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Micro-elemental analysis of statoliths as a tool for tracking tributary
origins of sea lamprey
Stuart Ludsin
Collaborators
J. Ellen Marsden (U of Vermont)
Brian J. Fryer (U of Windsor)
Executive Summary
During
2004, collections of larval, parasitic, and adult lamprey were made from
numerous tributaries of Lake Huron, as well as the open lake, by our partners
(US Fish and Wildlife Service and Canadian Department of Fisheries and
Oceans). Larvae (n = 15 per stream) from ~30 Lake Huron streams have been
processed using laser-ablation inductively coupled plasma mass spectrometry
(LA-ICPMS). Our preliminary results, which were presented in a poster
at the annual American Fisheries Society meeting in Anchorage, AK, suggest
that site discrimination will be possible for many streams, based upon
statolith micro-elemental composition, but not for others. Further analysis,
however, is needed to determine which sites can be discriminated. Similar
larval, parasitic, and adult sea lamprey collections were made by the
USFWS and DFO during 2005, which will be analyzed during FY06.
Scientific Rationale
The
analysis of otolith micro-elemental composition has been a valuable tool
for differentiating between local spawning populations, and identifying
origins of recruits to the fishery. Building on two pilot investigations
conducted in lakes Champlain and Huron, we will determine whether trace
elements incorporated into sea lamprey statoliths during larval stream
residence can be used to discriminate among local populations in Lake
Huron. Assuming discrimination is possible, we will begin to identify
origins of parasites and spawners in Lake Huron, focusing on tributaries
(e.g., St. Marys River) important to US and Canadian agencies. We also
will develop predictive relationships between water and statolith chemistry,
which eventually could help eliminate the need to sample larvae annually
to develop stream-specific signatures. By using statolith elemental “signatures”
as a natural environmental tag, we aim to provide an alternative means
to identify spawning origins of spawners and parasites, which currently
can only be acquired using labor-intensive tagging studies that yield
low tag returns. Beyond helping assess whether recent sea lamprey control
efforts (e.g., St. Marys River) have been successful, an ability to identify
natal origins would pave the way for studies aimed at understanding attributes
(e.g., stream-specific growth rates, sex ratios, and movement) that promote
survival to parasitic and spawning stages, as well as help better target
streams for control measures.
Background
Of the numerous anthropogenic perturbations experienced by the Laurentian
Great Lakes during the past century, arguably the most costly, from both
ecological and economic standpoints, has been invasion of the sea lamprey
(Petromyzon marinus). By parasitizing large-bodied native fishes,
sea lampreys have greatly altered the ecology of the Laurentian Great
Lakes, as well as other coolwater systems within the surrounding basin
(e.g., Lake Champlain). Of primary importance has been the reduction of
native salmonines, including lake trout (Salvelinus namaycush)
and lake whitefish (Coregonus clupeaformis), which underwent dramatic
declines during the 1940s and 1950s (Smith and Tibbles 1980). In fact,
sea lamprey predation played a major role in causing the extirpation of
lake trout in lakes Ontario, Erie, Huron, and Michigan (Cornelius et al.
1995, Elrod et al. 1995, Eshenroder et al. 1995, Holey et al. 1995).
Large-bodied fishes such as lake trout and lake whitefish are of vital
ecological, economic, and cultural importance in the Great Lakes. As such,
the Great Lakes Fishery Commission (GLFC), in partnership with both U.S.
and Canadian resource management agencies, has developed an integrated
sea lamprey management program designed to suppress sea lamprey populations
in the Great Lakes, and ultimately allow for the rehabilitation and continued
protection of these and other important salmonines (GLFC 2001). Critical
to the success of this effort is development of a control program that
minimizes the costs expended for sea lamprey control relative to the economic,
ecological, and social benefits associated with protecting fish from sea
lamprey predation (GLFC 2001). To make progress toward development of
a cost-effective control program, a major effort has been undertaken to
understand sea lamprey life history (physiology, ecology, and behavior),
as well as describe the geographical distribution of sea lamprey production
in the Great Lakes.
Historically, sea lamprey control efforts have focused on blocking upstream
migration of adults to spawning sites using barriers, and application
of lampricides (e.g., TFM, Bayluscide) to spawning tributaries. These
chemicals kill sea lamprey as larvae, before they metamorphose into parasites
(Smith and Tibbles 1980). Overall, the use of barriers and lampricides
helped reduce sea lamprey populations by as much as 90% in the Great lakes
between the 1940s and 1970 (Young et al. 1996). In turn, these efforts,
combined with stocking programs and restrictions on harvest, allowed for
re-establishment of lake trout populations in all of the Great Lakes (Cornelius
et al. 1995, Elrod et al. 1995, Eshenroder et al. 1995, Hansen et al.
1995, Holey et al. 1995).
