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Rapid Expansion of Banded Killifish Fundulus diaphanus across Northern Illinois: Dramatic Recovery or Invasive Species?

ABSTRACT.--The distribution of the Illinois state-threatened Banded Killifish Fundulus diaphanus has remained largely unchanged in Illinois from 1880 to 2000, being restricted mainly to the northeastern corner of the state. One population identified as Western Banded Killifish F. d. menona has remained stable in the glacial lakes region along the southeastern Wisconsin-northeastern Illinois border. Starling in 2001 a second population began to spread and become more common along the Lake Michigan shoreline. From there the population expanded through the Chicago Area Waterway System into the Des Plaines River and eventually the Illinois River. Historical museum specimens from this region are identified as F. d. menona, but recent specimens are identified as hybrids between F. d. menona and Eastern Banded Killifish F. d. diaphanus. Subsequently, a third population appeared in the Mississippi River near the mouth of the Rock River in 2009 and has spread from there. These individuals are also identified as F. d. menona, The rapid expansion of Banded Killifish from bake Michigan to the Illinois River appears to be an invasion of the Eastern subspecies F. d. diaphanus and the subsequent hybridization with the native Western subspecies. It is unknown where the Banded Killifish in the Mississippi River originated, possibly from populations 160+ kilometers upstream or human introductions. As the Illinois River and Mississippi River populations continue to expand their ranges, their ecological impacts are unknown at this time.


Rapid changes in a species distribution involving the increasing range of an invasive species (e.g., Irons el al., 2006; Kolar el al., 2007) or the decreasing range of a sensitive species (Metzke et al., 2012) are known to occur. Far less common is the increasing range of a sensitive species, especially when there is no known intentional stocking or translocation by natural resource agencies. Un-aided range expansion of sensitive species raises the hope that perhaps they are responding to improvements in environmental quality. If this is indeed the case, then there is great interest in attempting similar environmental improvements elsewhere to expand the ranges of these sensitive species even further. A situation like this is currently developing in the state of Illinois where Banded Killifish Fundulus diaphanus have been listed as state-threatened since 1989 (Mankowski, 2012).

The species as a whole is considered secure over its entire range, which extends from the Dakotas eastward through the Great Lakes to Newfoundland, and then south along the Atlantic coast to South Carolina (Scott and Crossman, 1973; Page and Burr, 2011). The eastern half of the range is generally considered to be inhabited by the subspecies Eastern Banded Killifish Fundulus diaphanus diaphanus, whereas the western half is inhabited by the subspecies Western Banded Killifish Fundulus diaphanus menona (Hubbs and Lagler, 1947; based on Shapiro, 1947). Genetic divergence between die two subspecies, according to mitochondrial D-Loop, is 2% and dates the split between subspecies to approximately 570,000 y ago (April and Turgeon, 2006). Introgression between the two subspecies is known (April and Turgeon, 2006), as well as hybridization with other species, such as die Mummichog Fundulus heteroclitus (Weed, 1921; Hubbs et al., 1943; Griffith, 1968, 1972; Fritz and Garside, 1974; Dawley, 1992; Dawley et at, 1999, 2000; Hernandez Chavez and Turgeon, 2007; Merette et al, 2009).

Illinois is along the southern edge of the Western Banded Killifish range, with historical records primarily in northeast Illinois and two in the center of the state (Forbes and Richardson, 1919, 1920; Smith, 1979). Fish biologists in recent years are collecting more Banded Killifish and from locations where they have not previously been recorded. The goals of this manuscript are to (1) document changes in the distribution of Banded Killifish in Illinois from 1880 to the present and (2) assess reasons that may be driving these distributional changes. This information is critical for the successful management of Banded Killifish that is designated a Species in Greatest Conservation Need (e.g., state-threatened) and whose recovery is a goal of the Illinois Wildlife Action Plan (Illinois Department of Natural Resources, 2016).

