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Article

Endemic and Invasive Species: A History of Distributional Trends in the Fish Fauna of the Lower New River Drainage

by
Stuart A. Welsh
1,*,
Daniel A. Cincotta
2,†,
Nathaniel V. Owens
2 and
Jay R. Stauffer, Jr.
3,4
1
U.S. Geological Survey, West Virginia Cooperative Fish and Wildlife Research Unit, West Virginia University, Morgantown, WV 26506, USA
2
West Virginia Division of Natural Resources, Elkins, WV 26241, USA
3
Ecosystem Science and Management, The Pennsylvania State University, 432 Forest Resources Building, University Park, PA 16802, USA
4
South African Institute for Aquatic Biodiversity, Makhanda 6140, South Africa
*
Author to whom correspondence should be addressed.
†
Retired.
Water 2025, 17(2), 221; https://doi.org/10.3390/w17020221
Submission received: 21 November 2024 / Revised: 7 January 2025 / Accepted: 13 January 2025 / Published: 15 January 2025
(This article belongs to the Section Water, Agriculture and Aquaculture)
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Abstract

Invasive species are often central to conservation efforts, particularly when concerns involve potential impacts on rare, endemic native species. The lower New River drainage of the eastern United States is a watershed that warrants conservation assessment, as the system is naturally depauperate of native fish species and it is nearly saturated with non-native fish species: there are 31 natives, including at least nine endemic taxa, and 63 non-natives. For endemic taxa, we examined temporal distribution shifts (range expansions or contractions) based on percent change in the occupied watershed area. We contrasted these findings with time series analyses on distribution trends of non-native minnows (Leuciscidae) and darters (Percidae) based on growth curve models of the cumulative sum of the total area of occupied 12-digit hydrologic unit codes. We documented range reductions for six of nine endemic taxa. We determined that 11 of 18 non-native minnows and 6 of 8 non-native darters were invasive based on range expansions and associated invasion curve models. The endemic taxa are of conservation concern given the limited distribution ranges and documented population declines. Although among-species comparisons of range shifts do not support causal inference, documentation of changes in distribution ranges of endemic and invasive species is critical to inform conservation efforts.

1. Introduction

Reductions in ranges and associated declines in populations of riverine fishes may lead to imperilment of species and are of global conservation concern [1,2,3]. An understanding of the anthropogenic causes of range reduction is seldomly straightforward, as fishes in aquatic systems are often influenced by multiple stressors [4,5,6,7,8]. Although habitat degradation of riverine systems has generally been considered as a leading cause of range reduction and imperilment [9,10], invasive species are also considered to be common contributors [11,12]. Causal factors of range reductions can be problematic to parse, as cumulative impacts are often attributed to both habitat degradation and invasive species [13,14,15,16,17].
A non-native species is considered to be invasive when a population becomes established and undergoes rapid expansion [18,19,20,21,22]. The stages of invasion often follow a sigmoidal growth curve with three defined phases: a lag period (zero to limited expansion), a period of exponential growth (rapid expansion), and an asymptotic period of reduced expansion [23,24]. The lag and exponential growth phases of the invasion curve have many influences, such as propagule pressure, life history or dispersal traits of species, traits of the community or environment, or interactions of species [25,26,27,28]. The lag period can be defined by the x-axis intercept (λ) of the tangent line to the inflection point of the curve [29] (Figure 1). For the exponential growth phase, the maximum growth rate is represented as the slope of the tangent line at the inflection point of the curve [30,31]. The asymptotic phase of the curve may represent a period of saturation, where an environmental carrying capacity is approached by the invading species [24]. Other influential points on the curve include the maximum acceleration point (MAP), maximum deceleration point (MDP), and the asymptotic deceleration point (ADP), all of which can be determined from roots of the curve’s third- and fourth-order derivatives (Figure 1). Additionally, our understanding of invasion curves, as determined by sigmoidal growth models fit to time series of species distribution data, will be influenced by the detection of species and other qualities of the input data [32,33].
Invasive species are often successful in riverine systems that are naturally depauperate of native species diversity [34,35,36], which may lead to homogenization of faunas [37,38]. Invasive species may increase the overall diversity within a watershed, but can lead to imperilment of native species through predation [39,40], competition [41,42,43], or hybridization [44,45,46]. Although impacts via hybridization can be clearly defined [12] other implied impacts are often anecdotal [47], owing to the correlative nature of observational field data. Further, invasive species impacts can be difficult to measure or separate from cumulative impacts of a suite of anthropogenic stressors, yet conservation agencies are often tasked with curtailment or management of the problem. An evaluation of the invasion curve provides insights into potential management strategies, as the lag period is the best time for removal or control measures. A paradox, however, is that to model an invasion curve requires that the species is already established and has undergone rapid range expansion; post invasion assessments can assist understanding of future invasive species [48].
Rare and endemic species, particularly those with narrow ranges, are often of conservation concern because of vulnerability to imperilment [49,50]. Endemic fishes that occupy watersheds with a high percentage of non-native or invasive species are particularly vulnerable [51]. The New River system of the eastern United States is an example of a watershed with a high percentage of both endemic and non-native species [52,53,54,55]. The lower New River system is naturally depauperate of native species and it is nearly saturated with non-native fish species: there are 31 natives, including at least nine endemic taxa, and 63 non-natives [55]. Endemic fishes include New River Shiner (Miniellus scabriceps), Bigmouth Chub (Nocomis platyrhynchus), Kanawha Minnow (Phenacobius teretulus), Bluestone Sculpin (Cottus cf. carolinae), Buckeye Creek Cave Sculpin (Cottus cf. carolinae), Kanawha Sculpin (Cottus kanawhae), Candy Darter (Etheostoma osburni), and Appalachia Darter (Percina gymnocephala). Additionally, the Rosyface Shiner (Notropis cf. rubellus) in the New River drainage has been recognized recently as an undescribed endemic species [56,57,58]. Tracy et al. [59] referred to this undescribed form as the “Kanawha” Rosyface Shiner. Given the relatively high proportion of both endemic and non-native species in the lower New River fish fauna, further evaluations of the temporal distribution ranges of these taxa are warranted for conservation assessment.
The study objectives were to (1) quantify the reduction or expansion in distribution ranges of endemic fish species and (2) model range expansions and invasion curves of non-native minnows (Leuciscidae) and darters (Percidae) in the lower New River drainage. The high percentage of both endemic species and non-native species make the New River drainage a model system for investigations into the ecology of invasive species. Our aim was not to determine causal factors of expansion or reduction in species’ ranges, but rather to document distributional changes in endemic and non-native species for use in conservation planning.

