Endangered and Threatened Wildlife and Plants; 12-Month Finding on a Petition To List the American Eel as Threatened or Endangered, 4967-4997 [07-429]
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Federal Register / Vol. 72, No. 22 / Friday, February 2, 2007 / Proposed Rules
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Dated: January 26, 2007.
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Quality.
[FR Doc. E7–1726 Filed 2–1–07; 8:45 am]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife
and Plants; 12-Month Finding on a
Petition To List the American Eel as
Threatened or Endangered
Fish and Wildlife Service,
Interior.
ACTION: Notice of 12-month petition
finding.
AGENCY:
SUMMARY: We, the U.S. Fish and
Wildlife Service (USFWS), announce
our 12-month finding on a petition to
list, under the Endangered Species Act
of 1973, (Act) as amended, the
American eel (Anguilla rostrata) as a
threatened or endangered species
throughout its range. After a thorough
review of all available scientific and
commercial information, we find that
listing the American eel as either
threatened or endangered is not
warranted at this time. We ask the
public to continue to submit to us any
new information that becomes available
concerning the status of or threats to the
species. This information will help us to
monitor and encourage the ongoing
conservation of this species.
DATES: The finding in this document
was made on February 2, 2007.
ADDRESSES: Data, information,
comments, or questions regarding this
finding should be sent by postal mail to
Martin Miller, Chief, Division of
Endangered Species, Region 5, U.S. Fish
and Wildlife Service, 300 Westgate
Center Drive, Hadley, Massachusetts
01035–9589; by facsimile to 413–253–
8428; or by electronic mail to
AmericanEel@fws.gov.
FOR FURTHER INFORMATION CONTACT:
Heather Bell, at the street address listed
in ADDRESSES (telephone 413–253–8645;
facsimile 413–253–8428). Persons who
use a telecommunications device for the
deaf (TDD) may call the Federal
Information Relay Service (FIRS) at
800–877–8339, 24 hours a day, 7 days
a week.
SUPPLEMENTARY INFORMATION: The
complete administrative file for this
finding is available for inspection, by
appointment and during normal
business hours, at the street address
listed in ADDRESSES. The petition
finding, the status review for American
eel, related Federal Register notices,
and other pertinent information, may be
obtained online at https://www.fws.gov/
northeast/ameel/.
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Background
Section 4(b)(3)(B) of the Act, as
amended (16 U.S.C. 1531 et seq.),
requires that, for any petition to revise
the Lists of Endangered and Threatened
Wildlife and Plants that contains
substantial scientific and commercial
information that listing may be
warranted, we conduct a status review
and make a finding within 12 months of
the date of receipt of the petition
(hereafter referred to as a 12-month
finding) on whether the petitioned
action is (a) not warranted, (b)
warranted, or (c) warranted but the
immediate proposal of a regulation
implementing the petitioned action is
precluded by other pending proposals to
determine whether any species is
threatened or endangered, and
expeditious progress is being made to
add or remove qualified species from
the Lists of Endangered and Threatened
Wildlife and Plants.
On May 27, 2004, the Atlantic States
Marine Fisheries Commission (ASMFC),
concerned about extreme declines in the
Saint Lawrence River/Lake Ontario
(SLR/LO) portion of the species’ range,
requested that the USFWS and the
National Oceanic and Atmospheric
Administration’s National Marine
Fisheries Service (NMFS) conduct a
status review of the American eel. The
ASMFC also requested an evaluation of
the appropriateness of a Distinct
Population Segment (DPS) listing under
the Act for the SLR/LO and Lake
Champlain/Richelieu River portion of
the American eel population, as well as
an evaluation of the entire Atlantic coast
American eel population (see Finding
for definition of DPS) (ASMFC 2004a, p.
1). The USFWS responded to this
request on September 24, 2004; our
response stated that we had conducted
a preliminary review regarding the
potential DPS as described by the
ASMFC, and determined that the
American eel was not likely to meet the
discreteness element of the policy
requirements due to lack of population
subdivision (further analysis is provided
under Finding). Rather, the USFWS
agreed to conduct a rangewide status
review of the American eel in
coordination with NMFS and ASMFC
(USFWS 2004, p. 1).
On November 18, 2004, the USFWS
and the NMFS received a petition, dated
November 12, 2004, from Timothy A.
Watts and Douglas H. Watts, requesting
that the USFWS and NMFS list the
American eel as an endangered species
under the Act. The petitioners cited
destruction and modification of habitat,
overutilization, inadequacy of existing
regulatory mechanisms, and other
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natural and man-made factors (such as
contaminants and hydroelectric
turbines) as the threats to the species.
On July 6, 2005, in response to the
petition, the USFWS issued a 90-day
finding on the petition (70 FR 38849),
which found that the petition presented
substantial information indicating that
listing the American eel may be
warranted. The finding noted concern
that the dramatic decrease in
recruitment of American eel noted at the
Moses-Saunders Dam in Canada (on the
St. Lawrence River), coupled with the
significant decline seen in the European
eel (ASMFC 2000, pp. 12–14), could
indicate a decline in the American eel.
Information on possible reasons for this
suggested decline included the
following threats: Commercial harvest,
habitat loss and degradation (primarily
the loss of wetlands and upper tributary
habitat), hydropower turbine mortality,
and inadequacy of existing regulatory
mechanisms. Other potential threats,
such as seaweed harvest, benthic (sea or
lake bottom) habitat destruction,
alterations of stream flow, disease,
predation, and contaminants, were not
fully addressed or supported by the
information presented in the petition.
Further analysis of oceanic variations
(such as changes in the Gulf Stream)
were recommended in the 90-day
finding, particularly in light of the scant
direct evidence and the potential for
oceanic variations to be compounding
or confounding the impact of other
threats. Additionally, the 90-day finding
concluded that the complex life history
and the incompleteness of historical
data (abundance, stock composition, life
stage mortality rates, and exploitation
rates) made it challenging to understand
the potential influence of multiple
threats to the American eel (USFWS
2005a, p. 38860).
In response to our 90-day finding’s
request for information for use in the
species’ status review, we received
comments and information on American
eel from the majority of the State fish
and wildlife agencies within the range
of the eel; State universities; State and
university museums; the U.S. Forest
Service (USFS); National Park Service
(NPS); U.S. Geological Survey (USGS);
Army Corp of Engineers (ACOE); the
Department of Defense; the ASMFC; the
Great Lakes Fisheries Commission;
Department of Fisheries and Oceans
(Canada); Tribal Nations; academics and
researchers from the United States,
Canada, Japan, and several European
countries; hydropower and fishing
industries; nongovernmental
organizations; private citizens; and
other entities. Additionally, we
coordinated with the USFWS’s
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International Affairs Program (IAP) to
obtain information on international
trade and with State and Federal law
enforcement officials on illegal trade.
Although all countries where the
American eel is native were contacted
regarding information, there was no
available data on eel distribution,
habitat use, habitat degradation or loss,
or other threats (other than international
harvest data) from Central or South
America. Distribution information was
provided by some Caribbean Islands.
Therefore, the status review focused on
where data is available within the North
American Continent.
A status review allows for additional
collection, clarification, and
interpretation of information on the
status of the species by the USFWS. The
resulting status review, from which the
12-month finding is based, relied on our
extensive review of the existing
literature, data resulting from the 90-day
finding request for information, and
new information obtained during the
status review period. Among the new
information we received, the documents
most relevant to the status review
include the recently completed stock
assessments for the Atlantic coast
(ASMFC 2006a and b), the American eel
data assembled for the Canadian stock
assessment (Cairns et al. 2005), and
recently completed research on life
history and potential threats to the
American eel (van den Thillart et al.
2005; Oliviera in USFWS 2006; Machut
2006; Lamson et al. 2006; Devarut et al.
2006; Knights et al. 2006).
Also, because of the large body of
literature and the uncertainty
surrounding several threats, we hosted
two scientific workshops with over 25
scientific experts. The goal of the
workshops was to insure that the
USFWS properly utilized the best and
most current scientific and commercial
data available in conducting the status
review. To reach this goal, each of the
experts was asked a series of facilitated
questions to assess the presented
information (which included multiple
factual inputs, data, models,
assumptions, etc.), including the
completeness of the literature selected,
and to comment on the relevance and
quality of the literature for purposes of
our status review (see workshop
summaries Web site at https://
www.fws.gov/northeast/ameel/). The
USFWS recorded each expert’s
individual assessments and the basis for
those assessments in a compendium
(cited in the finding as USFWS 2005b
and 2006). Workshop objectives
included determining the following:
Utility of the information; life history
stages vulnerable to certain threats; the
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geographic scope of the threats; the
immediacy of the threats; and
uncertainties in the available
information and the potential
implications of those uncertainties in
making a status determination.
The selection of the expert panelists
was based on recommendations from
within and outside of the USFWS and
NMFS (the Services). The panelists
selected represented a broad and diverse
range of scientific perspectives relevant
to the status review of the American eel
coming from State and Federal agencies,
fishery commissions, Tribes, academia,
domestic and foreign research
institutions (Canada, Japan, and
England), industry organizations, and
nongovernmental organizations.
Participating individuals had expertise
on threats or life history characteristics
associated with threats to the American
eel.
Therefore, in addition to the
published literature, our review
considered: (1) Each expert panelist’s
characterization of the threat (the life
stages acted upon by the threat, the
severity of the threat, and the timing of
the threat) based on their own and other
published and unpublished research on
the species; (2) the basis for each expert
panelist’s assessments of the literature
in the context of a rangewide status
review; and, (3) each expert panelist’s
assessments of the implications of the
uncertainty in the information. This
finding therefore builds on, clarifies,
reinterprets, and, in some cases,
supersedes information presented in the
90-day finding.
In conducting our 12-month finding
for American eel, we considered all
scientific and commercial information
on the status of American eel that we
had in our files. Parallels in life history
traits that are unknown for the
American eel are drawn from other
species of Anguilla.
Evolution and Population Structure
The American eel is one of 15 ancient
species (evolving circa 52 million years
ago) of the worldwide genus Anguilla,
whose members spawn in ocean waters,
migrate to coastal and inland
continental waters to grow, and then
return to ocean spawning areas to
reproduce and die—a life history
strategy known as catadromy (McCleave
2001a, p. 800; Avise 2003, p. 31;
Knights et al. 2006, pp. 2–3).
The North Atlantic is home to two,
closely related, recognized species of
Anguilla—the American eel and the
European eel (A. anguilla) (Avise 2003,
p. 31). Genetic research indicates that
the American eel lacks appreciable
phylogeographic population structure,
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meaning that American eels are one,
well-mixed, single breeding population,
termed panmixia or panmictic (Avise
2003, pp. 34–35). This likely occurs
from a combination of the random
distribution of the eel’s larval stage
when they reach continental waters and
random mating among all adults
throughout the species’ range. This is in
contrast to many anadromous species
(which, even though they have an
oceanic phase, return to their rivers of
origin to spawn), where mating is
within separate populations that are
geographically or temporally isolated.
This panmictic life history strategy
maximizes adaptability to changing
environments and is well suited to
species that have unpredictable larval
dispersal to many habitats (Stearns 1977
in Helfman et al. 1987, p. 52).
Additionally, by not exhibiting
geographic or habitat-specific
adaptations, eels have the ability to
rapidly colonize new habitats and to recolonize disturbed ones over wide
geographical ranges (McDowall 1996 in
Knights et al. 2006, p. 7).
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Life History
In brief, the life history of the
American eel begins in the Sargasso Sea,
where eggs hatch into a larval stage
known as ‘‘leptocephali.’’ These
leptocephali are transported by ocean
currents to the Atlantic coasts of North
America and upper portions of South
America. They enter coastal waters,
where they may stay, or they may move
into estuarine waters or migrate up
freshwater rivers, where they grow as
juveniles and mature. Upon nearing
sexual maturity, these eels begin
migration toward the Sargasso Sea,
completing sexual maturation en route.
Spawning occurs in the Sargasso Sea.
After spawning, the adults die; a species
with this life history trait is known as
a semelparous species. For a detailed
description of the life cycle and other
life history characteristics, see McCleave
2001a, Tesch 2003, and Cairns et al.
2005. Aspects of the species’ life history
most relevant to this finding are
discussed in more detail below.
Egg and Larval Life History Stage
The egg and larval stage of the
American eel occur in the Atlantic
Ocean, the Sargasso Sea, ocean currents,
and Continental Shelf waters.
Sargasso Sea. The Sargasso Sea is part
of the North Atlantic Ocean, lying
roughly between the West Indies and
the Azores. The Sargasso Sea is part of
the western half of a large clockwise
gyre (circular pattern of ocean
circulation). It is here that American eel
eggs hatch into a larval stage known as
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‘‘leptocephali.’’ The leptocephali are
distributed in the upper 300 meters (m)
of the ocean and are subject to transport
from surface currents in the Sargasso
Sea. These surface currents can be
complex due to the fronts that form in
the Subtropical Convergence Zone
(where equatorial and temperate waters
meet) primarily in the winter and
spring, and the eddies that are likely
present year round.
Ocean current transport. The Sargasso
Sea includes a powerful western
boundary current, the Florida Current
and Gulf Stream, which flows to the
north and northeast along the Atlantic
coast of North America. The Florida
Current is the southern half of this flow,
from the Straits of Florida to Cape
Hatteras (Schott et al. 1988 in Miller
2005, p. 3). The Florida Current
transports water from the Caribbean,
Gulf of Mexico, and more distant
regions through the Straits of Florida. It
then combines with Gulf Stream
recirculation water from the Sargasso
Sea as it flows north of the Bahamas
(Marchese 1999, pp. 29, 549), and forms
the Gulf Stream off Cape Hatteras, North
Carolina. Once past Cape Hatteras, the
Gulf Stream (which is at least 48 km or
30 miles offshore but more typically 160
km or 100 miles or greater offshore)
usually has pronounced meanders,
which, if large enough, can get
separated and cast off to the north into
the continental slope water (a water
mass found in the permanent
thermocline between the Gulf Stream
and the continental shelf north of Cape
Hatteras (35 °N)). The flow of the Gulf
Stream continues to the northeast,
mostly paralleling the Atlantic coast,
towards Europe and becomes the North
Atlantic Current (Miller 2005, pp. 3–4).
The majority of the leptocephali enter
the Florida Current just south of Cape
Hatteras (just south of where the Florida
Current enters the Gulf Stream) directly
from the Sargasso Sea. The remainder
may enter the Florida Current by a more
southern route (e.g., transported on the
Caribbean Current through the Yucatan
Straights (Kleckner and McCleave 1985,
p. 89), to the Gulf Loop Current and
then to the Florida Current, which
would be the route most likely taken for
Gulf of Mexico recruitment) (Kleckner
and McCleave 1982, p. 329–330; Miller
2005, p. 3).
The distribution of American eel
leptocephali in the Florida Current was
first described by Kleckner and
McCleave (1982, pp. 334–337; 1985, pp.
73–77). Additionally, they found
evidence of westward movement of
leptocephali across the current toward
the coastal waters. Because the
distances of transport, to southern and
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northern points along the Atlantic coast,
differ by thousands of kilometers, it has
been suggested that the timing of
metamorphosis from leptocephali to the
next life history stage may determine
where individuals arrive in Continental
Shelf waters.
Other than likely current transport,
we know very little about the American
eel leptocephali. Recent studies on other
species have indicated that leptocephali
may feed on marine snow or specific
detrital particles, such as discarded
larvacean (planktonic tunicates that
secrete a gelatinous house) houses and
zooplankton fecal pellets (Otake et al.
1993, pp. 28–32; Mochioka and
Iwamizu 1996, p. 447).
Continental shelf waters. The
American eel undergoes metamorphosis
twice. The first occurs when the
leptocephali enter the Continental Shelf
waters (the area of shallow seas just off
the coast to the area of marked increase
in slope to greater depths); the second
is during sexual maturation. The
leptocephalis’ leaf-like, laterally
compressed shape transforms during
metamorphosis into a reduced,
characteristically eel-like shape, as they
become transparent ‘‘glass’’ eels.
Leptocephali are unusual fish larvae
that are filled with a transparent
gelatinous energy storage material, and
they can swim either forwards or
backwards (Miller and Tsukamoto 2004
in Miller 2005, pp. 1–2); this may be an
important aspect in detraining from
(getting off of) the Gulf Stream.
According to Miller (2005), this
directional swimming appears to be the
only way that leptocephali can cross
and detrain from the Gulf Stream system
and cross the Continental Shelf waters,
due to the lack of any persistent oceanic
transport mechanism that can account
for the large-scale transport of millions
of larvae across the current.
Juvenile Life History Stage
Arrival in coastal waters. When
juvenile eels arrive in coastal waters,
they can arrive in great density and with
considerable yearly variation (ICES
2001, p. 2). Arriving juvenile eels
(unpigmented ‘‘glass eels’’ and
pigmented ‘‘elvers’’) have been collected
and recorded for 10 years from two sites
in North Carolina in the Beaufort
estuary. Densities as high as 13.5–14.0
eels/100m3 and as low as 1.5 eels/100m3
have been recorded (Powles and Warlen
2002, p. 301). In the East River, Canada,
Jessop (2000, p. 520) had daily counts
of 30,000 elvers entering the mouth of
the river. Between May and August
200,000 elvers were recorded by trap
method, and a population estimate of
960,000 elvers was conducted by mark-
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recapture (Jessop 2000, pp. 518–520).
Variation in recruitment between years
can be quite significant. In the 9 years
of records between the years 1982 to
1999, estimated recruitment to the Petite
`
´
riviere del la Trinite varied roughly
four-fold, from a low of 14,014 to a high
of 61,308 (ICES 2001, p. 36). Some
arrivals remain in brackish (estuarine)
or marine (salt) waters, others migrate
up rivers to a variety of fresh water
habitats, and still others, as they mature,
will show inter-habitat movement
patterns (Jessop et al. 2002, pp. 217–
218; Morrison et al. 2003, pp. 90–92;
Cairns 2006a, p. 2; Thibault et al. 2005,
p. 36; Lamson et al. 2006, p. 1567;
Daverat et al. 2006, p. 2).
Juvenile mortality. Information on
mortality rates for all of the life stages
is limited. In Jessop (2000, p. 514), the
recruitment of elvers to the East River,
Chester, Nova Scotia, during May
through July was estimated by markrecapture population estimates to be
960,000 elvers. The population size
following migration to recapture sites
about 1.3 kilometers (km) upstream
during late July–October was 2,894
elvers. These data indicate high juvenile
mortality rates, in this case at a rate of
99 percent. This high mortality was
attributed to the effects of low pH (4.7–
5.0), high initial elver density (4.7
elvers/m2) (which may lead to
predation, including cannibalism,
starvation, and competition for space),
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predation and starvation as size
increases.
Juvenile diet. The enormous dietary
breadth of eels reflects their great
adaptability with respect to nearly all
conditions of water bodies. Yellow eels
are opportunistic, consuming nearly any
live prey that can be captured. Smaller
eels eat benthic invertebrates; larger eels
include mussels, fish, and even other
eels in their diet. Yellow eels also adapt
to seasonal changes, decreasing intake
or ceasing to eat during the winter. Eels
can also respond to local abundances of
appropriately sized prey through the
seasons (Tesch 2003, pp. 152–163). This
adaptable diet allows for resource
partitioning as well as the ability to
withstand changes in local
environmental conditions and the
ability to occupy a geographically wide
variety of habitats.
Density-dependent dispersion. As
young eels begin to grow, densitydependent competition promotes eels to
disperse into less crowded areas
(Feunteun et al. 2003, pp. 201–204;
Ibbotson et al. 2002 in Knights et al.
2006, p. 10). Aggressive interactions at
high density inhibit feeding and growth,
but stimulate dispersive swimming
activity in smaller eels (Knights 1987 in
Knights et al. 2006, p. 10), the latter
likely as a defense against predation. As
size differences in these juveniles
increase, cannibalism can also be an
important cause of mortality (Knights
1987 in Knights et al. 2006, p. 10).
Density dependent dispersion ensures
wider distributions, further minimizing
intra-specific competition. Benefits of
density dependent dispersion include
selection of optimal habitat productivity
and temperature, lower predation risks,
rapid colonization or re-colonization of
habitats, and avoidance of inter-specific
competition. Larger individuals farther
upstream tend to become more
sedentary and occupy territories,
densities of eels decline, and females
predominate (Feunteun et al. 2003, p.
201).
Distribution clines. It has been
suggested that there are latitudinal
clines in eel distribution related to river
typologies. For example, the American
eel tends to extend farther inland in
southerly lowland drainages compared
to distributions in the shorter and
steeper post-glacial stream systems in
the Northeast (Jessop et al. 2004 in
Knights et al. 2006, p. 11). Smogor et al.
(1995, p. 799) and Knights (2001 in
Knights et al. 2006, p. 8) have
documented decreases in densities with
increasing distance from the Continental
Shelf in a predictable pattern, likely as
a result of density dependant dispersion
and mortality due to predation.
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Although mean watershed densities
decrease by an order of magnitude with
distance inland from the Continental
Shelf, mean biomass only declines by
about 50 percent because mean body
weight and eel length increase (and
hence relative fecundity). This,
according to Knights et al. (2006, p. 10),
helps maintain biomass relative to
carrying capacity. Machut (2006, p. 13)
indicates that as barrier intensity
increases, so does eel growth above the
barrier. Recent research (Knights et al.
2006, pp. 11–13) has documented that
as eel density decreases, the proportion
of females increases, which, assuming
females are the limiting sex, would be,
according to Knights et al. (2006, p. 13),
a compensatory mechanism during
times or in areas of low density.
Sexually Maturing Life History Stage
Sex determination. There are no
morphologically differentiated sex
chromosomes in the American eel
(McCleave 2001a, p. 803). Prior to
sexual differentiation, eels are
intersexual, meaning they can develop
into either sex. It is only when yellow
eels reach a length of about 20–35 cm
that it is possible to distinguish males
from females visually, and there is
considerable variation in age and size at
differentiation. The determination of sex
is likely influenced by environmental
factors, including eel densities (Tesch
2003, pp. 43–46). Studies indicate that
as the density of eels in a particular area
increases the number of male eels
increases; decreasing density favors
more females. It has been argued by
Knights et al. (2006, p. 13), that an
advantage of this life history strategy is
that when recruitment declines, so will
density and tendencies to migrate far
upstream in rivers. In turn, this will
lead to relative increases in the number
of (larger) females and hence
compensatory increases in fecundity.
This may take a number of generations
(and hence decades) to manifest itself,
but this strategy confers enormous
benefits in the face of threats, past,
present and future, such as tectonic
events and changes in ocean currents
and climate (Knights et al. 2006, p. 13).
Silvering. After a number of years, the
yellow eels begin metamorphosis.
Beginning at 3 years old and up to 24
years, with the mean becoming greater
with increasing latitude (e.g., 6–16 years
in the Chesapeake Bay region; Helfman
et al. 1987, pp. 44–45; and 8–23 years
in Canada; Cairns et al. 2005, p. 11),
yellow eels metamorphose into ‘‘silver
eels’’ (Cairns et al. 2005, p. 13). This
metamorphosis from bottom-oriented
yellow eels to silver eels (termed
‘‘silvering’’) is a key physiological event
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preparing these future spawners for
oceanic migration and reproduction
(van den Thillart et al. 2005, p. 12).
Environmental factors may play a role
in the triggering of silvering. Habitat
conditions, such as food availability and
temperature, will influence the size and
age of silvering eels via growth
conditions. Thus, variation in length
and age at maturity can occur in
different habitats (e.g., freshwater
habitat versus estuarine habitat) within
a restricted geographic range and over
larger geographic scales as well.
The length of the growing season and
the temperature are negatively
correlated with latitude, so age at
maturity is strongly correlated with
latitude (McCleave 2001a, p. 803).
Characteristics of silver eels vary across
the species’ range. Eels from northern
areas, where migration distances are
great, show slower growth and greater
length, weight, and age at migration,
preparing them, it could be assumed, for
the longer migration.
Indeed, favorable growth conditions
cause eels to silver more rapidly
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this size increasing with age (thus,
rapidly growing eels will silver at
smaller sizes than slow-growing eels). In
males, silvering happens at a very early
stage, at a size typically greater than 35
centimeters (cm). In females, silvering
happens at a size greater than 40 to 50
cm (Goodwin and Angermeier 2003, p.
530; van den Thillart et al. 2005, pp. 31,
55).
Actual metamorphosis is a gradual
process occurring during the summer,
and in the fall eels metamorphosing in
preparation for migration back to the
spawning grounds have a silvery body
color, enlarged eyes and nostrils, and a
more visible lateral line (Dave et al.
1974; Lewander et al. 1974; Pankhurst
1983; and Barni et al. 1985 in van den
Thillart 2005, p. 12). As the structure
and metabolism of the liver changes, the
swim-bladder also changes, allowing for
increased gas deposition rates and
decreased loss of gas (McCleave 2001a,
p. 804).
A drop in temperature appears to
trigger the final events of
metamorphosis (gut regression and
cessation of feeding), which will lead to
migratory movements under the
appropriate environmental conditions.
It is theorized that responding to a drop
in temperature would help to
synchronize out-migrating eels, thus
increasing their chances of reaching the
Sargasso Sea simultaneously.
Conversely, increasing temperatures,
delays in migration, or possibly low fat
content will cause eels to start feeding
again and to revert to a yellow resident
stage. This would happen in the natural
environment if eels did not reach the
sea before the end of the migrating
season. It has been observed that even
after eggs and sperm have developed,
eels are capable of gut regeneration and
feeding (Fontaine et al. 1982, Dollerup
and Graver 1985, in van den Thillart et
al. 2005, p. 56). Van den Thillart et al.
(2005, p. 56) confirmed that silvering
may occur more than once in the
lifetime of an eel. It has been said that
this phenomenon would explain the
extreme variability in age and size of
silver eels. It has been hypothesized that
conditions encountered during oceanic
migration, such as the high pressure
they would experience at depth in the
open ocean, may complete the sexual
maturation of eels (Fontaine et al. 1985
in van den Thillart et al. 2005, p. 13).
Outmigration Life History Stage
Energy requirements. To successfully
complete the migration from the
continent to the Sargasso Sea (outmigration), great endurance and an
extensive fat reserve are required.
Larger, fatter eels have an advantage
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over smaller eels in reaching the
Sargasso Sea and having sufficient
energy stores to reproduce. Eels are very
efficient swimmers (eels swim
approximately four to six times more
efficiently than salmonids), and larger
eels appear more efficient than smaller
eels (van den Thillart et al. 2005, pp.
106–107). Also, larger eels usually have
larger fat stores per body weight. Silver
eels have ceased feeding, and use their
stored fat for energy during their
migration and for completing gonadal
growth. In a study conducted on
European eel, the most recent estimate
of necessary energy (fat) needed to
successfully complete the migration to
the Sargasso Sea from Europe and
spawn is 20 percent fat reserves, of
which 13 percent is for transport, and
an additional 7 percent for completing
gonadal growth. In European silver eel,
about 50 percent of the eels studied had
a fat percentage of 20 percent (van den
Thillart et al. 2004 in van den Thillart
et al. 2005, p. 109).
It is unknown if American eels
require 20 percent fat reserves.