Materials and Methods

Locality data were based primarily on vouchered specimens in fish collections at The Field Museum (FMNH), Chicago and the Illinois Natural History Survey (INHS), Champaign [museum acronyms based on Sabaj (2016)]. Included with these records were significant amounts of field observation data from the Illinois Department of Natural Resources--Aquatic Nuisance Species Program conducting surveys in Illinois rivers. The Illinois Department of Natural Resources--Lake Michigan Program provided field observation data from five annual beach surveys (south to north; Jackson-Outer Harbor, Farwell Avenue, Tower Road, Waukegan, and North Point) starting in 1979. Supplemental data included personal observations from several of the authors and colleagues, as well as literature records that are considered reliable (e.g, Meek and Hildebrand, 1910; Rivera et al., 2013; Tiemann et al., 2015; Hrabik, 2016; Lamer et al., 2016; Schmidt, 2016).

These types of data are considered 'presence only' and do not explicitly take into consideration absences from locations where Banded Killifish are historically known to occur. However, some of the data sources, such as the Illinois Department of Natural Resources--Lake Michigan Program annual beach surveys and the Long-Term Illinois River Fish Population Monitoring Program (also known as Long Term Electrofishing), do include repeated surveys at the same sites over long time periods. In other instances there are examples of areas that were the focus of short-term intensive surveys, such as Will County (Willink and Veraldi, 2009) and Lake Calumet (Greenfield and Rogner, 1984). Survey methods, effort, and reporting have also changed over the past 130+ y. There will always be gaps in our understanding, but based on the considerable number of fish surveys that have been conducted in Illinois, we believe the general changes in distribution patterns of Banded Killifish in Illinois are real and not simply artifacts of sampling error, changes in methodology, or increased sampling effort. Because fish survey methods have changed considerably over the years, it was not possible to determine a standardized catch per unit effort with this data.

Identification of preserved specimens in the collections of the FMNH and INHS was based on lateral row scales, which were counted according to Hubbs and Lagler (1947). Although the character is variable, it has been demonstrated to be informative over the entire geographic range of the species (Shapiro, 1947, in Hubbs and Lagler, 1947; April and Turgeon, 2006; Hernandez Chavez and Turgeon, 2007). Other characters, such as fin ray counts and pigmentation, were explored. However, normal fin ray counts between these subspecies differ on average by one fin ray, with considerable overlap in the range of counts between them. Pigmentation is difficult due to changes during ontogeny.

Data for specimens identified as the Western subspecies F. d. menona from Minnesota, Iowa, South Dakota, Illinois, and Indiana were combined because of considerable overlap of lateral row scale counts among regions. These values were consistent with what has previously been reported in the literature (Shapiro, 1947, in Hubbs and Lagler, 1947). All museum specimens examined were collected prior to 1957, well before any purported changes in distribution patterns.

Specimen data for individuals identified as the Eastern subspecies F. d. diaphanus were based on specimens from Maine as well as mean lateral row scale values reported from Hernandez Chavez and Turgeon (2007). The literature values were included because only a few museum specimens of F. d. diaphanus were readily available, and Hernandez Chavez and Turgeon (2007) corroborated their results with genetic data.



The distribution of Banded Killifish in Illinois is fairly consistent from 1880-2000 (Fig. 1). There are two records from the center of the state, but both date to 1880. They have not been reported from the Sangamon watershed since and are considered extirpated (Smith, 1979). The rest of the records are restricted to northeastern Illinois, with population centers in the glacial lakes region near the Wisconsin border and the Calumet region adjacent to Indiana (Forbes and Richardson, 1919, 1920; Smith, 1979; Retzer and Batten, 2005; Willink, 2009).

Banded Killifish were common in the mouths of Lake Michigan tributaries in the late 1800s (Nelson, 1876; Jordan, 1878) but disappeared through most of the 20th century (Smith, 1979). However, they began appearing in standardized Illinois Department of Natural Resources Lake Michigan Program annual beach surveys in 2001 (Fig. 2). They have been present ever since and catch rates have increased over time. Locations are scattered along the entire length of the Lake Michigan shoreline in Illinois (Fig. 1).

