Rapid Colonization of the Potomac River Drainage by
the Rainbow Darter (Etheostoma caeruleum) Following
Introduction
Jason F. Cessna, Richard L. Raesly, Jay V. Kilian, Daniel A. Cincotta, and Robert H. Hilderbrand
Northeastern Naturalist, Volume 21, Issue 1 (2014): 1–11
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J.F. Cessna, R.L. Raesly, J.V. Kilian, D.A. Cincotta, and R.H. Hilderbrand
2014
1
2014 NORTHEASTERN NATURALIST 21(1):1–11
Rapid Colonization of the Potomac River Drainage by
the Rainbow Darter (Etheostoma caeruleum) Following
Introduction
Jason F. Cessna1,*, Richard L. Raesly2, Jay V. Kilian3, Daniel A. Cincotta4,
and Robert H. Hilderbrand1
Abstract - Etheostoma caeruleum (Rainbow Darter) was first collected from the Potomac
River drainage in 1977 and has subsequently expanded its range to include nearly all
major tributaries in the system. Analysis of the chronology of Rainbow Darter collection
data demonstrates the explosive nature of this range expansion. Using this chronology, we
assert that Rainbow Darter is not native to the drainage and likely occurs as a result of human-
induced interbasin transfer. Although the ecological impact of non-native Rainbow
Darters in the Potomac is unknown, the potential exists for competition with native species.
The presence of numerous instream blockages prohibiting colonization of portions
of the drainage make this system well suited to study the impacts of introduced darters on
native ichthyofauna.
Introduction
Etheostoma caeruleum Storer (Rainbow Darter) (Teleostei: Percidae) is a
small benthic fish, widely distributed in streams and small rivers in central North
America and generally known from the Mississippi River and Great Lakes drainages.
The species was not known historically from Atlantic Slope drainages, but
now occurs in the Hudson and Potomac rivers (Fuller and Neilson 2013, Stauffer
et al. 1978). The recent appearance of Rainbow Darters in these systems presents
an opportunity to observe factors influencing the dispersal and range expansion of
small benthic fishes.
Introduced darters have demonstrated rapid range expansion in several North
American river drainages, including the Genesee, Susquehanna, Potomac, New, and
Colorado rivers (Jenkins and Burkhead 1994, Neely and George 2006, Raesly et al.
1990, Smith 1985, Walford and Bestgen 2008). In these systems, darters have colonized
the majority of the drainage within a few decades of introduction. For example,
Etheostoma blennioides Rafinesque (Greenside Darter) was first collected from the
Susquehanna River drainage in 1962 and had colonized most of the drainage by 2005
(Neely and George 2006). Similarly, the Rainbow Darter was first collected from the
Genesee River drainage in 1953 (Carlson and Daniels 2004), where a basin-wide
1University of Maryland Center for Environmental Sciences, Appalachian Laboratory, 301
Braddock Road, Frostburg, MD 21532. 2Frostburg State University, Department of Biology,
101 Braddock Road, Frostburg, MD 21532. 3Maryland Department of Natural Resources,
580 Taylor Avenue C-2, Annapolis, MD 21401. 4West Virginia Division of Natural Resources,
Wildlife Resources Section, PO Box 67, Elkins, WV 26241. *Corresponding author
- Jcessna@umces.edu.
Manuscript Editor: Jay Stauffer
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J.F. Cessna, R.L. Raesly, J.V. Kilian, D.A. Cincotta, and R.H. Hilderbrand
2014 Vol. 21, No. 1
survey in 1926 had not revealed its presence (Moore 1927). By the early 1980s, the
Rainbow Darter was considered to be in a stage of explosive range expansion in the
Genesee and was commonly captured in parts of the drainage (Smith 1985).