American eels travel a shorter distance
to reach the Sargasso Sea than do
European eels. Actual distances, routes,
and depths of migration for adult eels
are unknown. Distances traveled by
migrating silver American eels likely
vary from under 1,500 km to over 4,500
km, shorter than the 5,000 km to 7,000
km likely traveled by European eels. An
American eel maturing in the
Mississippi River, Louisiana, would
travel a distance of over 2,200 km; from
South Carolina, 1,440 km; from
Chesapeake Bay, Virginia, 1,550 km;
from Newfoundland, Canada, over 2,800
km (McCleave 2001a, p. 805); and from
western Lake Ontario, over 4,500 km.
Silver eels, it has been hypothesized by
Knights (2003, p. 240), may follow the
deep currents (for American eel, the
Deep Western Boundary Current) to
return to the Sargasso Sea. However,
others believe the American eel migrates
in the upper portions of the ocean (see
van Ginneken and Maes 2005, pp. 385–
387; Tesch 2003, pp. 206–207).
Fecundity. Fecundity also varies with
size. Fecundity increases exponentially
with length, ranging from about 0.6
million to almost 30 million eggs
depending on the size of the female
(McCleave 2001a, p. 804). As an
example, in the lower Potomac
watershed, the average silver female
length of 734 mm would produce 2.7
million eggs; farther up the watershed
the average silver female length of 870
mm would produce 5.2 million eggs
(Goodwin and Angermeier 2003, p.
533). Fecundity is also linked to the
habitat which the eel occupies. In an eel
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farm growth experiment, favorable
nutrition was one of two factors (the
other being genetic heterozygosity,
where 2 different alleles are at one loci)
producing eels with a high reproductive
capacity (van den Thillart et al. 2005, p.
232). This high fecundity is thought to
compensate for very high larval
mortality (reported by Knights et al.
2006, p. 4, as most probably well in
excess of 99 percent).
Spawning. Spawning takes place in
the Sargasso Sea (Schmidt 1922 in
¨
Boetius and Harding 1985, p. 122). Here,
in the area where northern and southern
waters meet, it has been hypothesized
that there is some unidentified feature
of the surface water (perhaps the abrupt
horizontal temperate change of the
frontal zone located within the
subtropical convergence) that serves as
a cue for migrating adults to cease
migration and begin spawning (Kleckner
et al. 1983, p. 289; Kleckner and
McCleave 1988, pp. 647–648; Tesch and
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Wegner 1991 in Miller 2005, p. 1).
Spawning has not been witnessed by
humans, but the assumption is that
adult eels die after spawning.
Range
The extensive range of the American
eel includes all accessible river systems
and coastal areas having access to the
western North Atlantic Ocean and to
which oceanic currents would provide
transport. These drainages and coastal
areas are along more than 50 degrees of
latitude (from 5° to 63°) of the western
North Atlantic Ocean coastline, from
Northern Brazil/Venezuela to southern
Greenland (Scott and Crossman 1973,
pp. 624–625; Tesch 2003, pp. 92–97;
Helfman et al. 1987, p. 42), including
most Caribbean Islands and Bermuda,
the eastern Gulf of Mexico and
associated drainages including the
extensive Mississippi River watershed
(e.g., Mississippi River, Ohio River,
Tennessee River, Arkansas River, and
Missouri River) as far north as
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Minnesota, the Gulf of St. Lawrence and
the associated rivers, and Lake Ontario
and associated drainages. It is believed
that the eel was absent from the waters
of Lakes Erie, Huron, and Superior
before the completion of the Welland
Canal in 1829 (Patch 2006, p. 2). In
1878, the Michigan Fish Commission
planted young eels in southern
Michigan waters, and for more than a
decade, beginning in 1882, the Ohio
Fish Commission released young eels
throughout Ohio, including drainages to
Lake Erie (Trautman 1981, pp. 192–193)
(Figure 1). This extensive range should
provide the American eel with a buffer
against adverse conditions, as spawners
would still be coming from areas not
experiencing adverse conditions, and
would, due to random dispersal and
relatively homogeneous genetic
structure, be capable to successfully recolonize areas once the threat has
abated.
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It has been reported in other
¨
documents that Boetius and Harding
(1985) estimated that the American eel
range covers more than 10,000 km of
coastline; however, we could not locate
this information. Utilizing current
mapping technology, our estimate of the
available coastline (including barrier
islands) from Maine to Texas (Atlantic
and Gulf coast) is 29,612 km
(Castiglione 2006, p. 1).
As a result of oceanic currents, the
majority of the American eel population
is located along the Atlantic seaboard of
the United States and Canada. The
historic and current distribution of the
American eel within its extensive
continental range is well documented
along the United States and Canadian
Atlantic coast, and the SLR/LO. The
distribution is less well documented
and likely rarer, again due to currents,
in the Gulf of Mexico, Mississippi
watershed, and Caribbean Islands, and
least understood in Central and South
America.
Habitat
The American eel is said to have the
broadest diversity of habitats of any fish
species (Helfman et al. 1987, p. 42) by
occupying multiple aquatic habitats.
From an evolutionary standpoint, this
generalist use of habitats is favored in
fluctuating environments, while
specialists excel under constant or
slowly changing environmental
conditions (Richmond et al. 2005, pp.
279–280).
During their spawning and oceanic
migrations, eels occupy saltwater, and
in their continental phase, they use all
salinity zones: Fresh, brackish, and
marine (for detailed habitat use by life
stage, see Cairns et al. 2005). Eels occur
in waters highly productive to fish
species and those that are not, and from
waters of near tropical temperatures to
waters that are seasonally ice-covered
(McCleave 2001a, p. 800).
Growing eels are primarily benthic,
utilizing substrate (rock, sand, mud) and
bottom debris such as snags and
submerged vegetation for protection and
cover (Scott and Crossman 1973, p. 627;
Tesch 2003, pp. 181–183). In Canadian
waters, American eels hibernate in mud
during the winter. Wintering areas
include fresh water, brackish estuaries,
and bays with full strength salt water
(Cairns et al. 2005, p. 3.4.6).
Barring impassable natural or humanmade barriers, eels occupy all
freshwater systems, including large
rivers and their tributaries, lakes,
reservoirs, canals, farm ponds, and even
subterranean springs. The anquillid (eelshaped) body form allows for climbing
when at young stages and under certain
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conditions (e.g., rough surfaces),
allowing it to pass up and over some
barriers encountered during upstream
migrations in freshwater streams (Craig
2006, pp. 1–4). Eels are able to survive
out of water for an exceptionally long
time (eels can meet virtually all their
oxygen needs through their skin), as
long as they are protected from drying
(for which their ability to produce
mucus is of great adaptive significance),
and eels have been seen using overland
routes (while moist) when they
encounter a barrier, explaining their
entrance into landlocked waters (Tesch
2003, pp. 184–185) and their presence
above numerous dams and weirs
(USFWS 2005b, pp. 16–18).
Abundance. Abundance (density) and
distribution of eels within habitats may
be a function of distance from the ocean
and may not be related to habitat
features (Smogor et al. 1995, pp. 796–
797) (see also Density-Dependant
Dispersion). According to Smogor et al.
(1995, p. 799) when examining Virginia
streams, they found little connection
between habitat features and the
distribution and abundance of American
eels at least at a large scale. Their
results, they suggest, demonstrate a
diffusion pattern of eel occurrence. This
lack of eel-habitat relations (at least at
a large scale) within freshwater systems
suggests that comparison of abundance
for purposes of identifying quality
habitat would be misleading. Rather, it
has been suggested (USFWS 2006, pp.
13–14, 22) that the reproductive
contribution of an area to the total
American eel population would be the
best manner of identifying quality
habitat; however, reproductive
contribution estimates from throughout
the range of the American eel are not
available. Examples of densities
provided below are to illustrate the
variation of densities, not for
comparison of habitat importance.
Machut (2006) summarized freshwater
and brackish water density research and
standardized to eel densities per 100m2.
In Lake Champlain, Vermont, densities
ranged from 2.32–6.36 eel/100m2 (LeBar
and Facey 1983 in Machut 2006, p. 50).
In a tidal creek, Georgia, densities
ranged from 1.82–2.32 eel/100m2
(Bozeman et al. 1985 in Machut 2006, p.
50). A Massachusetts salt marsh yielded
densities of 8.46–9.28/100m2 (Ford and
Mercer 1986 in Machut 2006, p. 50). In
Machut’s own study in the Hudson
River freshwater tributaries densities
ranged from 0.28–155.06/100m2
(Machut 2006, p. 50), while in brackish
waters Morrison and Secor (2003 in
Machut 2006, p. 50) reported densities
of 0.03–0.24/100m2 . In four Maine
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freshwater rivers, densities ranged from
1.80–35.40/100m2 (Oliveira and
McCleave 2000, p. 144). Recent
population estimates of juvenile eels
(mostly elvers) on the South Anna River
in Virginia were 1.88 eels/100m2. On
the North Anna River, where the eels
were smaller, the population estimate
was greater at 4.48/100m2 (Odenkirk
2006, p. 1). No estimates of abundance
or density are yet available for marine
waters.
Habitat associations at a finer scale,
such as areas within a lake, have
recently been researched by Cudney
(2004). In her studies, she was able to
associate certain short-term habitat
conditions, such as non-stagnant waters
and to a lesser extent long-term habitat
features such as water depth and
percent organic matter, to a higher
probability of eel capture (Cudney 2004,
pp. 57–60).
Facultative Catadromy. Contrary to
the earlier dominating paradigm that the
eel growth phase is restricted to fresh
water, it has been suggested that
brackish (or estuarine) waters produce
eels that grow faster, mature earlier, and
emigrate as silver eels sooner than eels
in fresh water, and that some eels
complete their life cycle in brackish or
marine waters without ever entering
fresh water. Facultative catadromy,
therefore, refers to migrations into fresh
water as not being obligatory
(Tsukamoto and Arai 2001, p. 2651).
Morrison et al. (2003, p. 94) found
annual growth rates in brackish water
were two times higher than growth rates
of eels that resided entirely in fresh
water. The mechanism for this higher
growth in brackish water is not well
understood. Possible causes include an
increase in quality or quantity of food,
increase in habitat quality (Helfman et
al. 1987 in Morrison et al. 2003, p. 94),
lower resting metabolism resulting from
living in near-isoosmotic (same salinity
within the eel as the external
environment) conditions, increased
water temperature (which reduces the
amount of time that eels are dormant
during winter) (Walsh et al. 1983 in
Morrison and Secor 2003, p. 1499),
reduced effects from parasites,
decreased predation, or decreased intraor inter-specific competition. Morrison
and Secor (2003, p. 1499) hypothesized
that the higher brackish-water eel
growth measured on the Hudson River
is general to most large North American
estuaries.
Two other studies became available
during our status review, which
provided data on use by eels of marine
habitats during the eel growth phase
(Daverat et al. 2006; Lamson et al. 2006).
The first study, by Daverat et al. 2006,
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looked at habitat plasticity in the
American, European, and Japanese eel
(A. japonica;) the second, by Lamson et
al. (2006), at American eel in Canadian
waters. In the first study, habitat use
consisted of either residency in one
habitat (fresh, brackish, or marine) or
movements between habitats. Seasonal
or minor (1 or 2) movement patterns
were seen from brackish water to fresh
water and vice versa. Single habitat
switch events occurred, usually between
3 and 5 years of age. ‘‘Nomadic’’
movement between water masses of
different salinity was common; the
differences in productivity between
freshwater and brackish habitats (and
the resulting lower growth of eels in
temperate freshwater sites), the authors
state, might explain this phenomenon.
Occurrence of eels with no freshwater
experience was demonstrated, but such
eels accounted for a smaller proportion
of the overall sample than did eels with
some (even brief) freshwater experience.
Another interesting result was that eels
tend to prefer brackish and marine
habitats for feeding at the northern
extremes of their range. The authors also
suggest that this high degree of habitat
use plasticity suggests a remarkable ‘‘bet
hedging’’ strategy for angullids as a
group (Daverat et al. 2006, p. 11). In the
second study, conducted on American
eels in Canada, marine (saltwater)
resident eels were the dominant
migratory contingent of eels in saltwater
bays (85 percent). Resident eels were
established in salt and freshwater
habitats by the year after their arrival in
continental waters. Eels that shifted
between habitats increased their rate of
inter-habitat shifting with age. This
study also showed that plasticity of
habitat usage is the norm among eels,
and that the American eel life cycle can
be completed in marine waters (Lamson
et al. 2006, p. 1572). A study of Japanese
eel found that estuarine (43 percent) and
marine (40 percent) eels contributed
more spawners than did eels from
freshwater areas (17 percent), with some
seasonal differences. Additionally, the
study noted that eels from all three
habitats began their marine spawning
migration at about the same time. The
implication here is that eels from all
habitats can mix together during
spawning migration and potentially
contribute to the next generation
(Kotake et al. 2005, p. 220). In
Tsukamoto et al’s evolutionary
perspective, the authors hypothesize,
based on Inoue 2001, that molecular
evidence might suggest that
catadromous Anguillidae come from
deep-sea eels, with a migration loop that
extended to coastal waters and
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incidentally visited estuaries; these eels
may have eventually obtained a
reproductive advantage because of
higher food availability in estuaries than
in freshwater (Tsukamoto et al. 2002 in
Miller 2005, p. 2).
According to Lamson et al. (2006, p.
´
´
1568), Edeline and Elie (2004) reported
that European glass eels have distinct
individual salinity preferences. This
implies that young eels separate into
migratory contingents upon arrival on
the coast, with salt-seeking eels
remaining in marine waters while freshseekers ascend into fresh waters.
The benefits of facultative catadromy
include resource partitioning, by
minimizing intra-specific competition
between life stages and cannibalism of
young by adults. Additionally, there are
growth-temperature benefits, as shallow
brackish and fresh waters (especially
still waters) will heat up faster in the
spring and summer than marine waters.
Although not tested by any large-scale
quantitative distribution data, the
effective reproductive contribution of
brackish/marine habitats may be
substantial (Tsukamoto and Arai 2001,
p. 275; Jessop 2002, p. 228; Kotake et al.
2005, p. 220; Knights et al. 2006, pp.
12–13; Cairns 2006a, p. 1). Densities
may be relatively low in coastal waters,
but for European eel in England and
Wales, Knights et al. (2001 in Knights et
al. 2006, p. 13) calculated that estuarine
and shallow coastal waters (estimated at
5,000 km2) exceed that of freshwater
(1,035 km2).
Clinal Variations. American eels show
clinal variation (gradual changes over a
geographic area) in their growth rates
and size at maturity between the
southern and northern portions of their
range. Although mostly a warm water
species, Anguillids are eurythermal
(tolerant of a wide range of
temperatures) and can survive extremes
by migratory and cryptic behaviors.
Even so, growth seasons inevitably
shorten with increasing latitude. This
produces clines as you move north of
slower growth rates and larger size at
maturity, thus retaining relative
fecundity with increasing latitude
(Knights et al. 2006, p. 6).
Population Status
Typically an evaluation of population
status for a 12-month finding would
include a rangewide estimate of
population size and information on the
demographic structure of the population
and subpopulations as well as
population trend information in context
with historical data, and possibly an
evaluation of the long-term viability of
the current population through a
population viability analysis model.
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No rangewide estimate of abundance
exists for the American eel. Information
on demographic structure is lacking and
difficult to determine because the
American eel is a single population
(panmixia) with individuals randomly
spread over an extremely large and
diverse geographic range, with growth
rates and sex ratios environmentally
dependent. Because of this unique life
history, site-specific information on eels
must be evaluated in context with its
significance to the entire population.
Determining population trends is
challenging because the relevant
available data is limited to a few
locations that may or may not be
representative of the species’ range and
little information exists about key
factors such as mortality and
recruitment which could be used to
develop an assessment model.
Furthermore, the ability to make
inferences about species’ viability based
on available trend information is
hampered without an overall estimate of
eel abundance. Despite these challenges
we have determined the species
currently appears stable, as we explain
below.
The Stock Assessment Committee of
the ASMFC recently assessed the ‘‘stock
status’’ of the American eel (ASMFC
2006a), and this assessment was
subsequently reviewed by an
independent panel of scientists (ASMFC
2006b). The Stock Assessment
Committee concluded that the status of
the stock is uncertain as a result of
insufficient data. Their conclusion was
based on the review of nine indices, two
were fisheries-dependent and seven
were fisheries-independent. Of these
indices, one index shows an upward
trend over time, one shows no trend,
and the remaining seven show a
downward trend (ASMFC 2006a, p. x).
The committee hypothesized that the
indices exhibiting a downward trend
suggest that the stock is at or near
documented low levels. The glass eel
data from two Atlantic Coast sites were
not used, and the panelists who
reviewed the stock status felt that these
indices were a valuable asset. These
panelists interpreted the absence of a
declining trend in glass eel abundance
in either series over the last 14 to 15
years as the only positive indicator that
recruitment, at least to the glass eel
stage to these portions of the coast, had
not declined in concert with some of the
yellow eel indices (ASMFC 2006b, p. 4).
The ASMFC stock status assessment has
limited value in the 12-month finding
because the purpose of the ASMFC
stock status assessment is to inform
management of the commercial
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American eel fishery by determining
allowable harvest, not to look
specifically at long-term viability of the
species.
Recently Canada completed its review
of the American eel status within
Canadian waters as part of the
Committee on the Status of Endangered
Wildlife in Canada’s (COSEWIC) review
for possible listing under their version
of the Endangered Species Act, known
as Species At Risk Act (SARA). This
review also was more similar to a stock
status assessment than a population
viability analysis. They determined that
indicators of the status of the total
Canadian component of this species
were not available. Their evaluation of
the data (indices of abundance in the
upper SLR/LO declined by
approximately 99 percent since the
1970s and four out of five time series
from the lower St. Lawrence River and
Gulf of St. Lawrence declined) led them
to apply the Special Concern
designation (COSEWIC 2006, p. III).
Because the COSEWIC review focuses
on the status of American eels in
Canadian waters, the report also
discussed the ‘‘rescue effect.’’ In the
hypothetical scenario where the
American eel became depleted or
extirpated within Canadian waters
external components would ‘‘rescue’’
the species in Canada. These external
components refer to the young eels from
the Sargasso Sea that are from American
eels whose parents originated from U.S.
waters, and experience random
dispersal due to oceanic currents which
would continue to deposit leptocephali
into Canadian waters (COSEWIC 2006,
p. 43).
Together, however, these reports
provide a more recent presentation of
the individual data sets than was
available in the stock status report by
the International Council for the
Exploration of the Sea or ICES (2001,
pp. 51–52), which was the only stock
assessment available at the time of the
90-day finding published on July 6,
2005 (70 FR 38849). As a result of these
factors, our assessment of the American
eel population status will utilize the
available information to: (1) Provide
context of historical reports and current
landings data as a surrogate for absolute
abundance estimates; (2) evaluate the
data from each different life stage and
the significance of that life stage when
evaluating the population status of the
species including trend data in specific
geographic areas and each area’s
significance to the population status of
the species; and (3) evaluate the data to
determine if there is a sustained
downward trend in a location or
locations that would be considered
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representative of the entire range.
Together these will provide the basis for
our assessment of whether the species is
currently being impacted by threats to
the degree that the American eel meets
the definition of threatened or
endangered. In addition, in the 12month finding we also take into account
the species’ life history characteristics
and compensatory mechanisms (see
Background and for further discussion).
not captured) of yellow and silver phase
eels is greater than 15 million within the
areas fished. Given that not all areas
within the range of the eel are fished,
this number would represent a
minimum. These calculations are not
intended to be used as a formal estimate
of population size, but simply to
provide the context that large American
eels, throughout their range, likely
number in the many millions.
(1) Historical and Current Information
Historically eels were a significant
winter food source for Native Americans
(see Casselman 2003, for a compilation
of prehistoric and historic information
from the United States and Canada) and
later for European settlers. However,
qualitative rather than quantitative
information is all that is available from
these early times. In the early 1900s,
records from commercial fisheries began
to appear. For example, weirs at Oneida
Lake, Canada, caught 100 metric tons
(220,000 pounds) annually of emigrating
eels (Adams and Hankinson 1928 in
Casselman 2003, p. 260). Casselman
cites the subsequent construction of
dams and canals, which restricted
access to the lake as the reason for its
eventual extirpation from Lake Oneida.
Given the size of the harvest, Casselman
concludes that recruitment immigration
in the past was much more extensive
and probably much greater than in
recent times.
Although the current status of
American eels cannot be described in
absolute terms because rangewide
estimates of abundance do not exist
(ASMFC 2006a, p. viii; ASMFC 2006b,
pp. 3, 13), we provide below recent
ASMFC and COSEWIC landings data
(long-term fishery independent indices
do not exist) that indicate that the order
of magnitude of yellow and silver phase
eel abundance is probably in the many
millions. In the past decade, commercial
fisheries in the United States and
Canada have landed approximately 800
metric tons (1.8 million pounds) of
yellow and silver phase American eels
annually (ASMFC 2006a, p. 82). These
landings data provide a general sense of
eel abundance if we make assumptions
about the size and relative proportion of
eels that are landed. Specific data on the
size of eels harvested were not available,
but 45 cm was considered a reasonable
estimate (Cairns 2006b, p. 1). The
average weight of American eels 45 cm
long is 156 grams (g) (Cairns 2006b, p.
1), which indicates that 800 metric tons
is equivalent to over 5 million eels.
Assuming a high capture efficiency of
25 percent for the eel fisheries (Caron et
al. 2003, p. 235) suggests that the postfishery abundance (i.e., 75 percent are
(2) Trend Data From Different Life
Stages and Locations
Trends in American eel abundance
from fishery-independent indices (e.g.,
data from surveys and research) varied
among locations and life stages during
the past 10–25 years. Data from yellow
eels (which may include silver eels) and
glass eels (and elvers) are presented
below.
Yellow eel. Four indices (including
Maritime rivers in Canada and a
standardized U.S. coastwide yellow eels
abundance index) did not exhibit trends
(ASMFC 2006b, p. 3). Indices from
freshwater and tidal sites distributed
from the mid-Atlantic region north to
Canada and the St. Lawrence River
indicated a statistically significant
declining trend in yellow eel abundance
at three sites. Two of these indices, Lake
Ontario and the Chesapeake Bay index,
had strong and statistically significant
declining trends over the recent 1994 to
2004 time period, with 10-year declines
in the order of 50 percent in the
Chesapeake Bay index to 99 percent in
the Lake Ontario indices (ASMFC
2006b, p. 3). Smaller declines (15
percent) were reported in the St.
Lawrence estuary (COSEWIC 2006, p.
vi). Recent data suggest that declines
may have ceased in some Canadian
locations; but the positive trends in
some indicators for the Gulf of St.
Lawrence are, the COSEWIC report
states, too short to provide strong
evidence of an increasing trend
(COSEWIC 2006, p. 58).
It should be mentioned that yellow
eel indices may reflect local or regional
impacts, such as impacts from harvest
or turbine mortality (see Factors B and
E for further discussion). Additionally,
yellow eels have not yet been subject to
mortality that may occur during their
oceanic outmigration to the Sargasso
Sea. Therefore, yellow eel indices are
not the best indicator for estimating
annual reproductive success.
Evaluation of the Significance of
Upper SLR/LO. The extreme decline in
eels migrating up to the upper SLR/LO,
as tallied at the Moses-Saunders eel
ladder, has focused attention on the
potential impact of that decline to the
overall status of the American eel;
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however, COSEWIC states that a
rigorous way to quantify this impact to
the overall population has yet to be
developed (COSEWIC 2006, p. 35). The
suggestion is that the reproductive
contribution to the overall American eel
population from the upper SLR/LO may
be disproportionately larger than from
other freshwater portions of the range
because the American eels in the upper
SLR/LO are almost exclusively female
and highly fecund (producing many
eggs) due to their large size, and the
watershed is of considerable size. Two
methods for estimating the relative
reproductive contribution were
presented in the COSEWIC report (2006,
pp. 35–41), but both methods, they
state, are based upon questionable
assumptions and large uncertainties that
reduce confidence in the results.
Additionally, contributions from marine
and estuarine waters were not
considered in the analysis. According to
COSEWIC some sources of uncertainty
suggest that it is more probable that the
methods overestimate, rather than
underestimate, the reproductive
contribution of the St. Lawrence River
basin (COSEWIC 2006, p. 41).
Glass eels. Indices of glass eel
recruitment at the only two U.S. sites
with long-term data (North Carolina and
New Jersey) did not exhibit a declining
trend over the last 14–15 years (ASMFC
2006b, p. 4). Recruitment estimates into
Canadian rivers are available for two
Nova Scotian sites. The East River,
Sheet Harbour, abundance series is the
longest elver series available for the
species. Annual recruitment varied
without any upward or downward trend
from 0.1 to 0.5 million elvers between
1989 and 1999 (Jessop 2003a in
COSEWIC 2006, p. 28). In the East
River, Chester, the total run of elvers
peaked at 1.7 million in 2002. Since the
overlap periods of the two series are
strongly correlated, a combined index of
13 years was interpreted in the
COSEWIC report. Elver recruitment
showed inter-annual variability, but no
indication of decline between 1989 and
2002 (COSEWIC 2006, p. 28).
Glass eel counts, also called
recruitment indices, are the best
measure we have to annual reproductive
success (see section immediately
below).
(3) Evaluation of Trend Information
Of the available index data for the
different American eel life history
stages, we have determined that glass
eel indices best represents the species
status rangewide. Although we do not
have glass eel indices from the entire
range, the random nature of the
leptochephali dispersal allows us to
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consider these data representative of the
reproductive success of the species. As
described above, there is not evidence of
a sustained downward trend of these
glass eel indices; therefore, we conclude
that the American eel is not undergoing
a sustained downward trend at a
population level.
In summary, the best available
scientific and commercial information
indicates that despite a population
reduction over the past century, eels
remain very abundant and occupy
diverse habitats over an exceptionally
broad geographic range. Because of the
species’ unique life history traits areas
which have experienced depletions may
experience a ‘‘rescue effect’’ allowing
for continued occupation of available
areas without concern for genetic
fitness. Trends in abundance over recent
decades vary among locations and life
stages, showing decreases in some areas,
and increases or no trends in other
areas. Limited records of glass eel
recruitment do not show declines that
would signal recent declines in annual
reproductive success or the effect of
new or increased threats. Taken as a
whole, a clear trend cannot be detected
in species-wide abundance during
recent decades, and while
acknowledging that there have been
large declines in abundance from
prehistoric and historic times, we have
determined the species currently
appears stable.
Summary of Background
The American eel is an extremely
wide ranging species, continuing to
occupy most of its historic range. This
species is highly plastic in both its
behavior and physiology, being able to
occupy habitats ranging from sea water
to freshwater lakes. This species also
exhibits adaptive behaviors such as
switching between habitats and diets.
These life history characteristics
provide the American eel with the
ability to withstand a wide range of, and
changing, environmental conditions.
The best available scientific and
commercial information does not
indicate any sustained declining trend
in the American eel population.
Previous Federal Actions
On July 6, 2005, we published a 90day finding (70 FR 38849) which found
that the petition to list the American eel
presented substantial scientific and
commercial information indicating that
listing the American eel may be
warranted. That document initiated a
status review to determine if listing the
species was warranted. This 12-month
finding provides the results of that
status review.
