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Fish Movement Among Lakes: Are Lakes Isolated?
Robert A. Daniels, Richard S. Morse, James W. Sutherland, Robert T. Bombard, and Charles W. Boylen

Northeastern Naturalist, Volume 15, Issue 4 (2008): 577–588

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2008 NORTHEASTERN NATURALIST 15(4):577–588 Fish Movement Among Lakes: Are Lakes Isolated? Robert A. Daniels1,*, Richard S. Morse1, James W. Sutherland2, Robert T. Bombard2, and Charles W. Boylen3 Abstract - The concept of a lake as an isolated unit is a central theme in research and management of freshwater systems. Support is based on direct observations of lake communities. Studies undertaken in the last several decades lend tacit support because the methods used in both research and management often do not question the underlying notion that lake communities are essentially isolated. In a study of fish assemblages in interconnected lakes, we noted movement of tagged fish among lakes. We also found that species introduced to one lake were later captured in neighboring lakes. We found that fish species in lake assemblages did not differ from those in inlet and outlet stream assemblages; although relative abundance varied, species richness and composition did not. This finding suggests that fish assemblages in lakes are not isolated. Rather, immigration and emigration from streams and other lakes occurs. Although few individuals migrated to new lakes, any movement can affect population structure (e.g., through recolonization, gene flow) and management goals (e.g., spread of disease). Consequently, we suggest that methods commonly used to assess fish assemblages in lakes and the concept of the lake as a management unit may need to be reconsidered. Rather than be treated as isolated populations, fishes in lake communities may be better treated as a watershed-wide metapopulation. Introduction The belief that a lake is an island, initially presented by Forbes in 1887 (c.f. Forbes 1925), is a fundamental tenet of limnology (Magnuson et al. 1998). For over a century, this concept has guided the work of limnological researchers and lake managers, who often treat lakes as individual, isolated units. The belief that lake-dwelling organisms make up a distinct community is an appealing concept in that it greatly simplifies the way limnological and fisheries data are collected and interpreted. The concept further provides a framework for how such data are ultimately used in managing those communities. However, inherent in the concept are assumptions that are becoming increasingly difficult to support. Simply stated, the concept holds that lakes are habitat islands surrounded by different and disconnected habitats. In this scheme, organisms are confined to a single lake; organisms not currently in the lake would not occur there unless they were introduced. Because a lake is functionally isolated, it is implicitly assumed that the species in the lake community must be relatively sedentary. Movement within the lake is recognized as possible, but 1New York State Museum, CEC 3140, Albany, NY 12230. 2New York State Department of Environmental Conservation, 625 Broadway, Albany, NY 12233. 3Darrin Freshwater Institute, Rensselaer Polytechnic Institute, Troy, NY 12180-3590. *Corresponding author - rdaniels@mail.nysed.gov. 578 Northeastern Naturalist Vol. 15, No. 4 emigration from the lake is not. This viewpoint ignores characteristics of one prominent component of many lake communities—fish. North American fishes, with few exceptions, are stream dwellers; few are obligate lake dwellers (Moyle and Cech 1996). Moreover, the long-held notion that fish are relatively sedentary (Gerking 1959) is becoming less defensible (Fausch and Young 1995, Gowan et al. 1994, Rodríguez 2002). With a growing body of information, the concept of the isolated, individual lake has been questioned, and many recognize the connectivity that exists among lakes, streams, and landscapes (Magnuson and Kratz 2000). We report results here that support this contention. Several kinds of studies depend upon and support the notion that distinct fish assemblages exist in lakes (e.g., Magnuson et al. 1998). The basic design of such studies is similar. Lakes are either sampled with several types of gear for a relatively short period, or a single lake is sampled repeatedly over a long period (Magnuson and Kratz 2000). Several