2. Materials and Methods

The New River drainage, which represents the upper reaches of the ancient Teays River, is delineated by Kanawha Falls, a well-documented zoogeographic barrier to fish distributions [51,60]. Kanawha Falls is located on the Kanawha River approximately 2.3 km downstream of the confluence of the Gauley and New Rivers in West Virginia, and zoogeographers have generally considered the drainage area upstream of Kanawha Falls as the New River system [52,61,62]. The New River drainage begins in North Carolina, flows through western Virginia, and ends in southern West Virginia. Drainage areas by state are approximately 1951 km2 (North Carolina), 7939 km2 (Virginia), and 11,944 km2 (West Virginia). The New River drainage of West Virginia, hereafter called the lower New River drainage, includes 115 12-digit hydrologic watershed units (HUC-12s) based on watershed boundaries of the National Watershed Boundary Dataset [63] (Figure 2). The mean area of HUC-12s is 103.5 km2 with a range of 43.6–245.0 km2. Tributaries of the lower New River system are the Bluestone, Gauley, and Greenbrier Rivers, including two smaller watersheds (East River and Indian Creek) near the West Virginia/Virginia border (Figure 2). The lower New River drainage includes three large dams: the Hawks Nest and Bluestone Dams are located on the New River approximately 13 and 108 km upstream of Kanawha Falls, respectively, and Summersville Dam is on the lower Gauley River approximately 56 km upstream of the Gauley/New River confluence. The Bluestone, Hawks Nest, and Summersville Dams, constructed during 1941–1949, the early 1930s, and 1960–1966, respectively, restrict upstream fish migration. Downstream fish passage is possible for the run-of-the-river reservoirs and associated spillway designs of the Bluestone and Hawks Nest Dams, but it is less likely for the deep, oligotrophic Summersville reservoir and the high-pressure bottom release of Summersville Dam. Additionally, Sandstone Falls of the New River (approximately 91 km upstream of Kanawha Falls) is a partial barrier to fish passage.
We examined species occurrence data at the HUC-12 watershed scale based on 2132 locality records from the lower New River drainage ranging from 1897 to 2023. Collection methods included electrofishing (70%), seining (25%), and using rotenone (5%). Most species occurrence data were temporally distributed from 1928 to 2023 (Figure 3). Periods of fish sampling lulls occurred during the mid-1930s to mid-1940s, 1980s, and 2020s. The species occurrence records, which are maintained by the West Virginia Division of Natural Resources (WVDNR), included data from a literature review, state and federal agency fish surveys (including surveys of the authors), and regional museum specimens. Data from the literature review included species occurrence records from multiple sources [52,64,65,66,67,68,69]. We confirmed species identification with museum records by visiting the American Museum of Natural History (AMNH), Cornell University (CU), North Carolina Museum of Natural Sciences (NCSM), Ohio State University Museum (OSUM), Pennsylvania State University (PSU), University of Michigan (UMMZ), Frostburg State University (USMF), the US National Museum (USNM), and the Wildlife Resources Section (WVWR collection) of the WVDNR. The origin status of species (native vs. non-native) was determined following the classification by Cincotta and Welsh [55].
For native endemic taxa, we examined temporal changes in distribution ranges through visual assessment of occurrence records and by calculating the percent change in the total area of occupied HUC-12s. A species’ total area of occupied HUC-12s was considered as a measure of the species’ extent of occurrence. Initially, we used ArcGIS Pro 3.3.0 to plot the extent of occurrence and visually depict range reductions in endemic taxa based on three time periods: ≤1960, >1960 and ≤1990, and >1990 and ≤2023. The number of sites sampled for the three time periods were uneven: ≤1960, n = 304; >1960 and ≤1990, n = 799; and >1990 and ≤2023, n = 1029. Thus, to further examine the temporal distributional change in all nine endemic taxa, we reduced our comparison to two time periods with a similar sample size: 1928–1990 (n = 1103) and 1991–2023 (n = 1029). The numbers of represented HUC-12s were also similar between the two time periods (113 and 109, respectively). We calculated the % change in the total area of occupied HUC-12s between the two time periods. For example, if sums of the areas (km2) of occupied HUC-12s for a species were 1500 (period 1) and 1000 (period 2), then the % change was calculated as (1000 − 1500)/1500 = −0.33.
For non-native fishes, we examined the time series data of occurrence records for 18 minnows (Leuciscidae) and 8 darters (Percidae). We examined the range expansion of these non-native species by calculating the cumulative sum of the HUC-12 areas (km2) occupied by each species across the years. We also calculated the proportion of cumulative area occupied to total watershed area, a value that represented the percent of the occupied area within the lower New River watershed. Time series plots of the year (x-axis) by the cumulative sum of the occupied area (y-axis) were expected to depict the range expansion of an invasive species as a sigmoidal curve [23,24], which can be modeled effectively with growth curves, such as the three-parameter logistic and Gompertz models. These growth curves have lag, growth, and asymptotic phases. We used JMP 17.2.0 to fit three-parameter logistic and Gompertz growth curves to the cumulative extent of the occurrence data of minnow and darter species. Prediction equations are as follows, where a = asymptote, b = growth rate, c = inflection point, Y = cumulative area, x = year, and ∈ = random error:
Gompertz:     Y i = a e − e − b x − c + ∈ i
Logistic:     Y i = a / 1 + e − b x − c + ∈ i
We used AICc model selection to determine the best approximating growth curve model, and we also reported an R2 value of model fit. The best approximating model was used for inference and parameter estimation, where parameters included asymptote, growth rate, and inflection point. We also calculated the first–fourth-order derivatives of the growth curve. The first-order derivative of the inflection point (i.e., the slope of the tangent line) represented the maximum rate of range expansion, a value useful for among-species comparisons. To calculate lag time, we first found the x-axis intercept (λ) of the tangent line to the inflection point of the curve, and then subtracted the year of the first species occurrence record from λ [29,70]. Comparison of lag times was useful for understanding among-species differences relative to the early stages of invasion. The third- and fourth-order derivatives were used to determine three points on the growth curve: the maximum acceleration point (MAP), the maximum deceleration point (MDP), and the asymptotic deceleration point (ADP) (see Figure 1). The section of the curve between λ and ADP represented the period of range expansion with an exponential increase between the MAP and MDP. The ADP represented a stabilization point proximate to the asymptote [71], where range expansion becomes asymptotic. Additional periods of time between influential points of the curve provide further among-species comparisons, including the lengths of time during the growth phase to reach either the maximum rate of range expansion (c-λ) or the asymptotic phase (ADP-λ). An invasion curve is incomplete if roots of derivatives are missing for the MAP, MDP, or ADP.