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Summary of Factors Affecting the
Species
Section 4 of the Act (16 U.S.C. 1533),
and implementing regulations at 50 CFR
424, set forth procedures for adding
species to the Federal Lists of
Endangered and Threatened Wildlife
and Plants. In making this finding,
information regarding the status and
threats to this species in relation to the
five factors provided in section 4(a)(1) of
the Act is summarized below. We
examined each of these factors as they
relate to the current distribution of
American eel.
Regional information was more
obtainable from the Atlantic coast,
likely due to the economic interest in
the American eel. We have divided the
range of the American eel into seven
areas for purposes of discussion: (1) The
Gulf of Mexico (from south Texas to the
southern tip of Florida); (2) The
Mississippi watershed (Lake Itasca in
Minnesota to the Gulf of Mexico); (3)
The U.S. Atlantic coast (the southern tip
of Florida north to Maine’s border with
Canada); (4) The Canadian Atlantic
coast (Canadian border north to
Labrador, and including the Gulf of the
St. Lawrence); (5) The St. Lawrence
River and Lake Ontario (from the Gulf
of the St. Lawrence River to and
including Lake Ontario, abbreviated as
SLR/LO); (6) The Caribbean Islands
(Antigua, Barbuda, Bahamas, Cuba,
Dominica, the Dominican Republic,
Saint Kitts and Nevis, Saint Lucia, Saint
Vincent and the Grenadines, and
Bermuda); and (7) Central/South
America (Atlantic coasts of northern
Mexico; south through Guyana,
Suriname, and Venezuela; to northern
Brazil).
Addressing Uncertainties
The life history of American eels
presents unique challenges to
understanding the biological and
environmental processes influencing
eels at the species level. The eel’s
panmictic nature, wide geographic
range, oceanic spawning, and
segregation into freshwater, estuarine,
and marine environments all contribute
to the complexity of assessing status,
threats, and whether listing is
warranted. With many species,
population dynamics modeling can
inform listing determinations, but the
current understanding of American eel
population dynamics is rudimentary
due to its complex life history and the
paucity of data available for many key
parameters, such as recruitment,
growth, and mortality. A useful
conceptual framework for a population
dynamics model has recently been
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developed by a group of eel experts
(Angermeier 2005), but quantitative
analysis has been precluded due to a
lack of data.
As discussed below in the five factor
analysis, much speculation exists on
factors that could negatively affect eels,
often based on effects seen on other
species but with little supporting data
for eels. Much of the uncertainty exists
because decreased fitness would be
realized during life stages that are
currently not possible to assess,
specifically, the time between adult
spawning migration and the return of
glass eels to coastal streams. For
example, contaminants and swimbladder parasites may compromise the
health of silver eels during migration.
Contaminants could also contribute to
significant early life history mortality,
but these effects are not directly
observable.
We considered a number of questions
when reviewing the available
information and potential threats to
American eel. What is the population
status of American eel and how much
caution is warranted? What is the
species’ ability to withstand threats and
changing environmental conditions?
Would all eels throughout the widely
distributed range of the panmictic
population be affected by a given threat?
Is there evidence that indicates a threat
has caused significant population
effects, or are effects only speculative?
Has there been a reduction in juvenile
(glass eel) recruitment (which would
signal population-level effects)? And if
so, does it correlate in time (temporal
correlation) to the appearance of a
particular threat or threats? Answers to
these and other questions are important
to making a listing determination.
When addressing uncertainty (not
having complete, or in some cases any,
data on one or more of the questions
listed above), we employed a multi-step
approach. The first step was to review
all available data on the American eel
with regard to uncertainty and
determine, for example, if the data we
have regarding an impact at a local or
regional level implies an impact at a
population level, and if so, what the
likely response of the population is and
in what given time period. If data for
American eel is lacking, then we
reviewed data for other Anguillid
species, such as the European and
Japanese eel, and determined if the
application of that data was appropriate
to the analysis. If uncertainty still
remained high, then we requested
individual assessments from experts
regarding the probable implications to
the species given the uncertainties.
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In making this finding we examined
all the relevant data on threats, life
history characteristics (such as
resiliency and vulnerabilities), and
distribution information. We explored
all reasonable conclusions and
examined information to support and
refute theories on population level
effects, looking at whether the species
was currently showing the effects of any
population level threats. A population
level effect is defined for purposes of
this finding as an effect that is acting in
a way which puts the persistence of the
entire species at risk. Population-level
effects would be demonstrated by a
sustained downward trend in glass eel
abundance (recruitment) observed at
index sites that represent a substantial
portion of the range. Our five-factor
analysis follows.
Factor A. The Present or Threatened
Destruction, Modification, or
Curtailment of the Species’ Habitat or
Range
In analyzing these threats we
assessed: (1) The relative importance to
reproductive contribution of the various
habitats occupied by the American eel
during its life stages (such as spawning
habitat in the Sargasso Sea, oceanic
migration habitats, fresh water,
estuarine and marine habitats),
including which habitats are more likely
to produce males or females, various
growth rates, and levels of fecundity; (2)
the threats to these habitats; and (3) the
availability of that habitat to the
American eel. Much of the information
on the habitats other than freshwater
was not available for the 90-day finding,
and the new information has had a
significant effect on our assessment of
the status of the American eel.
Spawning and Ocean Migration Habitat
American eels spawn only in the
Sargasso Sea, and the young produced
from that spawning utilize ocean
currents to migrate to continental
habitats where they will grow to
maturity before again entering oceanic
habitats to migrate back to the Sargasso
Sea to spawn. Therefore, the spawning
and ocean migration habitats are of vital
importance to the persistence of this
species.
Seaweed harvest was indicated as a
possible threat to the American eel in
the ASMFC’s Interstate Fisheries
Management Plan for the American eel
(FMP) (2000, pp. 6, 34). The seaweed
Sargassum is commonly found floating
in the Sargasso Sea and drifting with
currents along the Atlantic coast from
Florida to Massachusetts. Harvesting
Sargassum, it was proposed, would
affect eggs and leptocephali, if
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harvesting occurs where eggs and
leptocephali are present.
After analysis of the available data,
we conclude that Sargassum harvest is
not a threat to American eel either in the
Gulf Stream current or in the Sargasso
Sea because first, studies of larval and
juvenile fishes associated with
Sargassum found no American eel
larvae (Settle 1993 in SAFMC 2002, pp.
20–23), and second, according to the
South Atlantic Fishery Management
Council (SAFMC), there has been no
commercial harvest of Sargassum
reported in U.S. waters since 1997. Any
future Sargassum harvest will be highly
regulated because in November 2002,
the SAFMC finalized the revised
Fishery Management Plan (FMP) for
Pelagic Sargassum Habitat of the South
Atlantic Region. This plan specifies
maximum and optimum sustainable
Sargassum yield and sets total allowable
catch limits, which severely limit
Sargassum harvest (SAFMC 2002, pp.
vi, viii). As such, we have concluded
that U.S. commercial Sargassum harvest
is not a threat to the American eel.
Furthermore, there is no information
indicating any other threat to the
Sargasso Sea or ocean migration habitats
(see Factor E for Oceanic Conditions),
and these habitats remain abundantly
available to the American eel.
Estuarine and Marine Habitat
Estuarine. The importance of
estuarine habitat is described by
Helfman et al. (1984, p. 135), Jessop et
al. (2002, pp. 84, 228), Morrison et al.
(2003, pp. 93–95, 97), and Knights et al.
(2006, pp. 12–13). An estuary is a semienclosed coastal body of water which
has a free connection with the open sea
and within which sea water is
measurably diluted with fresh water
derived from land drainage tributaries.
Estuarine habitat appears to not only be
habitat in which eels may choose to
remain during their continental phase,
but it is used by freshwater residents for
weight gain. According to Knights et al.
(2006, p. 25), inshore coastal and
estuarine mean net primary productivity
(the transformation of chemical or solar
energy to biomass) is greater than that
of rivers and lakes. Females inhabiting
estuarine waters, therefore, can provide
a greater reproductive contribution.
Estuarine habitat includes a mix of
males and females. Because eels grow
faster in estuarine waters than fresh
water, the average age of a female within
estuarine waters preparing to spawn is
much younger (9 years of age) than
females leaving lake habitats (24 years
of age in Lake Ontario). Variation in
maturation age benefits the population
by allowing different individuals of a
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given year class to reproduce over a
period of many years, which increases
the chances of encountering
environmental conditions favorable to
spawning success and offspring
survival. Jessop et al. (2002, p. 228)
provides an interesting perspective on
the relative production of silver eels by
comparing elvers that spend 1 to 4 years
in the estuary versus elvers that entered
the river shortly after continental
arrival. The authors suggested that the
relative production of silver eels was
380 times higher for juvenile eels that
spent 1 or more years in estuarine water,
due possibly to lower mortality rates in
the estuary than in fresh water (see
Background, Facultative Catadromy).
Helfman et al. (1984, p. 135), even as
early as 1984, recognized the value of
estuarine habitat where annual growing
conditions were more favorable.
Maximum size was greater in fresh
water, but lengths at a given age were
greater in estuaries. Morrison et al.
(2003, pp. 94–95) found that annual
growth rates were approximately 2 fold
higher in brackish water when
compared to annual growth rates in
fresh water. The theory is that eels
which grow faster, emigrate to spawn
earlier.
Although there have been historic
losses and degradation of estuarine
habitat (from, e.g., contaminants, low
dissolved oxygen, etc.), current rates of
estuarine habitat loss (nationwide) are
now estimated at 0.9 percent (averaging
5,540 acres annually) (Dahl 2006, p. 16).
The results of the most recent Status
and Trends of Wetlands in the
Conterminous United States from 1998–
2004 became available during the status
review. In summary, coastal wetlands
are still being lost but at a slower rate
than in prior reports. Human-caused
loss of deep salt water in coastal
Louisiana accounts for much of the
recent coastal wetland loss (Dahl 2006,
p. 16). Hurricanes can also transform
coastal habitats, but the effects of this
transformation of habitats on the
American eel have not been studied. A
U.S. Geological Service (USGS 2006, pp.
1–2) preliminary wetland loss estimate
for southeastern Louisiana from
hurricanes Katrina and Rita, which is
not included in the status and trends
report, is the transformation of some
64,000 acres of marsh to open water.
From the 1950s to 1970s, substantial
amounts of estuarine wetlands were
dredged and filled extensively for
residential and commercial
development and for navigation (Hefner
1986 in Dahl 2006, p. 48). Since the mid
1970s, however, many of the nation’s
shoreline habitats have been protected
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either by State or Federal regulations or
public ownership (Dahl 2006, p. 48).
Channel dredging and overboard spoil
disposal are common throughout the
Atlantic coast, and changes in salinity
as a result of dredging projects could
alter the distribution of American eels.
Additionally, dredging associated with
whelk and other fisheries may damage
benthic habitat for this species (ASMFC
2000, p. 42). Although it is likely that
dredging and overboard spoil disposal
at least temporarily degrade benthic
habitat, we were not aware of any
analysis indicating that these activities
are a threat to the American eel.
The two largest estuaries in North
America are both on the eastern
seaboard and support American eels:
The Chesapeake Bay and the AlbemarlePamlico Sound. The Chesapeake Bay
and its tidal tributaries have over 11,000
miles of shoreline; this is more than the
entire West coast. The AlbemarlePamlico Sound, located in North
Carolina, is the second largest estuary
with 1.5 million acres of brackish
estuarine waters (EPA 2006, pp. 3–4).
Although there are limitations to the
following data, as they include areas
outside the range of the American eel,
the status and trends report estimated
that in 2004, there were slightly more
than 5.3 million acres (2.1 million
hectares) of marine and estuarine
wetlands in the conterminous United
States. Eighty-six percent of that total
area was vegetated wetland (Dahl 2006,
p. 48).
Significant estuarine areas remain
from Maine to Texas. Therefore, this
important habitat remains available to
American eels, and there is
documentation of distribution of the
yellow stage of American eels within
estuarine areas from commercial harvest
data (Weeder and Hammond, in press,
pp. 1, 6), surveys, and research data
(Helfman et al. 1984, p. 135; Morrison
et al. 2003, pp. 91–92).
Marine. New information on marine
or saltwater habitat became available
during the status review (Daverat et al.
2006, see Background, Facultative
Catadromy). The relative importance of
marine habitat is not well understood,
and the use of marine habitat by
American eel for growth and maturity
has only been recently confirmed. There
was earlier confirmation in Japanese
and European eel. We do not know what
percent of the eel population inhabits
strictly marine habitats, but eels in this
habitat have high growth potential
(Knights et al. 2006, pp. 6, 10–11), there
is a predominance of females, and
extensive habitat is available. Sasal et al.
(2001 in Knights et al. 2006, p. 12)
found the female–male ratio to be 4:1 for
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Japanese eel caught in the East China
Sea from 1952–1999. Knights et al.
(2006, p. 13) calculates that for the
European eel in England and Wales the
combined estuarine and marine
contribution to reproduction probably
exceeds that of fresh water. Others have
also suggested that the percent of the
American eel population living in
estuarine and marine waters,
particularly those that will contribute to
future generations, may be quite high
(Cairns 2006a, p. 1). Although there is
no available data on the distribution of
the American eels in marine waters
throughout their range, the estimated
totaled nearshore habitats (tidal fresh
areas, through mixing areas, to seawater)
are substantial. In the United States
nearshore habitats have been estimated
at 5,379 km2 for the North Atlantic,
20,298 km2 for the Mid Atlantic, 12,172
km2 for the South Atlantic, and 30,604
km2 for the Gulf of Mexico (ASMFC
2000, p. 35; NOAA 2006, pp. 1–3); this
amounts to a total of 68,453 km2. No
threats to the American eel in marine
habitats are known to exist.
Freshwater Habitat
Lacustrine Habitat. Lacustrine, or
lake, habitat has historically been
considered among the most important
habitats for eel because some very wellknown lake habitats, such as Lake
Ontario, produce exclusively large,
highly fecund females (Castonguay et al.
1994a, p. 481; Casselman 2003, p. 255).
Studies by Oliveira et al. (2001, pp.
947–948) showed that the greater the
amount of lake habitat within a
watershed, the more the sex ratio favors
females. There are numerous lakes
within the distribution of the American
eel, many of which have likely been
impacted by water quality issues or
exotic species invasions, and American
eels have been denied access to some
historical lake habitats due to barriers
(see Riverine Habitat below for more
discussion of barrier impacts) such as
dams constructed in the past. We are
not aware of new dam construction
activities that are likely to threaten the
American eel. Below we will present the
information on two lakes, Lake
Champlain and Lake Ontario that are in
the Saint Lawrence River drainage. It
has been suggested in the literature that
a cause of declines of American eels in
these lakes was barriers.
The significance of Lake Ontario’s
reproductive contribution to the
American eel was presented and
discussed at a workshop (Casselman
2006, pp. 1–8 in USFWS 2006, pp. 8–
10) and presented in the recently
released COSEWIC Assessment and
Status Report on the American Eel
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(2006, pp. 35–41) (see Background,
Population Status for further
discussion).
Access to Lake Ontario and other
Great Lakes by American eel was
restricted to a degree by the building of
hydroelectric facilities on the St.
Lawrence River; however, the building
of canals also opened new avenues and
even provided passage past the natural
barrier of Niagara Falls. Eels migrating
into the Great Lakes and Finger Lakes
basin in New York historically had one
route through the Gulf of St. Lawrence
and up the St. Lawrence River to Lake
Ontario. Once in Lake Ontario, the eels
could access a large number of
tributaries in the United States or
Canada, but were blocked from Lake
Erie and the upper Great Lakes by the
natural barrier at Niagara Falls. With the
opening of the Erie Canal in 1825, and
later, the New York State Barge Canal in
1928, a second route up the Hudson
River and through the canal system was
created, allowing eels another access
route to Lake Ontario and the Finger
Lakes (Patch 2006, p. 2).
Although the building of the
Beauharnois Dam blocked American
eels from passing directly up the St.
Lawrence River for 70 years, many eels
were able to continue their migration
through the adjacent canal—the St.
Lawrence Seaway. Two ladders were
recently constructed on the Beauharnois
Dam, increasing the opportunities for
upstream eel passage at that site. A
second large hydroelectric dam, the
Moses-Saunders Dam, is located 40
miles upstream from the Beauharnois
Dam. From 1959 until 1974, eels were
able to pass upstream of the MosesSaunders dam only through the WileyDondero Canal (Verdon and Desrochers
2003, p. 140–141). In 1974, an eel ladder
was constructed on the Canadian side of
the Moses-Saunders Dam, allowing
American eels to again migrate directly
up the St. Lawrence to Lake Ontario
(Casselman et al. 1997, p. 163), and a
ladder on the U.S. side of the MosesSaunders Dam was completed in 2006.
These historical and recently
constructed fish ladders are likely to
benefit American eels in the SLR/LO by
providing them with multiple
opportunities to access to this drainage.
Lake Champlain also produces
predominately female eels. Declines in
Lake Champlain were noted in the
fishery in the Richelieu River (the river
carrying about 3 percent of the fresh
water from the lake to the St. Lawrence
River). The decline has been mainly
related to the rebuilding of two old
cribwork dams on the Richelieu River in
the 1960s (Verdon et al. 2002, p. 2) that
impeded access to Lake Champlain by
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young up-migrating eels. In 1997, a
ladder was retrofitted on the Chambly
Dam to enhance eel recruitment, and in
2001, the Saint-Ours dam, downstream,
was retrofitted with a similar eel ladder
(Verdon et al. 2002, p. 11–12). In 1997,
the total population at the foot of the
dam was estimated at 19,650
individuals, and minimum ladder
efficiency was estimated at
approximately 57 to 68 percent. Access
to Lake Champlain, having been
reestablished, now allows American eel
access to 1,200 km2 of habitat (Verreault
et al. 2004, p. 5).
Although we are not aware of a
rangewide analysis of the remaining
amount of lacustrine habitat available to
the American eel, according to the
NatureServe data a significant amount
of lacustrine habitat remains available to
the American eel. A survey of 203
randomly selected lakes in eight states
in the northeast United States showed
American eel as being present in at least
20 percent of the lakes sampled (Wittier
et al. 2001, p. 1).
Also, efforts are being undertaken in
the two large lake systems described
above to increase American eel
densities. A 10-year annual transfer to
Lake Champlain of 0.5 to 1 million
elvers from the Bay of Fundy (New
Brunswick, Canada) is underway as an
effort to improve abundance within
Lake Champlain (Dumont et al. 2006,
pp. 1–2). In Lake Ontario, 50,000 young
eels were recently stocked as a first step
in a Canadian multi-year plan to restore
the American eel to greater numbers in
Lake Ontario (CNEWS 2006, p. 1).
Riverine Habitat. Riverine habitat
within the range of the American eel is
highly variable with respect to water
depth, temperature, and flow, and
habitats available. Therefore, yearly
reproductive contributions vary among
river systems. The amount of habitat,
rather than specific types of habitat
within the river, primarily determines
how many eels a river can support
(Oliveira and McCleave 2000, p. 148–
149). Both males and females are
produced; densities of eels apparently
determine the sex of individual eels,
rather than habitat type (see
Background, Sex Determination).
Loss of access to riverine habitat has
been put forward as a threat to the
American eel (ASMFC 2000, pp. 35–39)
by both decreasing distribution and
abundance. However, most of the loss of
access to riverine habitat occurred prior
to 1960 and we have no information of
future water development projects that
threaten the American eel. Below we
will discuss effects of the construction
of dams to the eel’s distribution first.
Busch et al. (1998, pp. 1–3) conducted
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a preliminary analysis of stream habitat
availability for diadromous fish in
Atlantic coast watersheds. They
reported that from Maine to Florida,
15,115 dams have the potential to
hinder or prevent upstream and
downstream movement of fish such as
eels, resulting in a restriction or loss of
access to 84 percent of the stream
habitat within the Atlantic coastal
historic range. This constituted a
potential reduction from 345,359 miles
(556,801 kilometers) to 56,393 miles
(90,755 kilometers) of stream habitat.
However, only 35 percent (5,387) of the
dams from Maine to Florida are over 25
feet in height. The majority (65 percent
or 9,728) are, therefore, less than 25 feet
in height. Regional analysis of two
watersheds in the South Atlantic area
noted that eels remained present over
many barriers, until those barriers
reached 50 feet in height (Cantrell 2006,
pp. 4–5). Of the 15,115 dams, only 7
percent are for hydroelectric power
(Busch et al. 1998, p. 3).
Most barriers are thought to have been
in place before the 1960s. Castonguay et
al. (1994a, p. 484) reviewed major
habitat modifications as a potential
cause for the extreme decline of
American eels in the Lake Ontario and
Gulf of St. Lawrence ecosystems.
Anthropogenic (human-caused) habitat
modifications in the Lake Ontario and
St. Lawrence River ecosystem occurred
mostly before the 1960s, whereas the eel
upstream migration decline noted at the
Moses-Saunders Dam started only in the
early to mid 1980s. Castonguay et al.
(1994a, pp. 484, 486) proposed that the
lack of temporal correspondence
between permanent habitat
modifications and the start of the
regional decline evident in the SLR/LO
argues against the role of habitat loss in
the decline, as the decline should have
been evident earlier than the 1980s.
This assessment was tempered by the
brief mention that American eels may be
slower to respond to impacts than other
fish species.
Riverine habitats within the range of
the eel can be highly degraded through
contaminants (see Factor E,
Contaminants) and changes in
temperature, pH, and biological
communities. The effect, if any, on eel
is an increase in susceptibility in eels to
disease, likely decreased growth
(Machut 2006, p. 152; USFWS 2006, p.
27), increased elver mortality (Jessop
2000, pp. 523–524), and changes in
behavior (USFWS 2006, pp. 9–10).
Stream flow velocities can affect the
upstream migration of elvers (Jessop
2000, pp. 515, 520) due to their weak
swimming ability. However, reduced
velocities due to seasonal or operational
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changes of managed flows have likely
provided periods when velocities are
passable for migration. The elver’s
ability to find paths around these
velocity barriers has also been
documented (elvers have strong
climbing abilities and can negotiate
vertical barriers) (Jessop 2000, p. 520;
Craig 2006, pp. 2–4).
Impacts of barriers on distribution:
When discussing impacts of barriers on
distribution, we will cover impacts at
three levels: (1) Rivers, (2) watersheds,
and (3) the American eel’s entire range.
At the level of individual rivers, the
impact of barriers can range from very
little impact to local or regional
extirpation. This is because the effect of
barriers on eel upstream migration
appears to be site-specific. For example,
a steep vertical barrier has a different
effect on elvers, which can climb, than
on yellow eel, which do not have the
same climbing ability. Therefore, the
location of the barrier along the river
and in the watershed will dictate its
impact (USFWS 2005b, p. 16).
Additionally, the level of impact is also
affected by the type of barrier (i.e.,
hydroelectric dam, weir, old mill dam,
or dam for recreation, water supply, or
navigation), as well as how the barrier
is operated (if there is spill water), its
general condition (those in poor repair
are more likely to have rough areas or
spillage, both better for eel), whether it
was equipped with eel or other fish
passage, and other site specific
conditions (Goodwin and Angermeier
2003, pp. 532–533; USFWS 2005b, pp.
16–19). Indeed Busch et al. (1998, p. 3)
originally suggested that site-specific
assessments would be required when
further analyzing the impacts of barriers
to the American eel, and that their
estimate of 84 percent loss of freshwater
habitat for the American eel was a gross
estimate, provided as a starting point for
future scientific studies.
Our additional research into eel
distribution shows that eels remain
widely distributed within most of the
watersheds historically inhabited by the
American eel. For example, Jacobs et al.
(2004, pp. 325, 330), in a Connecticut
watershed survey, verifies the presence
of American eel above barriers and a
current extensive distribution.
American eel were the most ubiquitous
species of all fish species sampled in the
Connecticut River drainage, present in
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97 percent of all sites sampled and
common in both the main stem rivers
and tributary streams (Jacobs et al. 2004,
p. 325). Machut (2006, p. 49), in his
study of Hudson River tributaries, found
that American eels are the most
numerous fish within the tributaries
surveyed.
To better understand the impacts of
historically constructed barriers on eel
upstream migration and potential loss of
habitat we analyzed three watersheds
we think are representative of the U.S.
range of the species.
The Mississippi Watershed. The
American eel persists in the Mississippi
watershed (Mississippi River and the
tributaries of the Missouri, Arkansas,
Ohio, and Tennessee Rivers), albeit
having likely declined in abundance
during the past half century (Becker
1983, p. 258). Very little data exists on
the abundance of the American eel
within the Mississippi watershed (Ickes
et al. 2005, p. 4), both historically and
currently, as eels are not typically
targeted during studies and are likely
underestimated. The Long-Term
Resource Monitoring Program (LTRMP)
conducted by the Upper Mississippi
Environmental Sciences Center
(UMESC) observed 75 eels out of nearly
four million fish collected from 1993–
2002 (Ickes et al. 2005, p. 9).
The distribution of the American eel
remains widespread in the Mississippi
watershed even though it was
anticipated by Coker (1929, p. 173) that
the American eel, in time, would cease
to exist in areas of Minnesota,
Wisconsin, and Iowa, due to the
construction in 1913 of the Keokuk
Dam, or Lock and Dam 19, in Keokuk,
Iowa (River Mile 364). The barriers on
the Mississippi River mainstem are
mainly navigation locks and dams in the
upper portion of the river. These
navigation locks and dams were built to
hold back water and form deeper
navigation ‘‘pools’’ while allowing for
barge passage through the locks.
Presumably, these lock and dam
complexes allow for eel passage when
barges pass (Cochran 2005, p. 2) or eels
pass during high water stages, as
American eel are still found above
Keokuk Dam today. The Keokuk Dam is
currently the tenth dam eel encounter
during their upstream migration on the
Mississippi River.
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South Atlantic-Pee Dee River and
Santee River Basins, North Carolina and
South Carolina. American eels continue
to be distributed throughout the lower
areas of these watersheds, indicating
they are able to negotiate certain barriers
and persist within this historic habitat.
Of the six dams in the Santee and Pee
Dee River basin, eels are able to pass
four (Cantrell 2006, p. 3). They are
prevented from reaching their extreme
headwaters where they had historically
been reported as ‘‘everywhere common’’
by Jordan (1889, p. 139). Large (over 50
feet) hydroelectric and other dams likely
impede upstream movements of elvers
and subadult eels to these historic
habitats.
Androscoggin and Kennebec River
Basins, Maine and New Hampshire. Our
knowledge of current distribution of
American eel for the Androscoggin and
Kennebec watersheds of Maine and New
Hampshire is based on a systematic
survey in 2002 and 2003, and
supplemental electrofishing survey data
(Yoder et al. in preparation, pp. 1–7).
Presence of fishways on dams; dam
leakage, height, configuration, materials,
and location up the river relative to the
size of eel; water quality issues; and
presence of lakes (which may be of more
interest to eels due to odor cues) are
thought, by Wippelhauser, to play a role
in the distribution differences within
the two watersheds and explain why
eels are more abundant in the Kennebec
watershed (2006a, p. 1).
The American eel remains present
above the first dams encountered
inland, as well as subsequent barriers,
up to the Gulf Island Dam on the
Androscoggin (approximately 52 river
miles) and the Wyman Dam on the
Kennebec (approximately 122 river
miles), with anecdotal information
indicating that abundance has decreased
(Adams 1992, p. 86).
