Endangered and Threatened Wildlife and Plants; Final Listing Determinations for Two Distinct Population Segments of Atlantic Sturgeon (Acipenser oxyrinchus oxyrinchus, 5914-5982 [2012-1950]
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Federal Register / Vol. 77, No. 24 / Monday, February 6, 2012 / Rules and Regulations
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
Administration
50 CFR Part 224
[Docket No. 090219208–1762–02]
RIN 0648–XN50
Endangered and Threatened Wildlife
and Plants; Final Listing
Determinations for Two Distinct
Population Segments of Atlantic
Sturgeon (Acipenser oxyrinchus
oxyrinchus) in the Southeast
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce.
ACTION: Final rule.
AGENCY:
We, NMFS, issue a final
determination to list the Carolina and
South Atlantic distinct population
segments (DPSs) of Atlantic sturgeon
(Acipenser oxyrinchus oxyrinchus) as
endangered under the Endangered
Species Act (ESA) of 1973, as amended.
We have reviewed the status of the
species and conservation efforts being
made to protect the species, considered
public and peer review comments, and
we have made our determination that
the Carolina and South Atlantic DPSs
are in danger of extinction throughout
their ranges, and should be listed as
endangered, based on the best available
scientific and commercial data.
DATES: This final rule is effective April
6, 2012.
ADDRESSES: Assistant Regional
Administrator for Protected Resources,
NMFS, Southeast Regional Office, 263
13th Avenue South, St. Petersburg, FL
33701–5505.
FOR FURTHER INFORMATION CONTACT:
Kelly Shotts, NMFS, Southeast Regional
Office (727) 824–5312 or Lisa Manning,
NMFS, Office of Protected Resources
(301) 427–8466.
SUPPLEMENTARY INFORMATION:
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SUMMARY:
Background
We first identified Atlantic sturgeon
as a candidate species in 1991. On June
2, 1997, NMFS and U.S. Fish and
Wildlife Service (USFWS; collectively,
the Services) received a petition from
the Biodiversity Legal Foundation
requesting that we list Atlantic sturgeon
in the United States, where it continues
to exist, as threatened or endangered
and designate critical habitat within a
reasonable period of time following the
listing. A notice was published in the
Federal Register on October 17, 1997,
stating that the Services had determined
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substantial information existed
indicating the petitioned action may be
warranted (62 FR 54018). In 1998, after
completing a comprehensive status
review, the Services published a 12month determination in the Federal
Register announcing that listing was not
warranted at that time (63 FR 50187;
September 21, 1998). We retained
Atlantic sturgeon on the candidate
species list (and subsequently
transferred it to the Species of Concern
List (69 FR 19975; April 15, 2004)).
Concurrently, the Atlantic States Marine
Fisheries Commission (ASMFC)
completed Amendment 1 to the 1990
Atlantic Sturgeon Fishery Management
Plan (FMP) that imposed a 20- to 40year moratorium on all Atlantic
sturgeon fisheries until the Atlantic
Coast spawning stocks could be restored
to a level where 20 subsequent year
classes of adult females were protected
(ASMFC, 1998). In 1999, pursuant to
section 804(b) of the Atlantic Coastal
Fisheries Cooperative Management Act
(ACFCMA) (16 U.S.C. 5101 et seq.), we
followed this action by closing the
Exclusive Economic Zone (EEZ) to
Atlantic sturgeon retention. In 2003, we
sponsored a workshop in Raleigh, North
Carolina, with USFWS and ASMFC
entitled, ‘‘The Status and Management
of Atlantic Sturgeon,’’ to discuss the
status of sturgeon along the Atlantic
Coast and determine what obstacles, if
any, were impeding their recovery. The
workshop revealed mixed results in
regards to the status of Atlantic sturgeon
riverine populations, despite the
coastwide fishing moratorium. Some
populations seemed to be recovering
while others were declining. Bycatch
and habitat degradation were noted as
possible causes for continued
population declines.
Based on the information gathered
from the 2003 workshop on Atlantic
sturgeon, we decided that a new review
of Atlantic sturgeon status was needed
to determine if listing as threatened or
endangered under the ESA was
warranted. The Atlantic sturgeon status
review team (ASSRT), consisting of four
NMFS, four USFWS, and three U.S.
Geological Survey (USGS) biologists
prepared a draft status review report.
The draft report was then reviewed and
supplemented by eight state and
regional experts who provided their
individual expert opinions on the
scientific facts contained in the report
and provided additional information to
ensure the report provided the best
available data. Lastly, the report was
peer reviewed by six experts from
academia. A Notice of Availability of
the final status review report was
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published in the Federal Register on
April 3, 2007 (72 FR 15865). On October
6, 2009, we received a petition from the
Natural Resources Defense Council to
list Atlantic sturgeon as endangered
under the ESA. As an alternative, the
petitioner requested that the species be
delineated and listed as the five DPSs
described in the 2007 Atlantic sturgeon
status review report (ASSRT, 2007):
Gulf of Maine, New York Bight,
Chesapeake Bay, Carolina, and South
Atlantic DPSs, with the Gulf of Maine
and South Atlantic DPSs listed as
threatened, and the remaining three
DPSs listed as endangered. The
petitioner also requested that critical
habitat be designated for Atlantic
sturgeon under the ESA. We published
a Notice of 90-Day Finding on January
6, 2010 (75 FR 838), stating that the
petition presented substantial scientific
or commercial information indicating
that the petitioned actions may be
warranted. On October 6, 2010, we
published a proposed rule (75 FR
61904) to list the Carolina and South
Atlantic DPSs, the two DPSs that spawn
in the NMFS Southeast Region, as
endangered. We originally solicited
written public comments via email, fax,
and letter on the proposed listing rule
for 90 days and extended it for an
additional 30 days by public request.
We also accepted written and verbal
comments at two public hearings in
Wilmington, North Carolina, and
Atlanta, Georgia, in December 2010. A
separate proposed rule (75 FR 91872)
was published on October 6, 2010, for
the three DPSs of Atlantic sturgeon that
spawn in the NMFS Northeast Region.
Listing Determinations Under the
Endangered Species Act
We are responsible for determining
whether Atlantic sturgeon are
threatened or endangered under the
ESA (16 U.S.C. 1531 et seq.). To be
considered for listing under the ESA, a
group of organisms must constitute a
‘‘species,’’ which is defined in section 3
of the ESA to include ‘‘any subspecies
of fish or wildlife or plants, and any
distinct population segment of any
species of vertebrate fish or wildlife
which interbreeds when mature.’’ On
February 7, 1996, the Services adopted
a policy describing what constitutes a
DPS of a taxonomic species (61 FR
4722). The joint DPS policy identified
two elements that must be considered
when identifying a DPS: (1) The
discreteness of the population segment
in relation to the remainder of the
species (or subspecies) to which it
belongs; and (2) the significance of the
population segment to the remainder of
the species (or subspecies) to which it
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belongs. As stated in the joint DPS
policy, Congress expressed its
expectation that the Services would
exercise authority with regard to DPSs
sparingly and only when the biological
evidence indicates such action is
warranted.
Section 3 of the ESA defines an
endangered species as ‘‘any species
which is in danger of extinction
throughout all or a significant portion of
its range’’ and a threatened species as
one ‘‘which is likely to become an
endangered species within the
foreseeable future throughout all or a
significant portion of its range.’’ The
statute requires us to determine whether
any species is endangered or threatened
as a result of any one or a combination
of the following five factors: (A) The
present or threatened destruction,
modification, or curtailment of its
habitat or range; (B) overutilization for
commercial, recreational, scientific, or
educational purposes; (C) disease or
predation; (D) the inadequacy of
existing regulatory mechanisms; or (E)
other natural or manmade factors
affecting its continued existence
(section 4(a)(1)(A)(E)). Section 4(b)(1)(A)
of the ESA requires us to make listing
determinations based solely on the best
scientific and commercial data available
after conducting a review of the status
of the species and after taking into
account efforts being made to protect
the species. Accordingly, we have
followed a stepwise approach in making
our listing determination for Atlantic
sturgeon. Considering biological
evidence, such as the separation
between river populations during
spawning and the possibility of multiple
distinct interbreeding Atlantic sturgeon
populations, we evaluated whether
Atlantic sturgeon population segments
met the DPS Policy criteria. We then
determined the status of each DPS (each
‘‘species’’) and identified the factors and
threats contributing to their status per
section 4(a)(1) of the ESA. Finally, we
assessed efforts being made to protect
the species, determining if these efforts
are adequate to mitigate impacts and
threats to the species’ statuses. We
evaluated ongoing conservation efforts
using the criteria outlined in the Policy
for Evaluating Conservation Efforts
(PECE; 68 FR 15100; March 28, 2003) to
determine their certainties of
implementation and effectiveness.
Finally, section 4(b)(1)(B) of the ESA
requires us to give consideration to
species which: (1) Have been designated
as requiring protection from
unrestricted commerce by any foreign
nation or pursuant to an international
agreement; or (2) have been identified as
in danger of extinction, or likely to
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become so within the foreseeable future,
by any state agency or by any agency of
a foreign nation.
Peer Review and Public Comments
In December 2004, the Office of
Management and Budget (OMB) issued
a Final Information Quality Bulletin for
Peer Review establishing minimum
standards for peer review. Similarly, a
joint NMFS/FWS policy (59 FR 34270;
July 1, 1994) requires us to solicit
independent expert review from at least
three qualified specialists. We solicited
peer review comments on the proposed
listing rule from three peer reviewers,
two from academia and one from a
Federal resource agency, with expertise
on Atlantic sturgeon. Written public
comments were received from 59
commenters and 7 commenters
provided verbal comments at the public
hearings. Peer review comments are
treated in the next section. In the
following sections of the document, the
public comments are categorized in the
following areas: (1) The delineation of
DPSs; (2) abundance and trends; (3)
differences between the proposed rule
and the conclusions in the 2007 and
1998 status reviews; (4) the need to list
Atlantic sturgeon under the ESA and
consequences of listing; (5) the analysis
of threats (habitat modification and
destruction, overutilization, disease and
predation, the inadequacy of regulatory
mechanisms, other natural and
manmade factors); (6) recovery; (7)
critical habitat; and (8) adequacy of the
public hearing. Many comments were
complex and had multiple inferences,
and thus individual statements are
addressed in multiple comments and
responses below. Information and data
provided by commenters supported or
did not conflict with our findings for the
Carolina and South Atlantic DPSs. Some
information submitted by commenters
as ‘‘new’’ information was information
already included and evaluated in our
proposed listing rule determination.
Some commenters asked us to consider
information, such as increased
compliance responsibilities and
economic costs on agencies and the
public, that the ESA and its
implementing regulations prohibit us
from considering in making listing
determinations. Many commenters
stated that NMFS should postpone a
listing determination until the results of
recent research are available, further
research can be undertaken, state and
Federal moratoria on the harvest and
possession of Atlantic sturgeon have
been in effect for the full planned
duration, and/or until non-listing
alternatives (e.g., entering into multiagency partnerships and expanding
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existing programs) have been explored.
Because we were petitioned to list the
Atlantic sturgeon, we cannot delay an
assessment of the status of Atlantic
sturgeon. We were required to evaluate
the status of the species and the threats
it is currently facing and make a finding
on whether the petitioned action was
warranted within 12 months, which
resulted in our proposed listing
determination of endangered for the
Carolina and South Atlantic DPSs of
Atlantic sturgeon. We believe the
current body of information on the
declines of Atlantic sturgeon, the failure
of their population numbers to rebound
despite harvest prohibitions, and the
ongoing impacts from bycatch, habitat
modification, and the inadequacy of
existing regulatory mechanisms or
protective efforts to control or mitigate
for these impacts, warrant listing the
Carolina and South Atlantic DPSs as
endangered. The information provided
in the peer review and public comments
did not provide a basis for revising our
evaluation of the status of Atlantic
sturgeon, the nature and significance of
the threats and impacts they face, or our
listing determinations. In the following
sections of the document, we
summarize the comments pertaining to
the proposed listing rule for the
Carolina and South Atlantic DPSs and
provide our responses to those
comments. Complete copies of the peer
review comments, the written public
comments, and transcripts of the public
hearings are available on the Internet at
www.regulations.gov.
Peer Review Comments
In this section, we refer to peer
reviewers 1, 2, and 3, which correspond
to the way the peer reviewers are
identified on https://www.
regulations.gov.
Comment 1 (definitions of endangered
and threatened): Two of the three peer
reviewers disagreed, all or in part, with
our proposed listing of the Carolina and
South Atlantic DPSs as endangered.
Each peer reviewer provided their own
definitions of endangered and
threatened.
Peer reviewer 1 believed that a DPS
warranted an endangered listing only if
no single historical spawning river
within the DPS sustained an abundant
and regularly reproducing Atlantic
sturgeon population. Peer reviewer 1
stated that no substantive biological
justification or new evidence is
presented in the proposed listing of the
Carolina and South Atlantic DPSs as
endangered to change the conclusions
presented in the 2007 status review,
which concluded that the Carolina DPS
should be listed as threatened and made
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no conclusion with regard to the South
Atlantic DPS due to lack of information
to allow a full assessment of
subpopulations within the DPS. Peer
reviewer 1 stated that an endangered
listing would be appropriate if no single
historical spawning river within that
DPS appeared to sustain both a
relatively abundant and simultaneously
regularly-reproducing Atlantic sturgeon
population.
Based on the available information on
abundance, reproduction, and the
presence of early life history stages, the
reviewer stated that the Carolina DPS
comes closest to conforming to the
standard of an endangered species. The
reviewer cited data from the proposed
listing rule that two of the original three
major spawning populations (the
Roanoke and Santee-Cooper
populations) in the Carolina DPS appear
to remain functional, and not
particularly vulnerable to extinction.
The reviewer also stated the proposed
listing of the South Atlantic DPS did not
appear to be supported by the best
available scientific information, since
there is evidence of at least one viable,
reproducing, and increasing Atlantic
sturgeon population in the South
Atlantic DPS, the Altamaha River
population (Schueller and Peterson,
2006, 2010). The reviewer further cited
both the Savannah River and the ACE
(Ashepoo, Combahee, and Edisto Rivers)
Basin systems as appearing to support
reproducing Atlantic sturgeon
populations, and stated Atlantic
sturgeon appear to be abundant in the
ACE system. The reviewer questioned
whether the remaining South Atlantic
DPS river populations in the smaller
and less well-studied Ogeechee and
Satilla rivers together constitute a
significant portion of the species’ range
over which extinction is probable in
order to justify an ‘‘endangered’’
designation for the entire DPS. The
reviewer noted that the 2007 status
review report deferred from such a
designation and that it appears the
South Atlantic DPS does not closely
conform to the standard of being
endangered. Based on the available
scientific evidence concerning
population size and reproduction in the
historically most important populations,
the resilience of sturgeons to
extirpation, and their capacity for repopulation from small effective
population size, the reviewer believed
the appropriate ESA designation for
both DPSs would seem to be threatened.
The reviewer suggested that the
threatened status would provide
protection for the species from direct
take of any kind and a basis for habitat
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restoration, while providing greater
flexibility for scientific sampling, tissue
analyses, and experimental
manipulation than would endangered
status. The reviewer stated the
downside is that threatened status
would provide a lower level of legal
leverage relative to the larger industrial
impacts, e.g., dams and bycatch, either
of which may represent an
insurmountable impasse to sturgeon
recovery. The reviewer offered that
under existing direct harvest
prohibitions, threatened status has
worked effectively for Gulf sturgeon
recovery in rivers where dams and
bycatch are not significant issues. It has
not worked effectively where dams and
bycatch are significant issues (e.g., the
Pearl, Pascagoula, and Apalachicola
rivers), although none of those
populations seem in danger of
extinction.
Peer reviewer 2 stated that implicit in
the definition of ‘‘endangered’’ is that
the species must be on a significant
downward trend, or at least there is
cause to believe that such a trend is
happening now, or will happen soon,
and concluded that is not the case on
the Altamaha River in Georgia.
However, this reviewer also commented
that every single Atlantic sturgeon
population has been decimated by
overfishing and habitat degradation and
that we have very little quantified
evidence that the species as a whole has
recovered, despite 14 years of the
protection afforded under the current
moratorium on harvest and possession.
Peer reviewer 2 recommended that a
‘‘threatened’’ listing would seem
appropriate for almost every Atlantic
coast river, including the St. Marys,
Satilla, Ogeechee, and Savannah Rivers
in Georgia, with the Altamaha being the
one exception, and an endangered
listing would be difficult to support.
Response: We must rely on the
definition of ‘‘endangered’’ and
‘‘threatened’’ species provided in
section 3 of the ESA, the implementing
regulations, and case law in applying
the definitions to marine and
anadromous species. Section 3 of the
ESA defines an endangered species as
one that is in danger of extinction
throughout all or a significant portion of
its range, and a threatened species as
one that is likely to become endangered
within the foreseeable future. Recent
case law (In Re Polar Bear Endangered
Species Act Listing and § 4(d) Rule
Litigation, D.D.C. WL 2601604 (June 30,
2011 Order); 748 F.Supp.2d 19 (D.D.C.
2010)) regarding USFWS’s listing of the
polar bear as threatened provides a
thorough discussion of the ESA’s
definitions and the Services’ broad
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discretion to determine on a case by
case basis whether a species is in danger
of extinction. Upon listing the polar
bear as threatened, USFWS’s rule was
challenged by a number of parties who
claimed that the polar bear was in
danger of extinction and should have
been listed as endangered, and by others
who conversely argued that the bear did
not warrant listing even as threatened.
The Court determined that neither the
ESA nor its legislative history compels
the interpretation of ‘‘endangered’’ as a
species being in ‘‘imminent’’ risk of
extinction, finding instead that the
phrase ‘‘in danger of extinction’’ is
ambiguous. The Court held that there is
a temporal distinction between
endangered and threatened species in
terms of the proximity of the ‘‘danger’’
of extinction, noting that the definition
of ‘‘endangered species’’ is phrased in
the present tense, whereas a threatened
species is ‘‘likely to become’’ so in the
future. Thus, in the context of the ESA,
the Services interpret an ‘‘endangered
species’’ to be one that is presently at
risk of extinction. A ‘‘threatened
species,’’ on the other hand, is not
currently at risk of extinction, but is
likely to become so. In other words, a
key statutory difference between a
threatened and endangered species is
the timing of when a species may be in
danger of extinction, either now
(endangered) or in the foreseeable future
(threatened). The Court concluded,
however, that the distinction is not
based ‘‘solely and unambiguously’’ on
the imminence of the species’
anticipated extinction,’’ and that
Congress delegated responsibility to the
Services to determine whether a species
is presently ‘in danger of extinction’ in
light of the five ESA section 4(a)(1)
factors and the best available science for
that species. The Court ruled that
although imminence of harm is clearly
one factor that the Services weigh in
their decision-making process, it is not
necessarily a limiting factor. In many
cases, the Services might appropriately
find that the imminence of a particular
threat is the dispositive factor that
warrants listing a species as ‘threatened’
rather than ‘endangered,’ or vice versa.
The Services have broad discretion to
decide that other factors outweigh the
imminence of the threat. In conclusion,
the Court confirmed that the Services
have flexibility to determine
‘‘endangerment’’ on a case-by-case basis.
Congress did not intend to make any
single factor controlling when drawing
the distinction between endangered and
threatened species, nor did it seek to
limit the applicability of the endangered
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category to only those species facing
imminent extinction.
Thus, contrary to the peer reviewers’
comments, there is no per se
requirement that a species be
experiencing current or imminent
significant downward trends, or that
there are no single historical spawning
river populations within the DPSs that
are relatively abundant and
simultaneously regularly-reproducing,
in order to be listed as endangered (we
discuss the status and data on the
Altamaha River population in more
detail in Comment 2 below). Our
determination of endangerment for the
Carolina and South Atlantic DPSs is
based on the exercise of our expert
professional judgment on the basis of
the best available information for each
DPS. In addition, we agree with the
USFWS’ judgment, discussed in its
supplemental explanation filed in the
polar bear litigation, that to be listed as
endangered does not require that
extinction be certain or probable, and
that it is possible for a species validly
listed as ‘‘endangered’’ to actually
persist indefinitely.
We determined that the Carolina and
South Atlantic DPSs of Atlantic
sturgeon are currently in danger of
extinction throughout their ranges, on
the basis of precipitous declines to
population sizes, the protracted period
in which sturgeon populations have
been depressed, the limited amount of
current spawning, and the impacts and
threats that have and will continue to
prevent population recovery.
Populations of Atlantic sturgeon
declined precipitously decades ago due
to directed commercial fishing. The
failure of Atlantic sturgeon numbers
within the Carolina and South Atlantic
DPSs to rebound even after the
moratorium on directed fishing was
established in 1998 indicates that
impacts and threats from limits on
habitat for spawning and development,
habitat alteration, and bycatch are
responsible for the risk of extinction
faced by both DPSs. In addition, the
persistence of these impacts and threats
points to the inadequacy of existing
regulatory mechanisms to address and
reduce habitat alterations and bycatch.
As described in the proposed listing
rule, the Carolina DPS is estimated to
number less than 3 percent of its
historical population size; the South
Atlantic DPS is estimated to number
less than 6 percent of its historical
population size, with all river
populations except the Altamaha
estimated to be less than 1 percent of
historical abundance. There are an
estimated 343 adults that spawn
annually in the Altamaha River and less
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than 300 adults spawning annually
(total of both sexes) in the river systems
where spawning still occurs for each
DPS (not all of the river systems
occupied by the two Southeast DPSs
currently support spawning, or effective
spawning leading to recruitment).
In light of threats and impacts, the
low population numbers of every river
population in the Carolina and South
Atlantic DPSs suggests that the DPSs are
currently in danger of extinction
throughout their ranges; none of the
populations are large or stable enough to
alone or in combination provide any
level of certainty for continued
existence of either DPS, and thus, the
peer reviewer’s suggestion that these
DPSs may not be endangered rangewide
or in a significant portion of their ranges
is erroneous. While the directed fishery
that originally drastically reduced the
numbers of Atlantic sturgeon has been
closed, recovery of depleted populations
is an inherently slow process for a latematuring species such as Atlantic
sturgeon, and they continue to face a
variety of other threats that contribute to
their risk of extinction. Their late age at
maturity (5 to 19 years in the Southeast)
provides more opportunities for
individual Atlantic sturgeon to be
removed from the population before
reproducing. While a long life-span also
allows multiple opportunities to
contribute to future generations, it
increases the timeframe over which
exposure to the multitude of threats
facing the Carolina and South Atlantic
DPS can occur.
Based on available information, we
determined that to be viable, the
Carolina and South Atlantic DPSs
require multiple stable riverine
populations, and we have added
discussion to the final determination to
better explain our reasoning. The
importance of having multiple stable
riverine spawning populations within
each DPS and the need to maintain
suitable habitat to support the various
life functions (spawning, feeding,
growth) of Atlantic sturgeon is best
understood by looking at the concept of
metapopulations. Each DPS, made up of
multiple river populations, is analogous
to a metapopulation, which is a
‘‘population of populations’’ (Levins,
1969), a group of spatially separated
populations of the same species which
interact at some level. Separation into
metapopulations is expected by
sturgeon and other anadromous fishes.
While recolonization of northern rivers
following post-Pleistocene deglaciation
likely occurred following a steppingstone sequential model (Waldman et al.,
2002), genetic analyses reveal that
currently, there are very low rates of
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exchange between river populations.
The amount and effectiveness of
movement separates a metapopulation
from a single large, patchy population.
Low rates of connectivity through
dispersal, with little to no effective
movement, allow individual
populations to remain distinct as the
rate of migration between local
populations is low enough not to have
an impact on local dynamics or
evolutionary lineages and distinguishes
a metapopulation from a patchy
population (Harrison 1994).
Metapopulation persistence depends
on the balance of extinction and
colonization in a static environment
(Hanski 1996). If habitat remains
suitable following local extirpation,
recolonization via immigrants into nowempty habitat may replace at least some
of those losses (Thomas, 1994).
However, if the cause of extinction is a
deterministic population response to
unsuitable conditions (e.g., lack of
suitable spawning habitat, poor water
quality, or disturbance of substrates
through repeated dredging), the local
habitat is likely to remain unsuitable
after extinction and be unavailable for
effective recolonization (Thomas, 1994).
Therefore, recolonization is dependent
upon both immigration from adjacent,
healthy populations and habitat
suitability. Because the DPSs are groups
of populations, the stability, viability,
and persistence of individual
populations affects the persistence and
viability of the larger DPS. The loss of
any population within a DPS will result
in: (1) A long-term gap in the range of
the DPS that is unlikely to be
recolonized, or recolonized only very
slowly; (2) loss of reproducing
individuals; (3) loss of genetic
biodiversity; (4) potential loss of unique
haplotypes; (5) potential loss of adaptive
traits; and (6) reduction in total number.
The loss of a population will negatively
impact the persistence and viability of
the DPS as a whole as fewer than two
individuals per generation currently
spawn outside their natal rivers (Wirgin
et al., 2000; King et al., 2001; Waldman
et al., 2002).
The persistence of individual
populations, and in turn the DPS,
depends on successful spawning and
rearing within the freshwater habitat,
the immigration into marine habitats to
grow, and then the return of adults to
natal rivers to spawn. Information on
Atlantic sturgeon spawning within the
Carolina and South Carolina DPSs is
limited. In the proposed listing rule, we
presumed spawning was occurring if
young-of-the-year (YOY) were observed
or mature adults were present in
freshwater portions of the system.
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Within the Carolina DPS, we concluded
that spawning is occurring, or occurred
in the recent past, in the following
rivers based on these data:
1. Roanoke River—collection of 15
YOY (1997–1998); single YOY (2005).
2. Tar and Neuse Rivers—one YOY
(2005).
3. Cape Fear—upstream migration of
adults in the fall, carcass of ripe female
upstream in mid-September.
4. Winyah Bay—running ripe male in
Great Pee Dee River (2003).
Within the South Atlantic DPS, we
concluded that spawning is occurring,
or has occurred in the recent past, in the
following rivers based on these data:
1. ACE Basin—1,331 YOY (1994–
2001); gravid female and running ripe
male in the Edisto (1997); 39 spawning
adults (1998).
2. Savannah River—22 YOY (1999–
2006); running ripe male (1997).
3. Ogeechee River—age-1 captures,
but high inter-annual variability (1991–
1998); 17 YOY (2003); 9 YOY (2004).
4. Altamaha River—74 captured/308
estimated spawning adults (2004); 139
captured/378 estimated spawning adults
(2005).
5. Satilla River—4 YOY and spawning
adults (1995–1996).
These data indicate that spawning
occurs within the Carolina and South
Atlantic DPSs; they do not indicate the
frequency of annual spawning events or
the degree to which spawning in these
systems leads to population growth,
persistence, or viability. The extent and
effectiveness of spawning events is
unknown and likely precarious in many
rivers, given ongoing threats that limit
population size and spawning success,
such as water quality and restricted
access to upstream spawning areas (75
FR 61904). Peer reviewer 1 stated that
data from the proposed listing rule
indicate the spawning populations in
the Santee-Cooper system appear to
remain functional and not particularly
vulnerable to extinction; however, in
the proposed listing rule, we noted our
determination that spawning may occur
in the Santee and/or the Cooper Rivers,
but it may not result in successful
recruitment. Lack of access to historical
spawning habitat due to dams restricts
spawning to areas just below the dam.
The proximity of these spawning areas
to salt water may result in very high
mortality to any larvae spawned in
those systems.
In addition to spawning success, it is
difficult to quantify spawning potential
within the two DPSs, given the lack of
population estimates. Currently, the
number of Atlantic sturgeon in the
Carolina DPS is estimated to be
3 percent of historical population size
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and the South Atlantic DPS is estimated
to be 1 percent of historical population
size, with the exception of the Altamaha
River population, estimated to be at 6
percent of historical population size.
Although the largest impact that caused
the precipitous decline of the species
has been curtailed (directed fishing), the
population size has remained relatively
constant at these greatly reduced levels
for approximately 100 years.
In response to comments about
divergence from the status review
report’s listing conclusions for the
Carolina and South Atlantic DPSs,
NMFS’ Protected Resources Divisions
have the responsibility to make listing
recommendations to the Assistant
Administrator. Status review reports are
an important part of the information
base for such recommendations, but
NMFS must independently review the
information in status review reports and
apply the ESA’s listing determination
requirements in accordance with
regulations, case law, and agency
guidance. The Atlantic Sturgeon Status
Review Report states that ‘‘risks of
extinction assessments are performed to
help summarize the status of the
species, and do not represent a decision
by the Status Review Team on whether
the species should be proposed for
listing as endangered or threatened
under the ESA’’ (page 106; ASSRT,
2007). Subsequent to the status review
report, we conducted a comprehensive
assessment of the combined impact of
the five ESA section 4(a)(1) factors
across the Carolina and South Atlantic
DPSs in classifying extinction risk for
each DPS. We focused on evaluating
whether the DPSs are presently in
danger of extinction or the danger of
extinction is likely to develop in the
future. In our proposed rules to list 5
DPSs of Atlantic sturgeon, we
determined that each DPS was at greater
risk of extinction than determined by
the 2007 ASSRT. While the ASSRT did
discuss and consider how multiple
threats might act in concert on a given
subpopulation, they ultimately
classified extinction risk using the
highest single threat score on an
individual population within a DPS, or
within what they considered to be a
significant portion of a DPS’s range
(pages 108–109; ASSRT, 2007). We
evaluated the overall stability and
viability of the DPSs as a whole based
on the combined statuses of the
component river populations and the
impacts of threats and impacts across
the DPS, when determining extinction
risk of each DPS, because, as discussed
above, the Carolina and South Atlantic
DPSs require multiple stable river
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populations. In addition, because of the
lapse in time between the development
of the status review report (ASSRT,
2007) and the publication of the
proposed listing rule (75 FR 61904,
October 6, 2010), new information on
bycatch (ASMFC, 2007) and water
quality (USEPA, 2008), as well as
climate change (IPCC, 2008) and
drought (e.g., USGS, 2007), became
available to us, and we incorporated this
information into our listing
determinations.
Since publication of the proposed
rules, a Federal District Court has
thoroughly reviewed and considered the
distinction between the definitions of
threatened and endangered species in
the ESA, explained by the USFWS in
litigation challenging their
determination to list the polar bear as
threatened and not endangered, as
discussed above (In re. Polar Bear
Endangered Species Act Litigation).
Prompted by this decision and the
comments received by the Services
requesting further explanation of the
divergence of our proposed listing
statuses and the conclusions of the
ASSRT, we have reviewed our
determinations and concluded that all
the proposed listings of specific DPSs as
‘‘threatened species’’ or ‘‘endangered
species,’’ respectively, satisfy the
requirements of the relevant ESA
definition. Thus, we have not changed
these classifications in the final rules.
We found that the Carolina and South
Atlantic DPSs are presently in danger of
extinction, and thus, listing them as
endangered is warranted.
As discussed above, because a DPS is
a group of populations (a
metapopulation), the stability, viability,
and persistence of individual
populations affects the persistence and
viability of the larger DPS. The
persistence of individual populations,
and in turn the DPS, depends on
successful spawning and rearing within
the freshwater habitat, the immigration
into marine habitats to grow, and then
the return of adults to natal rivers to
spawn. While the directed fishery that
originally drastically reduced the
numbers of Atlantic sturgeon has been
closed, modification and curtailment of
Atlantic sturgeon habitat resulting from
dams, dredging, and degraded water
quality are inhibiting spawning and
population rebounding throughout both
DPSs, and contributing to their
endangered statuses. Existing water
allocation issues will likely be
compounded by human population
growth and potentially by climate
change as well. Climate change is
predicted to elevate water temperatures
and exacerbate nutrient-loading,
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pollution inputs, and lower dissolved
oxygen (DO), all of which are currently
negatively impacting the Carolina and
South Atlantic DPSs. Continued
overutilization of Atlantic sturgeon from
bycatch in multiple commercial
fisheries in both their marine and
freshwater habitats is another ongoing
impact to the Carolina and South
Atlantic DPSs that is contributing to
their endangered status. Atlantic
sturgeon taken as bycatch may suffer
immediate mortality. In addition, stress
or injury to Atlantic sturgeon taken as
bycatch but released alive may result in
increased susceptibility to other threats,
such as poor water quality (e.g.,
exposure to toxins and low DO). This
may result in reduced ability to perform
major life functions, such as foraging
and spawning, or may even result in
post-capture mortality. Several of the
river populations in the South Atlantic
DPS (e.g., the Ogeechee and the Satilla)
are stressed to the degree that any level
of bycatch could have an adverse impact
on the status of the DPS (ASSRT, 2007).
The Carolina and South Atlantic DPSs
are in danger of extinction now, due to
precipitous declines from historical
abundances to population sizes that are
low and potentially unstable throughout
the DPSs. As discussed above, both
DPSs exhibit sporadic spawning with
uncertain effectiveness. Population
rebuilding and recovery in both DPSs is
being inhibited by impacts due to
habitat curtailment and degradation,
and due to capture as bycatch in
commercial fisheries. The current low
levels of abundance noted previously in
combination with the high degree of
threat to the two Southeast DPSs put
them in danger of extinction throughout
their ranges; none of the populations
making up the DPSs are large or stable
enough to provide any level of certainty
for continued existence of either DPS.
Regarding the conclusion that the
Carolina and South Atlantic DPSs
should be listed as threatened, peer
reviewer 1 incorrectly stated that listing
as threatened provides protection from
direct take of any kind. The ESA’s
prohibition against take contained in
section 9 only applies to endangered
species, unless a section 4(d) rule is in
place to extend the take prohibition to
a threatened species. If we determine
that the Carolina and South Carolina
DPSs meet the ESA’s definition of
endangered, then we cannot list the
species as threatened for the purposes of
providing flexibility for scientific
sampling, tissue analyses, and
experimental manipulation. We also
cannot list the DPSs as endangered to
obtain legal leverage relative to the
larger industrial impacts, e.g., dams and
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bycatch, as suggested by the reviewer.
Rather, we must make our listing
determination based on application of
the statutory factors.
Comment 2 (new information on
Altamaha River population): Peer
reviewer 2 presented data on the
estimated abundance of age-1, river
resident Atlantic sturgeon in the
Altamaha River from 2004 to 2010,
which showed large estimated increases
in this age group in 2009 and 2010. The
peer reviewer also stated that he and
other researchers are beginning to detect
slower growth in age-1 Atlantic sturgeon
in the Altamaha and he attributed this
to density-dependent factors that are
beginning to limit available resources.
The reviewer stated that a few more
years of data are needed to determine if
the increasing trend is real, but none of
the other variables tested (e.g., river
flows or temperature) explain the trend.
The peer reviewer attributed the
apparent increases in juveniles in the
Altamaha to the moratorium on the
harvest of adults. The peer reviewer
stated that data are not available to
determine whether this trend is
occurring in other spawning
populations. The reviewer stated that
catch per unit effort (CPUE) data are
worthless without calibration or
validation and we do not have historical
abundance data to know what
abundance should be on any of the river
systems, though there is general
agreement that populations are a
fraction (less than 1 to 10 percent) of
historical abundance. The reviewer
recommended that long-term
monitoring of recruitment using markrecapture of age-1 juveniles be
implemented on key river systems.
Response: We are encouraged by the
apparent increases in juvenile Atlantic
sturgeon estimated by the peer
reviewer’s research in the Altamaha
River and appreciate the contribution of
this information for our consideration in
our listing determination. We revised
the relevant discussion in the text from
the proposed listing rule to include this
information. We agree that additional
years of data are necessary to confirm
this trend in the Altamaha and that we
cannot determine whether similar
trends may be occurring in other river
populations. This information is
consistent with information we
provided in the proposed listing rule,
which refers to the Altamaha River as
having a larger and healthier Atlantic
sturgeon population than any other river
in the Southeast. The proposed listing
rule also stated that juvenile Atlantic
sturgeon from the Altamaha are
relatively more abundant in comparison
to other rivers in the region.
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5919
Peer reviewer 2 noted that densitydependent factors may be starting to
limit available resources. We are
interested in working with the reviewer
and other researchers to determine
whether habitat modification, which we
describe in detail in the proposed listing
rule, is a contributing factor to the
limitation of resources in addition to the
increase in numbers of juvenile Atlantic
sturgeon utilizing the resources. While
water quality in the Altamaha River is
good at this time, the drainage basin is
dominated by silviculture and
agriculture, with two paper mills and
over two dozen other industries or
municipalities discharging effluent into
the river. Nitrogen and phosphorus
concentrations are increasing, and
eutrophication and loss of thermal
refugia are growing concerns for this
and other rivers in the South Atlantic
DPS. The Altamaha is one of the rivers
with current and pending water
allocation issues. We are currently
funding a project through the ESA
section 6 program to map habitats in
four Georgia rivers, including the
Altamaha, and this may be a valuable
step in answering this question.
We agree that CPUE data should be
used in the proper context and that
historical abundance data, other than
data from commercial fisheries in the
late 19th century, are not available.
However, as required by section
4(b)(1)(A) of the ESA, we must make our
listing determination based on the best
scientific and commercial data
available. When only CPUE and other
fishery-dependent data were available to
us, we clarified and acknowledged the
constraints of the data, and we conclude
that we used them in a valid manner.
This is further addressed in our
responses to several public comments
on specific sections of the proposed
listing rule (e.g., comments 19, 23, 24,
25, and 29).
Comment 3 (import of the 2003
workshop): Peer reviewer 1 stated the
proposed listing rule appeared to
dismiss any evidence of an increase in
Carolina DPS populations of Atlantic
sturgeon, citing a statement in the
proposed listing rule (page 61904–
61905) that ‘‘the [NMFS-sponsored
2003] workshop revealed mixed results
in regards to the status of Atlantic
sturgeon populations, despite the coastwide fishing moratorium. Some
populations seemed to be recovering
while others were declining.’’ The
reviewer stated that at the time of the
2003 workshop, the moratorium on
direct harvest and possession of Atlantic
sturgeon had only been in effect for 4
years and this was not sufficient time
for populations to increase in response
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to the protective measures. The reviewer
concluded the observations available at
the time of the 2003 workshop do not
provide a scientific basis for listing the
Carolina DPS as endangered.
Response: The information we
evaluated in making our proposed
listing determination of endangered for
the Carolina and South Atlantic DPSs of
Atlantic sturgeon was not confined to
the results of the 2003 workshop. As
stated in the proposed listing rule, the
information gathered at the 2003
workshop, including the equivocal
evidence that some populations
appeared to be recovering while others
were declining, prompted us to
complete a new review of Atlantic
sturgeon status, which was published in
2007. Since the ASSRT’s completion of
its status review, we obtained and
evaluated additional information on
threats to Atlantic sturgeon (see our
response to comment 1). Our evaluation
of this information indicates that the
moratorium on directed fisheries has
not and will not be sufficient to address
the impacts that are preventing sturgeon
populations from recovering (including
lack of access to required habitat, and
habitat quality issues). Section
4(b)(1)(A) of the ESA stipulates that
listing decisions be made using the best
available scientific and commercial
information, therefore we used
information from the 2007 status review
report (which incorporated information
from the 2003 workshop) and new
information in forming our
determination. Our responses to
comments from the public further detail
our use of information available at the
time of the proposed listing rule, as well
as our consideration of new information
submitted during the public comment
period.