Stauffer et al. (1978) first collected Rainbow Darters in the upper South Branch
Potomac River in 1977. These authors suggested this species was native and had
apparently been overlooked by previous collectors due to its isolated location. Esmond
and Stauffer (1983) presented morphological and meristic data in support of
this indigenous notion. Since this time, this darter has greatly expanded in abundance
and distribution. The extent to which the Potomac system had previously
been sampled without collecting this species (Davis 1974, Jenkins and Burkhead
1994, Lee et al. 1976, Manseuti 1957, Schwartz 1965, Truitt et al. 1929) raises uncertainty
as to whether Rainbow Darter is native to the drainage or occurs as a result
of interbasin transfer. Due to its rapid spread, regional investigators have doubted
the indigenous status of Rainbow Darters in the basin, but few have addressed this
speculation in the literature (Christmas et al. 1998).
Herein we review the current status and distribution of Rainbow Darter in the Potomac
drainage. Prior to this study, the distribution and range expansion of the species
within this river system have not been well documented. We use the dispersal behavior
of the species and historical sampling data to suggest that Rainbow Darter should
not be considered part of the native ichthyofauna of the Potomac River.
Methods
We sampled 54 localities in Pennsylvania, Maryland, and Virginia between
2010 and 2012 for the presence of Rainbow Darters. We sampled using a backpack
electrofisher for a minimum of 600 seconds beyond the final new species
detection at each site, to ensure a representative sampling of all fishes.
We queried numerous sources for Rainbow Darter records, including regional
ichthyologists, unpublished survey reports, and the scientific literature, as well as
the following state and federal agencies: The West Virginia Department of Natural
Resources (WVDNR), the Maryland Department of Natural Resources Maryland
Biological Stream Survey (MDNR MBSS), the Virginia Department of Game and
Inland Fisheries (VDGIF), the Pennsylvania Department of Environmental Protection
(PADEP), the Pennsylvania Fish and Boat Commission (PFBC), the US
Environmental Protection Agency Mid-Atlantic Integrated Assessment Program
(EPA MAIA), the US Geological Survey National Water Quality Assessment Program
(USGS NAWQA), and the National Park Service Inventory and Monitoring
Program (NPS IMP). Records were considered valid if corroborated by additional
nearby collections or the presence of preserved specimens.
We used Arc GIS to spatially and temporally analyze the point-locality data. To
generate a temporal chronology of Rainbow Darter range expansion, we divided
observations into four time intervals from the date of initial discovery: 0–10 years,
11–20 years, 21–30 years, and 31–36 years. We also analyzed the full extent of colonization
across temporal intervals in Sideling Hill Creek, a fourth-order tributary to
the upper Potomac.
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J.F. Cessna, R.L. Raesly, J.V. Kilian, D.A. Cincotta, and R.H. Hilderbrand
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Results and Discussion
We obtained 202 accounts of Rainbow Darter in the Potomac River drainage
(Table 1). Three records have been previously noted in the scientific literature (Esmond
and Stauffer 1983; Stauffer et al. 1978, 1995), 174 existed in unpublished
sources, and 25 are new records arising from our electrofishing efforts. One record
obtained from VDGIF was discounted (Quantico Creek) because we could not collect
specimens from there or nearby streams.
Stauffer et al. (1978) collected the first Rainbow Darters from the Potomac River
drainage in 1977, from the North Fork and upper South Branch of the Potomac
River, during a state-wide survey of West Virginia. Once established, Rainbow
Darters expanded rapidly. Within 25 years of initial detection, this darter’s range
expanded over 400 river kilometers (rkm) downstream and had been collected from
the lower reaches of every major unimpounded tributary in the system.