Rangewide our analysis of the impacts
of barriers was limited to the
information available, that of North
America. An update of NatureServe’s
distribution map (Figure 2) includes the
American eel freshwater distribution
information we received from most
States within the species’ historic range
as well as from Canada and a few of the
Caribbean Islands, along with
NatureServe’s existing database.
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At the scale analyzed, the American
eel remains distributed over roughly 75
percent of its historic native range
within U.S. watersheds (Castiglione
2006, pp. 1–5). Figure 2 represents the
historic (291,416,355 hectares) and
current distribution (163,781,049
hectares) of the American eel within its
native freshwater habitat in the United
States. Additionally, Figure 2 identifies
the area where the eel was introduced
and is considered currently present, an
addition of 2,921,343 hectares
(Castiglione 2006, pp. 1–5).
The watershed examples provided
earlier are indicative of the relationship
of barriers and eel distribution
throughout the species’ range in North
America. From these examples, and the
data from NatureServe, we conclude
that not all structures (natural or
human-made) considered barriers to
other fish species should be thought of
as barriers to the eel. We also conclude
that there are dams, other human-made
structures, and some natural features
that are complete barriers to American
eel. In the case of human-made
structures, those structures have
reduced the historical range of the
American eel.
The fate of eels that are unsuccessful
in passing a barrier is unknown, but it
has been speculated that eels may find
alternative habitat, that overcrowding
below the barrier may increase the
likelihood the eels will become male,
and that below the dams there is likely
increased competition, reduced food
availability negatively affecting growth
rates, and predation (USFWS 2005b, p.
19; Machut 2006, p. 53).
Impacts of barriers on density:
Whereas general fish surveys can
provide American eel distribution data,
few studies address the changes in eel
density (also called abundance) due to
barriers. Goodwin and Angermeier
(2003, p. 533) found that dams can
exacerbate the decline in eel density;
however, this is clearly the case for only
one in three dams within their study
area. Machut (2006, p. 51) found in the
Hudson River watershed, where there
are almost 800 barriers, that the first
barrier encountered dramatically
reduces eel densities, but did not
necessarily result in local extirpation.
Densities were highest below barriers,
while age, growth (in length), and the
number of females increased above
barriers.
Two aspects of the eel’s life history
add complexity to understanding the
true impact that decreased density may
have on eel reproductive contribution.
Densities decrease naturally with
distance from the Continental Shelf (see
Background), while relative female
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fecundity increases with lower density
(see Background). Based on these
factors, we conclude that low upstream
abundance is a natural phenomenon
exacerbated to varying degrees
geographically by human-made
structures and natural barriers, but that
relative reproductive contribution is not
lost in direct proportion to the decrease
in density (see Background, Distribution
Clines). Additionally, we conclude that
when taking into consideration or trying
to quantify the impact of barriers on the
American eel, site-specific information
on the barrier is critical, as is analyzing
the historic sex ratio of an area, the
dynamic between lower abundance and
the higher probability that females will
be produced, density-dependant growth
relationships, and length-fecundity
relationships. Unfortunately, the
information to conduct this
comprehensive analysis is not available.
The availability of riverine habitat can
be seen in Figure 2, and also be looked
at in terms of kilometers of riverine
habitat unimpeded. Unimpeded
freshwater habitat (riverine kilometers
downstream of terminal dams, the dams
closest to the ocean) in each river also
remains available to the American eel.
In the United States alone, from Texas
to Maine (not including the Great
Lakes), there remains over 590,000 km
of freshwater habitat available to
American eels downstream of terminal
dams or within rivers that do not have
significant barriers (such as the
Delaware River). An example of this
downstream available habitat on a
watershed basis is the 1,153 river miles
available on the Connecticut River
downstream of the terminal dam,
including both the mainstem and
tributaries (Castiglione 2006, p. 1–2).
In our analysis, we found that the
distribution of the American eels has
not been significantly reduced by
barriers, as many barriers do not
preclude upstream migration of the
American eel. Some dams appear to
form a complete barrier to upstream
migration, potentially responsible for
the reduction in available freshwater
habitat of approximately 25 percent.
Further, distribution is far less affected
by barriers than is density. If there were
population level effects from this
decrease in American eel distribution or
density in maturation habitats, there
would be corresponding declines in the
recruitment of juvenile eels; however,
this is not the case (see Background,
Population Status).
Summary of Factor A
Spawning and ocean migration
habitats are essential to the persistence
of the American eel; there are no
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apparent human-caused or significant
threats to these habitats; and, they
remain available and occupied by the
American eel.
Estuarine, marine, and freshwater
habitats provide maturation habitat for
the American eel, and new information
verifies that some portion of the
American eel population completes its
lifecycle without ever entering fresh
water. Of these maturation habitats,
freshwater habitat has been the most
impacted by human-caused actions such
as barriers (i.e., dams constructed for
hydroelectric, water supply, and
recreation purposes), most of which we
would consider historic losses; in which
case population level impacts have
likely been mostly realized. We are not
aware of future dam construction which
is likely to cause significant impact to
the American eel. We have concluded
that although some dams appear to form
a complete barrier to upstream
migration and likely caused the regional
extirpations seen in 25 percent of the
eel’s historic freshwater habitat,
American eels are able to negotiate
many barriers. This has allowed the
American eel to remain well-distributed
throughout roughly 75 percent of its
historic freshwater range, mainly in the
lower reaches of watersheds. American
eel abundance has been affected by
barriers to a greater degree than has
distribution; however, there is no
evidence that the reduction in densities
has resulted in a population level effect,
such as a reduction in glass eel
recruitment. Analyses of local and
regional declines in abundance do not
temporally correlate with the loss of
access to habitat.
The status of the American eel and
the effects of freshwater habitat loss
must be examined in light of the
American eel’s habitation in fresh,
estuarine, and marine habitats. Highly
fecund females continue to be present in
extensive areas of fresh water (lacustrine
and riverine) and estuarine and marine
habitats; males also continue to be
present in these habitats. Recruitment of
glass eels continues to occur in these
habitats with no evidence in reduction
in glass eel recruitment. For these
reasons, we believe the available
freshwater, estuarine, and marine
habitats are sufficient to sustain the
American eel population.
Factor B. Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes
In analyzing the threat of
overutilization, we focused primarily on
recreational and commercial fisheries
on the U.S. Atlantic coast and in Canada
because these fisheries are the most
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active. We will briefly characterize these
two fisheries and discuss recent
changes, summarizing the pertinent
scientific and commercial information.
For detailed descriptions of United
States and Canadian fisheries (e.g.,
harvest restrictions by State), see the 90day finding (July 6, 2005, 70 FR 38849)
or ASMFC 2006a (pp. 11–20) and for
Canada’s fishery, see the COSEWIC
report (2006, pp. 46–48). We will begin,
however, with a short discussion of the
factors that drive the commercial
harvest of Anguillid eel.
Commercial Fishery (Including Bait
Fishery)
Eels (most notably Japanese and
European eels) are popular seafood in
Europe and Asia, particularly Japan, and
to a much lesser degree in North
America. At this time, fish culturists
have not been able to provide the
conditions necessary for eels to
reproduce and mature in captivity;
therefore all eels consumed or used as
bait are taken from the wild. Some of
the eels taken from the wild as glass eels
or elvers are grown out to maturity in
aquaculture facilities.
The commercial eel harvest both here
or in other countries is driven in large
part by the international demand for eel
(see Pawson et al. 2005 for discussion of
international eel market), yet American
eel represent but a fraction of the total
international trade in eels. China
appears to be setting the world price by
both buying eels on the international
market and producing eels in extensive
aquaculture facilities (Dekker 2005, p.
2). According to TRAFFIC, a joint
program of the World Wildlife Fund and
the World Conservation Union (IUCN),
over 90 percent of the world’s eel
aquaculture yield takes place in the
Asian countries of Japan, Taiwan, and
mainland China (TRAFFIC 2002, pp.
11–12). Between 1998 and 2004, China
supplied two-thirds (i.e., approximately
130,000 metric tons) of the world’s
cultured eel production. The species
used in aquaculture in Asian countries
consists primarily of European and
Japanese eel. According to the United
Nations’ Food and Agriculture
Organization (FAO), even with
increasing dependence on European and
American glass eels for aquaculture
purposes with the decline of Japanese
eels (TRAFFIC 2002, pp. 13–14),
American eels represent only about 5
percent of the overall worldwide yield
of Anguillid eels (OLE 2004, p. 1; FAO
in Dekker 2005, p. 3). The insignificant
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contribution to the worldwide eel trade
indicates that the American eel harvest
is unlikely to be appreciably affected by
changes in international markets.
Commercial harvest of the American
eel in North America occurs mostly
along the Atlantic coast of the United
States and Canada. In the United States,
the commercial fishery occurs mainly in
the Chesapeake Bay with smaller
fisheries scattered throughout other
States. All continental life stages are
harvested commercially, but regulations
restrict harvest so that exploitation of
life stages differs geographically.
American eel fisheries are unevenly
distributed within Canada. In some
regions, there are intensive fisheries,
while in other regions, eels are
unexploited. All continental stages are
harvested commercially in Canada, but
the stages that are exploited vary
geographically (COSEWIC 2006, pp. 46–
47). Limited commercial fisheries exist
in Mexico and some Caribbean islands
(ASMFC 2006a, p. 14). No glass eel or
elver fishery exists in the Gulf of Mexico
(ASMFC 2000, p. 18).
Exploitation rates (the percent of
mortality associated with harvest) vary
with the life stage, fishing gear, and
other factors. Glass eels and elvers are
typically harvested as they ascend rivers
and estuaries. One study suggests an
exploitation rate of 30–50 percent of
arriving elvers (Jessop 2000, p. 523). If
there was no density-dependent change
in sex ratio, growth, survival, or
emigration rate in subsequent stages, the
reduction in egg production due to the
elver fishery would be equivalent to the
percent elver exploitation described
above. However, such densitydependent effects are believed to occur
(ICES 2001, p. 34). In other words, the
relatively high exploitation rate for glass
eels and elvers does not translate to that
level of reproduction loss because the
glass eels and elvers that are not
harvested have a greater potential for
survival and, therefore, reproduction.
Elver fisheries, it has been suggested by
Jessop (2000, p. 523), may be
biologically justified to a greater degree
in Nova Scotian streams with low pH,
given the abundance of elvers entering
these streams and the high mortalities
that occur during their first summer in
fresh water (rather than in more
productive streams with higher pH
values).
Silver eels are exploited in rivers
mainly in weir fisheries and in coastal
waters with eel pots. In the St. Lawrence
estuary silver eel fishery, mark-
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recapture experiments estimated
exploitation rates of 19 percent in 1996,
and 24 percent in 1997 (Caron et al.
2003, p. 239).
In the Chesapeake Bay, the estimated
exploitation rate is something less than
25 percent. The data collected did not
separate exploitation rates for yellow
eels harvested in the pot fishery from
eels that naturally emigrated from the
area. This combined fishing mortality
and emigration was estimated at 25
percent, significantly lower than the
Prince Edward Island fishery presented
below (ICES 2001, p. 34).
Data from Prince Edward Island,
Canada, were used by the authors of the
ICES report (2001) to calculate yellow
eel exploitation rates. They estimated an
approximately 50 percent rate of
exploitation in estuary and tidal waters
(ICES 2001, p. 41). The authors also
estimated how this rate of exploitation
would be expressed in loss of
reproductive contribution, but based on
some significant assumptions, they
consider the estimate preliminary. They
suggest the effect on reproduction
would be a decrease of approximately
90 percent, based on the premise that
the largest, and hence most fecund,
females are targeted. However, they also
note that the estimated reduction in
reproduction for the entire Prince
Edward Island area would be less than
this value, because there is no eel
fishery in non-tidal waters, and there is
minimal fishing effort in the central and
western portions of the Northumberland
Strait, which amount to about one third
of the Prince Edward Island coastline
(ICES 2001, pp. 34–35).
Exploitation rates are lacking for most
of the range where the American eel is
harvested, but the above examples show
how complex estimating exploitation
rates is, given that factors, such as areas
unfished, need to be accounted for
when evaluating harvest effects on a
species rangewide.
The American eel fishery has changed
over time. Harvest, or landings, were
significantly higher in the 1970s (Figure
3), presumably as a result of demand for
glass eels for the newly emerging
aquaculture industry in China (St. Pierre
1998, p. 1), which inflated prices and
made eel fishery profitable. Landings
have declined in the United States and
Canada since then; however, the reason
for the decline in landings appears
multifaceted.
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The price per pound fluctuates
considerably for American eel, thereby
affecting landings. For instance, the
Chinese aquaculture market still
requires glass eels to maintain the
established aquaculture business
(Moriarty and Dekker 1997 in ASMFC
2006a, p. 6), but when available, the
Chinese buy Japanese glass eel, which is
the eel preferred by Asians.
Consequently, the price for American
eel has dropped. ASMFC (2006, p. 7,
12–13, 43) also lists poor market
conditions as likely responsible for
more recent reductions in all
commercial eel fisheries. Since 1998,
glass eel market prices have fluctuated
from $300 per pound (1998), to $10–$15
per pound in 1999, to $105–300 per
pound in 2005, to $60 per pound in
2006 (Wippelhauser 2006b, p. 1).
License requirements and Stateregulated size and catch limits have also
played a role in the decline seen in
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landings (ASMFC 2006, p. 43). In 2000,
the ASMFC (the agency regulating
harvest along the U.S. Atlantic coast),
responding to the concerns of fishers,
scientists, and resource managers that
American eel had declined from historic
levels and that assessment data was
limited, implemented a Fishery
Management Plan that required States to
establish minimum size limits for
commercial eel fisheries.
Trends in Canadian eel fishery. In
Canada, there has been a trend towards
increasingly restrictive fishing
regulations in the last several decades,
especially in the Atlantic Provinces, and
especially since 2000 (Cairns et al. 2005
submitted in COSEWIC 2006, p. 48).
This could translate, we believe, to a
decline seen in Canadian landings data.
Changes include shortening of seasons,
increases of minimum size, caps on the
number of fishing gear that can be
deployed, and freezes on development
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of any new American eel fisheries
(COSEWIC 2006, p. 48). There was a
buy-out of 50 percent of commercial
licenses at Lake St. Pierre, the fishery in
the Richelieu River was closed in 1998,
and the fishery in the upper SLR/LO
was closed in 2004 (OMNR 2004, p. 1).
Glass eel and elver fishery only exists in
the Scotia-Fundy area of the Maritime
Provinces and occurs during narrow
time windows (COSEWIC 2006, pp. 46–
47).
Trends in United States glass eel and
elver eel fishery. During the lucrative
early 1970s, Florida, North Carolina,
South Carolina, Virginia, Massachusetts,
and Maine developed glass eel and elver
fisheries. By 2002, all Atlantic coast
States except Maine and South Carolina
had restrictions on harvestable eel size
or fishing gear that restricted glass eel
and elver fishery (ASMFC 2006a, pp.
12–18). One of those remaining States,
Maine, began in 1999 to limit glass eel
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and elver harvest through emergency
legislation with a limited entry system,
restrictions in fishing gear, restrictions
on locations, and a reduced length of
the season (March 15–June 15). This
later requirement allows for one or more
months in winter when glass/elvers are
not harvested. The emergency
legislation reduced fishing effort in
Maine by at least 79 percent (ASMFC
2005, p. 18), ensuring that a significant
run remains in Maine waters. Maine
was the only State reporting glass eel
and elver landings in 2004, at
approximately 0.5 metric tons, down
from 7.53 metric tons in 1995, and 9.98
metric tons in 1977. South Carolina and
Florida permit glass eel fishery, but it is
not active (ASMFC 2005, pp. 5, 14).
Trends in United States yellow and
silver eel fishery. Currently a yellow and
silver eel fishery exists to varying
degrees in all States and jurisdictions
along the Atlantic coast except
Pennsylvania and the District of
Columbia. South of Maine, the yellow
and silver eel fishery seems to be
primarily coastal pot fisheries, and
different States have varying
regulations, if any, imposed on this
fishery. In Maine, the yellow and silver
fishery occurs in both inland and tidal
waters (ASMFC 2006a, pp. 19–20). The
Maine fishery has declined since 1998
because of legislation and poor market
conditions, with prices paid declining
from $3–$4 per pound to $1.25–$1.75
per pound. Harvesters report that the
low prices are due to eels being grown
out in aquaculture facilities in Canada
(Knights 2003, p. 242). Eels grown out
in an aquaculture facility, a fish
company representative suggests, are
better suited to smoking, due to their
high fat content and uniform size and
shape. The uniform size is better suited
for the current mechanized processing
(Feigenbaum 2005, p. 12). The decline
in effort may encompass other areas
along the Atlantic coast as well (ASMFC
2006a, pp. 13–14). For example, on the
northern shores of New Jersey, the
number of active fishers has declined
from 16 in 1980s to 0 in 2004
(Feigenbaum 2005, p. 6).
In characterizing the future impact of
harvest, the literature supports the
prediction that 1970s harvest levels are
unlikely to occur again due to the
changes in the market (Pawson et al.
2005, p. 6; Dekker 2005, p. 2), including
the interest in eels raised in aquaculture
facilities rather than wild caught eel,
due to ease of processing (Feigenbaum
2005, p. 12); the implementation of
harvest regulations (ASMFC 2006a, p.
43); and the retirement of eel fishers
(Wippelhauser 2006b, p. 1).
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Population level impacts. In assessing
population level impacts of commercial
fishing on American eels, we took into
account both the species’ resiliencies
and vulnerabilities, and levels of
exploitation, including a review of
fished versus unfished areas in the
species’ range, and whether there is
evidence of a population level impact.
Resiliencies include the following: (1)
The wide range of the species, which
leaves many areas without fishing
pressure (USFWS 2005b, pp. 69–70, 76;
COSEWIC 2006, pp. 46–47, 53; Cairns
2006c, pp. 1–3); (2) harvesting within an
area is unlikely to substantially affect
the replenishment of the area through
recruitment (to the degree it might with
fish species that have river specific
stocks) because of the random nature of
recruitment (see Background section
and Factor E Ocean Conditions); (3)
harvesting will not affect genetic
variability because the species is a
single population; (4) eels have
relatively high fecundity rates; and (5)
the species possesses general plasticity
and robustness (Knights 2005 in USFWS
2005b, pp. 50–59); also see Background
for further explanation and citations).
Conversely, vulnerabilities include the
following: (1) All eel harvest takes place
before the species has had an
opportunity to spawn, and American eel
only spawn once; (2) all continental life
stages and multiple year classes are
subjected to harvest in some portions of
the species’ range; and (3) harvest of
large individuals unequally affects
females (eels below 40 cm in length are
either male or female, but almost all eels
greater than 40 cm are female) (ASMFC
2000, p. 2; USFWS 2005b, p. 75).
Although we have data on landings
(harvest) of American eel, we lack
specific data on fished versus unfished
areas over the range of the American eel.
Recent mapping by Cairns and others
(2006c, p. 3) has begun to identify (but
not yet quantify) fished versus unfished
areas in Canada, but initial results
suggest that much of the Canadian range
of the American eel is unfished
(COSEWIC 2006, pp. 46–47, 53). In
Canada, there is little eel fishing effort
in the Gulf of Nova Scotia, and none in
most fresh waters of the southern Gulf
of the St. Lawrence River. Many rivers
and coastal areas in the Scotia-Fundy
area of the Maritime Provinces are
unfished and Newfoundland and
Labrador have rivers which are not
exploited. Additionally, there are the
areas of harvest closure including the
Richelieu River and Lake Ontario
(Cairns 2006c, pp. 1–3).
Although we do not have similar
mapping in the United States, there are
considerable areas within the species’
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range that are not subject to harvest.
Commercial eel harvest is either
prohibited (such as in Tennessee, Todd
2006, p. 1) or at low levels in States
within the Mississippi watershed
(Keuler 2006, p. 1) and the U.S. portion
of the Great Lakes (Lutz 2006, p. 1).
Although the ASMFC was unable to
provide fished versus unfished areas
along the Atlantic coast, a fish company
representative who works with the
fishers was able to confirm that there are
areas along the Atlantic coast which
support eels and are not now being
exploited (Feigenbaum 2006, p. 6).
Modeling exercises have indicated
that harvest has depleted the abundance
of eels in the Chesapeake Bay, where
approximately 50 percent of the U.S.
yellow eel landings occur (Weeder and
Uphoff, in press, pp. 6–7). Modeling
conducted by BEAK (2001, pp. 31, 5.1,
5.7) for the purposes of prioritizing
factors influencing eel abundance,
ranked fishing mortality on yellow and
silver eels as the number one factor with
regards to American eel abundance in
the upper SLR/LO. The upper SLR/LO
was an area of substantial harvest
beginning in the 1970’s, with a peak in
1978 of 230 metric tons (Robitaille et al.
2003, p. 258). Commercial harvest in the
upper SLR/LO closed in 2004.
At a population level, however, one
must take into account existing
regulations and exploitation rates that
allow for: (1) A level of individuals who
are not subjected to fishing pressure; (2)
the theory that fishing of glass eels and
elvers does not necessarily represent a
substantial loss to reproductive capacity
of the species; (3) the vast areas that
remain unfished; and, (4) the lack of
evidence that there is a reduction in
glass and elver recruitment rangewide
(which would be the indicator of
overharvest) (see Background,
Population Status). Taking all these
factors into account, we have
determined that commercial harvest
currently affects the American eel only
at a local or regional level.
Recreational Fishery
Recreational harvest is either limited
or nonexistent throughout most of the
range of American eel. Eels are likely
purchased or caught by recreational
fishermen for use as bait for larger
gamefish such as striped bass (USFWS
2005b, p. 74; ASMFC 2005, p. 6), and
the remainder is mostly catch and
release (ASMFC 2005, pp. 5–6). The
NMFS Marine Recreational Fisheries
Statistics Survey (MRFSS), which has
surveyed recreational catch in ocean
and coastal waters since 1981, shows a
declining trend in the recreational catch
of eels during the latter part of the
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1990s. In 2003, total recreational catch
was 156,381 eels, and in 2004, 112,001
eels. In 2004, the combined catch from
New Jersey and Delaware represented
40 percent of the recreational American
eel catch, and the combined catch from
New York and Delaware represented 62
percent of the recreational American eel
harvest. About 79 percent of the eels
caught were released alive by the
anglers in 2004 (ASMFC 2005, p. 6).
To protect American eel from
unregulated recreational harvest, all
ASMFC member States were required to
establish uniform size (6 inches) and
possession limits (maximum 50 eels per
person per day) for recreational
fisheries, and recreational fishermen are
not permitted to sell eels without a State
license that specifically authorizes this
activity (ASMFC 2006a, p. 17). After a
review of the best available scientific
and commercial information, it does not
appear that recreational harvest poses a
significant threat to American eel.
There is little information in the
literature on subsistence harvest and
bycatch. But according to Laney (2006,
p. 1) and others (USFWS 2005b, p. 14,
79), bycatch of eels in marine waters,
during harvest for other targeted fish
species, does not appear to be of
concern for the American eel. This is
likely due to the fishing gear used in
these other fisheries (Laney 2006, p. 1).
Fisheries utilizing trawl gear may catch
eels, depending on the size of the
netting. Netting of a 1⁄2 inch and 1 inch
used in the late 1960s did catch eel, but
only a handful (Wenner 1973, p. 1).
Modern netting size is more specific to
the targeted fish species in an attempt
to limit bycatch.
Summary of Factor B
In conclusion, there are no data to
suggest that subsistence harvest,
bycatch, and recreational harvest are
having a significant impact on American
eel regionally or rangewide. Future
commercial harvest of American eel is
not anticipated to reach 1970s levels,
and we find it unlikely that American
eel landings will increase significantly
by future changes in the international
market.
Commercial harvest has had a strong
influence on eel densities in some local
and regional areas, but we see no
evidence that commercial harvest is
having an effect at a population level. A
population level impact would be seen
in declines in juvenile recruitment
rangewide, yet this is not in evidence.
It is probable that: (1) The random
dispersal of the larval stage enables the
species to successfully recruit to other
areas, including extensive unfished
areas, throughout its range, thereby
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buffering the effects of harvest; (2) the
compensatory mechanism of the
increasing probability of glass eel and
elver survival, or of undifferentiated
eels becoming female, as densities
decrease provide this species with some
level of resilience; and, (3) current
exploitation rates and regulations insure
that substantial numbers of eels remain
unfished. These factors are likely
sufficient enough to maintain the
species as a whole even under
foreseeable fishing pressure. As such,
we have determined that harvest is not
a significant threat to the American eel
at a population level.
Factor C. Disease or Predation
In our analysis of diseases and
predation, we focused on the diseases
and types of predation that were most
likely to affect the American eel at a
population level.
Predation
We evaluated changes in predation as
a result of human-caused activities. It
had been suggested in the 90-day
finding that American eels blocked or
delayed at upstream barriers could
experience higher than normal mortality
rates due to predation, because birds of
prey and piscivorous fish often
congregate at the base of dams to prey
on other fish species (USFWS 2005b, p.
20). However, we found nothing more
than anecdotal information on this
topic, and therefore we were unable to
quantify the impact of predation as a
result of barriers. Natural predation
rates are likely very high for elvers upon
entering freshwater (see Background,
Juvenille Mortaltiy and Jessop 2000, p.
522), but there is no evidence to
indicate that natural rates of predation
have risen, or that eel population
numbers are approaching a diminished
level where natural predation rates pose
an increased risk to the eel rangewide
(USFWS 2005b and 2006).
Disease
We analyzed whether the spread of
fish diseases, and in particular parasites,
has accelerated due to human activities,
including global transport of fish for
aquaculture, and whether the threat of
disease presented a risk to the American
eel at a population level.
Parasites. The parasite of most
concern is the nonindigenous nematode
Anguillicolla crassus, a parasite with
five life stages that becomes sexually
mature in the swimbladder of the eel.
The only other parasite found in the eel
swimbladder is another nematode,
Daniconema anguillae (Moravec and
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proportion of oxygen in the
swimbladder of adult eels by
approximately 60 percent when
compared to uninfected eels. Simulated
swimming experiments in European eel
indicate the impact of heavily
parasitized eels (20 or more parasites)
results in a decrease in swim efficiency
and possibly reduced buoyancy. Heavily
infected eels were not able to swim
longer than a few months. Parasites
cause the swimbladder to shrink,
resulting in higher costs of transport
(van den Thillart et al. 2005, p. 105). In
addition, heavy infection causes
deterioration of the swimbladder
function due to severe permanent
damage.
According to van den Thillart et al.
(2005, pp. 233, 236) a damaged
swimbladder interferes with the
buoyancy control, resulting in poor or
absent vertical navigation capacity in
the open ocean and a decrease in swim
efficiency which, they hypothesize,
prevents the completion of the
spawning migration. The likely result is
death en route to the spawning grounds
in the Sargasso Sea.
There is a significant level of
speculation about the impact of A.
crassus on the American eel during
outmigration and spawning, neither of
which can be easily studied under
natural conditions. A level of
uncertainty is therefore, inherent in our
analysis. Also unknown is whether
contaminants may act synergistically
with parasites, possibly magnifying the
impact on the species (USFWS 2006,
pp. 7, 26).
