2012 NORTHEASTERN NATURALIST 19(4):653–664 Ecology and Status of the Redbreast Sunfish, Lepomis auritus, in Yoho Lake, New Brunswick Mark D. Gautreau1,* and R. Allen Curry1 Abstract - In Canada, Lepomis auritus (Redbreast Sunfish) is only found in southwestern New Brunswick, which is the northern limit for the species. In 1989, The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) listed it as a species of special concern due to its limited distribution, and changed the status to data deficient in 2008. The purpose of our study is to begin to build a database of ecological information for the Redbreast Sunfish in New Brunswick. The most northern population occurs in Yoho Lake. Population estimates averaged 386 from 2005 to 2007. The average total length and weight were 12.5 ± 3.1 cm and 42.6 ± 27.3 g, respectively. The length-frequency distributions were consistent over the years, suggesting a stable population structure. Most adult and juvenile sunfish were observed in areas of dense vegetation and large woody debris. Snorkeling surveys were conducted from late June to early July 2005–2007 to assess timing and habitat preference of spawning Redbreast Sunfish. The nests averaged 52.4 ± 10.1 cm in diameter and were 43.9 ± 12.2 cm deep. Nest substrate was sandy with small gravel, and only 33% of the nests were associated with a physical structure in the water column. Overall, the population of Redbreast Sunfish in Yoho Lake appears to be healthy and stable. Introduction In Canada, Lepomis auritus L. (Redbreast Sunfish) are only found in southwestern New Brunswick (Fig. 1), which is the northernmost occurrence of the species (Schultz 2004, Scott and Crossman 1998). The Redbreast Sunfish is distinguished from other sunfishes by its long opercular flap (no wider than its eye) and the absence of a colored border (Scott and Crossman 1998). The species is found from Florida to New Brunswick, where it inhabits both rivers and lakes. There has been some debate about the Redbreast Sunfish in New Brunswick. Kendall (1914) listed the freshwater species of Maine by river watershed, but did not indicate that Redbreast Sunfish were observed or reported in the most eastern rivers adjacent to New Brunswick. Scott and Crossman (1959) also indicated that there is uncertainty about the observations of the species in the earliest records of fish distributions in New Brunswick. Lepomis gibbosus L. (Pumpkinseed) was present in New Brunswick in the earliest records (Perley 1852 as reported in Scott and Crossman 1959). Both species co-exist today in New Brunswick and they are often confused. If Redbreast Sunfish were not present at the time of the earliest samples, they would have arrived only via the transfer and stockings of Micropterus dolomieu Lacépède (Smallmouth Bass) in the province (Catt 1949); however, those stocking records do not explain the current distribution of 1Canadian Rivers Institute, New Brunswick Cooperative Fish and Wildlife Research Unit Department of Biology, University of New Brunswick, Fredericton, NB, Canada E3B 6E1. *Corresponding author - mgautrea@unb.ca. 654 Northeastern Naturalist Vol. 19, No. 4 Redbreast Sunfish because there are a number of lakes where Redbreast Sunfish and Pumpkinseed occur, but not Smallmouth Bass, e.g., George and Oram lakes. Given that Pumpkinseed are reported in the earliest records for the same waters, the species co-exist today in these waters, and there is no apparent vector of human introductions, the logical conclusion is that earliest reports of Redbreast Sunfish in New Brunswick are accurate. This presumption is consistent with the conclusions of the Committee on the Status of Endangered Wildlife in Canada (COSEWIC 2008). In 1989, COSEWIC listed the Redbreast Sunfish as a species of concern due to its limited distribution in the country and the potential for impact from anthropogenic influences (Houston 1989). The COSEWIC designation was changed to data deficient in 2008, due to a lack of information pertaining to the sizes and trends in the Redbreast Sunfish populations in Canada. Despite the change in designation, there is still little or no published information on this species in Canada. Globally, it is listed as G5 