Unfortunately, however, in recent decades sea lamprey populations in the
Great Lakes have begun to increase, especially in Lake Huron, to levels
on par with those before control efforts were established (Eshenroder
et al. 1995, Young et al. 1996). Much of this increase in parasitic-phase
sea lamprey is apparently due to production from the St. Marys River,
the connecting corridor between Lake Superior and Lake Huron, where ~66%
of all suitable spawning habitat in the Great Lakes exists (Eshenroder
et al. 1987, Young et al. 1996). Although habitat rehabilitation in the
St. Marys River may have played a role in this increase in sea lamprey
in the northern basin (Ferreri et al. 1995), the documented increase since
the late 1970s is more likely due to enhanced abundance of potential prey
(e.g., lake trout, bloater chub Coregonus hoyi) for recently transformed
sea lamprey (Young et al. 1996). Owing to the continued presence of sea
lamprey in the Great Lakes, mortalities due to sea lamprey predation remain
high in most lakes (Hansen et al. 1995, Holey et al. 1995, Schneider et
al. 1996), especially in northern Lake Huron where sea lamprey are responsible
for a larger proportion of lake trout mortality than all other sources
of mortality combined (Sitar et al. 1999).
Given the persistence of sea lamprey populations in the Great Lakes, continuation
of control efforts is essential, which is problematic because of high
associated costs (e.g., lampricide applications cost ~$15 million annually).
These economic concerns, as well as concerns about the potential for development
of resistance to lampricides, potential effects of lampricides on non-target
species, and the negative stigma associated with applying large quantities
of a chemical pesticide to tributaries, have prompted the GLFC to seek
ways to reduce the use of lampricides (GLFC 2001). Toward this end, efforts
have been undertaken to optimize control strategies so that lampricide
treatments (and other more experimental control efforts, such as sterile
male release) are used where and when they would be most effective. A
significant step forward has been development of the Integrated Management
of Sea Lamprey process, which ultimately generates a ranked list of tributaries
for lampricide treatment based on cost-effectiveness (i.e., dollars expended
to kill each larva). Although numerous tributary-specific variables are
considered in these rankings (e.g., amount of lampricide and labor needed
for treatment, amount of spawning habitat, estimated economic-injury levels),
a major factor determining which spawning locations receive treatment
is the estimate of larval sea lamprey abundance in each location. Thus,
a tributary with a large estimated larval sea lamprey population would
be ranked as a higher priority for control than a tributary with a small
population.
There are two major drawbacks to relying heavily on larval abundance for
these prioritizations. First, Quantitative Assessment Survey (QAS) sampling,
which is conducted by both the U.S. Fish and Wildlife Service (USFWS)
and the Canadian Department of Fisheries and Oceans (DFO) to estimate
sea lamprey population sizes, is operationally difficult in large rivers
(e.g., St. Marys, Spanish, and Mississagi rivers). Consequently, confidence
in estimates of larval sea lamprey population sizes in large rivers is
not great, and hence, an understanding of the potential contributions
of parasites from these rivers remains largely unknown (M. Steeves, DFO,
pers. comm.). Second, prioritizing tributaries according to larval abundance
estimates makes the implicit assumption that the number of transformers
(and ultimately future parasitic- and spawning-stage lamprey) that emerge
from a tributary is correlated with larval abundance such that tributaries
with a large larval population will contribute more individuals to parasitic
populations than those with a small number of larvae. In other words,
this approach assumes that survival rates of larvae to the parasitic and
adult life stages are equal across all tributaries. Because tributary-specific
survival rates are largely unknown, and quite possibly vary among tributaries,
owing to habitat differences, this approach seems limited. Realistically,
two tributaries with very different larval sea lamprey populations could
contribute equal numbers of parasites due to factors such as competition
(high larval production x high mortality = low larval production x low
mortality). Thus, there is a strong need to develop alternative methods
to prioritize tributaries for control efforts that are not constrained
by assumptions concerning larval survival. Such methods ideally would
have the potential to provide a definitive understanding of which spawning
locations contribute both parasites and spawners, as well as be cost-effective.
Clearly, until a true understanding of the relative contribution of each
tributary to the parasitic population is obtained, the GLFC cannot be
assured that its control measures are being implemented in the most cost-effective
manner possible.
Herein, we propose to continue development of a rather novel technological
approach, statolith microchemistry, as a tool to determine which local
spawning tributaries contribute to parasitic- and spawning-stage sea lamprey
populations in Lake Huron. First, we seek to provide a definitive answer
as to whether statolith elemental concentrations can be used as a tool
to identify sources of sea lamprey in Lake Huron, which no study has currently
done. Given the geographical scope of our study and technologies available
in Fryer’s laboratory, we are confident in our ability to do this.