For the American eel, the number of
nematodes per infected eel (mean
intensities) is an important aspect in
evaluating the potential impact of this
nematode on American eel, as is
understanding the depths at which
American eels outmigrate back to the
Sargasso Sea, the length of that
migration, and further understanding of
what proportion of the American eel
completes its life cycle in salt and
brackish water where infection rates
may be significantly lower.
Unfortunately much of this information
is not available.
Mean intensities in American eels
have been found to be significantly
different among sites, including being
significantly lower in brackish water
when compared to fresh water,
(Morrison and Secor 2003, p. 1492). The
majority of studies of American eels
have shown fairly moderate levels of
intensity of infection. North Carolina
had a mean ranging from 2.0 to 12.3
nematodes per eel, depending on the
river (Moser et al. 2001, p. 851). Mean
intensities of infection of eels from the
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Hudson River in early studies were 1.0
to 1.7, increasing over time to 3.2 and
23.7, depending on the site (Morrison
and Secor 2003, p. 1491). Low to
moderate mean intensities of 2.6 to 9.0
were reported in the Chesapeake Bay
(Barse et al. 2001, p. 1366). It is
unknown if these relatively moderate
mean intensities would have the same
impact on American eels under natural
conditions as was reported by the recent
laboratory research by van den Thillart
et al. (2005, p. 105) on European eels
where higher densities of parasites
caused a decrease of the optimal swim
speed and increased the energetic cost
of swimming.
We remain cautious in extrapolation
of these preliminary laboratory studies
with regard to rangewide implications
given the absence of evidence for
population-level effects, such as
reduced recruitment of glass eels (which
would be an indicator of decreased
outmigration survival). This being said,
we acknowledge the statement by the
International Council for the
Exploration of the Sea (ICES 2001, p. 6)
that due to the fairly recent invasion of
the U.S. by A. crassus and the long-lived
nature of at least a portion of the
American eel population, the impact of
A. crassus on American eel may not yet
have been fully realized. ICES (2001, p.
6) concluded that, for the European eel,
the occurrence of this parasite does not
match the timeline for when the decline
in recruitment for European eel
occurred. Given the extensive research
on the European eel and the reasons for
its apparent decline this statement
should be given due consideration.
In summary, indigenous parasites are
not known to be of significant concern
to American eel at a population level.
During the status review, we were
provided with new information on the
nonindigenous parasite A. crassus,
including the northern extent of
invasion. The literature details the
impacts to individual European eels by
A. crassus in a laboratory setting, and
puts forward the hypothesis that these
impacts reduce an individual’s chance
of successful spawning. However,
similar research in the American eels
has yet to be undertaken and several
factors pertaining to the American eel
may indicate less potential impact from
A. crassus: (1) The mean intensities
reported for American eels appear to be
moderate; (2) the American eel has a
shorter outmigration distance to the
Sargasso Sea than European eels; (3)
some areas currently are free from A.
crassus infection (Canada, and possibly
Central and South American and the
Caribbean Islands); and (4) areas remain
where A. crassus is found, that are still
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producing uninfected outmigrating
individuals.
Pathogens. Viruses such as EVA (Eel
Virus—America) and bacteria are
present in the American eel, and
periods of stress, such as
metamorphosis, may activate viruses
and bacteria. Although mortality from
viruses may occur, there is no
information available about virus
prevalence and impact on American eel
at a population level.
Van den Thillart et al. (2005, p. 7)
found that European eels infected with
the rhabdovirus EVEX (Eel Virus
European X), a virus widely spread in
the European eel population, developed
hemorrhage and anemia during
simulated migration in large swim
tunnels and died after swimming for
1,000 to 1,500 km (estimated European
eel outmigration to the Sargasso Sea is
5,500 km). The resting group of eels did
not develop the disease, although they
were also infected with the virus. This
supports the theory that stress, such as
completing metamorphosis and
migrating, may activate the virus.
Because none of the infected swimming
eels survived the swim test, the authors
concluded that virus infections may
adversely affect the spawning migration
of eels. The virus infection appeared
more severe than the infection with the
swimbladder parasite, A. crassus (van
den Thillart et al. 2005, p. 7). In a report
on the presence of viruses in eel
populations from various geographic
regions and countries, the samples taken
from the United States (Virginia) and
Canada (St. Lawrence River) were
negative for EVEX virus (van Ginneken
et al. 2004, p. 270). Disease screening for
glass eels used in recent stocking
programs have also been free of EVEX
virus. Other pathogens, such as
Aeromonas salmonicida, a bacterium
known to cause furunculous lesions,
exist in cultured American eel
(Hayasaka and Sullivan 1981, p. 658),
but neither rates of infection in the wild
nor population level impacts have been
established.
In summary, pathogens such as EVEX
virus appear to have a significant impact
on eels in a laboratory setting; however,
the prevalence of this virus, or any other
virus or bacteria, in the American eel
population is not documented.
Summary of Factor C
We conclude that predation is not a
threat to the American eel at the
population level, nor are disease and
pathogens. We acknowledge that there
is a high level of uncertainty with
regards to the impacts on individual
silver American eels infested with A.
crassus during outmigration. However,
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given the absence of information for
population-level effects, such as
reduced recruitment of glass eels, and
given that there remain uninfected eels
for spawning and extensive areas of the
species range which are not currently
invaded by A. crassus or infection levels
are low to moderate, we have
determined that the current information
does not indicate that A. crassus is a
threat to the American eel at a
population level.
Because outmigration occurs in the
open ocean, direct study of the effect of
A. crassus under natural conditions will
continue to be difficult. This
emphasizes the need for data collection
and analysis designed to differentiate
between population fluctuations
responding to natural phenomena, such
as oceanic conditions, and those that are
human-caused. We support the
continuation and expansion of the
coastwide monitoring program started
several years ago, and the ongoing
research being conducted by the
scientific community.
Factor D. Inadequacy of Existing
Regulatory Mechanisms
Under this factor we will briefly
describe and address whether existing
regulatory mechanisms are adequate or
inadequate to conclude that the
American eel is not endangered or
threatened. As part of our analysis of
threats under Factors A, B, and E, we
describe how certain existing regulatory
mechanisms directly or indirectly
reduce these threats (we are unaware of
regulatory mechanisms that would
directly reduce the threats discussed in
factor C). Based on this analysis, we
conclude that Sargassum harvest,
freshwater and estuarine benthic habitat
destruction, streamflow alteration,
harvest, passage barriers, turbines, and
contaminants are not significant threats
to the American eel at the population
level and that additional protection is
not necessary to determine that listing
the species is not warranted. Because
we found no threat that, individually or
in combination with other threats, is
significant at a population level, there is
no instance in which the protections
provided by existing regulatory
mechanisms are inadequate such that
listing as endangered or threatened
would be necessary.
Seaweed Harvest
The status of the American eel with
regard to Sargassum harvest is
influenced by the effect of the following
regulation, and therefore, we describe in
this section how the existing regulatory
mechanisms directly or indirectly
reduces this threat. During the status
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4989
review, we evaluated the harvest
restrictions outlined in the second
revised Fishery Management Plan for
Pelagic Sargassum Habitat of the South
Atlantic Region. The specified
maximum and optimum harvest of
Sargassum severely limit Sargassum
harvest, and American eel larvae have
not been found in the Sargassum. We
concluded during the status review that
the commercial harvest of Sargassum is
not a threat to the American eel (see
Factor A), and therefore we find that the
regulations governing Sargassum
harvest are more than adequate for the
protection of American eel larvae.
Habitat Degradation
The status of the American eel with
regard to habitat degradation is
influenced by the effect of the following
regulations, and therefore, we describe
in this section how certain existing
regulatory mechanisms directly or
indirectly reduce this threat.
Stream Flow and Benthic Habitat.
During the status review, we evaluated
Federal and State and local regulations
that afford levels of protection and
regulate benthic habitat destruction and
stream flow alteration. The Clean Water
Act (33 U.S.C. 1251 et seq.) is the
primary Federal law, enacted at Federal
and State levels that restricts the
degradation of benthic habitats and flow
alteration. The Fish and Wildlife
Coordination Act, as amended (16
U.S.C. 661 et seq.), has been the
principal authority for incorporating
fish and wildlife conservation measures
into water development projects. The
River and Harbors Act of 1938 (Pub. L.
75–685) provided for wildlife
conservation to be given ‘‘due regard’’ in
planning Federal water resources
projects. The Federal Power Act, as
amended (16 U.S.C. 791a et seq.),
contains requirements to incorporate
fish and wildlife concerns into
licensing, relicensing, and exemption
procedures. The original Federal Power
Act provides for cooperation between
the Federal Energy Regulation
Commission (FERC) and other Federal
agencies, including resource agencies,
in licensing and relicensing power
projects.
Many States have specific laws and
regulations that limit benthic habitat
destruction and flow alterations. Some
mirror or implement Federal clean
water law regarding water quality
standards, including designated uses,
criteria, and an antidegradation policy,
which can provide a sound legal basis
for protecting wetland resources,
including benthic habitats for American
eels, through State water quality
management programs. In most of the
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eastern United States and Canada, the
riparian doctrine provides some
protection for maintenance of instream
flows. The riparian doctrine generally
affords some protection for off-stream
uses of water, while flow alterations
usually must conform to some
minimum standard.
Estuarine habitat. Laws, such as the
Estuary Protection Act (16 U.S.C. 1221
et seq.), the Estuaries and Clean Waters
Act of 2000 (33 U.S.C. 2901 et seq.), and
the Coastal Barrier Resources Act (16
U.S.C. 3501 et seq.), provide financial
incentives for estuary habitat protection
and restoration. Additionally, the Rivers
and Harbors and the Federal Power Act
described above would also address
impacts within estuarine waters.
During the status review, we
concluded that habitat degradation is
not a significant threat to the American
eel (see Factor A) and therefore we find
that the regulations governing activities
such as estuarine and benthic habitat
degradation and stream flow alteration
are adequate for the protection of
American eel.
Contaminants
In general, before the 1960s there
were no Federal environmental laws
regulating pollution. Concerns began to
mount with regard to the threat of
pollution to environmental resources
and were first addressed in 1965 with
the Solid Waste Disposal Act and the
Water Resources Planning Act. In 1970
the U.S. Environmental Protection
Agency (US EPA) was established to
‘‘protect human health and safeguard
the natural environment’’. Currently
there are numerous International,
Federal, and State regulations that
reduce the threats of contaminants to
environmental resources such as the
American eel. The 1972 Great Lakes
Water Quality Agreement was signed
between the U.S. and Canada to ‘‘restore
and maintain the chemical, physical,
and biological integrity of the waters of
the Great Lakes Basin Ecosystem’’. In
addition, Canada also has authority to
manage water resources and control
pollution under two primary acts, the
Ontario Water Resources Act and the
Environmental Protection Act. Federal
regulations that address environmental
contaminants include the Water
Pollution Control Act and the Federal
Insecticide, Fungicide and Rodenticide
Act of 1972, Safe Drinking Water Act of
1974, Resource Conservation and
Recovery Act of 1976, Clean Water Act
and the Soil and Water Resources
Conservation Act of 1977,
Comprehensive Environmental
Response Compensation and Liability
Act of 1980, and the Oil Pollution Act
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of 1990. Under the Clean Water Act, the
U.S. EPA can delegate many of the
permitting and regulatory aspects of the
law to state governments. In accordance
with the Clean Water Act and state
statutory authority, individual states
have developed water quality
regulations that are comparable to and
often more stringent than the Federal
regulations.
We concluded during the status
review that contaminants are not a
significant threat to the American eel
(see Factor E), and therefore we find that
the regulations governing contaminants
are adequate for the protection of the
American eel.
Fish Passage
The status of the American eel with
regard to barriers and turbines are
influenced by the effect of the following
regulations, and therefore, we describe
in this section how certain existing
regulatory mechanisms directly or
indirectly reduce these threats.
During the status review, we
evaluated section 18 of the Federal
Power Act (16 U.S.C. 791a et seq.).
Section 18 is the regulatory mechanism
that specifically provides for fish
passage prescriptions by the Secretary of
Interior (as exercised by the USFWS)
and the Secretary of Commerce (as
exercised by NMFS) for dams regulated
by FERC. Most States within the range
of the American eel in the United States
have specific fish passage laws, and
those State resource agencies often work
closely with the USFWS or NMFS when
creating fish passage facilities.
Sometimes fish passage is incorporated
in the 401 Water Quality Certificate
issued by the States under the Clean
Water Act (33 U.S.C 1251 et seq.).
Along the Atlantic coast, most fish
passage facilities are prescribed under
section 18 of the Federal Power Act or
recommended under section 10(j) of the
Federal Power Act administered
through FERC at hydroelectric facilities.
On the mainstem of the upper
Mississippi River /Illinois Waterway,
the Army Corp of Engineers (ACOE)
owns and operates a series of navigation
locks and dams for the Federal 9-Foot
Channel Project. However, other than
recommendations made by resource
agencies under provisions of the Fish
and Wildlife Coordination Act (16
U.S.C. 661 et seq.), there is no specific
regulatory mechanism requiring the
ACOE to provide fish passage (Wege
2006, p. 6). There may be opportunities
in the future for fish passage under the
proposed Federal Navigation and
Ecological Sustainability Program,
which requires Congressional
authorization and funding. Many of the
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large reservoirs in the Midwest were
constructed by the ACOE and remain
under its jurisdiction. In the Tennessee
River Valley, the Tennessee Valley
Authority owns and operates 49
developments for flood control,
navigation, and hydroelectric
development; none of these facilities is
operated specifically for fish passage,
although some upstream and
downstream passage is likely through
those mainstem dams with locks (Wege
2006, pp. 5–6). Recent records of
American eels from the Tennessee and
Cumberland River are few (Etnier and
Starnes 1993, p. 120).
Thousands of small dams that were
constructed over the last several
hundred years for water power to run
grist mills, saw mills, and textile mills,
as well as for water storage for drinking
water and other industrial and
municipal purposes, are exempted from
most modern regulatory mechanisms
except for State dam safety codes.
Thousands of dams in the Mississippi
River watershed and along the Atlantic
coast fall under this category. However,
as these structures age, funding is often
not available to bring them up to State
dam safety codes, which provides an
opportunity for their removal (Wege
2006, p. 5).
The Energy Policy Act of 2005 (Pub.
L. 109–58) amended the Federal Power
Act amended section 18 of the Federal
Power Act and calls for administrative
hearings when the material facts of an
agency-prescribed fishway measure can
be challenged by the dam owner or
other party to the proceeding. The
alternative fishway measure presented
by the dam owner or other party can be
adopted if it is as effective in purpose
and economically beneficial to the dam
owner. The burden of proof, of both the
benefit and need for the fish passage,
has been somewhat shifted from the
private sector (i.e., dam owner) to the
public sector (i.e., agency personnel).
Additionally, the agency is now to
consider the economic impact of a
fishway prescription to the dam owner.
While the process to consider
alternative fishways is new, the agencies
(USFWS and NMFS) have received and
considered alternatives from license
parties as a regular practice, and have
revised preliminary conditions and
prescriptions as new information was
received (Hoar 2006, p. 2; DOI 2005, p.
69808). It is yet to be seen whether these
amendments to the Federal Power Act
will have an effect on eel passage
implementation.
In Canada, there is no licensing or
regulatory system comparable to FERC
for hydroelectric dams. Canadian
resource agencies must rely on various
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fisheries laws that can be invoked, and
they must often negotiate the
construction of fishway facilities rather
than require them.
We have concluded that barriers limit,
and in some watersheds eliminate,
access to inland portions of the
American eel’s range in North America,
but that there is no indication that the
roughly 25 percent restriction of access
to historic freshwater areas is
significantly impacting the American
eel at a population level (see Factor A).
We have also concluded that turbines
can cause regional impacts to
abundance of American eels within the
watershed, but there is no evidence that
turbines are affecting the species at a
population level (for full discussion of
turbine impacts see Factor E). Therefore
we find that the regulations governing
fish passage are adequate for the
protection of American eel.
Harvest and Trade
The status of the American eel with
regard to harvest and trade are
influenced by the effect of the following
regulations, and therefore, we describe
in this section how certain existing
regulatory mechanisms directly or
indirectly reduce these threats.
During the status review, we
reexamined the ASMFC’s mechanism
for regulating the commercial and
recreational harvest of American eel
along the Atlantic coast States (see
Factor B. Overutilization) and ASMFC’s
flexibility in responding to changing
stock status. The American Eel Fisheries
Management Plan (FMP) requires that
member States establish uniform size
limits and other regulations for
commercial harvest. In 2005 and 2006,
the ASMFC underwent a public process
for potential changes to the FMP. In
2006, the ASMFC adopted Addendum I
to their American Eel FMP (ASMFC
2006c, p. 1; ASMFC 2006d, pp. 1–3)
which requires a reporting system.
Addendum 1 recommends the
implementation of a specific eel
harvester permit or license for each
State. Under this addendum, each
license requires reporting of trip-level
catch and effort, or States can choose to
implement an eel dealer permit and
reporting system. The American Eel
Technical Committee under the ASMFC
stated that this improved monitoring
system will assist in future stock
assessments. The permit or license
should be required for all eel harvesters,
including those who harvest eels for use
as bait. The American Eel Technical
Committee also recommended a specific
eel report from dealers and a license or
permit for dealers, including bait
dealers. Harvester and dealer reports
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must differentiate between the amount
of eels used or sold for food and the
amount of eels used or sold for bait. The
Addendum responds to concerns
regarding the lack of accurate catch and
effort data, and the critical need for
these data for stock assessment purposes
(ASMFC 2006a, p. 2). Although silver
eel fishery and seasonal closures were
options presented during the public
process (ASMFC 2004b, p. 7), no further
harvest restrictions, other than those
already laid out in the ASMFC’s FMP in
2000, have been implemented at this
time.
In Canada, harvest restrictions are
under the purview of the federal
government unless the authority has
been passed to the Provinces.
Restrictions and closures are already in
effect for certain areas in response to the
decline in the upper SLR/LO (see Factor
B. Overutilization). Provincial
management programs in Ontario and
Quebec have imposed license and
season restrictions, and reduced quotas,
in some cases to zero catch (Mathers
and Stewart 2005, p. 1). The federal
government of Canada retains authority
within the Maritime Provinces.
New information was gained on the
lack of restrictions in harvest from
responding countries outside U.S. and
Canadian waters, and the lack of import
restrictions in the responding European
countries (see Factor B). Our
determination, based on the analysis of
commercial harvest during the status
review, is that although abundance of
eels is likely affected locally and
regionally by commercial harvest,
commercial harvest is not a significant
threat to the American eel (see Factor
B).
To protect American eel from
unregulated recreational harvest, all
ASMFC member States were required to
establish uniform size (6 inches) and
possession limits (maximum 50 eels per
person per day) for recreational
fisheries, and recreational fishermen are
not permitted to sell eels without a State
license that specifically authorizes this
activity (ASMFC 2006a, p. 17). During
the status review recreational harvest
was determined not to be a significant
threat to the American eel at a
population level (see Factor B).
In summary, because we conclude
that Sargassum harvest is not a threat to
the American eel, and habitat
degradation, harvest, and fish passage,
including turbines, were not significant
threats to the American eel at the
population level, it is reasonable to
conclude that current regulatory
mechanisms governing habitat
degradation, harvest and fish passage,
including turbines, are adequate to the
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4991
extent that listing under the Act is not
necessary.
Factor E. Other Natural or Manmade
Factors Affecting the Species’
Continued Existence
Hydropower Turbines
During the status review, we
examined the extensive body of
literature on the impacts of turbines to
eels. Specifically, we looked at: (1)
Types of turbine impacts; (2) variations
in mortality and injury rates and
possible causes; (3) uncertainties and
information gaps; and, (4) impacts of
turbines on the American eel at a
population level.
During outmigration, as eels swim
downriver, where hydroelectric
facilities are present, some eels become
entrained and enter the turbines. Of the
eels that enter the turbines, some
survive and others are injured or die
(EPRI 2001, p. 3–1). Smaller turbines
and turbines that rotate faster pose the
greatest threat to eels. The degree of
injury and mortality increases with
larger eels (EPRI 2001, p. 3–8),
suggesting that mortality rates of large
female eels may be disproportionately
higher than mortality rates of males.
Turbine mortality to eels has also been
shown to be affected by dam size,
turbine type, load, and specific
operating conditions (including
nighttime versus daytime operation,
because eels tend to outmigrate during
the night; peak versus off peak power
production, and level of spill), and the
behavior of the eels (EPRI 2001, pp. 3–
4—3–10; USFWS 2005b, pp. 30–33).
There is only limited data on sublethal
effects to eels and their impact on
outmigration and reproductive viability
of the population. Sublethal effects
include injuries that may result in loss
of fitness (USFWS 2005b, pp. 34–36),
increased risk of predation, and delayed
migration (as observed in Anguillid
species native to New Zealand) (Watene
et al. 2002 in EPRI 2001, pp. 2–18).
The Electric Power Research Institute
report compiled data on eel mortality
through turbines and found that not all,
but most, eels go through turbines due
to migration behavior. For eels that go
through the turbines, the mortality level
was highly variable, depending on
turbine design, size of eels, and
operational conditions. For example, for
survival rates estimated at Moses—
Saunders and Beauharnois hydropower
facilities on the St. Lawrence River,
Francis turbines were found to result in
mortality rates of approximately 15
percent (85 percent survival), and fixedblade propeller turbines were found to
result in mortality rates of
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approximately 25 percent (75 percent
survival) (COSEWIC 2006, pp. 45–46;
see EPRI 2001, pp. 3–1—3–11 for more
details on the impacts to eels from
turbines). Higher mortality rates have
ˆ
also been reported. Monten (1985 in
McCleave 2001b, p. 593) reviewed
literature through the early 1970s on
injury and mortality on European eel
during turbine passage. He reported
injury rates, where injury likely resulted
in death, of 40 to 100 percent in 73-cm
eels passing through Kaplan turbines
under various operating conditions.
According to Hadderingh (1990 in
ASMFC 2000, p. 40) and McCleave
(2001b, p. 611), if American eels have to
pass through turbines in their
downstream migration, mortality rates
range from 5 to 60 percent.
Cumulative mortality refers to the
estimated combined mortality within a
watershed, and is thought to cause
significant reductions in that
watersheds’ eel reproductive
contribution to the population.
Verreault and Dumont (2003, p. 247)
estimated combined mortality rates of
40 percent for Lake Ontario s
outmigrating female eels from the
Moses—Saunders and Beauharnois
hydroelectric facilities on the St.
Lawrence River. The cumulative impact
of multiple hydroelectric projects
within a watershed, as simulated by
McCleave (2001b, p. 602), indicates
substantial decrease in overall eel
reproductive contribution from a
watershed, even when survival rates of
eel passage were high through each
successive turbine or dam project. The
simulated cumulative mortality within
the watershed was approximately 60
percent (40 percent survival) of overall
reproductive contribution when
mortality per dam was 20 percent (80
percent survival). McCleave states,
however, that his model is meant as a
tool to compare results based on
different inputs, not a definitive
statement about cumulative mortality
within the watershed. Based on the data
available, we can reasonably assume
that where American eels encounter one
hydropower facility during
outmigration, there is a typical mortality
rate in the range of 25 to 50 percent, and
when one or more turbines are
encountered, the range of mortality rate
increases to 40 to 60 percent for that
watershed. This still leaves escapement
values (the percent of individuals who
survive to continue outmigration) of a
minimum of 40 percent and a maximum
of 75 percent. Even if the mortality rate
has been underestimated, there are still
eels in freshwater areas that are
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unaffected by turbines, and eels that
survive passage in spillover.
We have updated Busch et al.’s (1998)
data on the percentage of dams with
turbines on the Atlantic coast and have
added the Gulf Coast. Out of the 33,663
dams, 1,511 (or 4.5 percent) are for
hydropower and, we assume, are fitted
with turbines. Of these only a small
percentage (2.06 percent) are on
terminal dams (Castiglione 2006, p. 1).
Terminal dams (dams closest to the
ocean) fitted with turbines affect
American eels throughout the watershed
as they outmigrate, but dams fitted with
turbines farther up in the watershed
impact only eels outmigrating from
tributaries and the mainstem of the river
above the dam, not outmigrating eels
from tributaries or mainstem river
habitats below the dam. Mapping also
showed that hydroelectric facilities
appear clustered in the Northeast and
Great Lakes area (Castiglione 2006, p. 2).
Still, we do not have the percent of eels
subject to turbines. This number could
be relatively small given that: (1) The
species’ range is extensive (see
Background, Range); (2) not all Atlantic
coast watersheds have multiple
hydroelectric turbines (USFWS 2005b,
p. 31); (3) dams that have turbines are
likely large dams (more then 50 feet
high), which often limit upstream
passage of eels in these watersheds
because of their height, and therefore
limit the risk of turbine mortality or
injury at maturity (see Factor A); and,
(4) there are tributaries to the Gulf of
Mexico that have limited impacts from
hydroelectric turbines, including the
Mississippi watershed (which has few
hydroelectric facilities) (Wege 2006, pp.
5–6).
The impacts from turbines to the
American eel, experts have suggested,
could result in a decrease in local or
regional abundance, as well as a
population skewed toward smaller and
younger females and more males, and
together these changes in the population
could ultimately result in a decline in
recruitment (USFWS 2005b, p. 34). In
analyzing the effects of turbines on the
American eel, however, we also took
into account that turbines principally
affect freshwater inhabitants, leaving the
portion of the population that inhabits
estuarine and marine waters largely
unaffected (USFWS 2005b, p. 3). As a
consequence, a decline resulting
specifically from turbine mortality may
be buffered by the spawning input from
eels residing in unaffected freshwater
habitats, or the estuarine or marine
habitats throughout its wide range.
It was also suggested by experts that
the importance of turbines as a
population threat can be assessed only
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in the context of a general
understanding of distribution and
dispersal patterns of the eel.
Specifically, a watershed’s specific
reproductive contribution rates and size
distribution of females needs to be
accounted for in determining the impact
of turbines on anything larger than a
watershed level basis (USFWS 2005b, p.
31). Currently there is no such
rangewide estimate.
In lieu of this rangewide estimate, we
can look at whether there has been an
impact to the American eel population,
and if so, if it relates to the construction
of hydropower facilities. As is discussed
under Population Status, there does not
appear to be a rangewide decline in
recruitment of juvenile eels; therefore,
we can draw no connection between
turbine mortality and population level
impacts. Additionally, according to
Castonguay et al. (1994a, p. 486), the
timing of the 1980s decline of the
American eel in the upper SLR/LO does
not correlate with the human-caused
changes that occurred on the St.
Lawrence River prior to 1965.
In summary, turbines, particularly
multiple turbines within a watershed or
turbines on terminal dams, can cause
substantial mortality within those
watersheds. However, turbines are
present on a small portion of the dams
within the Atlantic coast and are absent
from most of the barriers encountered in
the Mississippi Watershed, and there
remains a percentage of successful eel
passage through turbines or with spill
over the top of dams. Additionally,
there is no evidence of a population
level effect from turbine mortality. We
conclude that turbines are responsible
for decreases in abundance on a local or
regional scale, but turbine mortality is
not a significant threat to the American
eel at a population level.