3. Results

Range reductions occurred for most endemic taxa and were visually determined using temporal distribution maps, e.g., New River Shiner, Kanawha Minnow, Candy Darter, and Appalachia Darter (Figure 4). Based on a comparison of the two time periods (1928–1990 and 1991–2023), one of the largest reductions in range was documented for the New River Shiner, where the % change in the total area of occupied HUC-12s was −72.3 (Table 1). Further, the number of HUC-12s occupied by the New River Shiner was reduced from 35 to 9. The percent changes in the extent of occurrence of the Kanawha Minnow, Candy Darter, and Appalachia Darter were −37.1, −31.8, and −36.9 with reductions in occupied HUCs of 8–5, 37–25, and 16–10, respectively. A percent change of −36.5 in the extent of occurrence was calculated for the “Kanawha” Rosyface Shiner, where the number of occupied HUC-12s decreased from 75 to 47. The extent of occurrence for the Bigmouth Chub was similar between the two periods with occupied areas of 8572.4 and 8223.1 km2, a % change of −4.1, and a reduction in occupied HUC-12s of 80 to 73. The Kanawha Sculpin expanded its range by 61.6% with a shift in the extent of occurrence of 1484.4 to 2399.0 km2. The range of the narrowly distributed Bluestone Sculpin in the West Virginia waters of the Bluestone River drainage increased from 134.4 km2 (one HUC-12) to 278.5 km2 (two HUC-12s). The Buckeye Creek Cave Sculpin is likely extinct, with a single occurrence record from one site locality in 1967.
The New River drainage of West Virginia has a long history of introductions of fishes and subsequent range expansions, including minnows (n= 18) and darters (n = 8). Based on first occurrence records for minnow and darter species, the timeline of introductions included the 1920s (two species), 1930s (four), 1940s (one), 1950s (two), 1960s (three), 1970s (seven), 1980s (two), 1990s (two), 2000s (one), and 2010s (two) (Table 2). Sampling effort was low during the early part of the time series (1897–1930), but 68.7% (8243.6 km2) of the total watershed area was sampled by 1935 based on the cumulative sum of areas of HUC-12s. The sampled area of the total watershed reached 90.6% by 1970 and 97.5% by 1990 (Figure 5). Species occurrence data for 115 HUC-12s from 1928 to 2023 had a mean of 18.5 samples per HUC-12.
Range expansions have occurred for many non-native minnows and darters based on the time series data of the cumulative sum of occupied 12-digit HUC areas (km2) (Figure 5). We considered 11 of 18 non-native minnows and 6 of 8 non-native darters as invasive species based on their extent of range expansion, as defined by the cumulative sum of HUC-12 areas (km2) occupied by each species. The 11 invasive minnows represented cumulative areas ranging from 1473 to 8334 km2 or 12.3 to 69.5 percent of the total watershed area, whereas the remaining 7 non-native minnows occupied smaller areas (cumulative area of 107–626 km2 and percent total area of 0.89–5.2) (Table 2). Range expansion of the six invasive darters was supported by cumulative areas ranging from 803 to 6461 km2 or 6.7 to 53.9 percent of the total watershed area, whereas the remaining two non-native darters had smaller ranges (cumulative area of 251–278 km2 and percent total area of 2.1–2.3) (Table 2). Given the recency of the first occurrence records for Swallowtail Shiner (2019) and Redline Darter (2019), these species may be in a lag phase period prior to range expansion.
Growth curve analyses of 17 invasive species of minnows and darters provided insights into among-species differences in lag phase duration and maximum rates of range expansion (Figure 6, Figure 7 and Figure 8). The AICc-selected growth rate models included the logistic (n = 12) and Gompertz (n = 5) models (Table 3), although both models were good fits to the data of all species. The lengths of lag periods (years) were highest for Mountain Redbelly Dace (44.5), Rosyside Dace (33.1), and White Shiner (31.1). Lag periods were between 20 and 25 years for Whitetail Shiner, Longnose Dace, Rainbow Darter, and Johnny Darter, and between 9 and 17 years for Bluehead Chub, Golden Shiner, Variegate Darter, Logperch, and Roanoke Darter. Lag times less than four years occurred for Spottail Shiner, Striped Shiner, Telescope Shiner, Fathead Minnow, and Tennessee Darter (Table 3). Maximum rates of range expansion (km2/yr) were highest for the Whitetail Shiner (504.7), Telescope Shiner (224.0), and Striped Shiner (239.0), and ranged between 150 and 200 for White Shiner, Longnose Dace, Rainbow Darter, and Variegate Darter (Table 3). The Rosyside Dace and Roanoke Darter had maximum rates of range expansion between 100 and 150, whereas those of all other species were < 100, including the two lowest rates for species commonly sold by commercial suppliers (Golden Shiner, 36.1, and Fathead Minnow, 39.7) (Table 3).
Species differed in lengths of time periods between influential points of the invasion curve (Table 3). The time to reach the maximum rate of range expansion (c-λ) ranged from 20.3 to 26.0 years for most minnows with exceptions for Whitetail Shiner (5.5 years), Spottail Shiner (8.8 years), Striped Shiner (10.6 years), Bluehead Chub (30.7 years), and Rosyside Dace (38.8 years). Values of c-λ were similar for four of six invasive darters (20.6–21.3 years) but lower values occurred for Variegate Darter (6.4 years) and Tennessee Darter (2.2 years). The total period of range expansion (ADP-λ), which represented the time (years) to reach the asymptotic phase, was longest for Johnny Darter (58.3) and Longnose Dace (55.1), and shortest for Whitetail Shiner (11.9) and Variegate Darter (13.7). The period of exponential increase (MDP-MAP), which ranged from 20 to 40 years for most species, was longest for Golden Shiner (50.2) and shortest for Whitetail Shiner (7.1), Variegate Darter (8.4), and Tennessee Darter (13.2). Invasion curves were incomplete for eight species, including missing values for the MAP (Spottail Shiner), MDP (Rosyside Dace and Bluehead Chub), and ACP (Rosyside Dace, Mountain Redbelly Dace, Bluehead Chub, Golden Shiner, Rainbow Darter, Logperch, and Roanoke Darter) (Table 3, Figure 6, Figure 7 and Figure 8).