assumptions about the resulting data are adopted, but not often tested. Studies assume that (1) the temporally-limited data fairly represent the species composition of the lake and (2) with these data as a basis, future changes in the species composition of the lake can be explained. Olden et al. (2006) examined these assumptions and, using a long-term data set from lakes in Wisconsin, found general support for both assumptions. They noted, however, that assessing community structure in lakes can be complicated by the environmental factors that affect composition and the spatial and temporal extent of the data. Both assumptions can be supported as demonstrated by Olden et al. (2006), and are reasonable if the lake is truly isolated. However, possible confounding issues include immigration or emigration of fish or the presence of rare species (Magnuson et al. 1994), so results of studies using single-year community assessments need to be carefully assessed and can perpetuate the notion that a lake assemblage is isolated. Since 1995, we have monitored fish populations in connected lakes in the Adirondack Mountains in Herkimer County, NY. Short streams link five of our study lakes (Fig. 1). We examined fish movement among lakes and between streams and lakes in several ways. If lakes are isolated, individual fish should not move among lakes or between lakes and streams. If our results show that fish are not confined to a single lake, then we should reject this hypothesis. Rejection would necessitate adjustments to many current fisheries-management practices and research protocols. Researchers and managers would need to view lakes, and the biota found therein, as being part of a broader landscape and biotic community. Methods A tagging program was initiated in 1995 to examine annual and seasonal changes in the population size of fishes in the study lakes: Rondaxe, Dart, and Moss. All are located in the Moose River watershed within the Saint 2008 R.A. Daniels, R.S. Morse, J.W. Sutherland, R.T. Bombard, and C.W. Boylen 579 Lawrence River drainage. All are relatively small (mean surface area = 63 ha) and are situated in a landscape dominated by mixed conifer and deciduous forest. Each lake has inlet and outlet streams and the distance between lakes averages 3.5 km. Two other lakes, Big Moose and Cascade Lakes, are also connected to these three lakes by short stream segments; we also sampled these two lakes. Other characteristics of the streams and lakes are included in Figure 1. Fish were captured in trapnets, identified to species, and counted. Fish were measured (standard length [SL], in mm), and individuals in five species over a minimum length were tagged with sequentially numbered anchor tags and released. The five species that we tagged were: Ameiurus nebulosus (Lesueur) (Brown Bullhead, over 80 mm SL), Catostomus commersonii (Lacepède) (White Sucker, over 90 mm SL), Ambloplites rupestris (Rafinesque) (Rock Bass, over 75 mm SL), Lepomis gibbosus (Linnaeus) (Pumpkinseed, over 75 mm SL), and Perca flavescens (Mitchill) (Yellow Perch, over 90 mm SL). Other species were identified, counted, measured, and released. Sampling was done in spring and autumn so that fish would be handled in cooler temperatures and mortality minimized (Stickney 1983). Tagged and recaptured individuals provided one basis for evaluating fish movements among lakes and between lakes and streams. In general, anchor tags are retained well (Wydoski and Emory 1983) and were embedded in the musculature of each of these species equally well. Figure 1. Lakes in the upper Moose River watershed, Adirondack Mountains, Hamilton and Herkimer counties, NY. At each lake, the surface area of the lake (ha), the elevation (m), and the area of the drainage basin (km2) are listed. For each stream segment, the average gradient (m/km) and segment length (km) are listed. 