Contaminants
During the status review, we
developed a summary of the current
American eel contaminant literature
(Roe 2006, pp. 1–26), and analyzed the
impacts of: (1) Existing contaminants on
the American eel life cycle, including
levels of uncertainty, and particularly
the inability to successfully raise eels
and consequently study the impacts of
contaminants on any of the eel life
stages; (2) new and emergent
contaminants; (3) other persistent
contaminants, such as genotoxic
polycyclic aromatic hydrocarbons
(PAHs); (4) non-persistent
contaminants, such as pharmaceutical
chemicals and pesticides; (5) complex
mixtures of contaminants; (6) vitamin
deficiency related to diet; and (7)
combined threats, such as disease,
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parasites, and contaminants, on eel
health.
(1) Existing Contaminants
Concentrations of polychlorinated
biphenyls (PCBs), PAHs,
polychlorinated diphenyldioxins/
polychlorinated diphenyl furans
(PCDDs/PCDFs), pesticides such as
mirex and
dichlorodiphenyltrichloroethane (DDT),
and metals such as mercury were
reported in yellow and silver American
eel tissues from eastern U.S. and
Canadian waters. However, much
uncertainty exists with regard to the
population’s rangewide contaminant
load since environmental contaminant
data were only available from a small
portion of the species’ range; therefore,
the contaminant loads within American
eel throughout its entire population
range are unknown.
The contaminant concentrations
reported in American eel tissues are
within the range of concentrations
associated with impacts that have been
documented in other fish species. These
environmental contaminants have been
shown to have biochemical,
immunological, genotoxic (chemicals
toxic to DNA), growth, survival, and
reproductive impacts on various fish
species. We believe that contaminants
therefore have the potential to also
impact the American eel (Roe 2006, pp.
5–8). Interestingly, American eels
survive with these contaminant loads at
concentrations that would be toxic to
other fish species. There is, however, a
potential for the impacts to be fully
expressed during critical periods of
their life cycle such as metamorphosis,
hatching, and larval development
(Robinet and Feunteun 2002, pp. 267,
270–272), all of which occur at sea and
therefore are currently impossible to
research under natural conditions
(USFWS 2006, p. 24–27). Because of
this species’ unique life history, caution
was suggested in utilizing surrogate
species data in determining impacts of
contaminants on eels (USFWS 2006, p.
24).
Inability to successfully study
contaminants on all American eel life
stages. To date, researchers have not
been able to successfully complete the
eel life cycle in the laboratory
(Penderson 2003 pp. 324, 336–337;
Palstra et al. 2005, pp. 533–534).
Research has also not been conducted
on the impacts of contaminants on eel
embryos and leptocephali, or during
metamorphosis from the yellow to silver
eel stage, or during outmigration and
reproduction. Two recent laboratory
studies on the reproductive capacity of
European eels by van den Thillart et al.
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(2005, pp. 110, 169) and Palstra et al.
(2006, pp. 147–148) indicated that
preliminary studies of PCB and dioxinlike contaminant impacts to maturation
and fertilization showed negative
impacts on egg quality and embryonic
development. However, artificial
hormone inducement of maturation in
European eels is complicated by high
female adult mortality rates and high
rates of embryo death after fertilization
(Pedersen 2003, pp. 336–337; Knights
submitted, pp. 1–2). Therefore, it is
difficult to be certain whether the
mortality rates are associated with
artificial maturation or fertilization
techniques or with exposure to
contaminants (Knights submitted, p. 2).
Unless or until the issue of embryo
death can be attributed exclusively to
the presence of contaminants, the data
is still inconclusive with regard to the
determination of the impacts of PCB and
dioxin-like contaminants at a
population level in the American eel.
(2) New and Emergent Contaminants
The impacts of new and emergent
chemical contaminants in fish are
unclear and not available for the
American eel. An example of new and
emergent contaminants presented
during the workshop (USFWS 2006)
was polybrominated diphenyl ethers
(PBDEs), a group of chemicals used as
flame retardants in a multitude of
consumer products (Agency for Toxic
Substances and Disease Registry or
ATSDR 2004, pp. 11–12). PBDEs are
similar to PCBs in that they are
lipophilic, persistent in the
environment, and bioaccumulate in
organisms. However, the impacts to fish
and other aquatic organisms have not
been completely defined in the
scientific literature. There is evidence
that PBDEs cause enzyme activity
alterations and delayed embryonic
hatching in fish, and they result in
behavioral alterations (Timme-Laragy et
al. 2006, pp. 1098–1103).
Concentrations of PBDEs have been
measured in European eels (de Boer
1990, pp. 315–318; Covaci et al. 2004,
pp. 3851–3855) and in other species
(Lebeuf et al. 2004, pp. 2973–2976);
however, the impacts of PBDEs to eels
were not discussed. Therefore any
impacts to the American eel at a
population level would be purely
speculative.
(3) Impacts of Genotoxic Contaminants
The impacts of genotoxic PAHs on the
eel remain uncertain. There is
considerable evidence that indicates a
causal relationship between exposure to
PAHs and genotoxic impacts such as
tumor frequency, deformities, and other
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lesions in fish, particularly bottom
feeding fish (Black 1983, pp. 328–333;
Metcalfe et al. 1990, pp. 133–139;
Baumann and Harshbarger 1995, pp.
168–170; Baumann et al. 1996, pp. 131–
149; Johnson et al. 1998, pp. 125–134).
Couillard et al. (1997, pp. 1918–1926)
documented the occurrence of
precancerous lesions in liver tissues
from migrating American eels from the
St. Lawrence River. The prevalence of
the lesions in the eel liver tissue was
reported to be correlated with increasing
contamination in eels, and the authors
concluded that PAHs may have been the
cause (Couillard et al. 1997, p. 1924).
Recent research in American eels
(Schlezinger and Stegeman 2000, pp.
378–384) and European eels (Doyotte et
al. 2001, pp 1317–1320; Bonacci et al.
2003, pp. 470–472; Mariottini et al.
2003, pp. 94–97) has shown that
induction of enzyme activity has also
been used as a biomarker for exposure
to PAHs and similar contaminants.
Genotoxic PAHs may be impacting
successful outmigration, but impacts of
lesions and tumors have not been
researched under natural conditions or
within the laboratory.
(4) Non-Persistent Contaminants
Short-term exposure to non-persistent
contaminants during critical American
eel life stages may be of concern
(USFWS 2006, p. 25), but uncertainty
remains. The literature has shown that
endocrine disrupting environmental
contaminants such as 4-nonylphenol
(which is formed during the industrial
synthesis of detergents), and pesticides
such as atrazine and diazinon, cause
physiological changes, inhibit growth,
and therefore inhibit the survival of
wild Atlantic salmon (Salmo salar)
along the Canadian Atlantic coast
(Moore and Waring 1996, p. 758;
Fairchild et al. 1999, p. 349; Brown and
Fairchild 2003, p. 146; Arsenault et al.
2004, p. 255; Waring and Moore 2004,
p. 93). American eels are sporadically
exposed to relatively high
concentrations of non-persistent
contaminants during their migration
through the St. Lawrence River to the
Sargasso Sea (Pham et al. 2000, p. 78).
For example, the largest primary physiochemical municipal sewage treatment
plant in North America is located in
Montreal, and treated effluent is
discharged to the St. Lawrence River
(Environment Canada 2006, pp. 1–3;
USFWS 2006, p. 25). At this location,
there is evidence of endocrine
disruption in other aquatic organisms
exposed to the effluent from 50 km
upstream to 50 km downstream of the
plant (Aravindakshan et al. 2004, pp.
´
156–164; Gagne et al. 2004, pp. 33–43).
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However, currently there is no
information within the literature on the
sensitivity of eels to short-term exposure
to these potentially endocrine
disrupting non-persistent contaminants.
(5) Exposure to Complex Mixtures of
Contaminants
The cumulative impacts of complex
mixtures of contaminants on eel species
are unknown. Fish and other wildlife
are not exposed to just one single
contaminant in the aquatic
environment. Contaminants mixed
together may interact and have additive
(Dioxin-like contaminants: Safe 1990,
pp. 71–73; Van den Berg et al. 1998, pp.
775–776) or synergistic (PAHs:
Wassenberg and Di Giulio 2004, p.
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(6) Vitamin Deficiency Related To Diet
In addition to contaminant-induced
impacts discussed above, decreased
concentrations of antioxidant vitamins
may also be impacting American eel
survival, but this remains uncertain.
Deficiences of antioxidant vitamins,
such as thiamine, vitamin B1, and
astaxanthin (a precursor to vitamin A),
have been associated with increased
early mortality in salmon and trout
species (Fitzsimons 1995a, p. 267;
Fitzsimons 1995b, pp. 286–288;
Vuorinen et al. 1997, pp. 1151–1163;
Fitzsimons et al. 2001, p. 229). It has
been suggested that the occurrence of
the early mortality syndrome in Lake
Ontario lake trout is related to alewife
(Alosa pseudoharengus) and their high
thiaminase content (Fitzsimons 1995b,
p. 288). Thiaminase are a group of
enzymes that break down thiamine in
the body and Alewife is a common food
item for young trout. Because alewife
are also consumed by American eels it
has been hypothesized that American
eels in Lake Ontario may be
experiencing effects from reduced levels
of thiamine. However, because this
hypothesis has yet to be tested this
theory remains speculative.
(7) Impacts of Combined Threats
Finally, contaminants can impact the
immune system and therefore increase
the organism’s susceptibility to other
threats such as diseases, parasites, and
bacterial and viral infections (Arkoosh
et al. 1996, pp. 1154–1161, Arkoosh et
al. 1998, p. 182; Grassman et al. 1996,
p. 829; Couillard et al. 1997, p. 1916;
Johnson et al. 1998, p. 125; Van Loveren
et al. 2000, p. 319; Zelikoff et al. 2000,
p. 325), but the effect on the American
eel remains uncertain. The cumulative
stress of the complex mixtures of
environmental contaminants and other
threats may potentially lead to increased
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mortality. Field studies have
documented susceptibility to infections
in European and North American fish
species (Arkoosh et al. 1998, pp. 188–
189; Van Loveren et al. 2000, pp. 322–
323; Zelikoff et al. 2000, pp. 325–330),
which would make these fish more
susceptible to disease. Bacterial
pathogens have been isolated in
American eels, and the authors
suggested that increased prevalence of
these pathogens may potentially be
related to stress and subsequent
decreased immune resistance (Hayasaka
and Sullivan 1981, p. 658; Davis and
Hayasaka 1983, pp. 559, 561; see Factor
C).
In summary, contaminants may
impact early life stages of the American
eel, but we remain cautious in
extrapolation of these preliminary
laboratory studies with regard to
rangewide implications without specific
information. A correlation between the
contamination of the upper SLR/LO and
the timing of the 1980s decline of
American eel in the upper SLR/LO is
not evident (Castonguay et al. 1994a, pp.
482–483), and current environmental
laws and regulations have significantly
decreased the discharge of many
persistent environmental contaminants.
Given the absence of evidence for
population-level effects, such as
reduced recruitment of glass eels (which
would be an indicator of decreased
outmigration survival, or egg or
leptochephali survival as a result of the
impacts of contamination), we believe
that the available information on
contaminants does not indicate a
significant threat to the American eel at
a population level.
Because spawning and egg and
leptochephali maturation occurs in the
open ocean, directly study of the effects
of contaminants under natural
conditions will continue to be difficult.
This emphasizes the need for data
collection and analysis designed to
differentiate between population
fluctuations responding to natural
phenomena such as oceanic conditions
and those that are human-caused. We
support the continuation and expansion
of the coastwide monitoring program
started several years ago, and the
ongoing research being conducted by
the scientific community.
Oceanic Conditions
During the status review, we explored
the relationship between oceanic
conditions and the recruitment of
leptocephali to coastal and riverine
habitats both hypothetically and
through correlative data. Additionally,
we investigated and describe briefly
here the types of oceanic conditions that
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have the potential to impact American
eels. Finally, we analyzed the potential
for oceanic conditions to impact the
American eel at a population level.
Variations in oceanic conditions have
been linked to wide-ranging and longterm changes in many fish, invertebrate,
and zooplankton species. General
ecological responses to oceanic
variations encompass changes in timing
of reproduction, egg viability, timing of
food availability, larval growth and
mortality, population sizes, spatial
distribution, and inter-specific
relationships (such as competition and
predator-prey relationships), by
affecting temperature, salinity, vertical
mixing, circulation patterns, and ice
formation. However, the relationships
are complex, usually non-linear, and
operate through complex mechanisms
through several trophic levels over the
ecosystem, and over a broad range of
time and spatial scales (Colbourne 2004,
p. 16). Further, a population’s response
is likely to vary in different regions
(Ottersen et al. 2001, pp. 1–14; Attrill
and Power 2002, pp. 275–278; Hurrell et
al. preprint, p. 10, 22–25, 38; Perry et al.
2005, p. 1–4; Weijerman et al. 2005
abstract and appendix 2, p. 3).
Oceanic conditions likely play a
significant role in the population
dynamics of American eel (Knights et al.
2006, p. 2), but the relationships
between specific oceanic conditions and
eel recruitment remain almost entirely
hypothetical. Changes in oceanic
conditions have previously been
thought not to be correlated with the
decline in the upper SLR/LO
(Castonguay et al. 1994b, p. 6; ICES
2001, p. 5). To better understand this
complex relationship given the scant
available literature, we requested
assistance from oceanic and eel experts.
Part of the assistance was a summary of
all available literature, entitled
American Eel Leptocephali-Larval
Ecology and Possible Vulnerability to
Changes in Oceanographic Conditions,
by M. Miller of the Ocean Research
Institute at the University of Tokyo
(cited as Miller 2005). Additionally, we
examined published and unpublished
data on the topic (Knights, Friedland,
Casselman, Miller, Kritzer, and Govoni
in USFWS 2005b, pp. 50–65).
The types of oceanic conditions that
have the potential to affect eels in the
North Atlantic include: (1) Changes to
sea surface temperatures (SSTs); (2)
changes to mixed layer depth (MLD); (3)
deflections of the Gulf Stream at the
Charleston Bump and Cape Hatteras;
and (4) other changes. Changes of SSTs
include inhibition of spring mixing, and
nutrient recirculation and productivity,
which may influence leptocephali food
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abundance. MLD (the depth to which
mixing is complete, relative to the layer
of ocean water beneath it) changes
include changes in size and depth of
leptocephali habitat, which would affect
leptocephali abundance, survival, or
transport. Changes in the Gulf Stream
could interrupt migration by slowing or
removing leptocephali from the Gulf
Stream, and any transport and
subsequent recruitment problems might
be accentuated at the extremes of the
species’ range. The ‘‘other’’ category
included changes to other aspects of the
Gulf Stream, such as the formation of
eddies, which may spin leptocephali off
of the main current (USFWS 2005b, p.
53).
Variation in oceanic conditions is
often depicted by the North Atlantic
Oscillation Index (NAOI). The NAOI is
a measure of oceanic-climate changes,
expressed as the difference in
atmospheric pressure measured between
Greenland and the Azores. The NAOI
has phases (positive and negative) that
have important oceanographic effects.
For example, a positive (high) NAOI is
indicated by periods of stronger winds,
greater surface-water mixing, reduction
of the Gulf Stream, shift of the Gulf
Stream in a northeast direction, and
increases in deep water formation and
water mass formation in the Labrador
Sea (and, it is hypothesized, weak eel
recruitment); a negative NAOI shifts the
Gulf Stream south and increases the
transport in the Labrador Current (the
western boundary current of the North
Atlantic subpolar gyre) (and it is
hypothesized, a strong eel recruitment).
These oscillations correlate with other
oceanic factors such as MLD, SST
anomalies, and position of the North
Wall (a steep water temperature
gradient) of the Gulf Stream (for further
discussion of NAOI see Weijerman et al.
Appendix 2, pp. 3, 9).
The NAOI has received considerable
attention because of its strong negative
correlation with recruitment of
European eels (glass eels recruited to
den Oever, Netherlands) (ICES 2001, p.
5) and a similar, but weaker, negative
correlation with recruitment of
American eels (juvenile eels recruited to
the St. Lawrence River) (ICES 2001, p.
5; Cairns et al. 2005, Table 9.2, p. 66).
From the mid 1950s to 1978/1979
winter the NAOI was in a 24 year
negative phase. From 1979/1980 winter
to 1994/1995 winter the NAOI was in a
positive phase (Weijerman et al.
Appendix 2, pp. 3, 9) and this positive
phase may have continued until
recently. During this prolonged positive
(high) phase European eel recruitment
had been correspondingly low (ICES
2002, p. 2). The last few winters,
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however, have not been strongly
positive (Hurrell et al. preprint, p. 4),
which may indicate that the NAOI is
beginning to shift to a negative phase,
which would benefit eels (USFWS
2005b, p. 66). A shift to a negative phase
would be consistent with the
observation that the NAOI seems to
follow 7- to 8-year cycles, superimposed
on 20- to 30-year cycles (Knights 2003,
p. 238).
The correlation between NAOI and
recruitment suggests that oceanic
conditions are currently the most
influential variable affecting
recruitment. As noted earlier, efforts to
model the population dynamics of
American eel are inherently limited by
sparse or nonexistent data. Nonetheless,
sensitivity analysis of one modeling
effort indicated that oceanic conditions
had greater eel population effects than
fishing, dams, or other habitat impacts
(BEAK 2001, pp. 5.10–5.11).
In summary, oceanic conditions
influence growth, recruitment, and
distribution of many marine species.
The interactions between the marine
environment and production of marine
species, however, are exceedingly
complex. Although the interactions are
not completely understood, the success
of early eel life stages and subsequent
recruitment to fresh water is dependant
on oceanic conditions, which are
subject to natural variation. Natural
conditions can, when a species is
significantly reduced in range or
abundance, be considered a threat.
However, there is no indication that the
American eel is suffering this level of
reduction in either abundance or range.
Therefore, because oceanic conditions
are within normal variations, the
American eel is evolutionarily adapted
to oceanic variations, and there is no
indication that the American eel is at a
reduced level where this natural oceanic
variation would significantly affect the
species, we have concluded that oceanic
conditions are not now, and there is no
information indicating oceanic
conditions should be in the future, a
significant threat to the American eel at
a population level.
Summary of Factor E
In conclusion, hydropower turbines
are a source of ongoing mortality. This
mortality has affected, and will continue
to affect, regional presence and
abundance of eels. However, the current
information does not provide evidence
to support turbines as a significant
threat to the American eel at a
population level. There is substantial
uncertainty on the effects of
contaminants on the American eel and
more research is needed. However, after
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examination, the literature does not
support a population level impact from
contaminants. Oceanic conditions are
highly variable and cyclical. They
determine recruitment to the continent,
and therefore they have a substantial
influence on the presence and
abundance of eels on the continent,
particularly in freshwater habitats.
Oceanic conditions are a naturally
occurring influence on the American eel
during its early life history, and are not
a significant threat to the American eel.
In sum, given the absence of evidence
for population-level effects, such as
reduced recruitment of glass eels, we
have concluded that there is not
supporting data to indicate other natural
or manmade factors as a significant
threat to the American eel.
Finding
The Act defines the term ‘‘threatened
species’’ as any species (or subspecies
or, for vertebrates, distinct population
segment) that is likely to become an
endangered species within the
foreseeable future throughout all or a
significant portion of its range. The term
‘‘endangered species’’ is defined as any
species that is in danger of extinction
throughout all or a significant portion of
its range. The principal considerations
in the determination of whether a
species does or does not warrant listing
as a threatened or endangered species
under the Act are the threats that
confront the species, as discussed in the
five factor analysis above.
In reviewing the status of the
American eel, we make the following
findings. The species has been
extirpated from some portions of its
historical freshwater habitat over the
last 100 years or so, mostly as a result
of dams built by the late 1960s. There
is also evidence that the species’
abundance within freshwater habitats,
and to some degree estuarine habitats,
has declined in some areas (e.g., upper
SLR/LO and the Chesapeake Bay) likely
as a result of harvest or turbine
mortality, or a combination of factors.
However, the species remains widely
distributed over the majority of its
historical range. Based on information
from the ASMFC stock assessment and
peer review and the COSEWIC
Assessment and Status Report, an
indication of decline exists in yellow eel
abundance, but recent glass eel
recruitment trends, although variable
from year to year, appear stable over the
past 15 years. The American eel is a
highly resilient species, with the ability
to occupy the broadest range of habitats
within freshwater, as well as estuarine
and marine waters, and it remains a
widely distributed fish species. The lack
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of population subdivision (i.e.,
panmixia) in the American eel provides
resilience to genetic problems that can
result from decline and isolation of
subpopulations.
Although roughly 25 percent of the
American eel’s historical freshwater
habitat is now inaccessible due to dams,
the loss of this habitat does not threaten
the species’ long-term persistence. This
is because a large amount of freshwater
habitat still remains (roughly 75 percent
of historic freshwater habitat in the
United States remains available and
occupied by the American eel), from
which both males and females
outmigrate, and because a portion of
American eels complete their life cycle
in estuarine and marine waters without
entry into freshwater. Although the
significance of the estuarine and marine
eel contribution to reproduction is
considered speculative by some, a
growing number of researchers think the
contribution could be substantial
(Tsukamoto and Arai 2001, p. 275;
Jessop 2002, p. 228; Kotake et al. 2005,
p. 220; Cairns 2006a, p. 1; Knights et al.
2006, pp. 12–13), and there is no doubt
that substantial amounts of estuarine
and marine waters remain available to
and are occupied by the American eel
throughout its range.
The threat of Sargassum harvest is no
longer considered a threat due to new
information indicating that the
American eel larvae do not utilize
Sargassum, and due to regulations
restricting its harvest. Recreational and
commercial eel harvests are no longer
factors of concern at a population level
due to economics, the species’
resilience, and existing regulatory
mechanisms. Although mortality during
outmigration due to parasites and
contaminants, and the potential effects
of contaminants on early life stages,
remain a concern, we have no
information indicating that these threats
are currently causing or are likely to
cause population level effects to the
American eel. We have no information
indicating that predation or competition
with nonnatives or mortality from
turbines are causing population-level
effects. Recruitment success of the
American eel is dependent on ocean
conditions, and variation in ocean
conditions causes fluctuation in
recruitment. However, because the
available information indicates that the
species remains widely distributed and
glass eel recruitment trends appear
stable over the past 15 years, observed
ocean conditions do not threaten the
current population status of the
American eel. Also, we have no
information to indicate that ocean
conditions are likely to threaten the
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American eel at a population level in
the future.
In reviewing the status of the
American eel, we also considered
whether there was any area where the
species is threatened or endangered
throughout a significant portion of its
range. We considered threats to its
spawning, migratory, and growth
habitats (see discussion under Factor A
and Ocean Conditions in Factor E) and
found no area where the species is
threatened or endangered throughout a
significant portion of its range. The
Sargasso Sea, where the American eel
spawns, is for that reason a significant
portion of the range, but we identified
no threats to this habitat. Similarly, the
open ocean migratory habitat of the
American eel is also a significant
portion of the range, but we identified
no threats to this habitat either.
The American eel’s growth habitat
consists of those areas, apart from its
spawning and migratory habitats, where
the species’ growth primarily takes
place. We evaluated whether the upper
SLR/LO, an area of the American eel’s
growth habitat that has experienced an
extreme decline in American eel
abundance, is a significant portion of
the range. The American eel is
panmictic, genetically homogeneous,
and capable of occupying a diversity of
growth habitats. It currently occupies a
number of growth habitats, each of
which is similar in habitat
characteristics. Therefore no one growth
habitat would be a significant portion of
the range unless it was significant in
terms of eel reproductive contribution.
Although it has been suggested that the
upper SLR/LO historically contributed a
disproportionately larger amount of
reproduction than other freshwater
areas of similar size, significant
uncertainties have been identified
regarding this analysis (COSEWIC 2006,
pp. 35–41). Even if the upper SLR/LO
had historically contributed a
disproportionately larger amount of
reproduction than other freshwater
areas of similar size (see Population
Status in Background section), our
consideration of the data on facultative
catadromy (the ability to grow and
become sexually mature in estuarine
and marine waters in addition to
freshwater) suggests that the total
reproductive contribution from the rest
of the range (including other freshwater
and all estuarine and marine waters)
outside the upper SLR/LO is
substantially greater than the historical
reproductive contribution from the
upper SLR/LO (see Population Status in
Background section). Consequently, any
historical additional reproductive
contribution from the upper SLR/LO
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does not make this area significantly
more important than if its historical
reproductive contribution was similar to
that of other similarly sized areas within
the range of the species. Because the
upper SLR/LO area does not contain any
unique or particularly high-quality
habitat, does not contribute to any
genetic differences, contains
substantially less than 50 percent of the
growth habitat for the eel, and does not
appear to contribute greatly to the longterm persistence of the species, we have
determined that it is not a significant
portion of the range. In addition, even
if the SLR/LO were to be considered a
significant portion of the range we find
from the record before us that the eel is
not threatened or endangered in the
SLR/LO because eels will likely persist
there into the foreseeable future (for
discussion of this ‘‘rescue effect’’ see
Background, Population Status). The
American eel is panmictic and
substantial reproductive contribution
comes from outside the upper SLR/LO.
We believe that the upper SLR/LO will
likely continue to receive eels and,
therefore, extirpation of eels from the
upper SLR/LO is unlikely.
In addition, we considered whether
there are any segments of the population
of American eel that would qualify as
distinct population segments (DPSs)
under the USFWS’s Policy Regarding
the Recognition of Distinct Vertebrate
Population Segments Under the
Endangered Species Act (DPS Policy)
(USFWS 1996). To be identified as a
DPS, a population must satisfy both the
discreteness and significance tests of the
DPS Policy. Because the species is
panmictic (a single inter-breeding
population), no part of the species’
population meets the discreteness test of
the DPS policy. Because no discrete
populations can be identified, there are
no populations for which we could
evaluate significance. Therefore, no
American eel DPSs can be recognized.
Due to the concerns about the status
of the American eel in Canada, we
considered delineation of a Canadian
DPS using the international border.
However, we determined that the
Canadian population of American eels
would not satisfy the significance test.
There is no evidence to suggest that eels
in Canada are genetically different from
eels in other parts of the species’ range,
that eels in Canada inhabit a unique
ecological setting, that loss of eels in
Canada would result in a significant gap
in the range of the species, or that the
Canada population of eels otherwise
could be considered significant under
the DPS policy. Also, because the
species is panmictic and juveniles are
distributed randomly over a wide range,
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and because substantial reproductive
contribution occurs over most of the
range, Canada will likely continue to
receive eels despite any reduction in
yellow eel abundance in Canada.
Therefore, the Canadian population
would not be considered endangered or
threatened and as a result would not
qualify as a DPS under the DPS policy.
In summary, we find that the
American eel remains widely
distributed over their vast range
including most of their historic
freshwater habitat, eels are not solely
dependent on freshwater habitat to
complete their lifecycle utilizing marine
and estuarine habitats as well, they
remain in the millions, that recruitment
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trends appear variable but stable, and
that threats acting individually or in
combination do not threaten the species
at a population level. On the basis of the
best available scientific and commercial
information, we conclude that the
American eel is not likely to become an
endangered species within the
foreseeable future throughout all or a
significant portion of its range and is not
in danger of extinction throughout all or
a significant portion of its range.