4. Discussion

Species occurrence data supported range reductions for most endemic fishes, where six of nine endemic taxa had reduced ranges, two taxa exhibited range expansion, and one taxon had a relatively stable range. Most endemic fishes of the New River drainage, which are considered of conservation importance by the WVDNR, are managed under the state’s wildlife action plan (SWAP) as Species in Greatest Need of Conservation (SGCN). The SGCN of primary conservation concern are assigned a Priority 1 status [72]. Three of the species for which we reported reduced ranges are recognized by the WVDNR as Priority 1 SGCN: New River Shiner, Kanawha Minnow, and Candy [72]. The Candy Darter also has federal protection status under the Endangered Species Act [73]. The % change (−72.3) in the range of the New River Shiner greatly exceeded that of the Kanawha Minnow (−37.1) and Candy Darter (−31.8). Further, the area of occupied HUC-12s during the 1991–2023 period for the New River Shiner (998 km2) and Kanawha Minnow (528.0 km2) were considerably less than the area occupied by the Candy Darter (2658.6 km2). Although we elected to use two time periods (1928–1990 vs. 1991–2023) to quantify distribution change, range reductions for these species are apparent in the last 10 years, particularly regarding hybridization-related range reductions for the Candy Darter in the Greenbrier River drainage [74]. Additionally, we documented a reduced range for the Appalachia Darter (% change = −36.9), a result that is consistent with the existing SGCN conservation status assigned by the WVDNR [72] but that also could support evaluation of Priority 1 status. We also documented distribution changes for the Kanawha Sculpin, a priority 1 SGCN that has recently expanded its range from an occupied area of 1484.4 to 2399.0 km2 (% change = 61.6). Although range expansion is encouraging from a conservation perspective, this species remains narrowly distributed in occupying an area representing 20% of the total area of the lower New River drainage. Additionally, we documented a distribution change for the narrowly distributed Bluestone Sculpin. This species occurs in the Bluestone River drainage and is primarily restricted to the upper watershed in [52]. The known extent of occurrence for this species in West Virginia waters increased from one to two HUC-12s with a corresponding area increase of 134.4 to 278.5 km2, a change that could represent a range expansion or may be an artifact of the sampling effort.
Three endemic taxa of the lower New River system are not currently considered by the WVDNR to be of conservation concern. The “Kanawha” Rosyface Shiner has only recently been recognized as an undescribed endemic species, but it could be considered for further conservation evaluation. For comparison, we documented a % change (−36.5) in the range for the “Kanawha” Rosyface Shiner that was similar to that for Kanawha Minnow (−37.1), Candy Darter (−31.8), and Appalachia Darter (−36.9). Despite a range reduction of this taxon in the lower New River, the total area of occupied HUC-12s during 1991–2023 was 5213.8 km2, an area of occupancy representing a wider distribution range than most other endemic species. The Bigmouth Chub had the widest and most stable distribution range of all endemic species of the lower New River with similar areas of occupied HUC-12s during 1928–1990 (8572.4 km2) and 1991–2023 (8223.1 km2). This species has not been considered as an SGCN, a status supported by and consistent with our finding of a temporally stable distribution range. The Buckeye Creek Cave Sculpin, which had a range reduction from one to zero HUC-12s, has not been given SGCN consideration due to its presumed extinct status [55] Further sampling efforts for this sculpin in the Greenbrier River system are warranted given the difficulty of detecting species occupancy in karst habitats.
Bait-bucket introduction of aquatic species is well documented [12,75,76,77], and it is the likely vector for most non-native and invasive minnows and darters in the lower New River drainage [52,69,78,79]. Propagule pressure contributing to non-native species in the lower New River drainage is difficult to discern as some species may have one or two introduction events followed by species dispersal, whereas others may have been introduced on three or more occasions. Many non-native species of the lower New River system have had at least two introduction events based on occurrence records from the upper Gauley River drainage after 1966, with the time frame following the completion of an upstream dispersal barrier, i.e., Summersville Dam. The only invasive species not found in the upper Gauley River system were Whitetail Shiner, Rainbow Darter, Tennessee Darter, Variegate Darter, and Roanoke Darter. The Variegate Darter’s absence from the upper Gauley River is critical for Candy Darter conservation, as its presence would likely lead to hybridization and population decline of the Candy Darter as has occurred in the Greenbrier River drainage [74,80]. We expect multiple introduction events for Golden Shiner and Fathead Minnow, two species commonly available from commercial bait dealers in the region.
First occurrence records of non-native minnows and darters often align with popular fishing areas. The original location of a species introduction, however, may not be at or near the coordinates of first occurrence, as non-native species may disperse from the origin site of introduction. The first occurrence records in West Virginia of many non-native species are from the New River or tributaries near the state line with Virginia (Whitetail Shiner, Cutlip Minnow, Spottail Shiner, White Shiner, Rosefin Shiner, Swallowtail Shiner, Bluehead Chub, Emerald Shiner, Telescope Shiner, Tennessee Darter, and Redline Darter). These species may have been introduced by bait bucket in Virginia waters of the upper New River drainage and dispersed downstream to the lower drainage. The first occurrence record of the Roanoke Darter (mouth of Greenbrier River) was from 1970, but a 1972 record is from East River near the West Virginia/Virginia state line, suggesting that this species may have also dispersed downstream from Virginia waters. Jenkins and Burkhead [52] noted first occurrence records for species in Virginia waters that predate West Virginia records, such as Whitetail Shiner (1954), Cutlip Minnow (1956), Spottail Shiner (1948), White Shiner (1867), Rosefin Shiner (1922), Swallowtail Shiner (1971), Bluehead Chub (1880s), Telescope Shiner (1958), Roanoke Darter (1963), and Tennessee Darter (1976 and 1987 in Bluestone and New Rivers, respectively), The first occurrence record of the Emerald Shiner, however, may be from a local introduction associated with Bluestone Reservoir, as Jenkins and Burkhead [52] did not report records of this species from the New River, Virginia. The first occurrence records of other species from the lower New River or its small tributaries include Johnny Darter, Variegate Darter, and Blackside Darter. The first occurrence records of Mountain Redbelly Dace, Rosyside Dace, Common Shiner, and Rainbow Darter are from the Greenbrier River drainage adjacent to or within a historic resort property. Other species first noted from the Greenbrier River drainage include Striped Shiner (Anthony Creek), Golden Shiner (Seneca Lake), and Fathead Minnow (Sinking Creek), all locations of which are popular fishing areas. The Southern Redbelly Dace, Longnose Dace, and Logperch were first documented from the Gauley River drainage, a system where live bait fishing is common for many gamefishes.
Most non-native minnows (11 of 18) and darters (6 of 8) of the lower New River system were considered as invasive species, a designation based on their modeled invasion curve depicting a period of exponential range expansion and often a lag phase. Crooks and Soule [25] discussed two types of lag periods: lags in population increases at the local level, and lags that precede the exponential growth expansion of a species’ distribution range. We did not have adequate data to address abundance and population size, so we restricted our analysis to range expansion. The time lag that precedes exponential growth, however, is difficult to predict [81], as it is influenced by many biotic and abiotic factors, but its measurement and documentation provide conservation agencies with knowledge to inform management strategies and policies. A caveat to the estimation of lag time is the uncertainty regarding the timing of introduction, as the date of the first occurrence record may not closely approximate the date of first introduction [25,32]. Additionally, the end point of the lag phase could be overestimated if species detection is low owing to sampling methodologies, environmental conditions, or species traits [33,82,83].
Lag times for the 17 invasive species in our study were highly variable, ranging from 0.0 to 44.5 years. Of seven species with lag periods > 20 years, five had the earliest years for first occurrence records (1928–1935): Mountain Redbelly Dace, Rosyside Dace, White Shiner, Johnny Darter, and Longnose Dace. This result may reflect relatively low rates of both sampling efforts and species detection during the early years of the time series. Alternatively, it could be associated with poor habitat and water quality during the first half of the 20th century (see [64,84,85]). Crooks [55] discussed the oft-cited relationship between successful invaders and degraded habitat, but also emphasized an increase in invasive success and range expansion following improvements to habitat quality. Many of the invasive species in our study moved from the lag phase to exponential range expansion phase following the environmental water laws of the 1970s, a period when population recovery occurred for many fish species in the Ohio River drainage [55].
Longer lag times could be determined by a non-native species’ traits that influence dispersal or the ability to establish a population. A first occurrence record obtained during an unsuccessful introduction event (i.e., where a population was not established) followed years later by a successful introduction and second occurrence record would give the appearance of an extended lag time. As an example, the first and second occurrence records of the Mountain Redbelly Dace were from the Greenbrier River drainage in 1933 and 1957, a time difference that could reflect separate introduction events. Small benthic fishes could have longer lag times than that of pelagic fishes based on dispersal abilities [86,87], but rapid range expansion of non-native benthic darters has been well demonstrated [88,89,90]. In our study, the lag period for the benthic Rainbow Darter (23.2 years) was similar to that of the Whitetail Shiner (22.4 years), where both species subsequently underwent rapid range expansion resulting in extents of occurrence of 6461 and 5611 km2, respectively.
