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Movement Patterns of the Threatened Blackside Dace, Chrosomus cumberlandensis, in Two Southeastern Kentucky Watersheds
Jason E. Detar and Hayden T. Mattingly

Southeastern Naturalist, Volume 12, Special Issue 4 (2013): 64–81

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J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 64 Vol. 12, Special Issue 4 Movement Patterns of the Threatened Blackside Dace, Chrosomus cumberlandensis, in Two Southeastern Kentucky Watersheds Jason E. Detar1,2 and Hayden T. Mattingly1,* Abstract - Chrosomus cumberlandensis (Blackside Dace) is a threatened stream fish endemic to the upper Cumberland River drainage in Kentucky and Tennessee. Little is known about the movement patterns of this species. Acquiring an understanding of baseline dispersal patterns is necessary to inform management and recovery actions. We tagged 653 Blackside Dace with visible implant elastomer injections in the Big Lick Branch and Rock Creek watersheds of southeastern Kentucky to determine the frequency, spatial extent, directionality, and environmental correlates of dace movements. We recaptured dace from February 2003 through March 2004 using baited minnow traps. Most tagged dace (81% in Big Lick Branch and 58% in Rock Creek) were recaptured within the same 200-m stream reach where tagging occurred. However, several individuals moved considerable distances from the original tagging site, including the first documented intertributary movement for this species. Mean (± SD) distances moved upstream in Big Lick Branch (148 ± 138 m) and Rock Creek (733 ± 1259 m) were not significantly different from mean distances moved downstream (77 ± 29 m and 314 ± 617 m, respectively). However, the mean overall distance moved was greater in Rock Creek, a longer stream than Big Lick Branch. The spatial arrangement of traps in both watersheds likely produced a distance-weighted bias such that we slightly overestimated the frequency of short-distance movements and underestimated the frequency of long-distance movements. Our results for Blackside Dace are consistent with a number of other studies that found stream fish populations composed of a large sedentary group and a smaller mobile group. The demonstrated ability of Blackside Dace to move into and between tributaries will remain vital for long-term population viability, and emphasizes the importance of maintaining suitable corridors within and among Cumberland River tributary streams. Introduction Chrosomus cumberlandensis (Starnes and Starnes) (Blackside Dace) is a small stream fish of the minnow family, Cyprinidae, endemic to the upper Cumberland River drainage in southeastern Kentucky and northeastern Tennessee (Eisenhour and Strange 1998). Anthropogenic activities, including coal mining, road construction, agriculture, and poor logging techniques, have led to the decline of the Blackside Dace and resulted in its federal listing as a threatened species (O’Bara 1990, L.B.Starnes and W.C. Starnes 1981, W.C. Starnes and L.B. Starnes 1978, USFWS 1988). The entire Blackside Dace range lies within a region of rich coal and timber reserves (Starnes 1981), which has been and continues to be subjected to the impacts of mining and logging. 1Department of Biology, Box 5063, Tennessee Technological University, Cookeville, TN 38505. 2Current Address - Pennsylvania Fish and Boat Commission, 450 Robinson Lane, Bellefonte, PA 16823. *Corresponding author - hmattingly@tntech.edu. Ecology and Conservation of the Threatened Blackside Dace, Chrosomus cumberlandensis 2013 Southeastern Naturalist 12(Special Issue 4):64–81 65 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 Relatively few studies have been conducted on the Blackside Dace, and none have focused on determining their movement patterns. Movement and dispersal are key determinants of population structure and function (Skalski and Gilliam 2000) that allow stream fishes to be resilient to habitat alteration and stochastic events. Additionally, periodic long-range movements may enable small-bodied species to respond to variation in resources over space and across a variety of habitats (Freeman 1995). In general, stream fish movements are important for maintaining population connectivity, reducing the potential for local extinctions, and predicting the rate of colonization or recolonization of suitable habitats (Albanese et al. 2009, Larson et al. 2002, Woolnough et al. 2009). Therefore, a basic understanding of Blackside Dace population dynamics, including movement patterns, is vital to the prudent management of this threatened species. As the extraction of coal and timber resources continues within watersheds containing Blackside Dace populations, it is critical that management agencies know whether the species is relatively sedentary or mobile. Quantitative estimates of movement probabilities are needed to better evaluate population viability under different scenarios of natural