Therefore, listing of the American eel as
threatened or endangered under the Act
is not warranted.
Author
The primary author of this finding is
Heather Bell, Fisheries Biologist, Region
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5, USFWS, 300 Westgate Center Drive,
Hadley, Massachusetts, 01035.
References Cited
A complete list of all references cited
is available on request from the U.S.
Fish and Wildlife Service’s Region 5
Regional Office (see ADDRESSES section
above).
Authority: The authority for this action is
the Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Dated: January 23, 2007.
Kevin Adams,
Acting Director, U.S. Fish and Wildlife
Service.
[FR Doc. 07–429 Filed 2–1–07; 8:45 am]
BILLING CODE 4310–55–P
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Agencies
[Federal Register Volume 72, Number 22 (Friday, February 2, 2007)]
[Proposed Rules]
[Pages 4967-4997]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 07-429]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife and Plants; 12-Month Finding
on a Petition To List the American Eel as Threatened or Endangered
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of 12-month petition finding.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (USFWS), announce our
12-month finding on a petition to list, under the Endangered Species
Act of 1973, (Act) as amended, the American eel (Anguilla rostrata) as
a threatened or endangered species throughout its range. After a
thorough review of all available scientific and commercial information,
we find that listing the American eel as either threatened or
endangered is not warranted at this time. We ask the public to continue
to submit to us any new information that becomes available concerning
the status of or threats to the species. This information will help us
to monitor and encourage the ongoing conservation of this species.
DATES: The finding in this document was made on February 2, 2007.
ADDRESSES: Data, information, comments, or questions regarding this
finding should be sent by postal mail to Martin Miller, Chief, Division
of Endangered Species, Region 5, U.S. Fish and Wildlife Service, 300
Westgate Center Drive, Hadley, Massachusetts 01035-9589; by facsimile
to 413-253-8428; or by electronic mail to AmericanEel@fws.gov.
FOR FURTHER INFORMATION CONTACT: Heather Bell, at the street address
listed in ADDRESSES (telephone 413-253-8645; facsimile 413-253-8428).
Persons who use a telecommunications device for the deaf (TDD) may call
the Federal Information Relay Service (FIRS) at 800-877-8339, 24 hours
a day, 7 days a week.
SUPPLEMENTARY INFORMATION: The complete administrative file for this
finding is available for inspection, by appointment and during normal
business hours, at the street address listed in ADDRESSES. The petition
finding, the status review for American eel, related Federal Register
notices, and other pertinent information, may be obtained online at
https://www.fws.gov/northeast/ameel/.
Background
Section 4(b)(3)(B) of the Act, as amended (16 U.S.C. 1531 et seq.),
requires that, for any petition to revise the Lists of Endangered and
Threatened Wildlife and Plants that contains substantial scientific and
commercial information that listing may be warranted, we conduct a
status review and make a finding within 12 months of the date of
receipt of the petition (hereafter referred to as a 12-month finding)
on whether the petitioned action is (a) not warranted, (b) warranted,
or (c) warranted but the immediate proposal of a regulation
implementing the petitioned action is precluded by other pending
proposals to determine whether any species is threatened or endangered,
and expeditious progress is being made to add or remove qualified
species from the Lists of Endangered and Threatened Wildlife and
Plants.
On May 27, 2004, the Atlantic States Marine Fisheries Commission
(ASMFC), concerned about extreme declines in the Saint Lawrence River/
Lake Ontario (SLR/LO) portion of the species' range, requested that the
USFWS and the National Oceanic and Atmospheric Administration's
National Marine Fisheries Service (NMFS) conduct a status review of the
American eel. The ASMFC also requested an evaluation of the
appropriateness of a Distinct Population Segment (DPS) listing under
the Act for the SLR/LO and Lake Champlain/Richelieu River portion of
the American eel population, as well as an evaluation of the entire
Atlantic coast American eel population (see Finding for definition of
DPS) (ASMFC 2004a, p. 1). The USFWS responded to this request on
September 24, 2004; our response stated that we had conducted a
preliminary review regarding the potential DPS as described by the
ASMFC, and determined that the American eel was not likely to meet the
discreteness element of the policy requirements due to lack of
population subdivision (further analysis is provided under Finding).
Rather, the USFWS agreed to conduct a rangewide status review of the
American eel in coordination with NMFS and ASMFC (USFWS 2004, p. 1).
On November 18, 2004, the USFWS and the NMFS received a petition,
dated November 12, 2004, from Timothy A. Watts and Douglas H. Watts,
requesting that the USFWS and NMFS list the American eel as an
endangered species under the Act. The petitioners cited destruction and
modification of habitat, overutilization, inadequacy of existing
regulatory mechanisms, and other
[[Page 4968]]
natural and man-made factors (such as contaminants and hydroelectric
turbines) as the threats to the species.
On July 6, 2005, in response to the petition, the USFWS issued a
90-day finding on the petition (70 FR 38849), which found that the
petition presented substantial information indicating that listing the
American eel may be warranted. The finding noted concern that the
dramatic decrease in recruitment of American eel noted at the Moses-
Saunders Dam in Canada (on the St. Lawrence River), coupled with the
significant decline seen in the European eel (ASMFC 2000, pp. 12-14),
could indicate a decline in the American eel. Information on possible
reasons for this suggested decline included the following threats:
Commercial harvest, habitat loss and degradation (primarily the loss of
wetlands and upper tributary habitat), hydropower turbine mortality,
and inadequacy of existing regulatory mechanisms. Other potential
threats, such as seaweed harvest, benthic (sea or lake bottom) habitat
destruction, alterations of stream flow, disease, predation, and
contaminants, were not fully addressed or supported by the information
presented in the petition. Further analysis of oceanic variations (such
as changes in the Gulf Stream) were recommended in the 90-day finding,
particularly in light of the scant direct evidence and the potential
for oceanic variations to be compounding or confounding the impact of
other threats. Additionally, the 90-day finding concluded that the
complex life history and the incompleteness of historical data
(abundance, stock composition, life stage mortality rates, and
exploitation rates) made it challenging to understand the potential
influence of multiple threats to the American eel (USFWS 2005a, p.
38860).
In response to our 90-day finding's request for information for use
in the species' status review, we received comments and information on
American eel from the majority of the State fish and wildlife agencies
within the range of the eel; State universities; State and university
museums; the U.S. Forest Service (USFS); National Park Service (NPS);
U.S. Geological Survey (USGS); Army Corp of Engineers (ACOE); the
Department of Defense; the ASMFC; the Great Lakes Fisheries Commission;
Department of Fisheries and Oceans (Canada); Tribal Nations; academics
and researchers from the United States, Canada, Japan, and several
European countries; hydropower and fishing industries; nongovernmental
organizations; private citizens; and other entities. Additionally, we
coordinated with the USFWS's International Affairs Program (IAP) to
obtain information on international trade and with State and Federal
law enforcement officials on illegal trade. Although all countries
where the American eel is native were contacted regarding information,
there was no available data on eel distribution, habitat use, habitat
degradation or loss, or other threats (other than international harvest
data) from Central or South America. Distribution information was
provided by some Caribbean Islands. Therefore, the status review
focused on where data is available within the North American Continent.
A status review allows for additional collection, clarification,
and interpretation of information on the status of the species by the
USFWS. The resulting status review, from which the 12-month finding is
based, relied on our extensive review of the existing literature, data
resulting from the 90-day finding request for information, and new
information obtained during the status review period. Among the new
information we received, the documents most relevant to the status
review include the recently completed stock assessments for the
Atlantic coast (ASMFC 2006a and b), the American eel data assembled for
the Canadian stock assessment (Cairns et al. 2005), and recently
completed research on life history and potential threats to the
American eel (van den Thillart et al. 2005; Oliviera in USFWS 2006;
Machut 2006; Lamson et al. 2006; Devarut et al. 2006; Knights et al.
2006).
Also, because of the large body of literature and the uncertainty
surrounding several threats, we hosted two scientific workshops with
over 25 scientific experts. The goal of the workshops was to insure
that the USFWS properly utilized the best and most current scientific
and commercial data available in conducting the status review. To reach
this goal, each of the experts was asked a series of facilitated
questions to assess the presented information (which included multiple
factual inputs, data, models, assumptions, etc.), including the
completeness of the literature selected, and to comment on the
relevance and quality of the literature for purposes of our status
review (see workshop summaries Web site at https://www.fws.gov/northeast/ameel/). The USFWS recorded each expert's individual
assessments and the basis for those assessments in a compendium (cited
in the finding as USFWS 2005b and 2006). Workshop objectives included
determining the following: Utility of the information; life history
stages vulnerable to certain threats; the geographic scope of the
threats; the immediacy of the threats; and uncertainties in the
available information and the potential implications of those
uncertainties in making a status determination.
The selection of the expert panelists was based on recommendations
from within and outside of the USFWS and NMFS (the Services). The
panelists selected represented a broad and diverse range of scientific
perspectives relevant to the status review of the American eel coming
from State and Federal agencies, fishery commissions, Tribes, academia,
domestic and foreign research institutions (Canada, Japan, and
England), industry organizations, and nongovernmental organizations.
Participating individuals had expertise on threats or life history
characteristics associated with threats to the American eel.
Therefore, in addition to the published literature, our review
considered: (1) Each expert panelist's characterization of the threat
(the life stages acted upon by the threat, the severity of the threat,
and the timing of the threat) based on their own and other published
and unpublished research on the species; (2) the basis for each expert
panelist's assessments of the literature in the context of a rangewide
status review; and, (3) each expert panelist's assessments of the
implications of the uncertainty in the information. This finding
therefore builds on, clarifies, reinterprets, and, in some cases,
supersedes information presented in the 90-day finding.
In conducting our 12-month finding for American eel, we considered
all scientific and commercial information on the status of American eel
that we had in our files. Parallels in life history traits that are
unknown for the American eel are drawn from other species of Anguilla.
Evolution and Population Structure
The American eel is one of 15 ancient species (evolving circa 52
million years ago) of the worldwide genus Anguilla, whose members spawn
in ocean waters, migrate to coastal and inland continental waters to
grow, and then return to ocean spawning areas to reproduce and die--a
life history strategy known as catadromy (McCleave 2001a, p. 800; Avise
2003, p. 31; Knights et al. 2006, pp. 2-3).
The North Atlantic is home to two, closely related, recognized
species of Anguilla--the American eel and the European eel (A.
anguilla) (Avise 2003, p. 31). Genetic research indicates that the
American eel lacks appreciable phylogeographic population structure,
[[Page 4969]]
meaning that American eels are one, well-mixed, single breeding
population, termed panmixia or panmictic (Avise 2003, pp. 34-35). This
likely occurs from a combination of the random distribution of the
eel's larval stage when they reach continental waters and random mating
among all adults throughout the species' range. This is in contrast to
many anadromous species (which, even though they have an oceanic phase,
return to their rivers of origin to spawn), where mating is within
separate populations that are geographically or temporally isolated.
This panmictic life history strategy maximizes adaptability to
changing environments and is well suited to species that have
unpredictable larval dispersal to many habitats (Stearns 1977 in
Helfman et al. 1987, p. 52). Additionally, by not exhibiting geographic
or habitat-specific adaptations, eels have the ability to rapidly
colonize new habitats and to re-colonize disturbed ones over wide
geographical ranges (McDowall 1996 in Knights et al. 2006, p. 7).
Life History
In brief, the life history of the American eel begins in the
Sargasso Sea, where eggs hatch into a larval stage known as
``leptocephali.'' These leptocephali are transported by ocean currents
to the Atlantic coasts of North America and upper portions of South
America. They enter coastal waters, where they may stay, or they may
move into estuarine waters or migrate up freshwater rivers, where they
grow as juveniles and mature. Upon nearing sexual maturity, these eels
begin migration toward the Sargasso Sea, completing sexual maturation
en route. Spawning occurs in the Sargasso Sea. After spawning, the
adults die; a species with this life history trait is known as a
semelparous species. For a detailed description of the life cycle and
other life history characteristics, see McCleave 2001a, Tesch 2003, and
Cairns et al. 2005. Aspects of the species' life history most relevant
to this finding are discussed in more detail below.
Egg and Larval Life History Stage
The egg and larval stage of the American eel occur in the Atlantic
Ocean, the Sargasso Sea, ocean currents, and Continental Shelf waters.
Sargasso Sea. The Sargasso Sea is part of the North Atlantic Ocean,
lying roughly between the West Indies and the Azores. The Sargasso Sea
is part of the western half of a large clockwise gyre (circular pattern
of ocean circulation). It is here that American eel eggs hatch into a
larval stage known as ``leptocephali.'' The leptocephali are
distributed in the upper 300 meters (m) of the ocean and are subject to
transport from surface currents in the Sargasso Sea. These surface
currents can be complex due to the fronts that form in the Subtropical
Convergence Zone (where equatorial and temperate waters meet) primarily
in the winter and spring, and the eddies that are likely present year
round.
Ocean current transport. The Sargasso Sea includes a powerful
western boundary current, the Florida Current and Gulf Stream, which
flows to the north and northeast along the Atlantic coast of North
America. The Florida Current is the southern half of this flow, from
the Straits of Florida to Cape Hatteras (Schott et al. 1988 in Miller
2005, p. 3). The Florida Current transports water from the Caribbean,
Gulf of Mexico, and more distant regions through the Straits of
Florida. It then combines with Gulf Stream recirculation water from the
Sargasso Sea as it flows north of the Bahamas (Marchese 1999, pp. 29,
549), and forms the Gulf Stream off Cape Hatteras, North Carolina. Once
past Cape Hatteras, the Gulf Stream (which is at least 48 km or 30
miles offshore but more typically 160 km or 100 miles or greater
offshore) usually has pronounced meanders, which, if large enough, can
get separated and cast off to the north into the continental slope
water (a water mass found in the permanent thermocline between the Gulf
Stream and the continental shelf north of Cape Hatteras (35 [deg]N)).
The flow of the Gulf Stream continues to the northeast, mostly
paralleling the Atlantic coast, towards Europe and becomes the North
Atlantic Current (Miller 2005, pp. 3-4).
The majority of the leptocephali enter the Florida Current just
south of Cape Hatteras (just south of where the Florida Current enters
the Gulf Stream) directly from the Sargasso Sea. The remainder may
enter the Florida Current by a more southern route (e.g., transported
on the Caribbean Current through the Yucatan Straights (Kleckner and
McCleave 1985, p. 89), to the Gulf Loop Current and then to the Florida
Current, which would be the route most likely taken for Gulf of Mexico
recruitment) (Kleckner and McCleave 1982, p. 329-330; Miller 2005, p.
3).
The distribution of American eel leptocephali in the Florida
Current was first described by Kleckner and McCleave (1982, pp. 334-
337; 1985, pp. 73-77). Additionally, they found evidence of westward
movement of leptocephali across the current toward the coastal waters.
Because the distances of transport, to southern and northern points
along the Atlantic coast, differ by thousands of kilometers, it has
been suggested that the timing of metamorphosis from leptocephali to
the next life history stage may determine where individuals arrive in
Continental Shelf waters.
Other than likely current transport, we know very little about the
American eel leptocephali. Recent studies on other species have
indicated that leptocephali may feed on marine snow or specific
detrital particles, such as discarded larvacean (planktonic tunicates
that secrete a gelatinous house) houses and zooplankton fecal pellets
(Otake et al. 1993, pp. 28-32; Mochioka and Iwamizu 1996, p. 447).
Continental shelf waters. The American eel undergoes metamorphosis
twice. The first occurs when the leptocephali enter the Continental
Shelf waters (the area of shallow seas just off the coast to the area
of marked increase in slope to greater depths); the second is during
sexual maturation. The leptocephalis' leaf-like, laterally compressed
shape transforms during metamorphosis into a reduced,
characteristically eel-like shape, as they become transparent ``glass''
eels. Leptocephali are unusual fish larvae that are filled with a
transparent gelatinous energy storage material, and they can swim
either forwards or backwards (Miller and Tsukamoto 2004 in Miller 2005,
pp. 1-2); this may be an important aspect in detraining from (getting
off of) the Gulf Stream. According to Miller (2005), this directional
swimming appears to be the only way that leptocephali can cross and
detrain from the Gulf Stream system and cross the Continental Shelf
waters, due to the lack of any persistent oceanic transport mechanism
that can account for the large-scale transport of millions of larvae
across the current.
Juvenile Life History Stage
Arrival in coastal waters. When juvenile eels arrive in coastal
waters, they can arrive in great density and with considerable yearly
variation (ICES 2001, p. 2). Arriving juvenile eels (unpigmented
``glass eels'' and pigmented ``elvers'') have been collected and
recorded for 10 years from two sites in North Carolina in the Beaufort
estuary. Densities as high as 13.5-14.0 eels/100m\3\ and as low as 1.5
eels/100m\3\ have been recorded (Powles and Warlen 2002, p. 301). In
the East River, Canada, Jessop (2000, p. 520) had daily counts of
30,000 elvers entering the mouth of the river. Between May and August
200,000 elvers were recorded by trap method, and a population estimate
of 960,000 elvers was conducted by mark-
[[Page 4970]]
recapture (Jessop 2000, pp. 518-520). Variation in recruitment between
years can be quite significant. In the 9 years of records between the
years 1982 to 1999, estimated recruitment to the Petite rivi[egrave]re
del la Trinit[eacute] varied roughly four-fold, from a low of 14,014 to
a high of 61,308 (ICES 2001, p. 36). Some arrivals remain in brackish
(estuarine) or marine (salt) waters, others migrate up rivers to a
variety of fresh water habitats, and still others, as they mature, will
show inter-habitat movement patterns (Jessop et al. 2002, pp. 217-218;
Morrison et al. 2003, pp. 90-92; Cairns 2006a, p. 2; Thibault et al.
2005, p. 36; Lamson et al. 2006, p. 1567; Daverat et al. 2006, p. 2).
Juvenile mortality. Information on mortality rates for all of the
life stages is limited. In Jessop (2000, p. 514), the recruitment of
elvers to the East River, Chester, Nova Scotia, during May through July
was estimated by mark-recapture population estimates to be 960,000
elvers. The population size following migration to recapture sites
about 1.3 kilometers (km) upstream during late July-October was 2,894
elvers. These data indicate high juvenile mortality rates, in this case
at a rate of 99 percent. This high mortality was attributed to the
effects of low pH (4.7-5.0), high initial elver density (4.7 elvers/
m\2\) (which may lead to predation, including cannibalism, starvation,
and competition for space), and predation by resident, presumably
older, eels. V[oslash]llestad and Jonsson's (1988 in Jessop 2000, p.
523) research indicates that eel mortality in fresh waters is density-
dependent when elver numbers exceed a certain abundance. Although it is
not certain if early juvenile mortality is this high throughout the
range of the species, this supports the observation, according to
Jessop, that oceanic conditions may deliver relatively high quantities
of elvers to rivers, such as those along the south shore of Nova Scotia
(Jessop 1998 in Jessop 2000, p. 523), even to the point that elver
abundances too great for habitat capacity can occur (Jessop 2000, p.
523). Surviving juvenile eels mature into fully pigmented ``yellow
eels.''
Mortality rates likely decrease with size. One study in Prince
Edward Island, Canada, calculated loss from the population due to
mortality and emigration. Estimates of loss in American yellow eels
from the Prince Edward Island study are reported at 22 percent, with
mortality rates decreasing to 12 to 15 percent as the juvenile yellow
eels age (Anonymous 2001 in Morrison and Secor 2003, p. 1498), likely
due to lower mortality from predation and starvation as size increases.
Juvenile diet. The enormous dietary breadth of eels reflects their
great adaptability with respect to nearly all conditions of water
bodies. Yellow eels are opportunistic, consuming nearly any live prey
that can be captured. Smaller eels eat benthic invertebrates; larger
eels include mussels, fish, and even other eels in their diet. Yellow
eels also adapt to seasonal changes, decreasing intake or ceasing to
eat during the winter. Eels can also respond to local abundances of
appropriately sized prey through the seasons (Tesch 2003, pp. 152-163).
This adaptable diet allows for resource partitioning as well as the
ability to withstand changes in local environmental conditions and the
ability to occupy a geographically wide variety of habitats.
Density-dependent dispersion. As young eels begin to grow, density-
dependent competition promotes eels to disperse into less crowded areas
(Feunteun et al. 2003, pp. 201-204; Ibbotson et al. 2002 in Knights et
al. 2006, p. 10). Aggressive interactions at high density inhibit
feeding and growth, but stimulate dispersive swimming activity in
smaller eels (Knights 1987 in Knights et al. 2006, p. 10), the latter
likely as a defense against predation. As size differences in these
juveniles increase, cannibalism can also be an important cause of
mortality (Knights 1987 in Knights et al. 2006, p. 10). Density
dependent dispersion ensures wider distributions, further minimizing
intra-specific competition. Benefits of density dependent dispersion
include selection of optimal habitat productivity and temperature,
lower predation risks, rapid colonization or re-colonization of
habitats, and avoidance of inter-specific competition. Larger
individuals farther upstream tend to become more sedentary and occupy
territories, densities of eels decline, and females predominate
(Feunteun et al. 2003, p. 201).
Distribution clines. It has been suggested that there are
latitudinal clines in eel distribution related to river typologies. For
example, the American eel tends to extend farther inland in southerly
lowland drainages compared to distributions in the shorter and steeper
post-glacial stream systems in the Northeast (Jessop et al. 2004 in
Knights et al. 2006, p. 11). Smogor et al. (1995, p. 799) and Knights
(2001 in Knights et al. 2006, p. 8) have documented decreases in
densities with increasing distance from the Continental Shelf in a
predictable pattern, likely as a result of density dependant dispersion
and mortality due to predation. Although mean watershed densities
decrease by an order of magnitude with distance inland from the
Continental Shelf, mean biomass only declines by about 50 percent
because mean body weight and eel length increase (and hence relative
fecundity). This, according to Knights et al. (2006, p. 10), helps
maintain biomass relative to carrying capacity. Machut (2006, p. 13)
indicates that as barrier intensity increases, so does eel growth above
the barrier. Recent research (Knights et al. 2006, pp. 11-13) has
documented that as eel density decreases, the proportion of females
increases, which, assuming females are the limiting sex, would be,
according to Knights et al. (2006, p. 13), a compensatory mechanism
during times or in areas of low density.
Sexually Maturing Life History Stage
Sex determination. There are no morphologically differentiated sex
chromosomes in the American eel (McCleave 2001a, p. 803). Prior to
sexual differentiation, eels are intersexual, meaning they can develop
into either sex. It is only when yellow eels reach a length of about
20-35 cm that it is possible to distinguish males from females
visually, and there is considerable variation in age and size at
differentiation. The determination of sex is likely influenced by
environmental factors, including eel densities (Tesch 2003, pp. 43-46).
Studies indicate that as the density of eels in a particular area
increases the number of male eels increases; decreasing density favors
more females. It has been argued by Knights et al. (2006, p. 13), that
an advantage of this life history strategy is that when recruitment
declines, so will density and tendencies to migrate far upstream in
rivers. In turn, this will lead to relative increases in the number of
(larger) females and hence compensatory increases in fecundity. This
may take a number of generations (and hence decades) to manifest
itself, but this strategy confers enormous benefits in the face of
threats, past, present and future, such as tectonic events and changes
in ocean currents and climate (Knights et al. 2006, p. 13).
Silvering. After a number of years, the yellow eels begin
metamorphosis. Beginning at 3 years old and up to 24 years, with the
mean becoming greater with increasing latitude (e.g., 6-16 years in the
Chesapeake Bay region; Helfman et al. 1987, pp. 44-45; and 8-23 years
in Canada; Cairns et al. 2005, p. 11), yellow eels metamorphose into
``silver eels'' (Cairns et al. 2005, p. 13). This metamorphosis from
bottom-oriented yellow eels to silver eels (termed ``silvering'') is a
key physiological event
[[Page 4971]]
preparing these future spawners for oceanic migration and reproduction
(van den Thillart et al. 2005, p. 12).
Environmental factors may play a role in the triggering of
silvering. Habitat conditions, such as food availability and
temperature, will influence the size and age of silvering eels via
growth conditions. Thus, variation in length and age at maturity can
occur in different habitats (e.g., freshwater habitat versus estuarine
habitat) within a restricted geographic range and over larger
geographic scales as well.
The length of the growing season and the temperature are negatively
correlated with latitude, so age at maturity is strongly correlated
with latitude (McCleave 2001a, p. 803). Characteristics of silver eels
vary across the species' range. Eels from northern areas, where
migration distances are great, show slower growth and greater length,
weight, and age at migration, preparing them, it could be assumed, for
the longer migration.
Indeed, favorable growth conditions cause eels to silver more
rapidly (V[oslash]llestad and Jonsson1988 in Jessop 2000, p. 522;
V[oslash]llestad 1988 and 1992 in van den Thillart 2005, p. 56; De Leo
and Gatto 1995 in van den Thillart 2005, p. 56) such as is the case in
aquaculture, under experimental conditions (Tesch 1991 and Beullens et
al. 1997 in van den Thillart et al. 2005, p. 56), or in brackish water
and at low latitudes (Lee 1979 and Fernandez-Delgado et al. 1989 in van
den Thillart et al. 2005, p. 56). For example, Morrison et al. (2003,
p. 95-96) found annual growth rates in brackish water were two times
higher than growth rates of eels that resided entirely in fresh water.
Also American eels in U.S. southern Atlantic coast waters develop into
silver eels about 5 years sooner than northern populations (Hansen and
Eversole 1984, p. 4; Helfman et al. 1984, p. 139), likely as a result
of warmer, more stable water conditions (Helfman et al. 1984, p. 138).
Variation in maturation age benefits the population by allowing
different individuals of a given year class to reproduce over a period
of many years, which increases the changes of encountering
environmental conditions favorable to spawning success and offspring
survival. For example, variability in the maturation age of eels born
in 2006 may result in spawners throughout 2010-2030, during which time
favorable environmental conditions are likely to be encountered at
least once.
Additionally, males and females differ in the size at which they
begin to silver. Eels appear to need to reach a certain size to begin
the silvering process, with this size increasing with age (thus,
rapidly growing eels will silver at smaller sizes than slow-growing
eels). In males, silvering happens at a very early stage, at a size
typically greater than 35 centimeters (cm). In females, silvering
happens at a size greater than 40 to 50 cm (Goodwin and Angermeier
2003, p. 530; van den Thillart et al. 2005, pp. 31, 55).
Actual metamorphosis is a gradual process occurring during the
summer, and in the fall eels metamorphosing in preparation for
migration back to the spawning grounds have a silvery body color,
enlarged eyes and nostrils, and a more visible lateral line (Dave et
al. 1974; Lewander et al. 1974; Pankhurst 1983; and Barni et al. 1985
in van den Thillart 2005, p. 12). As the structure and metabolism of
the liver changes, the swim-bladder also changes, allowing for
increased gas deposition rates and decreased loss of gas (McCleave
2001a, p. 804).