The extremes for low lag times from our study could be influenced by sampling effort and species detection. The low lag estimates for Striped Shiner and Spottail Shiner may reflect non-detection of these species in earlier surveys. The first three collection records for Striped Shiner occurred in 1955 from separate HUC-12s in the Greenbrier River watershed. The Spottail Shiner was distributed across multiple HUC-12s shortly after its first encounter during the 1970s, and its invasion curve is incomplete without a value for the MAP. Thus, population establishment and range expansion were in progress at the time of the first record of occurrence for both species.
Propagule pressure, which may influence lag time and range expansion, includes both propagule size (the number of introduced individuals) and propagule number (the repeat introduction of individuals over time) [27]. Relatively low lag times were observed for two common bait minnows: Golden Shiner (11.8 years) and Fathead Minnow (1.3 years). Repeat introductions of these two species are expected to be frequent, given common use by anglers and availability for purchase at bait stores in the region. Thus, the lag times and range expansion for these two minnows may not be influenced by either the traits of species or of the community/environment, but rather by propagule number. The maximum rates of range expansion (m) for Fathead Minnow and Golden Shiner, however, were less than all other invasive minnows. Also, we assumed that lag times would be shorter for those species that dispersed from the upper to the lower New River drainage, as the leading edge of an expanding species distribution could have a higher propagule pressure. Another possibility is long-distance dispersal of a few individuals from the leading edge of an expanding range [27,91], a scenario equivalent to a bait-bucket introduction of a few individuals.
Invasive species differed widely in the maximum rate of range expansion (m), which is a value at the inflection point of the exponential growth phase of the invasion curve. Understanding the causal factors of these differences is problematic given a wide range of potential influences such as propagule pressure, life history or dispersal traits of species, traits of the community or environment, or species interactions. The highest maximum rates of range expansion in the lower New River, which were not family or genus specific, were those of minnows representing three genera (Whitetail Shiner, Telescope Shiner, Striped Shiner, and White Shiner), and one species of darter (Rainbow Darter). The invasive impact of these species is undocumented, but the Variegate Darter’s impact via hybridization with the Candy Darter is well defined [74,80,92]. The range expansion of the benthic Variegate Darter, with a lag period of 13.2 years, only achieved an extent of occurrence of 2027 km2, but it reached its maximum rate of range expansion (m = 157.9) in 6.4 years, a much shorter time period than that of most invasives. The two lowest maximum rates of range expansion were by Golden Shiner and Fathead Minnow, two common bait minnows sold by regional commercial bait dealers [93,94], whose range expansions were likely driven by propagule pressure from multiple bait-bucket releases.
The presence of an invasion curve’s asymptote and associated ADP may indicate that the species is approaching an equilibrium state or saturation point [23,24]. Invasion curves were incomplete with missing values for ADP for seven species (Rosyside Dace, Mountain Redbelly Dace, Bluehead Chub, Golden Shiner, Rainbow Darter, Logperch, and Roanoke Darter), an indication of continuing range expansion. In contrast, asymptotes were well defined for Whitetail Shiner, Spottail Shiner, White Shiner, Striped Shiner, Longnose Dace, and Variegate Darter. The extents of occurrence for White Shiner (7158 km2), Striped Shiner (7135 km2), and Longnose Dace (8174 km2) are approaching the total watershed area, a factor that may contribute to slowed range expansion. Other species in an asymptotic phase had narrower extents of occurrence, where multiple explanations are possible including traits of the community or environment, or species interactions.
Non-native species have an increased success of becoming invasive in watersheds with depauperate native faunas [34,35], a scenario that may contribute to the high proportion of invasive species in the lower New River system. Additionally, invader–invader and invader–native species relationships can influence invasive species success [95], and it is likely that species traits and species interactions influence some of the invasion curves from our study. As an example, invasive species that modify habitat (i.e., ecosystem engineers) may influence lag periods [27,96] or aid in the range expansion of other invasive species (i.e., “invasional meltdown” (sensu [97,98])). In the lower New River system, the Bluehead Chub is an ecosystem engineer, as males build spawning gravel mounds that are also used by native and non-native “nest associate” minnows [52]. This is an unusual case where an invasive fish (Bluehead Chub) may benefit the native species, but it also may reduce lag times and aid in the range expansion of other invasive minnows. Minnow species in the New River drainage of the genera Chrosomus, Clinostomus, Luxilus, Lythrurus, Notropis, and Rhinichthys are all known to spawn on Bluehead Chub mounds [99,100]. In another example from the lower New River system, Keplinger [101] documented mixed-species shoaling between two invader–native pairs (Telescope Shiner and New River Shiner, and Whitetail Shiner and Spotfin Shiner Cyprinella spiloptera), supporting species interactions that could aid in the range expansion of invasive species.
Although most non-native minnows and darters of the lower New River have become invasive, nine species were either not able to establish populations or have been restricted in dispersal or range expansion. The presence of the Southern Redbelly Dace (Chrosomus erythrogaster) [55] and the exotic Rudd (Scardinius erythrophthalmus) [55,102] are based on single occurrence records, indicating that low propagule pressure may have prevented population establishment or restricted their distributions. The Swallowtail Shiner and Cutlip Minnow are also represented by single occurrence records and are species which likely dispersed downstream, as multiple records support established populations in Virginia waters [52]. A population of the Common Shiner persists in one tributary of the Greenbrier River drainage that flows through a resort hotel property, but range expansion of this species has not been documented. The Rosefin Shiner, which was first documented from Indian Creek of the lower New River system in 1966, expanded its range to 626 km2 (six HUC-12s), but has not been observed since 2011. The Blackside Darter was first documented in 1979 in the New River gorge, which is a high-gradient canyon area of the watershed and is where an established population is currently confined to a single tributary. Range expansion of the Blackside Darter beyond the local area of introduction has not occurred, which may be due to species traits or environmental restrictions to dispersal. The Redline Darter is a recent addition to the fauna [55], representing a non-native species that may be in a lag phase subsequent to rapid range expansion within the lower New River system.
A similar study was conducted by Buckwalter et al. [54] using a 77-year dataset (1938–2014) of fish occurrence records from the Virginia and North Carolina portion of the New River drainage, which allowed for comparison of fish distribution trends between the lower and upper regions of the watershed. Methods of analysis differed between the two studies, as distribution changes were based on the cumulative area of occupied HUC-12s (our study) versus the number of detections per sampled HUC-12s. Also, the origin status (native vs. non-native) of several species differed as we followed the designations of Cincotta and Welsh [55], whereas Buckwalter et al. [54] followed those of Jenkins and Burkhead [52]. Relative to distribution trends, Buckwalter et al. [54] classified species as “strong spreaders”, “weak spreaders”, “decliners”, and “stable”. For endemic species, distribution trends for Bigmouth Chub (“stable”) and “Kanawha” Rosyface Shiner (“decliner”) were consistent with our findings from the lower New River system. A “stable” status for four species (Kanawha Minnow, New River Shiner, Candy Darter, and Kanawha Sculpin) and a “weak spreader” classification for Appalachia Darter were inconsistent with our findings for the lower New River drainage.
Not all species considered as invasive in our study were given a similar designation in the study by Buckwalter et al. [54]. The Rainbow Darter had the highest maximum rate of range expansion of all invasive darters in our study, a finding consistent with the “strong spreader” designation of Buckwalter et al. [54]. Cessna et al. [89] also reported the rapid range expansion of the Rainbow Darter following its introduction to the Potomac River drainage of the Chesapeake Bay. Buckwalter et al. [54] documented three minnows as “strong spreaders” (Mountain Redbelly Dace, Rosyside Dace, and Bluehead Chub) and considered them as native, whereas we considered these species as invasive. Rapid range expansions of these species in the lower and upper New River drainage are consistent with the expected distribution trends of invasive species. For the lower New River system, maximum rates of range expansion and extents of occurrence were high for Whitetail Shiner and Telescope Shiner, moderate for Roanoke Darter, and low for Johnny Darter, whereas all four species were considered as “weak spreaders” in the upper New River. Distribution trends of White Shiner, Striped Shiner, Golden Shiner, Fathead Minnow, Longnose Dace, and Logperch were “stable” in the upper New River system, whereas we considered these six species as invasive with expanding distribution ranges in the lower New River system. The between-study comparison demonstrated that trends in endemic and invasive species can differ between lower and upper regions of a watershed. Within-drainage differences are particularly important when the lower and upper regions of a watershed represent a political division under jurisdiction of separate management agencies, demonstrating and supporting the need for state-level data for conservation assessments.