580 Northeastern Naturalist Vol. 15, No. 4 Inlet and outlet streams (Fig. 1) were sampled in summer 1998, 2001, and 2004 with either a backpack electroshocker or 6-m bag seine (bag with 3 mm bar, wings with 5 mm bar). Stream sampling was conducted in reaches from 50 to 100 m in length and that included riffle, run, and pool habitats. All fish in stream collections were identified to species and counted. Most fish were returned to the stream. Individuals not returned alive to the lake or streams are vouchered at the New York State Museum. In addition, unanticipated fish introductions occurred during our monitoring studies that allowed us to test the fish movement hypothesis using a second, independent source of information. Micropterus salmoides (Lacepède) (Largemouth Bass) was introduced into Lake Rondaxe in stocking events in 1998 and 1999. This species is not native to Adirondack lakes and was absent from the system until the stocking events occurred. The lake association reported that 500 fingerling fish were stocked in the spring each of the two years. Two other species were stocked into the system in the mid-1990s: Osmerus mordax (Mitchill) (Rainbow Smelt) was introduced into Moss Lake by 1995, and Micropterus dolomieu (Lacepède) (Smallmouth Bass) was stocked into Lake Rondaxe in 1997. The introduction of Rainbow Smelt was not sanctioned; our information rests on the date of first appearance of the species in catches and/or is gleaned from anecdotal information from local sources. Results Recapture of tagged fish Recapture locations of previously tagged fish provided information on movement within lakes and among them. During the study, we tagged 33,875 fish in Moss, Rondaxe, and Dart lakes and recaptured 4649 fish at least once. Several individuals were recaptured more than once, making the recapture rate 20.4%. Between 1995 and 2006, 43 individual fish moved among lakes. Fifteen White Suckers, 6 Brown Bullheads, and one each of Rock Bass, Pumpkinseed, and Yellow Perch were tagged in a downstream lake and moved upstream. Thirteen White Suckers, one Brown Bullhead, and one Yellow Perch moved downstream from one of the upstream lakes. Three White Suckers and one each of Yellow Perch and Brown Bullhead were tagged in one upstream lake, moved downstream to Lake Rondaxe, and were recaptured in the other upstream lake (Table 1). The shortest time between tagging in one lake and recapture in a different lake was 21 days; this by two White Suckers tagged in late spring 1997. Other fish recaptured in a lake different from the one where they were originally tagged were taken from 117 to 1808 days after tagging. Fish tagged in each of the years from 1995 to 2005 were recaptured in a lake different than the one in which it was tagged originally between 1996 and 2006. Of all recaptured fish, 0.9% emigrated from the lake in which it was tagged initially. White Sucker emigrated most frequently, and Pumpkinseed and Rock Bass least frequently. Table 1 provides a summary of fish movement among lakes. 2008 R.A. Daniels, R.S. Morse, J.W. Sutherland, R.T. Bombard, and C.W. Boylen 581 Table 1. Movement of tagged fish among Moss, Rondaxe, Dart, and Big Moose lakes, Herkimer County, NY, 1995–2005. Direction arrows refer to upstream (↑) and downstream (↓) movement; movement between Moss and Dart lakes required both upstream and downstream (see Fig. 1). Catostomus Ameiurus Perca Lepomis Ambloplites Source lake - commersonii nebulosus flavescens gibbosus rupestris Total Destination lake Direction (White Sucker) (Brown Bullhead) (Yellow Perch) (Pumkinseed) (Rock Bass) Mean days Range (days) Fish Moss - Rondaxe ↓ 8 1 729 117–1537 9 Rondaxe - Moss ↑ 2 2 1 1307 369–3251 5 Rondaxe - Dart ↑ 13 3 1 1 500 21–1076 18 Dart - Rondaxe ↓ 5 1160 371–1808 5 Dart - Big Moose ↑ 1 481 481 1 Dart - Moss ↓↑ 2 1 1 512 357–862 4 Moss - Dart ↓↑ 1 340 340 1 Total 31 8 2 1 1 43 Mean days 710 862 371 369 713 715 Range 21–1808 255–3251 368–373 369 713 21–3251 Total fish recaptured in Moss Lake 599 543 788 71 0 2001 Total fish recaptured in Dart Lake 451 231 534 39 179 1434 Total fish recaptured in Lake Rondaxe 315 359 338 101 101 1214 582 Northeastern Naturalist Vol. 15, No. 4 Fish movement subsequent to a fish introduction In spring 1998, 500 Largemouth Bass were released into Lake Rondaxe; an additional 500 fish were stocked in spring 1999. In autumn 1998, we began to catch Largemouth Bass in Lake Rondaxe, and it since has become an important component of the lake assemblage (Fig. 2). In 2000, Largemouth Bass was first collected in the upstream Dart Lake, and the number and relative abundance of Largemouth Bass in Dart Lake has continued to increase (Fig. 2). Largemouth Bass was first collected in Moss Lake in 2003, and the number of individuals and relative abundance increased in subsequent years (Fig. 2). Several size classes have been caught in both lakes, which suggest annual recruitment, although the extent of in-lake reproduction in either upstream lake is not known. Annual