The WVDNR documented expansion into the South Fork of the South Branch
by 1982, some 40 rkm from the initial detection. The following year, this agency
collected the species 110 rkm downstream of the initial detection, only 25 rkm from
the South Branch’s confluence with the North Branch (Fig. 1). Several more collections
of the species were made in the upper South Branch in the subsequent few
years. In 1994, Bennett (1997) collected Rainbow Darter in Sideling Hill Creek,
a tributary to the middle Potomac, 212 rkm from the point of initial discovery. In
1997, Raesly (1998) documented it much further downstream, near the mouths
of Israel Creek, Catoctin Creek, Little Monocacy River, and Broad Run in Maryland
(Fig. 2). That same year, MDNR MBSS quantitatively sampled 3 of these 4
tributaries less than 3 rkm upstream of Raesly’s sites without collecting Rainbow
Darter, suggesting that the establishment of the species in these tributaries was
quite recent. Additionally, VDGIF documented a 1998 record from Goose Creek in
northern Virginia. The Goose Creek and Broad Run records are approximately 400
rkm from the initial Stauffer et al. (1978) collection and represented the furthest
downstream collections of the species until 2012.
Table 1. Sources of unpublished Rainbow Darter records.
Source Records
MDNR MBSS 75
WVDNR 46
Raesly et al. (2004) 31
MDNR collection permit database 7
EPA MAIA 6
Raesly (1998) 4
VDGIF 2
USGS NAWQA 2
NPS IMP 1
PADEP 1
Bennett (1997) 1
Jennings (1998) 1
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J.F. Cessna, R.L. Raesly, J.V. Kilian, D.A. Cincotta, and R.H. Hilderbrand
2014 Vol. 21, No. 1
By the mid-2000s, Rainbow Darter had expanded into many lower North Branch
and upper Potomac tributaries, including Evitts, Town, Licking, Conococheague,
and Antietam creeks in Maryland and Sleepy, Back, and Opequon creeks in the
Eastern Panhandle of West Virginia. Rainbow Darter also expanded into Mill and
Lunice creeks, tributaries to the upper South Branch (Fig. 2).
The mid-2000s through early 2010s has seen Rainbow Darter populations continuing
to colonize portions of the drainage where it is not blocked by dams (Fig. 3);
expanding into south-central Pennsylvania in Sideling Hill, Tonoloway, and Licking
creeks and into northern Virginia in Back, Opequon, and Catoctin creeks. In
2012, we documented Rainbow Darter in Cabin John Creek, a tributary just west
of the District of Columbia, a few kilometers upstream of the tidal interface. This
site represents the known extent of downstream range expansion and was quantitatively
sampled in 1995 by Starnes (2002), in 1997 by MDNR MBSS, and in 2004
by Raesly et al. (2004), all efforts that revealed no Rainbow Darters. At present,
the Rainbow Darter does not appear to have successfully colonized tributaries
downstream of the tidal interface. All major Maryland tributaries to the tidal-fresh
Potomac have been sampled by MDNR MBSS within the last 10 years, and we surveyed
many Virginia tidal-fresh tributaries in 2012. It is unknown if the Rainbow
Darter will be capable of colonizing habitats downstream of the tidal interface.
Figure 1. Records of Rainbow Darter in the Potomac River drainage from 1977 to 1986 and
from 1987 to 1996.
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However, introduced populations of the Greenside Darter have apparently been
unable to do so (Starnes et al. 2011).
Analysis of collection points in Sideling Hill Creek (PA and MD), a fourthorder
tributary to the upper Potomac, presents an interesting case study of Rainbow
Darter colonization of a watershed (Fig. 4). This watershed experienced sufficient
sampling intensity to document the extent of colonization at several temporal
junctures. Rainbow Darters were not collected in the system during basin-wide
sampling conducted by Lebo (1983). By 1994, Bennett (1997) documented the
presence of Rainbow Darters extending to 13.5 rkm upstream of the mouth of
Sideling Hill Creek. Jennings (1998) documented the extent of colonization an additional
8 rkm upstream in 1996. In 2002, PADEP collected this darter an additional
6 rkm upstream, very near localities that were sampled in 1985 by the Pennsylvania
Fish and Boat Commission and in 1994 by PADEP (Pennsylvania Department of
Environmental Resources 1994). In 2010, we documented this species in all major
tributaries to Sideling Hill Creek and found the extent of colonization to be an additional
22.5 rkm upstream—some 50 rkm upstream of the mouth. These values
correspond with a mean upstream colonization rate of 2.6 ± 1.5 rkm per year.