Although Banded Killifish were known to occur in the Calumet Region since record keeping started, this population appeared to increase in this area around 2010. From there they spread down the Calumet Sag Channel, the Chicago Sanitary and Shipping Canal, the Des Plaines River, and into the upper Illinois River (Fig. 1). There is no evidence (e.g., from Illinois Department of Natural Resources routine basin surveys) for Banded Killifish from the Illinois or lower Des Plaines rivers moving up the Fox or Des Plaines rivers to the glacial lakes along the Wisconsin border. As a side note, the Forest Preserve District of DuPage County transplanted Banded Killifish from northern Illinois to a site along the West Branch of the DuPage River in 2010, but the fish have not been seen since at the site.

The first record for Banded Killifish in the Illinois River was 2006 near Starved Rock State Park in Utica, LaSalle County (Fig. 1). Around 2010 the population expanded downstream to the Marshall--Peoria County line. From 2001 to 2010, the average rate of range expansion of Banded Killifish from Lake Michigan to the Marshall-Peoria County line on the Illinois River was 25.4 river km/y.

Conversely, the first record for Banded Killifish along the Illinois border of the Mississippi River was in 2009 in Davenport, Iowa, (John Olson, Ben Hucka, and Jerad Strieker, Iowa Department of Natural Resources, pers. comm.), which is opposite Rock Island County, Illinois (Fig. 1). Around 2010 the Mississippi River population expanded downstream and moderately up the Rock River. From 2006 to 2015, the average rate of expansion down the Mississippi River from Davenport, Iowa to West Alton, Missouri, was 55.1 river km/y.

Subspecies and hybrid identification.--Lateral row scales for Western Banded Killifish, including specimens from Illinois collected prior to 1957, ranged from 36 to 44, with all but one value between 38-43 (n = 62) (Fig. 3). Eastern Banded Killifish had 48-51 lateral row scales (n = 7).

Specimens collected in the Calumet Region in 2010 and 2011 had 43 to 46 lateral row scales, with six of the seven vouchers with either 45 or 46 lateral row scales (n = 7) (Fig. 3). These values are intermediate between the Western and Eastern values. Hybrids between Western Banded Killifish and Eastern Banded Killifish exhibit a similar intermediate pattern elsewhere (Shapiro, 1947, in Hubbs and Lagler, 1947; April and Turgeon, 2006; Hernandez Chavez and Turgeon, 2007); therefore, we are tentatively identifying the recent Calumet individuals as hybrids between the Western and Eastern subspecies. Specimens collected in the Calumet Region prior to 1937 had 39-44 lateral row scales with a mean of 42 (n = 23).

Specimens collected in the Mississippi River proper in 2013 had 39 and 44 lateral row scales, which identify them as Western Banded Killifish (n = 2). Specimens collected in Coon Creek (the most upstream Rock River locality) in 2013 had 42 and 47 lateral row scales (n = 2). The individual with 42 scales is consistent with the other Western Banded Killifish identified from the Mississippi watershed. The individual with 47 lateral row scales is difficult to identify because it is just below values for Eastern Banded Killifish, and conceivably within the range of hybrids.


Populations in the distribution of Banded Killifish remained unchanged for over a century (Smith, 1979). Most glacial lakes populations appeared stable, although there may have been a decline in the Fox Chain O'Lakes, which is a subset of lakes that the Fox River flows directly through that has been heavily impacted by development. Known populations in Wolf Lake (Willink, 2009) and Powder Horn Lake (Retzer and Batten, 2005) in the Calumet Region also have persisted over time.

Banded Killifish abundance increased along the shoreline of Lake Michigan at the beginning of the 21st century and eventually cascaded down the Illinois River. Environmentally, the Great Lakes are continually exposed to different conditions and stressors, with introduced species being a common item of concern. Invasive Zebra Mussels Dreissena polymorpha and Quagga Mussels Dreissena rostriformis bugensis became established in the 1980s and 1990s (Nalepa et al., 2001), filtering plankton from the water column, therefore increasing water clarity. A consequence of increased water clarity is increased macrophyte growth. This benefits Banded Killifish because they use vegetation to spawn (Richardson, 1939). Concordant with the increase in mussel numbers was a decrease in invasive Alewife Alosa pseudoharengus (Bunnell et al., 2015; Warner et al., 2015), which are known to have negative impacts on numerous Lake Michigan fish species via predation on larvae and smaller individuals (Madenjian et al, 2008).