5. Conclusions

Most of the endemic species of the lower New River drainage are of conservation concern, where range reductions have occurred concomitantly with the range expansion of many non-native fishes. Most of the non-native minnows and darters in the lower New River system were considered invasive based on the fit of range expansion data to invasion curve models. It was not our intent to speculate or infer causal relationships between distribution trends of invasive and endemic species. As discussed by Ricciardi et al. [103], there is no guarantee of a corresponding increase between both the impact of an invasive species and its extent of occurrence. Given the skewed proportion of non-native to native species in the lower New River, it would be reasonable to expect some level of invasive impact by the non-native fauna. Range reductions of endemic species, however, could be associated with cumulative impacts of habitat-related stressors and invasive [13,104], a combination of causal factors that are often not well understood [17,35]. In the lower New River, the only clear invasive species impact is that of the Variegate Darter on the endemic Candy Darter, where hybridization has extensively reduced the Candy Darter’s range within the Greenbrier River drainage [74]. Impacts of the other 16 invasive species are unmeasured, but given their potential for impacts to the native fauna, our documentation of the lag times, exponential range expansions, and other derived variables from invasion curves of these species could provide state conservation agencies with knowledge to aid management strategies. We concur with Crooks ([32], p. 317), who stated that “examining why lags occur and when they end will help improve predictive ability as well as point to limitations in our efforts to decide what invaders will do in the future”. The same logic applies to examining attributes of all phases of the invasion curve, as we have carried out for invasive minnows and darters of the lower New River system.

Author Contributions

Conceptualization, S.A.W.; data curation, S.A.W., D.A.C., N.V.O. and J.R.S.J.; formal analysis, S.A.W.; writing—original draft, S.A.W.; writing—review and editing, S.A.W., D.A.C., N.V.O. and J.R.S.J.; project administration, S.A.W. All authors have read and agreed to the published version of the manuscript.

Funding

Partial support was provided through grants to the West Virginia Cooperative Research Unit from the National Park Service (1434-03HQRU1563), U.S. Forest Service (1434-07HQRU1563), and West Virginia Division of Natural Resources (1434-17HQRU1563).

Institutional Review Board Statement

Portions of this study were performed under the auspices of West Virginia University IACUC protocols 04-1005, 06-0707, and 2304064841.

Data Availability Statement

The datasets presented in this article were not available at the time of publication. Data on endemic fishes are not readily available because of species conservation status in West Virginia. Requests to access datasets should be directed to N.V.O.