or anthropogenic changes in stream habitat conditions. If individuals exhibit extensive movement, it may be possible for them to vacate their present stream reaches to avoid mining, logging, or other disturbance, and to repopulate areas if habitat conditions improve following the disturbance. Furthermore, a better understanding of movement patterns can lend insight into the degree of connectivity between seemingly isolated populations within and among upper Cumberland River watersheds. Thus, the primary objective of this study was to determine the frequency, spatial extent, directionality (upstream vs. downstream), and environmental correlates of Blackside Dace movement to aid in the conservation and management of the species. Study Area Movement patterns were monitored in two southeastern Kentucky watersheds, Big Lick Branch (including an adjacent unnamed tributary, hereafter Dace Branch) in Pulaski County and Rock Creek in southeastern McCreary County (Fig. 1). Big Lick Branch is a small, second-order stream that is a direct tributary to Lake Cumberland below Cumberland Falls. Big Lick Branch generally flows southeastward to its confluence with Lake Cumberland, and there are no perennial tributaries to the stream. The Blackside Dace population in Big Lick Branch is presumably isolated from other substantial populations in the area because of the reservoir. The Big Lick Branch watershed encompasses 7.1 km² and is comprised of mostly Daniel Boone National Forest land and a smaller amount of privately owned land. The majority of the watershed is undisturbed and well-forested. The stream maintains good flow all year. A dirt and gravel forest road parallels the lower two-thirds of the stream. Aside from the forest road, there is currently no other development within the watershed. Dace Branch is a small, first-order tributary to Lake Cumberland which was a tributary to Big Lick Branch before the Cumberland River was impounded in J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 66 Vol. 12, Special Issue 4 1952. After 1952, Lake Cumberland inundated the Big Lick Branch watershed upstream of the historic confluence of these two streams, thus severing their lotic connection. The entire watershed of this small stream is well-forested and lies within Daniel Boone National Forest. A steep, 2-m-high cascade creates a barrier near the mouth of Dace Branch when Lake Cumberland is drawn down to winter pool, which likely restricts migration into this stream during the winter months. Rock Creek is a moderate-sized, second-order stream that is a tributary of the considerably larger third-order Jellico Creek (upstream from Cumberland Falls), and its watershed drains 17 km² of both private and public land. A dirt and gravel forest road parallels the majority of the stream. Other than the forest road and a few seasonal cabins, there is currently no other development within the watershed. Most of the floodplain and lower mountainsides on the upper 6.5 km of Rock Creek are privately owned, and timber harvest occurs there occasionally. The ridges and upper mountainsides are mostly Daniel Boone National Forest land. The majority of the lower 2.5 km of Rock Creek is on Daniel Boone National Forest land. The upper portion of Rock Creek can become intermittent during dry summers, but groundwater seepage appears to keep water temperatures cool in the intermittent pools. The gradient is considerably higher in the Figure 1. Map of the Big Lick Branch and Rock Creek watersheds in southeastern Kentucky depicting locations of Blackside Dace tagging sites (black dots). Distances between sites, study periods, number of tagged dace, and other details are provided in the text and Table 1. 67 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 lower 2 km of the stream, and a 3-m-high waterfall creates a barrier approximately 1 km upstream from its mouth, restricting fish migration into Rock Creek. Stream-gradient profiles for Rock Creek and Big Lick Branch are provided by Detar (2004). Rock Creek has several relatively undisturbed first-order tributaries, four flowing southward and one flowing northward (Fig. 1). During summer months, discharge in these tributaries was substantially reduced, but some Blackside Dace remained in these streams all year. Similar to the upper portion of Rock Creek, the small tributaries apparently have sufficient groundwater connections during summer to keep water cool enough in intermittent pools for dace to persist. Methods Tagging and trapping Blackside Dace were initially captured in Big Lick Branch, Dace Branch, and Rock Creek and its tributaries using a generator-powered variable-voltage AC backpack electrofisher (in-house design similar to a Coffelt BP-1C). Seining was relatively ineffective for capturing this species due to coarse substrate, and we did not have access to a DC electrofisher when the study was initiated. Upon recovery from electronarcosis, Blackside Dace were anesthetized using a 40-mg/L concentration