Comment 4 (viability of small Atlantic
sturgeon populations): The estimated
343 spawning adult Atlantic sturgeon in
the Altamaha River exceeds the number
of spawning adults in the ‘‘very viable’’
Yellow River Gulf sturgeon population,
according to peer reviewer 1. Peer
reviewer 1 stated that information
presented in Schueller and Peterson
(2010) suggests a very robust
reproductive response to protection of
adult spawners under the Atlantic
sturgeon moratorium. From these data,
the reviewer stated that it seems highly
improbable that the Altamaha River
population is at risk of extinction and a
listing of endangered does not seem
applicable to the Altamaha population
within the South Atlantic DPS. The
reviewer stated that if the Altamaha
population follows the model of the
Suwannee River Gulf sturgeon
population after harvest was banned,
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then overall population growth in the
next decade will be exponential until
density-dependent population controls
come into play. Peer reviewer 1 also
stated that the ‘‘less than 300 spawning
adults’’ criterion in the proposed listing
rule for classifying a river population as
vulnerable to extinction sets a ‘‘very
high, probably unrealistic, bar,’’ and one
not conforming to scientific literature
documenting sturgeon population
recovery from much smaller effective
breeding population sizes (20–80
spawning females, based on examples
provided by the reviewer). Peer
reviewer 1 stated that sturgeon species
have the documented ability to
establish/re-establish viable populations
over a short timeframe (10 to 20 years),
starting from ‘‘a few tens’’ of spawning
adults without negative fitness impacts
from low genetic diversity.
Response: As explained above, NMFS
does not agree with peer reviewer 1’s
premise that an endangered listing
would only be appropriate if no single
historical spawning river within that
DPS appeared to sustain both a
relatively abundant and simultaneously
regularly-reproducing Atlantic sturgeon
population. We note that the Yellow
River population of Gulf sturgeon
referred to as ‘‘very viable’’ by the
reviewer is listed as threatened under
the ESA. While the number of spawning
adult Atlantic sturgeon in the Altamaha
River may be larger than that of Gulf
sturgeon in the Yellow River, the peer
reviewer noted that the Yellow River
population is one that has rebounded
since it was listed. The Altamaha River
population of Atlantic sturgeon is
estimated to be at only 6 percent of its
historical abundance. While there is a
moratorium on harvest and possession
of Atlantic sturgeon, the species is not
currently afforded the protections of
section 9 of the ESA, nor do they benefit
from the consultation and permitting
responsibilities of ESA sections 7 and
10, that apply to the listed Gulf
sturgeon. Information provided by peer
reviewers 1 and 2 indicated recent
(2009–2010) increases in the estimated
number of juveniles in the Altamaha
River. We are encouraged by this and
hope that the Altamaha River
population does exhibit exponential
growth, as the Suwannee River Gulf
sturgeon population did following
listing. However, our listing
determination is based on the best
information currently available to us,
and we do not feel that the information
provided on increases in juvenile
Atlantic sturgeon in the Altamaha River
or the comparison to Gulf sturgeon
populations in the Suwannee and
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Yellow Rivers provides a basis for
revising our proposed determination
that the South Atlantic DPS be listed as
endangered.
In response to the reviewer’s
comment that the ‘‘less than 300
spawning adults criterion’’ sets a ‘‘very
high, probably unrealistic, bar’’, we
clarify that the 300 spawning adults per
year was an estimate of the relative sizes
of Atlantic sturgeon river spawning
populations, based on the available
information on the annual spawning
adult abundance measured in the
Altamaha River (343 spawning adults)
and the fact that it is the largest
population in the Southeast, combined
with qualitative and quantitative
anecdotal information from the other
river systems. The 300 spawning adults
per year estimate does not constitute a
criterion or a bar for listing and/or
recovery as a general matter. Rather, the
estimate is evaluated in the specific
context of the Atlantic sturgeon river
populations and the impacts and threats
they face. These populations likely have
the capacity to recover, as the reviewer
suggested, if existing and future impacts
and threats are alleviated. The low
estimated population numbers in each
of the river systems within the DPSs
(1–6 percent of historical abundance),
combined with the ongoing impacts and
threats from habitat modification and
bycatch, indicate that the populations
are small and vulnerable, and the DPSs
they comprise are in danger of
extinction.
Comment 5 (sturgeon ability to
recolonize systems; genetic exchange):
Peer reviewer 1 stated that sturgeon
species are resilient and capable of
repopulating an extirpated river, or
colonizing a new river, if habitat
remains available, dams do not block
spawning ground access, water quality
is satisfactory, and a competing sturgeon
population is not already established. A
natal river population, well-established
over a long span of geological time and
highly adapted to its respective natal
river, would not realize success in
colonizing another river already
populated by a second population better
adapted to its respective natal river than
a potential colonist. The reviewer stated
that the low rate of genetic exchange
displayed among adjacent sturgeon
populations does not reflect the
incapacity of the species to colonize, but
the competitive advantage held by a preestablished natal river population facing
migrant individuals. The reviewer
provided examples of recolonization by
Atlantic sturgeon in bays and rivers
from New England to Labrador and
Newfoundland within a span of 10,000
years following deglaciation. The
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reviewer stated the northernmost and
most genetically conservative Atlantic
sturgeon population re-colonized over
1,500 miles of coastline within 40
generations (and probably much fewer)
in addition to undertaking a successful,
essentially instantaneous, 3,300 mile
migration to colonize the Baltic Sea
1,200 years ago. The reviewer also
provided an example of Gulf sturgeon
rebounding in Gulf Coast river systems.
Peer reviewer 1 stated that following
state harvest prohibitions on the Gulf
sturgeon and its listing as threatened
under the ESA, some river populations
have rebounded (the Yellow,
Choctawhatchee, and Suwannee river
populations). The reviewer concluded
the logic in the proposed listing rule is
not compelling that if one of the DPSs
were to be completely extirpated, it
would remain so over a long span of
time. Peer reviewer 3 stated, in
reference to the genetic analyses
showing fewer than two individuals per
generation spawn outside their natal
rivers, that this reflects the average
number of individuals and noted it
would be useful to compare this to
straying determined from tagging data.
Response: We agree with the peer
reviewer’s comment that the low rate of
genetic exchange displayed between
Atlantic sturgeon river populations may
reflect the competitive advantage held
by pre-established natal river
populations facing migrant individuals.
We revised the relevant discussion in
the text from the proposed listing rule
to include this information. However, as
stated in the proposed listing rule, we
do not expect Atlantic sturgeon that
originate from other river systems to
recolonize extirpated systems and
establish new spawning populations,
except perhaps over a long time frame
(i.e., many Atlantic sturgeon
generations). Though the reviewer
provided an example of Atlantic
sturgeon colonizing the Baltic Sea 1,200
years ago after a single migration, other
examples of recolonization provided
took 40 generations (approximately
1,000 years, based on a 25-year
generation period) to 10,000 years,
which is consistent with our statement
in the proposed listing rule. Further,
recolonization occurred in the absence
of present-day human impacts, such as
habitat modifications and mechanized
fishing.
We noted the reviewer’s comment
that sturgeon species are resilient and
capable of repopulating an extirpated
river or colonizing a new river if habitat
remains available, dams do not block
spawning ground access, and water
quality is satisfactory. As discussed
extensively in the proposed listing rule
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and in our responses to comments in
this document, Atlantic sturgeon in the
Carolina and South Atlantic DPSs and
the rivers within their ranges are
affected by habitat modification and
destruction, blocked access to spawning
grounds, downstream habitat impacts
caused by dams, and water quality (and
quantity) issues. Thus, the commenter’s
stated conditions for expecting
recolonization by Atlantic sturgeon are
not met. Atlantic sturgeon from the
Carolina and South Atlantic DPSs are
also taken as bycatch in fisheries.
Regarding the statement that following
state harvest prohibitions on the Gulf
sturgeon and its listing as threatened
under the ESA, some river populations
have rebounded (the Yellow,
Choctawhatchee, and Suwannee river
populations), the rivers that have
rebounded have two factors in common
versus those which have not rebounded
(e.g., the Apalachicola, Pascagoula, and
Pearl river populations): (1) No
mainstem dam on the natal river
limiting Gulf sturgeon access to upriver
spawning grounds or YOY access to
riverine feeding habitat; and, (2) no
major commercial fishery causing Gulf
sturgeon bycatch mortality in the natal
river, natal river estuary, or adjacent
marine waters. Assessing the impacts of
these two factors may be equally as
important to sturgeon population
recovery as is protection from all other
impacts, now that direct harvest has
been stopped. We agree with these
comments by the peer reviewer and also
believe that these threats associated
with dams, habitat, water quality, and
bycatch would hamper and slow
recolonization of extirpated river
systems. One reviewer acknowledged
that rivers, watersheds, and coastal
habitats inhabited by Atlantic sturgeon
have been drastically modified and
impacted by human activities (dammed,
channelized, de-watered, diverted,
dredged, mined, sedimented, polluted,
deforested, developed, populated by
introduced species, etc.) and that it
would be remarkable to achieve
recovery to even 10–30 percent of the
1890 carrying capacity of individual
sturgeon rivers.
In reference to peer reviewer 3’s
suggestion about comparing the degree
of straying from tagging studies to the
estimate of straying from the genetic
studies, we agree this could be a
valuable exercise in the future when we
have the necessary information on river
of origin (based on genetic analyses) and
the degree of straying (from tagging and
relocation studies). While the estimate
of less than 2 individuals spawning in
rivers outside their natal system is a
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5921
measure of successful transfer of genetic
information from a fish originating from
another system, the analysis suggested
by the peer reviewer would provide us
with knowledge of how many fish
actually stray into another system and
potentially attempt to spawn. This
could also provide insight into the
comments by the first peer reviewer that
lack of gene flow between river
populations is due to reduced success
from competition and not from lack of
attempts at migrant spawning.
Comment 6 (issues with estimating
sturgeon abundance): According to one
peer reviewer, targeted Atlantic
sturgeon population studies in the
Roanoke River and Santee-Cooper
system, as well as most other river
systems, have been limited in duration,
intensity, and continuity such that
population estimates may be
substantially underestimated. Peer
reviewer 1 noted that sturgeon species
are cryptic fish found in deep, mainstem
rivers. They are rarely observed
visually, not typically sampled in many
commercial river fisheries targeting
other fish species (with the exception of
the shad gill net fishery), and are rarely
caught by recreational anglers. The
reviewer stated that this illustrates that
presence and abundance of sturgeon
cannot be based on incidental catches
from commercial fisheries or scientific
sampling not specifically targeting
sturgeon. The reviewer stated that in the
past, sturgeon abundance has often been
vastly underestimated until an
appropriate and dedicated reporting or
sampling program was undertaken. The
reviewer recommended that only
continuous, standardized markrecapture efforts spanning sufficient
time (a minimum of 3 years, but
realistically greater than 5 years) can
provide reliable preliminary abundance
estimates.
Response: The majority of the data
presented in the proposed listing rule
came from studies targeting Atlantic
sturgeon or from fisheries that are
known to have a high incidence of
interaction with Atlantic sturgeon (i.e.,
gillnet fisheries). As much as possible,
we clarified the data collection methods
and constraints, and any assumptions
we made. This is also discussed in our
response to comment 2. We have used
the best available commercial and
scientific information to evaluate the
status of the Carolina and South
Atlantic DPSs, but we agree with the
reviewer that long-term, continuous,
standardized studies of Atlantic
sturgeon abundance are needed.
Comment 7 (viable population sizes
and sturgeon genetics): Peer reviewer 1
stated the minimum viable population
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sizes of several hundreds to several
thousands of individuals advanced in
the literature are not particularly
instructive with respect to sturgeon
species based on new genetic
information (Kreiger et al., 2006). The
reviewer commented that sturgeon are
polyploid and the significance of
polyploidy upon genetic diversity has
just emerged. Most fishes are diploid
with 40–50 chromosomes, a number
similar to most vertebrates. However, all
sturgeons are polyploid, having
approximately 120 chromosomes
(tetraploid, 4N), 240 chromosomes
(octoploid, 8N) or more, including
species with 12N or 16N ploidy.
Polyploidy allows for multiple alleles
(not just two as in diploid species) at a
given gene locus, allowing for intraindividual genetic variation (Kreiger et
al. 2006). The reviewer suggested that
this might explain the high degree of
plasticity displayed by sturgeon
populations and the documented ability
of sturgeons to repopulate from very few
spawning adults without apparent
inbreeding depression. He concluded
that until we gain a deeper
understanding of the genetics of
polyploidy and the implications
regarding sturgeon population
dynamics, any discussion of minimum
viable population size for sturgeon
populations cannot be phrased in terms
of what we know about inbreeding
depression in diploid mammal
populations. Thus, the 50/500 rule of
thumb cited in the proposed listing rule
may be an inappropriate criterion by
which to assess viability of sturgeon
populations, and we do not know how
few polyploid sturgeons are too few to
sustain a viable population.
Response: We appreciate the peer
reviewer’s input on the polyploid nature
of Atlantic sturgeon and how this
genetic characteristic may affect our
evaluation of minimum viable
population sizes in our listing
determination. We revised the relevant
discussion in the text from the proposed
listing rule to include this information.
As noted by the reviewer, we need a
deeper understanding of the genetics of
polyploidy and the implications
regarding sturgeon population
dynamics. We are not sure how
polyploidy in Atlantic sturgeon will
affect their recovery, but even if it
allows the species to repopulate from
relatively fewer individuals without
inbreeding depression, there is no
assurance that this will occur. Other
polyploid Acipenser species have
required listing under the ESA, such as
shortnose sturgeon (listed as endangered
in 1967), Gulf sturgeon (listed as
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threatened in 1991), and green sturgeon
(listed as threatened in 2006). In the
case of the shortnose sturgeon, recovery
has not been achieved even though it
has been protected for almost 45 years.
Further, the polyploid nature of Atlantic
sturgeon may further support the need
for protection under the ESA. Southern
populations of Atlantic sturgeon exhibit
high diversity and many low frequency
(and sometimes private) haplotypes
(Grunwald et al., 2008). Allendorf and
Leary (1988) noted that in polyploid
cutthroat trout, alleles constituting the
majority of the variation in the species
are found in only one or two local
populations, but they often occur at
high frequencies in those populations.
They concluded preserving the genetic
variation in cutthroat trout entails
preserving as many local populations as
possible. Finally, a polyploid nature
may not be sufficient to promote
recovery in Atlantic sturgeon
populations, even if it is indicative of
smaller viable population sizes, given
the nature and number of ongoing
impacts and threats to sturgeon and
their habitats.
Comment 8 (ACE Basin populations):
Peer reviewer 1 commented that the
statement in the proposed listing rule
that ‘‘the low population numbers of
every river population in the Carolina
and South Atlantic DPSs put them in
danger of extinction throughout their
ranges; none of the populations are large
or stable enough to provide with any
level of certainty for continued
existence of Atlantic sturgeon in this
part of its range,’’ seems contrary to the
data from recent Atlantic sturgeon
sampling results for the Altamaha,
Savannah, and ACE Basin. This
reviewer asserts that collection of 3,000
juvenile Atlantic sturgeon from the ACE
Basin in seven years of sampling is not
a low number.
Response: The proposed listing rule
stated that 3,000 juvenile Atlantic
sturgeon were collected in the ACE
Basin (consisting of the Ashepoo,
Combahee, and Edisto Rivers) between
1994 and 2001. While the reviewer did
not believe this is a low number, we
disagree. The ACE Basin and every
system in the South Atlantic DPS, with
the exception of the Altamaha River, is
estimated to be at 1 percent of its
historical abundance and to have less
than 300 adult Atlantic sturgeon
spawning per year (the Altamaha is
estimated to be at 6 percent of its
historical abundance and have 343
spawning adults per year). However, the
statement from the proposed listing rule
referred to by the peer reviewer was not
referring strictly to population size, but
rather to the restrictive effects of low
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population numbers in all component
river populations on the DPSs’ ability to
respond to threats. This statement was
taken from a section of the proposed
listing rule addressing viable population
size, and the statement was meant to be
taken in the context of the statements
that preceded it: ‘‘The concept of a
viable population able to adapt to
changing environmental conditions is
critical to Atlantic sturgeon, and the low
population numbers of every river
population in the Carolina and South
Atlantic DPSs put them in danger of
extinction throughout their ranges.’’
Low population numbers hamper
recovery by making the populations less
resilient to the dangers they continue to
face from being taken as bycatch and
from the loss, reduction, and
degradation of habitat resulting from
dams, dredging, and changes in water
quality parameters (such as depth,
temperature, velocity, and dissolved
oxygen). Because these DPSs are groups
of populations, the stability, viability,
and persistence of individual
populations affects the persistence and
viability of the larger DPS. In the
example of the ACE Basin, the capture
of 3,000 juvenile Atlantic sturgeon
between 1994 and 2001 (an average of
375 Atlantic sturgeon juveniles per year)
alone is not sufficient to indicate that
the DPS can persist, given the low
population numbers in each of the river
systems in the DPS and the existing
threats to the species (e.g., bycatch,
habitat degradation), some of which
may worsen as a result of water
allocation issues and climate change.
Comment 9 (relevance of historical
abundance estimates): Peer reviewer 1
commented on the statements in the
proposed listing rule that the Carolina
DPS is estimated to number less than 3
percent of its historical population size;
the Altamaha River is suspected to be
less than 6 percent of its historical
abundance; and the abundances of the
remaining river populations within the
South Atlantic DPS are estimated to be
less than 1 percent of what they were
historically. This describes the depleted
status of these populations, and
provides a reference point from which
to gauge re-population. Peer reviewer 1
commented that caution should be
exercised in using 1890s fisheries
abundance as the recovery target, and
similarly as a metric against which
population recovery can be measured.
Rivers, watersheds, and coastal habitats
inhabited by Atlantic sturgeon have
been drastically modified and impacted
by human activities (dammed,
channelized, de-watered, diverted,
dredged, mined, sedimented, polluted,
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deforested, developed, populated by
introduced species, etc.) and it would be
remarkable to achieve recovery to even
10–30 percent of the 1890 carrying
capacity of individual sturgeon rivers.
The reviewer believed the remaining 1–
6 percent of the historical population
numbers represents a good foundation
for population recovery at the beginning
of an unprecedented era of harvest
prohibition, habitat restoration, and
conservation awareness.
Response: The discussion in the
proposed listing rule of current
population size relative to historical
levels was not meant to imply those
levels would be recovery targets.
Relative population size was intended
as a metric of the depth of the DPS’
decline over time. The reviewer’s
observation that permanent habitat
modifications have reduced potential
population levels by 70–90 percent
underscores the significance of the
multiple habitat threats facing Atlantic
sturgeon.
Comment 10: Peer reviewer 1 took
issue with the statement in the proposed
listing rule that ‘‘recovery of depleted
populations is an inherently slow
process for a late-maturing species such
as Atlantic sturgeon.’’ This reviewer
stated this thesis is fundamentally faulty
for sturgeon and other species. Late
maturity in a species has little to do
with speed of population increase
beyond the initial lag period of one
generation span or less, after which
reproduction is continuous. As per the
theory of Malthus, the reviewer stated
that any population of any species in
nature, whether of mice or elephants,
will increase geometrically, as long as
resources are not limiting. For a
sturgeon population depleted by
overfishing, once subadults are
permitted to mature and spawn without
being harvested first, recovery can be
quite rapid if other human impacts have
not removed or severely restricted
essential resources.
Response: We have considered the
peer reviewer’s comment. However, we
continue to find that the Atlantic
sturgeon’s life history traits are
hindering its recovery in several ways,
as supported by scientific literature. For
example, Meyers and Worm (2005)
state, ‘‘from the land it is well known
that large species with high ages at
maturity are particularly vulnerable to
extinction (Purvis et al., 2000). There is
no reason to believe that this may be
different in the ocean (Myers & Mertz,
1998; Hutchings, 2001; Dulvy and
Reynolds, 2002; Dulvy et al., 2003).’’
Specifically regarding the Atlantic
sturgeon, Balazik et al. (2010) state that
‘‘the Atlantic sturgeon’s life history
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(high age of maturation and 2–5 years
between female broods) probably
inhibits population recovery (Boreman,
1997; Smith and Clugston, 1997).’’
Gardmark et al. (2003) states that ‘‘small
populations are sensitive to stochastic
effects, especially so if not all mature
individuals reproduce,’’ and as noted in
the proposed listing rule (as well as
Balazik et al., 2010, above), adult
Atlantic sturgeon do not reproduce
every year.
There are several ways the Atlantic
sturgeon’s life history traits may be
hampering recovery. The species’ late
age at maturity provides more
opportunities for individuals to be
removed from the population before
reproducing. The limited ability of
small populations with non-annual
spawning adults to respond to
stochastic effects could greatly affect
Atlantic sturgeon recovery, and human
population increases and climate
change are likely to exacerbate existing
water quality and quantity problems.
Based on their life history, Atlantic
sturgeon populations are more sensitive
to fishing (bycatch) mortality than other
coastal fish species. Like other Kselected species (which have large body
size, long life expectancy, and produce
fewer offspring, versus r-selected
species, which are characterized as
having high fecundity, small body size,
early maturity onset, short generation
time, and the ability to disperse
offspring widely), Atlantic sturgeon are
long-lived, have an older age at
maturity, and have lower maximum
fecundity values, with 50 percent of the
lifetime egg production for Atlantic
sturgeon occurring later in life
(Boreman, 1997). That species with
K-selected life history traits, such as
Atlantic sturgeon, exhibit greater
sensitivity to bycatch mortality is also
supported by Baskett et al. (2006):
‘‘fisheries have a greater long-term
negative impact on species with lower
population growth rates, later
maturation, larger organism size, and
greater longevity than on species with
faster production (Jennings et al., 1998;
Heino and God<, 2002).’’
We agree with the peer reviewer’s
comments that any species with discrete
generations or distinct breeding seasons
will increase geometrically, ‘‘as long as
resources are not limiting.’’ We also
agree that Atlantic sturgeon can recover
if fisheries mortality is reduced,
allowing sub-adults to recruit to the
spawning population, and ‘‘if other
human impacts have not removed or
severely restricted essential resources.’’
We stated in the proposed listing rule
that the species’ ‘‘long life-span also
allows multiple opportunities to
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5923
contribute to future generations
provided the appropriate spawning
habitat and conditions are available.’’
However, we believe that even though
prohibitions on direct harvest and
possession of Atlantic sturgeon have
been in place for years, their life history
characteristics, small population sizes,
and the continued threats associated
with bycatch and habitat modification
are hampering the recovery of Atlantic
sturgeon.
Comment 11: Peer reviewer 3
questioned why the use of samples from
YOY and mature adult Atlantic sturgeon
in the genetic analysis by Wirgin and
King (2006) ensured that the samples
came from fish originating in the
sampled river system. The reviewer
stated this implies that intermediate size
fish stray more than adults. The
reviewer also asked if the adults
sampled were running ripe adults.
Response: Whether all of the adults
utilized in the study were running ripe
(i.e., were making a spawning run) is
unclear. However, adults generally only
enter freshwater to spawn and the vast
majority of Atlantic sturgeon spawn in
their natal river (with estimates of less
than 2 individuals per generation
spawning outside their natal system).
Therefore, the use of genetic samples in
this study from adults captured in the
freshwater portion of a river would
indicate that the fish originated from
that river and had returned to spawn.
Similarly, Atlantic sturgeon spend the
first year of their life in their natal river.
Therefore, using genetic samples from
YOY in a river system ensures that the
fish originated in that river. Subadult
(fish older than 1 to 2 years old) Atlantic
sturgeon, as well as non-spawning
adults, are known to make extensive
coastal migrations. Subadults may use
multiple estuarine or riverine areas for
refuge, foraging, and nursery habitat,
while non-spawning adults make
extensive marine migrations. These life
stages were excluded from the study
because the river of origin cannot be
determined from the location the fish
are captured.
Comment 12: Peer reviewer 3 noted
that 88 percent average accuracy in
determining a sturgeon’s natal river of
origin was high and questioned whether
the 94 percent average accuracy in
assigning a sturgeon to one of the 5
DPSs was significantly better. The
reviewer asked if the variance around
the 88 and 94 percent figures is known.
The proposed listing rule stated that the
loss of either the Carolina or the South
Atlantic DPS would constitute an
important loss of genetic diversity for
the Atlantic sturgeon. The reviewer
commented that additional context on
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the amount of genetic diversity within
river populations, among river
populations within a DPS, and between
the 5 DPSs would better support that the
loss of a DPS would represent a
significant loss of diversity.
Response: The overall accuracy in
assigning an Atlantic sturgeon to its
natal river ranged from 60 to 94.8
percent (60 to 91.7 percent for
southeastern rivers), while the overall
accuracy in identifying a sturgeon to
one of the 5 DPSs ranged between 88.1
and 95.9 percent (91.7 to 95.9 percent
for the two southeastern DPSs). The
peer reviewer’s point is well-taken that,
while there is higher accuracy in
identifying a sturgeon to its DPS
because of clearer genetic differences
between the DPSs, the accuracy in
identifying a sturgeon to its natal river
is also quite high. We also agree with
the peer reviewer that the broader
context of the amount of genetic
diversity exhibited by Atlantic sturgeon,
within a DPS as well as among DPSs,
provides additional support for our
conclusion that the loss of a DPS would
constitute a significant loss of genetic
diversity. The high accuracy (60 to 92
percent) in utilizing genetic differences
to assign Atlantic sturgeon in the
Southeast to their natal rivers indicates
that there is a significant amount of
genetic diversity among rivers within a
DPS, as well as between the two
Southeast DPSs. Grunwald et al. (2008)
reports that southern Atlantic sturgeon
river populations have high diversity
and many low frequency (and
sometimes private) haplotypes. The
information from Grunwald et al. (2008)
indicates that each river population
within a DPS makes unique
contributions to the genetic diversity of
the DPS as a whole and lends greater
support to our determination that the
loss of a DPS represents a significant
loss of genetic diversity.
Comment 13: Peer reviewer 3 asked if
the statement in the proposed listing
rule that ‘‘with the exception of the
Waccamaw River population, all river
populations sampled within each
population segment along the entire
East Coast were geographically
adjacent’’ was intended to mean that,
with one exception, the genetic results
are consistent with geography. In
reference to the statement that the
sample size from the Waccamaw River
population was small (21 fish), the
reviewer asked what the sample size
was for the remaining river populations
utilized in the genetic analysis.
Response: The peer reviewer
interpreted the statement in the
proposed listing rule correctly. In
reference to the genetic sample sizes for
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rivers other than the Waccamaw, they
ranged from 35 to 115. However, it is
also important to note that genetic
samples used in the analysis for the
other river populations were taken from
YOY and adult Atlantic sturgeon only to
ensure that the fish were spawned in the
river they were captured in. The genetic
samples from Atlantic sturgeon
captured in the Waccamaw River, in
addition to being small in number, were
taken from only juvenile Atlantic
sturgeon, as those were the only
samples available. As stated previously,
juveniles may utilize multiple systems
for foraging and nursery habitat,
therefore the fish captured in the
Waccamaw River and used in the
genetic analysis were not necessarily
spawned in that system. We are revising
information in this final rule to indicate
that the genetic samples from the
Waccamaw River all came from juvenile
Atlantic sturgeon.
Public Comments
Comments on the Delineation of the
DPSs
Comment 14: Multiple comments
were received either disagreeing with
the listing of DPSs or disagreeing with
the way populations were grouped into
DPSs. One commenter stated that DPS is
not a scientific term and that the DPS
policy is arbitrary. The commenter also
stated that the decision to list five DPSs
results from the lack of NMFS’ scientific
ability to support the listing of the
species as a whole. Several comments
were received, some citing Grunwald et
al. (2008), that all riverine populations
of Atlantic sturgeon are genetically
distinct. Another commenter stated that
populations should either be evaluated
on a drainage-specific basis or as a
single unit south of Cape Hatteras
because current DPS delineations
combine high abundance rivers with
rivers that have low abundance or
unknown population status, are
extirpated, or exist at the margins of the
historical range. Comments were
received that the entire Carolina DPS
does not warrant listing as a unit and
that only populations from river systems
that would be afforded further
protection by an ESA listing should be
listed. Multiple commenters were
concerned that incorrect delineation of
DPSs could result in negative impacts to
Atlantic sturgeon.
Response: The ESA, as amended in
1978, included in the definition of
‘‘species’’ ‘‘any distinct population
segment of any species of vertebrate fish
or wildlife which interbreeds when
mature.’’ On February 7, 1996, the
USFWS and NMFS adopted a joint
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policy (61 FR 4722) regarding the
recognition of distinct population
segments (DPSs) under the ESA. We
agree with the commenter that this is
not a scientific term, which is
acknowledged in the policy itself: ‘‘the
authority to list a ‘species’ as
endangered or threatened is thus not
restricted to species recognized in
formal taxonomic terms, but extends to
subspecies, and for vertebrate taxa, to
distinct population segments (DPSs)’’
and ‘‘the term is not commonly used in
scientific discourse, although
‘population’ is an important term in a
variety of contexts.’’ The DPS policy is
not arbitrary, and has been upheld by
numerous courts as a rational and
permissible interpretation of the statute
by the Services. The policy formalizes
the criteria that must be met in order to
consider a subset of a species a DPS,
and those criteria are based on scientific
principles. The Services determined
that the listing, delisting, and
reclassification of DPSs of vertebrate
species would consider the discreteness
and significance of the population
segment in relation to the remainder of
the species to which it belongs.
We did not delineate the DPSs based
on population abundance information
and lumping high and low abundance
rivers. We do not agree that the best
available scientific information supports
listing other population segments in the
Southeast, such as on a drainagespecific basis or as a single DPS south
of Cape Hatteras. In accordance with the
DPS policy, we determined that two
DPSs of Atlantic sturgeon exist in the
Southeast based on genetic information
that indicates the DPSs as delineated
constitute cohesive ecological and
evolutionary units, on each DPS’
persistence in unique ecological
settings, and on the conclusion that the
loss of either population segment would
result in a significant gap in the range
of the species as a whole. In the
proposed rule, we stated that Atlantic
sturgeon studies consistently
demonstrated the species to be
genetically diverse and that between
seven and ten Atlantic sturgeon
population groupings can be statistically
differentiated range-wide (e.g., King et
al., 2001; Waldman et al., 2002; Wirgin
et al., 2002; Wirgin et al., 2005;
Grunwald et al., 2008).
Given a number of key differences
among the studies (e.g., the analytical
and/or statistical methods used, the
number of rivers sampled, and whether
samples from subadults were included),
it is not unexpected that each reached
a somewhat different conclusion. In the
proposed listing rule, we specifically
evaluated and discussed the information
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presented by Grunwald et al. (2008) and
concluded that, though they used
additional samples, some from fish in
the size range (less than 130 cm)
excluded in the analysis we relied on
because they were smaller than fish
considered to be mature adults, the
results of the genetic analyses in
Grunwald et al. (2008) and in the
proposed listing were qualitatively the
same and did not invalidate our DPS
structure. We agree that Atlantic
sturgeon from different riverine
spawning populations can be
distinguished genetically. However,
genetic distances and statistical analyses
(bootstrap values and assignment test
values) used to investigate significant
relationships among, and differences
between, Atlantic sturgeon river
populations, formed the basis of our
judgment that the DPSs as proposed
constitute cohesive ecological and
evolutionary units that are appropriate
for listing under the ESA and the DPS
policy. In our judgment, the groupings
of river populations into the DPSs as
proposed, incorporates likely patterns of
Atlantic sturgeon dispersal between
drainages.
We believe all river populations
within the DPSs will be afforded greater
protection by an ESA listing, and listing
the DPSs as proposed will not result in
negative impacts to Atlantic sturgeon.
Any action funded, authorized, or
undertaken by a Federal agency that
may affect Atlantic sturgeon from either
DPS would require consultation with
NMFS under section 7 of the ESA.
Those analyses will focus initially on
the impact of an action on the spawning
population(s) to which affected sturgeon
belong and then consider the
significance of those impacts to the
DPS(s).
Comment 15: Several commenters
said use of the genetic data that are
available for the designation of DPSs
may be unreliable due to limited sample
sizes, spatial, temporal, and ontogenetic
differences in collection, and lack of
samples from all river systems.
Commenters also said our review of the
literature was based on techniques used
rather than the samples used to derive
the conclusions. A commenter stated
(citing Grunwald et al., 2008) that
genetic analyses should have been
restricted to samples from spawning
adults. The commenter cited several
studies (Grunwald et al., 2008; Wirgin
and King, 2006; Wirgin et al., 2005;
Wirgin et al., 2000) as indicating that
the north-to-south clustering of Atlantic
sturgeon river populations into DPSs is
not valid. The commenter believed
NMFS relied on genetic studies to say
that there are genetic differences among
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populations but then ignored the actual
results of the studies. The commenter
stated that the Wirgin and King data
were not peer reviewed and should be
given less consideration. The
commenter also stated that genetic
information needs to be integrated with
ecological and behavioral data in order
to draw appropriate conclusions.
Commenters stated that more data are
needed to list DPSs and that although
the peer reviewed studies have
described a high degree of genetic
separation with good classification
success, there are problems when the
papers are reviewed and considered for
management. Several commenters noted
that genetic samples for adult sturgeon
will be collected in upcoming years
through federally funded projects along
the Atlantic Coast.
Response: We agree with commenters
that sample sizes, spatial, temporal, and
ontogenetic differences in collection,
and lack of samples from all river
systems create uncertainty in the
Atlantic sturgeon genetic data. However,
in our judgment the available data show
genetic separation of the Carolina and
South Atlantic DPSs from northern
populations and from each other.
Results showed 92 and 96 percent
accuracy in correctly classifying a
sturgeon from four sampled river
populations (the Albemarle Sound,
Savannah, Ogeechee, and Altamaha
River populations) to two groupings of
river populations (Albemarle Sound and
Savannah/Ogeechee/Altamaha Rivers).
Contrary to the commenters’ assertion,
in reviewing the literature and
evaluating the available genetics data in
our consideration of DPSs we looked at
both technique and the samples used.
As stated in Grunwald et al. (2008), due
to the potential for subadult and adult
Atlantic sturgeon to undertake extensive
migration between systems, specimens
certain to be spawned within a system
(and candidates for use in genetic
studies of spawning populations)
include spawning adults or juveniles
less than two-years-of-age. When
possible, the genetic analyses we relied
on in the 2007 status review report and
in the proposed listing rule limited the
samples utilized to those collected from
spawning adults and YOY, which is
consistent with (and more restrictive
than) what Grunwald et al (2008)
described. Where genetic samples from
adult and YOY were missing, we
reported the results of other analyses
utilizing juvenile Atlantic sturgeon and
clarified this in the proposed listing
rule.
The commenter presented a
comparison of river groupings (UPGMA
trees) derived from genetic cluster
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5925
analyses from the cited studies and
asserted that the various groupings
conflicted with the DPS structure
proposed in the listing, stating that a
north-to-south clustering of river
populations is not valid. However, there
is no difference in the river groupings
resulting from the genetic analyses
presented in Grunwald et al. (2008;
Figure 3) and the river groupings
resulting from genetic analyses
presented in the 2007 status review
report (Wirgin and King, 2006; Figure
16) for the southern populations. The
river groupings presented in Wirgin et
al. (2000) differ from our results, likely
due to the inclusion of samples from
subadults which may have originated
from a system other than where they
were collected. Wirgin et al. (2000) did
find a pronounced latitudinal cline in
the number of composite mtDNA
haplotypes and in haplotypic diversity,
which increased from north to south.
The researchers ascribed the greater
genetic diversity within and among
southern populations to the persistence
of these populations through the
Pleistocene and to the faster mutation
rates associated with their shorter
generation times. The genetic results
referred to by the commenter in Wirgin
et al. (2005) were for shortnose
sturgeon, not Atlantic sturgeon.
While the genetic analysis by Wirgin
and King presented in the 2007 status
review report was not previously
published, it was peer reviewed as part
of the status review and as part of the
proposed rule. The status review report
was peer reviewed by six experts from
academia, and the proposed listing rule
was peer reviewed by three experts, two
from academia (including an Atlantic
sturgeon genetics expert) and one from
a Federal resource agency.
We agree with the comment that
genetic information needs to be
integrated with ecological and
behavioral data in order to draw
appropriate conclusions. We relied on
behavioral information (i.e., the
migratory nature of subadults and nonspawning adults) to determine the
appropriate life stages (i.e., YOY and
spawning adults) to use for the genetic
analysis. We also used behavioral and
ecological information in conducting
our DPS analysis per the Services’ joint
DPS policy. We considered the species’
behavior in that the majority of Atlantic
sturgeon return to their natal rivers to
spawn. We also considered ecological
issues, such as the fact that the DPSs
persist in unique ecological settings and
that the loss of a DPS would represent
a significant gap in the range of the
species.
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Additional genetic analyses will
improve our understanding of Atlantic
sturgeon and their population structure,
and we eagerly await the results of
upcoming and ongoing genetic analyses,
some of which we are funding through
our Species Recovery Grant Program
under section 6 of the ESA. However,
we believe that the currently available
data support the discreteness and
significance of the Carolina and South
Atlantic DPSs. Because we have
integrated the genetic data with other
sources of Atlantic sturgeon
information, such as the behavioral and
ecological information noted above, we
do not believe listing DPSs will create
management problems.
Comment 16: One commenter
disagreed that the DPSs persist in
unique ecological settings, citing a study
by the Institute for Ocean Conservation
(2010) that Atlantic sturgeon tagged in
the Hudson traveled from Nova Scotia
to Georgia. The commenter also
disagreed that the loss of a DPS would
result in the loss of important genetic
diversity, citing Quattro et al. (2002)
that dispersal is sufficient to prevent
deep divergence over long evolutionary
scales and Peterson et al. (2008) that
Atlantic sturgeon are resilient to genetic
bottlenecks.
Response: The proposed listing rule
states multiple times that Atlantic
sturgeon mix extensively in the marine
environment, which is consistent with
the citation provided by the commenter.
However, we disagree with the
commenter that the Carolina and South
Atlantic DPSs do not persist in unique
ecological settings. The vast majority of
Atlantic sturgeon return to their natal
river to spawn, and the spawning
habitat of each DPS is found in a
separate and distinct ecoregion as
identified by The Nature Conservancy
(TNC) based on the habitat, climate,
geology, and physiographic differences
for terrestrial and marine ecosystems
throughout the range of the Atlantic
sturgeon.. The unique ecological
characteristics of the ecoregions the
Carolina and South Atlantic DPSs
originate from are described in detail in
the proposed listing rule.