Physical aspects of Lake Michigan are shifting as well. Water temperature, for example, is increasing with climate change, with a particularly large jump of about 2.5 C during the 1997-1998 El Nino event (Gronewold and Stow, 2014). Pollution is on the decline [e.g., drops in PCB levels in salmon (Rasmussen et al., 2014)] attributable to decades of increased environmental regulations, although pollution is certainly still present in the region. Similarly, the upper Illinois River was so heavily polluted it became devoid of freshwater mussels (Starrett, 1971). However, after the passage of the Clean Water Act, conditions improved and mussels recolonized the area (Sietman et al, 2001).

As to whether any of these biological or environmental factors, an unknown factor(s), or combination of factors played a role in the expansion of Banded Killifish in Illinois is unclear. Several could have improved environmental conditions for Banded Killifish and facilitated population increases, even if they were not the root cause of the increase. One reason it is difficult to determine which factors may have contributed to the range expansion is because many of the Banded Killifish in Illinois today do not appear to be the same subspecies of Banded Killifish present in Illinois in the past.

Prior to 1989 Banded Killifish in Illinois appeared to belong to the Western subspecies. Recent records (1989-today) are sparse because it is listed as state-threatened in Illinois, hence collection of vouchers is strongly discouraged and regulated by permits. The few specimens that have been preserved recently in museum collections have lateral row scale counts intermediate between the Western subspecies and the Eastern subspecies. This is a common introgression pattern among fishes, with hybrids often intermediate between the two parent species (Hubbs, 1955).

Hybrids have been documented between Banded Killifish subspecies. Hubbs and Lagler (1947) based on data from Shapiro (1947) reported an intergrade zone between the two subspecies in Lake Ontario and the St. Lawrence River. The thought was the subspecies occupied separate refugia during the last glaciation, with the Eastern subspecies occupying an area along the Atlantic coast and the Western subspecies in the Mississippi River (Bailey and Smith, 1981). As the glaciers retreated, each subspecies expanded their respective ranges until they made secondary contact and introgressed. Genetic patterns among Banded Killifish populations along the St. Lawrence are consistent with this history (Rey and Turgeon, 2007). This theory was validated by April and Turgeon (2006) who analyzed genetics and morphology. However, they found the 'hybrid zone' to extend into Ohio along Lake Erie, farther than people realized. The western edge of this 'hybrid zone' was identified by mitochondrial DNA from the Eastern subspecies introgressing into the Western subspecies. Farther east, the morphology and nuclear DNA were intermediate between parent subspecies.

Part of the explanation for the biogeographic patterns evident in Illinois today may be that Eastern Banded Killifish or hybrids somehow made it to southern Lake Michigan and spread from there. This is consistent with identifications based on lateral row scales and the westward expansion pattern documented by April and Turgeon (2006) in the eastern Great Lakes. As to why these killifish are expanding their range now, when the Western Banded Killifish previously present in Illinois did not, is unclear. Trautman (1981) reported Eastern Banded Killifish appeared to be more tolerant of pollution and were not as reliant on vegetation for spawning; therefore, it may be that they are better suited to the prevalent habitats in the region than are the original Western Banded Killifish. The native range of Eastern Banded Killifish also extends farther south than that of the Western Banded Killifish. It may be that increasing water temperatures due to climate change are also benefiting these hybrids if Eastern Banded Killifish tolerances are expressed (Hayhoe et al, 2010; Gronewold and Stow, 2014).