upstream migrations may continue. Smallmouth Bass was captured in Rondaxe Lake in 1997, 1998, 2001, and 2005 but none has been taken in either Dart or Moss Lakes. Rainbow Smelt was collected initially in Moss Lake in 1995. This species was captured in Moss Lake every autumn since 1997. Relative abundance has varied between 0.1 and 0.8%. In 2000, an individual (SL = 106 mm) was captured in Rondaxe Lake. Stream fish assemblage Seventeen fish taxa inhabit inlet and outlet streams associated with each of the study lakes. Eighteen taxa have been collected in the three lakes (Table 2). One species, Culaea inconstans (Kirkland) (Brook Stickleback), has Figure 2. Relative abundance of Largemouth Bass in three upland lakes in the Moose River watershed, Adirondack Mountains, NY, 1998–2006. Based on catches from summer and fall samples in Rondaxe, Moss, and Dart lakes. 2008 R.A. Daniels, R.S. Morse, J.W. Sutherland, R.T. Bombard, and C.W. Boylen 583 Table 2. Relative abundance of species in fish assemblages in Lake Rondaxe, Moss Lake, Dart Lake and their inlet and outlet streams. The outlet streams of Moss and Dart lakes are also the inlet streams of Lake Rondaxe. Relative abundance of fish in each of the lakes represents a total abundance based on samples from 1995 to 2004. Annual and seasonal variation in the lakes is masked in these numbers, but dominant species are relatively consistent across years. Numbers with “*” represent species that clearly dominate the assemblage. + indicates presence in small numbers. A. n. = Ameiurus nebulsosu, C. c. = Catostomus commersonii, N. c. = Notemigonus crysoleucas, R. a. = Rhinichthys atratulus, L. c. = Luxilus cornutus, S. a. = Semotilus atromaculatus, L. x S. = Luxilus x Semotilus, P. e. = Phoxinus eos, S. f. = Salvelinus fontinalis, S. n. = Salvelinus namaycush, O. m. = Osmerus mordax, U. l. = Umbra limi, F. d. = Fundulus diaphanus, C. i. = Culaea inconstans, A. r. = Ambloplites rupestris, L. g. = Lepomis gibbosus, M. d. = Micropterus dolomieu, M. s. = Micropterus salmoides, and P. f.= Perca flavescens. Number of Location samples A. n. C. c. N. c. R. a. L. c. S. a. L. x S. P. e. S. f. S. n. O. m. U. l. F. d. C. i. A. r. L. g. M. d. M. s. P. f. Moss Lake 11.8 12.5 3.9 0.1 48.0* 0.9 + 0.1 0.1 + 0.3 + + 8.5 0.2 13.6 Dart Lake 7.4 15.2 4.8 1.5 1.5 + 0.1 + 0.5 6.5 6.4 0.1 0.3 55.6* Lake Rondaxe 17.2 18.1 9.8 5.6 0.8 + + + + + 5.8 15.5 + 0.5 26.6 Moss inlet 1998 2 0.5 33.0 10.7 14.1 1.5 5.3 34.5 0.5 2001 2 2.2 0.9 7.6 0.9 4.2 0.4 2.2 0.4 80.7* 0.4 2004 1 2.3 30.7 8.0 3.4 3.4 48.9* 1.1 1.1 1.1 Dart inlet 1998 1 1.9 32.7 53.8* 1.9 9.6 2001 1 2.6 23.1 5.1 69.2* 2004 0 Moss outlet 1998 2 1.0 5.0 4.0 0.5 24.6 63.8* 1.0 2001 2 1.2 1.2 1.2 4.9 4.9 22.0 9.8 14.6 40.2* 2004 1 14.3 4.8 28.6 33.3 14.3 4.8 Dart outlet 1998 1 75.0 25.0 2001 2 9.5 14.3 28.6 19.0 14.3 9.5 4.8 2004 1 86.7* 13.3 Rondaxe outlet 1998 1 12.7 0.8 11.9 7.6 3.4 9.3 3.4 16.9 2.5 31.4* 2001 1 6.8 2.9 23.3 10.7 8.7 19.4 1.0 27.2 2004 1 1.5 19.4 4.5 6.0 11.9 17.9 38.8* Present in streams x x x x x x x x x x x x x x x x x Present in lakes x x x x x x x x x x x x x x x x x x 584 Northeastern Naturalist Vol. 15, No. 4 been taken in stream samples only and two (Rainbow Smelt and Salvelinus namaycush (Walbaum) [Lake Trout]), have been collected only in lakes. The species that compose the stream and lake assemblages are similar. Relative abundance varies, however. In the lakes, Brown Bullhead, White Sucker, Luxilus cornutus (Mitchill) (Common Shiner), Pumpkinseed, and Yellow Perch were the species most frequently taken. These species were rarely taken in streams, where Umbra limi (Kirtland) (Central Mudminnow), Semotilus atromaculatus (Mitchill) (Creek Chub), Rhinichthys atratulus (Hermann) (Eastern Blacknose Dace), and Salvelinus fontinalis (Mitchill) (Brook Trout) dominated upstream sites and where the centrarchids and Yellow Perch were more common downstream of Lake Rondaxe (Table 2). Neither black bass species was collected at any stream sites in 1998. By 2001, Largemouth Bass was taken downstream of Lake Rondaxe and in the North Branch, Moose River between Lake Rondaxe and Dart Lake. By 2004, Largemouth Bass was taken downstream of Lake Rondaxe, between Lake Rondaxe and Moss Lake and upstream of Moss Lake. Discussion Three different types of information support the contention that Adirondack lake fish assemblages are not isolated either from the interconnecting streams or nearby lakes: recapture of tagged fish, capture data on fish movement subsequent to introductions, and the composition of the lake and stream assemblages. Overall, 0.9% of recaptured fish were taken in lakes other than the one in which they were initially caught (Table 1). White Sucker was the most likely species to be recaptured in a different lake (2.7%). The 43 fish with catch histories detailed in Table 1 emigrated from one lake to another. Although a relatively small percentage of the total number of recaptured tagged fish emigrate, the presence of any emigration demonstrates that at least a small part of each population regularly disperses, which also has been noted in other fish populations (Petty and Grossman 2004, Smithson and Johnston 1999). Fish movement is key to rejecting the idea of lake-specific fish assemblages. Researchers recognize that some fish move, although the behaviors often are identified as specialized as noted in Gowan et al. (1994). For example, fish are known to migrate to feed or spawn (Josephson and Youngs 1996, Raney and Webster 1942) or minimize threats (Fraser et al. 2006). Certain species move in response to seasonal changes (Josephson and Youngs 1996, Meyers et al. 1992). Our data indicate, however, that the movement behavior of at least some individuals of some species does not fall clearly into these categories. White Sucker, for example, is a fish that migrates upstream to spawn (Raney and Webster 1942), and because it was the species that emigrated most frequently in our study, its movement perhaps might be explained as part of its spawning behavior. However, two individuals clearly were tagged after the completion of the spring spawning run and were recaptured in a different lake before the beginning of the next run. Furthermore, 13 individuals 2008 R.A. Daniels, R.S. Morse, J.W. Sutherland, R.T. Bombard, and C.W. Boylen 585 migrated downstream rather than upstream, which is unexpected if the emigration was accidentally the result of a spawning run. Thus, almost half of the individuals in the sample that were documented to have moved between lakes did not behave in a way typical of spawning White Sucker. Rainbow Smelt migrate from lakes into tributaries to spawn (Langlois 1935). The individual migrant was taken downstream of the source lake— a behavior not associated with a spawning run. Pumpkinseed, Rock Bass, and Brown Bullhead, the other species that migrated between lakes, spawn in nests in lakes (Smith 1985). None of our marked fish was young, so the observed behavior was not associated with out-migration of young fish, a search for nursery or rearing habitat, or any movement associated with early life history. In fact, all the individual fish that immigrated to a second lake were adults when tagged. Although information to the contrary is mounting (e.g., Gowan and Fausch 1995, Neely and George 2006), fisheries biologists and managers often accept that fish, particularly as adults, spend their lives in relatively small areas. This assumed sedentary nature of freshwater fish (Gerking 1959), implicit in the belief that lakes are isolated, has influenced management strategies. Our data corroborate other studies that suggest that lengthy movements of individual fish, even if only a small part of the population, are not unusual (Coombs and Rodríguez 2007, Gowan et al. 1994). Reports of fish movement among lakes have been noted for over a century. For example, what was believed to have been a single point introduction of Yellow Perch in the Moose River watershed, NY, was followed by a rapid expansion into all neighboring lakes within a decade (Mather 1886). Mather (1886) also reported the multi-lake expansion of Smallmouth Bass from a single-lake introduction during the same period. Inter-lake movement of 0.9% of the individuals of a community would seem to have little effect on community structure and the population ecology of any of the study lakes. Other aspects of the biology of the populations involved, however, are affected. For example, dispersal among lakes can maintain genetic similarity among neighboring populations and dispersing fish can act as vectors in dispersal of diseases and parasites. Dispersing fish can replenish declining or extirpated populations, or, as in the case of Largemouth Bass described here, can serve as the vanguard of an invasion of an exotic species. The presence of Largemouth Bass in upstream lakes and the presence of