We disagree with the comment that
the loss of a DPS would not result in the
loss of important genetic diversity.
Grunwald et al. (2008) note that, while
northern populations of Atlantic
sturgeon have low genetic diversity,
southern populations exhibit high
genetic diversity with many low
frequency haplotypes. The loss of
genetic diversity associated with the
loss of either the Carolina or South
Atlantic DPS would reduce the ability of
Atlantic sturgeon as a subspecies to
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adapt to new selective pressures, such
as climate change or shifts in available
resources. We also disagree with the
commenter’s assertion that Peterson et
al. (2008) supports a conclusion that
Atlantic sturgeon populations are
resilient to bottlenecks. Peterson et al.
(2008) reported ‘‘pronounced cropping’’
of genetic diversity in the Altamaha
River Atlantic sturgeon population. The
researchers expressed surprise over this
result ‘‘given the resiliency to genetic
bottlenecks previously reported in other
studies of remnant Atlantic and
shortnose sturgeon populations (Quattro
et al., 2002; Waldman et al., 2002).’’
Grunwald et al. (2008) also stated that
‘‘current populations from the Hudson
River northward represent step-wise
recolonizations with a bottlenecking
effect.’’
Comment 17: One commenter stated
that the proposed listing rule suggested
the number of Atlantic sturgeon
spawning in locations other than their
natal rivers (‘‘outmigrants’’) is not
dependent on population size. The
commenter believed the rate of
outmigration is much higher than stated
and should be presented as a
percentage, but that some level of
mixing should be considered. Another
commenter stated that recolonization of
a basin would be slow regardless of
whether adjacent populations are low or
robust due to the low rate of
outmigration and genetic transfer
between basins. The commenter noted
that there are greater distances between
rivers within the Carolina DPS than
between the Carolina and South
Atlantic DPSs. This commenter stated
that if outmigration is limited and most
likely occurs between adjacent
populations, this refutes the DPS
structure.
Response: The number of Atlantic
sturgeon outmigrants (less than 2 per
generation) included in the proposed
listing rule was estimated from genetic
analyses by the studies we cited. We did
not relate outmigration of Atlantic
sturgeon to population size in the
proposed listing rule, and we do not
have available data to present
outmigration as a percentage of
population size; however, we agree with
the commenter that rates of
outmigration may increase with
increasing population size. We agree
that recolonization of a system from
adjacent populations would be slow,
which is consistent with statements in
the 2007 status review report (page 97)
and in the proposed listing rule (page
61912). The distances separating rivers
within and between the Carolina and
South Atlantic DPSs do not account for
the extremely low level of outmigration
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in Atlantic sturgeon spawning
populations. Adult (and subadult)
Atlantic sturgeon are known to make
extensive movements between systems
along the East Coast range of the
species. Though the exact cues are not
known, it is a life history characteristic
of Atlantic sturgeon that the vast
majority spawn in their natal river
system. The low level of outmigration
does not refute the DPS structure; as we
stated above, the groupings of river
populations into the DPSs as proposed,
incorporates patterns of Atlantic
sturgeon dispersal among drainages.
The evidence supporting the structure
of the Carolina and South Atlantic DPSs
is presented in the proposed listing rule
and in our responses to comments
14–16 above.
Comment 18: Several commenters
stated that there were no specific
geographic boundaries or coordinates
listed to delineate the five DPSs and
believed this should be addressed in the
final rule, since conservation and other
management measures will likely be
implemented based on the delineation
of the DPSs. The commenters also had
concerns that the rivers and tributaries
listed in each DPS are not all-inclusive
and could potentially create loopholes
for management and conservation
measures. Another commenter stated
that the extensive mixing of Atlantic
sturgeon in the marine environment will
make conservation and management of
the DPSs difficult to impossible.
Response: We do not believe
additional geographic boundaries or
coordinates delineating the DPSs are
necessary or that there are any
loopholes for management or
conservation. As stated in the proposed
rule text, each of the DPSs is defined to
include fish that spawn in the range of
watersheds encompassed by the DPS.
Our intent was that all fish spawned in
such watersheds would also be included
in the listing throughout their life
cycles. Thus, fish spawned in one river,
but using an adjacent river as nursery or
subadult feeding habitat, are included in
the listing. We have refined the text
descriptions of the Carolina and South
Atlantic DPSs in the final listing rule to
more clearly reflect this issue. The
modifications to the text clarify the
riverine ranges of the DPSs but do not
change the populations making up each
of the Southeast DPSs.
As noted by commenters, Atlantic
sturgeon from each riverine watershed
and DPS may be found in multiple
riverine, estuarine, and marine
environments at various life stages. We
agree that the extensive mixing of
Atlantic sturgeon will make
conservation and management of the
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DPSs challenging. As we stated in the
proposed listing rule, this extensive
mixing of Atlantic sturgeon in the
marine environment, as well as in
multiple riverine and estuarine systems,
can expose Atlantic sturgeon of a given
DPS to a variety of threats at various life
stages and in multiple locations. We
discuss management challenges and
potential strategies for dealing with
them in the sections of the proposed
and final listing rules entitled
‘‘Identifying the DPS(s) Potentially
Affected by an Action During Section 7
Consultation.’’
Species Data and Information Supplied
by Commenters
Comment 19: Commenters from North
and South Carolina state agencies and
other commenters supplied data and
information for the Carolina DPS. One
comment stated that there was an
observed increase in abundance of
Atlantic sturgeon in Albemarle Sound
between 2005 and 2008. The commenter
also stated there was a slight increase in
abundance of juveniles and subadults in
Pamlico Sound, while river surveys
showed a slight decrease in abundance.
Commenters also included data from
late 2010 indicating there is a fall
spawning run in the Roanoke River.
Based on anecdotal angler reports from
North Carolina, some commenters
asserted that Atlantic sturgeon are
persisting, though there has been little
improvement in the size and age
distributions of the Carolina DPS
relative to historical levels. They also
noted sampling efforts directed toward
sturgeon have been sparse and limit
ability to accurately characterize
existing populations. Comments from
South Carolina noted that Atlantic
sturgeon were captured in most nets set
in Winyah Bay from April to July in
2007 to 2009, including sites far upriver,
and that sonar sampling indicated
several hundred Atlantic sturgeon at the
confluence of the Sampit River and
Winyah Bay in 2009. A commenter
stated that fishery surveys conducted as
a requirement of the Federal Energy
Regulatory Commission (FERC) license
for the Yadkin-Pee Dee Hydroelectric
Project resulted in the capture of a
running ripe male in the Pee Dee River
in October of 2003, indicating spawning
activity. Large fish believed to be
Atlantic sturgeon were sighted during
electrofishing from 2002 to 2003. The
commenter stated that this and other
research (Collins and Smith, 1997;
Collins et al., 2003; Gibbons and Post,
2009) suggest that there may be a
sizeable Atlantic sturgeon population
present in the Pee Dee River and the
Winyah Bay system. State agency
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comments noted that there have been
few encounters with Atlantic sturgeon
in the Santee River and there are
anecdotal reports of breaching sturgeon
in the Cooper River.
Response: We reviewed the specific
information supplied for Atlantic
sturgeon from the Carolina DPS and
have added it to the ‘‘Distribution and
Abundance’’ section of the final listing
rule; however, this information does not
require a change in our listing
determination. The Independent Gill
Net Survey (IGNS) data supplied by the
North Carolina Department of
Environment and Natural Resources
(NCDENR) does show an increase in
CPUE between 2005 and 2008 in
Albemarle Sound. Based on Table 1 and
Figure 2 included in NCDENR’s
comments, the CPUE in 2005 was 0.012,
and increased in each successive year
until 2008, when it reached 0.031.
However, the data supplied by NCDENR
for Albemarle Sound dates back to 1990
and continues to 2009. The 1990–2009
CPUE data as a whole shows a great deal
of fluctuation, with no increasing trend,
but rather periodic increases and
decreases. In 2009, the CPUE dropped
back down to 0.015, the level recorded
in 2006. While 2008 was the highest
CPUE observed since 2002, the CPUEs
recorded for 1990 (0.081), as well as for
2000 and 2001 (0.032 both years), were
actually the highest recorded in the
1990–2009 dataset for Albemarle Sound
provided by NCDENR. The lowest CPUE
levels recorded in the 1990s (0.005 to
0.010 in 1992, 1993, 1995, and 1996)
were observed again in 2002, 2003, and
2004 (0.005 to 0.007). The commenter
stated that there has been an increase in
juveniles and subadults in Pamlico
Sound since 2001. Based on IGNS data
provided by NCDENR (Table 4, Figure
8), the CPUE for Pamlico Sound was 0
in 2001, and greater than 0 for 2002
through 2009. While all CPUEs for
Pamlico Sound are greater than that
recorded in 2001, there is no apparent
increasing trend in the data. While the
highest CPUEs were observed between
2004 and 2007 (0.016 to 0.066), the
highest being recorded in 2005, the
CPUE has decreased since 2005. The
level observed in 2009 (0.003), the
lowest CPUE in this dataset, was also
observed in 2002 and 2003. Similarly,
the river surveys of the Pamlico, Pungo,
and Neuse Rivers showed a peak CPUE
in 2005, with very low numbers
observed in the other years within the
survey period of 2000 to 2009. The
information provided by the commenter
on spawning in the Roanoke River
supports information included in the
proposed listing rule. While the
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Roanoke was determined to be an active
spawning river within the Carolina DPS
in the proposed listing rule, information
supporting that a fall spawning run
occurs there will greatly aid in the
conservation and management of the
species. We agree with the commenters’
statement that Atlantic sturgeon are
persisting, though there has been little
improvement in the size and age
distributions of the Carolina DPS
relative to historical levels. The failure
of Atlantic sturgeon populations to
rebound, even with the moratorium on
harvest and possession and other efforts
to recover the species, is the primary
reason we are proposing to list the
species as endangered. In 1901, the
Atlantic sturgeon fishery collapsed
when less than 10 percent of the U.S.
1890 peak landings were reported. The
landings continued to decline
coastwide, reaching about 5 percent of
the peak in 1920. Coastwide landings
remained between 1 and 5 percent of
the 1890 peak levels until the Atlantic
sturgeon fishery was closed by ASMFC
in 1998. Atlantic sturgeon populations,
estimated to be 1 to 6 percent of their
historical levels, have remained
relatively unchanged since the initial
collapse caused by the Atlantic sturgeon
fishery of the late 19th century. We
agree that sampling efforts need to be
increased to effectively characterize
populations and we are making efforts
to see that it happens.
The South Carolina Department of
Natural Resources provided information
(SCDNR) for Winyah Bay that Atlantic
sturgeon were encountered in most nets
set from April to July (2007 to 2009) and
that a researcher using sonar observed
several hundred Atlantic sturgeon in
Winyah Bay near the confluence of the
Sampit River in 2009. We contacted Dr.
Hightower, the researcher conducting
the sonar work in Winyah Bay, to get
further information on his observations.
Dr. Hightower provided additional
information via email on July 7, 2011,
that he and fellow researchers were
conducting ‘‘pilot trials without a
specific survey protocol, so we have not
tried to generate density estimates. One
of the issues that must be resolved
before using the side-scan files in a
quantitative way is to estimate the
probability of identifying (detecting) a
sturgeon, given that it is present in the
area surveyed by the side-scan sonar.
We are still working on that question,
but results to date suggest that the
detection probability depends on fish
size, position in the water column, and
possibly orientation relative to the
sonar. Thus, we could come up with a
density estimate for fish above some
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size threshold, but we would not be able
to reliably estimate how many of those
were Atlantic sturgeon. Some of the
large fish on those images are clearly
Atlantic sturgeon and many others are
likely to be sturgeon. The statement that
several hundred were in that area is a
reasonable description of what the sidescan data show but we are not at the
point of being able to estimate the
density with confidence.’’ Dr.
Hightower also remarked that ‘‘we have
done pilot survey work in the Roanoke,
Neuse, Cape Fear, and Pee Dee river
systems. The side-scan images from the
Pee Dee (Winyah Bay) suggest markedly
higher densities than in the other
rivers.’’ If all fish detected by Dr.
Hightower were Atlantic sturgeon, the
possibility that there were hundreds in
Winyah Bay does not conflict with our
estimate of less than 300 spawning
adults per year in each spawning river.
The sonar study was conducted in
August 2009. Due to the time of year
and location, it is unlikely this was a
spawning aggregation and there is no
way of knowing what age classes were
present. It is possible that some of these
fish were juvenile Atlantic sturgeon,
which are known to utilize multiple
riverine and estuarine systems other
than their natal system. The information
provided regarding the surveys
conducted on the Yadkin-Pee Dee as a
requirement of a FERC license is not
new information, as it was included in
the proposed listing rule. The
information on the Santee-Cooper
system is noted, and it is consistent
with the proposed listing rule. The
information for Atlantic sturgeon in the
Carolina DPS presented by commenters,
when considered as part of our listing
determination, does not change our
determination that the Carolina DPS
warrants listing as endangered. In our
judgment, none of the river populations
in the DPS are large or stable enough to
provide with any level of certainty for
the continued existence of the DPS in
the face of threats currently acting on
the species. In our judgment, the
Carolina and South Atlantic DPSs
require multiple stable spawning
populations.
Comment 20: Commenters from state
agencies supplied data and information
for the South Atlantic DPS. South
Carolina Department of Natural
Resources (SCDNR) supplied data from
the Edisto, where 3,661 Atlantic
sturgeon were captured between 1994
and the present; their population
models estimate between 20,000 to
70,000 sturgeon. Between 1997 and
1999, SCDNR captured 118 adults in the
Edisto River during spring and fall
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spawning runs, but netting ceased once
that number was reached. They believed
if they had continued netting activities,
they would have captured more than
300 spawning adults. SCDNR also noted
approximately 20 adults were captured
in one to two months during surveys
targeting other species. In 2010, four
adults tagged in the 1990’s as age 0+
were recaptured, which they believe
indicates the moratorium is having the
desired effect of allowing fish to recruit
to the broodstock population. In the
Savannah River, the SCDNR captured
369 Atlantic sturgeon between 1997 and
2010. SCDNR commented that there is
not enough data to support the
contention that the Altamaha has the
largest population in the southeast and
that other rivers have less than 300
spawning adults per year. The Georgia
Department of Natural Resources
(GADNR) commented that there is new
information on the potential increase of
Atlantic sturgeon in the Altamaha, and
additionally, the Satilla River has been
found to contain a substantial number of
fish, where few to none were thought to
exist in the past. Citing Peterson et al.
(2008), GADNR stated the Altamaha
may be recovering, though absence of
adults older than age 17 suggests the
effects of overfishing are still evident.
According to Georgia’s recent
compliance reports to the ASMFC, the
2009 and 2010 estimates of age-1
Atlantic sturgeon in the Altamaha River
were two and five times the estimates
from the 2004–2008 period,
respectively. In the most recent
compliance report to ASMFC,
University of Georgia (UGA) researchers
collected more than 200 Atlantic
sturgeon in the Satilla River in less than
2 years of sampling. They concluded
that the presence of juvenile fish
measuring less than 50 cm indicates this
is likely a self-sustaining, spawning
population.
Response: We reviewed the specific
information supplied by the states for
Atlantic sturgeon from the South
Atlantic DPS and have added it to the
‘‘Distribution and Abundance’’ section
of the final listing rule. However, the
additional information does not require
a change in our listing determination.
SCDNR stated that in the 16-year period
since 1994, they captured 3,661 juvenile
(one- to three-year-old) Atlantic
sturgeon in the Edisto River. This
updates information we included in the
proposed listing rule that over 3,000
juvenile Atlantic sturgeon were
collected in the ACE Basin between
1994 and 2001, including 1,331 YOY.
SCDNR used Lincoln-Peterson and
Schnabel models to derive Atlantic
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sturgeon population estimates from
these data, which resulted in estimates
of 70,000 and 20,000 Atlantic sturgeon
in the Edisto River, respectively. SCDNR
commented that the models’ results
suggest increasing trends in abundance.
Both models rely on mark-recapture
data and assume a closed population
(there are no births, deaths, or
immigration/emigration between the
initial capture and the recapture period)
and that all individuals have an equal
chance of being captured (Nichols,
1992; Lindeman, 1990; Chao, 1987). We
note that there is great uncertainty in
the population estimates resulting from
the two models, as evident in the great
disparity between the two results
(20,000 versus 70,000). The reliability of
the population models used depends on
the validity of the assumptions of those
models. The primary assumption of
these two models, that each individual
has an equal probability of capture, is
likely unattainable in natural
populations (Chao, 1987; Carothers,
1973). The assumption of a closed
population is probably violated for any
estimate calculated using the Schnabel
or Lincoln-Petersen method on data
collected over several weeks or months,
and it is surely violated when data from
one or more active seasons are used
(Lindeman, 1990). SCDNR indicated
they are currently completing an open
system model (which is based on
survival probabilities, as well as capture
probabilities) to better assess the
Atlantic sturgeon population in the
Edisto River. Because the closed system
models used by SCDNR provide
estimates of juvenile abundance only
and do not account for other population
dynamics (birth, mortality, immigration/
emigration), the estimates provided by
the models likely represent an
overestimate of juvenile abundance, do
not provide an estimate of how many
juveniles likely mature into spawning
adults, and do not provide any
information that undermines our use of
the estimate of less than 300 spawning
adults per year in the system. Atlantic
sturgeon do not reproduce every year;
females reproduce on the order of once
every 2 to 5 years, males every 1 to 5
years. Small numbers of fish spawning
can reduce the likelihood of successful
spawning and the amount of genetic
variation introduced into the next
generation.
SCDNR commented that we do not
have enough data to support the belief
that the Altamaha River has the largest
spawning population in the Southeast
and that all other rivers have less than
that. However, we relied on the best
available information in arriving at the
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estimate, and the information supplied
by commenters, including the data
provided by SCDNR, actually supports
the estimate. The Altamaha is believed
to have the largest Atlantic sturgeon
spawning population in the Southeast,
based on the absence of dams impeding
access to appropriate spawning habitat,
the lack of heavy development in the
watershed, and relatively good water
quality. The information supplied by
GADNR showed an increase in age-1
Atlantic sturgeon from the Altamaha
River in 2009 and 2010 over 2004 to
2008 levels. This was also reported by
peer reviewer 2 and discussed in our
response to comment 2. The information
provided for the Satilla River is
consistent with information in the
proposed listing rule that the Satilla
River has a resident spawning
population of Atlantic sturgeon. The
information for Atlantic sturgeon in the
South Atlantic DPS presented by
commenters, when considered as part of
our listing determination, does not
change our determination that the South
Atlantic DPS warrants listing as
endangered. In our judgment all river
populations in the DPS are too small to
be stable and self-sustaining.
Comment 21: In response to our
request in the proposed listing rule for
information on the mixing of Atlantic
sturgeon populations, the petitioner
cited Erickson et al. (2011) stating that
out of 15 Atlantic sturgeon tagged in the
Hudson River, one was relocated in
Georgia, which supports extensive, long
range mixing of sturgeon. The petitioner
also cited Laney et al. (2007) that
Atlantic sturgeon from the Hudson
River represent approximately
44 percent of those in North Carolina
overwintering habitat.
Response: We appreciate the
information provided by the petitioner.
These studies support our assertion in
the proposed listing rule that extensive
mixing of the DPSs outside their natal
rivers occurs during non-spawning
phases. We are continuing to seek
information on the degree of mixing of
the different river populations,
including through our funding of the
project to determine seasonal and
spawning migration patterns and
incidences of inter-basin transfer for
adult Atlantic (and shortnose) sturgeon
in southeastern rivers in North Carolina,
South Carolina, and Georgia.
Comments on Abundance and Trends
Comment 22: Many comments were
received stating that the abundance
estimate of 300 spawning adults per
year is not supported by data. Many of
these comments stated that the
proposed listing rule is not valid
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without stock assessments of Atlantic
sturgeon populations. One commenter
stated that the estimate of 300 spawning
adults per year is misleading in regards
to total population abundance since
Atlantic sturgeon do not spawn every
year and the total population abundance
is likely much higher. Another
commenter, citing Schueller and
Peterson (2010), stated that we should
consider juveniles rather than spawning
adults. A comment was received that
the statements on page 61920 of the
proposed listing rule about spawning
populations being less than the 500
recommended by Thompson (1991)
conflict with the statement that total
population abundances for the Carolina
and South Atlantic DPSs are not
available. Additionally, a comment was
received that based on modeling,
populations in the Winyah Bay system
and the ACE Basin have more
individuals than Thompson (1991)
recommended as minimum viable
population sizes for short-term and
long-term population fitness.
Response: In response to comments
on lack of stock assessments being a bar
to listing determinations, we note that
section 4(b)(1)(A) of the ESA provides
that the Secretary shall make required
listing determinations solely on the
basis of the best scientific and
commercial data available to him at the
time of the determination, after
conducting a review of the status of the
species and taking into account efforts
to protect the species. Even if a formal
stock assessment of the species has not
been conducted, if the best available
information indicates the species
warrants listing, as it does for Atlantic
sturgeon, then we are required to list the
species. Lack of formal stock
assessments is not an unusual
circumstance for species that have
drastically declined, are at very low
population numbers, or whose ranges
have constricted, such that they are the
subject of petitions to list them as
threatened or endangered. Though we
do not have stock assessments, we
believe the current body of information
on the declines of Atlantic sturgeon, the
failure of their population numbers to
rebound despite harvest prohibitions,
the small relative magnitude of riverine
spawning populations, and the ongoing
impacts and threats from bycatch and
habitat modification, warrant listing the
Carolina and South Atlantic DPSs as
endangered.
In the Southeast, the Altamaha is the
only river where abundance has been
directly surveyed. While traditional
stock assessments from other Southeast
rivers in the species’ U.S. range are not
available, we nevertheless relied on the
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5929
best available data to produce a relative
estimate of the number of Atlantic
sturgeon in the remaining spawning
populations. Based on a comprehensive
review of the available data, the
literature, and information provided by
local, state, and Federal fishery
management personnel (both
documented in the 2007 status review
report and in comments received on this
rule), it is our judgment that the
Altamaha River has the largest Atlantic
sturgeon spawning population in the
Southeast. The larger size of this
population relative to the other river
populations in the Southeast is likely
due to the absence of dams that impede
access to appropriate spawning habitat,
the lack of heavy development in the
watershed, and relatively good water
quality, as Atlantic sturgeon
populations in the other rivers in the
Southeast have been affected by one or
more of these factors. Though
abundance estimates from stock
assessments are not available for the
other river populations, because the
Altamaha spawning population is the
largest, we believe it is reliable to
estimate the size of other spawning
populations in the Southeast Region as
no more than 300 adults spawning per
year. Further, data supplied by
managers and researchers (and
discussed in the previous section of
responses to comments), support an
estimate of less than 300 spawning
adults per year in the other Southeast
rivers.
The use of annually spawning adults
is not intended to be misleading. We
agree with the commenter that total
riverine population numbers of Atlantic
sturgeon are higher than the number of
annually spawning adults. However, the
only quantitative abundance estimate
available to us when the proposed rule
published was the number of annually
spawning adults in the Altamaha River,
not total population numbers or the
total number of juveniles, as suggested
by another commenter citing Schueller
and Peterson (2010).
Schueller and Peterson (2010) stated
that quantified methods of assessing
sturgeon recruitment are essential for
evaluating population trends, but that
early life stages of most sturgeon species
are notoriously difficult to sample, and
their study on the Altamaha River
provides the first quantified recruitment
data describing a juvenile Atlantic
sturgeon population in a southern river.
They conducted their research during
the summers of 2004 to 2007 and
estimated that juvenile abundance
ranges from 1,072 to 2,033 individuals
in the Altamaha River, with age-1 and
age-2 individuals comprising greater
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than 87 percent of the population. Based
on modeling, estimated apparent
survival and per capita recruitment
indicate that the juvenile population
experiences high annual turnover:
Apparent survival rates are low (less
than 33 percent), and per capita
recruitment is high (0.82–1.38).
However, the authors noted that their
mark–recapture methods were not
capable of providing separate estimates
of annual survival and out-migration,
yet these rates are critical in
understanding recruitment processes for
the species. They noted future studies
are needed to obtain quantified
recruitment data using alternative
methods, such as biotelemetry and
known-fates modeling approaches.
Schueller and Peterson (2010)
concluded that future studies of
subadult and adult life stages are
needed, but quantified assessment of
river resident juveniles can provide
fisheries managers with the data
necessary for evaluating population
trends.
The statement in the proposed listing
rule that spawning populations are less
than the 500 recommended by
Thompson (1991) as a minimum viable
population size for long-term
population fitness does not conflict with
the statement that total population
abundances for the Carolina and South
Atlantic DPSs are not available. As we
stated in this response, we do not have
direct estimates of total population
numbers for any of the Southeast
spawning populations. Based on data
from Schueller and Peterson (2006), we
were able to present an estimate of the
number of annually spawning adults in
the Altamaha River. Although survey/
stock assessment data on total
population numbers or annually
spawning adults are not available for the
remaining Southeast river populations,
based on information that the Altamaha
is the largest population in the
Southeast and data from the remaining
rivers, we estimate in comparison that
the other spawning populations have no
more than 300 spawning adults per
year.
In response to the comment that based
on observations and modeling, the
Winyah Bay system and ACE Basin have
more individuals than Thomas (1990)
recommended as minimum viable
population sizes for short-term and
long-term population fitness, we note
that Thomas (1990) offered a population
size of 5,500 as ‘‘a useful goal,’’ but
suggested that where uncertainty
regarding a species’ population
dynamics, changing environmental
conditions, and the species’ reaction to
the changing environmental conditions
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is extreme (as it is for Atlantic sturgeon)
‘‘we should usually aim for population
sizes from several thousand to ten
thousand.’’ Information provided for the
Winyah Bay and ACE Basin does not
provide an estimate of total population
size in either system. Because annual
spawning adults was the only
quantitative population metric we had
for any southern river population at the
time of the proposed listing, we looked
at estimated annual spawning adult
population sizes in comparison to
various viable population sizes
suggested in the literature. We now have
additional information on juvenile
abundance in the Altamaha River and
some preliminary modeling of juvenile
abundance in the Edisto River; however,
this information is lacking for most river
systems, and the population trends are
not certain from the data we have.
Although the Carolina and South
Atlantic DPSs, made up of multiple
river populations of Atlantic sturgeon,
were determined to be interbreeding
population units, the vast majority of
Atlantic sturgeon return to their natal
rivers to spawn, with fewer than two
migrants per generation spawning
outside their natal system. We looked at
the number and size of each riverine
spawning population within each DPS
when considering the effects of a small
population size on the extinction risk
for the DPS as a whole. We do not
believe that information presented by
the commenters provides a basis to
revise our evaluation of the status of the
Carolina and South Atlantic DPSs of
Atlantic sturgeon.
Comment 23: Several commenters
stated that historical commercial
landings do not accurately reflect
abundance and are not a good indicator
of status. One commenter stated that
Secor (2002) should not be used as the
basis for estimating historical
abundances of Atlantic sturgeon. The
commenter stated that due to the
nearshore location of the fisheries in the
latter part of the 19th century, the data
would include Atlantic sturgeon from
multiple populations and represent a
gross overestimate of historical
abundance. A comment was received
that population modeling should have
been used to analyze the trajectory of
the species.
Response: Section 4(b)(1)(A) of the
ESA states that the Secretary shall make
listing determinations solely on the
basis of the best scientific and
commercial data available. Historical
abundance data is not available.
However, we believe that the historical
landings data and the sharp downward
decline observed in landings throughout
the 20th century are a valid indicator of
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the declines in abundance experienced
by Atlantic sturgeon. Secor (2002)
represents the best available data on the
estimated historical abundances of
Atlantic sturgeon, as does the U.S. Fish
Commission data on historical landings,
which the Secor (2002) publication was
based on and which we reviewed
ourselves for clarification in preparing
the making our listing determinations.
We agree that it is impossible to
conclusively determine whether
historical landings data potentially
represents Atlantic sturgeon from
multiple river systems and multiple
DPSs. In the proposed listing rule, we
reported historical abundances of
Atlantic sturgeon from Secor (2002) as
state-wide estimates of spawning
females for North Carolina, South
Carolina, and Georgia. Though not
stated directly in the proposed listing
rule, this infers that multiple river
populations and DPSs are represented
in these estimates, since each state
contains multiple river systems, both of
the DPSs in the Southeast encompass
multiple states, and in the case of South
Carolina, both DPSs include river
populations originating in that state.
Therefore, our use and presentation of
the data in the proposed listing rule was
appropriate and not inconsistent with
the commenter’s statement. Due to the
lack of data (e.g., abundance,
recruitment, natural mortality, bycatch
mortality) on Atlantic sturgeon
throughout most of the species’ range,
reliable population modeling at the
species/DPS level is not possible.
However, as detailed in the proposed
listing rule, we believe that the
trajectory observed in the commercial
landings from the late 19th century
through the 20th century, combined
with information from recent and
ongoing surveys of Atlantic sturgeon
populations and information on threats
to the species from habitat modification
(e.g., dams, dredging, water quality and
quantity) and bycatch clearly
demonstrates that Atlantic sturgeon
population abundances have shown
little improvement since their initial
declines and continue to face a degree
of threat that warrants listing the
Carolina and South Atlantic DPSs of
Atlantic sturgeon as endangered.
Comment 24: A commenter stated that
the 1990–2003 increasing trend in
Atlantic sturgeon abundance in the
Cape Fear River should not have been
discounted in the status review.
Response: We did not discount
information in the proposed listing rule
on trends in Atlantic sturgeon
abundance in the Cape Fear River
between 1990 and 2003, as reported by
Moser et al. (1998) and Williams and
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Lankford (2003). We presented different
interpretations of the data that the
researchers noted themselves in their
research publications. In the proposed
listing rule, we stated ‘‘abundance of
Atlantic sturgeon below Lock and Dam
#1 in the Cape Fear River seemed to
have increased dramatically during the
1990–1997 surveys (Moser et al., 1998),
as the CPUE of Atlantic sturgeon was up
to eight times greater during 1997 than
in the earlier survey years. Since 1997,
Atlantic sturgeon CPUE doubled
between the years of 1997 and 2003
(Williams and Lankford, 2003).
However, it is unknown whether this is
an actual population increase reflecting
the effects of North Carolina’s ban on
Atlantic sturgeon fishing that began in
1991, or whether the results were
skewed by one outlier year. There was
a large increase observed in 2002,
though the estimates were similar
among all other years of the 1997 to
2003 study.’’ The commenter stated that
the 2007 status review report should not
have discounted the increase in
sturgeon abundance in 2002 as an
outlier year for the reason that it was a
flood year. Williams and Lankford
(2003) stated that CPUE is used to
indicate a population size, but if
environmental conditions affect the
susceptibility of fish to being captured
in gillnets, then the data may show a
change in population size when
environmental conditions actually
caused the change in CPUE. Williams
and Lankford (2003) further stated that,
‘‘although previous years have
documented relatively similar catchper-unit-efforts, the summer of 2002
yielded twice the CPUE of any season
since 1997. This also happens to be the
lowest flow conditions experienced
during this survey. Although catch-perunit-effort increased greatly during
these low flow conditions, previous
years with low flow summers did not
have the same resulting increases in
CPUE. Future surveys should
investigate river flow and other
environmental conditions that may
impact the Atlantic sturgeon’s use of the
Lower Cape Fear River.’’ The
researchers acknowledged ambiguity in
whether these results represent
increases in Atlantic sturgeon
abundance or whether environmental
conditions affected CPUE. Therefore,
the information we presented in the
proposed listing rule on trends in the
Cape Fear River is consistent with what
the researchers presented. Further, even
if the data in the Cape Fear River do
represent an actual increase in Atlantic
sturgeon, data provided by NCDENR
during the public comment period on
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the proposed listing rule did not show
increasing trends in Atlantic sturgeon
populations in other North Carolina
rivers.
Comment 25: A commenter asked if
the historical data on pounds of Atlantic
sturgeon landed in South Carolina (page
61907 of the proposed listing rule) can
be converted to CPUE.
Response: The majority of the
landings data for South Carolina
referred to by the commenter cannot be
converted into CPUE. However, the data
were taken from Smith et al. (1984),
which did provide CPUE for the time
period 1973 to 1982, and provided
anecdotal data about the level of fishing
effort for earlier time periods. The
objective of the research conducted by
Smith et al. (1984) was to obtain
baseline information on the Atlantic
sturgeon fishery in South Carolina. At
the time their research commenced,
South Carolina accounted for 55 percent
of the total U.S. landings of Atlantic
sturgeon, but little information on the
characteristics of the fishery was
available. Figure 2 in Smith et al. (1984)
shows license data for the Atlantic
sturgeon fishery in South Carolina.
From 1960 to 1982, the number of
fishermen licensed for sturgeon
remained relatively constant, averaging
21 individuals (ranging between 15 and
30) per year. Smith et al. (1984) noted
that fishermen possessing certain other
types of fishing licenses (e.g., a shrimp
fishing license) were permitted to fish
for Atlantic sturgeon without having a
specific sturgeon license. Based on field
observations, they estimated that there
were two to three times the number of
recorded licensed sturgeon fishermen
active in these fisheries. No data on the
amount of gear fished were available for
the period of most active exploitation of
the fishery (pre 1910), but from 1925 to
1970, the number of licensed units of
fishing gear was also relatively constant
and averaged 17.8 (ranging between 11
and 26). This suggests that landings data
are representative of relative abundance,
since fishing pressure remained
constant.
There was a dramatic increase in
fishing effort in the 1970 to 1982 time
period, with the number of licensed nets
at record levels for the time. The
number of licensed nets in 1970 was
less than 30, but by 1982, it was around
140. Smith et al. (1984) calculated CPUE
data for 1973 to 1982 based on reported
total landings and number of net
licenses, as well as field observations
and verbal information provided by
fisherman. They noted several
limitations of the license and landings
data for calculating CPUE: (1) Though
individual gear were required to be
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5931
licensed, the license was not based on
type or length; (2) the license data
included gear fished in the northern
(Winyah Bay) and southern (Edisto,
Coosawhatchie, and Combahee Rivers)
fisheries, whereas the landings data
only included fish from the northern
fisheries; (3) field observations
indicated that not all nets were licensed,
nor landings reported; and, (4) pre-1973
data included landings of shortnose
sturgeon in addition to Atlantic
sturgeon. Figures 6 and 7 in Smith et al.
(1984) show landings, effort, and CPUE.
Landings rose from about 20,000 to
42,000 kilograms (kg) between 1973 and
1982, while the number of licensed nets
increased from 36 to 133 during the
same time period, resulting in a slight
declining trend in CPUE (Figure 6).
Observations of fishermen on the
Winyah Bay jetties between 1978 and
1982 (Figure 7) also showed a decline in
CPUE during the time period, with
Atlantic sturgeon landings declining
even with effort increasing. Smith et al.
(1984) concluded that a definitive
analysis of the fishery was not possible
because of the limitations of the data,
but they stated that ‘‘fishing effort has
substantially increased without a
concomitant increase in landings’’, and
though ‘‘the fate of this fishery in South
Carolina is not clear, it appears likely
that intensive fishing effort will
adversely affect local populations of
these long-lived fish.’’
Comments on the 2007 Status Review,
the 1998 Status Review, and Difference
Between the Status Reviews and the
Proposed Listing Rule
Comment 26: Commenters disagreed
with NMFS’ proposal to list both of the
DPSs in the Southeast as endangered,
when the 2007 status review report
concluded that the Carolina DPS should
be listed as threatened and did not make
a listing conclusion for the South
Atlantic DPS due to lack of information
to allow a full assessment of
subpopulations within the DPS. Several
of these commenters stated that there
was no new scientific information
presented justifying the proposed listing
of the Carolina and South Atlantic DPSs
differently from the conclusions reached
in the 2007 status review report. Similar
comments were received that no new
data has been collected, and no changes
in the level of threats have been
documented, since the 1998 status
review, which concluded that listing
was not warranted at that time. One
commenter said the proposed listing
rule does not sufficiently explain why
the conclusion in the 1998 status review
report that the existing moratorium on
fishing for Atlantic sturgeon and the
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listing of the shortnose sturgeon was
adequate to protect Atlantic sturgeon is
no longer valid.
Response: Regarding comments about
divergence from the 2007 status review
report’s listing conclusions for the
Carolina and South Atlantic DPSs, see
our response to peer reviewer comment
1 above.
In 1998, NMFS and USFWS (Services)
determined that an ESA listing of
Atlantic sturgeon throughout its range
was not warranted at that time (63 FR
50187, September 21, 1998). The
Services cited eight reasons for the
negative determination: (1) Evidence
that the historical range of the species
has not been substantially reduced and
that its current range is not likely to be
significantly reduced in the foreseeable
future; (2) persistence of at least 14
spawning populations; (3) the expected
efficacy of existing prohibitions on
harvest and possession in all 15 states
comprising the species’ U.S. range; (4)
detailed evaluation of current habitat
conditions and threats to habitat
showing that conditions are adequate to
sustain the species and are likely to
remain so in the foreseeable future; (5)
lack of substantial information
indicating that overutilization for
commercial, recreational, scientific or
educational purposes is currently
significantly affecting the species; (6)
lack of information indicating that
disease or predation are causing
significant losses of individuals of the
species; (7) existing regulatory
mechanisms which provide adequate
protection and further the conservation
of the species; and, (8) lack of
information indicating that artificial
propagation is currently posing a threat
to the species.
Section 4(b)(1)(A) of the ESA requires
that listing decisions be made using the
best available scientific and commercial
information at the time of the decision,
after conducting a review of the status
of the species and considering the
conservation efforts of states and foreign
nations.
Information provided in the 2007
status review report and the 2010
proposed listing rule explain why we no
longer believe all of the eight
conclusions in the 1998 status review
report are valid, particularly as applied
to DPSs of Atlantic sturgeon.
Specifically: (1) Reductions in the
historical range of Atlantic sturgeon
have occurred, as evidenced by
extirpations of several spawning
populations in both Southeast DPSs and
limited access to historical river reaches
and habitats above dams (detailed in the
‘‘Distribution and Abundance’’ and
‘‘Conservation Status’’ sections of the
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proposed and final listing rules); (2) no
spawning populations in the DPSs are
large or stable enough to provide with
any level of certainty for the continued
existence of the DPS in the face of
threats currently acting on the species;
(3) existing prohibitions on harvest and
possession of Atlantic sturgeon in all
East Coast states do not alleviate other
significant threats to Atlantic sturgeon
(i.e., bycatch and habitat destruction/
modification, a point discussed in
further detail in the discussion on those
threats); (4) habitat destruction and
modification (from dams, dredging,
degraded water quality and quantity,
etc.) is a significant threat to Atlantic
sturgeon river populations and DPSs, as
discussed below and in our responses to
comments 39–45; (5) information on
overutilization of Atlantic sturgeon as
bycatch suggests that this is also a
significant threat to Atlantic sturgeon
populations, as discussed below and in
our responses to comments 46 and 47;
and, (7) existing regulatory mechanisms
have proven inadequate at controlling
the threats to Atlantic sturgeon from
habitat modification/destruction and
bycatch, as discussed in our responses
to comments 49 and 50. Evidence for
these conclusions and detailed
responses to the comments received on
these conclusions is presented in the
following text.