These scenarios may explain the increase in the Lake Michigan-Illinois River population and the relative stasis of the glacial lakes population, but they do not explain the appearance and subsequent expansion of the Mississippi River population. There were pre-existing Banded Killifish populations in the Mississippi watershed, notably central Wisconsin, Minnesota, and northwestern Iowa (Becker, 1983; Page and Burr, 2011). Each of these is 160+ km away from the Illinois records in the Mississippi. The largest Illinois population appears to be in Pool 19. This impoundment is heavily vegetated with very low flow (Tazik et al., 1993), acting more like a lake than a river. This is consistent with the habitat preferences of the species. Although, it is possible there was a downstream colonization event from Wisconsin, Minnesota, or Iowa, perhaps associated with recent flooding, we have no intervening records between these upstream populations and the recent Illinois population to support this scenario.

There is an aquatic connection between the Illinois River watershed and the Mississippi watershed via the Hennepin Canal. However, the canal has been defunct for years and it would have been difficult for the fishes to move through the multiple closed locks. Furthermore, the Lake Michigan-Illinois River individuals are putative hybrids, whereas most of the Mississippi River individuals appear to be Western Banded Killifish.

Another possible explanation for the Mississippi River population is an accidental or intentional release by someone. There are documented occurrences of people moving killifish between waterbodies [e.g., western Pennsylvania (Raney, 1938) and in Newfoundland (Mitchell and Purchase, 2014)] but no specific evidence surrounding this possibility for Banded Killifish in the Midwest.

There is still a considerable amount of ambiguity surrounding this rapidly changing range of Banded Killifish in Illinois. Genetic data are required to develop a clearer picture. Due to the potential for directional introgression, mitochondrial and nuclear markers would be necessary. It would even be beneficial to incorporate microsatellites to analyze finer biogeographic patterns within this rapid range expansion, as well as to determine potential source populations.

Nonetheless, there are already important management implications based on the findings reported in this publication. First, in Illinois the listing of a species as state-threatened or state-endangered is decided at the species level. Subspecies are not considered as separate taxonomic units. Hence it does not matter if the Banded Killifish in Illinois are of the Western subspecies, Eastern subspecies, or hybrids. They are all considered equally in determining listing status.

Second, there appear to be at least three management units, each with its particular history, requirements, and potential future trajectories. One is the Lake Michigan to Illinois River population that is made up of putative hybrids. This population could conceivably be divided into sub-populations (e.g., Illinois River, Lake Michigan, etc.). The second is the recent Mississippi River population that continues to expand. The third is the glacial lakes population. There is no indication the rapid expansion occurring elsewhere in the state has reached this region along the Wisconsin border. Expansion here would require movement of fish past dams in the Fox and Des Plaines rivers, as well as their tributaries. Additionally, the connections that occurred between glacial lakes and streams of this area have been disrupted due to development. For the time being, this area appears to be isolated and may be the last remaining original stock of Western Banded Killifish in the state.

One implication of there being different Banded Killifish management units involves stocking considerations. There have been two recent introductions of Banded Killifish authorized by the Illinois Department of Natural Resources--Prairie Crossing in 1998 and the West Branch DuPage River in 2010 mentioned above. Each involved stock populations from glacial lakes, and both of the introduced populations were close to the source populations in northeast Illinois. The Prairie Crossing appears to have worked (Schaeffer et al., 2012; Bland, 2013), whereas there is no evidence to date that the West Branch DuPage River has worked. In regards to future translocations and to preserve original genetics, there needs to be increased scrutiny regarding source populations and stocking locations.

A third management issue is the potential ecological impacts, especially in areas where Banded Killifish had not been found in the past. For example further expansion throughout the Mississippi River system could potentially lead to "unnatural" competitive pressures on species that have not encountered this killifish before. Close monitoring of the expansion is needed.

Determining whether the range expansion represents dramatic recovery or an invasive species is more complicated than originally realized, and it depends upon the taxonomic level used to address the question. At the species level, Banded Killifish as a whole was stable in Illinois, possibly declining slowly, until recent events. From that perspective this is a dramatic recovery. At the subspecies level, if Illinois is on the forefront of the westward expansion of the Eastern Banded Killifish, then it could be considered an invasion, as there are no historical records for Banded Killifish for the Illinois portion of the Mississippi River. This could also be considered an invasion if human intervention is involved, even though this population appears to be from regional stock.