or secure, and N5 (secure) in the United States (NatureServe 2010). Yoho Lake lies approximately 30 km southwest of Fredericton, NB, Canada, with a well-known, and what appears to be the northernmost established population of Redbreast Sunfish (Fig. 1). The purpose of this research is to: 1) investigate the ecology of the Redbreast Sunfish in Yoho Lake as a starting point in understanding the species at its northern limit, and 2) provide fisheries managers with detailed information needed to manage and appropriately classify this species in a provincial and national context. Figure 1. Location of Yoho Lake and the distribution of Redbreast Sunfish in southwestern New Brunswick, Canada. Shaded watersheds indicate locations with specific water bodies as stars. Shaded stars are the unconfirmed locations. 2012 M.D. Gautreau and R.A. Curry 655 Methods Yoho Lake is 131 ha in size with a mean depth of 3.66 m. Annual water quality analyses for Yoho Lake from 1975 to 1991 indicate that it is well-oxygenated and that it has a median pH of 6.6 (NB Department of Environment 1991). The lake supports a recreational fishery for Smallmouth Bass, stocked landlocked Atlantic Salmon (Salmo salar L.), and Brook Trout (Salvelinus fontinalis (Mitchill)) as well as the Pumpkinseed and nine cyprinid species (R.A. Curry , unpubl. data). Mark-recapture sampling was conducted in mid-September of each year. Sampling gear consisted of 2 fyke nets and 4 Windermere traps. Fyke nets were made of 0.6-cm knotless netting, with a 0.91- x 1.22-m front frame, five 0.91-m diameter hoops with funneled throats, a 0.91- x 22.9-m lead, and two 0.91- x 7.6-m wings. The nets were deployed perpendicular from shore, with the lead attached to shore. Windermere traps are large pod traps, 0.75 m in length, 0.50 m wide, and 0.40 m in height. One end has a funnel into the pod with an opening 0.08 m in diameter. The gear was set for 4 consecutive nights and moved each night to ensure equal catchability of marked and unmarked fish. In 2007, a single effort was conducted with 5 fyke nets and 4 Windermere traps, deployed for 1 night. In 2005 and 2006, the Schnabel Method for multiple censuses was used for estimating the population size; the Peterson Method was used in 2007, with marked individuals from the previous year (Ricker 1975). The captured fish were measured for fork and total lengths (cm), were weighed (g), had several scales removed for later aging, and were given an upper caudal fin mark (hole punch - Guy et. al. 1996) before being released. Condition (K) was calculated using the Fulton condition factor, K = (weight in g / length in mm3) x 100,000 (Anderson and Neumann 1996). In 2005, 10 individuals were retained, and their stomach contents identified and enumerated. Scales from 61 fish were mounted on slides and aged using a dissecting scope and/or microfiche reader. Digital images were taken of 27 scales using a LEICA MZ95 dissecting scope and LEICA DFC 320 camera. These images were used to age the fish, and the distance from the focus to each annulus was measured using the LEICA IM50 software. The scales and images were blindly aged 3 times by the author. Direct proportion back calculation was applied to calculate the fish total length at each age interval (Devries and Frie 1996). Nest surveys were conducted via snorkeling in late June to early July 2005, 2006, and 2007. Shoreline surveys of the entire lake were conducted by boat. The nests were shallow circular holes dug in the substrate and were readily visible. When a nest was observed, the snorkeler entered the water to determine if it was a Redbreast Sunfish nest, confirmed by the presence of the guarding male. These surveys were conducted for 4 to 5 days after the initial Redbreast Sunfish nest was located, covering a different section of lake each day. Three measurements of diameter were taken at each nest using a meter stick and then averaged (cm). Excavated depth (cm) was calculated by subtracting the water depth at the edge from the deepest point in the nest. When present, the distance to structures, i.e., rocks or logs, and other nests, was measured (cm). Qualitative observations of substrate type, surrounding vegetation, presence of eggs