of clove oil as described by Detar and Mattingly (2005), measured to the nearest mm total length (TL) , and tagged with a visible implant elastomer (VIE; Northwest Marine Technologies, Inc., Shaw Island, WA) injection. Blackside Dace were then placed in an aerated water bucket until fully recovered from anesthesia and released back into one or two pools located approximately in the middle of the site (sites are described below). Dace were batch-tagged with different colors or tag locations at each site so that movement could be detected among sites. The four possible locations used for tagging were on the dorsal surface of the dace (i.e., left and right pre-dorsal-fin origin, left and right postdorsal- fin origin). The second component of monitoring Blackside Dace movement patterns required that fish be captured multiple times; therefore, an efficient yet minimally invasive technique was needed. A pilot study was conducted in July 2002 using minnow traps to capture Chrosomus erythrogaster (Rafinesque) (Southern Redbelly Dace). Baited minnow traps produced higher catch rates than unbaited traps, and thus baited traps were used for capturing Blackside Dace during our movement study. On each monitoring occasion, we counted the number of Blackside Dace while also recording the presence of other fish species in the minnow traps. Number of Blackside Dace per trap was divided by the number of hours the trap was fished to calculate catch per unit effort (CPUE). Recaptured individuals with tags were given a second VIE injection in March 2003 through January 2004 sampling events. These uniquely tagged individuals provided the opportunity to document individual movement histories. Big Lick Branch watershed. In November 2002, Blackside Dace were collected via backpack electrofishing in each of the four 200-m sites in Big Lick Branch J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 68 Vol. 12, Special Issue 4 and tagged with a single VIE injection (Table 1, Fig. 1). Movement was monitored in Big Lick Branch every 6 to 8 wk from February 2003 through March 2004. Twenty-four baited Gee’s model G-40 galvanized wire minnow traps (420 x 190 mm, 20-mm opening, 6-mm mesh) were set throughout the length of Big Lick Branch (two below, two within, and two above each of the four tagging sites). Traps set below and above tagging sites were placed approximately 50–100 m from the downstream and upstream margins of the sites. Each trap was baited with one slice of white bread enclosed in a 0.5-mm-mesh screen and one medium dog biscuit (MILK-BONE ®) broken in half. The mesh screen was used to protect the bread from consumption by fish and to minimize the particles floating downstream of the trap which encouraged fish to enter the trap and increased catch rates. Minnow traps were set starting at Site 1 working upstream to Site 4. Traps were fished for 4.5 to 7.8 hr on each occasion and were checked in the same order in which they were set. In February 2003, Blackside Dace were collected via backpack electrofishing from a 250-m reach of Dace Branch (Table 1, Fig. 1). Blackside Dace were tagged in Dace Branch in an attempt to detect movement from this stream across Table 1. Summary of Blackside Dace (dace) mark-recapture efforts at sites in the Big Lick Branch and Rock Creek watersheds. Dace at each site were batch-marked with visible implant elastomer tags using a different tag color or body location. Minnow traps were used to recapture dace unless noted otherwise. Distance in stream kilometers from the mouth of each stream to mainstem tagging sites or tributary confluences is provided in the third column (site locations are illustrated in Fig. 1). Maximum movement distances (nearest 50 m) that could be detected in a downstream (-) or upstream (+) direction from each site are given in the fourth column. In the last two columns are the number of trapping events and number of dace initially tagged per site. Stream km = stream km from mouth. Events = number of trapping events. Dace = number of dace tagged. Max. detectable Study site Study duration Stream km movement (m) Events Dace Big Lick Branch watershed Big Lick Branch 1 Nov 2002–Mar 2004 0.60 -100, +2400 8 50 Big Lick Branch 2 Nov 2002–Mar 2004 1.45 -950, +1550 8 50 Big Lick Branch 3 Nov 2002–Mar 2004 2.05 -1550, +950 8 50 Big Lick Branch 4 Nov 2002–Mar 2004 2.90 -2400, +100 8 50 Dace Branch* Feb 2003–Mar 2004 n/a n/a 8 30 Rock Creek watershed Rock Creek 1 Feb 2003–Mar 2004 2.75 -100, +5950 7 24 Litton Branch Mar 2003–Mar 2004 3.05 -450, +5600 6 39 John Anderson Branch Feb 2003–Mar 2004 4.35 -1850, +4350 7 110 Rock Creek 2 Feb 2003–Mar 2004 5.75 -3100, +2950 7 50 Sid Anderson Branch Mar 2003–Mar 2004 6.20 -3600, +2550 6 115 Rock Creek 3 Feb 2003–Mar 2004 8.25 -5600, +450 7 50 Lot Hollow Branch Mar 2003–May 2003 8.40 -5800, +350 1 5 Rock Creek 4 Feb 2003–Mar 2004 8.60 -5950, +100 7 30 Total (both watersheds) 653 *Seining was used instead of minnow traps at Dace Branch; movements within Dace Branch itself were not monitored. 