Comments stated that no new data has
been collected and no changes in the
level of threats have been documented
since the 1998 status review. However,
studies not available at the time of the
1998 status review report on bycatch
(discussed here) and habitat quality
(discussed later in this section) have
been reviewed by NMFS as part of our
current listing determination. The 1998
status review report determined that
estimated levels of mortality associated
with bycatch on the Delaware and
Hudson Rivers indicated that bycatch
was not a significant threat to the
species survival but could impede
recovery, and recommended that efforts
be made to better quantify data on
bycatch levels, fishing effort, and river
population levels to ensure that
assumptions made using Hudson and
Delaware River information are valid for
other river populations. Since 1998, the
ASMFC (2007) produced a bycatch
report providing estimates of Atlantic
sturgeon bycatch, as did Stein et al.
(2004), a bycatch report used by the
2007 ASSRT. The reports documented
mean bycatch mortality rates of 13.8
percent and 22 percent, respectively.
However, the ASMFC (2007) report
noted that the estimates of bycatch
utilized in the analysis are likely to be
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underestimates of true bycatch and
mortality levels, since they rely only on
reported bycatch from the observer
program (there is limited observer
coverage in fisheries potentially
capturing Atlantic sturgeon in Federal
waters from North Carolina to Florida),
and delayed mortality is not accounted
for in their estimates. Further, the 1998
status review report did not consider the
effects of bycatch and degraded habitat
working in combination on greatly
reduced Atlantic sturgeon populations,
which are at 1 to 6 percent of historical
levels.
In response to the comments that the
proposed listing rule does not
sufficiently explain why the conclusion
in the 1998 status review report that the
existing moratorium on fishing was
adequate to protect Atlantic sturgeon is
no longer valid, at the time of the 1998
determination, we note that the ASMFC
moratorium on retention of Atlantic
sturgeon had recently gone into effect.
Because this eliminated directed fishing
for Atlantic sturgeon (NMFS followed
this with a 1999 closure of the EEZ to
fishing for Atlantic sturgeon), which
was considered the primary threat to the
continued existence of the species at the
time, the moratorium factored heavily in
the Services’ decision not to list the
species at the time. However, since
implementation of the moratorium,
additional bycatch information (Stein et
al., 2004; ASMFC, 2007) has become
available and indicates that Atlantic
sturgeon are vulnerable to bycatch in
commercial fisheries, and that the
current rate of bycatch is unsustainable
in the long term (ASMFC, 2007).
Further, the proposed listing rule
described in detail why the existing
moratorium on directed capture of
Atlantic sturgeon has not eliminated the
incidence of sturgeon bycatch in other
fisheries and also does not address
threats associated with the destruction
and modification of their habitat.
Comments were also received that the
proposed listing rule does not
sufficiently explain why the conclusion
in the 1998 status review report that the
listing of the shortnose sturgeon was
adequate to protect Atlantic sturgeon is
no longer valid. While Atlantic sturgeon
have benefited from some of the
protections afforded the endangered
shortnose sturgeon due to their shared
presence in some rivers, shortnose
sturgeon do not coexist in all rivers
within the Atlantic sturgeon’s range and
shortnose sturgeon do not use the
coastal and marine environments used
extensively by Atlantic sturgeon.
Additionally, there is often spatial and
temporal separation of riverine habitat
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use by the Atlantic and shortnose
sturgeon. Adults of both species use
similar habitats for spawning in the
riverine environment, but they are
known to use them at slightly different
times of the year. As stated in the 1998
recovery plan for the shortnose
sturgeon, spawning begins in freshwater
from late winter/early spring in
southern rivers. The 2007 Atlantic
sturgeon status review report stated that
spawning adults generally migrate
upriver in the spring/early summer
(February to March in southern
systems). Further, the 2007 Atlantic
sturgeon status review report noted that
other life stages of Atlantic and
shortnose sturgeon may use different
sections and/or different depths within
the same river system. Therefore, the
threats facing each species are not
identical and protections for shortnose
sturgeon cannot be expected to fully
alleviate threats to Atlantic sturgeon.
Comment 27: Several comments were
received on differences in the 1998 and
2007 status reviews in the evaluations of
the threat to Atlantic sturgeon from
habitat modification. A commenter
noted that the 1998 status review report
denied the petitioner’s claims that dam
blockages, degraded water quality, and
dredging significantly contributed to
low Atlantic sturgeon abundances, but
NMFS has not provided any evidence
supporting a reversal of this conclusion.
Another commenter specifically asked
what changed between the 1998 and
2007 status reviews to warrant the
‘‘moderately high (4)’’ ranking of threats
from dams on the Cape Fear River in the
2007 status review. The commenter also
asked if the recommendations on page
91 of the 1998 status review report have
been followed. The commenter
requested we provide the baseline data
on spawning and nursery habitat,
including locations, depths, flows,
substrates, carrying capacity or optimal
population, that was recommended as
‘‘contributing to and accelerating the
ongoing recovery or enhancement of
Atlantic sturgeon’’ in the 1998 status
review. Several commenters also cited
the 1998 status review’s statements that
water quality has been improving since
the 1970s, dredging activities are
increasingly rare and have minimal
effects on sturgeon, and successful
shortnose restoration is indicative of
future rebounding of Atlantic sturgeon
stocks. One of the commenters
referenced Table 9 in the 2007 Status
Review, which shows a 2004 U.S.
Environmental Protection Agency
(USEPA) water quality grade in the
Southeast as ‘‘B’’, then questioned the
extinction risk ratings in Table 13 of the
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2007 Status Review, which rates water
quality in most of the Southeast rivers
as having a moderate risk of causing
extinction.
Response: In reaching our 1998 not
warranted determination, we did not
consider the loss of habitat due to dams
to be a significant threat. Page 31 of the
status review report states, ‘‘In the
southern region of the U.S. Atlantic
coast, the fall line is commonly much
farther inland (322 river kilometers or
rkm on the Savannah River, South
Carolina-Georgia border) or almost
nonexistent (St. Johns River, Florida).
This potentially provided more
freshwater (spawning) habitat than in
many northern rivers. However,
historical records of the amount of
habitat actually used by Atlantic
sturgeon are lacking. Thus, for most
rivers, it is not possible to determine
how much habitat was lost due to dam
construction for southern rivers.’’ As
stated above, the 1998 analysis included
the amount of spawning habitat
available to the species across its range.
Since that time, we have determined the
amount of habitat lost on each of the
rivers due to dams (see Table 7 of the
2007 status review). We also have
additional information on spawning
locations for some rivers. The 1998
status review report cited the Savannah
River as an example of a river with a fall
line far inland and the 2007 status
review report also stated that 92 percent
of the habitat on the Savannah is
unimpeded by dams. While both of
these facts are true, the historical
primary spawning habitat for Atlantic
sturgeon (and only shoal habitat on the
Savannah River), the Augusta Shoals, is
not accessible to Atlantic sturgeon
because it lies above the New Savannah
Bluff Lock and Dam (Wrona et al., 2007;
Marcy et al., 2005; Duncan et al., 2003;
USFWS, 2003). Regarding the comment
on the ranking for the Cape Fear River,
the 1998 status review report did not
have an estimate for how much
spawning habitat was blocked by Lock
and Dam #1. The 2007 status review
report included the following
information and provides insight into
the ‘‘moderately high’’ ranking for the
threat of dams on that river (page 51):
‘‘Historical spawning locations are
unknown in the Cape Fear River;
therefore, it is assumed that the fall line
is the upper limit of spawning habitat.
Using the fall line as guide, only 33
percent of the historical habitat is
available to Atlantic sturgeon (96 km of
292 km). In some years, the salt water
interface reaches the first lock and dam;
therefore, spawning adults in the Cape
Fear River either do not spawn in such
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5933
years or spawn in the major tributaries
of the Cape Fear River (i.e., Black River
or Northeast Cape Fear rivers) that are
not obstructed by dams.’’
Dredging activities are far from rare.
NMFS routinely conducts section 7
consultations on listed species for
dredging projects within the range of
Atlantic sturgeon. Statistics on hopper
dredging, the form of dredging most
likely to take aquatic species (such as
sea turtles and Atlantic sturgeon), can
be found on the U.S. Army Corps of
Engineers’ (USACE) ‘‘Sea Turtle Data
Warehouse’’ Web site (https://el.erdc.
usace.army.mil/seaturtles/index.cfm).
The Charleston, Jacksonville, Savannah,
and Wilmington Districts have
completed 307 hopper dredging
projects, removing over 220 million
cubic yards of material from federally
maintained navigation channels in 307
projects since 1991. The number of
private dredging projects permitted by
USACE would increase that number
considerably. Further, these numbers do
not include other dredging methods
(e.g., cutterhead and mechanical) used
by Federal and private entities that are
less likely to directly interact with
sturgeon species, but can modify and
degrade sturgeon habitat.
While water quality has generally
improved since the 1970s due to
numerous Federal, state, and local laws,
including the Clean Water Act of 1972,
water quality continues to be an issue
for Atlantic sturgeon due to human
population expansion and a variety of
agricultural, industrial, and commercial
activities in the coastal zone. Table 9 in
the 2007 status review report cites the
USEPA’s National Coastal Condition
Report (NCCR) II (2005) in grading the
Southeast water quality as a B. The
NCCR II also assigned water quality a
numerical score of 4 (where 1 is poor
and 5 is good), ranking it as ‘‘good to
fair.’’ It is important to note that the
water quality index in the NCCR II was
based on a combination of several
parameters, the most important of
which to Atlantic sturgeon is dissolved
oxygen (DO). The DO range considered
‘‘good’’ in the NCCR II was greater than
5 mg/L while a DO range of 2 to 5
mg/L was considered ‘‘fair.’’ As stated
in the proposed listing rule, sturgeon are
more highly sensitive to low DO than
other fish species and ‘‘low’’ DO was
defined as less than 5 mg/L (Niklitschek
and Secor, 2009a, 2009b). A DO of 2 mg/
L (the lower end of the ‘‘fair’’ scale in
the NCCR II report) would be
considered very poor for an Atlantic
sturgeon, likely lethal to early life stages
(Niklitschek and Secor, 2009a, 2009b;
Niklitschek and Secor, 2005; Secor and
Gunderson, 1998). The USEPA
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published the NCCR III in 2008 and
downgraded water quality in the
Southeast from a 4 to a 3, ranking it as
‘‘fair’’ rather than ‘‘good to fair.’’ It also
showed that the portion of the Southeast
that had a ‘‘poor’’ water quality index
ranking increased slightly from 5
percent to 6 percent. While other
condition indicators for the Southeast in
the NCCR III showed improvement over
the NCCR II levels (the benthic index
was upgraded from a 3 to a 5 in the
Southeast) or remained the same (the
coastal habitat index remained a 3), the
sediment quality index was downgraded
from a 4 to a 3, and the fish tissue
contaminant index was downgraded
from a 5 to a 4. This resulted in a
decrease from 3.8 to 3.6 in the overall
condition of the Southeast. The results
of the NCCR III report do not support
the commenters’ assertion that water
quality has continually improved since
the 1970’s. Water quality was
downgraded to ‘‘fair’’, and DO levels
included under a ‘‘fair’’ rating may be
less than adequate for Atlantic sturgeon,
particularly early life stages. Further,
the percentage of geographic areas in the
Southeast with ‘‘poor’’ water quality
increased between NCCR II and III.
NMFS and other partners involved in
the conservation of Atlantic sturgeon
(such as the ASMFC, USFWS, and state
agencies) continue to work on
monitoring, research, and other
activities, including those outlined in
the 1998 status review, to recover
Atlantic sturgeon. However, these are
long-term, ongoing efforts, and the
objectives outlined in the 1998 status
review report are not complete. We do
not have all of the data requested by the
commenter, but what is available is
included in the 2007 status review, the
proposed listing rule, and the references
cited therein. Once Atlantic sturgeon are
listed, NMFS will have a greater
opportunity to prioritize and
standardize Atlantic sturgeon research,
as recommended for recovery and
conservation of Atlantic sturgeon in the
1998 status review.
Comment 28: Several comments
stated that the 2007 status review report
was developed with little or no input
from state agency experts and that state
agency comments should be weighed
heavily. One commenter stated that the
results of the 2003 workshop that
preceded the 2007 status review report
were not publicly available.
Response: Eight state and regional
experts from six state agencies provided
their individual expert opinions on the
information contained in the 2007 status
review report and provided additional
data to ensure the status review report
included the best available. Many of the
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comments, data, and information
presented in this document originated
from state agencies. As stated in the
status review report and the proposed
listing rule, information obtained at the
2003 workshop prompted the initiation
of the status review. Information from
the workshop was incorporated into the
2007 status review. In addition, the
2003 workshop was held in conjunction
with a meeting of the ASMFC Atlantic
Sturgeon Technical Committee and
some of the proceedings of the
workshop are published in various
meeting summaries, reports, and
documents on the ASMFC’s Atlantic
sturgeon Web site (https://
www.asmfc.org).
Comments on the Need To List Atlantic
Sturgeon Under the ESA
Comment 29: Comments were
received stating that Atlantic sturgeon
should not be listed because their
populations are stable, sufficiently large,
and/or increasing. Commenters cited to
Grunwald et al. (2008) for statements
that the Altamaha and Edisto appear to
have large, multiple year class
populations that exhibit high annual
reproductive success. The State of
Georgia commented that, in order to list
as threatened or endangered under the
ESA, the condition must exist in ‘‘all or
a significant portion of its range.’’ The
commenter stated the Altamaha River
represents a significant portion of the
South Atlantic DPS’s range due to the
large population of Atlantic sturgeon in
that river and the area of the watershed.
They also stated populations are
persisting in other systems, and
therefore, they do not believe Atlantic
sturgeon are threatened or endangered
throughout a significant portion of their
range. Other commenters stated that
Atlantic sturgeon have been observed in
most South Carolina coastal rivers
during the last two decades, although it
is not known if all rivers support a
spawning population. Currently, the
only long term data set available for
Atlantic sturgeon in South Carolina is
on the Edisto River, where the Atlantic
sturgeon population seem to be
relatively stable based on fishery
independent sampling efforts by the
South Carolina Department of Natural
Resources. A commenter stated that
abundance and distribution presented
in the proposed listing rule is
inconclusive, citing increasing
incidental take in Albemarle Sound gill
nets, increases in average length of
Atlantic sturgeon captured off North
Carolina between 1986 and 2003,
suspected spawning activity on the Pee
Dee River during the Fall of 2003, and
the doubling of CPUE of Atlantic
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sturgeon from annual surveys
conducted in the Cape Fear River
between 1997 and 2003. One
commenter stated that for the Savannah
River, conclusions were incorrectly
drawn in the proposed listing rule that
the greater catch of shortnose sturgeon
than Atlantic sturgeon, as cited in
Collins et al. (1996), was not a reflection
of lower than expected catch of Atlantic
sturgeon, but rather that they were
fishing in areas/habitat not preferred by
juvenile Atlantic sturgeon.
Response: The information presented
by commenters stating that Atlantic
sturgeon should not be listed does not
provide a basis for revising our
proposed listing rule determination of
endangered for the Carolina and South
Atlantic DPSs of Atlantic sturgeon.
Grunwald et al. (2008) stated that
‘‘among southeastern populations, those
in the Altamaha (Peterson et al. in press)
and Edisto appear to be large, with
multiple year classes and high annual
reproductive success.’’ Grunwald et al.
(2008) continued that ‘‘others range
from small (Ogeechee and Savannah) to
possibly extirpated (Satilla).’’ This is
consistent with information we
presented in the proposed listing rule
that, at the 2003 workshop, we
determined some populations seemed to
be recovering while others were
declining, prompting our initiation of
the 2007 status review. This comment is
also consistent with our description in
proposed listing rule of the Altamaha
population as larger and more robust
than other populations in the Southeast.
We received information from SCDNR
(presented in the previous section of
comments) that they have captured
3,661 Atlantic sturgeon in the Edisto
since 1994. If all of these were spawning
adults, then this represents an average
of approximately 230 spawning adults
per year since 1994, which is consistent
with our estimate of less than 300
spawning adults per year for this
system. The low number of annually
spawning adults estimated for Atlantic
sturgeon in the Southeast (343 for the
Altamaha River and less than 300 for
the remaining spawning populations)
factored heavily in our determination
that the Carolina and South Atlantic
DPSs warranted listing as endangered.
In the proposed listing rule, we did not
define which rivers constitute a
significant portion of the species’ range
because we concluded that the Carolina
and South Atlantic DPSs are endangered
throughout their entire ranges. The
presence of multiple spawning
populations does not negate the need for
listing. As discussed above, we do not
believe that any of the riverine
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populations within either DPS, alone or
in combination, are viable and stable
enough to constitute a significant
portion of either DPS’s range.
We acknowledged uncertainty in the
abundance and distribution information
we presented. However, we believe that
a conservative evaluation of the
information the commenter referred to
as ‘‘inconclusive’’ supports our
endangered listing determination. As
the commenter noted, we stated that
catch records for Albemarle Sound, as
well as the Roanoke River, indicate that
this population seemed to be increasing
until 2000, when recruitment began to
decline. We also indicated the existence
of catch records and observations from
other river systems in North Carolina
(e.g., the Tar, Neuse, and Cape Fear
Rivers), but, based on the relatively low
numbers of fish caught, we stated it was
difficult to determine whether the
populations in those systems are
declining, rebounding, or remaining
static. However, the fact alone that low
numbers of fish were caught does not
logically lead to a conclusion that
populations are increasing. The
commenter’s interpretation of data on
increases in average length of Atlantic
sturgeon caught off North Carolina
between 1986 and 2003 (from Laney et
al., 2007) is incorrect. While Figure 5 in
Laney et al. (2007) showed an increase
in average length of fish caught from
1988 to 2006, the commenter suggested
this is due to a reduction in commercial
harvest of larger sturgeon. Commercial
harvest of Atlantic sturgeon was
completely prohibited in 1999. The
trend of increasing size was linear over
the full time period and the rate of
increase showed no association with the
time period during which the
moratorium was active. Laney et al.
(2007) did not draw any conclusions
about the increase in average size over
the study period. However, they did
conclude from the length data that all
but five of the Atlantic sturgeon
captured were juveniles. They attributed
the low numbers of adults to either the
age distribution of the population (i.e.,
low numbers of adults in the population
because of pre-moratorium fishing) or
the ability of adults to more successfully
evade capture in nets. As we discussed
in our response to a previous comment,
it is possible that the increases in
Atlantic sturgeon observed in the Cape
Fear River surveys were due to
environmental conditions rather than
actual population increases. As we also
stated above, the same data the
commenter states shows an increased
population in the Cape Fear River
would have to be interpreted to show no
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increase in Atlantic sturgeon in other
North Carolina Rivers, and as previously
stated, neither DPS can be judged not in
danger of extinction based on any single
river population within the DPS.
We do not agree that we incorrectly
interpreted the lower catch of Atlantic
versus shortnose sturgeon in the
Savannah River, as reported in Collins
et al. (1996). Researchers conducted
surveys in both the lower river (rkm 45–
75) and upper river (rkm 160–299). No
Atlantic sturgeon were captured in the
upper river, while 14 Atlantic sturgeon
(and 189 shortnose) were captured in
the lower Savannah River. As stated in
Collins et al. (1996), juvenile Atlantic
sturgeon in the size range likely to be
captured in the shad fishery (and the
size range observed in this study) occur
in estuarine and tidally influenced
portions of the river. According to the
New Georgia Encyclopedia, the
Savannah River is tidally influenced up
to Clyo, Georgia, 61 miles (98 rkm)
upriver. Therefore, the lower river study
area was within the area Collins et al.
(1996) expects juvenile Atlantic
sturgeon to occur.
Comment 30: Several commenters
recommended that NMFS implement
alternative actions instead of listing
Atlantic sturgeon. One commenter
suggested that NMFS designate Atlantic
sturgeon as a Species of Concern and
conduct another status review in 2017.
Some commenters believed that, in lieu
of listing Atlantic sturgeon, NMFS
should enter into multi-state, multiagency partnerships to obtain the
information they believe is necessary to
support management actions. A
commenter specifically requested that
we provide information on any
cooperative efforts NMFS is engaged in.
One commenter suggested that
increased fishing regulations, including
the development of habitat reserves, as
well as area and seasonal closures, are
warranted instead of listing. Another
commenter stated that NMFS should
expand the 1965 Anadromous Fish
Conservation Act in order to protect
Atlantic sturgeon and said that
imposing a listing is a poor substitute
for restoring habitat and water quality.
Response: We made our proposed
listing determinations for the Carolina
and South Atlantic DPSs of Atlantic
sturgeon by carefully analyzing the
declines in population abundance,
available information on the current
status of riverine spawning populations,
and the threats facing the species, and
whether their status or the threats are
adequately addressed by existing
regulatory mechanisms or protective or
conservation mechanisms. Though
moratoria on harvest and possession of
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5935
Atlantic sturgeon were enacted by the
ASMFC, NMFS, and several states,
populations have not rebounded and the
moratoria do not control bycatch. We
believe continued overutilization of
Atlantic sturgeon from bycatch in
commercial fisheries is an ongoing
impact to the Carolina and South
Atlantic DPSs that is contributing to
their endangered status. Because
Atlantic sturgeon mix extensively in
marine waters and may access multiple
river systems, they are subject to being
caught in multiple fisheries throughout
their range. While some of the threats to
the Carolina and South Atlantic DPSs
have been reduced through the existing
regulatory mechanisms, such as the
moratoria on directed fisheries for
Atlantic sturgeon, bycatch is currently
not being addressed through existing
mechanisms. Further, water quality
continues to be a problem even with
existing controls on some pollution
sources and water withdrawal, and
dams continue to curtail and modify
habitat, even with the Federal Power
Act. Since our evaluation of the
Carolina and South Atlantic DPSs
resulted in our determining that both
DPSs are in danger of extinction
throughout their ranges (i.e., meet the
definition of endangered), we cannot list
the DPSs as threatened or continue to
designate Atlantic sturgeon as a species
of concern.
Section 4(b)(3)(B) of the ESA requires
us to make a finding within 12 months
of receiving a petition as to whether the
petitioned action is warranted. Section
4(b)(6)(A) of the ESA requires that a
final listing determination be made
within 12 months of publication of the
proposed listing rule. Because we
received a petition to list Atlantic
sturgeon from the Natural Resources
Defense Council (NRDC) on October 6,
2009, that established mandatory
deadlines under the ESA for
determining whether listing of the
species is warranted, and for associated
rules. As described above, the best
available scientific and commercial
information on the status of, and threats
to, Atlantic sturgeon is sufficient to
warrant listing of the Carolina and
South Atlantic DPSs of Atlantic
sturgeon as endangered under the ESA.
Therefore, listing cannot be postponed,
and we cannot enter into multi-state,
multi-agency partnerships or increase
fishery regulations to address Atlantic
sturgeon conservation issues in lieu of
listing. However, once listed, fishery
regulations, such as the development of
habitat reserves or seasonal/area
closures, could be considered as a
means to reduce threats to Atlantic
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sturgeon from being taken as bycatch. If
this was determined to be necessary for
the conservation of Atlantic sturgeon, it
would be done in collaboration with all
the stakeholders, including the affected
fishing community.
We currently work with multiple
agencies in multiple states to improve
our knowledge of the species and to
enhance conservation efforts. In fact,
our efforts and exchange of knowledge
with our multi-agency, multi-state
partners factored into our decision that
listing the Carolina and South Atlantic
DPSs of Atlantic sturgeon as endangered
is warranted. In 1999, pursuant to
section 804(b) of the Atlantic Coastal
Fisheries Cooperative Management Act
(16 U.S.C. 5101 et seq.), we supported
the ASMFC’s moratorium on Atlantic
sturgeon by closing the Exclusive
Economic Zone (EEZ) to Atlantic
sturgeon retention. In 2003, we
sponsored a workshop with the USFWS
and ASMFC to discuss the status of
sturgeon along the Atlantic Coast and
determine what obstacles, if any, were
impeding their recovery. Based on the
information gathered from the 2003
workshop, we decided that a new
review of Atlantic sturgeon status was
needed to determine if listing as
threatened or endangered under the
ESA was warranted. The ASSRT was a
multi-agency team consisting of NMFS,
USFWS, and USGS biologists. Also, as
described in the example given in the
response above, we have entered into
multi-state, multi-agency partnerships
to conduct research.
The projects described in the previous
response to document seasonal and
spawning migrations of sturgeon,
identify interbasin migrations, develop
genetic aging techniques, and map
habitat were all funded through the
Species Recovery Grants Program
(‘‘section 6 program’’) in 2010. Section
6 of the ESA provides a mechanism for
cooperation between NMFS and states
in the conservation of threatened,
endangered, and candidate species.
Under section 6, NMFS is authorized to
enter into agreements with any state that
establishes and maintains an ‘‘adequate
and active’’ program for the
conservation of endangered and
threatened species. Once a state enters
into such an agreement, NMFS is
authorized to assist in, and provide
Federal funding for, implementation of
the state’s conservation program.
Federal funding, provided in the form of
grants, can be used to support
management, outreach, research, and
monitoring projects that have direct
conservation benefits for listed species,
recently delisted species, and candidate
species that reside within that State.
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Each of the states occupied by the
Carolina and South Atlantic DPSs has a
section 6 agreement with NMFS. In
addition to the multi-year, multi-state,
multi-agency projects funded in 2010,
various research projects by multiple
agencies involving the Carolina and
South Atlantic DPSs were funded
through the section 6 program in prior
years (NMFS, 2009), including
evaluations by GADNR of Atlantic
sturgeon populations and habitat in the
Altamaha River (2003, 2004, and 2006)
and the St. Mary’s and Satilla Rivers
(2008), and studies by SCDNR of
Atlantic sturgeon growth, diet, and
genetics (2003, 2005).
The Anadromous Fish Conservation
Act (AFCA) of 1965 is another source of
collaboration between Federal and state
partners. Projects funded under this act
are conducted for the conservation,
development, and enhancement of
anadromous fishery resources and must
be cleared with the fishery agency of the
state that the work is carried out in.
Many projects funded under AFCA are
critical elements of larger programs to
manage, restore, or enhance
anadromous resources. In 1998, SCDNR
was awarded $176,837 for a 3-year
project to collect life history data on
juvenile Atlantic sturgeon and
determine seasonal habitat utilization,
movements, and growth. SCDNR was
also awarded $116,926 in 2001 for a 3year period to continue work on the
previous project funded through the
AFCA, as well as look at the effects of
fisheries, such as shad gillnet fisheries,
on sturgeon. Research publications
resulting from these projects were
evaluated in the proposed listing. AFCA
funding for research in the Southeast
Region is generally around $104,000 per
year, though the program has not
received funding for the past 3 years.
We do not believe the listing of the
Carolina and South Atlantic DPSs is a
substitute for restoring habitat and water
quality. Rather, the need to list the two
DPSs of Atlantic sturgeon in the
Southeast as endangered highlights the
need to restore water quality and their
habitat, because as we outlined in the
proposed listing, habitat modification
and poor water quality are significantly
contributing to the endangered status of
Atlantic sturgeon.
Comment 31: Commenters both
supporting and opposed to the proposed
listing believed that additional
information on Atlantic sturgeon, such
as abundance, movement, life history
information, habitat usage, response to
threats, etc., is necessary. Commenters
supporting the proposed listing believed
this information is important to address
threats to the species and determine
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recovery actions. Fisheries and Oceans
Canada provided information on current
Atlantic sturgeon studies planned or
underway and expressed their interest
in exploring potential areas of
collaboration to enhance our mutual
understanding of Atlantic sturgeon.
Commenters opposed to the proposed
listing believed that NMFS should not
pursue listing before more information
on abundance, movement, genetics,
threats, etc., is obtained. A comment
was received that NMFS is proposing
listing the Atlantic sturgeon without
dedicating funding to collecting
necessary information on the species.
Some commenters believed that a final
listing determination should be
postponed until the results of recently
commenced studies on Atlantic
sturgeon are available. Several
commenters also stated that NMFS
should implement the measures listed
in the 1998 amendment to the ASMFC’s
FMP for Atlantic sturgeon and address
the monitoring and data needs in it
before making a listing determination.
Response: As described in the
proposed listing rule and in the
previous response, section 4(b)(3)(B) of
the ESA requires us to make a finding
within 12 months of receiving a petition
as to whether the petitioned action is
warranted, on the basis of the best data
available at the time of the
determination. Because we determined
the Carolina and South Atlantic DPSs of
Atlantic sturgeon warranted listing as
‘‘endangered,’’ we published a proposed
listing rule in the Federal Register. The
ESA requires that we publish final
listing rules within one year from the
date that we publish proposed rules to
list species. The best available scientific
and commercial data on the historical
declines of Atlantic sturgeon, the
species’ failure to rebound even with
the prohibition on directed captured
and possession, the information on the
status of current spawning populations,
the information on the level of threats
to the species from bycatch, habitat
modification and curtailment, and the
failure of existing regulatory
mechanisms to protect the species
indicate that listing of the Carolina and
South Atlantic DPSs of Atlantic
sturgeon as endangered under the ESA
is warranted. Therefore, we cannot
postpone a listing determination until
the results of recently commenced
studies are available. However, we agree
with commenters that additional
information on Atlantic sturgeon
concerning abundance, movement, life
history information, habitat usage, and
response to threats is critical to fully
recovering the species.
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Section 4(b)(1)(A) of the ESA requires
us to make listing determinations solely
on the basis of the best scientific and
commercial data available, whether
research funding for the species is
available or not. However, while
Atlantic sturgeon were considered a
‘‘species of concern’’ and a candidate
species, NMFS dedicated funding to
Atlantic sturgeon in order to gain
knowledge necessary for conservation
and recovery of the species. NMFS is
currently funding a multi-year, multistate, multi-agency project to document,
through telemetry, seasonal and
spawning migration patterns and
incidences of inter-basin transfer for
adult Atlantic (and shortnose) sturgeon
in southeastern rivers in North Carolina,
South Carolina, and Georgia, as well as
develop, test, and implement a genetic
aging technique. We are also funding
research to map habitat in four Georgia
rivers that will complement this study,
as it overlaps with the area where the
telemetry work is being conducted.
These studies also address components
of the monitoring and data needs
outlined in the ASMFC’s Atlantic
sturgeon FMP. We will continue to
conduct and fund Atlantic sturgeon
research as funds become available in
the future. We look forward to working
with the ASMFC, Fisheries and Oceans
Canada, our state partners, and other
stakeholders in the conservation and
recovery of Atlantic sturgeon, including
obtaining the necessary research to fill
in the gaps in our knowledge.
Comment 32: One commenter stated
that NMFS relied on non-peer reviewed,
agency-based opinion rather than
scientific fact and stated that future
management steps would also be driven
by conjecture rather than science.
Another commenter stated that the
proposed listing rule was politically
motivated instead of scientifically
warranted. A comment was received
that NMFS is rushing to list Atlantic
sturgeon to gain leverage in FERC
relicensing activities underway, such as
the Santee-Cooper Hydroelectric Project.
Response: Section 4(b)(1)(A) of the
ESA requires us to make listing
determinations solely on the basis of the
best scientific and commercial data
available, not conjecture or political
motivation. However, the ESA’s best
available data standard does not require
us to limit the information we consider
to published, peer-reviewed scientific
literature. Our listing determination is
consistent with the Services’
Interagency Cooperative Policy on
Information Standards (59 FR 24271;
July 1, 1994). The majority of the
literature cited in the status reviews and
the proposed listing rule consists of
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peer-reviewed publications. As required
by the regulations and agency policy for
implementing the ESA and by the
Information Quality Act, status reviews
and listing decisions themselves are
peer reviewed. The proposed listing rule
for the Carolina and South Atlantic DPS
was peer reviewed by three experts. The
list of peer reviewers, with their
affiliations, and the peer review
comments in their entirety, are posted at
www.regulations.gov and https://www.
cio.noaa.gov/Policy_Programs/prplans/
ID184.html. Our responses to the peer
review comments are stated in this
document. NMFS is not rushing to list
Atlantic sturgeon to gain leverage in
FERC relicensing activities; as discussed
previously, section 4(b) of the ESA
dictates strict timelines for making
determinations and publishing rules in
response to a petition to list a species
as threatened or endangered.
Comments on the Consequences of the
Proposed Listing Rule
Comment 33: Several comments were
received stating that listing will not
eliminate the impacts to Atlantic
sturgeon (e.g., it will not result in the
removal of locks and dams).
Response: The commenters are correct
that listing will not eliminate all
impacts to Atlantic sturgeon. However,
section 4(b)(1)(A) of the ESA states that
the Secretary shall make listing
determinations solely on the basis of the
best scientific and commercial data
available to him after conducting a
review of the status of the species and
taking into account efforts to protect the
species. Based on our review of the best
available information on the status of
the Carolina and South Atlantic DPSs of
Atlantic sturgeon and the efforts
currently in place to protect the DPSs,
we concluded that both DPSs should be
listed as endangered. Our reasoning is
outlined in the proposed listing rule and
supplemented by our responses to the
public comments in this document.
While listing a species does not
automatically remove all threats, the
ESA does provide tools for greater
protection of listed species. When this
final rule takes effect, the prohibition on
‘‘take’’ in section 9 of the ESA will
apply. Also, any action funded,
authorized, or undertaken by a Federal
agency that may affect Atlantic sturgeon
from either DPS will require
consultation between that Federal
agency and NMFS under section 7 of the
ESA. Once listed, section 4 of the ESA
also requires that we develop and
implement a recovery plan that must, in
part, identify objective, measurable
criteria which, when met, would result
in a determination that the species may
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be removed from the list; this standard
inherently requires that recovery plans
propose methods to address impacts
and threats to the species. In the
example given by the commenter for
locks and dams, during section 7
consultation, NMFS would work with
the operating and/or authorizing agency
(e.g., USACE or FERC) to minimize the
effects on Atlantic sturgeon and their
habitat. This could result in a variety of
conservation measures to allow passage
of Atlantic sturgeon upstream of the
lock or dam and to control any
downstream effects from the structures.
The installation of fish passage, dam
breaching, and even lock/dam removal
have been undertaken in the past to
restore natural flows and allow access to
habitat for anadromous species.
Comment 34: Comments were
received stating there will be negative
consequences to various stakeholders
associated with the listing. One
commenter stated the Federal listing
would increase regulations and
potentially affect parties that do not
have significant impacts on Atlantic
sturgeon survival. Several commenters
stated that there will be impacts to
fisheries if additional restrictions are
placed on them due to the listing, even
if the interactions with Atlantic
sturgeon do not cause significant
mortality. Several comments were
received that the listing process will
hold up the issuance of new FERC
licenses in the range of the two DPSs,
which contain measures that they
believe would benefit sturgeon.
Commenters were concerned that there
will be impacts to commerce if ship
strikes result in speed restrictions and
could be more far-reaching than the
right whale protection zone.
Commenters also believed there will be
further restrictions on dredging, such as
at large ports, that could have economic
consequences for ports and commercial
shipping interests. Commenters
suggested that the cost to Federal and
state entities associated with increased
permitting needs should be considered
in the listing. Another commenter stated
that NMFS is using the listing to force
regulators to impose requirements on
third parties (e.g., hydropower
licensees) through the ESA consultation
process, and the impact will affect
society for decades. The commenter said
that the costs of recovery should be
leveled equitably among all parties,
including NMFS, by allocating funding
to collecting data needed for
management. One commenter stated
that economics should not be
considered in the listing.
Response: As explained in the
response above, section 4(b)(1)(A) of the
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ESA states that the Secretary shall make
listing determinations based solely on
the best scientific and commercial data
available to him after conducting a
review of the status of the species and
taking into account efforts to protect the
species. The regulations implementing
the ESA at 50 CFR 424.11(b), consistent
with case law interpreting the ESA and
its legislative history, state that the
listing determination will be made
without reference to possible economic
or other impacts of such determination.
We cannot consider the potential
consequences (e.g., increased economic
costs or regulatory responsibilities) to
the various stakeholders in our listing
determination. Through the ESA section
7 consultation process, measures to
reduce the effect of impacts on Atlantic
sturgeon may be required for federally
funded or permitted projects that
adversely affect fish from the Carolina
or South Atlantic DPS, but the listing
will not affect entities or activities that
do not affect Atlantic sturgeon. NMFS is
not using the listing to force regulators
to impose requirements on third parties
(e.g., hydropower facility licensees) and
we are working with FERC to ensure
that the listing of Atlantic sturgeon does
not hold up the issuance of new
licenses. For example, where we had
already been engaged in section 7
consultation regarding a proposed
relicensing’s effects on the endangered
shortnose sturgeon, we began
‘‘conference consultations’’ on the
effects of such projects on Atlantic
sturgeon once the species was proposed
to be listed. Such ‘‘conference opinions’’
can be promptly adopted without
reinitiating consultation on a project, if
a species’ listing is finalized as
proposed. The listing determination,
prompted by the 2007 status review
report and the 2010 NRDC petition, is
based solely on the status of the species
and its current level of protection from
impacts and threats.
NMFS currently dedicates funding to
the recovery of listed species (and
species of concern) through a variety of
channels; we provide funds to the
NMFS Science Centers, to academic
institutions, and our state partners
doing research. We currently have a
multi-state effort to tag and track
Atlantic sturgeon, and a simultaneous
habitat mapping project in a portion of
the area where the tagging/tracking is
occurring, funded through our ESA
section 6 grant program (Species
Recovery Grants). However, successful
recovery of the species will require the
actions of entities other than NMFS.
Section 4(f)(2) of the ESA states that the
Secretary, in developing and
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implementing recovery plans, may
procure the services of appropriate
public and private agencies and
institutions, and other qualified
persons. Section 7(a)(1) of the ESA
charges all Federal agencies to utilize
their authorities in furthering the
purposes of the ESA by carrying out
programs for the conservation of
threatened and endangered species.
Recovery may also be facilitated through
incorporating conservation measures
into activities that potentially affect
Atlantic sturgeon, for example, through
section 7(a)(2) consultation and section
10(a)(1)(B) permitting. Those processes
provide a means to tailor the required
conservation measures to the severity of
an activity’s impacts.