or larvae, and fish territoriality (i.e., male 656 Northeastern Naturalist Vol. 19, No. 4 present and guarding) were also recorded. When possible, the guarding male was captured using an aquarium net and measured for total length. Each individual was returned to the nest within 20 sec. In 2006, water temperature and velocity were measured at each nest using the SonTek Flow Tracker. Results A total of 311 Redbreast Sunfish were captured during the September sampling, 130 and 96, for 2005 and 2006, including 16 and 9 recaptures, respectively. In 2007, 85 were captured, including 8 marked fish from 2006. The majority were caught in the vicinity of large woody debris, particularly old Castor canadensis L. (Beaver) lodges, and areas close to dense, aquatic vegetation (water lilies, Nymphaea spp.). The substrate where they were captured was generally a vegetative/ silt mat, and a few sites were sandy or rocky . The average total length for fish captured in September was 12.5 ± 3.1 cm, weight was 42.6 ± 27.3 g, and condition factor was 1.8 ± 0.2 (Table 1). A lengthweight relationship was strong, weight = 0.0137 * weight 3.1092 (R2 = 0.99, n = 286; Fig. 2). The length-frequency distribution separated age classes up to 3+ and corresponded accurately with scale ages (Fig. 3). After age 3+, there is considerable overlap in fish length between years. The total length at age for fish captured in September was 72, 91, 114, 139, 150, and 161 mm for ages 1+ through 6+, respectively (n = 61; aged by scales; Fig. 4). Back-calculated total lengths from scale measurements averaged 36, 64, 88, 111, 127, and 141 mm for ages 1 through 6, respectively, where the total length at age 1 is the size at the end of the first growing season (n = 27; Fig. 5). The Schnabel estimates of population size were 325 (95% CL = 186–447) and 447 (95% CL = 235–1006) in 2005 and 2006, respectively. The Peterson estimate for 2006 using recaptures in 2007 was 744 (95% CL = 253–1233). Table 1. Summary of Redbreast Sunfish data collection from 2005 to 2007 for Yoho Lake, NB (± one standard deviation). Parameter 2005 2006 2007 All Nests (June/July) Number observed 43 62 39 144 Average diameter (cm) 51.4 ± 10.7 51.6 ± 10.3 55.7 ± 8.4 52.4 ± 10.1 Average depth (cm) 35.4 ± 5.5 46.5 ± 13.3 49.0 ± 10.9 43.9 ± 12.2 Excavated depth (cm) 8.4 ± 2.2 6.9 ± 3.1 6.9 ± 2.4 7.3 ± 2.7 Water velocity (cm/s) - 1.6 ± 0.9 - - % with structure 18.6 43.5 36.0 33 Average distance to structure (m) 0.7 ± 0.6 0.3 ± 0.6 0.2 ± 0.4 0.4 ± 0.5 % within 5 meters of other nests 44.2 83.9 75.5 73 Average distance to other nests (m) 2.2 ± 1.3 2.2 ± 1.3 1.9 ± 1.1 2.1 ± 1.2 Fish (September) Number observed 114 87 85 286 Average total length (cm) 12.1 ± 2.6 13.2 ± 3.3 12.3 ± 3.5 12.5 ± 3.1 Average weight (g) 37.7 ± 23.6 51.1 ± 28.0 40.7 ± 29.5 42.6 ± 27.3 Condition factor 1.8 ± 0.1 1.8 ± 0.1 1.8 ± 0.3 1.8 ± 0.2 2012 M.D. Gautreau and R.A. Curry 657 Figure 2. Length-weight relationship for Redbreast Sunfish captured in September from Yoho Lake, NB, Canada (2005 to 2007: weight = 0.0137 * length 3.1092; R2 = 0.99, n = 286 for all years combined). Figure 3. Length-frequency distribution of Redbreast Sunfish captured in Yoho Lake, NB, Canada in September 2005, 2006, and 2007. 658 Northeastern Naturalist Vol. 19, No. 4 Figure 4. Total length at age of Redbreast Sunfish from Yoho Lake, NB, Canada captured September 2005 and 2006 (aged by scales; age 1 = end of second growing season). Figure 5. Back-calculated total lengths at age (± one standard deviation) for Redbreast Sunfish in Yoho Lake sampled in June 2005 and back-calculated total lengths from Georgia and North Carolina. (age 1 = end of the first growing season). 