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. This work was
supported in part by EPA contract 68D20171.
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