Comment 35: Many commenters had
concerns over the time lag in getting
research permits to study Atlantic
sturgeon if they are listed as
endangered. Other commenters said that
in addition to creating a lengthy
research permitting process, listing will
lead to sampling constraints that would
invalidate long established sampling
protocols and will terminate long-term
indices of abundance, as a change in the
survey protocol is essentially the
initiation of a new survey. Several
commenters stated that the listing will
abolish all efforts presently being
undertaken to study the Atlantic
sturgeon, including research on captive
Atlantic sturgeon and studies conducted
by other Federal agencies, such as
USACE. One commenter suggested that
these issues be taken into account in
deciding whether to proceed with
listing as endangered versus threatened.
In addition to concerns over Atlantic
sturgeon research, commenters also
expressed concerns over impacts to
other fishery survey and sampling
programs that may encounter Atlantic
sturgeon, as these would also require
permitting. Commenters also expressed
concern over the ability to
opportunistically collect data from
incidental captures of Atlantic sturgeon
if they are listed as endangered. Several
commenters expressed concern that the
increased permitting workload
associated with an Atlantic sturgeon
listing would also cause a greater delay
in obtaining permits to conduct research
on other species, such as the shortnose
sturgeon.
Response: As explained in the
responses above, we cannot consider the
potential consequences to stakeholders,
including those conducting research on
Atlantic sturgeon that aids in the
management and conservation of the
species, in making listing
determinations. However, NMFS is
making every effort to ensure that
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Atlantic sturgeon research, including
ongoing care and study of captive fish,
can continue uninterrupted once they
are listed. Section 10(a)(1)(A) of the ESA
allows NMFS to issue permits
authorizing activities otherwise
prohibited by section 9 of the ESA for
the purpose of scientific research on
listed species. The NMFS Office of
Protected Resources, Permits Division
contacted known Atlantic sturgeon
researchers, at the time the proposed
listing rule published in the Federal
Register, requesting information on
planned research activities so that an
expedited permitting process could be
put in place. Twelve applications for
research permits for Atlantic sturgeon
have been received and are undergoing
review, and the steps necessary to
comply with the National
Environmental Policy Act and section
7(a)(2) of the ESA are already underway.
Section 10(a)(1)(B) of the ESA allows
NMFS to issue permits authorizing
incidental take of listed species during
the course of otherwise legal activities,
such as fishery survey and sampling
programs targeting species other than
Atlantic sturgeon. If the activities are
Federal actions, section 7 consultations
can also provide incidental take
authorization.
In March 2010, NMFS published ‘‘A
Protocol for Use of Shortnose, Atlantic,
Gulf, and Green Sturgeons’’ (Kahn and
Mohead, 2010; available at https://www.
nmfs.noaa.gov/pr/pdfs/species/kahn_
mohead_2010.pdf). Section 10 permits
will likely require that the protocol be
followed during Atlantic sturgeon
research. The goal of the protocol is
standardization of research practices to
benefit the recovery of sturgeon species,
including the Atlantic sturgeon, while
also minimizing potentially negative
impacts of research.
These protocols were developed from
a comprehensive review of the best
available scientific information at the
time of publication, including peer
reviewed journals, technical
memoranda, interviews with
researchers, and empirical evidence
provided by researchers. Some
researchers expressed concern that
sampling constraints associated with
such a protocol would invalidate longestablished sampling protocols and will
terminate long-term indices of
abundance. However, the protocol was
developed with input from researchers
and will serve to standardize research in
the future. Any variation from previous
research methods can likely be
accounted for when comparing results.
It is common in research, including
Atlantic sturgeon research, for methods
and equipment to evolve as experience
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and technology in the field of research
grows. Further, there is flexibility built
into the protocol. For example, the
introduction to the document states,
‘‘When researchers or managers have
reason to exceed recommendations in
this document using less known or
riskier techniques, NMFS recommends
first using surrogate Acipenserids or
hatchery-reared sturgeon. When
researchers or managers feel nonrecommended methods must be
conducted on wild listed or candidate
species, the researchers should consult
with the appropriate permitting agency
in order to justify why their
methodology is necessary to provide
information for the recovery of these
species.’’
Comment 36: Flagler County, Florida,
commented that they do not believe
Atlantic sturgeon or habitat supporting
sturgeon exists in their county and
requested that they be excluded from
regulatory jurisdiction. Oconee County,
Georgia, requested an exemption for
previously permitted public water
supply projects.
Response: Section 4(b)(5)(a)(ii) of the
ESA requires that we notify each county
where Atlantic sturgeon are believed to
occur and invite their comment.
Because we do not know all of the exact
locations where Atlantic sturgeon may
occur, and to ensure all counties
potentially affected by the proposed
listing were contacted, we used a GIS
database to generate a list of all counties
within the watersheds of rivers with
current or historical spawning
populations of Atlantic sturgeon. This
resulted in over 200 counties for the
Carolina and South Atlantic DPSs.
Flagler County, Florida, is part of the St.
Johns River watershed. The St. Johns
River is used by Atlantic sturgeon as
nursery habitat. We realize that not all
of the counties we contacted have
Atlantic sturgeon present; however,
upstream projects can have effects on
Atlantic sturgeon downstream, and we
chose to be more inclusive to give
adequate opportunity for
communication between NMFS and
potentially affected counties. Moreover,
Atlantic sturgeon may reoccupy areas of
their former ranges once their
populations begin to recover, or when
impediments to their migration are
removed. Areas where Atlantic sturgeon
do not exist and where activities that
could potentially affect Atlantic
sturgeon, directly or indirectly, are not
occurring, will not be affected by the
listing of Atlantic sturgeon. We cannot
grant exemptions for projects that may
affect Atlantic sturgeon once they are
listed as endangered. Oconee County
did not state whether they believe their
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permitted water supply projects will
have effects on Atlantic sturgeon. Once
listed as endangered, we will work with
such entities to protect Atlantic
sturgeon while still carrying out the
purpose of their projects, such as
providing water to the public.
Comments on Our Analysis of Threats
Comment 37: One commenter stated
that the extinction risk analysis assigns
arbitrary risk values to the level of threat
an activity poses for Atlantic sturgeon
populations in each river on a scale of
1 to 5. The commenter believed
statistically sound information would be
difficult to derive from this analysis
when used to determine the status of a
species under the ESA.
Response: We believe this comment
misinterprets the purpose and utility of
the extinction risk analysis contained in
the Atlantic sturgeon status review.
However, that risk analysis was not
determinative to our proposed listing
because we did our own independent
extinction risk analysis, which we
determined was required to be
consistent with the ESA. The ASSRT
characterized their extinction risk
analysis as a ‘‘semi-quantitative’’
approach. It is not possible, nor did the
ASSRT or NMFS ever intend, to
conduct statistical analyses on the
results of the extinction risk analysis
contained in the status review. Further,
the status review report clarifies that the
intent of the extinction risk assessment
was to help summarize the status of the
species, and did not represent a
decision by the ASSRT on whether the
species should be proposed for listing as
endangered or threatened under the
ESA. In our proposed listing rule, we
considered the information contained in
the ASSRT’s extinction risk analysis as
part of our listing determination.
However, we also considered additional
threats (e.g., drought, water allocation
issues, and climate change) not
considered by the ASSRT. In addition to
evaluating the threats to the species, we
considered the effects of small
population size on the risk of extinction
of Atlantic sturgeon DPSs. We compared
estimated Atlantic sturgeon abundances
with minimum viable population sizes
discussed in relevant literature (see
‘‘Conservation Status’’ section in the
proposed listing rule).
Comments on Habitat Threats
Comment 38: Commenters supporting
the proposed listing rule emphasized
that Atlantic sturgeon are vulnerable to
habitat destruction, noting sensitivity to
low DO, pollution, and river-specific
threats from dams, dredging, and
development, and a summary of their
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comments are included here. Several
commenters noted that the Cape Fear
River is above permissible mercury
limits and all 13,123 waters in North
Carolina are in Category 5 (waters
impaired for one or more designated
uses by a pollutant(s)) on the state’s
2010 303(d) list (the list of impaired and
threatened waters that section 303(d) of
the Clean Water Act requires all states
to submit to the USEPA) for mercury
due to statewide fish consumption
advisories. Several commenters also
provided NMFS with information that a
proposed cement plant on the Cape Fear
River is requesting authorization to emit
263 pounds (119 kg) per year of mercury
and discharge 10–15 million gallons of
water a day (mgd). One commenter cited
an analysis by a marine chemist that
conditions are favorable in the Cape
Fear estuary to convert the mercury to
more dangerous forms. The chairman of
the North Carolina Marine Fisheries
Commission identified the proposed site
of the plant as a spawning area for
Atlantic sturgeon and five other
diadromous species. Commenters also
provided information on habitat threats
from other proposed projects, such as
the Cape Fear Skyway and the North
Carolina International Container
Terminal. A commenter encouraged
further studies on the effect of toxins on
all Atlantic sturgeon life stages.
Comments were also received
supplementing information in the
proposed listing rule on concentrated
animal feeding operations (CAFOs).
While there is a moratorium in North
Carolina limiting hog operations, a
commenter noted this does not apply to
the poultry industry, which is greatly
expanding in the state and poses a
significant water quality threat. The
commenter listed two processing plants,
one in the Neuse River basin and one
undergoing permitting in the TarPamlico basin, that are driving the
establishment of poultry CAFOs and
will result in increased nitrogen and
phosphorus loading in coastal waters.
Another commenter, citing NCDENR
Department of Water Quality (DWQ) as
the source of information, reported the
decline of 1,600 freshwater miles (50
percent of the total freshwater miles) in
the Neuse River basin and indicated that
runoff is a contributing factor. They
further cited NCDENR DWQ that this is
likely an underestimate of the true
number of miles affected by nonpointsource runoff. The commenter also
noted the ecological and water quality
benefits from undisturbed riparian
buffers and noted many instances in the
coastal counties where construction of
bulkheads and other shoreline
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stabilization activities has resulted in
the partial or complete loss of riparian
buffers. Comments were received that in
1999, 60 percent of surface water tested
in Georgia was too polluted for fishing,
swimming, or drinking compared to
national average of 40 percent. The
Savannah River was reported to be the
fourth most toxic river in the U.S., with
48 industrial outfalls over a 200 mile
stretch from Augusta to Savannah.
Comments included that the river has
high levels of mercury, low DO is likely
to worsen if the harbor deepening
project is approved, and temperature is
also a challenge, as cold water from the
J. Strom Thurmond Dam is discharged
75 feet below the lake surface,
disrupting the natural temperature
regime. Though the proposed listing
rule noted that water quality in the
Altamaha is relatively good, a
commenter provided information that
19 rivers and streams making up 192
miles of the Altamaha basin were on the
2002 303(d) list as not meeting their
designated uses. This is an area
dominated by silviculture and
agriculture, two paper mills, and
numerous other dischargers.
Information provided included that a
Federal Superfund site is contributing
chemicals (including mercury). A May
2009 report noted lesions on fish in the
river, linked to poor water quality and
bacteria present in floodwaters. A
commenter also noted the St. Mary’s
River is much warmer than the 70–75
degrees Fahrenheit ideal for sturgeon (it
reaches the 90s), DO levels drop to less
than 2 parts per million at times, and of
the coal power plants on the river, half
report releasing water in the summer
months (when high temperature and
low DO already a problem) at peak
temperatures of 100 degrees Fahrenheit
or more. The commenter concluded
thermal pollution can stress or kill any
fish present, and will be exacerbated by
poor water quality conditions in these
rivers.
Response: Additional information
provided by commenters on the threats
posed by the destruction, modification,
or curtailment of Atlantic sturgeon
habitat is consistent with our finding
that it poses a significant threat to the
Carolina and South Atlantic DPSs. We
will continue to work with our partners
and stakeholders through our existing
authorities to reduce or eliminate the
adverse effects of anthropogenic
activities on sturgeon and their habitat.
Comment 39: Comments stated that
water quality information presented in
the proposed listing rule was overly
generalized and should receive a more
comprehensive review. Some
commenters stated that water quality is
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good and/or improving, and disagreed
that water quality is affecting Atlantic
sturgeon. A commenter stated that after
more than 30 years of water quality
improvements associated with the Clean
Water Act, it is unreasonable to think
habitat is not of good quality. Another
commenter stated that water quality has
been improved through existing Federal
and state regulations and programs,
such as the mandate to implement water
quality improvement programs that are
consistent with Total Maximum Daily
Load (TMDL) plans. The Cape Fear
River was used as an example; the
comments maintained that while certain
areas are impaired, other areas used by
Atlantic sturgeon are in excellent
condition and fully support sturgeon
life functions, and the NCDENR DWQ’s
water quality data should be used.
NCDENR DWQ also submitted
comments on the proposed listing rule,
providing benthic macroinvertebrate
data for 1983 to 2010 and stated that
data from 12 river segments are fair to
excellent. Commenters also said fish
kills are not a good indicator of water
quality, as reporting varies by year and
location. One commenter stated that
NMFS failed to identify water quality
issues in the Cooper River, the Santee
River meets state DO standards greater
than 96 percent of the time, and NMFS
did not acknowledge increased
minimum flows associated with the new
Santee-Cooper license. One commenter
stated that NMFS did not present a
substantive analysis concerning the
sensitivity of sturgeon to water quality
parameters relative to water quality
conditions that currently exist in
‘‘critical habitat areas.’’ The commenter
provided a literature summary on
Atlantic sturgeon sensitivity to DO,
temperature, and salinity in the Cape
Fear River and a water quality database
from the Cape Fear River Estuary
Program and stated that an assessment
of these data would provide information
on the spatial and temporal distribution
of various sturgeon life stages,
sensitivities, and the likelihood of
exposure to potentially adverse water
quality conditions.
Response: As stated in our response to
comment 27 on the 1998 and 2007
status reviews, while water quality has
generally improved since the 1970s due
to numerous Federal, state, and local
laws, including the Clean Water Act of
1972, water quality continues to be an
issue for Atlantic sturgeon due to
human population expansion and a
variety of agricultural, industrial, and
commercial activities in the coastal
zone. The USEPA publishes the
National Coastal Condition Report and
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the NCCR II, published in 2005, graded
the Southeast’s water quality as a B. The
NCCR II also assigned water quality a
numerical score of 4 (where 1 is poor
and 5 is good), ranking it as ‘‘good to
fair.’’ The USEPA published the NCCR
III in 2008. It downgraded water quality
in the Southeast from a 4 to a 3, ranking
it as ‘‘fair’’ rather than ‘‘good to fair.’’ It
also showed that the portion of the
Southeast that had a ‘‘poor’’ water
quality index ranking increased slightly
from 5 percent to 6 percent. While other
condition indicators for the Southeast in
the NCCR III showed improvement over
the NCCR II levels (the benthic index
was upgraded from a 3 to a 5 in the
Southeast) or remained the same (the
coastal habitat index remained a 3), the
sediment quality index was downgraded
from a 4 to a 3, and the fish tissue
contaminant index was downgraded
from a 5 to a 4. This resulted in a
decrease from 3.8 to 3.6 in the overall
condition of the Southeast. It is also
important to note that the water quality
index in the NCCR is based on several
parameters, the most important of
which to Atlantic sturgeon is DO. The
DO level included within the ‘‘good’’
rating in the NCCR II was greater than
5 mg/L, while a DO range of 2 to 5 mg/
L is included in the ‘‘fair’’ rating. As
stated in the proposed listing rule,
sturgeon are more highly sensitive to
low DO than other fish species and
‘‘low’’ DO for sturgeon has been defined
as less than 5 mg/L (Niklitschek and
Secor, 2009a, 2009b). A DO of 2 mg/L
(the lower end of the ‘‘fair’’ scale in the
NCCR II report) would be considered
very poor for an Atlantic sturgeon and
is likely lethal to early life stages. The
USEPA also monitors TMDLs, a
calculation of the maximum amount of
a pollutant that a waterbody can receive
and still safely meet water quality
standards. Under section 303(d) of the
Clean Water Act, states, territories, and
authorized tribes are required to
develop lists of impaired waters. These
are waters that are too polluted or
otherwise degraded to meet the water
quality standards set by states,
territories, or authorized tribes. Based
on 2006 to 2010 data, each of the states
in the range of the Carolina and South
Atlantic DPSs had impaired waters
under section 303(d) of the Clean Water
Act: Florida (828 waterbodies), Georgia
(215 waterbodies), South Carolina
(1,060 waterbodies), North Carolina (902
waterbodies), and Virginia (2,534
waterbodies). Of the rivers and streams
assessed, 51 to 66 percent of these
waters were impaired in each of the
southeastern states. Between 24 and 84
percent of the lakes, reservoirs, and
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ponds assessed in each southeastern
state were listed as impaired, as were 22
to 95 percent of bays and estuaries
assessed. In the Cape Fear River basin,
the example used by the commenter,
205 sections of the river are listed as
impaired on the 303(d) list. As
suggested by the commenter, we
reviewed water quality information
from NCDENR DWQ. We reviewed the
most recent Water Quality Plan (October
2005) available for the Cape Fear River
basin (publicly available at https://h2o.
enr.state.nc.us/basinwide/draftCPFApril
2005.htm), which supplements the
TMDL data provided by the USEPA.
The plan (Chapter 27, Figure 31)
indicates ‘‘habitat degradation’’ and low
DO occur in over 140 miles of impaired
streams. Low DO is also estimated to
occur in approximately 6,500 acres of
impaired estuarine waters (Chapter 27,
Figure 32). Figures 37 and 38 note
various sources of stressors to streams
and estuarine waters, respectively.
While wastewater treatment, municipal
stormwater, agriculture, land clearing,
development, and impervious surfaces
are listed as potential sources, the
largest source affecting water quality in
impaired streams and estuarine waters
in the Cape Fear River basin is
‘‘unknown.’’
NCDENR DWQ commented on the
proposed listing rule, as well. They
stated that a review of benthic
macroinvertebrate data from the Cape
Fear mainstem demonstrates that the
river is supporting robust benthic
invertebrate communities. Benthic
invertebrate communities serve as prey
for foraging Atlantic sturgeon. NCDENR
DWQ stated that 6.2 percent of the
samples received ‘‘excellent’’
bioclassifications, and 31.2 percent each
received ‘‘good’’, ‘‘good to fair’’, and
‘‘fair’’ bioclassifications. There were no
samples receiving ‘‘poor’’
bioclassifications. However, with the
exception of one sample collected in
2003, the remaining samples were
collected in the 1980’s and 1990’s. Also,
benthic invertebrate communities are
only one of the many factors affecting
the quality and suitability of habitat for
Atlantic sturgeon. Regarding NCDENR
DWQ’s comment that fish kills were not
a good indicator of water quality and
that some of the fish kills on the Cape
Fear River are likely due to naturally
occurring low DO from blackwater
swamps, we also reported this in the
proposed listing rule. The comment that
fish kill reporting varies by year and
location, and is not a good indicator of
water quality, is also consistent with our
treatment of fish kill information in the
proposed listing rule. We did not
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compare fish kill information across
river systems with varying degrees of
monitoring and reporting effort, rather
we only included fish kill data as
anecdotal evidence of naturally
occurring low DO in the lower Cape
Fear River.
With regard to habitat modification
and curtailment in the Santee-Cooper
system, the majority of the discussion in
the proposed rule focused on the threats
to Atlantic sturgeon from dams. The
Clean Water Act 303(d) list of impaired
waters includes 21 waterbodies within
the Santee River basin and 34
waterbodies within the Cooper River
basin. The commenter stated that the
Santee River meets state DO standards
greater than 96 percent of the time but
did not provide data or a reference we
could evaluate. The list of 303(d) waters
in the Santee River basin lists 19
waterbodies that are listed as a result of
low DO. We also reviewed the South
Carolina State Water Assessment of the
Santee River Basin, prepared by SCDNR
(2009), which lists 9 waterbodies that
are partially supporting of aquatic life
and 19 waterbodies that are nonsupporting of aquatic life, based on DO.
The new license for the Santee-Cooper
Hydroelectric project has not yet been
issued, therefore the magnitude and
timing of implementation of required
increased minimum flows is unknown
at this time. Significant concerns still
exist over the inability of Atlantic
sturgeon to access over 60 percent of
historical habitat in the Santee-Cooper
system due to the presence of the dams,
though this would be partially
ameliorated by fish passage for sturgeon
that was prescribed in 2007 by NMFS
for the Santee and Cooper Rivers
pursuant to the Federal Power Act, if
these prescriptions are implemented.
A commenter stated that we did not
present an analysis of water quality in
critical habitat areas. NMFS has not
designated critical habitat, but the
proposed listing rule and responses
supplied in this document detail water
quality conditions and potential effects
of reduced water quality in habitat used
by the Carolina and South Atlantic DPSs
of Atlantic sturgeon. The literature
summary on Atlantic sturgeon
sensitivity to DO, temperature, and
salinity in the Cape Fear River and a
water quality database from the Cape
Fear River Estuary Program is consistent
with information in the proposed listing
rule. The literature reviewed by the
commenter was also cited in the 2007
status review report and/or the
proposed listing rule.
Comment 40: A commenter stated that
silviculture and forest manufacturing
facilities do not appear to have
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5941
significant implications for sturgeon or
their habitat, particularly when
compared to other land uses like
agriculture or development. The
commenter supplied information on
forestry best management practices,
sedimentation, the use of herbicides,
and urged NMFS to reconsider its
assertion that forest management
practices pose a significant threat to
biological diversity or to habitat for the
Atlantic sturgeon. The commenter
asserted that water quality has improved
and will continue to improve through
existing Federal and state regulations
and program. The commenter also
stated that implementation rates for
forestry best management practices
(BMPs) are high nationally, and there is
an extensive body of scientific literature
that confirms that forestry BMPs are
effective. The commenter also indicated
that state agencies and sustainable
forestry certification programs are
effective at educating the forest
management community about forestry
BMPs and encouraging their
implementation, and providing
reasonable assurance that forestry BMPs
are being implemented effectively. The
commenter concluded that sustainable
forest management that adheres to BMPs
does not pose a threat to terrestrial or
aquatic organisms, including Atlantic
sturgeon.
Response: The proposed listing rule
included silviculture and forestry
practices as potential threats to Atlantic
sturgeon. The proposed listing rule
stated that the spawning habitat of the
Carolina DPS occurs within the MidAtlantic Coastal Plain ecoregion. The
Nature Conservancy lists land
conversion (e.g., forests converted to
timber plantations) as one of several
significant threats in the ecoregion. The
South Atlantic DPS occurs within the
South Atlantic Coastal Plain ecoregion.
The Nature Conservancy described the
primary threats to biological diversity in
this ecoregion as silvicultural practices,
including conversion of natural forests
to highly managed pine monocultures
and the clear-cutting of bottomland
hardwood forests. The proposed listing
rule also noted that in the Altamaha
River, which has the largest spawning
population of Atlantic sturgeon in the
Southeast, water quality is good at this
time, but the drainage basin is
dominated by silviculture and
agriculture, with two paper mills and
over two dozen other industries or
municipalities discharging effluent into
the river. While we agree that some
existing programs are effective,
degraded water quality continues to
pose a threat to Atlantic sturgeon in
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many systems despite existing
regulatory mechanisms.
We appreciate the information
provided by the commenter on the
degree of threat to Atlantic sturgeon
from forestry activities, as well as
forestry BMPs and the efforts of the
industry to ensure successful BMP
implementation, including education
and monitoring. However, we do not
believe that our characterization of the
potential threat of forestry practices to
Atlantic sturgeon was overemphasized
or overstated in the proposed listing
rule, or was inconsistent with
information provided by the
commenter. While we do not disagree
with the comments regarding the
effective implementation of forestry
BMPs, we note that implementation of
the BMPs is voluntary in some cases,
and that while BMP implementation
nationally is high (89 percent), it is not
100 percent. The commenter also stated
that implementation rates for BMPs can
be used to understand trends and
identify areas where improvement is
necessary; however, BMP evaluations
are detailed reports of many on-site
practices, are designed to highlight
potential problems for post-harvest
monitoring, and are not a direct measure
of water quality impact. We look
forward to working with the commenter
and other industry representatives to
proactively evaluate and address
forestry impacts on Atlantic sturgeon.
Comment 41: We received multiple
comments supporting our evaluation of
the effects of dams on Atlantic sturgeon
and their habitat; some commenters
provided additional information on the
nature of the threat of dams to Atlantic
sturgeon. A commenter concerned about
the effects of dams on Atlantic sturgeon
recommended continued investigation
into ways to provide fish passage in
areas where barriers obstruct access to
essential habitat or where passage is
otherwise obstructed in a manner that
can injure and/or kill Atlantic sturgeon
and noted that effective sturgeon
passage does not exist. Another
commenter provided NMFS with
additional information on threats from
dams. For example, the commenter
detailed the effects of bed coarsening,
which can reduce the ability of Atlantic
sturgeon to forage for food, impair
nutrient and waste assimilation through
altered flow regimes and greater
evaporation from the presence of
reservoirs, and effect biodiversity as a
result of habitat loss. The commenter
also provided data on the presence of
dams in Georgia, which has the highest
density of dams in the Southeast. The
commenter provided information that
the number of dams listed in the
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National Dam Inventory (NDI) shows
4,423 reservoirs in Georgia but the
actual number is believed to be higher
based on studies conducted by UGA,
which estimates 68,000 reservoirs in
Georgia. The commenter stated that
American Rivers named the Altamaha
the 7th most endangered river in the
country based on its importance to
fisheries and multiple threats from five
proposed dams that would have severe
effects on fish species, including loss of
habitat and increased pollutant
concentrations, and noted that the
governor of Georgia urged legislative
action to build new reservoirs. The
commenter also noted that the
Savannah River is impacted by New
Savannah Bluff Lock and Dam and J.
Strom Thurmond Dam. The latter is the
largest reservoir east of the Mississippi
and Atlantic sturgeon are blocked from
habitat above Augusta where data shows
they previously occurred. The
commenter also noted loss of habitat
from dams in the St. Johns.
Response: Additional information
provided by commenters on the threats
posed by dams to Atlantic sturgeon and
their habitat is consistent with our
finding that dams pose a significant
threat to the Carolina and South
Atlantic DPSs.
Comment 42: We received multiple
comments disagreeing with our
evaluation of the effects of dams on
Atlantic sturgeon and their habitat. A
commenter stated that the proposed
listing rule failed to indicate the extent
to which Atlantic sturgeon access to
habitat has been lost on the Roanoke,
Tar-Pamlico, and Neuse River systems,
all of which have dams. Other
commenters disagreed with the
evaluation of dams in the proposed
listing rule. One stated that the majority
of Atlantic sturgeon habitat is available,
as 91 percent of historical spawning
habitat is unimpeded by dams, 27 of 35
rivers contain 100 percent of their
historical habitat (e.g., Pee Dee River),
and 32 have over 75 percent of the
historical range available. Another
commenter stated that NMFS has not
evaluated the quality of the remaining
91 percent of habitat available to
Atlantic sturgeon. One commenter
questioned whether the estimated 64
percent of historical habitat impeded by
Lock and Dam #1 on the Cape Fear was
accurate and provided his own estimate
of 30 percent. A comment was received
that the use of watershed miles as the
measure of habitat loss due to dams
suggests that the entire river system is
critical habitat and any reduction is a
reduction in sturgeon habitat. The
commenter contended that since critical
habitat has not been determined or
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designated, it is presumptuous to
assume every portion of the river is
appropriate habitat without an analysis
or evaluation. The commenter also
believed that the proposed listing rule
gave undue weight to restoration of
these habitats rather than prioritizing
actions that would have significant and
immediate benefits to Atlantic sturgeon
(e.g. reducing bycatch). A similar
comment was received that NMFS has
placed too much emphasis on restoring
historical habitat, which is poorly
defined and may be of questionable
importance to Atlantic sturgeon. The
commenter believed that there are lower
costs and larger near-term gains in
protecting, mitigating, and enhancing
currently accessible habitat than trying
to reconnect historical habitat in highly
developed and substantially modified
watersheds. Another commenter said
future habitat availability will increase
through fish passage efforts on the lower
Cape Fear River and through
hydropower flow enhancements on the
Pee Dee River, and similar flow
enhancements will occur on other rivers
through FERC relicensing projects. A
commenter stated that there is a lack of
knowledge about the exact location of
historical spawning habitat on the
Roanoke River. A commenter stated that
both the Gaston and Roanoke Rapids
hydroelectric facilities are located above
the fall line. Given that the Carolina
DPS is estimated to be less than 3
percent of the historical abundance and
lack of documentation of significant
spawning historically occurring
upstream of the fall line, it seemed
unlikely to this commenter that
restricted spawning habitat is limiting
restoration efforts. The commenter
stated that these hydroelectric facilities
have been modified to simulate more
natural flow during spawning season
and during the FERC relicensing,
measures to limit peaking operations
and enhance flows were put in place.
The commenter also said the facilities
adhere to North Carolina state water
quality standards for temperature and
DO except when flood control flows
from upstream at the Kerr Dam
overwhelm their ability to maintain the
water quality standards. A commenter
stated that the Cape Fear Lock and Dam
#1 has been in place since 1915 and
Atlantic sturgeon have obviously
adapted to it since they are still
spawning.
Response: In regard to the comment
that the proposed listing rule failed to
indicate the extent to which Atlantic
sturgeon access to habitat has been lost
on the Roanoke, Tar-Pamlico, and Neuse
River systems, Table 7 of the 2007 status
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review report estimates the percentage
of riverine habitat, in river kilometers,
available to Atlantic sturgeon in each
river system shows that access to 18
percent of the habitat on the Roanoke
River is blocked by the Roanoke Rapids
Dam. Table 7 shows no loss on the other
two rivers. The percentages of historical
habitat unimpeded by dams presented
by another commenter are mostly
consistent with Table 7 of the status
review. As documented in Table 7, the
91 percent of historical habitat available
to Atlantic sturgeon includes 36 rivers
(not 35), 2 of which are in Canada and
not included in the proposed U.S.
listings. Coast-wide, 25 U.S. rivers, plus
the 2 Canadian rivers, are listed as
having 100 percent of their historical
habitat accessible. As noted by the
commenter, an additional 5 rivers have
greater than 75 percent of their river
miles unimpeded by dams. However,
three rivers in the Southeast have 62 to
64 percent of their length inaccessible to
sturgeon due to the presence of dams.
Moreover, rivers without dams but
without spawning populations present,
may not provide habitat to sturgeon for
decades; because the vast majority of
Atlantic sturgeon spawn in their natal
river, they are not likely to seek out
spawning habitat in other rivers and
reduced spawning success due to lack of
appropriate habitat can greatly affect the
recovery potential of a spawning
population. In addition to preventing or
reducing the ability to spawn, dams can
have effects far downstream that reduce
the suitability of river habitat for other
sturgeon life functions. As identified in
the 2007 status review report and the
proposed listing rule, in addition to
blocking habitat upstream, dams also
degrade habitat downstream by altering
DO concentrations and temperature;
artificially destratifying the water
column; changing sediment load and
channel morphology; accelerating
eutrophication and changing nutrient
cycling; and contaminating water and
sediment. The suitability of riverine
habitat for Atlantic sturgeon spawning
and rearing also likely depends on
annual fluctuations in flow, which can
be greatly altered or reduced by the
presence of dams, as has been shown for
sturgeon species (Richter and Thomas,
2007; Pringle et al., 2000; Beamesderfer
and Farr, 1997). Activities associated
with dam maintenance, such as
dredging and minor excavations along
the shore, can release silt and other fine
river sediments that can be deposited in
nearby spawning habitat.
The estimate of 64 percent of
historical habitat on the Cape Fear River
blocked by Lock and Dam #1 was
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questioned by a commenter, who
provided his own estimate of 30 percent
of historical habitat blocked on the Cape
Fear River. The estimate for the Cape
Fear River included in Table 7 of the
status review report is accurate, and
potentially even an underestimate of the
amount of habitat blocked to Atlantic
sturgeon by Lock and Dam #1. The
estimate came from thesis research
(Oakley, 2003) that used regression
models based on river characteristics,
including total river length and distance
to the first dam, to help predict presence
of shortnose sturgeon within a river
system. BASINS 3.0, a GIS-based
program developed by the USEPA, was
used to estimate these physical
characteristics for each river modeled in
the study, including the Cape Fear.
Information from the thesis, presented
in Table 7, lists rkm 95 as the location
of Lock and Dam #1 and rkm 267 as the
fall line, which indicates 172 rkm (or
64.4 percent of the Cape Fear River) are
inaccessible to Atlantic sturgeon.
However, in ‘‘Rivers of North America’’
(Benke and Cushing, 2005) it is stated
that the fall line on the Cape Fear is
located at the confluence of the Deep
and Haw Rivers at rkm 313, which
would indicate 218 rkm (or 69.7 percent
of the Cape Fear River) are inaccessible
to Atlantic sturgeon due to Lock and
Dam #1. In addition, NCDENR’s Office
of Environmental and Public Affairs
notes that access to 160 miles (257 rkm)
of habitat has been blocked to
anadromous species on the Cape Fear
River (https://www.eenorthcarolina.org/
public/ecoaddress/riverbasins/cape
fear2.pdf).
In response to comments about the
use of river miles/kilometers as a
measure of habitat loss and availability
rather than habitat quality, we note that
Table 7 of the status review report states
‘‘river kilometers is only an estimate of
habitat availability and should not be
confused as a reference to habitat
suitability, as many factors can reduce
the quality of this available habitat (e.g.,
impeded by water flow, dredging, water
quality and other similar factors).’’ The
commenter is correct that we have not
designated critical habitat, and we are
not suggesting that the entire river is
necessary for spawning or other life
functions. Because we have little
historical or current information about
the exact locations of Atlantic sturgeon
habitat, the best available information
was the amount of habitat inaccessible
to sturgeon above dams. We agree that
habitat quality and its suitability for
different sturgeon life functions is a
necessary consideration in evaluating
the extent of accessible habitat. In fact,
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5943
the use of river kilometers below dams
as a measure of habitat availability is
potentially an overestimate of the
amount of spawning habitat available to
Atlantic sturgeon. For instance, Table 7
indicates that only 8 percent of
historical habitat on the Savannah River
is impeded by dams, based on the
location of the New Savannah Bluff
Lock and Dam (NSBL&D) at rkm 317
and the fall line at rkm 343. However,
the Augusta Shoals, the only rocky
shoal habitat on the Savannah River and
the former primary spawning habitat for
Atlantic sturgeon in the river (Wrona et
al., 2007; Marcy et al., 2005; Duncan et
al., 2003; USFWS, 2003), is located
above NSBL&D, and is inaccessible to
Atlantic sturgeon. While the status
review report states that 92 percent of
the historical habitat on the Savannah
River is still accessible (based on river
kilometers below NSBL&D), in actuality,
the remaining available spawning
habitat is likely far less. Additionally,
while spawning habitat may exist
downstream of many dams, the quality
of that habitat is often degraded, due to
fluctuations in water level, velocity, and
DO resulting from discharges from the
dam, as well as upstream migration of
the salt wedge, resulting from reduced
freshwater discharge from upstream
and/or channel modifications
downstream. Because Atlantic sturgeon
must spawn in freshwater and the
resulting offspring must have adequate
freshwater exposure for growth before
entering saltwater, the encroachment of
the salt wedge can reduce the
availability of spawning habitat and
even reduce the survival of YOY even
if spawning is successful.
One commenter felt that we did not
evaluate the quality of the 91 percent of
total undammed habitat available to
Atlantic sturgeon; however, the
proposed rule went into great detail
about dredging and water quality and
quantity issues existing below dams that
affect the suitability of spawning habitat
for the Carolina and South Atlantic
DPSs. While we have little historical or
current information about the exact
locations of Atlantic sturgeon habitat,
we are currently funding research to
document habitat utilization of Atlantic
sturgeon. We do not believe the
proposed listing rule gave undue weight
to the loss of access to habitat due to
dams or underestimated other threats,
such as Atlantic sturgeon bycatch. We
did not do a cost-benefit analysis on
potential conservation and recovery
efforts, as the ESA and its implementing
regulations prohibit this type of
consideration in listing determinations.
We are hopeful about pending efforts on
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the Cape Fear and Pee Dee Rivers, and
we will continue to work with FERC
and other stakeholders to improve
habitat quality and access during
relicensing activities. However, in our
listing determinations, we had to
evaluate the current status of, and
threats to, Atlantic sturgeon, and how
those are affected by existing regulatory
mechanisms and protective efforts.
Contrary to comments about the
Roanoke River, we do have information
suggesting spawning historically
occurred above the fall line in that
system (Kahnle et al., 1998; Armstrong
and Hightower, 2002). However, in the
proposed listing rule, we focused
primarily on downstream effects
associated with flow, water temperature,
and DO levels in the Roanoke River
from the Kerr Dam and the Gaston Dam/
Roanoke Rapids facilities. Consistent
with the comments received, we
acknowledged in the proposed listing
rule that there have been modifications
to facilities operations on the river to
simulate natural flows and that this has
likely benefited Atlantic sturgeon.
However, we also detailed the
continuing threat to Atlantic sturgeon
from hypoxic waters released from the
Kerr Dam on the Roanoke in the
summer, and the sensitivity of Atlantic
sturgeon to hypoxia coupled with high
temperature. Consistent with the
comments, the proposed listing rule
states that spawning populations occur
in the Roanoke and Cape Fear Rivers.
However, the failure of populations to
rebound does not signify their
adaptation to these conditions, but
rather suggests the threat posed by dams
to the Carolina and South Atlantic DPSs
is contributing to their status.
Comment 43: Several comments were
received on the effects of water
withdrawals on Atlantic sturgeon
habitat. A commenter supplemented
information included in the proposed
listing rule that demand for water for
consumption purposes in the Southeast
is not only going to increase with
increasing population, but also due to
increasing energy demands. The
commenter stated that power plants
withdraw an average of 40 billion
gallons of water every day, representing
65 percent of total water withdrawals.
The commenter also noted that there are
currently 25 interbasin transfers in
Georgia, involving 6 out of 14 of the
state’s river basins. One commenter
noted that there is substantial
information for water withdrawals in
North Carolina and permits are required
to some extent for agricultural
withdrawals. Another commenter stated
that the proposed listing rule discussed
permitted water quantities but did not
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provide data on the available volume of
water at each source or cite studies that
link permitted interbasin transfers to the
degradation of surface waters. A
commenter stated that conservation and
recovery decisions should not be based
on the assumption that most, possibly
all, subpopulations of Atlantic sturgeon
are at risk of entrainment and
impingement and that the impact from
water intakes should be further
evaluated according to the relationship
between the activity, river, and sturgeon
population.