2012 M.D. Gautreau and R.A. Curry 659 Stomach content analyses (n = 10 fish) revealed a diet of mainly aquatic invertebrates. Contents were mainly Amphipoda (50%: based on total numbers in gut), Trichoptera (20%), nematodes (11%), and chironomids (9%) (Table 2). Juvenile Lepomis spp., ants, and beetles were also found (Table 2). One fish accounted for 103 of the 114 amphipods observed. Redbreast Sunfish spawn in late June or early July in Yoho Lake when the littoral zone water temperatures approach 20 oC. Before spawning, there was an increase in number of individuals, presumably males, in the shallow spawning areas. During this period, the males excavate a bowl-shaped depression, but were not actively guarding and were easily frightened from the nest. Courtship and spawning rituals were not observed. Once a male successfully secured eggs in the nest, he strenuously guarded the nest from all other fishes until the larvae dispersed. This guarding period was 7–10 days. Table 2. Food items of Redbreast Sunfish ( n = 10) from Yoho Lake. Food items Number Percent by number Tricoptera, larvae (caddis fly) Phryganeidae 17 7.4 Dipseudopsindae 16 7.0 Lepidostomatidae 4 1.7 Polycentropodidae 1 0.4 Unknown 7 3.1 Total 45 19.7 Odonata, nymphs (dragonfly) Coenagrionidae 3 1.3 Unknown 3 1.3 Total 6 2.6 Ephemeroptera, nymphs (mayfly) Leptophlebiidae 3 1.3 Unknown 2 0.9 Total 5 2.2 Chironomid 20 8.7 Amphipoda (scuds) 114 49.8 Coleoptera (beetles) Gyrinidae 1 0.4 Cerambycidae 3 1.3 Unknown 2 0.9 Total 6 2.6 Nematoda 25 10.9 Hymenoptera Formicidae (ants) 4 1.7 Diptera Unknown 2 0.9 Fish, young of the year Lepomis spp. 2 0.9 Total 229 100 660 Northeastern Naturalist Vol. 19, No. 4 A total of 144 nests were observed during the 3 years (Table 1). The majority of nests (87%) were excavated through the dense vegetative mat, and the remainder in sandy areas. The surrounding substrate varied from sand to a vegetative mat (principally Eriocaulon aquaticum (Hill) Druce [Pipewort]). Nests were located in areas devoid of canopy cover and had an average water depth of 43.9 ± 12.2 cm (average + 1 standard deviation). The nests were in the open or near very sparse, submergent vegetation; only 33% were associated with a cover structure (e.g., rocks or logs). In those cases, the average distance to the structure was 0.4 ± 0.5 m. The average nest diameter was 52.4 ± 10.1 cm, and the average excavated depth was 7.3 ± 2.7 cm. The substrate in the nests was predominately fine gravel and sand, and several (n = 5) with sand and small cobble. Distance to other nests varied between years with 44, 84, and 76% of the nests constructed within 5 m of another nest (mean = 2.1 ± 1.2 m). This close proximity often enticed neighbors to dart at each other, leaving their nest undefended, and on several occasions, Pumpkinseed and non-spawning Redbreast Sunfish were observed taking advantage of the guarding male’s departure to prey on the eggs or larvae until he returned. Captured guarding males (n = 54) were 16.4 ± 1.3 cm TL, and their size was weakly correlated to nest depth (R2 = 0.18) and diameter (R2 = 0.02; Fig. 6). In 2006, velocity measurements for nests averaged 1.6 ± 0.9 cm/s, and temperature was 21.4 ± 0.9 °C. Figure 6. The total length of guarding males and nest diameter and depth for Redbreast Sunfish in Yoho Lake (2005, 2006, 2007; n = 54). 2012 M.D. Gautreau and R.A. Curry 661 Figure 6 shows the back-calculated total lengths at age for Lepomis auritus in Yoho Lake: 36, 64, 88, 111, 127, and 141 mm, respectively, for ages 1 through 6. These were smaller than those from North Carolina (47, 84, 122, 156, 168, and 211 mm, respectively, for ages 1 through 7; Carlander 1977) and Georgia (59, 90, 125, 153, 181, 205, and 222 mm, respectively, for ages 1 through 7; Sandow et al. 1974). Discussion Although often referred to as a “riverine species” preferring intermediate to large rivers (Bass and Hitt 1974), the Redbreast Sunfish is adaptable and is able to thrive in a variety of environmental conditions and habitat types (Houston 1989). In Yoho Lake, individuals were observed or captured in areas of dense, submergent vegetation or near large woody debris with substrates of silt/detritus and sand. The diet samples were few and from one season, but agreed with other populations in the South, i.e., Florida, North Carolina, and Georgia, which reported a diet of primarily aquatic and terrestrial invertebrates (Bass and Hitt 1974, Davis 1972, and Sandow et al. 1974). The large number of amphipods in the stomach of a single fish and the broad range of prey items indicate that this species is most probably an opportunistic feeder. Redbreast Sunfish spawned when water temperatures were approximately 20 °C in Yoho Lake, which is consistent with