Acknowledgments

We would like to thank the following people from the National Park Service: Jennifer Flippin and Jesse Purvis; U.S. Environmental Protection Agency: Lou Reynolds; U.S. Forest Service: Chad Landress and Mike Owens; U.S. Geological Survey: Terence Messinger and Doug Chambers; West Virginia Department of Environmental Protection: Jason Morgan and Ryan Pack; West Virginia Division of Natural Resources: Isaac Gibson, Walt Kordek, Kiernan O’Malley, Mark Scott, Dave Thorne, and Dave Wellman. Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the U.S. Government.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Conceptual growth curve model (A) as applied to the range expansion of an invasive species represented by time (x axis) and the cumulative area of species’ occurrence (y axis). Lag phase is determined by the x-axis intercept (λ) of the tangent line (red dash) to the inflection point (IP) of the curve. The growth phase is the period of exponential increase in the species’ range, where the slope of the tangent line at the IP represents the maximum rate of range expansion. The asymptotic phase is the saturation period of range expansion. Influential points can be determined by the curve’s first–fourth-order derivatives (B–E) and include the IP, maximum acceleration point (MAP), maximum deceleration point (MDP), and asymptotic deceleration point (ADP).
Figure 1. Conceptual growth curve model (A) as applied to the range expansion of an invasive species represented by time (x axis) and the cumulative area of species’ occurrence (y axis). Lag phase is determined by the x-axis intercept (λ) of the tangent line (red dash) to the inflection point (IP) of the curve. The growth phase is the period of exponential increase in the species’ range, where the slope of the tangent line at the IP represents the maximum rate of range expansion. The asymptotic phase is the saturation period of range expansion. Influential points can be determined by the curve’s first–fourth-order derivatives (B–E) and include the IP, maximum acceleration point (MAP), maximum deceleration point (MDP), and asymptotic deceleration point (ADP).
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Figure 2. Study area representing watershed boundaries and 12-digit hydrologic unit codes (HUCs) of the New River drainage in West Virginia. Emphasized watersheds include Gauley, Bluestone, East, and Greenbrier Rivers and Indian Creek. NC = North Carolina; VA = Virginia; WV = West Virginia.
Figure 2. Study area representing watershed boundaries and 12-digit hydrologic unit codes (HUCs) of the New River drainage in West Virginia. Emphasized watersheds include Gauley, Bluestone, East, and Greenbrier Rivers and Indian Creek. NC = North Carolina; VA = Virginia; WV = West Virginia.
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Figure 3. Temporal frequency of fish collections by year (1897–2023) for the New River drainage, West Virginia. Gray area indicates data (1928–2023) used for analysis.
Figure 3. Temporal frequency of fish collections by year (1897–2023) for the New River drainage, West Virginia. Gray area indicates data (1928–2023) used for analysis.
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Figure 4. Temporal dot distribution map of species occurrence records for visualizing range reductions in (A) Kanawha Minnow Phenacobius teretulus, (B) New River Shiner Miniellus scabriceps, (C) Candy Darter Etheostoma osburni, and (D) Appalachia Darter Percina gymnocephala) of the New River, West Virginia, based on three time periods.
Figure 4. Temporal dot distribution map of species occurrence records for visualizing range reductions in (A) Kanawha Minnow Phenacobius teretulus, (B) New River Shiner Miniellus scabriceps, (C) Candy Darter Etheostoma osburni, and (D) Appalachia Darter Percina gymnocephala) of the New River, West Virginia, based on three time periods.
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Figure 5. Range expansion of select non-native minnows (A) and darters (B) in the New River drainage, West Virginia, based on time series of the cumulative sum of occupied 12-digit hydrologic unit code (HUC) areas (km2). The thick gray line represents the cumulative area of sampled 12-digit HUC watersheds for comparison. The approximate total watershed area is 11,994 km2.
Figure 5. Range expansion of select non-native minnows (A) and darters (B) in the New River drainage, West Virginia, based on time series of the cumulative sum of occupied 12-digit hydrologic unit code (HUC) areas (km2). The thick gray line represents the cumulative area of sampled 12-digit HUC watersheds for comparison. The approximate total watershed area is 11,994 km2.
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Figure 6. Invasion curves based on growth curve models (Gompertz or logistic) fit to a time series of the cumulative sum of areas occupied by nine non-native minnows (Leuciscidae) of the New River drainage, West Virginia. The x-axis intercept of the tangent line at the inflection point (IP) of the curve defines the lag phase (λ) period. MAP = maximum acceleration point, MDP = maximum deceleration point, and ADP = asymptotic deceleration point.
Figure 6. Invasion curves based on growth curve models (Gompertz or logistic) fit to a time series of the cumulative sum of areas occupied by nine non-native minnows (Leuciscidae) of the New River drainage, West Virginia. The x-axis intercept of the tangent line at the inflection point (IP) of the curve defines the lag phase (λ) period. MAP = maximum acceleration point, MDP = maximum deceleration point, and ADP = asymptotic deceleration point.
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Figure 7. Invasion curves based on growth curve models (Gompertz or logistic) fit to a time series of the cumulative sum of areas occupied by six invasive darters (Percidae) of the New River drainage, West Virginia. The x-axis intercept of the tangent line at the inflection point (IP) of the curve defines the lag phase (λ) period. MAP = maximum acceleration point, MDP = maximum deceleration point, and ADP = asymptotic deceleration point.
Figure 7. Invasion curves based on growth curve models (Gompertz or logistic) fit to a time series of the cumulative sum of areas occupied by six invasive darters (Percidae) of the New River drainage, West Virginia. The x-axis intercept of the tangent line at the inflection point (IP) of the curve defines the lag phase (λ) period. MAP = maximum acceleration point, MDP = maximum deceleration point, and ADP = asymptotic deceleration point.
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Figure 8. Invasion curves based on growth curve models (Gompertz or logistic) fit to a time series of the cumulative sum of areas occupied by two commercially sold bait minnows (Golden Shiner and Fathead Minnow) of the New River drainage, West Virginia. The x-axis intercept of the tangent line at the inflection point (IP) of the curve defines the lag phase (λ) period. MAP = maximum acceleration point, MDP = maximum deceleration point, and ADP = asymptotic deceleration point.
Figure 8. Invasion curves based on growth curve models (Gompertz or logistic) fit to a time series of the cumulative sum of areas occupied by two commercially sold bait minnows (Golden Shiner and Fathead Minnow) of the New River drainage, West Virginia. The x-axis intercept of the tangent line at the inflection point (IP) of the curve defines the lag phase (λ) period. MAP = maximum acceleration point, MDP = maximum deceleration point, and ADP = asymptotic deceleration point.
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Table 1. Extent of occurrence and temporal distribution change in endemic fish taxa of the New River drainage, West Virginia, between two time periods: 1928–1990 and 1991–2023. Area (km2) = the total area of occupied 12-digit hydrologic unit codes (HUCs). % Area = the proportion of the total area of occupied 12-digit HUCs to the total area of the New River watershed, West Virginia. HUC (n) = the total number of occupied 12-digit HUCs. % Change = the percent of change in the total area of occupied 12-digit HUCs between the two time periods (negative values = range reduction, positive values = range expansion).
Table 1. Extent of occurrence and temporal distribution change in endemic fish taxa of the New River drainage, West Virginia, between two time periods: 1928–1990 and 1991–2023. Area (km2) = the total area of occupied 12-digit hydrologic unit codes (HUCs). % Area = the proportion of the total area of occupied 12-digit HUCs to the total area of the New River watershed, West Virginia. HUC (n) = the total number of occupied 12-digit HUCs. % Change = the percent of change in the total area of occupied 12-digit HUCs between the two time periods (negative values = range reduction, positive values = range expansion).
≤1990>1990
Endemic TaxaArea (km2)% AreaHUC