Given current trends, Banded Killifish may continue to expand its range. This expansion could incorporate other states in the region, most notably Indiana, Wisconsin, Iowa, and Missouri. A larger regional view will be needed to determine where the various taxonomic subunits originated and where they may be spreading. Although Banded Killifish historically split into two subspecies that were for a while evolving along their own independent trajectories to potentially becoming full distinct species in the future, they may now be gradually recombining back into a single species whose distribution could be continuing to expand.

Acknowledgments.--Access lo preserved voucher specimens was granted by Susan Mochel (The Field Museum), Kevin Swagel (The Field Museum), Caleb McMahan (The Field Museum), Chris Taylor (Illinois Natural History Survey), and Dan Wylie (Illinois Natural History Survey). Significant amounts of locality data were also provided by many staff associated with the Illinois Department of Natural Resources--Aquatic Nuisance Species Program and Lake Michigan Program annual beach surveys. Brian Zimmerman (Ohio State University--Museum of Biological Diversity) raised the potential that Eastern Banded Killifish could be in Illinois. Charles Knapp (Shedd Aquarium) and Rebecca Gericke (Shedd Aquarium) provided comments on drafts of the manuscript. Fieldwork was partially supported by Illinois Department of Natural Resources State Wildlife Grant # T-106-R-1 / United States Fish and Wildlife Service Grant # F15AF01082 and Shedd Aquarium.


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Submitted 8 September 2017

Accepted 8 January 2018

Philip W. Willink (1)

Daniel P. Haerther Center for Conservation and Research, Shedd Aquarium 1200 South Lake Shore Drive, Chicago, Illinois 60605

Tristan A. Widloe (2)

Aquatic Nuisance Species Program, Illinois Department of Natural Resources, 5931 Pox River Drive, Plano 60545

Victor J. Santucci Jr. (3) and Daniel Makauskas (4)

Lake Michigan Program, Illinois Department of Natural Resources, 9511 West Harrison Street, Des Plaines 60016

Jeremy S. Tiemann (5)

Illinois Natural History Survey, Prairie Research institute, University of Illinois at Urbana-Champaign, 1816 South Oak Street, Champaign 61820

Samantha D. Hertel (6)

Loyola University Chicago, Department of Biology, 1032 West Sheridan Road, Chicago, Illinois 60626

Daniel P. Haerther Center Jar Conservation and Research, Shedd Aquarium, 1200 South Lake Shore Drive,

Chicago, Illinois 60605

James T. Lamer (7)

Department of Biological Sciences, Western Illinois University, 372 Waggoner Hall, I University Circle, Macomb 61455


Joshua L. Sherwood (8)

Illinois Natural History Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, 1816 South Oak Street, Champaign 61820

(1) Corresponding author: e-mail:

(2) e-mail:

(3) e-mail:

(4) e-mail:

(5) e-mail:

(6) e-mail:

(7) e-mail:

(8) e-mail:

Caption Fig. 1.--Distribution of Banded Killifish Fundulus diaphanus in Illinois divided into two time periods

Caption Fig. 2.--Relative abundance (mean number of fish/seine haul) of Banded Killifish Fundulus diaphanus based on beach seine sampling at five locations along the Lake Michigan shoreline in Illinois, 1979-2016

Caption Fig. 3.--Lateral row scales of Banded Killifish Fundulus diaphanus from different regions and different time periods. Sample sizes are F. d. menona (N = 62), F. d. diaphanus (N = 7), Mississippi watershed (N = 4), and Calumet Region (N = 7)
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Article Details
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Author:Willink, Philip W.; Widloe, Tristan A.; Santucci, Victor J., Jr.; Makauskas, Daniel; Tiemann, Jeremy
Publication:The American Midland Naturalist
Article Type:Report
Geographic Code:1U3IL
Date:Apr 1, 2018
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