69 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 the embayment of Lake Cumberland and into Big Lick Branch or vice-versa. We seined the study reach in Dace Branch each time minnow traps were set in Big Lick Branch to recapture possible migrants from Big Lick Branch. We did not monitor movement of dace within Dace Branch itself because the more remote location of this stream precluded setting and checking minnow traps. Rock Creek watershed. In February 2003, we used backpack electrofishing to collect 154 Blackside Dace from the four Rock Creek mainstem sites plus 32 individuals from John Anderson Branch (Table 1, Fig. 1). In March 2003, we collected an additional 159 Blackside Dace in the other three southward-flowing first-order tributaries plus an additional 78 Blackside Dace in John Anderson Branch. Following the methodology used in Big Lick Branch, dace were batchtagged with VIE injections so that movement could be detected among sites in Rock Creek and its tributaries. Movement was monitored in the Rock Creek system every 6 to 8 wk from March 2003 through March 2004. Following the same approach used in Big Lick Branch, 24 baited minnow traps were set throughout the length of Rock Creek (two below, two within, and two above each of the four sites) during each sampling period. In addition, 3 to 4 traps were set in each of the four first order tributaries to Rock Creek. However, during the first monitoring period (March 2003), traps were only set in the Rock Creek mainstem and John Anderson Branch. Traps set below and above tagging sites were placed approximately 20–100 m from the downstream and upstream site margins. Traps were set starting at Rock Creek Site 1 working upstream (including tributaries as they were encountered) to Rock Creek Site 4, were fished for 4.6 to 8.3 hr on each occasion, and were checked in the same order in which they were set. Movement and environmental variables All distances moved by Blackside Dace in Big Lick Branch, except for the single documented intertributary movement, were measured to the nearest 1 m using a 50-m fiberglass tape measure. The distance of the intertributary movement was estimated to the nearest 10 m using Maptech Terrain Navigator digital topographic map software. Similarly, distances ≤615 m in Rock Creek were measured to the nearest m using the 50-m fiberglass tape measure, whereas distances >615 m were estimated to the nearest 10 m using the Maptech software. For double-tagged individuals, only the first movement distance was used for analysis. Because Blackside Dace were released in one or two pools located approximately in the middle of each 200-m site when they were originally tagged, distance moved was measured from the upstream margin of each site for upstream migrants and from the downstream margin of each site for downstream movement. Therefore, movement measurements are conservative because a dace would need to move a minimum of approximately 100 m in either direction from the center of a site to be considered a migrant. Six of the eight environmental variables were measured in only one location of the watershed on each sampling date for monitoring Blackside Dace J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 70 Vol. 12, Special Issue 4 movements (Table 2). Discharge was evaluated in a section of the stream that best exhibited a uniform, bowl-shaped channel. A Marsh-McBirney Flo-Mate model 2000 or model 201D portable flow meter and top-setting wading rod were used to measure water velocity at 60% of stream depth at multiple points along a transect perpendicular to flow. From these data, discharge was calculated using the velocity-area method outlined by Gallagher and Stevenson (1999). Dissolved oxygen, conductivity, and temperature were measured using a Yellow Springs Instrument (YSI) Model 85 meter. Turbidity was measured using a HF Scientific MicroTPI turbidimeter, and pH was documented using an Oakton Instruments pHTestr 3+ meter. The remaining two environmental variables, days since tagging and day length, were derived from basic calendar data. Statistical analyses Two nonparametric one-way ANOVAs (i.e., Kruskall-Wallace test; Kruskall and Wallace 1952) were used to determine if there was a significant difference in the distances moved upstream versus downstream in either Big Lick Branch or Rock Creek. A nonparametric two-way ANOVA (i.e., Friedman Test; Friedman 1937) was used to determine if there was a significant difference in the mean overall distance moved in Big Lick Branch versus Rock Creek. The two factors in the Friedman Test were (1) stream (Big Lick Branch, Rock Creek) and (2) movement direction (upstream, downstream). Table 2. Variables used in Pearson correlation analyses to determine if environmental variables were correlated with Blackside Dace (dace) movement variables in either the Big Lick Branch or Rock Creek watersheds. Three variables were only evaluated in Rock Creek and are identified