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Rapid Expansion of Banded Killifish Fundulus
diaphanus across Northern Illinois: Dramatic
Recovery or Invasive Species?
Author(s): Philip W. Willink, Tristan A. Widloe, Victor J. Santucci
Jr., Daniel Makauskas, Jeremy S. Tiemann, Samantha D. Hertel, ,
James T. Lamer, and Joshua L. Sherwood
Source: The American Midland Naturalist, 179(2):179-190.
Published By: University of Notre Dame
https://doi.org/10.1674/0003-0031-179.2.179
URL: http://www.bioone.org/doi/full/10.1674/0003-0031-179.2.179
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Am. Midl. Nat. (2018) 179:179–190
Rapid Expansion of Banded Killifish Fundulus diaphanus
across Northern Illinois: Dramatic Recovery or Invasive
Species?
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 Fox 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 for 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, 1 University Circle, Macomb
61455
AND
Joshua L. Sherwood
8
Illinois Natural History Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, 1816
South Oak Street, Champaign 61820
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. Starting 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
1
Corresponding author: e-mail: pwillink@sheddaquarium.org
2
e-mail: Tristan.Widloe@illinois.gov
3
e-mail: Vic.Santucci@illinois.gov
4
e-mail: Dan.Makauskas@illinois.gov
5
e-mail: jtiemann@illinois.edu
6
e-mail: shertel@luc.edu
7
e-mail: jt-lamer@wiu.edu
8
e-mail: jsherwo2@illinois.edu
179
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 Lake 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 160þkilometers 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.
INTRODUCTION
Rapid changes in a species’ distribution involving the increasing range of an invasive
species (e.g., Irons et al., 2006; Kolar et 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 the 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 the Mummichog Fundulus heteroclitus (Weed, 1921;
Hubbs et al., 1943; Griffith, 1968, 1972; Fritz and Garside, 1974; Dawley, 1992; Dawley et al.,
1999, 2000; Herna´ndez Cha´vez and Turgeon, 2007; M´erette 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
THE AMERICAN MIDLAND NATURALIST180 179(2)
[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 130þy. 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; Herna´ndez Cha´vez 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
Herna´ndez Cha´vez and Turgeon (2007). The literature values were included because only a
few museum specimens of F. d. diaphanus were readily available, and Herna´ndez Cha´vez and
Turgeon (2007) corroborated their results with genetic data.
RESULTS
DISTRIBUTION
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
2018 181WILLINK ET AL.: DRAMATIC RECOVERY OR INVASIVE SPECIES?
FIG. 1.—Distribution of Banded Killifish Fundulus diaphanus in Illinois divided into two time periods
THE AMERICAN MIDLAND NATURALIST182 179(2)
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
Stricker, Iowa Department of Natural Resources, pers. comm.), which is opposite Rock
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
2018 183WILLINK ET AL.: DRAMATIC RECOVERY OR INVASIVE SPECIES?
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 ¼62) (Fig. 3). Eastern Banded Killifish had 48–51 lateral row
scales (n ¼7).
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 ¼7) (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;
Herna´ndez Cha´vez 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 ¼23).
Specimens collected in the Mississippi River proper in 2013 had 39 and 44 lateral row
scales, which identify them as Western Banded Killifish (n ¼2). Specimens collected in Coon
Creek (the most upstream Rock River locality) in 2013 had 42 and 47 lateral row scales (n ¼
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.
FIG. 3.—Lateral row scales of Banded Killifish Fundulus diaphanus from different regions and different
time periods. Sample sizes are F. d. menona (N ¼62), F. d. diaphanus (N ¼7), Mississippi watershed (N ¼
4), and Calumet Region (N ¼7)
THE AMERICAN MIDLAND NATURALIST184 179(2)
DISCUSSION
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 Ni˜
no 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
2018 185WILLINK ET AL.: DRAMATIC RECOVERY OR INVASIVE SPECIES?
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
160þkm 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.
THE AMERICAN MIDLAND NATURALIST186 179(2)
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.
2018 187WILLINK ET AL.: DRAMATIC RECOVERY OR INVASIVE SPECIES?
Acknowledgments.—Access to 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.