The Potomac drainage was extensively surveyed prior to 1977 with no mention
of Rainbow Darter (Lee et al. 1976, Schwartz 1965, Truitt et al. 1929, and references
therein). As populations continue to grow and spread, it is unlikely that a native
Figure 2. Records of Rainbow Darter in the Potomac River drainage from 1997 to 2006.
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2014 Vol. 21, No. 1
species could have remained undetected until 1977 and then become abundant and
widespread within a few decades. Stauffer et al. (1978) suggested that Rainbow
Darter was likely overlooked by previous collectors, and Esmond and Stauffer
(1983) suggested that this population was geographically isolated and taxonomically
distinct from populations in adjacent drainages, asserting that the Potomac
population was therefore native. However, by 1983, the year of the Esmond and
Stauffer (1983) publication, Rainbow Darter had already significantly extended its
range 110 rkm downstream in the South Branch. Additionally, Schwartz (1965)
shows a collection point within a few hundred meters of the 1977 collection, which
apparently yielded no Rainbow Darters.
We believe Rainbow Darter is not native to the Potomac and occurs as a
result of human-induced interbasin transfer; most likely a bait-bucket introduction.
Many studies have demonstrated the probablilty of introduction events
occurring via this vehicle (DiStefano et al. 2009, Kilian et al. 2012, LoVullo
and Stauffer 1993, Ludwig and Leitch 1996). Additionally, the first several collections
of this darter were made within a stocked trout fishery, where the use of
live bait is permitted.
The rapid spread of the Rainbow Darter in the Potomac drainage is characteristic
of darter invasions in other North American river systems, including the Susquehanna,
Genessee, New, and upper Colorado rivers (Jenkins and Burkhead 1994,
Figure 3. Records of Rainbow Darter in the Potomac River drainage from 2007 to 2012.
Northeastern Naturalist Vol. 21, No. 1
J.F. Cessna, R.L. Raesly, J.V. Kilian, D.A. Cincotta, and R.H. Hilderbrand
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Neely and George 2006, Raesly et al. 1990, Smith 1985, Walford and Bestgen
2008). The spread of the species throughout the drainage over the last 35 years and
the speed with which populations have grown makes it exceedingly unlikely that
isolated populations went undetected for decades. It is equally unlikely that such
growth is a result of pervasive changes in the ecology of the drainage.
The morphological and meristic distinctness of the Potomac River Rainbow
Darter populations found by Esmond and Stauffer (1983) may be a founder effect,
similar to that reported by Raelsy (1991). Raesly (1991) noted significant morphological
distinctness from nearby populations in a recently introduced population
of Etheostoma zonale (Cope) (Banded Darter) in the Susquehanna drainage, and
genetic analysis revealed a bottleneck consistent with a founder event. Recent
range-wide genetic analyses have been conducted for Rainbow Darter (Haponski
et al. 2009, Ray et al. 2006), but no studies have evaluated genetic material from
Potomac populations.
Figure 4. Rainbow Darter colonization of Sideling Hill Creek, a fourth-order tributary to
the upper Potomac River. Points represent the known extent of colonization in 1994, 1996,
2002, and 2010.
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J.F. Cessna, R.L. Raesly, J.V. Kilian, D.A. Cincotta, and R.H. Hilderbrand
2014 Vol. 21, No. 1
Multiple factors may have influenced the rapid dispersal of Rainbow Darter
in the Potomac drainage. In addition to introducing this species to the drainage,
humans may have relocated it within the drainage, particluarly as it became more
abundant. Basin-wide flooding likely also contributed to rapid downstream range
expansion. The flood of 1996 was the Potomac’s largest since 1972, and record
flood stages were exceeded in several localities. In the ensuing two years, Rainbow
Darter was documented in low abundances over 100 rkm further downstream than
previously known. Comparable range expansion after massive flooding has been
observed for introduced populations of both Banded Darter and Greenside Darter
in the adjacent Susquehanna River drainage (Denoncourt and Stauffer 1976, Neely
and George 2006, Raesly et al. 1990). Basin-wide flooding in 1985 may have
also contributed to downstream range expansion; however, only one collection of
Rainbow Darter was made between 1985 and 1992, which is insufficient sampling
intensity to document such expansion.