with an asterisk (*). Environmental variables Movement variables Days since tagging Total number of dace moved Discharge Number of dace moved upstream Dissolved oxygen Number of dace moved downstream pH Number of resident dace Day length Movement ratio (number of migrants / number of residents) Specific conductivity Dace captured per trap hr (Blackside Dace CPUE) Turbidity Percentage of tagged dace in each sample (number of tagged dace Water temperature captured / total number of dace captured) Mean overall distance moved Mean distance moved upstream Mean distance moved downstream Coefficient of variation (CV) of overall mean distance moved CV of mean distance moved upstream CV of mean distance moved downstream *Number of dace moved into the first-order tributaries from Rock Creek *Number of dace moved out of the four first-order tributaries into Rock Creek *Tributary ratio (number of dace moved in or out of tributaries / total number of dace moved) 71 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 Pearson correlation analyses were used to determine if movement variables were correlated with environmental variables in either of the Big Lick Branch (21 total variables) or Rock Creek (24 total variables) watersheds (Table 2). Significance was determined at α = 0.05, and the sequential Bonferroni technique was used to reduce experiment-wise error (Holm 1979). Results Trapping Catch per unit effort averaged 2.00 ± 0.83 (mean ± SD) Blackside Dace per trap hour (range = 0.83–3.13 fish per trap hour) in Big Lick Branch and 2.82 ± 0.87 Blackside Dace per trap hour (range = 1.19–3.80 fish per trap hour) in Rock Creek (Fig. 2). In Big Lick Branch, CPUE generally increased from February 2003 to July 2003, when it reached a maximum and then began to decline, whereas CPUE generally increased over time in Rock Creek. The percentage of tagged Blackside Dace in each minnow-trapping event (i.e., number of tagged Blackside Dace captured in the trapping event divided by the total number of Blackside Dace captured in the trapping event) averaged 5.8 ± 1.6% (range = 4.0–7.9%) in Big Lick Branch and 2.8 ± 0.5% (range = 1.75–3.3%) in Rock Creek. Big Lick Branch Most tagged dace (81%) were recaptured within the 200-m stream reach of their original tagging (Fig. 3, Table 3). However, five individuals were recaptured Figure 2. Summary of Blackside Dace (BSD) catch per unit effort (CPUE; number of fish per trap hour) in the Big Lick Branch and Rock Creek watersheds from February 2003 to March 2004. Eight sampling trips were made in Big Lick Branch and seven in Rock Creek using baited minnow traps to capture fish. J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 72 Vol. 12, Special Issue 4 downstream and 17 were recaptured upstream of their original site of tagging. Of these migrants, several individuals moved considerable distances from their original site (Fig. 4). Three individuals moved >400 m upstream, while two Figure 3. Summary of individual recapture locales of tagged Blackside Dace in the Big Lick Branch and Rock Creek watersheds from February 2003 to March 2004. Recapture locales were scored as either (1) upstream from the original tagging site (upstream migrants; white stacked bars), (2) within the original 200-m-long tagging site (marked residents; gray stacked bars), or (3) downstream from the original tagging site (downstream migrants; black stacked bars). 73 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 moved >100 m downstream. The mean distance moved upstream (148 ± 138 m) in Big Lick Branch was more variable but not statistically different (H = 2.34, P = 0.13) from the mean distance moved downstream (77 ± 29 m) (Fig. 4). The longest upstream movement recorded was 465 m, and the longest downstream movement was 104 m. In addition to the within-stream movements, one individual apparently moved downstream out of Dace Branch and traveled across the embayment of Lake Cumberland and then upstream into Big Lick Branch where it was recaptured. The total length of this movement was approximately 600 m. This observation represents the first intertributary movement documented for this species. Rock Creek While greater movement was documented in the Rock Creek watershed, most (58%) Blackside Dace were still recaptured within the 200-m stream reach of their original tagging (Table 3, Fig. 3). Dace were observed moving in and out of the four southward-flowing first-order tributaries to Rock Creek and nine individuals migrated >1400 m upstream, while two individuals moved >1900 m downstream. Twenty-three individuals were recaptured downstream and 27 were captured upstream of their original site of tagging. The mean distance moved upstream in Rock Creek (733 ± 1259 m) was not statistically different (H = 0.78, P = 0.38) than the mean distance moved downstream (314 ± 617 m) (Fig. 4). However, the mean overall distance moved was significantly greater (F = 17.03, P = 0.0001) in Rock Creek than in Big Lick Branch. The longest upstream