Response: Additional information
provided by commenters on the threats
posed by water withdrawals to Atlantic
sturgeon and their habitat is consistent
with our finding that these activities
pose a significant threat to the Carolina
and South Atlantic DPSs. A commenter
noted that there is substantial
information on water withdrawals in
North Carolina and permits are required
to some extent for agricultural
withdrawals. This is consistent with
information we presented in the
proposed listing rule on permitted water
withdrawals. A commenter stated that
we did not provide data on the available
volume of water at each source or cite
studies linking permitted interbasin
transfers to the degradation of surface
waters. Real-time water data for the
United States is publicly available on
the USGS Web site (https://waterdata.
usgs.gov/usa/nwis/rt). However, as we
stated in the proposed listing rule,
categories of potentially large water
withdrawals in several states do not
require permits and are therefore not
easily quantifiable. While river and
stream flow data is monitored and
recorded, we do not know how much
non-permitted water withdrawals
account for reductions in flow, and
often we do not have data on the
historical (i.e., unimpaired) flow
regimes in most rivers to quantify the
degree to which flow volumes are
currently reduced (Fisher et al., 2003).
The proposed listing rule included
citations from studies describing the
impacts of water withdrawals, permitted
and non-permitted, on water quantity
and quality parameters important to
Atlantic sturgeon (e.g., UGA, 2002; CBO,
2006; Georgia Water Coalition, 2006).
The Congressional Budget Office (CBO,
2006) directly quantified the effects of
water withdrawal on other ESA-listed
species. CBO stated that among the 663
species listed as ‘‘threatened’’ or
‘‘endangered’’ in 1995, 141 were
affected by the diversion or drawdown
of surface water, 82 by water-level
fluctuation, 26 by water-level
stabilization, 61 by water temperature
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alteration, 103 by reservoirs, 71 by the
drawdown of groundwater, and 14 by
alteration of water’s salinity. In addition
to the citations included in the section
on water allocation, many of the
citations in the remainder of the ‘‘Water
Quality’’ section of the proposed listing
rule specifically address the effects of
alteration of DO, temperature, and
pollutant assimilation (potential effects
associated with water withdrawals) on
Atlantic sturgeon (e.g., Niklitschek and
Secor, 2005, 2009a, 2009b; Secor and
Gunderson, 1998; Secor, 1995).
The proposed listing rule stated that
the withdrawal of water from rivers that
support Atlantic sturgeon populations
was considered to pose a threat as a
result of impingement and entrainment
of eggs, larvae, and small juvenile
sturgeon; however, data are lacking to
determine the overall impact of this
threat on sturgeon populations, as
impacts are dependent on a variety of
factors (e.g., the species, time of year,
location of the intake structure, and
strength of the intake current). Of the
three extant studies on direct impacts to
Atlantic sturgeon from water
withdrawals, only one was conducted in
the Southeast at the Edwin I. Hatch
Nuclear power plant, which withdraws
from, and discharges to, the Altamaha
River. Pre-operational drift surveys were
conducted and only two Acipenser
larvae were collected. Entrainment
samples were collected for the years
1975, 1976, and 1980, and no Acipenser
species were observed in the samples
(Sumner, 2004). As stated in the
proposed listing rule, the migratory
behavior of larval sturgeon may allow
them to avoid intake structures, since
migration is active and occurs in deep
water (Kynard and Horgan, 2002). The
2007 status review report ranked the
threats from impingement and
entrainment as low for both DPSs, and
we concurred. If additional information
becomes available on impingement and
entrainment of Atlantic sturgeon, that
information will be evaluated on a level
appropriate to the activity, the river, and
the sturgeon population.
Comment 44: Comments were
received about the effects of dredging on
Atlantic sturgeon and their habitat. One
commenter pointed to a 2007 study that
Atlantic sturgeon in the St. Lawrence
River avoided areas created by
displaced sediments from dredging
activities and that those sites have lower
value as juvenile benthic feeding habitat
as compared to control sites. In contrast,
USACE commented that a 2009 study
showed dredging operations did not
impede movement or utilization of
habitat by Atlantic sturgeon, and that
direct take of sturgeon by hopper
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dredging between 1990 and 2005 was
observed to be 0.6 fish per year. A
commenter noted the 1998 status review
report listed dredging on spawning
grounds as a stressor, but that all
dredging in the Cape Fear River occurs
in saltwater, so the commenter believed
the only habitat being affected is
nursery habitat. The commenter
requested NMFS provide information on
dredging in the freshwater portion of the
Cape Fear River and whether there are
any known effects to shortnose sturgeon
from dredging by the Corps in the past
10 years. Another commenter noted
frequent maintenance dredging occurs
in the Savannah and St. Johns Rivers.
One commenter was concerned that
different types of dredging (new,
maintenance, marine mining, etc.) in
different environments (small portion of
river versus entire navigation channel;
narrow, shallow sections versus wide,
deep sections) were treated the same in
the proposed rule and that a listing
could inappropriately curtail or
eliminate all maintenance dredging.
Several commenters believed that
additional research on the effects of
dredging on Atlantic sturgeon habitat
should be undertaken. One commenter
recommended that the identification of
spawning, nursery, foraging, and
overwintering habitats be given top
priority in rivers with existing Atlantic
sturgeon populations where there is
significant current or proposed dredging
or port expansion activity.
Response: Additional information
provided by commenters on the threats
posed by dredging to Atlantic sturgeon
and their habitat is consistent with our
finding that these activities pose a
significant threat to the Carolina and
South Atlantic DPSs. A commenter
questioned the level of threat to
sturgeon from dredging in the Cape Fear
River, and requested information on
effects to shortnose sturgeon from
dredging. As cited in the 2007 status
review, Dickerson (2005) reported
observed takings of sturgeon from
dredging activities conducted by
USACE between 1990 and 2005.
Overall, 24 sturgeon (2 Gulf, 11
shortnose, and 11 Atlantic sturgeon)
were taken by dredges during those
years. Of the 24 sturgeon captured, 15
(62.5%) were reported as dead. In 2006–
2008, the South Atlantic Division (North
Carolina to Florida) of USACE reported
a single take of a 125 cm Atlantic
sturgeon (categorized in the incidental
take report as ‘‘fresh dead’’) during
dredging of the Savannah Harbor
entrance channel. Relocation trawling
for the same project captured and
moved eight Atlantic sturgeon. Though
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dredging is a source of mortality, and
therefore a concern to NMFS, we believe
the most significant potential threats to
Atlantic sturgeon from dredging are
associated with effects to their habitat.
In response to the commenter requesting
information on dredging in freshwater
on the Cape Fear River, we do not know
of specific examples. However, we have
significant concerns over dredging in
the portions of the river Atlantic
sturgeon can access (i.e., habitat below
Lock and Dam #1), which includes both
spawning and nursery habitat. As noted
in the proposed listing rule, dredging
operations (including the blasting of
rock) on the lower Cape Fear River,
Brunswick River, and port facilities at
the U.S. Army’s Sunny Point Military
Ocean Terminal and Port of Wilmington
are extensive. Moser and Ross (1995)
found that some of the winter holding
sites favored by sturgeon in the lower
Cape Fear River estuary also support
very high levels of benthic infauna and
may be important feeding stations. The
Shortnose Sturgeon Recovery Plan also
notes that, in addition to direct effects,
dredging operations may also impact
shortnose sturgeon by destroying
benthic feeding areas, disrupting
spawning migrations, and filling
spawning habitat with resuspended fine
sediments. A commenter noted that
frequent maintenance dredging occurs
in the Savannah and St. Johns River,
which was also noted in the proposed
listing rule.
The proposed listing rule did not
include a detailed evaluation of the
different forms and locations of
dredging. Rather, we focused on the
effects of dredging that pose the greatest
threat to Atlantic sturgeon and their
habitat, including the disturbance or
removal of benthic fauna, elimination of
deep holes, and alteration of rock
substrates, as well as the creation of
turbidity/siltation, contaminant
resuspension, noise/disturbance, and
alterations to hydrodynamic regime and
physical habitat. We have a large body
of knowledge on potential effects to
habitat from our ESA section 7
consultations with USACE on dredging
in Gulf sturgeon habitat, as well as in
habitat on the East Coast for shortnose
sturgeon. It is unlikely that listing
Atlantic sturgeon would inappropriately
curtail or eliminate all maintenance
dredging, as maintenance dredging is a
common occurrence in areas inhabited
by ESA-listed Gulf and shortnose
sturgeon. However, through our ESA
consultations with USACE and other
action agencies, we may recommend or
require conservation measures that
reduce or eliminate potential impacts to
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Atlantic sturgeon and their habitat. We
agree that additional research on the
effects of dredging on Atlantic sturgeon
habitat and on the locations of
spawning, nursery, foraging, and
overwintering habitat in relation to
potential dredging activities would be
useful. We are constantly working to
expand our knowledge on the effects of
dredging on ESA-listed (and candidate)
species and their habitat, which
includes Atlantic sturgeon.
Comment 45: A commenter concerned
about the effects of climate change
recommended additional research and
monitoring with respect to the impacts
and synergistic effects of climate change
on Atlantic sturgeon subpopulations.
Another commenter stated that climate
change will be the single largest driver
of changes in biodiversity by the end of
the 21st century and that
disproportionate effects will be
experienced in the Southeast, which is
the most vulnerable region due to its
long low-lying coastline and high
biodiversity. The commenter noted that
the South will be drier, with climate
models predicting decreases in
precipitation in the summer combined
with higher temperatures, resulting in
increased evaporation. The commenter
also noted the Carolinas and Georgia
have already shown significant trends of
increasing drought from 1958 to 2007. A
commenter noted that drought occurred
in North Carolina during the same time
frame drought occurred in South
Carolina and Georgia, which further
supports the threat to Atlantic sturgeon
from such occurrences. In addition to
habitat threats from climate change
outlined in the proposed listing rule, a
commenter provided information and a
presentation from a NCDENR climate
change symposium that included
potential effects to the North Carolina
coast and noted that habitat for the
Carolina DPS is almost exclusively in
this area. The presentation discussed
threats of sea level rise, increasing
storms, and resultant property
protection activities, such as beach
renourishment and installation of hard
structures. The presentation stated there
will be detrimental effects to sounds,
rivers, and estuaries utilized by the
Carolina DPS. In contrast, a commenter
stated that even with gradual climate
change and warming, it is likely that
Atlantic sturgeon populations will
continue to increase over most of their
range, as the species has survived more
significant climate and temperature
regimes in its evolutionary past.
However, the commenter acknowledged
that genetic diversity of the species may
be important to assure its survival.
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Several commenters cautioned that
climate change models do not provide
information appropriate for making
management decisions regarding
Atlantic sturgeon. One commenter
cautioned against using the most
extreme scenarios modeled by the
International Panel on Climate Change
(IPCC) and also noted that climate
change may negatively impact species
in one area, but benefit the species in
others, and both positive and negative
impacts should be considered. Two
commenters noted that the proposed
listing rule incorrectly stated the two
Southeast DPSs are in a region the IPCC
predicts will experience decreases in
precipitation, which could exacerbate
low oxygen, and that increases in
precipitation are actually predicted.
Response: Additional information
provided by commenters on the threats
posed by climate change to Atlantic
sturgeon and their habitat is consistent
with our finding that it poses a
significant threat to the Carolina and
South Atlantic DPSs. We agree that
additional research and monitoring of
impacts and synergistic effects on
Atlantic sturgeon are necessary. As we
noted in the proposed listing rule, we
are particularly concerned about the
exacerbation through climate change of
existing water quality issues and
increasing water demands due to human
population increases in the Southeast.
While Atlantic sturgeon may have
experienced different climate and
temperature regimes over their
evolutionary history, they have not had
to persist with the combination of
threats they face now, and we do not
agree with the commenter that Atlantic
sturgeon populations will increase
without addressing these threats. Their
populations were rapidly depleted by
1901 as a result of fishing. Even though
directed fishing was abolished, Atlantic
sturgeon continue to be taken as bycatch
in various fisheries. Dams block access
to habitat and affect downstream habitat
quality, as does dredging. Water
quantity and quality is affected by a
variety of watershed activities. These
threats are predicted to increase as
population in the Southeast increases,
and climate change is expected to
further exacerbate water quality and
quantity issues. We agree with the
commenter that genetic diversity (and
larger population sizes) will be
necessary for Atlantic sturgeon to
recover in the face of these increasing
threats.
We agree with the comment that the
most extreme scenarios modeled by the
IPCC are not appropriate for making
management decisions associated with
our listing of the Carolina and South
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Atlantic DPSs. While the IPCC modeled
many scenarios and reported results
with varying degrees of certainty, we
only reported the most conservative
results, the scenarios that were ‘‘very
likely’’ to occur and which the IPCC
projected with ‘‘high confidence.’’ In
addition, our discussion of climate
change focused on the ways in which it
was likely to exacerbate existing threats,
which we do feel warranted
consideration in our listing
determination. We did not use the
IPCC’s most extreme climate change
model scenarios to make predictions
about potential future threats to Atlantic
sturgeon or factor those scenarios into
our proposed listing determination.
While we agree in theory that climate
change could have both positive and
negative effects, our review of the IPCC
information did not reveal any aspects
of climate change that would have
positive effects on the Carolina and
South Atlantic DPSs in the Southeast
and the comment did not include
specific examples of positive effects for
our consideration.
We appreciate the commenters noting
that we incorrectly stated the two
Southeast DPSs are in a region that the
IPCC predicts will experience decreases
in precipitation, exacerbating low DO.
Overall, the Southeast is predicted to
experience increases in precipitation.
However, evaporation is also predicted
to increase with increasing temperatures
and the net effect for the Southeast is
predicted to be overall drying. Further,
conservative seasonal predictions for
the summer show either a slight
increase in precipitation or a slight
decrease. Decreased precipitation or
even a slight increase, offset by
increased summer temperatures and
evaporation, would exacerbate low DO
when temperatures are highest. As
discussed in the proposed listing rule,
Atlantic sturgeon are particularly
vulnerable to low DO when combined
with high temperatures. Also, overall
decreased water availability due to
increased temperature and longer
periods of time between rainfall events
is predicted for the Southeast, even
though individual rainfall events are
predicted to be more extreme, leading to
the increased precipitation estimates.
We have corrected this information in
the section of the final rule that
addresses climate change.
Comments on Bycatch
Comment 46: Many comments were
received from parties concerned about
the impacts of Atlantic sturgeon bycatch
in both commercial fisheries and
scientific surveys, and several
commenters provided suggested
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solutions. One commenter stated that
over 1,000 Atlantic sturgeon are taken
annually as bycatch. Another
commenter cited Munro et al. (2007)
that bycatch likely has more detrimental
effects in habitats that are limited in
area and where certain life stages of
Atlantic sturgeon tend to congregate,
such as early juvenile habitats in the
estuarine transition zone and the
subadult/adult habitat in the nearshore
oceanic zone. The commenter also
stated that protecting juvenile marine
stage Atlantic sturgeon from bycatch
mortality in aggregation areas is likely
the key to restoring Atlantic sturgeon
populations given that the intrinsic rate
of population increase for long-lived
species like Atlantic sturgeon is most
sensitive to changes in juvenile survival.
The commenter noted that while little
direct mortality is reported for trawl
fisheries, within aggregation areas it is
not uncommon to catch ten or more
Atlantic sturgeon in a single 20 minute
tow, and that with longer trawl times in
commercial fisheries, fish released alive
may die days after. A commenter was
concerned that bycatch of Atlantic and
shortnose sturgeon has been occurring,
citing data from the Santee River. A
comment was received recommending
research to determine the impacts of
bycatch and bycatch mortality on
Atlantic sturgeon populations,
identification of the spatial and
temporal distribution of bycatch
throughout the species range, and
development of measures that could be
implemented to reduce bycatch and/or
bycatch mortality. Several commenters
stated that NMFS has not taken
adequate steps to reduce or stop the use
of gillnets and other gears to protect
sturgeon. Comments were received that
the moratorium has not prevented
bycatch, and gill nets should be banned
in order to recover Atlantic sturgeon.
One commenter asked if NMFS had
solicited or received advice from
commercial fishermen on limiting
bycatch mortality in gillnets. Citing
Dunton et al. (2010), a commenter stated
that because previous Atlantic sturgeon
management has not resulted in
significant improvements to
populations, recovery efforts should
now focus on establishing marine
reserves or implementing area closures
to protect essential habitat and to reduce
fishing mortality on juveniles (Collins et
al., 2000). The commenter stated that
the primary juvenile habitat and
juvenile migrations are limited to
narrow corridors in waters less than 20
meters deep and this is conducive to a
seasonal or permanent closure to gillnet
and trawl fisheries. The commenter
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believed that by focusing immediate
efforts on the protection of these
hotspots and corridor pathways, bycatch
mortality will be reduced effectively
through protection of habitat. One
commenter was concerned about
mortality levels in scientific surveys and
recommended that scientific sampling
be banned in the Cape Fear River.
Response: Additional information
provided by commenters on the impacts
of Atlantic sturgeon bycatch is
consistent with our finding that it poses
a significant threat to the Carolina and
South Atlantic DPSs. As we continue to
work to reduce Atlantic sturgeon
bycatch, we will consider suggestions
provided by commenters, such as the
importance of protecting juvenile
marine stage Atlantic sturgeon,
identifying hotspots and migratory
corridors, investigating the
establishment of marine reserves or
closed areas, and working with gillnet
fisheries to reduce the level of Atlantic
sturgeon bycatch. We do not feel that
banning scientific sampling in the Cape
Fear River would benefit Atlantic
sturgeon, and we recently published ‘‘A
Protocol for Use of Shortnose, Atlantic,
Gulf, and Green Sturgeons’’ (Kahn and
Mohead, 2010; available at https://www.
nmfs.noaa.gov/pr/pdfs/species/kahn_
mohead_2010.pdf) that can be followed
to better ensure the safety of sturgeon
during research, including during
capture using gillnets. We will continue
to work with our partners and
stakeholders through our existing
authorities to reduce or eliminate the
effects of bycatch on Atlantic sturgeon.
Comment 47: Several commenters
questioned how listing Atlantic
sturgeon will result in a greater
reduction in bycatch than is already
being realized by closing the
commercial fishery for Atlantic
sturgeon. A commenter stated that there
has been a significant reduction in
vessels and effort in the shad gillnet and
shrimp trawling fisheries over the last
10 years. Other commenters listed a
number of commercial fishery
regulations (i.e., harvest seasons, gill net
mesh size, and quantity restrictions),
some associated with other fisheries
(e.g., striped bass, American shad) that
also reduce the potential for gill net
interactions with Atlantic sturgeon.
Commenters also noted significant
reductions in pound net and haul seine
use have occurred during recent
decades in the Albemarle Sound area,
further reducing potential interactions
between sturgeon and commercial
fisheries. Two North Carolina state
agencies reported that out of more than
3,000,000 yards of large and small mesh
gill nets observed since 2001, overall
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bycatch mortality was 6 percent (with
an annual range of 0 to 12 percent),
which is lower than the 13.8 percent
estimated by the ASMFC and cited in
the proposed listing rule. The agencies
also reported that mortality in the
Albemarle and Pamlico Sound IGNS
had overall Atlantic sturgeon mortalities
of 3 and 10 percent, respectively, and
mortality in the Pamlico, Pungo, and
Neuse Rivers IGNS was 12 percent. The
agencies commented that mortalities
were high in the Cape Fear River IGNS
(35 percent), and that mortality was less
than 13.8 percent in Cape Fear River
and near shore Atlantic Ocean Fishery
Independent Assessment Program. In
reference to the 35 percent mortality in
the Cape Fear IGNS, commenters
(including NCDENR) said that these
results cannot be extrapolated to
commercial fisheries because of gear
and seasonal restrictions in place for
those fisheries that do not allow them to
be operated in the same time, place, or
with the same gear. These agencies also
noted that bycatch has been
documented for over 958 tows
conducted by commercial shrimp
trawlers working in North Carolina with
no Atlantic sturgeon reported and that
no Atlantic sturgeon have been captured
in the 528 blue crab trawl tows
examined since 1990. They also stated
that the White and Armstrong Fishery
Resource Grant study (2000) conducted
in the Albemarle Sound was used in the
listing documents because of a high
collection rate; however, targeting of
Atlantic sturgeon may have occurred
since the design of the study was to
estimate survival of sturgeon captured
in commercial flounder nets. White and
Armstrong (2000) also noted no
mortality of Atlantic sturgeon collected.
GADNR commented that less than 10
fish per year were estimated to have
been captured in the Altamaha River
anchored gillnet fishery during a 3-year
study. All fish were juveniles and no
injury or mortality was documented.
Georgia also noted the season for
gillnetting shad occurs while adults are
at sea and juveniles are in the lower
parts of the estuary. Since the 2007
status review, which ranked bycatch as
a moderate threat in the Altamaha, the
State of Georgia commented that recent
action by the Board of Natural
Resources has prohibited the use of
gillnets for shad fishing in a large
portion of the Altamaha. Two
commenters disagreed with the use of
Stein et al. (2004) in relation to bycatch
in the Southeast, stating that offshore
fisheries with long soak times should
not be used as a proxy for inshore
fisheries, and though mixing of sturgeon
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populations occurs in marine areas,
most of the fish captured as bycatch
would be fish from northern DPSs. A
comment was received that listing
Atlantic sturgeon could require changes
to gear design or fishery regulations for
fisheries that encounter Atlantic
sturgeon as bycatch, and that while
bycatch mortality estimates are
unknown for many species, they are
believed to be low with the exception of
sink gillnet fisheries with long soak
times. One commenter suggested that
the South Atlantic DPS was not subject
to the same level of bycatch as the
Carolina DPS.
Response: Listing the Carolina and
South Atlantic DPSs as endangered
could result in a further reduction in
fishing mortality, beyond the
commercial harvest moratoria, if
conservation measures implemented
pursuant to the ESA lead to reductions
in bycatch, for example through section
10 permits or section 7 biological
opinions. While the moratoria on
harvest and possession have greatly
reduced the effects of fisheries on
Atlantic sturgeon, fish from these DPSs
are still being taken as bycatch in many
fisheries. Once listed as endangered,
bycatch of Atlantic sturgeon would be
considered ‘‘take’’, defined in section 3
of the ESA as ‘‘to harass, harm, pursue,
hunt, shoot, wound, kill, trap, capture,
or collect, or attempt to engage in any
such conduct.’’ Section 7 consultation
would be required for federally
authorized fisheries that take Atlantic
sturgeon as bycatch. During
consultation, NMFS would evaluate the
anticipated level of take associated with
the fishery, evaluate whether it would
jeopardize the continued existence of
the species, and determine reasonable
and prudent measures that would
reduce the anticipated effects of the
incidental take on the species. A section
10(a)(1)(B) permit would be required for
fisheries authorized by states that result
in Atlantic sturgeon bycatch. A section
10(a)(1)(B) would require the
development of a conservation plan that
details the impact to the species, the
steps that will be taken to minimize and
mitigate the impacts, alternative actions
considered and why they were not
implemented, and any other measures
required by NMFS to benefit the
species.
Even with reductions in gillnet and
trawl vessels and fishing effort, and the
implementation of other seasonal and
gear restrictions, there are still large
numbers of participants in fisheries
using these gears. Every year, NMFS
publishes a list of commercial fisheries
and classifies them into categories
according to the level of interactions
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with marine mammals. Based on the
latest list, published on November 8,
2010 (75 FR 68468), fisheries using
gillnet and trawl gear and the number of
participants in those fisheries in the
range of the Carolina and South Atlantic
DPSs include the following: MidAtlantic gillnet fishery, 5,495
participants; the North Carolina inshore
gillnet fishery, 2,250 participants; the
Southeast Atlantic gillnet fishery, 779
participants; the Southeastern U.S.
Atlantic shark gillnet fishery, 30
participants; the Mid-Atlantic bottom
trawl fishery, 1,182 participants; and,
the Southeast U.S. Atlantic, Gulf of
Mexico shrimp trawl fishery, 4,950
participants (though this includes Gulf
of Mexico participants). However, we
note that the number of participants
listed here is potentially an overestimate
of the number of participants interacting
with Atlantic sturgeon. For example, in
the gillnet fisheries, the number of
participants includes fishermen using
non-sink gillnets, which have fewer
interactions with Atlantic sturgeon. In
addition, all fishery participants may
not be operating at times or in areas
where they are likely to encounter
Atlantic sturgeon. Further, based on
available bycatch data, which suggests
sturgeon are primarily caught in waters
less than 50 meters deep, commercial
and recreational fisheries using trawl
and gillnet gear in waters greater than
50 meters deep may not have Atlantic
sturgeon bycatch. Estimates for Atlantic
sturgeon bycatch in these fisheries is
largely unavailable, as bycatch is
underreported in state waters and there
is limited observer coverage in fisheries
potentially capturing Atlantic sturgeon
in the South Atlantic (North Carolina to
Florida) Federal waters.
We have added information on
bycatch provided by North Carolina and
Georgia to section ‘‘B. Overtutilization
for Commercial, Recreational, Scientific,
or Educational Purposes’’ of the final
listing determination. Regarding bycatch
data supplied by the State of North
Carolina, the lack of recorded Atlantic
sturgeon bycatch in commercial shrimp
trawls and blue crab trawls in North
Carolina is consistent with information
presented in the proposed listing rule
that trawl gear is not believed to be a
significant threat to Atlantic sturgeon.
Data reported for the Albemarle and
Pamlico Sound IGNS, as well as IGNS
in the Pamlico, Pungo, Neuse, and Cape
Fear Rivers, show overall (i.e., mortality
over all survey years combined) Atlantic
sturgeon capture mortality in gillnets
ranging from 3 to 35 percent. With the
exception of the highest morality rate,
which was observed in the Cape Fear
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River IGNS, North Carolina commented
that all of the observed mortality rates
were less than the 13.8 mortality cited
in the proposed listing rule, but that the
majority of the results cannot be
extrapolated to commercial fisheries
due to gear and seasonal harvest
restrictions under which they operate.
Based on the data supplied by the state,
capture mortality of Atlantic sturgeon
varied greatly by year, by month, and by
the gillnet mesh size used during the
survey. For instance, mortality during
individual survey years in the
Albemarle Sound IGNS ranged from 0 to
19 percent during 1990 to 2009.
Mortality by month ranged from 0 to 7
percent, with the highest mortalities
recorded in April (3 percent), May
(7 percent), and November (5 percent).
Mortality ranged from 0 to 100 percent
in mesh sizes ranging from 2.5- to
10-inch stretched mesh (ISM), with
fairly consistent levels: 2 to 4 percent
for mesh sizes 2.5 to 5.5 ISM, 9 percent
for 6.5 ISM, and 100 for 10 ISM
(representing 1 Atlantic sturgeon).
Similar variability was seen in the
Pamlico Sound IGNS data. During 2001
to 2008, 0 to 17 percent mortality was
observed in the Pamlico Sound IGNS,
with 100 percent in 2009, based on 1
Atlantic sturgeon. Mortality ranged from
0 to 25 percent by month, with peak
mortalities occurring in June (25
percent), August (17 percent), and
November (17 percent). The Pamlico
Sound IGNS used mesh sizes ranging
from 3 to 6.5 ISM. Mortality by mesh
size ranged from 0 to 25 percent, with
the highest mortalities observed in the
3 ISM (25 percent), 3.5 ISM (20 percent),
and 6.5 ISM (20 percent). While the
State of North Carolina commented that
the IGNS data should not be
extrapolated to estimate a mortality rate
for commercial fisheries, it does show
that time of year and gear type factor
heavily into Atlantic sturgeon bycatch
mortality. As stated by North Carolina,
as well as in the proposed listing rule,
other factors, such as gillnet soak time,
affect mortality rates. Overall mortality
rates in all North Carolina surveys (with
the exception of the Cape Fear River
IGNS) may be below the 13.8 percent
estimate reported in the proposed listing
rule; however, mortality rates during
individual survey years, during certain
survey months, and for specific gillnet
mesh sizes used often exceeded 13.8
percent. While North Carolina provided
fishery-dependent survey data from
their observer program, observer
coverage in fisheries potentially
capturing Atlantic sturgeon is very
limited for the remainder of the
Southeast range occupied by the
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Carolina and South Atlantic DPSs. High
levels of bycatch underreporting are
suspected. Further, even if bycatch
mortality is lower than the 13.8 percent
estimate reported in the proposed listing
rule, total population abundances for
the Carolina and South Atlantic DPSs
are not available and we do not know
what portion of the Carolina and South
Atlantic DPSs are subject to being taken
as bycatch. As cited in the proposed
listing rule, Boreman (1997) calculated
a sustainable fishing (bycatch) mortality
rate of 5 percent per year for adult
Atlantic sturgeon, indicating they can
only tolerate relatively low levels of
bycatch mortality.
Fisheries known to incidentally catch
Atlantic sturgeon occur throughout the
marine range of the species and in some
riverine waters as well. Because Atlantic
sturgeon mix extensively in marine
waters and may access multiple river
systems, they are subject to being caught
in multiple fisheries throughout their
range. Atlantic sturgeon taken as
bycatch may suffer immediate mortality.
In addition, stress or injury to Atlantic
sturgeon taken as bycatch but released
alive may result in increased
susceptibility to other threats, such as
poor water quality (e.g., exposure to
toxins and low DO). This may result in
reduced ability to perform major life
functions, such as foraging and
spawning, or may even result in postcapture mortality. Several of the river
populations in the South Atlantic DPS
(e.g., the Ogeechee and the Satilla) are
stressed to the degree that any level of
bycatch could have an adverse impact
on the status of the DPS (ASSRT, 2007).
Therefore, the information supplied by
the State of North Carolina does not
provide a basis for revising our
evaluation of the threat of bycatch to
Atlantic sturgeon populations or our
determination that the Carolina and
South Atlantic DPSs warrant listing as
endangered. For the same reasons, the
information supplied by the State of
Georgia does not provide a basis for
revising our evaluation of the threat of
bycatch to Atlantic sturgeon. The state
documented less than 10 fish per year
taken as bycatch in the Altamaha River
gillnet fishery, with no observed
mortality during a 3-year study. Georgia
also commented that the shad gillnet
season occurs while adults are at sea
and juveniles are in the lower part of the
estuary and that the state now prohibits
shad gillnetting in a large portion of the
Altamaha. However, the Altamaha River
has the largest and healthiest population
of Atlantic sturgeon in the Southeast
and bycatch occurring in systems with
smaller, more greatly stressed
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populations (such as the Ogeechee and
Satilla; ASSRT, 2007) may have adverse
impacts. We commend the state for their
efforts in reducing the threat of bycatch
in the Altamaha River, but we believe
bycatch still represents a significant
threat to the Carolina and South
Atlantic DPSs of Atlantic sturgeon. The
bycatch information for Atlantic
sturgeon in the Carolina and South
Atlantic DPSs provided by North
Carolina and Georgia, when considered
as part of our listing determination, does
not change our determination that the
two DPSs warrant listing as endangered.
The White and Armstrong (2000)
study was not considered in the
proposed listing rule for the reason
suggested by the commenter, i.e., due to
the high collection rate of Atlantic
sturgeon. We cited this study as one of
the only fishery-dependent bycatch
surveys of Atlantic sturgeon from either
the Carolina or South Atlantic DPSs
available to us. Contrary to the
commenters’ assertion that targeting of
Atlantic sturgeon may have occurred,
the research publication states in the
‘‘Methods’’ section that ‘‘southern
flounder (not Atlantic sturgeon) were
the target species, and the incidence of
Atlantic sturgeon captures in the catch
was expected to be representative of
normal bycatch rates.’’ The publication
also stated that ‘‘survival rates were
inestimable, the apparently healthy
condition of incidentally captured
Atlantic sturgeon is consistent with low
release mortality.’’
While commenters disagreed with our
use of offshore fisheries data in relation
to bycatch in the Southeast (e.g., Stein
et al., 2004; ASMFC, 2007), we used the
best data available to us in the proposed
listing rule and clarified its utility. We
noted in the proposed listing rule that
any estimate of bycatch from the NMFS
ocean observer dataset will be an
underestimate, because bycatch is
underreported in state waters and there
is no observer coverage in the South
Atlantic (North Carolina to Florida)
Federal waters. We are updating
information in this section of the final
rule to reflect that there is limited
observer coverage in Federal waters in
the Southeast for gear types that
potentially capture Atlantic sturgeon.
The shark drift gillnet program, which
operates primarily off the southern
Atlantic Coast of Florida and North
Carolina, observes a relatively small
fishery (25–30 vessels) targeting coastal
shark species, as well as king and
Spanish mackerel, little tunny, bluefish,
and Atlantic croaker. There is also an
observer program for the Southeastern
shrimp trawl fishery, which covers
approximately 1 percent of the fishery
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in the South Atlantic. This information
does not change our conclusion that
bycatch is underreported in state and
Federal waters. In addition to
immediate mortality, bycatch mortality
estimates do not account for postcapture mortality and may further
underestimate the mortality rate in sink
gillnets in the Carolina and South
Atlantic DPSs because bycatch survival
is greater in colder water temperatures
of the north compared to warmer
southern waters occupied by these
DPSs.
Comments on Disease and Predation
Comment 48: One commenter stated
that the need and ability to regulate the
aquarium trade should not have been
discounted in the proposed listing rule.
The commenter believed importation of
non-native sturgeon is a greater threat to
native sturgeon than any other factor
because non-natives potentially outcompete native fish and introduce
disease.
Response: We agree that the ability to
regulate the aquarium trade should not
have been discounted in the proposed
listing rule, and we are removing that
text in the final rule. However, we do
not have information that suggests the
aquarium trade is a current threat to
Atlantic sturgeon. We disagree that the
importation of non-native sturgeon is a
greater threat to native sturgeon than
any other threat. We included
information in the proposed listing rule
that there were only five known Atlantic
sturgeon commercial aquaculture
operations in the Southeast, one in
North Carolina and four in Florida.
These operations all cultured Atlantic
sturgeon originating from Canadian
stock, with the exception of the North
Carolina operation that acquired
Siberian sturgeon (A. baerii) in 2006
after obtaining an addendum to their
permit from the ASMFC. Additionally,
we obtained information on the culture
of other sturgeon species. Commercial
U.S. culture of meat and caviar is
currently taking place in three states:
California, Idaho, and Florida (Monterey
Bay Aquarium, 2007). Four facilities
(Evans Farm, Mote Marine Laboratory,
Rokaviar, and Sturgeon AquaFarms,
LLC) in Florida, the only state in the
range of Atlantic sturgeon culturing
non-native species, conduct tank culture
of the following species: Siberian
sturgeon (A. baerii), Russian sturgeon
(A. gueldenstaedti), Stellate sturgeon (A.
stellatus), Sterlet sturgeon (A. ruthenus),
Adriatic sturgeon (A. naccarii), beluga
sturgeon (Huso huso), and the hybrid
Bester sturgeon (H. huso x A.
rutheni)(M. Berrigan, FDACS, pers.
comm.). The nature of current
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containment practices and the reported
record of total escape prevention for the
Florida facilities that presently culture
non-native sturgeons suggest currently
low exposure for wild sturgeon stocks to
the ecological risks of farmed fish
escapes (Monterey Bay Aquarium,
2007). We acknowledged in the
proposed listing rule that introduction
of non-native species could impact
native sturgeon populations. However,
we did not believe that this was a
significant threat based on the very low
occurrence of non-native Atlantic
sturgeon culture operations and the fact
that stock enhancement programs follow
culture and stocking protocols approved
by the ASMFC, which includes, ‘‘if nonnative or hybrid sturgeon are permitted
within a state, they should be restricted
to culture operations where escapement
and reproduction can and will be
controlled.’’ We also noted that
mechanisms are in place at all facilities
to prevent escapement of sturgeon;
facilities are all land based, and most
are not located in close proximity to any
Atlantic sturgeon rivers. All of the
facilities in Florida are periodically
screened for disease by University of
Florida Institute for Food and
Agricultural Science (IFAS)
veterinarian. None have reported
diseases. All facilities are above the 100year flood plain and have zero
discharge.
We received information during the
public comment period that indicates a
further reduction in the potential threat
of non-native sturgeon to the Carolina
and South Atlantic DPSs. The Florida
Department of Agriculture and
Commerce (FDACS), which certifies
aquaculture facilities and inspects those
facilities twice a year for compliance,
informed us that only one commercial
facility with Atlantic sturgeon is
currently operating in Florida, and they
only have one surviving fish. All other
Atlantic sturgeon held in Florida
aquaculture facilities died in captivity.
Additional information supplied by
FDACS on Florida aquaculture facilities
is included in our response to comment
53.
Comments on the Inadequacy of
Existing Regulatory Mechanisms
Comment 49: Several commenters
provided us with additional examples of
the inadequacy of regulatory
mechanisms. One commenter believed
the ASMFC’s failure to end the harvest
of overfished stocks (e.g., winter
flounder and weakfish) and North
Carolina’s request for an exemption to
the law that fishery management plans
have a 50 percent probability of
recovering depleted stocks exemplify
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inadequacies of existing regulatory
mechanisms to protect species and
highlight the need for ESA listing.
Several commenters noted the lack of
permitting programs in southeastern
states for water withdrawals, including
interbasin transfers, and the lack of
regulation of instream flows. One
commenter noted that while there is a
blanket prohibition against water
transfers into metro Atlanta, adjacent
counties are joining the district where
Atlanta is incorporated in order to avoid
the prohibition. Another commenter
stated that North Carolina coastal
counties currently seeking interbasin
transfers have been exempted from 2007
amendments regulating interbasin
transfer and the North Carolina
Department of Water Quality is seeking
to create regulatory changes to the
current buffer rules. The commenter
also stated that, for the second year in
a row, North Carolina passed legislation
allowing any existing permit or finding
to extend until after 2011 without
having to reapply or renew as a way to
mitigate the economic downturn. The
extension is applicable to several types
of permits and applications that could
affect the Carolina DPS, including:
Findings of no significant impact;
approvals of an erosion and
sedimentation control plan; permits for
major developments or minor
developments under the State’s Coastal
Area Management Act; water or
wastewater permits; building permits;
stream origination certifications; water
quality certifications; air quality
permits; and city and county site
specific development plans. A comment
was also received regarding Senate Bill
778, which became law in North
Carolina in August of 2010. The bill was
drafted as a response to litigation
regarding the proposed Titan Cement
plant and created a loophole that any
project such as Titan, which may have
a significant environmental impact, can
bypass the State Environmental Policy
Act (SEPA) by structuring a contract on
the basis of incentives. While the
legislation does not retroactively exempt
Titan Cement from SEPA, it ensures that
a roadmap exists for any similar projects
in the future to avoid the environmental
review process established in SEPA.
Response: In the proposed listing rule,
we concluded that the inadequacy of
regulatory mechanisms to fully address
the threats of bycatch and habitat
modification are contributing to the
endangered status of the Carolina and
South Atlantic DPSs of Atlantic
sturgeon. The information provided by
these commenters supports this
conclusion. We will continue to
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investigate these issues and ways to
ameliorate any effects they are having
on Atlantic sturgeon.