(n)		Area (km2)% AreaHUC (n)% Change
Miniellus scabriceps (New River Shiner)3600.030.035998.08.39−72.3
Nocomis platyrhynchus (Bigmouth Chub)8572.471.5808223.168.673−4.1
Notropis cf. rubellus (Rosyface Shiner) 8208.168.4755213.843.547−36.5
Phenacobius teretulus (Kanawha Minnow)839.57.08528.04.45−37.1
Cottus cf. carolinae (Bluestone Sculpin)134.41.11278.52.32107.2
Cottus cf. carolinae (Buckeye Creek Cave)222.61.910.00.00−100.0
Cottus kanawhae (Kanawha Sculpin)1484.412.4142399.020.01961.6
Etheostoma osburni (Candy Darter)3898.532.5372658.622.225−31.8
Percina gymnocephala (Appalachia Darter)1720.114.3161085.09.010−36.9
Table 2. Non-native minnows (Leuciscidae) and darters (Percidae) of the New River drainage, West Virginia, based on the study by Cincotta and Welsh [55]. Year = the year of the first occurrence record. Area = the sum of the areas of occupied 12-digit hydrologic unit codes (HUCs). % Area = the proportion of the sum of the area of occupied 12-digit HUCs to the total watershed area of the New River drainage in West Virginia. HUC (n) = the total number of occupied 12-digit HUCs. * = commercially sold bait minnow.
Table 2. Non-native minnows (Leuciscidae) and darters (Percidae) of the New River drainage, West Virginia, based on the study by Cincotta and Welsh [55]. Year = the year of the first occurrence record. Area = the sum of the areas of occupied 12-digit hydrologic unit codes (HUCs). % Area = the proportion of the sum of the area of occupied 12-digit HUCs to the total watershed area of the New River drainage in West Virginia. HUC (n) = the total number of occupied 12-digit HUCs. * = commercially sold bait minnow.
SpeciesYearArea (km2)% AreaHUC (n)
Chrosomus erythrogaster (Rafinesque) (Southern Redbelly Dace)20031070.891
Chrosomus oreas Cope (Mountain Redbelly Dace)1933373031.137
Clinostomus funduloides Girard (Rosyside Dace)1928637653.257
Cyprinella galactura (Cope) (Whitetail Shiner)1972561146.850
Exoglossum maxillingua (Lesueur) (Cutlip Minnow)19721531.31
Hudsonius hudsonius (Clinton) (Spottail Shiner)1972147312.312
Luxilus albeolus (Jordan) (White Shiner)1935715859.763
Luxilus chrysocephalus Rafinesque (Striped Shiner)1955713559.567
Luxilus cornutus (Mitchill) (Common Shiner)19851090.91
Lythrurus ardens (Cope) (Rosefin Shiner)19666265.26
Miniellus procne (Cope) (Swallowtail Shiner)20191251.01
Nocomis leptocephalus (Girard) (Bluehead Chub)1972362630.234
Notemigonus crysoleucas (Mitchill) (Golden Shiner) *1948190915.918
Notropis atherinoides Rafinesque (Emerald Shiner)19354623.94
Notropis telescopus (Cope) (Telescope Shiner)1972833469.576
Pimephales promelas Rafinesque (Fathead Minnow) *1964179314.913
Rhinichthys cataractae (Valenciennes) (Longnose Dace)1928817468.272
Scardinius erythrophthalmus (Linnaeus) (Rudd)19911511.31
Etheostoma caeruleum Storer (Rainbow Darter)1959646153.958
Etheostoma nigrum Rafinesque (Johnny Darter)1935242120.222
Etheostoma tennesseense Powers and Mayden (Tennessee Darter)19998036.77
Etheostoma variatum Kirtland (Variegate Darter)1982202716.918
Nothonotus rufilineatus (Cope) (Redline Darter)20192782.32
Percina caprodes (Rafinesque) (Logperch)1965220818.419
Percina maculata (Girard) (Blackside Darter)19792512.12
Percina roanoka (Jordan and Jenkins) (Roanoke Darter)1970466238.941
Table 3. Parameter estimates and standard errors of asymptote (a), growth rate (b), and inflection point (c) from three-parameter growth curve models M is Gompertz (G) and logistic (L). Models were fit to a time series of the cumulative area occupied by non-native minnows (Leuciscidae) and darters (Percidae) of the New River drainage, West Virginia. The first derivative (m) is the slope of the tangent line to the inflection point of the growth curve, a value representing the maximum rate of range expansion. Lag (years) was calculated by subtracting the first year of species occurrence from λ, where λ is the x-axis intercept of the tangent line to the inflection point of the growth curve. The maximum acceleration point (MAP), maximum deceleration point (MDP), and asymptotic deceleration point (ADP) of the curve are determined by roots of the third- and fourth-order derivatives. MAP-MDP represents the period (years) of exponential range expansion. c-λ and ADP-λ are periods (years) during the growth phase to reach the maximum rate of range expansion and the asymptotic phase, respectively. A dash (–) indicates a missing value from an incomplete invasion curve. * = commercially sold bait minnow.
Table 3. Parameter estimates and standard errors of asymptote (a), growth rate (b), and inflection point (c) from three-parameter growth curve models M is Gompertz (G) and logistic (L). Models were fit to a time series of the cumulative area occupied by non-native minnows (Leuciscidae) and darters (Percidae) of the New River drainage, West Virginia. The first derivative (m) is the slope of the tangent line to the inflection point of the growth curve, a value representing the maximum rate of range expansion. Lag (years) was calculated by subtracting the first year of species occurrence from λ, where λ is the x-axis intercept of the tangent line to the inflection point of the growth curve. The maximum acceleration point (MAP), maximum deceleration point (MDP), and asymptotic deceleration point (ADP) of the curve are determined by roots of the third- and fourth-order derivatives. MAP-MDP represents the period (years) of exponential range expansion. c-λ and ADP-λ are periods (years) during the growth phase to reach the maximum rate of range expansion and the asymptotic phase, respectively. A dash (–) indicates a missing value from an incomplete invasion curve. * = commercially sold bait minnow.
SpeciesMa b c m λLagMAPMDPADPMDP-MAPc-λADP-λ
Chrosomus oreas (Mountain Redbelly Dace)L4712.7
(155.9)		0.081
(0.003)		2002.1
(1.0)		95.91977.544.51985.92018.2–32.324.6–
Clinostomus funduloides (Rosyside Dace)L8883.1
456.1)		0.052
(0.002)		1999.9
(2.3)		114.31961.133.11974.3–––38.8–
Cyprinella galactura (Whitetail Shiner)L5584.5
(48.3)		0.366
(0.017)		1999.9
(0.15)		504.71994.422.41996.32003.42006.37.15.511.9
Hudsonius hudsonius (Spottail Shiner)G1338.4
(36.9)		0.113
(0.015)		1977.6
(0.73)		55.61968.8−3.2–1986.11993.5–8.824.7
Luxilus albeolus (White Shiner)L7502.4
(159.2)		0.096
(0.005)		1987.0
(0.75)		179.41966.131.11973.02000.72010.727.720.944.6
Luxilus chrysocephalus (Striped Shiner)G6934.1
(57.1)		0.094
(0.004)		1965.9
(0.27)		239.01955.30.31955.71976.21985.120.510.629.8
Nocomis leptocephalus (Bluehead Chub)L5615.8
(605.7)		0.065
(0.005)		2011.9
(3.5)		91.51981.29.21991.7–––30.7–
Notemigonus crysoleucas (Golden Shiner) *G2556.8
(144.0)		0.038
(0.003)		1985.8
(1.8)		36.11959.811.81960.72010.9–50.226–
Notropis telescopus (Telescope Shiner)L9430.2
(364.6)		0.095
(0.007)		1995.1
(1.16)		224.01974.12.11981.32009.02019.227.72145.1
Pimephales promelas (Fathead Minnow) *G2187.2
(103.5)		0.049
(0.004)		1985.6
(1.3)		39.71965.31.31964.51999.52015.03520.349.7
Rhinichthys cataractae (Longnose Dace)L8278.5
(98.7)		0.078
(0.002)		1974.6
(0.54)		161.01948.920.91957.61991.52004.033.925.755.1
Etheostoma caeruleum (Rainbow Darter)L7737.5
(375.0)		0.10
(0.007)		2000.6
(1.4)		187.51980.0211987.12014.2–27.120.6–
Etheostoma nigrum (Johnny Darter)G2781.7
(143.5)		0.048
(0.005)		1980.6
(1.5)		49.11959.824.81960.62000.72018.140.120.858.3
Etheostoma tennesseense (Tennessee Darter)L904.9
(43.1)		0.198
(0.019)		2010.1
(0.69)		44.51999.90.92003.52016.72021.613.22.221.7
Etheostoma variatum (Variegate Darter)L2017.9
(31.5)		0.315
(0.02)		2001.6
(0.27)		157.91995.213.21997.42005.82008.98.46.413.7
Percina caprodes (Logperch)L2637.9
(114.1)		0.095
(0.005)		2003.0
(1.2)		62.31981.816.81957.61991.5–33.921.2–
Percina roanoka (Roanoke Darter)L5606.1
(292.4)		0.094
(0.007)		2001.1
(1.48)		131.51979.89.81987.12015.1–2821.3–
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MDPI and ACS Style

Welsh, S.A.; Cincotta, D.A.; Owens, N.V.; Stauffer, J.R., Jr. Endemic and Invasive Species: A History of Distributional Trends in the Fish Fauna of the Lower New River Drainage. Water 2025, 17, 221. https://doi.org/10.3390/w17020221

AMA Style

Welsh SA, Cincotta DA, Owens NV, Stauffer JR Jr. Endemic and Invasive Species: A History of Distributional Trends in the Fish Fauna of the Lower New River Drainage. Water. 2025; 17(2):221. https://doi.org/10.3390/w17020221

Chicago/Turabian Style

Welsh, Stuart A., Daniel A. Cincotta, Nathaniel V. Owens, and Jay R. Stauffer, Jr. 2025. "Endemic and Invasive Species: A History of Distributional Trends in the Fish Fauna of the Lower New River Drainage" Water 17, no. 2: 221. https://doi.org/10.3390/w17020221

APA Style

Welsh, S. A., Cincotta, D. A., Owens, N. V., & Stauffer, J. R., Jr. (2025). Endemic and Invasive Species: A History of Distributional Trends in the Fish Fauna of the Lower New River Drainage. Water, 17(2), 221. https://doi.org/10.3390/w17020221

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