Second, assuming that statolith microchemistry holds up to its potential,
we will seek to quantify the relative contribution of parasites and spawners
from spawning tributaries important to Great Lakes management agencies,
including the St. Marys River. This river is of specific interest because
historically it has been the primary source of sea lamprey production
in the Great Lakes (Young et al. 1996). In addition, a major control effort,
consisting of the application of granular Bayluscide, trapping of adult
sea lamprey, and release of sterile males (GLFC 2001), has recently been
undertaken in the St. Marys River. Although preliminary data suggest that
the larval population has been reduced by as much as 45% in this river
after initial (1999) application of Bayluscide, a true assessment of whether
the St. Marys River control effort has changed the relative contribution
of adult spawners and parasites in Lake Huron remains unknown. We fully
expect that estimation of relative contributions from other Lake Huron
tributaries, including difficult-to-sample large rivers (e.g., Spanish
and Mississagi; M. Steeves, DFO, pers. comm.), will be possible as well.
Finally, taking advantage of the stability of water chemistry in Lake
Huron tributaries (see below), we will attempt to quantify relationships
between water and statolith chemistry in hopes of eliminating any future
need for annual larval sampling from spawning tributaries. Ultimately,
this would reduce the expense associated with using this technology for
estimation of tributary contributions of sea lamprey to the open lake.

Inductively Coupled Plasma Mass Spectrometer (ICPMS)
2005 Accomplishments
Below, we describe our accomplishments for each of the three primary
objectives.
PROJECT OBJECTIVES: 1. Determine whether statolith elemental
signatures differ among larvae produced in different L. Huron streams.
Larval sea lamprey Collections
To develop characteristic statolith elemental signatures for Lake Huron
streams, larvae were collected from numerous Lake Huron tributaries (Figure
1) by the US Fish and Wildlife Service (Marquette and Ludington stations)
and the Department of Fisheries and Oceans (Sault Ste. Marie). Similar
collections in tributaries surrounding both Lake Michigan and Huron have
already begun for 2005. Importantly, these collections will allow us to
explore inter-annual variability in stream signatures, as well as fill
in gaps in sampling because not all Lake Huron streams were sampled during
2004.
We currently have analyzed larval sea lamprey from 30 streams surrounding
Lake Huron and Lake Michigan (Ford River, St. Mary's River, Silver Creek,
Mississagi River, Hessel Creek, Tawas River and Trout Creek) (n = 15 per
stream). We fully expect that all samples, from both 2004 and 2005, will
be processed and analyzed by January 2006.
Preliminary Results
Preliminary results from the seven streams (see Figure
1) processed thus far are encouraging. In total, 8 of 13 elements
analyzed were above detection limits (i.e., concentrations in statoliths
were sufficiently high to be reliably detected by our ICPMS). More importantly,
we found several elements (rubidium, manganese, strontium, and zinc) were
valuable for discriminating among streams (Table 1;
Figure 2). Using these 8 elements, we could reliably discriminate
most streams; 6 of 7 streams could be discriminated (using Linear Discriminant
Function Analysis, LDFA) with ? 85% accuracy, whereas the seventh stream
(Mississagi River) could only be discriminated from other with 50% accuracy
(Table 2). As we increase the number of streams
analyzed at any given time, it is likely that other elements will become
valuable for discrimination purposes. Further, as we increase sample sizes
for each stream-ideal sample sizes will be determined objectively, using
a rigorous bootstrapping analysis to look at how sample sizes within streams
influence characteristic elemental signatures-we are hopeful that our
discrimination abilities will increase (e.g., we will be able to discriminate
the Mississagi River from others as well).
Figure 1: Locations of sample sites, with processed
and analyzed sites highlighted. Unlabeled locations not yet processed.
2. Quantify relative contributions of parasitic and spawning
lamprey from important production tributaries.
We will address this objective once the preliminary work on the larvae
has been completed, since without results from our first objective, we
will have no basis for determining the origins of parasites and spawners.
However, to this point, we have secured both parasites and spawners from
Lake Huron from both commercial fishing operations and agent trapping
efforts (the USFWS, DFO, and USGS were instrumental in securing these
collections). We also will receive parasites and spawners from Lake Huron,
as well as Lake Michigan, during 2005. The Lake Michigan adults and parasites
will prove invaluable to (hopeful) future efforts to conduct a similar
study in Lake Michigan (per some discussions with Chuck Krueger).
One assumption we are making is that storage method has no effect on
elemental composition of statoliths, given that larvae are stored in ethanol
and adults/parasites are stored frozen. To explore the effects of storage
method on statolith elemental composition, we analyzed larvae (n = 14
stored frozen; n = 15stored in 95% ethanol) collected from Browns Bayou
in the Big Manistee River. Briefly, we found no significant differences
between frozen and ethanol preserved fish for magnesium, manganese, zinc,
barium, lead or strontium (two-sample t-tests; all p > 0.05). We did,
however, a significant, difference between storage methods for rubidium.