LITERATURE CITED
APRIL,J.AND J. TURGEON. 2006. Phylogeography of the Banded Killifish (Fundulus diaphanus): glacial races
and secondary contact. J. Fish Biol., 69(Supplement B):212–228.
BAILEY,R.M.AND G. R. SMITH. 1981. Origin and geography of the fish fauna of the Laurentian Great
Lakes basin. Can. J. Fish. Aquat. Sci., 38:1539–1561.
BECKER, G. C. 1983. Fishes of Wisconsin. The University of Wisconsin Press, Madison, Wisconsin. 1052 p.
BLAND, J. 2013. How do you spell success? The rare fish variety, that is. Am. Cur., 38(4):11–22.
BUNNELL, D. B., C. P. MADENJIAN,T.J.DESORCIE,M.J.KOSTICH,W.WOELMER,AND J. V. ADAMS. 2015. Status
and trends of prey fish populations in Lake Michigan, 2014. p 33–48. In: Compiled Reports to
the Great Lakes Fishery Commission of the Annual Bottom Trawl and Acoustics Surveys, 2014.
United States Geological Survey, Great Lake Science Center, Ann Arbor, Michigan.
DAWLEY, R. M. 1992. Clonal hybrids of the common laboratory fish Fundulus heteroclitus.P. Natl. Acad. Sci.
U.S.A.,89:2485–2488.
———, D. CHRZANOWSKI,K.L.PHIEL,K.BEAULIEU,AND K. A. GODDARD. 1999. Allozyme analysis of clonal
diversity among unisexual hybrids of the killifishes Fundulus diaphanus and F. heteroclitus.J.
Hered., 90:607–612.
———, A. M. YEAKEL,K.A.BEAULIEU,AND K. L. PHIEL. 2000. Histocompatibility analysis of clonal diversity
in unisexual hybrids of the killifishes Fundulus heteroclitus and Fundulus diaphanus.Can. J. Zool.,
78:923–930.
FORBES,S.A.AND R. E. RICHARDSON. 1919. Maps showing distribution of Illinois fishes to accompany a
report on the fishes of Illinois, 2nd ed. Illinois Natural History Survey, Champaign. 212 p.
——— AND ———. 1920. The fishes of Illinois, 2nd ed. Illinois Natural History Survey, Champaign. 357
p.
FRITZ,E.S.AND E. T. GARSIDE. 1974. Identification and description of hybrids of Fundulus heteroclitus and
F. diaphanus (Pisces: Cyprinodontidae) from Porters Lake, Nova Scotia, with evidence for
absence of backcrossing. Can. J. Zool., 52:1433–1442.
GREENFIELD,D.W.AND J. D. ROGNER. 1984. An assessment of the fish fauna of Lake Calumet and its
adjacent wetlands, Chicago, Illinois: past, present, and future. Trans. Ill. State Acad. Sci., 77:77–
93.
GRIFFITH, R. 1968. Analysis of serum protein in the Teleost genus Fundulus.Am. Zool., 8:806.
———. 1972. Studies on the physiology and evolution of killifishes of the genus Fundulus. Ph.D.
Dissertation, Yale University, New Haven, Connecticut.
GRONEWOLD,A.D.AND C. A. STOW. 2014. Water loss from the Great Lakes. Science,343:1084–1085.
HAYHOE, K., J. VANDORN,T.CROLEY II, N. SCHLEGAL,AND D. WUEBBLES. 2010. Regional climate change
projections for Chicago and the US Great Lakes. J. Great Lakes Res., 36:7–21.
HERNA
´NDEZ CHA
´VEZ,C.AND J. TURGEON. 2007. Asexual and sexual hybrids between Fundulus diaphanus and
F. heteroclitus in the Canadian Atlantic region. Mol. Ecol., 16:1467–1480.
HRABIK, R. A. 2016. Fish species found along Missouri’s border new to state. Mo. Cons., 77(3):7.
HUBBS, C. L. 1955. Hybridization between fish species in nature. Syst. Zool., 4:1–20.
——— AND K. F. LAGLER. 1947. Fishes of the Great Lakes region. Cran.Inst. Sci., 26:1–186.