both Largemouth and Smallmouth Bass in stream samples soon after their introduction into Rondaxe Lake is strong evidence of rapid out-migration from a single point source. Although unsanctioned stocking of upstream lakes is possible, the presence of Largemouth Bass in streams between the lakes suggests that migration between lakes is occurring and that any boundary between streams and lakes, if present at all, is porous (Jackson et al. 2001). In effect, the individual lake populations examined here can be considered local habitat patches of a metapopulation, i.e., discrete populations, 586 Northeastern Naturalist Vol. 15, No. 4 largely unaffected by each other, but with some inter-population interaction through inter-lake movement of individuals (Hanski 1999). In this definition of metapopulation, high extinction rate of local populations is not a defining characteristic, which makes the concept more useful in examining community relationships in aquatic sciences (Kritzer and Sale 2004). As an analytical approach to assess the dynamics of each local patch and also of the regional network, these lakes can serve as an effective metapopulation model because of the demonstrated low level of exchange among the local populations. The model will become increasingly valuable to managers as populations in patches (= lakes) become stressed by environmental disturbances, such as an invasive species or exotic diseases, and protection of individual populations becomes more dependent upon the patch network (= watershed). It is important that managers re-examine their treatment of lakes as management units. Evidence (e.g., the data in this report, Jackson et al. 2001) suggests that the convenience of accepting this approach does not offset the potential damage that can occur from its use. The rapid dispersal of the exotic Largemouth Bass into neighboring lakes and streams after an approved introduction into one lake demonstrates the need to consider the effect on the watershed when planning stocking, reclamation, or species protection projects. A second basic consideration is that, when dealing with management of drainage lakes, inlet and outlet streams need to be a part of the management plan. Ability to disperse through streams is dependent upon the characteristics of both the species and the stream (Olden et al. 2001), but effective management of the lake fishery is tied, in part, to the tributaries. Finally, data should be collected in a way that allows managers to make assessments at the watershed level. Although the need to use different devices to capture different species and life stages is widely recognized (e.g., Jackson and Harvey 1997), repeated sampling of the entire assemblage is also needed. In addition, lake surveys often do not include associated stream sampling. Repeated surveys of lake and stream sites across the watershed are useful and can provide the temporal and spatial detail needed for development of effective management plans. This additional sampling may not require additional resources (see Olden et al. 2006), but will require that available resources be used to maximize information obtained from appropriate sampling protocols. Our survey work demonstrates that some fish emigrate from the lake in which they were tagged to other lakes in the system. We have also followed the out-migration to neighboring lakes from point introductions of two exotic species. Finally, our data suggest that the stream and lake assemblages are generally composed of the same species and, presumably, individual fish move between the two macrohabitats in their inter-lake migrations. Future studies should be designed to incorporate methods that allow detection of fish movement, including unique marks on individual fish and repeated surveys. We suggest that the concept of the lake as an island, or more specifi- cally as a unique management unit, is too simple. Lake populations need to be treated as dynamic components of a metapopulation. 2008 R.A. Daniels, R.S. Morse, J.W. Sutherland, R.T. Bombard, and C.W. Boylen 587 Acknowledgments We thank all the volunteers, interns, students, and colleagues who participated in the field work over the past decade, particularly B.R. Weatherwax and D.A. Bloomquist. R.E. Schmidt, J.A. Tyler, and T.J. Sullivan reviewed drafts of this manuscript, and we appreciate their valuable comments and suggestions. 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