movement recorded in Rock Creek was 3990 m, and the longest downstream movement was 2400 m. Movement and environmental variables Blackside Dace movements were correlated with certain environmental variables in both Big Lick Branch and Rock Creek as indicated by Pearson correlation analyses. Six significant correlations (pre-correction P < 0.05) were identified in Big Lick Branch, and 15 were identified in Rock Creek (Detar 2004). The sequential Bonferroni technique reduced the number of significant Table 3. Number (n), percent (%), and location by distance category of tagged Blackside Dace recaptured in minnow traps in the Big Lick Branch and Rock Creek watersheds during 2003–2004. The original 200-m tagging site is represented by distance category “0 m”. Percentages for each distance category were rounded to nearest whole numbers. Big Lick Branch Rock Creek Distance category (m) n Percent n Percent 0 94 81 70 58 10–100 12 10 30 25 101–500 10 9 8 7 501–1000 0 0 2 2 >1000 0 0 10 8 Totals 116 100 120 100 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 74 Vol. 12, Special Issue 4 correlations from 6 to 2 in Big Lick Branch and from 15 to 3 in Rock Creek. Post-correction significant correlations in Big Lick Branch were (1) mean overall Figure 4. Summary of distances moved by Blackside Dace in Big Lick Branch and Rock Creek watersheds from February 2003 to March 2004. White bars indicate upstream migrants, gray bars represent individuals recaptured in their original 200-m-long tagging site, and black bars indicate downstream migrants. Mean ± SD distance moved is shown for each migrant group. 75 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 distance moved * specific conductivity (r = 0.91, P = 0.0041) and (2) movement ratio (number of migrants / number of residents) * day length (r = 0.84, P = 0.0095). Post-correction-significant correlations in Rock Creek were (1) number of dace moved out of the four first-order tributaries into Rock Creek * days since tagging (r = 0.94, P = 0.0017), (2) mean distance moved downstream * discharge (r = 0.92, P = 0.0033), and (3) number moved downstream * days since tagging (r = 0.92, P = 0.0034). Discussion Movement of individual fish maintains the connectivity of populations within watersheds, reduces the potential for local extinctions, provides a means for recolonization, and plays an important role in population genetics and community structure (Jackson et al. 2001, Larson et al. 2002). Documentation of dispersal and movement patterns of stream fishes (e.g., Fausch and Young 1995, Freeman 1995, Gowan and Fausch 1996, Gowan et al. 1994, Matheney and Rabeni 1995, Skalski and Gilliam 2000, Smithson and Johnston 1999, Todd and Rabeni 1989, Young 1994) has challenged the restricted-movement paradigm suggested by Gerking (1959) (i.e., resident stream fish tend to be sedentary and have limited home ranges). Several studies have shown that fish can be quite mobile in small stream systems (Gowan and Fausch 1996, Gowan et al. 1994, Matheney and Rabeni 1995, Todd and Rabeni 1989), whereas others have indicated that stream fish tend to have a small home range throughout the year (Chisholm et al. 1987, Freeman 1995, Heggenes et al. 1991, Hill and Grossman 1987, Skalski and Gilliam 2000, Smithson and Johnson 1999). Smithson and Johnston (1999) reported that a small portion of four fish species—Creek Chub, Fundulus olivaceus (Storer) (Blackspotted Topminnow), Lepomis megalotis (Rafinesque) (Longear Sunfish), and Lepomis cyanellus Rafinesque (Green Sunfish)—in an Arkansas stream completed regular roundtrip exploratory movements outside their original pool of capture. Similiarly, Larson et al. (2002) reported that a small percentage of five fish species—Rhinichthys atratulus (Hermann) (Blacknose Dace), Rhinichthys cataractae (Valenciennes) (Longnose Dace), Cottus bairdi Girard (Mottled Sculpin), Campostoma anomalum (Rafinesque) (Central Stoneroller), and Oncorhynchus mykiss (Walbaum) (Rainbow Trout)—were captured upstream of their home-range areas in a small Appalachian stream. Our results for Blackside Dace also do not fully conform to Gerking’s (1959) restricted-movement paradigm. Although the majority of dace were recaptured in their original tagging sites (81% in Big Lick Branch, 58% in Rock Creek), several individuals were recaptured considerable distances away. These results support a body of fish-movement research suggesting that many stream fish populations are comprised of a relatively large sedentary group and a smaller mobile group (Breen et al. 2009, Freeman 1995, Gowan and Fausch 1996, Heggennes et al. 1991, Rodriguez 2002, Skalski and Gilliam 2000, Smithson and Johnston 1999), although not all studies show support for two such distinct categories within populations (e.g., Alldredge et al. 2011). J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 76 Vol. 12, Special Issue 4 A common problem with mark-recapture movement studies is that recapture sites are often placed in or