THE AMERICAN MIDLAND NATURALIST188 179(2)
———, B. W. WALKER,AND R. E. JOHNSON. 1943. Hybridization in nature between species of American
Cyprinodont fishes. Con. Lab. Vert. Biol. Univ. Mich., 23:1–21.
ILLINOIS DEPARTMENT OF NATURAL RESOURCES. 2016. Illinois wildlife action plan: 2015 implementation
guide. Illinois Department of Natural Resources, Springfield. 292 p.
IRONS, K. S., M. A. MCCLELLAND,AND M. A. PEGG. 2006. Expansion of Round Goby in the Illinois waterway.
Am. Midl. Nat., 156:198–200.
JORDAN, D. S. 1878. A catalogue of the fishes of Illinois. Bull. Il. St. Lab. Nat. Hist., 1:37–70.
KOLAR, C. S., D. C. CHAPMAN,W.R.COURTENAY JR., C. M. HOUSEL,J.D.WILLIAMS,AND D. P. JENNINGS. 2007.
Bigheaded carps: a biological synopsis and environmental risk assessment. Am. Fish. Soc. Sp.
Pub., 33:1–204.
LAMER, J., E. RATCLIFF,AND K. RIVERA. 2016. Banded Killifish in the Upper Mississippi River. Up. Mi. Riv.
Cons. Comm. N., 2016(Summer):2.
MADENJIAN, C. P., R. O’GORMAN,D.B.BUNNELL,R.L.ARGYLE,E.F.ROSEMAN,D.M.WARNER,J.D.STOCKWELL,
AND M. A. STAPANIAN. 2008. Adverse effects of Alewives on Laurentian Great Lakes fish
communities. N.Am. J. Fish. Manage., 28:263–282.
MANKOWSKI, A. 2012. The Illinois endangered species protection act at forty: a review of the act’s
provisions and the Illinois list of endangered and threatened species. Illinois Endangered
Species Protection Board, Springfield. 152 p.
MEEK,S.E.AND S. F. HILDEBRAND. 1910. A syntopic list of the fishes known to occur within fifty miles of
Chicago. Publ. Field Mus. Nat. Hist. Zool. Ser., 142:223–338.
ME
´RETTE, D., A. BOURRET,AND J. TURGEON. 2009. Diversity and distribution of gynogenetic hybrids between
Fundulus diaphanus and Fundulus heteroclitus in Porter’s Lake (Nova Scotia) in relation to
salinity. J. Fish Biol., 75:1115–1122.
METZKE, B. A., L. C. HINZ JR., AND A. C. HULIN. 2012. Status revision and update for Illinois’ fish species in
greatest need of conservation. Ill. Nat. Hist. Sur.Tech. Rep., 2012(19):1–179.
MITCHELL,J.S.AND C. F. PURCHASE. 2014. Rapid colonization of a species at risk: a new eastern range limit
for Fundulus diaphanus (Banded Killifish), in Newfoundland. Northeast.Nat., 21:41–44.
NALEPA, T. F., D. W. SCHLOESSER,S.A.POTHOVEN,D.W.HONDORP,D.L.FANSLOW,M.L.TUCHMAN,AND G. W.
FLEISCHER. 2001. First finding of the amphipod Echinogammarus ischnus and the mussel Dreissena
bugensis in Lake Michigan. J. Great Lakes Res., 27:384–391.
NELSON, E. W. 1876. A partial catalogue of the fishes of Illinois. Bull. Il.Mus. Nat. Hist., 1:33–52.
PAGE,L.M.AND B. M. BURR. 2011. Peterson field guide to freshwater fishes of North America north of
Mexico. Houghton Mifflin Harcourt, Boston. 663 p.
RANEY, E. C. 1938. The distribution of the fishes of the Ohio drainage basin of western Pennsylvania.
Ph.D. dissertation, Cornell University. 102 p.
RASMUSSEN, P. W., C. SCHRANK,AND M. C. W. WILLIAMS. 2014. Trends in PCB concentrations in Lake
Michigan Coho and Chinook Salmon, 1975–2010. J. Great Lakes Res., 40:748–754.
RETZER,M.E.AND B. BATTEN. 2005. Fishes of the Chicago Region: a review of the Dennison and Illinois
Natural History Survey collections. Trans. Ill. State Acad. Sci., 98:63–73.