Comment 50: Several commenters
disagreed with NMFS’ finding in the
proposed listing rule that existing
regulatory mechanisms protecting
Atlantic sturgeon and their habitat are
inadequate. Numerous commenters
believed the proposed listing rule is
unnecessary because directed fishing
and retention of Atlantic sturgeon has
been prohibited by the moratoria
implemented by the ASMFC and NMFS,
as well as various prohibitions enacted
by individual states. A commenter
noted that South Carolina and North
Carolina initiated moratoria on harvest
and possession of Atlantic sturgeon in
1985 and 1991, respectively. Another
commenter noted that special concern
designations have been given to Atlantic
sturgeon by the states of Virginia and
Florida. Many commenters believe the
prohibitions are working and that no
listing action should be taken until the
moratoria have had sufficient time to
work. A commenter stated that
protections already in place for cooccurring endangered species are
sufficient to protect Atlantic sturgeon
and their habitat. Comments were
received that NMFS did not thoroughly
consider the benefits of existing
regulatory mechanisms addressing
bycatch and activities affecting Atlantic
sturgeon habitat (e.g., regulations
associated with construction,
demolition, and dredging), and that
existing regulations should be used to
protect Atlantic sturgeon populations.
The State of North Carolina commented
that the North Carolina Coastal Habitat
Protection Plan (NCCHPP) and
moratoria on construction, dredging,
and other habitat altering activities are
already managing habitat issues,
observer programs are expanding to
include more fisheries, gear
configurations and regulations have
been updated to reduce bycatch and
limit interactions with protected
species, and research is being funded
that will allow North Carolina and other
states to gain a better understanding of
the migratory patterns, spawning areas,
and distribution of Atlantic sturgeon
within the next few years. The NCCHPP
was adopted in 2005 and its stated goals
are: (1) Improving effectiveness of
existing rules and programs protecting
coastal fish habitats; (2) identifying,
designating, and protecting strategic
habitat areas (SHAs); (3) enhancing
habitat and protecting it from physical
impacts; and, (4) enhancing and
protecting water quality. The North
Carolina Marine Fisheries Commission
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(NCMFC) approved SHAs for Region 1
in North Carolina in January 2009, and
is currently evaluating SHAs for other
regions in North Carolina. According to
the commenter, SHAs represent priority
habitat areas for protection due to their
exceptional condition or imminent
threat to their ecological functions
supporting estuarine and coastal fish
and shellfish species and will be
incorporated into conservation and
restoration efforts. One SHA (Bellows
Bay to Knotts Island Bay) was identified
in part due to the nearshore ocean areas
that are important for Atlantic sturgeon
and striped bass and another SHA
(Chowan and Roanoke Rivers and
western Albemarle Sound) may include
one of the few Atlantic sturgeon
spawning habitats in North Carolina.
The State also commented that the
North Carolina Department of Marine
Fisheries (NCDMF) provides input to
federal and state regulatory agencies of
the location of habitats used by Atlantic
sturgeon. NCDMF and the North
Carolina Wildlife Resources
Commission have designated
Anadromous Fish Spawning Areas
(AFSA) through rules for their
respective jurisdictions.
Response: Though moratoria on
harvest and possession of Atlantic
sturgeon were enacted by the ASMFC,
NMFS, and several states, populations
have not rebounded and the moratoria
do not control bycatch. We believe
continued bycatch of Atlantic sturgeon
in commercial fisheries has an ongoing
impact upon the Carolina and South
Atlantic DPSs that is not adequately
addressed through existing regulatory
mechanisms and is contributing to their
endangered status. Because Atlantic
sturgeon mix extensively in marine
waters and may access multiple river
systems, they are subject to being caught
in multiple fisheries throughout their
range. Poor water quality also continues
to result in adverse effects to Atlantic
sturgeon even with existing controls on
some pollution sources and water
withdrawal, and dams continue to
curtail and modify habitat, even given
the provisions for establishing fishways
under the Federal Power Act.
As noted in the comments, Florida
has designated the Atlantic sturgeon as
a species of special concern. This
designation stipulates that no person
shall take, possess, transport, or sell any
species of special concern without a
permit. The comments also noted
Atlantic sturgeon was designated as a
species of special concern by Virginia,
which is described as a ‘‘watchlist’’ of
wildlife species with no other regulatory
or statutory requirements. Currently, the
state’s Wildlife Action Plan identifies
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Virginia’s Species of Greatest
Conservation Need (SGCN) and has
made the ‘‘special concern’’ designation
obsolete. The SGCN also has no
regulatory requirements, but requires
that Comprehensive Wildlife
Conservation Strategies be developed
that include 8 essential elements,
including key information on
distribution, abundance, threats,
descriptions of conservation actions,
and plans for species monitoring. While
states should be commended for
recognizing the need for conservation of
Atlantic sturgeon, these designations are
not enough to alleviate the threats to the
Carolina and South Atlantic DPSs of
Atlantic sturgeon or change our
evaluation of the species as meeting the
definition of endangered from section 3
of the ESA.
While there are a variety of other
Federal, state, and local laws and
programs (e.g., regulations governing
construction activities and gear
configurations that reduce bycatch) that
benefit Atlantic sturgeon, we believe
that threats from habitat modification
and bycatch are not sufficiently
managed through current regulatory
mechanisms in place. For instance,
seasonal restrictions governing
construction and dredging in North
Carolina may benefit Atlantic sturgeon
during critical time periods, as stated by
the commenter, but construction and
dredging during other times of the year
can still impact Atlantic sturgeon and
their habitat. Required gear
configurations may reduce Atlantic
sturgeon bycatch, but bycatch still
occurs. Further, the lack of bycatch data
does not allow us to evaluate the degree
to which bycatch is potentially reduced
by these measures. We reviewed the
information provided by the State of
North Carolina on the NCCHPP, SHAs,
and AFSAs, as well as additional
information on these programs on
NCDENR’s Web sites. While these
programs have excellent goals of
increasing enforcement of existing
regulations, identifying and protecting
habitat important to the species, and
monitoring these habitats, many of these
actions are still in the early stages and
it is not clear exactly what protections
will be given to areas designated as
SHAs or AFSAs. We are also including
an evaluation of these programs in the
section of the final listing rule
evaluating current protective efforts.
Comments on Other Natural and
Manmade Factors
Comment 51: The ASMFC
commented that states and jurisdictions
where ship strikes are an issue are
currently monitoring and working to
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minimize these impacts. Another
commenter was concerned that if ship
strikes increase, regulation may be
required, and the commenter requested
clarification on the ‘‘large number of
mortalities’’ cited in the proposed
listing rule. The commenter suggested
that if ship strikes have increased over
time, it could indicate the population of
Atlantic sturgeon in these areas has
increased.
Response: The ASMFC’s comment on
monitoring of ship strikes on Atlantic
sturgeon is noted. In response to the
commenter requesting clarification on
‘‘a large number of mortalities’’ cited in
the proposed listing rule, the full
statement on page 61924 is ‘‘a large
number of mortalities observed in these
rivers from potential ship strikes have
been of large adult Atlantic sturgeon.’’
The sentence is not indicating that there
are a large number of ship strike
mortalities, but rather a large percentage
of the mortalities resulting from ship
strikes are large adult fish. On the
following page (61925), we quantified
ship strikes in the one river in the
Southeast where they have been
documented (‘‘one ship strike per 5
years is reported for the Cape Fear River
within the Carolina DPS.’’). This section
of the proposed listing rule further
noted that, while it is possible that ship
strikes may have occurred and have
gone unreported or unobserved, the lack
of large ship traffic on narrow
waterways within the range of the DPS
may limit potential interactions. We
concurred with the ASSRT’s assessment
of the threat from ship strikes as low for
both the Carolina and South Atlantic
DPS and concluded that it was not
contributing to the endangered status of
the DPSs. An increase in ship strikes on
Atlantic sturgeon could result from a
variety of factors, including an increase
in Atlantic sturgeon populations, an
increase in shipping traffic, changes to
shipping channel characteristics (e.g.,
channel shallowing or narrowing), and
transit of larger vessels. If NMFS
receives new data showing that ship
strikes pose a significant threat to the
Carolina or South Atlantic DPS, we will
work with stakeholders, including the
shipping industry, to evaluate the best
options for minimizing impacts to
Atlantic sturgeon without unduly
hampering shipping activities.
Comment 52: A commenter agreed
with concerns expressed in the
proposed listing rule about the effects of
aquaculture and stock enhancement on
Atlantic sturgeon populations from
disease, escape, and out-breeding
depression, but believes these can be
minimized and that a responsible
stocking program using native
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broodstock is the best option for
reestablishing a population in extirpated
systems. The commenter stated that
there is evident disdain by the ASSRT
for stocking and enhancement activities
based on the discussion of the dangers
of stocking and categorizing the release
of cultured fish as a section 9 violation.
The commenter believed listing would
result in currently captive fish being
destroyed rather than used for
commercial or stocking purposes and
would provide no incentive for the
private sector to maintain the fish they
currently have. The commenter also
believed aquaculture and stock
enhancement afford research
opportunities and would afford a level
of protection for wild stocks from
poaching by providing a legal product to
the market. FDACS, which certifies
aquaculture facilities and inspects those
facilities twice a year for compliance,
commented on the proposed listing rule.
They stated that NOAA participated in
a cultured sturgeon risk analysis in 2000
that governs Florida sturgeon farming
and that the disposition of the Atlantic
sturgeon in aquaculture facilities is
known, contrary to information reported
in the proposed listing rule. FDACS
indicated that captive Atlantic sturgeon
in Florida are from a genetically distinct
population that is not being considered
for listing and were cultured in waters
outside those being defined as within
the South Atlantic DPS. The commenter
stated that sturgeon products sold by
Florida farms possess an Aquaculture
Certificate of Registration and are
exempt from the provisions of the ESA.
Response: Both the proposed listing
rule and the 2007 status review report
presented an objective discussion of
stocking and enhancement and did not
reflect disdain on the part of the agency
or the ASSRT for those activities. Both
documents state that artificial
propagation has the potential to be a
tool for recovery of the species, as well
as a threat. While collecting, handling,
releasing, and harming captive Atlantic
sturgeon were identified in the
proposed listing rule as potential
violations of the take prohibitions in
section 9 of the ESA, we also stated that
permits are available to lawfully
conduct these activities for purposes of
scientific research or to enhance the
propagation of the or survival of
Atlantic sturgeon DPSs. As stated in our
response to other comments above, we
must base listing determinations solely
on the best scientific and commercial
data available on the status of and
threats to the species. We cannot
consider the potential economic
consequences (or lack of economic
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incentives) to entities currently in
possession of captive Atlantic sturgeon.
However, in our response to comment
35, we describe the types of
authorizations available to conduct
activities such as artificial propagation
in full compliance with the ESA and we
encourage the affected parties to utilize
this option.
In response to comments from FDACS
that the disposition of all Atlantic
sturgeon acquired by Florida
aquaculture facilities is known, we
contacted FDACS to confirm current
holdings. FDACS informed us that
Evan’s Fish Farm is currently the only
facility in Florida with Atlantic
sturgeon, and they only have one
surviving fish. All other Atlantic
sturgeon held in Florida aquaculture
facilities died in captivity. We are
updating the final listing rule with this
information. As stated in the proposed
listing rule and confirmed by the
commenter, Atlantic sturgeon in
possession of Florida aquaculture
facilities originated from Canadian
populations and not from any of the
U.S. DPSs currently being proposed for
listing under the ESA. Therefore, the
remaining Atlantic sturgeon held by
Evan’s Fish Farm is not affected by the
listing.
Comment 53: A commenter provided
information on impingement of juvenile
Atlantic sturgeon at the Brunswick
Nuclear Power Plant on the lower Cape
Fear River. Plant modifications were
implemented in the early 1980s as part
of the NPDES permit. An average of 55
juvenile Atlantic sturgeon were
impinged per year from 1975 to 1981. A
fish diversion was installed in 1981 and
a fish return system was installed in
1983. Only 2 impinged juveniles were
observed between 1982 and 2010 and
were returned alive to the river.
Response: We appreciate the
information provided by the commenter
and we added this information to the
section of the final listing rule on
impingement and entrainment. As we
noted in the proposed listing rule, the
withdrawal of water from rivers that
support Atlantic sturgeon populations
was considered to pose a potential
threat of impingement and entrainment;
however, data are lacking to determine
the overall impact of this threat on
sturgeon populations, as impacts are
dependent on a variety of factors (e.g.,
the species, time of year, location of the
intake structure, and strength of the
intake current). Prior to receiving the
above information, we only had one
survey showing the direct impact of
water withdrawal on Atlantic sturgeon
in the Southeast. As stated in the
proposed listing rule, the Edwin I.
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Hatch Nuclear power plant, located 11
miles north of Baxley, Georgia,
withdraws water from, and discharges
to, the Altamaha River. Pre-operational
drift surveys were conducted and only
two Acipenser larvae were collected.
Entrainment samples at the plant were
collected for the years 1975, 1976, and
1980, and no Acipenser species were
observed in the samples (Sumner, 2004).
We concurred with the ASSRT’s
assessment of the threat from
impingement and entrainment as low
for both the Carolina and South Atlantic
DPS and concluded that it was not
contributing to the endangered status of
the DPSs. The information provided by
the commenter that two juvenile
Atlantic sturgeon were impinged at the
Brunswick Nuclear Power Plant
between 1982 and 2010, and both were
returned to the Cape Fear River alive,
does not change our conclusion.
Comment on Recovery
Comment 54: A comment was
received that a recovery plan for
Atlantic sturgeon should place a high
priority on research and gathering
sufficient information to define what it
means to both jeopardize and recover
Atlantic sturgeon and define the
allowable take authorized by the ESA.
Ecosystem dynamics and level of
anthropogenic activity vary in each
river, and recovery tasks should be
prioritized based on research into
potential impacts of the activities on
Atlantic sturgeon. The commenter
recommended an accelerated and
concentrated research effort prior to
development of a targeted restoration
strategy.
Response: Section 4(f) of the ESA
directs NMFS to develop and
implement recovery plans for
threatened and endangered species,
unless such a plan would not promote
conservation of the species. According
to the statute, these plans must
incorporate, at a minimum: (1) A
description of site-specific management
actions necessary to achieve recovery of
the species, (2) objective, measurable
criteria which, when met, would result
in a determination that the species be
removed from the list; and (3) estimates
of the time and costs required to achieve
the plan’s goal. NMFS agrees with the
commenter that research to fill
knowledge gaps in areas important to
recovery should be a priority. NMFS is
currently undertaking and funding a
variety of projects, including research
on abundance and to determine
movement and habitat utilization by
Atlantic sturgeon. In addition, through
years of section 7 consultations on
shortnose sturgeon, which share many
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of the same rivers as Atlantic sturgeon,
we have much information on
anthropogenic activities occurring in
those rivers. We will continue to seek
information on Atlantic sturgeon, their
habitat, and the threats they are facing
and use this information to prioritize
recovery actions. Once a draft recovery
plan is developed, we will submit it for
public review and comment before
finalizing it.
Comments on Critical Habitat
Comment 55: A commenter
recommended that confirmed and
potential nursery and spawning
locations in each river should be
designated as critical habitat for Atlantic
sturgeon, as well as known marine
migration corridors and aggregation
areas. The commenter provided
information and literature citations
identifying some of these areas and the
habitat characteristics potentially
preferred by Atlantic sturgeon.
Response: We appreciate the
information provided by the
commenter. Section 4(a)(3)(A) of the
ESA requires that critical habitat be
designated, to the maximum extent
prudent and determinable, concurrently
with a determination that a species is
endangered or threatened. When such a
designation is not determinable at the
time of final listing of a species, section
4(b)(6)(C)(ii) of the ESA provides for an
additional year to promulgate a critical
habitat designation. We have concluded
that critical habitat for the Carolina and
South Atlantic DPSs is not determinable
at this time. Through the status review
and public comment process on the
proposed listing rule, we have begun to
collect information on the location of
biological and physical and biological
features essential to the conservation of
the two DPSs. Throughout the next year,
we intend to gather and review current
and ongoing studies on the habitat use
and requirements of Atlantic sturgeon
from the two DPSs in the Southeast,
including an ongoing study with USGS
to compare sturgeon location data with
a variety of habitat parameters and a
study to map riverine habitat in four
Georgia rivers known to support the
South Atlantic DPS funded through
NMFS Section 6 program. We will also
gather and analyze information on the
benefits and impacts of a critical habitat
designation.
Comment 56: A comment stated that
critical habitat for the South Atlantic
DPS should be accurately defined. The
commenter noted that Figure 2 in the
proposed listing rule depicts habitat
well above the fall line and stated that
accurate delineation of critical habitat is
necessary so undue compliance costs
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are not placed on communities outside
the actual habitat utilized by the DPS.
Response: NMFS has not yet
designated critical habitat for the
Carolina and South Atlantic DPSs. The
shaded areas in Figure 2 in the proposed
listing rule encompass the rivers where
Atlantic sturgeon belonging to the
Carolina and South Atlantic DPSs may
occur. The shaded areas were made
sufficiently large so that no rivers or
tributaries potentially inhabited by fish
from the Carolina and South Atlantic
DPSs were excluded. The shaded areas
were meant to be a visual reference,
rather than a definitive indication of the
presence of Atlantic sturgeon, though
any sturgeon encountered in a location
within a shaded area would be from a
DPS being listed through this final rule
(as all Atlantic sturgeon on the East
Coast of the U.S. are from a DPS
currently proposed for listing as
threatened or endangered). We have
modified Figure 2, now Figures 2 and 3,
to more accurately reflect the text
descriptions of the Carolina and South
Atlantic DPSs. Because Atlantic
sturgeon are included in a DPS based on
the watershed in which they spawn or
were spawned, we have redrawn
Figures 2 and 3 using HUC 8 watershed
boundaries obtained from USGS.
Because this is only a visual
representation of where fish from the
Carolina and South Atlantic DPSs may
be encountered, it does not change the
entities being listed and does not
indicate that critical habitat may be
designated in a certain location. We
agree that critical habitat for the
Carolina and South Atlantic DPSs
should be accurately defined at the time
of designation to ensure compliance
with the ESA’s mandate at section
7(a)(2) that any activity authorized,
funded, or carried out by a Federal
agency is not likely to result in the
destruction or adverse modification of
critical habitat. Though activities
occurring outside designated critical
habitat can still affect critical habitat in
some instances, NMFS does not have
the authority or the intent to place
compliance burdens on entities engaged
in activities that would not adversely
affect Atlantic sturgeon or their
designated critical habitat.
Comment 57: A commenter stated that
NMFS has not designated critical
habitat for the endangered shortnose
sturgeon, which would improve habitat
protection for the Atlantic sturgeon due
to the substantial overlap in habitat
utilization between the two species. The
commenter stated NMFS should meet
the management objectives of the ESA
for shortnose sturgeon before taking on
the substantial administrative burden of
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a listing for Atlantic sturgeon and said
that species with critical habitat
designated are twice as likely to be
recovered as species without critical
habitat. Another commenter questioned
why NOAA failed to identify Essential
Fish Habitat (EFH) for Atlantic sturgeon
to support the proposed listing rule and
noted that EFH and Habitat Areas of
Particular Concern (HAPCs) have not
been designated for shortnose sturgeon
either.
Response: The shortnose sturgeon was
listed as endangered on March 11, 1967,
under the Endangered Species
Preservation Act of 1966, a predecessor
to the Endangered Species Act of 1973.
Shortnose sturgeon continued to meet
the listing criteria for endangered under
subsequent definitions specified in the
1969 Endangered Species Conservation
Act and remained on the list with the
inauguration of the ESA in 1973. NMFS
later assumed jurisdiction for shortnose
sturgeon under a 1974 government
reorganization plan (38 FR 41370).
Because the shortnose sturgeon was
listed prior to the amendments to the
ESA that made critical habitat
designations mandatory for newly listed
species, NMFS is not required to
designate critical habitat for the species
(designation is discretionary). However,
NMFS has undertaken a number of
activities to protect shortnose sturgeon
and their habitat, including publishing
a recovery plan for the species (63 FR
69613; December 17, 1998), funding
research on the species, and consulting
with Federal agencies under section 7 of
the ESA to ensure shortnose sturgeon
are not jeopardized by activities that
may harm the fish or their habitat. Some
of these efforts also benefit Atlantic
sturgeon, as noted in the proposed
listing rule. However, NMFS cannot
delay a listing determination or a
critical habitat designation for Atlantic
sturgeon until the recovery objectives
for shortnose sturgeon are met. Because
NMFS was petitioned to list the Atlantic
sturgeon, we were required to evaluate
the status of the species and the threats
it is facing and make a finding on
whether the petitioned action was
warranted within 12 months, which
resulted in our proposed listing rule
determination of endangered for the
Carolina and South Atlantic DPSs of
Atlantic sturgeon. NMFS works with the
regional fishery management councils to
identify EFH and HAPCs for federally
managed fishery species. Atlantic and
shortnose sturgeon are not federally
managed fishery species, therefore
NMFS did not identify EFH or HAPCs
for either species.
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5953
Comments on the Public Hearings
Comment 58: A commenter asked if
the ‘‘limited advertising’’ and one
public hearing met the minimum
statutory requirements for receiving
public comments on the proposed
listing rule, since it affects a large
geographic area, numerous counties,
cities, states, industries, etc. The
commenter stated the meeting was not
on the Southeast Region’s or the Office
of Protected Resources’ Web site. The
commenter noted that a legal notice was
placed in the local newspaper, but he
asked the paper to enlarge the notice
and to also include a separate article
about the public hearing.
Response: The notice and public
comment period on the proposed listing
rule for the Carolina and South Atlantic
DPSs of Atlantic sturgeon exceeded the
requirements established in section
4(b)(5) of the ESA. The proposed listing
rule established a 90-day comment
period (October 6, 2010, through
January 4, 2011), during which
comments were accepted electronically
via the Federal eRulemaking Portal
(https://www.regulations.gov), as well as
by mail, hand delivery, and facsimile.
We extended the comment period an
additional 30 days at the request of the
public and accepted comments through
February 4, 2011. In compliance with
section 4(b)(5)(A)(ii), we sent over 200
letters with a complete copy of the
proposed rule to each relevant state and
county agency where the Carolina and
South Atlantic DPS potentially occur,
inviting them to comment on the
proposed listing rule. Section 4(b)(5)(E)
of the ESA only requires that one public
hearing be held on a proposed listing
rule if it is requested by the public
within 45 days after the date of the
publication of the proposed listing rule
in the Federal Register. Though the
Southeast Region did not receive any
requests for a public hearing, we elected
to hold two public hearings, one each in
the areas occupied by the Carolina and
South Atlantic DPSs of Atlantic
sturgeon. Hearings were held in
Wilmington, North Carolina, on
December 6, 2010, and Atlanta, Georgia,
on December 7, 2010, to accept public
comments. In addition to publishing a
notice in the Federal Register (75 FR
69049; November, 10, 2010) announcing
the hearings, a notice was placed in the
legal section of a major newspaper in
each of the five states occupied by the
Carolina and South Atlantic DPSs on
November 15, 2010: the Florida TimesUnion (Florida), the Atlanta Journal
Constitution (Georgia), The State (South
Carolina), The Charlotte Observer
(North Carolina), and the Richmond
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Times-Dispatch (Virginia). As the
commenter noted, we also placed a
notice in the local paper, The StarNews, for Wilmington, North Carolina,
where the first hearing was held. An
announcement with a link to the
Federal Register notice for the hearings
was placed on the Southeast Regional
Office’s Web site on December 2, 2010.
Summary of Changes From the
Proposed Listing Rule
Based on the comments received and
our review of the proposed rule, we
made the changes listed below.
1. We refined the text descriptions of
the watersheds making up the ranges of
the Carolina and South Atlantic DPSs
and the individual fish that are included
in the DPSs. The modifications to the
text only clarify the riverine ranges of
the DPSs and do not change the
spawning populations making up each
of the Southeast DPSs.
2. We slightly extended the marine
range of the DPSs based on recent
tagging data. We also provided refined
maps showing the riverine ranges of the
Carolina and South Atlantic DPSs using
HUC 8 watershed boundaries.
3. We added information on
metapopulations and the importance of
multiple viable riverine populations to
the ‘‘Conservation Status’’ section, per
our response to comment 1 from peer
reviewers 1 and 2.
4. We added information on the role
of adaptation and competition in the
observed low rate of genetic exchange
between Atlantic sturgeon river
populations, per comment 5 submitted
by peer reviewer 1.
5. We added information on
polyploidy in Atlantic sturgeon and
potential effects on the evaluation of
minimum viable population size, per
comment 7 submitted by a peer
reviewer 1.
6. We added information on recent
estimated increases in juvenile Atlantic
sturgeon abundance in the Altamaha
River, Georgia, per comment 2
submitted by peer reviewer 2.
7. We added information about the
nature of the samples used in the
genetic analysis for the Waccamaw
River population, per comment 13
submitted by peer reviewer 3.
8. We added Atlantic sturgeon
location and abundance data provided
by the states (North Carolina, South
Carolina, and Georgia) for the Carolina
and South DPS to the ‘‘Distribution and
Abundance’’ section, per comments 19
and 20.
9. We revised the erroneous statement
in the section on climate change that the
Carolina and South Atlantic DPSs are
within a region the IPCC predicts will
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experience decreases in precipitation.
As noted in our response to comment
45, the Southeast is predicted to
experience increases in precipitation;
however, evaporation is also predicted
to increase with increasing temperatures
and the net effect for the Southeast is
predicted to be overall drying.
10. We added and updated Atlantic
sturgeon bycatch information in Section
B ‘‘Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes’’ with information provided by
the states of North Carolina and Georgia,
per comment 47.
11. We removed the statement that it
is unlikely the aquarium industry could
ever be effectively regulated, per
comment 48.
12. We updated information on the
current holdings of Atlantic sturgeon in
Florida aquaculture facilities.
13. We corrected the location of the
Bears Bluff National Fish Hatchery.
14. We added information on
impingement and entrainment of
juvenile Atlantic sturgeon at the
Brunswick Nuclear Power Plant on the
lower Cape Fear River.
15. We added an evaluation of North
Carolina’s NCCHPP and designation of
AFSAs to our evaluation of current
protective efforts.
16. We made minor corrections and
updates to information in the listing
rule based on recommendations from
peer reviewers, commenters, and our
own review of the proposed listing rule.
Our listing determination and
summary of the data on which it is
based, with the incorporated changes,
are presented in the remainder of this
document.
Taxonomy and Life History
There are two subspecies of Atlantic
sturgeon—the Gulf sturgeon (Acipenser
oxyrinchus desotoi) and the Atlantic
sturgeon (Acipenser oxyrinchus
oxyrinchus). Historically, the Gulf
sturgeon occurred from the Mississippi
River east to Tampa Bay. Its present
range extends from Lake Pontchartrain
and the Pearl River system in Louisiana
and Mississippi east to the Suwannee
River in Florida. The Gulf sturgeon was
listed as threatened under the ESA in
1991. The finding in this final rule
addresses the subspecies Acipenser
oxyrinchus oxyrinchus (referred to as
Atlantic sturgeon), which is distributed
along the eastern coast of North
America. Historically, sightings have
been reported from Hamilton Inlet,
Labrador, south to the St. Johns River,
Florida. Recently, a tagged Atlantic
sturgeon was tracked off Cape
Canaveral, Florida. Occurrences south
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of the St. Johns River, Florida, and in
Labrador may have always been rare.
Atlantic sturgeon is a long-lived, latematuring, estuarine-dependent,
anadromous species. Atlantic sturgeon
may live up to 60 years, reach lengths
up to 14 feet (ft; 4.27 meters (m)), and
weigh over 800 pounds (lbs; 363 kg).
They are distinguished by armor-like
plates and a long protruding snout that
is ventrally located, with four barbels
crossing in front. Sturgeon are
omnivorous benthic (bottom) feeders
and filter quantities of mud along with
their food. Adult sturgeon diets include
mollusks, gastropods, amphipods,
isopods, and fish. Juvenile sturgeon feed
on aquatic insects and other
invertebrates (ASSRT, 2007).
Vital parameters of Atlantic sturgeon
populations show clinal variation with
faster growth and earlier age at
maturation in more southern systems,
though not all data sets conform to this
trend. Atlantic sturgeon mature between
the ages of 5 and 19 years in South
Carolina (Smith et al., 1982), between
11 and 21 years in the Hudson River
(Young et al., 1988), and between 22
and 34 years in the St. Lawrence River
(Scott and Crossman, 1973). Atlantic
sturgeon likely do not spawn every year.
Multiple studies have shown that
spawning intervals range from 1 to 5
years for males (Smith, 1985; Collins et
al., 2000; Caron et al. 2002) and 2 to 5
years for females (Vladykov and
Greeley, 1963; Van Eenennaam et al.,
1996; Stevenson and Secor, 1999).
Fecundity of Atlantic sturgeon has been
correlated with age and body size, with
egg production ranging from 400,000 to
8 million eggs per year (Smith et al.,
1982; Van Eenennaam and Doroshov,
1998; Dadswell, 2006). The average age
at which 50 percent of maximum
lifetime egg production is achieved is
estimated to be 29 years, approximately
3 to 10 times longer than for other bony
fish species examined (Boreman, 1997).
Spawning adults migrate upriver in
the spring, which occurs during
February and March in southern
systems, April and May in mid-Atlantic
systems, and May and July in Canadian
systems (Murawski and Pacheco, 1977;
Smith, 1985; Bain, 1997; Smith and
Clugston, 1997; Caron et al., 2002). In
some southern rivers, a fall spawning
migration may also occur (Rogers and
Weber, 1995; Weber and Jennings, 1996;
Moser et al., 1998). Spawning typically
occurs in flowing water between the salt
front and fall line of large rivers, where
optimal flows are 18 to 30 inches (in)
per second (46 to 76 centimeters (cm)
per second) and depths are 36 to 89 ft
(11 to 27 m) (Borodin, 1925; Leland,
1968; Scott and Crossman, 1973; Crance,
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1987; Bain et al., 2000). The fall line is
the boundary between an upland region
of continental bedrock and an alluvial
coastal plain, sometimes characterized
by waterfalls or rapids. Sturgeon eggs
are highly adhesive and are deposited
on the bottom substrate, usually on hard
surfaces (e.g., cobble) (Gilbert, 1989;
Smith and Clugston, 1997). Hatching
occurs approximately 94 to 140 hours
after egg deposition at corresponding
temperatures of 68.0 to 64.4 degrees
Fahrenheit (20 to 18 degrees Celsius).
The newly emerged larvae assume a
demersal existence (Smith et al., 1980).
The yolksac larval stage is completed in
about 8 to 12 days, during which time
the larvae move downstream to rearing
grounds (Kynard and Horgan, 2002).
During the first half of their migration
downstream, movement is limited to
night. During the day, larvae use benthic
structure (e.g., gravel matrix) as refugia
(Kynard and Horgan, 2002). During the
latter half of migration, when larvae are
more fully developed, movement to
rearing grounds occurs both day and
night. Juvenile sturgeon continue to
move further downstream into brackish
waters and eventually become residents
in estuarine waters for months to years.
Recovery of depleted populations is
an inherently slow process for a latematuring species such as Atlantic
sturgeon. Their late age at maturity
provides more opportunities for
individuals to be removed from the
population before reproducing.
However, a long life-span also allows
multiple opportunities to contribute to
future generations provided the
appropriate spawning habitat and
conditions are available.
Distribution and Abundance
Historically, Atlantic sturgeon were
present in approximately 38 river
systems throughout their range, of
which 35 systems have been confirmed
to have had a historical spawning
population. More recently, presence has
been documented in 35 river systems
with spawning taking place in at least
18 rivers. Spawning has been confirmed
in the St. Lawrence, Annapolis, St. John,
Kennebec, Hudson, Delaware, James,
Roanoke, Tar-Pamlico, Cape Fear,
Waccamaw, Great Pee Dee, Combahee,
Edisto, Savannah, Ogeechee, Altamaha,
and Satilla rivers. Rivers with possible,
but unconfirmed, spawning populations
include the St. Croix, Penobscot,
Androscoggin, Sheepscot, York, Neuse,
Santee and Cooper Rivers.
Historical records from the 1700s and
1800s document large numbers of
sturgeon in many rivers along the
Atlantic Coast. Atlantic sturgeon
underwent significant range-wide
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declines from historical abundance
levels due to overfishing in the late
1800s, as discussed more fully below.
Sturgeon stocks were further impacted
through environmental degradation,
especially due to loss of access to
habitat and reduced water quality from
the construction of dams in the early to
mid-1900s. The species persisted in
many rivers, though at greatly reduced
levels (1 to 5 percent of their earliest
recorded numbers), and commercial
fisheries were active in many rivers
during all or some of the years 1962 to
1997 (Waldman and Wirgin, 1998;
Smith and Clugston, 1997). Many of
these contemporary fisheries resulted in
continued overfishing, which prompted
ASMFC to impose the Atlantic sturgeon
fishing moratorium in 1998 and NMFS
to close the EEZ to Atlantic sturgeon
retention in 1999.
Quantified abundance estimates of
Atlantic sturgeon obtained through
sampling surveys are currently only
available for the Hudson (NY) and
Altamaha (GA) rivers, where adult
spawning populations are estimated to
be approximately 870 and 343 fish per
year, respectively (Kahnle et al., 2007;
Schueller and Peterson, 2006). Surveys
from other rivers in the species’ U.S.
range are more qualitative, primarily
focusing on documentation of multiple
year classes and reproduction, as well as
the presence of very large adults and
gravid females, in the river systems. In
the Southeast Region, spawning has
been confirmed in 11 rivers (Roanoke,
Tar-Pamlico, Cape Fear, Waccamaw,
Great Pee Dee, Combahee, Edisto,
Savannah, Ogeechee, Altamaha, and
Satilla rivers), with possible spawning
occurring in 3 additional rivers (the
Neuse, Santee and Cooper Rivers).
Based on a comprehensive review of the
available data, the literature, and
information provided by local, state,
and Federal fishery management
personnel, the Altamaha River is
believed to have the largest population
in the Southeast (ASSRT, 2007). The
larger size of this population relative to
the other river populations in the
Southeast is likely due to the absence of
dams, the lack of heavy development in
the watershed, and relatively good water
quality, as Atlantic sturgeon
populations in the other rivers in the
Southeast have been affected by one or
more of these factors. Trammel net
surveys, as well as independent
monitoring of incidental take in the
American shad fishery, suggested that
the Altamaha population was neither
increasing nor decreasing. However,
recent studies by Schueller and Peterson
(2010) and Peterson (2011; UGA, pers.
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comm.) estimated large increases in
abundance of Atlantic sturgeon
juveniles from 2004–2010, particularly
during the 2009–2010 period. Schueller
and Peterson (2010) conducted their
research during the summers of 2004 to
2007 and estimated that juvenile
abundance ranged from 1,072 to 2,033
individuals in the Altamaha River, with
age-1 and age-2 individuals comprising
greater than 87 percent of the
population. Based on modeling,
estimated apparent survival and per
capita recruitment indicated that the
juvenile population experienced high
annual turnover: apparent survival rates
were low (less than 33 percent), and per
capita recruitment was high (0.82–1.38).
The numbers of juvenile Atlantic
sturgeon in the Altamaha River in 2009
and 2010 were between approximately
3,500 and 6,500. However, the authors
noted that their mark–recapture
methods were not capable of providing
separate estimates of annual survival
and out-migration, yet these rates are
critical in understanding recruitment
processes for the species. Though
quantitative abundance estimates
obtained through sampling surveys are
not available for the other river
populations, because the Altamaha
spawning population is the largest, we
believe a conservative estimate of the
other spawning populations in the
Southeast Region is no more than 300
adults spawning per year.
Historically, Atlantic sturgeon were
abundant in most North Carolina coastal
rivers and estuaries, with the largest
fisheries occurring in the Roanoke
River/Albemarle Sound system and in
the Cape Fear River (Kahnle et al.,
1998). Historical landings records from
the late 1800s indicated that Atlantic
sturgeon were very abundant within
Albemarle Sound (approximately
135,600 lbs or 61,500 kg landed per
year). Abundance estimates derived
from these historical landings records
indicated that between 7,200 and 10,500
adult females were present within North
Carolina prior to 1890 (Armstrong and
Hightower, 2002; Secor 2002). NCDMF
has conducted the Albemarle Sound
IGNS, initially designed to target striped
bass, since 1990. During that time, 842
YOY and subadult sturgeon have been
captured. Incidental take of Atlantic
sturgeon in the IGNS, as well as
multiple observations of YOY from the
Albemarle Sound and Roanoke River,
provide evidence that spawning
continues. Three adult Atlantic sturgeon
(2 males, 1 unknown) were tagged in the
Roanoke River during September 2010
and the fish were tracked out of the
river several weeks later, potentially
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suggesting a fall spawning run of
Atlantic sturgeon in the Roanoke River.
Catch records indicate that the Roanoke
River Atlantic sturgeon population
seemed to be increasing until 2000,
when recruitment began to decline. The
Albemarle IGNS data for 2006–2009
showed higher Atlantic sturgeon CPUEs
(0.015 to 0.031) than the 2002–2005
period, though they were still lower
than the 2000–2001 level (0.032) and
there is no overall trend in the overall
1990–2009 CPUE dataset. Catch records
and observations from other river
systems in North Carolina exist (e.g.,
Hoff, 1980, Oakley, 2003, in the Tar and
Neuse rivers; Moser et al., 1998, and
Williams and Lankford, 2003, in the
Cape Fear River) and provide evidence
for spawning, but based on the
relatively low numbers of fish caught, it
is difficult to determine whether the
populations in those systems are
declining, rebounding, or remaining
static. The Pamlico IGNS survey data
from 2001–2009 shows peak CPUE of
Atlantic sturgeon in 2005 (0.095), but no
decreasing or increasing trends are
apparent. River surveys in the Pamlico,
Pungo, and Neuse Rivers since 2000
have shown a slight decrease in Atlantic
sturgeon abundance. Also, large survey
captures during a single year are
difficult to interpret. For instance,
abundance of Atlantic sturgeon below
Lock and Dam #1 in the Cape Fear River
seemed to have increased dramatically
during the 1990–1997 surveys (Moser et
al., 1998) as the CPUE of Atlantic
sturgeon was up to eight times greater
during 1997 than in the earlier survey
years. Since 1997, Atlantic sturgeon
CPUE doubled between the years of
1997 and 2003 (Williams and Lankford,
2003). However, it is unknown whether
this is an actual population increase
reflecting the effects of North Carolina’s
ban on Atlantic sturgeon fishing that
began in 1991, or whether the results
were skewed by one outlier year. There
was a large increase observed in 2002,
though the estimates were similar
among all other years of the 1997 to
2003 study.