A drop in temperature appears to trigger the final events of
metamorphosis (gut regression and cessation of feeding), which will
lead to migratory movements under the appropriate environmental
conditions. It is theorized that responding to a drop in temperature
would help to synchronize out-migrating eels, thus increasing their
chances of reaching the Sargasso Sea simultaneously. Conversely,
increasing temperatures, delays in migration, or possibly low fat
content will cause eels to start feeding again and to revert to a
yellow resident stage. This would happen in the natural environment if
eels did not reach the sea before the end of the migrating season. It
has been observed that even after eggs and sperm have developed, eels
are capable of gut regeneration and feeding (Fontaine et al. 1982,
Dollerup and Graver 1985, in van den Thillart et al. 2005, p. 56). Van
den Thillart et al. (2005, p. 56) confirmed that silvering may occur
more than once in the lifetime of an eel. It has been said that this
phenomenon would explain the extreme variability in age and size of
silver eels. It has been hypothesized that conditions encountered
during oceanic migration, such as the high pressure they would
experience at depth in the open ocean, may complete the sexual
maturation of eels (Fontaine et al. 1985 in van den Thillart et al.
2005, p. 13).
Outmigration Life History Stage
Energy requirements. To successfully complete the migration from
the continent to the Sargasso Sea (out-migration), great endurance and
an extensive fat reserve are required. Larger, fatter eels have an
advantage over smaller eels in reaching the Sargasso Sea and having
sufficient energy stores to reproduce. Eels are very efficient swimmers
(eels swim approximately four to six times more efficiently than
salmonids), and larger eels appear more efficient than smaller eels
(van den Thillart et al. 2005, pp. 106-107). Also, larger eels usually
have larger fat stores per body weight. Silver eels have ceased
feeding, and use their stored fat for energy during their migration and
for completing gonadal growth. In a study conducted on European eel,
the most recent estimate of necessary energy (fat) needed to
successfully complete the migration to the Sargasso Sea from Europe and
spawn is 20 percent fat reserves, of which 13 percent is for transport,
and an additional 7 percent for completing gonadal growth. In European
silver eel, about 50 percent of the eels studied had a fat percentage
of 20 percent (van den Thillart et al. 2004 in van den Thillart et al.
2005, p. 109).
It is unknown if American eels require 20 percent fat reserves.
American eels travel a shorter distance to reach the Sargasso Sea than
do European eels. Actual distances, routes, and depths of migration for
adult eels are unknown. Distances traveled by migrating silver American
eels likely vary from under 1,500 km to over 4,500 km, shorter than the
5,000 km to 7,000 km likely traveled by European eels. An American eel
maturing in the Mississippi River, Louisiana, would travel a distance
of over 2,200 km; from South Carolina, 1,440 km; from Chesapeake Bay,
Virginia, 1,550 km; from Newfoundland, Canada, over 2,800 km (McCleave
2001a, p. 805); and from western Lake Ontario, over 4,500 km. Silver
eels, it has been hypothesized by Knights (2003, p. 240), may follow
the deep currents (for American eel, the Deep Western Boundary Current)
to return to the Sargasso Sea. However, others believe the American eel
migrates in the upper portions of the ocean (see van Ginneken and Maes
2005, pp. 385-387; Tesch 2003, pp. 206-207).
Fecundity. Fecundity also varies with size. Fecundity increases
exponentially with length, ranging from about 0.6 million to almost 30
million eggs depending on the size of the female (McCleave 2001a, p.
804). As an example, in the lower Potomac watershed, the average silver
female length of 734 mm would produce 2.7 million eggs; farther up the
watershed the average silver female length of 870 mm would produce 5.2
million eggs (Goodwin and Angermeier 2003, p. 533). Fecundity is also
linked to the habitat which the eel occupies. In an eel
[[Page 4972]]
farm growth experiment, favorable nutrition was one of two factors (the
other being genetic heterozygosity, where 2 different alleles are at
one loci) producing eels with a high reproductive capacity (van den
Thillart et al. 2005, p. 232). This high fecundity is thought to
compensate for very high larval mortality (reported by Knights et al.
2006, p. 4, as most probably well in excess of 99 percent).
Spawning. Spawning takes place in the Sargasso Sea (Schmidt 1922 in
Bo[euml]tius and Harding 1985, p. 122). Here, in the area where
northern and southern waters meet, it has been hypothesized that there
is some unidentified feature of the surface water (perhaps the abrupt
horizontal temperate change of the frontal zone located within the
subtropical convergence) that serves as a cue for migrating adults to
cease migration and begin spawning (Kleckner et al. 1983, p. 289;
Kleckner and McCleave 1988, pp. 647-648; Tesch and Wegner 1991 in
Miller 2005, p. 1). Spawning has not been witnessed by humans, but the
assumption is that adult eels die after spawning.
Range
The extensive range of the American eel includes all accessible
river systems and coastal areas having access to the western North
Atlantic Ocean and to which oceanic currents would provide transport.
These drainages and coastal areas are along more than 50 degrees of
latitude (from 5[deg] to 63[deg]) of the western North Atlantic Ocean
coastline, from Northern Brazil/Venezuela to southern Greenland (Scott
and Crossman 1973, pp. 624-625; Tesch 2003, pp. 92-97; Helfman et al.
1987, p. 42), including most Caribbean Islands and Bermuda, the eastern
Gulf of Mexico and associated drainages including the extensive
Mississippi River watershed (e.g., Mississippi River, Ohio River,
Tennessee River, Arkansas River, and Missouri River) as far north as
Minnesota, the Gulf of St. Lawrence and the associated rivers, and Lake
Ontario and associated drainages. It is believed that the eel was
absent from the waters of Lakes Erie, Huron, and Superior before the
completion of the Welland Canal in 1829 (Patch 2006, p. 2). In 1878,
the Michigan Fish Commission planted young eels in southern Michigan
waters, and for more than a decade, beginning in 1882, the Ohio Fish
Commission released young eels throughout Ohio, including drainages to
Lake Erie (Trautman 1981, pp. 192-193) (Figure 1). This extensive range
should provide the American eel with a buffer against adverse
conditions, as spawners would still be coming from areas not
experiencing adverse conditions, and would, due to random dispersal and
relatively homogeneous genetic structure, be capable to successfully
re-colonize areas once the threat has abated.
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It has been reported in other documents that Bo[euml]tius and
Harding (1985) estimated that the American eel range covers more than
10,000 km of coastline; however, we could not locate this information.
Utilizing current mapping technology, our estimate of the available
coastline (including barrier islands) from Maine to Texas (Atlantic and
Gulf coast) is 29,612 km (Castiglione 2006, p. 1).
As a result of oceanic currents, the majority of the American eel
population is located along the Atlantic seaboard of the United States
and Canada. The historic and current distribution of the American eel
within its extensive continental range is well documented along the
United States and Canadian Atlantic coast, and the SLR/LO. The
distribution is less well documented and likely rarer, again due to
currents, in the Gulf of Mexico, Mississippi watershed, and Caribbean
Islands, and least understood in Central and South America.
Habitat
The American eel is said to have the broadest diversity of habitats
of any fish species (Helfman et al. 1987, p. 42) by occupying multiple
aquatic habitats. From an evolutionary standpoint, this generalist use
of habitats is favored in fluctuating environments, while specialists
excel under constant or slowly changing environmental conditions
(Richmond et al. 2005, pp. 279-280).
During their spawning and oceanic migrations, eels occupy
saltwater, and in their continental phase, they use all salinity zones:
Fresh, brackish, and marine (for detailed habitat use by life stage,
see Cairns et al. 2005). Eels occur in waters highly productive to fish
species and those that are not, and from waters of near tropical
temperatures to waters that are seasonally ice-covered (McCleave 2001a,
p. 800).
Growing eels are primarily benthic, utilizing substrate (rock,
sand, mud) and bottom debris such as snags and submerged vegetation for
protection and cover (Scott and Crossman 1973, p. 627; Tesch 2003, pp.
181-183). In Canadian waters, American eels hibernate in mud during the
winter. Wintering areas include fresh water, brackish estuaries, and
bays with full strength salt water (Cairns et al. 2005, p. 3.4.6).
Barring impassable natural or human-made barriers, eels occupy all
freshwater systems, including large rivers and their tributaries,
lakes, reservoirs, canals, farm ponds, and even subterranean springs.
The anquillid (eel-shaped) body form allows for climbing when at young
stages and under certain conditions (e.g., rough surfaces), allowing it
to pass up and over some barriers encountered during upstream
migrations in freshwater streams (Craig 2006, pp. 1-4). Eels are able
to survive out of water for an exceptionally long time (eels can meet
virtually all their oxygen needs through their skin), as long as they
are protected from drying (for which their ability to produce mucus is
of great adaptive significance), and eels have been seen using overland
routes (while moist) when they encounter a barrier, explaining their
entrance into landlocked waters (Tesch 2003, pp. 184-185) and their
presence above numerous dams and weirs (USFWS 2005b, pp. 16-18).
Abundance. Abundance (density) and distribution of eels within
habitats may be a function of distance from the ocean and may not be
related to habitat features (Smogor et al. 1995, pp. 796-797) (see also
Density-Dependant Dispersion). According to Smogor et al. (1995, p.
799) when examining Virginia streams, they found little connection
between habitat features and the distribution and abundance of American
eels at least at a large scale. Their results, they suggest,
demonstrate a diffusion pattern of eel occurrence. This lack of eel-
habitat relations (at least at a large scale) within freshwater systems
suggests that comparison of abundance for purposes of identifying
quality habitat would be misleading. Rather, it has been suggested
(USFWS 2006, pp. 13-14, 22) that the reproductive contribution of an
area to the total American eel population would be the best manner of
identifying quality habitat; however, reproductive contribution
estimates from throughout the range of the American eel are not
available. Examples of densities provided below are to illustrate the
variation of densities, not for comparison of habitat importance.
Machut (2006) summarized freshwater and brackish water density research
and standardized to eel densities per 100m\2\. In Lake Champlain,
Vermont, densities ranged from 2.32-6.36 eel/100m\2\ (LeBar and Facey
1983 in Machut 2006, p. 50). In a tidal creek, Georgia, densities
ranged from 1.82-2.32 eel/100m\2\ (Bozeman et al. 1985 in Machut 2006,
p. 50). A Massachusetts salt marsh yielded densities of 8.46-9.28/
100m\2\ (Ford and Mercer 1986 in Machut 2006, p. 50). In Machut's own
study in the Hudson River freshwater tributaries densities ranged from
0.28-155.06/100m\2\ (Machut 2006, p. 50), while in brackish waters
Morrison and Secor (2003 in Machut 2006, p. 50) reported densities of
0.03-0.24/100m\2\ . In four Maine freshwater rivers, densities ranged
from 1.80-35.40/100m\2\ (Oliveira and McCleave 2000, p. 144). Recent
population estimates of juvenile eels (mostly elvers) on the South Anna
River in Virginia were 1.88 eels/100m\2\. On the North Anna River,
where the eels were smaller, the population estimate was greater at
4.48/100m\2\ (Odenkirk 2006, p. 1). No estimates of abundance or
density are yet available for marine waters.
Habitat associations at a finer scale, such as areas within a lake,
have recently been researched by Cudney (2004). In her studies, she was
able to associate certain short-term habitat conditions, such as non-
stagnant waters and to a lesser extent long-term habitat features such
as water depth and percent organic matter, to a higher probability of
eel capture (Cudney 2004, pp. 57-60).
Facultative Catadromy. Contrary to the earlier dominating paradigm
that the eel growth phase is restricted to fresh water, it has been
suggested that brackish (or estuarine) waters produce eels that grow
faster, mature earlier, and emigrate as silver eels sooner than eels in
fresh water, and that some eels complete their life cycle in brackish
or marine waters without ever entering fresh water. Facultative
catadromy, therefore, refers to migrations into fresh water as not
being obligatory (Tsukamoto and Arai 2001, p. 2651).
Morrison et al. (2003, p. 94) found annual growth rates in brackish
water were two times higher than growth rates of eels that resided
entirely in fresh water. The mechanism for this higher growth in
brackish water is not well understood. Possible causes include an
increase in quality or quantity of food, increase in habitat quality
(Helfman et al. 1987 in Morrison et al. 2003, p. 94), lower resting
metabolism resulting from living in near-isoosmotic (same salinity
within the eel as the external environment) conditions, increased water
temperature (which reduces the amount of time that eels are dormant
during winter) (Walsh et al. 1983 in Morrison and Secor 2003, p. 1499),
reduced effects from parasites, decreased predation, or decreased
intra- or inter-specific competition. Morrison and Secor (2003, p.
1499) hypothesized that the higher brackish-water eel growth measured
on the Hudson River is general to most large North American estuaries.
Two other studies became available during our status review, which
provided data on use by eels of marine habitats during the eel growth
phase (Daverat et al. 2006; Lamson et al. 2006). The first study, by
Daverat et al. 2006,
[[Page 4975]]
looked at habitat plasticity in the American, European, and Japanese
eel (A. japonica;) the second, by Lamson et al. (2006), at American eel
in Canadian waters. In the first study, habitat use consisted of either
residency in one habitat (fresh, brackish, or marine) or movements
between habitats. Seasonal or minor (1 or 2) movement patterns were
seen from brackish water to fresh water and vice versa. Single habitat
switch events occurred, usually between 3 and 5 years of age.
``Nomadic'' movement between water masses of different salinity was
common; the differences in productivity between freshwater and brackish
habitats (and the resulting lower growth of eels in temperate
freshwater sites), the authors state, might explain this phenomenon.
Occurrence of eels with no freshwater experience was demonstrated, but
such eels accounted for a smaller proportion of the overall sample than
did eels with some (even brief) freshwater experience. Another
interesting result was that eels tend to prefer brackish and marine
habitats for feeding at the northern extremes of their range. The
authors also suggest that this high degree of habitat use plasticity
suggests a remarkable ``bet hedging'' strategy for angullids as a group
(Daverat et al. 2006, p. 11). In the second study, conducted on
American eels in Canada, marine (saltwater) resident eels were the
dominant migratory contingent of eels in saltwater bays (85 percent).
Resident eels were established in salt and freshwater habitats by the
year after their arrival in continental waters. Eels that shifted
between habitats increased their rate of inter-habitat shifting with
age. This study also showed that plasticity of habitat usage is the
norm among eels, and that the American eel life cycle can be completed
in marine waters (Lamson et al. 2006, p. 1572). A study of Japanese eel
found that estuarine (43 percent) and marine (40 percent) eels
contributed more spawners than did eels from freshwater areas (17
percent), with some seasonal differences. Additionally, the study noted
that eels from all three habitats began their marine spawning migration
at about the same time. The implication here is that eels from all
habitats can mix together during spawning migration and potentially
contribute to the next generation (Kotake et al. 2005, p. 220). In
Tsukamoto et al's evolutionary perspective, the authors hypothesize,
based on Inoue 2001, that molecular evidence might suggest that
catadromous Anguillidae come from deep-sea eels, with a migration loop
that extended to coastal waters and incidentally visited estuaries;
these eels may have eventually obtained a reproductive advantage
because of higher food availability in estuaries than in freshwater
(Tsukamoto et al. 2002 in Miller 2005, p. 2).
According to Lamson et al. (2006, p. 1568), [Eacute]deline and
[Eacute]lie (2004) reported that European glass eels have distinct
individual salinity preferences. This implies that young eels separate
into migratory contingents upon arrival on the coast, with salt-seeking
eels remaining in marine waters while fresh-seekers ascend into fresh
waters.
The benefits of facultative catadromy include resource
partitioning, by minimizing intra-specific competition between life
stages and cannibalism of young by adults. Additionally, there are
growth-temperature benefits, as shallow brackish and fresh waters
(especially still waters) will heat up faster in the spring and summer
than marine waters. Although not tested by any large-scale quantitative
distribution data, the effective reproductive contribution of brackish/
marine habitats may be substantial (Tsukamoto and Arai 2001, p. 275;
Jessop 2002, p. 228; Kotake et al. 2005, p. 220; Knights et al. 2006,
pp. 12-13; Cairns 2006a, p. 1). Densities may be relatively low in
coastal waters, but for European eel in England and Wales, Knights et
al. (2001 in Knights et al. 2006, p. 13) calculated that estuarine and
shallow coastal waters (estimated at 5,000 km\2\) exceed that of
freshwater (1,035 km\2\).
Clinal Variations. American eels show clinal variation (gradual
changes over a geographic area) in their growth rates and size at
maturity between the southern and northern portions of their range.
Although mostly a warm water species, Anguillids are eurythermal
(tolerant of a wide range of temperatures) and can survive extremes by
migratory and cryptic behaviors. Even so, growth seasons inevitably
shorten with increasing latitude. This produces clines as you move
north of slower growth rates and larger size at maturity, thus
retaining relative fecundity with increasing latitude (Knights et al.
2006, p. 6).
Population Status
Typically an evaluation of population status for a 12-month finding
would include a rangewide estimate of population size and information
on the demographic structure of the population and subpopulations as
well as population trend information in context with historical data,
and possibly an evaluation of the long-term viability of the current
population through a population viability analysis model.
No rangewide estimate of abundance exists for the American eel.
Information on demographic structure is lacking and difficult to
determine because the American eel is a single population (panmixia)
with individuals randomly spread over an extremely large and diverse
geographic range, with growth rates and sex ratios environmentally
dependent. Because of this unique life history, site-specific
information on eels must be evaluated in context with its significance
to the entire population. Determining population trends is challenging
because the relevant available data is limited to a few locations that
may or may not be representative of the species' range and little
information exists about key factors such as mortality and recruitment
which could be used to develop an assessment model. Furthermore, the
ability to make inferences about species' viability based on available
trend information is hampered without an overall estimate of eel
abundance. Despite these challenges we have determined the species
currently appears stable, as we explain below.
The Stock Assessment Committee of the ASMFC recently assessed the
``stock status'' of the American eel (ASMFC 2006a), and this assessment
was subsequently reviewed by an independent panel of scientists (ASMFC
2006b). The Stock Assessment Committee concluded that the status of the
stock is uncertain as a result of insufficient data. Their conclusion
was based on the review of nine indices, two were fisheries-dependent
and seven were fisheries-independent. Of these indices, one index shows
an upward trend over time, one shows no trend, and the remaining seven
show a downward trend (ASMFC 2006a, p. x). The committee hypothesized
that the indices exhibiting a downward trend suggest that the stock is
at or near documented low levels. The glass eel data from two Atlantic
Coast sites were not used, and the panelists who reviewed the stock
status felt that these indices were a valuable asset. These panelists
interpreted the absence of a declining trend in glass eel abundance in
either series over the last 14 to 15 years as the only positive
indicator that recruitment, at least to the glass eel stage to these
portions of the coast, had not declined in concert with some of the
yellow eel indices (ASMFC 2006b, p. 4). The ASMFC stock status
assessment has limited value in the 12-month finding because the
purpose of the ASMFC stock status assessment is to inform management of
the commercial
[[Page 4976]]
American eel fishery by determining allowable harvest, not to look
specifically at long-term viability of the species.
Recently Canada completed its review of the American eel status
within Canadian waters as part of the Committee on the Status of
Endangered Wildlife in Canada's (COSEWIC) review for possible listing
under their version of the Endangered Species Act, known as Species At
Risk Act (SARA). This review also was more similar to a stock status
assessment than a population viability analysis. They determined that
indicators of the status of the total Canadian component of this
species were not available. Their evaluation of the data (indices of
abundance in the upper SLR/LO declined by approximately 99 percent
since the 1970s and four out of five time series from the lower St.
Lawrence River and Gulf of St. Lawrence declined) led them to apply the
Special Concern designation (COSEWIC 2006, p. III). Because the COSEWIC
review focuses on the status of American eels in Canadian waters, the
report also discussed the ``rescue effect.'' In the hypothetical
scenario where the American eel became depleted or extirpated within
Canadian waters external components would ``rescue'' the species in
Canada. These external components refer to the young eels from the
Sargasso Sea that are from American eels whose parents originated from
U.S. waters, and experience random dispersal due to oceanic currents
which would continue to deposit leptocephali into Canadian waters
(COSEWIC 2006, p. 43).
Together, however, these reports provide a more recent presentation
of the individual data sets than was available in the stock status
report by the International Council for the Exploration of the Sea or
ICES (2001, pp. 51-52), which was the only stock assessment available
at the time of the 90-day finding published on July 6, 2005 (70 FR
38849). As a result of these factors, our assessment of the American
eel population status will utilize the available information to: (1)
Provide context of historical reports and current landings data as a
surrogate for absolute abundance estimates; (2) evaluate the data from
each different life stage and the significance of that life stage when
evaluating the population status of the species including trend data in
specific geographic areas and each area's significance to the
population status of the species; and (3) evaluate the data to
determine if there is a sustained downward trend in a location or
locations that would be considered representative of the entire range.
Together these will provide the basis for our assessment of whether the
species is currently being impacted by threats to the degree that the
American eel meets the definition of threatened or endangered. In
addition, in the 12-month finding we also take into account the
species' life history characteristics and compensatory mechanisms (see
Background and for further discussion).
(1) Historical and Current Information
Historically eels were a significant winter food source for Native
Americans (see Casselman 2003, for a compilation of prehistoric and
historic information from the United States and Canada) and later for
European settlers. However, qualitative rather than quantitative
information is all that is available from these early times. In the
early 1900s, records from commercial fisheries began to appear. For
example, weirs at Oneida Lake, Canada, caught 100 metric tons (220,000
pounds) annually of emigrating eels (Adams and Hankinson 1928 in
Casselman 2003, p. 260). Casselman cites the subsequent construction of
dams and canals, which restricted access to the lake as the reason for
its eventual extirpation from Lake Oneida. Given the size of the
harvest, Casselman concludes that recruitment immigration in the past
was much more extensive and probably much greater than in recent times.
Although the current status of American eels cannot be described in
absolute terms because rangewide estimates of abundance do not exist
(ASMFC 2006a, p. viii; ASMFC 2006b, pp. 3, 13), we provide below recent
ASMFC and COSEWIC landings data (long-term fishery independent indices
do not exist) that indicate that the order of magnitude of yellow and
silver phase eel abundance is probably in the many millions. In the
past decade, commercial fisheries in the United States and Canada have
landed approximately 800 metric tons (1.8 million pounds) of yellow and
silver phase American eels annually (ASMFC 2006a, p. 82). These
landings data provide a general sense of eel abundance if we make
assumptions about the size and relative proportion of eels that are
landed. Specific data on the size of eels harvested were not available,
but 45 cm was considered a reasonable estimate (Cairns 2006b, p. 1).
The average weight of American eels 45 cm long is 156 grams (g) (Cairns
2006b, p. 1), which indicates that 800 metric tons is equivalent to
over 5 million eels. Assuming a high capture efficiency of 25 percent
for the eel fisheries (Caron et al. 2003, p. 235) suggests that the
post-fishery abundance (i.e., 75 percent are not captured) of yellow
and silver phase eels is greater than 15 million within the areas
fished. Given that not all areas within the range of the eel are
fished, this number would represent a minimum. These calculations are
not intended to be used as a formal estimate of population size, but
simply to provide the context that large American eels, throughout
their range, likely number in the many millions.
(2) Trend Data From Different Life Stages and Locations
Trends in American eel abundance from fishery-independent indices
(e.g., data from surveys and research) varied among locations and life
stages during the past 10-25 years. Data from yellow eels (which may
include silver eels) and glass eels (and elvers) are presented below.
Yellow eel. Four indices (including Maritime rivers in Canada and a
standardized U.S. coastwide yellow eels abundance index) did not
exhibit trends (ASMFC 2006b, p. 3). Indices from freshwater and tidal
sites distributed from the mid-Atlantic region north to Canada and the
St. Lawrence River indicated a statistically significant declining
trend in yellow eel abundance at three sites. Two of these indices,
Lake Ontario and the Chesapeake Bay index, had strong and statistically
significant declining trends over the recent 1994 to 2004 time period,
with 10-year declines in the order of 50 percent in the Chesapeake Bay
index to 99 percent in the Lake Ontario indices (ASMFC 2006b, p. 3).
Smaller declines (15 percent) were reported in the St. Lawrence estuary
(COSEWIC 2006, p. vi). Recent data suggest that declines may have
ceased in some Canadian locations; but the positive trends in some
indicators for the Gulf of St. Lawrence are, the COSEWIC report states,
too short to provide strong evidence of an increasing trend (COSEWIC
2006, p. 58).
It should be mentioned that yellow eel indices may reflect local or
regional impacts, such as impacts from harvest or turbine mortality
(see Factors B and E for further discussion). Additionally, yellow eels
have not yet been subject to mortality that may occur during their
oceanic outmigration to the Sargasso Sea. Therefore, yellow eel indices
are not the best indicator for estimating annual reproductive success.
Evaluation of the Significance of Upper SLR/LO. The extreme decline
in eels migrating up to the upper SLR/LO, as tallied at the Moses-
Saunders eel ladder, has focused attention on the potential impact of
that decline to the overall status of the American eel;
[[Page 4977]]
however, COSEWIC states that a rigorous way to quantify this impact to
the overall population has yet to be developed (COSEWIC 2006, p. 35).
The suggestion is that the reproductive contribution to the overall
American eel population from the upper SLR/LO may be disproportionately
larger than from other freshwater portions of the range because the
American eels in the upper SLR/LO are almost exclusively female and
highly fecund (producing many eggs) due to their large size, and the
watershed is of considerable size. Two methods for estimating the
relative reproductive contribution were presented in the COSEWIC report
(2006, pp. 35-41), but both methods, they state, are based upon
questionable assumptions and large uncertainties that reduce confidence
in the results. Additionally, contributions from marine and estuarine
waters were not considered in the analysis. According to COSEWIC some
sources of uncertainty suggest that it is more probable that the
methods overestimate, rather than underestimate, the reproductive
contribution of the St. Lawrence River basin (COSEWIC 2006, p. 41).
Glass eels. Indices of glass eel recruitment at the only two U.S.
sites with long-term data (North Carolina and New Jersey) did not
exhibit a declining trend over the last 14-15 years (ASMFC 2006b, p.
4). Recruitment estimates into Canadian rivers are available for two
Nova Scotian sites. The East River, Sheet Harbour, abundance series is
the longest elver series available for the species. Annual recruitment
varied without any upward or downward trend from 0.1 to 0.5 million
elvers between 1989 and 1999 (Jessop 2003a in COSEWIC 2006, p. 28). In
the East River, Chester, the total run of elvers peaked at 1.7 million
in 2002. Since the overlap periods of the two series are strongly
correlated, a combined index of 13 years was interpreted in the COSEWIC
report. Elver recruitment showed inter-annual variability, but no
indication of decline between 1989 and 2002 (COSEWIC 2006, p. 28).
Glass eel counts, also called recruitment indices, are the best
measure we have to annual reproductive success (see section immediately
below).
(3) Evaluation of Trend Information
Of the available index data for the different American eel life
history stages, we have determined that glass eel indices best
represents the species status rangewide. Although we do not have glass
eel indices from the entire range, the random nature of the
leptochephali dispersal allows us to consider these data representative
of the reproductive success of the species. As described above, there
is not evidence of a sustained downward trend of these glass eel
indices; therefore, we conclude that the American eel is not undergoing
a sustained downward trend at a population level.
In summary, the best available scientific and commercial
information indicates that despite a population reduction over the past
century, eels remain very abundant and occupy diverse habitats over an
exceptionally broad geographic range. Because of the species' unique
life history traits areas which have experienced depletions may
experience a ``rescue effect'' allowing for continued occupation of
available areas without concern for genetic fitness. Trends in
abundance over recent decades var