REY,O.AND J. TURGEON. 2007. Influence of historical events and contemporary estuarine circulation on
the genetic structure of the Banded Killifish (Fundulus diaphanus) in the St. Lawrence River
(Quebec, Canada). Can. J. Zool., 85:891–901.
RICHARDSON, L. R. 1939. The spawning behavior of Fundulus diaphanus (Le Sueur). Copeia,1939:165–167.
RIVERA, K. D., R. L. HAUN,C.A.ANDERSON,AND S. P. ROMANO. 2013. New distribution record for Fundulus
diaphanous (LeSueur), family Fundulidae in Illinois. Trans. Ill. State Acad. Sci., 106:57.
SABAJ, M. H. 2016. Standard symbolic codes for institutional resource collections in herpetology and
ichthyology: an online reference. Version 6.5 (16 August 2016). Electronically accessible at
http://www.asih.org/, American Society of Ichthyologists and Herpetologists, Washington, DC.
SCHAEFFER, J. S., J. K. BLAND,AND J. JANSSEN. 2012. Use of a storm water retention system for conservation of
regionally endangered fish. Fisheries,37:66–75.
SCHMIDT, K. 2016. Iowa’s Longear Sunfish mystery solved...maybe—Addendum. Am. Cur., 41(3):17–18.
SCOTT,W.B.AND E. J. CROSSMAN. 1973. Freshwater fishes of Canada. Bull. Fish. Res. Bd.Can., 184:1–966.
2018 189WILLINK ET AL.: DRAMATIC RECOVERY OR INVASIVE SPECIES?
SHAPIRO, S. 1947. Geographic variation in Fundulus diaphanus, a cyprinodontid fish. Ph.D. dissertation,
University of Michigan, Ann Arbor.
SIETMAN, B. E., S. D. WHITNEY,D.E.KELNER,K.D.BLODGETT,AND H. L. DUNN. 2001. Post-extirpation
recovery of the freshwater mussel (Bivalvia: Unionidae) fauna in the upper Illinois River. J.
Freshwater Ecol., 16:273–281.
SMITH, P. W. 1979. The fishes of Illinois. University of Illinois, Champaign. 314 pages.
STARRETT, W. C. 1971. A survey of the mussels (Unionacea) of the Illinois River: a polluted stream. Il. Nat.
Hist. Sur.Bull., 30:267–403.
TAZIK, P. P., R. V. ANDERSON,AND D. M. DAY. 1993. The development of an aquatic vegetation community
in Pool 19, Upper Mississippi River. J. Freshwater Ecol., 8:19–26.
TIEMANN, J. S., C. A. TAYLOR,D.WYLIE,J.LAMER,P.W.WILLINK,F.M.VERALDI,S.M.PESCITELLI,B.LUBINSKI,
T. THOMAS,R.SAUER,AND B. CANTRELL. 2015. Range expansions and new drainage records for
select Illinois Fishes. Trans. Ill. State Acad. Sci., 108:47–52.
TRAUTMAN, M. B. 1981. The fishes of Ohio. Ohio State University Press and Ohio Sea Grant Program
Center for Lake Erie Area Research, Columbus. 782 p.
WARNER, D. M., S. A. FARHA,R.M.CLARAMUNT,D.HANSON,AND T. P. O’BRIEN. 2015. Status of pelagic prey
fishes in Lake Michigan, 2014. p 47–58. In: Compiled Reports to the Great Lakes Fishery
Commission of the Annual Bottom Trawl and Acoustics Surveys, 2014. United States Geological
Survey, Great Lake Science Center, Ann Arbor, Michigan.
WEED, A. C. 1921. Fishes from Southern Wisconsin. Copeia,99:69–72.
WILLINK, P. W. 2009. A century of shifting fish assemblages in Wolf Lake, Illinois-Indiana. P. In.Acad. Sci.,
118:187–195.
——— AND F. M. VERALDI. 2009. The fishes of Will County, Illinois. Fieldiana Zool. N. S., 115:1–61.
SUBMITTED 8SEPTEMBER 2017 ACCEPTED 8JANUARY 2018
THE AMERICAN MIDLAND NATURALIST190 179(2)