans of glaciated North America. Canadian Field-

Naturalist 72(2):55-113. Bousfield, E.L. 1989. Revised morphological relationships within the amphipod genera Pontoporeia and

Gammaracanathus and the “glacial relict” significance of their postglacial distributions. Canadian Journal of Fisheries and Aquatic Sciences 46(10):1714–1725.

Bousfield, E.L. 1996. A contribution to the reclassification of Neotropical freshwater hyalellid

amphipods (Crustacea: Gammaridea, Talitroidea). Bolletino del Museo Civico di Storia Naturale de Verona 20:175–224.

Bundy, W.F. 1882. A list of the Crustacea of Wisconsin, with notes on some new or little known

species. Transactions of the Wisconsin Academy of Sciences, Arts and Letters 5:177-184. Chilton, E.W. 1990. Macroinvertebrate communities associated with three aquatic macrophytes

(Ceratophyllum dimersum, Myriophyllum spicatum, and Vallisneria americana) in Lake Onalaska, Wisconsin. Journal of Freshwater Ecology 5(4):455-466.

Clamp, J.C. 1987. Five new species of Lagenophrys (Ciliophora, Peritricha, Lagenophryidae) from the

United States with observations on their developmental stages. Journal of Protozoology 34(4): 382-392.

Cook, D.G., and M.G. Johnson. 1974. Benthic macroinvertebrates of the St. Lawrence Great Lakes.

Journal of the Fisheries Research Board of Canada 31(5):763-782. Crompton, D.W.T., and B.B. Nickol. 1985. Biology of the Acanthocephala. Cambridge University Press,

Cambridge. Dermott, R. 2001. Sudden disappearance of the amphipod Diporeia from eastern Lake Ontario, 1993–

1995. Journal of Great Lakes Research 27(4):423–433. Dermott, R., R. Bonnell, and P. Jarvis. 2005. Population status of the amphipod Diporeia in eastern

North American lakes with or without Dreissena. Verh. Internat. Verein. Limnol. 29:880-886. De Stasio, B. 2013. Aquatic Invasive Species Monitoring Project, Year 2013 Report to the Fox River

Navigation Authority. Unpubl. Technical Report. Lawrence University, Appleton. Dick, J.T.A., D. Platvoet, and D.W. Kelly. 2002. Predatory impact of the freshwater

invader Dikerogammarus villosus (Crustacea: Amphipoda). Canadian Journal of Fisheries and Aquatic Sciences 59(6):1078-1084.

Elstad, C.A. 1986. Macrobenthic distribution and community structure in the upper navigation pools of

the Upper Mississippi River. Hydrobiologia 136(1):85-100. Engel, S. 1985. Aquatic community interactions of submerged macrophytes. Technical Bulletin 156.

Bureau of Research, Wisconsin Department of Natural Resources, Madison. Gerhardt, A., L. Janssens de Bisthoven, A.M.V.M. Soares. 2004. Macroinvertebrate response to acid

mine drainage: community metrics and on-line behavioural toxicity bioassay. Environmental Pollution 130(2):263-274.

Gonzalez, E.R., and L. Watling. 2002. Redescription of Hyalella azteca from its type locality, Vera Cruz,

Mexico (Amphipoda: Hyalellidae). Journal of Crustacean Biology 22(1):173-183.

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Grigorovich, I.A., T.R. Angradi, E.B. Emery, and M.S. Wooten. 2008. Invasion of the upper Mississippi River system by saltwater amphipods. Fundamental and Applied Limnology 173(1):67-77.

Grigorovich, I.A., M. Kang, and J.J.H. Ciborowski. 2005. Colonization of the Laurentian Great Lakes by

the amphipod Gammarus tigrinus, a native of the North American Atlantic Coast. Journal of Great Lakes Research 31(3):333-342.

Grigorovich, I.A., A.V. Korniushin, D.K. Gray, I.C. Duggan, R.I. Colautti, and H.J. MacIsaac. 2003.

Lake Superior: an invasion coldspot? Hydrobiologia 499(1-3):191-210. Hilsenhoff, W.L. 1971. Changes in the downstream insect and amphipod fauna caused by an

impoundment with a hypolimnion drain. Annals of the American Entomological Society 64(3): 743-746.

Hilsenhoff, W.L. 1982. Using a biotic index to evaluate water quality in streams. Technical Bulletin

132. Wisconsin Department of Natural Resources, Madison, WI. Hogg, I.D., C. Larose, Y. de Lafontaine, and K.G. Doe. 1998. Genetic evidence for a Hyalella species

complex within the Great Lakes–St. Lawrence River drainage basin: implications for ecotoxicology and conservation biology. Canadian Journal of Zoology 76(6):1134-1140.

Holmes, S.J. 1910. Description of a new subterranean amphipod from Wisconsin. Transactions of the

Wisconsin Academy of Sciences, Arts and Letters 16(1):77–80. Holzinger, J.R. 1976. The freshwater amphipod crustaceans (Gammaridae) of North America. Second

printing. Water Pollution Control Research Series 18050 ELDO4/72. U.S. Environmental Protection Agency, Cincinnati, OH.

Hondorp, D.W., S.A. Pothoven, and S.B. Brandt. 2005. Influence of Diporeia density on diet

composition, relative abundance, and energy density of planktivorous fishes in southeast Lake Michigan. Transactions of the American Fisheries Society 134(3):588–601.

Hooper, A.E. 1993. Effects of season, habitat, and an impoundment on twenty-five benthic community

measures used to assess water quality. M.S. thesis. University of Wisconsin-Stevens Point, Stevens Point, WI.

Hou, Z., and B. Sket. 2016. A review of Gammaridae (Crustacea: Amphipoda): the family extent, its

evolutionary history, and taxonomic redefinition of genera. Zoological Journal of the Linnean Society 176(2):323-348.

Howmiller, R.P. 1971. The benthic macrofauna of Green Bay, Lake Michigan. Ph.D. thesis. University

of Wisconsin, Madison. Howmiller, R.P., and A.M. Beeton. 1971. Biological evaluation of environmental quality, Green Bay,

Lake Michigan. Journal (Water Pollution Control Federation) 43(1):123-133. Hubricht, L. 1943. Studies on the Nearctic freshwater Amphipoda, III. Notes on the freshwater

Amphipoda of eastern United States, with descriptions of ten new species. American Midland Naturalist 29(3):683–712.

Jackson, H.H.T. 1912. A contribution to the natural history of the amphipod, Hyalella knickerbockeri

(Bate). Bulletin of the Wisconsin Natural History Society 10:49-60. Jass, J., and B. Klausmeier. 1995. Amphipods (exclusive of Pontoporeiidae) of southeastern Wisconsin.

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Jass, J., and B. Klausmeier. 2003. Utilization on lotic, lentic, and temporary habitats by the amphipod Crangonyx gracilis. Journal of Freshwater Ecology 18(4):635-638.

Jass, J., and B. Klausmeier. 2011. Presence of the well-dwelling amphipod Stygobromus putealis

(Crustacea: Crangonyctidae) re-documented in Wisconsin. Journal of Freshwater Ecology 26(4):599–603.

Juday, C., and E.A. Birge. 1927. Pontoporeia and Mysis in Wisconsin lakes. Ecology 8(4):445-452. Kiehnau, E.L. 2015. The characterization of a vital Wisconsin waterway: a biological assessment of the

Lower Fix River from 2006-2014. Honors Projects Paper 72. Lawrence University, Appleton, WI. Available online at http://lux.lawrence.edu/cgi/viewcontent.cgi?article=1079&context=luhp.

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