near tagging sites, thereby making it easier to detect shorter-range movements. Albanese et al. (2003) refer to this bias as “distance weighting” and offer suggestions to reduce or eliminate such bias through better study designs regarding the spatial extent and arrangement of recapture sites. We analyzed trap arrangements for both watersheds to assess how our study design might have produced bias toward detecting movements of different distances. First, for any given trapping event, we determined the frequency of movement distances in both upstream and downstream directions that could be detected from each of the original four mainstem tagging sites in Big Lick Branch, as well as the original seven tagging sites in Rock Creek (excluding Lot Hollow Branch). From these distances, we then created a frequency histogram of detectable movement distances to evaluate distance weighting in each of the two watersheds (Fig. 5). We found that 62–66% of detectable movement distances in both streams were located in the lower two distance quartiles (including the 0-m category), while 34–38% were in the upper two quartiles, indicating that we were more likely to detect shorter-range than longer-range movements, particularly long-range distances >1800 m in Big Lick Branch or >4500 m in Rock Creek (Table 4). For short-range distances, detection of movements between 1–500 m in Rock Creek represented the most heavily weighted category (Fig. 5). Interestingly, we had no chance of detecting movements of certain distances in Big Lick Branch (201–400 m, 1001–1200 m, 1601–2000 m; Fig. 5), given the spatial arrangement of traps in that stream. Our study design was not especially biased toward collecting resident fishes in their original tagging sites, with only 5.6–8.3% of detectable movement distances located in the 0-m category (Table 4, Fig. 5). In sum, our study did contain distance-weighting bias, implying that we slightly overestimated the frequency of shorter-distance movements (10–500 m), especially in Rock Creek, and underestimated the frequency of longer-distance movements (>1000 m) reported Table 4. Frequency distribution of movement distances that could be detected during Blackside Dace recapture events. Distance categories are provided for original tagging sites (0 m) and streamspecific distance quartiles determined by recapture section lengths in Big Lick Branch (2500 m) and Rock Creek (6050 m). The frequency distribution for each watershed is illustrated at a finer resolution in Figure 5. Distance category Frequency (%) distribution of detectable movement distances Big Lick Branch watershed 0 m 8.3 1–600 m 25.0 601–1200 m 29.2 1201–1800 m 25.0 1801–2400 m 12.5 Rock Creek watershed 0 m 5.6 1–1500 m 29.3 1501–3000 m 31.0 3001–4500 m 19.0 4500–6000 m 15.1 77 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 Figure 5. Frequency distribution of movement distances that could be detected during trapping (recapture) events in the Big Lick Branch and Rock Creek watersheds. Distances include both upstream and downstream directions. Distance categories from left are 0 m, 1–200 m, 201–400 m, …, and 2201–2400 m in Big Lick Branch, and 0 m, 1–500 m, 501–1000 m, …, and 5501–6000 m in Rock Creek. The 0-m category represents traps within original tagging sites. J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist 78 Vol. 12, Special Issue 4 in Table 3 for both watersheds. We offer no corrections to Table 3, but encourage the reader to be aware of this slight bias present in our study design. Larson et al. (2002) used the small-mammal dispersal theory (Lidicker and Stenseth 1992) to interpret the movement of individual fish outside of their home-range areas in a small Appalachian stream. Under this theory, individuals that leave and occupy a new home range are classified as dispersers, whereas individuals that leave a home range and then return to it at a later time are considered explorers. Stenseth (1983) noted that dispersers and explorers are typically robust individuals but may be small animals that are healthy. In our study, we only recaptured 3 double-tagged individuals, which limited our ability to identify larger patterns of multiple movements by individuals. However, behaviors consistent with both disperser and explorer categories were present in our study. Blackside Dace that were recaptured long distances away from their original sites (i.e., >1000 m) and the one dace that moved 600 m downstream out of Dace Branch and up into Big Lick Branch could be considered dispersers. An individual from Big Lick Branch that moved upstream 110 m from its original tagging site and later was recaptured 200 m back downstream in its original site could be considered an explorer. The ratio of recaptured migrant to recaptured resident Blackside Dace was positively related to conductivity and day length in Big Lick Branch. In a Virginia stream network, Albanese et al. (2004) also noted a positive relation between the number of Chrosomus oreas Cope (Mountain Redbelly Dace) movements and day length, as well as temperature and flow events. Longer days generally are associated with warmer temperatures and consequently greater metabolic activity in ectotherms. However, we believe the relationship between Blackside Dace migrant ratio and conductivity represents a spurious correlation because Big Lick Branch had relatively low conductivity values that did not vary greatly over the course of our study. In Rock Creek, the number of Blackside Dace moving downstream and the movement out of tributaries were each positively correlated with time since tagging. Passage of time allows mobile individuals to spread away from tagging sites, thereby representing a fairly intuitive relationship. The downstream directionality of the movement is informed by the positive relation with discharge given that flow events might displace or otherwise encourage individuals to move in that direction. As mentioned above, Albanese et al. (2004) also observed a positive relation between flow events and the number of moving Mountain Redbelly Dace. Although we found a relation between the magnitude of flow and downstream distance moved, we could not assess how increasing or declining flows affected Blackside Dace movement. In short, our correlation analyses simply suggest a greater likelihood of dace movement when day lengths are longer, and further studies will be required to better understand other potentially important environmental associations. Density-dependent mechanisms as well as stochastic environmental events undoubtedly influence fish movement patterns (Freeman 1995). Freeman et al. (1988) suggested that periodic long-range movements may allow small stream fishes to respond to variation in resources over a large area and across a variety 79 J.E. Detar and H.T. Mattingly 2013 Southeastern Naturalist Vol. 12, Special Issue 4 of stream habitats. Migrant fish play important roles in adjusting the distributions and abundances of individuals in response to extrinsic and intrinsic factors that affect fitness levels and could be important to maintenance and recovery of imperiled species (Larson et al. 2002). The two Blackside Dace populations studied here, Big Lick Branch and Rock Creek, are among the most robust across the species’ range in Kentucky and Tennessee (Black et al. 2013 [this issue]). It remains uncertain whether the movement patterns observed in these two watersheds would transfer or apply to other populations because of differences in stream habitats among watersheds and possible density-dependent influences on movement behavior. For example, the greater movement distances we observed in Rock Creek might be expected because it is a longer stream than Big Lick Branch. Our recapture section in Rock Creek was 6.0 km compared to only 2.5 km in Big Lick Branch (Fig. 1). Albanese et al. (2009) found movement rate and abundance were the strongest predictors of recolonization and population recovery of fishes experimentally removed from headwater and mainstem sections of a southwestern Virginia stream. Certain Blackside Dace individuals exhibited exceptional movements in our study, which suggests that populations with sufficient densities should be competent colonists if suitable stream corridors are available (e.g., Roberts and Angermeier 2007). We observed dace readily traveling into, from, and between tributaries in both watersheds. Such tendencies are obvious prerequisites to reestablishment of populations after local extinction events. On the other hand, many individuals did not exhibit movement away from tagging sites. This sedentary tendency, coupled with low densities in many populations (Black et al. 2013 [this issue]), may render many populations susceptible to local extinction in the case of a stochastic event, poor year-class strength, or habitat degradation. Therefore, the sustainability of this species hinges on protecting stream habitat quality and promoting connectivity of suitable habitats within and among watersheds harboring Blackside Dace populations. Acknowledgments The US Fish and Wildlife Service provided research funding, and we are grateful for additional assistance provided by the Cookeville Field Office staff. Supplemental support was provided by the Center for the Management, Utilization, and Protection of Water Resources and Department of Biology at Tennessee Technological University (TTU). Completion of the manuscript was facilitated by a TTU Faculty Non-Instructional Assignment during 2011–2012. We wish to thank numerous graduate and undergraduate students that assisted with this project, especially Brena Jones and Anthony Smith. Christine Peterson assisted with manuscript preparation, and Charles Sutherland constructed Figure 1. The manuscript was improved by comments from D.L. Combs, S.B. Cook, the guest editor, and two anonymous reviewers. Literature Cited Albanese, B., P.L. Angermeier, and C. Gowan. 2003. 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