Importantly, however, the difference, although statistically significant,
was small (ethanol mean + 1 SD = 2.10+0.49 ppm; ethanol mean + 1 SD =
2.76+0.62 ppm) relative to the variation in rubidium that has been documented
across Lake Huron and Lake Michigan streams (range in Rb across the seven
streams processed thus far is 2.47 to 7.09 ppm). Thus, we are optimistic
that any biases associated with storage method will be negligible. We
will further assess the robustness of these results by performing a similar
suite of analyses on larvae collected in Lake Champlain this summer.
3. Develop relationships to predict statolith elemental chemistry
from water chemistry.
This objective will be met first with some preliminary work on Lake Champlain,
followed by comparative work on Lake Huron. Water and sediment samples
from Lake Champlain will be collected by Carrol Hand during the week of
June 13-18, brought back and analyzed at the lab in Windsor. Specifically,
Carrol will sample all 22 sea lamprey producing streams in Lake Champlain
for both water chemistry and sediments. Additional collections in these
same streams will be made again during fall 2005 and spring 2006 (after
snowmelt) to assess inter-annual variability in signatures. Collections
of water chemistry and sediment samples around Lake Huron also will be
made during summer 2005, which will supplement collections being made
by other agencies (USGS). Ultimately, we will compare our water and sediment
chemistry results to sea lamprey statolith micro-elemental composition
to develop partition coefficients.

Figure 2. Canonical analysis of sea lamprey larvae (only
Roots 1 and 2 presented, which explained a combined 78% of the variation
in the data). See Table 1 for an explanation of elemental concentrations,
and Figure 1 for an explanation of stream acronyms.
Table 1. Correlation of elemental concentrations with
Linear Discriminant Function Analysis axes (roots). Elements most highly
correlated with each axis are highlighted in gray, and indicate their
importance in discrimination. For example, Manganese (Mn) and Rubidium
(Rb) were most important for explaining variation along Root 1 (horizontal
axis; positive coefficients indicate that Mn and Rb increase from left
to right along Root 1; see Figure 2). Root 2 was positively correlated
with Strontium (Sr), indicating that individuals at the top of Figure
2 were higher in Sr than those at the bottom (see Figure 2). The percentage
of variation that each axis (root) explained is provided in the bottom
row.
| Element |
Root 1 |
Root 2 |
Root 3 |
Root 4 |
| Magnesium (Mg) |
0.20 |
-0.24 |
-0.33 |
-0.55 |
| Manganese (Mn) |
0.56 |
-0.22 |
-0.34 |
0.16 |
| Zinc (Zn) |
-0.07 |
0.35 |
-0.54 |
-0.56 |
| Rubidium (Rb) |
0.62 |
0.19 |
0.44 |
-0.58 |
| Strontium (Sr) |
0.11 |
0.73 |
-0.22 |
0.27 |
| Tin (Sn) |
-0.12 |
-0.06 |
0.03 |
-0.08 |
| Barium (Ba) |
0.15 |
0.12 |
-0.27 |
0.12 |
| Lead (Pb) |
-0.03 |
0.00 |
-0.04 |
-0.61 |
| Cumulative % variance explained |
0.63 |
0.78 |
0.91 |
0.96 |
Table 2: Classification matrix for larvae processed
at seven Lake Huron/Michigan sampling locations. Our ability to discriminate
among individuals (larvae) from different streams was quite good. With
the exception of Mississagi River (MISR) individuals, we could discriminate
among streams with > 85% accuracy. In the case of the Mississagi River,
individuals were misclassified as Hessel Creek (HESC) individuals in half
the cases (5 of 10). Individuals classified correctly are highlighted
in gray. See Figure 1 for stream acronym definitions.
|
Percent Correct |
FORR |
HESC |
MISR |
SILC |
STMR |
TAWR |
TROC |
Total |
| FORR |
88.9 |
8 |
0 |
0 |
0 |
1 |
0 |
0 |
9 |
| HESC |
86.7 |
0 |
13 |
1 |
0 |
0 |
0 |
1 |
15 |
| MISR |
50.0 |
0 |
5 |
5 |
0 |
0 |
0 |
0 |
10 |
| SILC |
90.0 |
1 |
0 |
0 |
9 |
0 |
0 |
0 |
10 |
| STMR |
90.0 |
1 |
0 |
0 |
0 |
9 |
0 |
0 |
10 |
| TAWR |
90.0 |
1 |
0 |
0 |
0 |
0 |
9 |
0 |
10 |
| TROC |
93.3 |
0 |
0 |
0 |
1 |
0 |
0 |
14 |
15 |
| Total |
84.8 |
11 |
18 |
6 |
10 |
10 |
9 |
15 |
79 |
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last updated: 2006-04-12 mbl
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