The Rainbow Darter is probably not the only darter recently introduced to the
drainage. The Greenside Darter was not collected in the Potomac until 1957 and
was also thought to be native (Lee et al. 1976, Miller 1968, Schwartz 1965, Stauffer
et al. 1978). However, its recent appearance and rapid range expansion are often cited
as evidence of introduction (Christmas et al. 1998, Jenkins and Burkhead 1994,
Starnes 2002, Starnes et al. 2011). Greenside Darter is now widespread throughout
the drainage, except the North Branch upstream of Cumberland, MD, where dams
have been maintained since the mid-1800s
Dams have also prevented Rainbow Darter colonization in several major portions
of the drainage (Fig. 3). Most notably, almost the entire 7600-km2 Shenandoah
River basin has been blocked by Millville Dam. Cacapon River has also apparently
been blocked by an old power-plant dam. Additionally, large portions of North
Branch, Evitts Creek, Town Creek, Conococheague Creek, Antietam Creek, and
Goose Creek have been blocked.
Darter species richness has been used in recent years as a favorable metric in
stream bioassessments. Since the presence of introduced darters may result in the
artificial inflation of indices of biological integrity (IBIs) when compared to localities
upstream of watershed impediments (e.g., waterfalls, dams), caution should be
used in aquatic community assessments. To avoid erroneous conclusions in darter
species richness metrics, we recommend that introduced darters be eliminated
from IBI calculations, and/or an inventory of stream blockages or connectivity be
incorporated into biological monitoring protocols. A complete inventory of stream
blockages in watersheds under evaluation would probably prevent an incorrect assessment
of darter and other species richness metrics. In Sideling Hill Creek, for
example, undocumented stream blockages have prevented Rainbow Darter colonization
in two of the four major tributaries (Fig. 4).
The ecological impact of non-native darters in the Potomac River drainage is unknown.
There exists potential for overlap in diet and habitat use between introduced
darters and other benthic fishes. Of particular concern are the remaining populations
of Percina notogramma (Raney and Hubbs) (Stripeback Darter), endangered
Northeastern Naturalist Vol. 21, No. 1
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in Maryland and known from only a handful of remaining localities in the Potomac
drainage. Given the limited data available on the diet of Stripeback Darter (Flemer
and Woolcott 1966) and the breadth of diet overlap of Rainbow Darter with other
sympatric species (Adamson and Wissing 1977, Hlohowskyj and White 1983, Paine
et al. 1982, van Snik Gray et al. 1997, Welsh and Perry 1998), there exists potential
for resource overlap. Competition with other benthic fishes is also possible. The numerous
locations where colonization by Rainbow Darter has been prevented by dams
reveals a convenient paired study design for observing changes in habitat utilization
among native assemblages of benthic fishes when a non-native is introduced.
Acknowledgments
We would like to thank the Pennsylvania Fish and Boat Commission, the Virginia Department
of Game and Inland Fisheries, and the Maryland Department of Natural Resources
for scientific collecting permits. Data were provided by many regional biologists: M. Pinder
provided VDGIF data; T. Shaw and B. Chalfant provided PA DEP data; M. Kashiwagi and R.
Bourquin provided MDNR data; D. Thorne provided WV DNR data; L. Reynolds provided
EPA MAIA data; and G. Sanders provided NPS IMP data. Additionally, G.K. Mack provided
a lead to the occurrence of Rainbow Darter in Cabin John Creek. J. Epstein, T. Miller, and
R. Rambo assisted with field collections.
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