southern populations (Davis 1972, Sandow et al. 1974). The nest characteristics of Yoho Lake Redbreast Sunfish appear to differ from other populations. Nests in Georgia, Florida, and North Carolina were larger in diameter and located in deeper water (Davis 1972, Helfrich and Nutt 1991, Lukas and Orth 1993, Sandow et. al 1974). Southern populations have larger body sizes, and this characteristic is the presumed explanation for the smaller nest sizes observed in Yoho Lake. Spawning substrate of fine gravel to sand is consistent with reports across the species’ range (Davis 1972, Helfrich and Nutt 1991, Lukas and Orth 1993). In Yoho Lake, Redbreast Sunfish did not select nest sites near structure, whereas, nests were almost always in the vicinity of a structure in river populations studied in the southern United States (Davis 1972, Bass and Hitt 1974, Helfrich and Nutt 1991, Lukas and Orth 1993, Sandow et al. 1974). Habitat suitability indices have suggested that a low, stable flow of around 0.9 cm/s was preferred for nest sites (Helfrich and Nutt 1991, Lukas and Orth 1993). Interestingly, the highest nest concentrations observed in Yoho Lake were at the mouth of two inflow streams and in the lake outflow area, which may have created the flowing water at nest sites we observed. The calculated total lengths at age for Redbreast Sunfish in Yoho Lake were smaller than those from North Carolina and Georgia (Sandow et al. 1974, Carlander 1977). This latitudinal difference widened with age and is most probably related to shorter growing seasons in the north as has been observed in many species including the Pumpkinseed (Conover and Present 1990, Copp et al. 2004, 662 Northeastern Naturalist Vol. 19, No. 4 Garvey and Marshall 2003). Body sizes were similar over the first 2 growing seasons regardless of latitude. This result suggests that Yoho Lake individuals grow more rapidly during their first growing season, likely in preparation for winter (Conover and Present 1990, Garvey and Marshall 2003). The Redbreast Sunfish has garnered an uncertainty about its status as a native species of New Brunswick because of its very restricted and isolated occurrence in Canada (COSEWIC 2008). In addition, Redbreast Sunfish and Pumpkinseed are the only native centrarchids east of central Maine (Curry and Gautreau 2010, Kendall 1914). As discussed earlier, our current evidence supports the opinion that Redbreast Sunfish are native to southwestern New Brunswick and therefore arrived with the other re-invading, freshwater species post-glaciation (Curry 2007). We still know very little about other populations in the area, for example, how they came to be in those places, i.e., natural, dispersed from an original population, or transferred, or the degree of their stability. Dispersal downstream from Yoho Lake is possible and may explain certain present-day occurrences in the Saint John River (Fig. 1). Our unpublished work from George Lake (Fig. 1), which is presumed to be a native occurrence, indicates this population is stable. For the Yoho Lake population, there is no indication that it is currently threatened. The Redbreast Sunfish is not sought after as recreational species. Yoho Lake is highly developed with summer and year-round homes, but the water quality has been stable for the last 10 years (New Brunswick Department of the Environment, Fredricton, NB, Canada, unpubl. data). The population size and structure were consistent between years, and overall our findings suggest that Yoho Lake Redbreast Sunfish population is stable and not at risk at this time. Acknowledgments We would like to thank the following people for field and laborat ory support throughout the project: E. Chernoff, S. Clark, C. Connell, S. Currie, R. Engelbertink, L. Gautreau, J. Hallet, K. Heard, J. Shaw, and R. Smedley. Funding was provided in part by the New Brunswick Wildlife Trust Fund. Literature Cited Anderson, R.O., and R.M. Neumann. 1996. Length, weight, and associated structural indices. Pp. 447–481, In B.R. Murphy and D.W. Willis (Eds.). Fisheries Techniques, 2nd edition. American Fisheries Society, Bethesda, MD. Bass, D.G., Jr. and V.G. Hitt. 1974. 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