Atlantic sturgeon were likely present
in many South Carolina river/estuary
systems historically, but it is not known
where spawning occurred. Secor (2002)
estimated that 8,000 spawning females
were likely present prior to 1890, based
on U.S. Fish Commission landing
records. Since the 1800s, however,
populations have declined dramatically
(Collins and Smith, 1997). Recorded
landings of Atlantic sturgeon in South
Carolina peaked at 481,050 lbs (218,200
kg) in 1897, but 5 years later, only
93,920 lbs (42,600 kg) were reported
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landed (Smith et al., 1984). Landings
remained depressed throughout the
1900s, with between 4,410 and 99,210
lbs (2,000 and 45,000 kg) of Atlantic
sturgeon reported annually between
1958 and 1982 (Smith et al., 1984).
During the last two decades, Atlantic
sturgeon have been observed in most
South Carolina coastal rivers, although
it is not known if all rivers support a
spawning population (Collins and
Smith, 1997). Sampling for shortnose
sturgeon (Acipenser brevirostrum)
conducted in Winyah Bay captured two
subadult Atlantic sturgeon in 2004.
SCDNR noted in comments on the
proposed listing rule that Atlantic
sturgeon were captured in most nets set
in Winyah Bay from April to July in
2007 to 2009, including sites far upriver.
Further, a researcher conducting pilot
sonar survey trials in Winyah Bay
potentially detected several hundred
fish, many of which could be Atlantic
sturgeon. The researcher has conducted
pilot sonar trials in the Roanoke, Neuse,
Cape Fear, and Pee Dee River systems
and believes the initial results suggest
Atlantic sturgeon densities in the Pee
Dee River (Winyah Bay) system are
higher than the other systems surveyed
(J. Hightower, pers. comm.) Captures of
age-1 juveniles from the Waccamaw
River during the early 1980s suggest that
a reproducing population of Atlantic
sturgeon may persist in that river,
although the fish could have been from
the nearby Great Pee Dee River (Collins
and Smith, 1997). Until recently, there
was no evidence that Atlantic sturgeon
spawned in the Great Pee Dee River,
although subadults were frequently
captured and large adults were often
observed by fishers. However, a fishery
survey conducted by Progress Energy
Carolinas Incorporated captured a
running ripe male in October 2003 and
observed other large sturgeon, perhaps
revealing a fall spawning run (ASSRT,
2007). There are no data available
regarding the presence of YOY or
spawning adult Atlantic sturgeon in the
Sampit River, although it did
historically support a population and is
thought to serve as a nursery ground for
local stocks (ASMFC, 2009).
The Santee-Cooper system had some
of the highest historical landings of
Atlantic sturgeon in the Southeast. Data
from the U.S. Fish Commission shows
that greater than 220,460 lbs (100,000
kg) of Atlantic sturgeon were landed in
1890 (Secor, 2002). The capture of 151
subadults, including age-1 juveniles, in
the Santee River in 1997 suggests that
an Atlantic sturgeon population still
exists in this river (Collins and Smith,
1997). The status review report
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documents that three adult Atlantic
sturgeon carcasses were found above the
Wilson and Pinopolis dams in Lake
Moultrie (a Santee-Cooper reservoir)
during the 1990s, and also states that
there is little information regarding a
land-locked population existing above
the dams. There is no effective fish
passage for sturgeon on the Santee and
Cooper Rivers, and the lowest dams on
these rivers are well below the fall line,
thus limiting the amount of freshwater
spawning and developmental habitat for
fish below the dams. In 2007, an
Atlantic sturgeon entered the fish lift (a
lock designed specifically for fish
passage) at the St. Stephen dam; it was
physically removed and translocated
downstream into the Santee River (A.
Crosby, SCDNR, pers. comm.) In 2004,
15 subadult Atlantic sturgeon were
captured in shortnose sturgeon surveys
in the Santee River estuary. The
previous winter, four juvenile (YOY and
subadults) Atlantic sturgeon were
captured from the Santee (one fish) and
Cooper (three fish) rivers. These data
support previous hypotheses that a fall
spawning run occurs within this system,
similar to that observed in other
southern river systems. However, the
status review report notes that SCDNR
biologists have some doubt whether
smaller sturgeon from the SanteeCooper are resident YOY, as flood
waters from the Pee-Dee or Waccamaw
Rivers could have transported these
YOY to the Santee-Cooper system via
Winyah Bay and the Intracoastal
Waterway (McCord, 2004). Resident
YOY could, however, be evidence of a
spawning population above the dams, as
is the case with shortnose sturgeon
(Collins et al., 2003)
From 1994 to 2001, over 3,000
juveniles have been collected in the
Ashepoo-Combahee-Edisto Rivers (ACE)
Basin, including 1,331 YOY sturgeon
(Collins and Smith, 1997; ASSRT,
2007). Specifically, SCDNR reports that
3,661 juvenile (one- to three-year-old)
Atlantic sturgeon were collected in the
Edisto River during the 16-year period
since 1994. Utilizing this data, SCDNR
used Lincoln-Peterson and Schnabel
models to derive Atlantic sturgeon
population estimates, which resulted in
estimates of 70,000 and 20,000 juvenile
Atlantic sturgeon in the Edisto River,
respectively. SCDNR commented that
the model results suggest increasing
trends in abundance. Both models rely
on mark-recapture data and assume a
closed population (there are no births,
deaths, or immigration/emigration
between the initial capture and the
recapture period) and that all
individuals have an equal chance of
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being captured (Nichols, 1992;
Lindeman, 1990; Chao, 1987). We note
that there is great uncertainty in the
population estimates resulting from the
two models, as evident in the great
disparity between the two numbers. The
reliability of the population models
depends on the validity of the
assumptions, and the primary
assumption of equal capture probability
is likely unattainable in natural
populations (Chao, 1987; Carothers,
1973). SCDNR indicated they are
currently completing an open system
model (which is based on survival
probabilities, as well as capture
probabilities) to better assess the
Atlantic sturgeon population in the
Edisto River. Sampling for adults began
in 1997, with two adult sturgeon
captured in the first year of the survey,
including one gravid female captured in
the Edisto River and one running ripe
male captured in the Combahee River.
The running ripe male in the Combahee
River was recaptured one week later in
the Edisto River, which suggests that the
three rivers that make up the ACE Basin
may support a single population that
spawns in at least two of the rivers.
Between 1997 and 1999, SCDNR
captured 118 adults in the Edisto River
during spring and fall spawning runs,
but netting ceased once that number
was reached, so the entire spawning run
was not sampled and more Atlantic
sturgeon may have been captured if
netting continued. SCDNR also noted
approximately 20 adults were captured
in the Edisto River over one to two
months during surveys targeting other
species. In 2010, four adults tagged in
the 1990’s as age 0+ were recaptured.
These captures show that a current
spawning population exists in the ACE
Basin, as both YOY and spawning
adults are regularly captured.
The Ashley River, along with the
Cooper River, drains into Charleston
Bay; only shortnose sturgeon have been
studied in these rivers. While the
Ashley River historically supported an
Atlantic sturgeon spawning population,
it is unknown whether the population
still exists. There has been little or no
scientific sampling for Atlantic sturgeon
in the Broad/Coosawatchie River. One
fish of unknown size was reported from
a small directed fishery during 1981 to
1982 (Smith and Dingley, 1984).
Prior to the collapse of the fishery in
the late 1800s, the sturgeon fishery was
the third largest fishery in Georgia.
Secor (2002) estimated from U.S. Fish
Commission landing reports that
approximately 11,000 spawning females
were likely present prior to 1890. The
sturgeon fishery was mainly centered on
the Altamaha River, and in more recent
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years, peak landings were recorded in
1982 (13,000 lbs, 5,900 kg). In Georgia,
Atlantic sturgeon are believed to spawn
in the Savannah, Ogeechee, Altamaha,
and Satilla rivers. Based on juvenile
presence and abundance, the Altamaha
River currently supports one of the
healthier Atlantic sturgeon populations
in the southeast (ASSRT, 2007). Atlantic
sturgeon are also present in the
Ogeechee River; however, the absence of
age-1 fish during some years and the
unbalanced age structure suggests that
the population is highly stressed (Rogers
and Weber, 1995). Sampling results
from the mid-1990s indicated that the
Atlantic sturgeon population in the
Satilla River was also highly stressed
(Rogers and Weber, 1995). Only four
spawning adults or YOY, which were
used for genetic analysis (Ong et al.,
1996), had been collected from this river
since 1995. In the most recent
compliance report to ASMFC,
University of Georgia (UGA) researchers
collected more than 200 Atlantic
sturgeon in the Satilla River in less than
2 years of sampling. The presence of
juvenile fish measuring less than 50 cm
supports this is likely a self-sustaining,
spawning population. The Savannah
River supports a reproducing
population of Atlantic sturgeon (Collins
and Smith, 1997). According to NOAA’s
National Ocean Service, 70 Atlantic
sturgeon have been captured since 1999
(ASSRT, 2007). Twenty-two of these
fish have been YOY. A running ripe
male was captured at the base of the
dam at Augusta during the late summer
of 1997, which supports the hypothesis
that spawning occurs there in the fall.
In the Savannah River, the SCDNR
captured 369 Atlantic sturgeon between
1997 and 2010.
Reproducing Atlantic sturgeon
populations are no longer believed to
exist south of the Satilla River in
Georgia. Sampling of the St. Marys River
in the early 1990s failed to locate any
sturgeon, which suggests that the
spawning population may be extirpated
(Rogers et al., 1994). In January 2010, 12
sturgeon, believed to be Atlantic
sturgeon, were captured at the mouth of
the St. Marys during relocation trawling
associated with a dredging project (J.
Wilcox, Florida Fish and Wildlife
Conservation Commission, pers.
comm.), the first capture of Atlantic
sturgeon in the St. Marys in decades.
However, because they were not YOY or
adults captured upstream, these trawlcaptured sturgeon do not provide new
evidence of a spawning population in
the St. Marys. Researchers captured a
total of 9 Atlantic sturgeon in the St.
Marys River in March and June of 2010,
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based on a final report submitted to
NMFS on a project funded through our
Species Recovery Grant Program under
section 6 of the ESA. The report stated
that, based on the sizes of these
individuals, the researchers concluded
that none of these individuals were
‘‘river-residents’’. Though there was no
definitive proof that these individuals
had immigrated from other rivers, the
report stated that the absence of small,
river-resident juveniles suggests that
Atlantic sturgeon in the St. Marys
experienced complete recruitment
failure from 2007–2010. There have
been reports of Atlantic sturgeon tagged
in the Edisto River (South Carolina)
being recaptured in the St. Johns River,
indicating this river may serve as a
nursery ground; however, there are no
data to support the existence of a
current spawning population (i.e., YOY
or running ripe adults) in the St. Johns
(Rogers and Weber, 1995; Kahnle et al.,
1998). In response to the proposed
listing rule, Florida Fish and Wildlife
Commission reported that 30 subadults
(1 meter in length) were captured in the
St. Marys River in 20 months and two
juveniles (approximately 50
centimeters, age-1 or 2) were captured
in the St. Johns River in February 2011,
though these captures do not provide
new evidence of spawning based on the
size/age classes of sturgeon caught.
Identification of Distinct Population
Segments
The ESA’s definition of ‘‘species’’
includes ‘‘any subspecies of fish or
wildlife or plants, and any distinct
population segment of any species of
vertebrate fish or wildlife which
interbreeds when mature.’’ The high
degree of reproductive isolation of
Atlantic sturgeon (i.e., homing to their
natal rivers for spawning; ASSRT, 2007;
Wirgin et al., 2000; King et al., 2001;
Waldman et al., 2002), as well as the
ecological uniqueness of those riverine
spawning habitats, the genetic
differentiation amongst subpopulations,
and the differences in life history
characteristics, provide evidence that
discrete reproducing populations of
Atlantic sturgeon exist, which led the
Services to evaluate application of the
DPS policy in its 2007 status review. To
determine whether any populations
qualify as DPSs, we evaluated
populations pursuant to the joint DPS
policy, and considered: (1) The
discreteness of any Atlantic sturgeon
population segment in relation to the
remainder of the subspecies to which it
belongs; and (2) the significance of any
Atlantic sturgeon population segment to
the remainder of the subspecies to
which it belongs.
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Discreteness
The joint DPS policy states that a
population of a vertebrate species may
be considered discrete if it satisfies
either one of the following conditions:
(1) It is markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors
(quantitative measures of genetic or
morphological discontinuity may
provide evidence of this separation) or
(2) it is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of Section
4(a)(1)(D) of the ESA.
Atlantic sturgeon throughout their
range exhibit ecological separation
during spawning that has resulted in
multiple, genetically distinct,
interbreeding population segments.
Tagging studies and genetic analyses
provide the evidence of this ecological
separation (Wirgin et al., 2000; King et
al., 2001; Waldman et al., 2002; ASSRT,
2007; Grunwald et al., 2008). As
previously discussed, though adult and
subadult Atlantic sturgeon originating
from different rivers mix in the marine
environment (Stein et al., 2004a), the
vast majority of Atlantic sturgeon return
to their natal rivers to spawn, with some
studies showing only one or two
individuals per generation spawning
outside their natal river system (Wirgin
et al., 2000; King et al., 2001; Waldman
et al., 2002). In addition, spawning in
the various river systems occurs at
different times, with spawning
occurring earliest in southern systems
and occurring as much as 5 months later
in the northernmost river systems
(Murawski and Pacheco, 1977; Smith,
1985; Rogers and Weber, 1995; Weber
and Jennings, 1996; Bain, 1997; Smith
and Clugston, 1997; Moser et al., 1998;
Caron et al., 2002). Therefore, the
ecological separation of the
interbreeding units of Atlantic sturgeon
results primarily from spatial separation
(i.e., very few fish spawning outside
their natal river systems), as well as
temporal separation (spawning
populations becoming active at different
times along a continuum from north to
south).
Genetic analyses of mitochondrial
DNA (mtDNA), which is maternally
inherited, and nuclear DNA (nDNA),
which reflects the genetics of both
parents, provides evidence of the
separation among Atlantic sturgeon
populations in different rivers (Bowen
and Avise, 1990; Ong et al., 1996;
Waldman et al., 1996a; Waldman et al.,
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1996b; Waldman and Wirgin, 1998;
Waldman et al., 2002; King et al., 2001;
Wirgin et al., 2002; Wirgin et al., 2005;
Wirgin and King, 2006; Grunwald et al.,
2008). Overall, these studies
consistently found Atlantic sturgeon to
be genetically diverse, and offered that
between seven and ten Atlantic sturgeon
population groupings can be statistically
differentiated range-wide (King et al.,
2001; Waldman et al., 2002; Wirgin et
al., 2002; Wirgin et al., 2005; ASSRT,
2007 (Tables 4 and 5); Grunwald et al.,
2008).
Given a number of key differences
among the studies (e.g., the analytical
and/or statistical methods used, the
number of rivers sampled, and whether
samples from subadults were included),
it is not unexpected that each reached
a different conclusion regarding the
number of Atlantic sturgeon population
groupings. Wirgin and King (2006)
refined the genetic analyses for Atlantic
sturgeon to address such differences in
prior studies. Most notably, they
increased sample sizes from multiple
rivers and limited the samples analyzed
to those collected from YOY and mature
adults (greater than 130 cm total length)
to ensure that the fish originated from
the river in which it was sampled. The
results of the refined analysis by Wirgin
and King (2006) are presented in the
status review report (ASSRT, 2007; e.g.,
Table 6 and Figure 17); both the mtDNA
haplotype and nDNA allelic frequencies
analyzed by Wirgin and King (2006)
indicated that Atlantic sturgeon river
populations are genetically
differentiated. The results of the mtDNA
analysis used for the status review
report were also subsequently published
by Grunwald et al. (2008). In
comparison to the mtDNA analyses of
the status review report, Grunwald et al.
(2008) used additional samples, some
from fish in the size range (less than 130
cm) excluded by Wirgin and King
because they were smaller than those
considered to be mature adults.
Nevertheless, the results were
qualitatively the same and demonstrated
that each of the 12 sampled Atlantic
sturgeon populations could be
genetically differentiated (Grunwald et
al., 2008).
Genetic distances and statistical
analyses (bootstrap values and
assignment test values) were used to
investigate significant relationships
among, and differences between,
Atlantic sturgeon river populations
(ASSRT, 2007; Table 6 and Figures 16–
18). Overall, the genetic markers used in
this analysis resulted in an average
accuracy of 88 percent for determining
a sturgeon’s natal river origin, but an
average accuracy of 94 percent for
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correctly classifying it to one of five
groups of populations (Kennebec River,
Hudson River, James River, Albemarle
Sound, and Savannah/Ogeechee/
Altamaha Rivers) when using
microsatellite data collected only from
YOY and adults (ASSRT, 2007; Table 6).
The overall accuracy in assigning an
Atlantic sturgeon to its natal river
ranged from 60 to 94.8 percent (60 to
91.7 percent for southeastern rivers),
while the overall accuracy in identifying
a sturgeon to one of the 5 DPSs ranged
between 88.1 and 95.9 percent (91.7 to
95.9 percent for the two southeastern
DPSs). A phylogenetic tree (a neighbor
joining tree) was produced from only
YOY and adult samples (to reduce the
likelihood of including strays from other
populations) using the microsatellite
analysis (ASSRT, 2007; Figure 17).
Bootstrap values (which measure how
consistently the data support the tree
structure) for this tree were high (the
lowest was 87 percent, and all others
were over 90 percent) (ASSRT, 2007).
Regarding sturgeon from southeast
rivers, this analysis resulted in a range
of 60 to 92 percent accuracy in
determining a sturgeon’s natal river
origin, but 92 and 96 percent accuracy
in correctly classifying a sturgeon from
four sampled river populations (the
Albemarle Sound, Savannah, Ogeechee,
and Altamaha River populations) to two
groupings of river populations
(Albemarle Sound and Savannah/
Ogeechee/Altamaha Rivers). These two
groupings exhibited clear separation
from northern populations and from
each other.
Genetic samples for YOY and
spawning adults were not available for
river populations originating between
the Albemarle Sound and the other
three rivers. However, nDNA from an
expanded dataset that included juvenile
Atlantic sturgeon was used to produce
a neighbor-joining tree with bootstrap
values (ASSRT, 2007; Figure 18). This
dataset included additional samples
from the Santee-Cooper, Waccamaw,
and Edisto populations in the Southeast.
Atlantic sturgeon river populations also
grouped into five population segments
along the U.S. East Coast in this
analysis. In the Southeast, Atlantic
sturgeon from the Santee-Cooper system
grouped with the Albemarle Sound
population, while the other two river
populations grouped with the
Savannah/Ogeechee/Altamaha River
population segment. With the exception
of the Waccamaw River population, all
river populations sampled within each
population segment along the entire
East Coast were geographically adjacent.
The Waccamaw River population
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grouped with the Edisto/Savannah/
Ogeechee/Altamaha River population
segment, even though it is
geographically located between
Albemarle Sound and the Santee and
Cooper Rivers. However, we attributed
this to the small sample size (21 fish)
from the Waccamaw River and the fact
that all samples came from juveniles,
which may be migrants from other
systems. From the seven Southeast river
populations included in the analysis,
we determined based on genetic
information that river populations from
the ACE Basin southward are a
genetically distinct, interbreeding
population segment and that river
populations between the Santee-Cooper
system and Albemarle Sound (Roanoke
River) are a genetically distinct,
interbreeding population segment.
The higher accuracy in identifying
Atlantic sturgeon to one of two
population groupings (Albemarle
Sound/Santee-Cooper Rivers and
Ogeechee/Savannah/Altamaha/Edisto
Rivers) compared to their natal rivers
supports the fact that these multipleriver population segments are discrete
from each other.
We have considered the information
on Atlantic sturgeon population
structuring provided in the status
review report and Grunwald et al.
(2008). The nDNA analyses described in
the status review report provide
additional genetic information and
include chord distances and bootstrap
values to support the findings for
population structuring of Atlantic
sturgeon within the United States.
Therefore, based on genetic differences
observed among certain river
populations and the assumption that
adjacent river populations are more
likely to breed with one another than
river populations from rivers that are
not adjacent to each other, five discrete
Atlantic sturgeon population segments
in the United States meet the DPS
Policy’s discreteness criterion, with two
located in the Southeast: (1) The
‘‘Carolina’’ population segment, which
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includes Atlantic sturgeon originating
from the Roanoke, Tar/Pamlico, Cape
Fear, Waccamaw, Pee Dee, and SanteeCooper Rivers, and (2) the ‘‘South
Atlantic’’ population segment, which
includes Atlantic sturgeon originating
from the ACE Basin (Ashepoo,
Combahee, and Edisto rivers),
Savannah, Ogeechee, Altamaha, and
Satilla Rivers.
Significance
When the discreteness criterion is met
for a potential DPS, as it is for the
Carolina and South Atlantic population
segments in the Southeast identified
above, the second element that must be
considered under the DPS policy is
significance of each DPS to the taxon as
a whole. The DPS policy cites examples
of potential considerations indicating
significance, including: (1) Persistence
of the discrete population segment in an
ecological setting unusual or unique for
the taxon; (2) evidence that loss of the
discrete population segment would
result in a significant gap in the range
of the taxon; (3) evidence that the DPS
represents the only surviving natural
occurrence of a taxon that may be more
abundant elsewhere as an introduced
population outside its historical range;
or, (4) evidence that the discrete
population segment differs markedly
from other populations of the species in
its genetic characteristics.
We believe that the Carolina and
South Atlantic population segments
persist in ecological settings unique for
the taxon. This is evidenced by the fact
that spawning habitat of each
population grouping is found in
separate and distinct ecoregions that
were identified by TNC based on the
habitat, climate, geology, and
physiographic differences for both
terrestrial and marine ecosystems
throughout the range of the Atlantic
sturgeon along the Atlantic coast (Figure
1). TNC descriptions do not include
detailed information on the chemical
properties of the rivers within each
ecoregion, but include an analysis of
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bedrock and surficial geology type
because it relates to water chemistry,
hydrologic regime, and substrate. It is
well established that waters have
different chemical properties (i.e.,
identities) depending on the geology of
where the waters originate.
Riverine spawning habitat of the
Carolina population segment occurs
within the Mid-Atlantic Coastal Plain
ecoregion, which is described as
consisting of bottomland hardwood
forests, swamps, and some of the
world’s most active coastal dunes,
sounds, and estuaries. Natural fires,
floods, and storms are so dominant in
this region that the landscape changes
very quickly. Rivers routinely change
their courses and emerge from their
banks. The TNC lists the most
significant threats (sources of biological
and ecological stress) in the region as:
Global climate change and rising sealevel; altered surface hydrology and
landform alteration (e.g., flood-control
and hydroelectric dams, inter-basin
transfers of water, drainage ditches,
breached levees, artificial levees,
dredged inlets and river channels, beach
renourishment, and spoil deposition
banks and piles); a regionally receding
water table, probably resulting from
both over-use and inadequate recharge;
fire suppression; land fragmentation,
mainly by highway development; landuse conversion (e.g., from forests to
timber plantations, farms, golf courses,
housing developments, and resorts); the
invasion of exotic plants and animals;
air and water pollution, mainly from
agricultural activities including
concentrated animal feed operations;
and over-harvesting and poaching of
species. Many of the Carolina
population segment’s spawning rivers,
located in the Mid-Coastal Plain,
originate in areas of marl. Waters
draining calcareous, impervious surface
materials such as marl are likely to be
alkaline, dominated by surface run-off,
have little groundwater connection, and
be seasonally ephemeral.
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The riverine spawning habitat of the
South Atlantic population segment
occurs within the South Atlantic Coastal
Plain ecoregion. TNC describes the
South Atlantic Coastal Plain ecoregion
as fall-line sandhills to rolling longleaf
pine uplands to wet pine flatwoods;
from small streams to large river
systems to rich estuaries; from isolated
depression wetlands to Carolina bays to
the Okefenokee Swamp. Other
ecological systems in the ecoregion
include maritime forests on barrier
islands, pitcher plant seepage bogs and
Altamaha grit (sandstone) outcrops. The
primary threats to biological diversity in
the South Atlantic Coastal Plain listed
by TNC are intensive silvicultural
practices, including conversion of
natural forests to highly managed pine
monocultures and the clear-cutting of
bottomland hardwood forests. Changes
in water quality and quantity, caused by
hydrologic alterations (impoundments,
groundwater withdrawal, and ditching),
and point and nonpoint pollution, are
threatening the aquatic systems.
Development is a growing threat,
especially in coastal areas. Agricultural
conversion, fire regime alteration, and
the introduction of nonnative species
are additional threats to the ecoregion’s
diversity. The South Atlantic DPS’s
spawning rivers, located in the South
Atlantic Coastal Plain, are primarily of
two types: brown-water (with
headwaters north of the Fall Line, siltladen) and black-water (with
headwaters in the coastal plain, stained
by tannic acids).
Therefore, the ecoregion delineations
support that the physical and chemical
properties of the Atlantic sturgeon
spawning rivers utilized by the Carolina
and South Atlantic DPSs are unique to
each population segment. Since
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reproductive isolation accounts for the
discreteness of each population
segment, the Carolina and South
Atlantic population segments of
Atlantic sturgeon are ‘‘significant’’ as
defined in the DPS policy given that the
spawning rivers for each population
segment occur in a unique ecological
setting.
The loss of either the Carolina or the
South Atlantic population segments of
Atlantic sturgeon would create a
significant gap in the range of the taxon.
The loss of the Carolina population
segment would result in a 475-mile
(764-kilometer (km)) gap between the
northern population segments and the
South Atlantic population segment. The
loss of the South Atlantic population
segment would truncate the southern
range of Atlantic sturgeon by greater
than 150 miles (241 km). Though
Atlantic sturgeon travel great distances
in the marine environment and may use
multiple river systems for foraging and
nursery habitat, the range occupied by
the Carolina and South Atlantic
population segments would likely not
be recolonized by a new, viable
spawning population if either
population segment was lost, except
over a long time frame. Genetic analyses
show that fewer than two individuals
per generation spawn outside their natal
rivers (Wirgin et al., 2000; King et al.,
2001; Waldman et al., 2002). However,
a caveat to this information is that a
natal river population, well-established
over a long span of geological time and
highly adapted to its respective natal
river, may not realize success in
colonizing another river already
populated by a second population better
adapted to its respective natal river than
a potential colonist. The low rate of
genetic exchange displayed among
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adjacent sturgeon populations may not
reflect the incapacity of the species to
colonize, but the competitive advantage
held by a pre-established natal river
population facing migrant individuals.
However, we do not expect Atlantic
sturgeon that originate from other
population segments to re-colonize
extirpated systems and establish new
spawning populations, except perhaps
over a long time frame (i.e., many
Atlantic sturgeon generations).
Therefore, the loss of either the Carolina
or South Atlantic population segments
would result in a significant gap in the
range of Atlantic sturgeon over a long
time frame, and negatively impact the
species as a whole because the loss of
either population segment would
constitute an important loss of genetic
diversity for the Atlantic sturgeon.
The information presented above
describes: (1) Persistence of the Carolina
and South Atlantic population segments
in ecological settings that are unique for
the Atlantic sturgeon as a whole; and (2)
evidence that loss of either population
segment would result in a significant
gap in the range of the taxon. Based on
this information, we conclude that the
Carolina and South Atlantic population
segments meet the discreteness and
significance criteria outlined in the DPS
policy. We hereafter refer to these DPSs
as the Carolina and South Atlantic
DPSs. Figures 2 and 3 show the Carolina
and South Atlantic DPSs. While the
entire marine range of both the Carolina
and South Atlantic DPSs extends from
the Hamilton Inlet, Labrador, Canada, to
Cape Canaveral, Florida, figures 2 and 3
only depict the portion of the marine
range directly adjacent to the riverine
portions of each DPS.
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Conservation Status
To determine the conservation status
of the two DPSs in the Southeast
Region’s jurisdiction, the Carolina and
South Atlantic DPSs, in relation to the
ESA’s standards for listing, we
evaluated whether each DPS meets the
definition of ‘‘endangered’’ or
‘‘threatened’’ as defined in section 3 of
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the ESA, and whether that status is a
result of one or a combination of the
factors listed under section 4(a)(1) of the
ESA. An endangered species is ‘‘any
species which is in danger of extinction
throughout all or a significant portion of
its range’’ and a threatened species is
one ‘‘which is likely to become an
endangered species within the
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foreseeable future throughout all or a
significant portion of its range.’’
The abundance of Atlantic sturgeon
has decreased dramatically within the
last 150 years. A major fishery for
Atlantic sturgeon developed in 1870
when a caviar market was established
(Smith and Clugston, 1997). Record
landings in the United States were
reported in 1890, with over 7,385,000
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lbs (3,350,000 kg) of Atlantic sturgeon
landed from coastal rivers along the
entire Atlantic Coast (Smith and
Clugston, 1997; Secor and Waldman,
1999). Ten years after peak landings, the
fishery collapsed in 1901, when less
than 10 percent (650,365 lbs, 295,000
kg) of the U.S. 1890 peak landings were
reported. The landings continued to
decline coastwide, reaching about 5
percent of the peak in 1920. During the
1950s, the remaining U.S. fishery
switched to targeting sturgeon for flesh,
rather than caviar, and coastwide
landings remained between 1 and 5
percent of the 1890 peak levels until the
Atlantic sturgeon fishery was closed by
ASMFC in 1998. None of the riverine
spawning populations in either DPS
have rebounded from the population
crashes to be large or stable enough to
provide with any level of certainty for
continued existence of Atlantic sturgeon
in this part of its range. All of the
spawning populations in each DPS are
subjected to threats and impacts that
have and will continue to prevent
population increases and recovery. We
must look at the status of river
populations across the whole of the
DPSs in making our listing
determinations.
The importance of having multiple
self-sustaining riverine spawning
populations within each DPS and the
need to maintain suitable habitat to
support the various life functions
(spawning, feeding, growth) of Atlantic
sturgeon is further highlighted by
looking at the concept of
metapopulations. Each DPS, made up of
multiple river populations, is analogous
to a metapopulation, which is a
‘‘population of populations’’ (Levins,
1969), a group of spatially separated
populations of the same species that
interact at some level. The
metapopulation concept is closely
linked with the processes of population
turnover, extinction, and establishment
of new populations, and the study of
metapopulation dynamics is essentially
the study of the conditions under which
these two processes are in balance and
the consequences of that balance to
associated processes (Hanski and
Gilpin, 1991). Separation into
metapopulations is expected by
sturgeon and other anadromous fishes,
given their likely stepping-stone
sequential model of recolonization of
northern rivers following postPleistocene deglaciation (Waldman et
al. 2002).
Metapopulation persistence depends
on the balance of extinction and
colonization in a static environment
(Hanski 1996). Models and empirical
observations suggest that very small
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populations are relatively likely to
´
become extinct (Soule, 1986; Lande,
1988; Simberloff, 1988; Thomas, 1990;
Kindvall and Alhlen, 1992), and many
local populations in remnant habitat
fragments will remain small. Under the
assumption that the environment does
not change greatly, many empirical
studies have shown that the expected
lifetime of a population increases with
its current size (Williamson 1981,
Diamond 1984, Schoener and Spiller
1987). However, for rare and declining
species, Thomas (1994) argues that: (1)
Extinction is usually the deterministic
consequence of the local environment
becoming unsuitable (through habitat
loss or modification, introduction of a
predator, etc.); (2) that the local
environment usually remains unsuitable
following local extinction, so
extinctions only rarely generate empty
patches of suitable habitat; and 3) that
colonization usually follows
improvement of the local environment
for a particular species. Therefore, if
habitat remains suitable following local
extirpation, recolonization via
immigrants into now-empty habitat may
replace at least some of those losses
(Thomas 1994). However, if the cause of
extinction is a deterministic population
response to unsuitable conditions (e.g.,
lack of suitable spawning habitat, poor
water quality, or disturbance of
substrates through repeated dredging),
the local habitat is likely to remain
unsuitable after extinction and be
unavailable for recolonization (Thomas
1994). Therefore, recolonization is
dependent upon both immigration from
adjacent, healthy populations and
habitat suitability. Because a DPS is a
group of populations, the stability,
viability, and persistence of individual
populations affects the persistence and
viability of the larger DPS. The loss of
any population within a DPS will result
in: (1) A long-term gap in the range of
the DPS that is unlikely to be
recolonized, or recolonized only very
slowly; (2) loss of reproducing
individuals; (3) loss of genetic
biodiversity; (4) potential loss of unique
haplotypes; (5) potential loss of adaptive
traits; and (6) reduction in total number.
The loss of a population will negatively
impact the persistence and viability of
the DPS as a whole as fewer than two
individuals per generation currently
spawn outside their natal rivers.
The persistence of individual
populations, and in turn the DPS,
depends on successful spawning and
rearing within the freshwater habitat,
the immigration into marine habitats to
grow, and then the return of adults to
natal rivers to spawn. Information on
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Atlantic sturgeon spawning within the
Carolina and South Carolina DPSs is
extremely limited. In the proposed
listing rule, we presumed spawning was
occurring if young-of-the-year (YOY)
were observed or mature adults were
present in freshwater portions of the
system. Within the Carolina DPS, we
concluded that spawning is occurring in
the following rivers based on these data:
1. Roanoke River—collection of 15
YOY (1997–1998); single YOY (2005).
2. Tar and Neuse Rivers—one YOY
(2005).
3. Cape Fear—upstream migration of
adults in the fall, carcass of ripe female
upstream in mid-September.
4. Winyah Bay—running ripe male in
Great Pee De River (2003).
Within the South Atlantic DPS, we
concluded that spawning is occurring in
the following rivers based on these data:
1. ACE (Ashepoo, Combahee, and
Edisto Rivers) Basin—1,331 YOY (1994–
2001); gravid female and running ripe
male in the Edisto (1997); 39 spawning
adults (1998).
2. Savannah River—22 YOY (1999–
2006); running ripe male (1997).
3. Ogeechee River—age-1 captures,
but high inter-annual variability (1991–
1998); 17 YOY (2003); 9 YOY (2004).
4. Altamaha River—74 captured/308
estimated spawning adults (2004); 139
captured/378 estimated spawning adults
(2005).
5. Satilla River—4 YOY and spawning
adults (1995–1996).
These data indicate that spawning
occurs within the Carolina and South
Atlantic DPSs; they do not indicate the
frequency of annual spawning events or
the degree to which spawning in these
systems leads to population growth,
persistence, or viability. The extent and
effectiveness of spawning events is
unknown and likely precarious in many
rivers given ongoing threats such as
water quality and restricted access to
upstream spawning areas (75 FR 61904).
In addition to spawning success, it is
difficult to quantify spawning potential
within the two DPSs given the lack of
population estimates. Currently, the
number of Atlantic sturgeon in the
Carolina DPS is estimated as at 3
percent of historical population size and
the South Atlantic DPS is estimated to
be at 1 percent of historical population
size, with the exception of the Altamaha
River population, estimated to be at 6
percent of historical population size.
Although the largest impact that caused
the precipitous decline of the species
has been curtailed (directed fishing), the
population size has remained relatively
constant at these greatly reduced levels
for approximately 100 years.
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The Carolina DPS includes all
Atlantic sturgeon that spawn or are
spawned in the watersheds (including
all rivers and tributaries) from
Albemarle Sound southward along the
southern Virginia, North Carolina, and
South Carolina coastal areas to
Charleston Harbor. The marine range of
Atlantic sturgeon from the Carolina DPS
extends from the Hamilton Inlet,
Labrador, Canada, to Cape Canaveral,
Florida. While Atlantic sturgeon exhibit
a high degree of spawning fidelity to
their natal rivers, multiple riverine,
estuarine, and marine habitats may
serve various life (e.g., nursery, foraging,
and migration) functions. Rivers known
to have current spawning populations
within the range of this DPS include the
Roanoke, Tar-Pamlico, Cape Fear,
Waccamaw, and Pee Dee Rivers.
However, in some rivers, spawning by
Atlantic sturgeon may not be
contributing to population growth
because of lack of suitable habitat and
other stressors on juvenile survival and
development. There may also be
spawning populations in the Neuse,
Santee and Cooper Rivers, though it is
uncertain. Historically, both the Sampit
and Ashley Rivers were documented to
have spawning populations at one time.
However, the spawning population in
the Sampit River is believed to be
extirpated and the current status of the
spawning population in the Ashley
River is unknown. Both rivers may be
used as nursery habitat by young
Atlantic sturgeon originating from other
spawning populations. This represents
our current knowledge of the river
systems utilized by the Carolina DPS for
specific life functions, such as
spawning, nursery habitat, and foraging.
However, fish from the Carolina DPS
likely use other river systems than those
listed here for their specific life
functions. The Carolina DPS also
includes Atlantic sturgeon held in
captivity (e.g., aquaria, hatcheries, and
scientific institutions) and which are
identified as fish belonging to the
Carolina DPS based on genetics
analyses, previously applied tags,
previously applied marks, or
documentation to verify that the fish
originated from (hatched in) a river
within the range of the Carolina DPS, or
is the progeny of any fish that originated
from a river within the range of the
Carolina DPS. NMFS has no records of
Atlantic sturgeon from the Carolina DPS
being held in captivity.
Historical landings data indicate that
between 7,000 and 10,500 adult female
Atlantic sturgeon were present in North
Carolina prior to 1890 (Armstrong and
Hightower, 2002; Secor, 2002). Secor
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(2002) estimates that 8,000 adult
females were present in South Carolina
during that same timeframe. Prior
reductions from the commercial fishery
and ongoing threats have drastically
reduced the numbers of Atlantic
sturgeon within the Carolina DPS.
Currently, the Atlantic sturgeon
spawning population in at least one
river system within the Carolina DPS
has been extirpated, with a potential
extirpation in an additional system. The
abundance of the remaining river
populations within the DPS, each
estimated to have fewer than 300
spawning adults, is estimated to be less
than 3 percent of what it was
historically (ASSRT, 2007). Though
directed fishing and possession of
Atlantic sturgeon is no longer legal, the
Carolina DPS continues to face threats
such as habitat alteration and bycatch.
The presence of dams has resulted in
the loss of access to over 60 percent of
the historical sturgeon habitat on the
Cape Fear River and in the SanteeCooper system. This has resulted in the
loss of important spawning and juvenile
developmental habitat and has reduced
the quality of the remaining habitat by
affecting water quality parameters (such
as depth, temperature, velocity, and DO)
that are important to sturgeon.
The South Atlantic DPS includes all
Atlantic sturgeon that spawn or are
spawned in the watersheds (including
all rivers and tributaries) of the ACE
Basin southward along the South
Carolina, Georgia, and Florida coastal
areas to the St. Johns River, Florida. The
marine range of Atlantic sturgeon from
the South Atlantic DPS extends from the
Hamilton Inlet, Labrador, Canada, to
Cape Canaveral, Florida. While Atlantic
sturgeon exhibit a high degree of
spawning fidelity to their natal rivers,
multiple riverine, estuarine, and marine
habitats may serve various life (e.g.,
nursery, foraging, and migration)
functions. Rivers known to have current
spawning populations within this DPS
include the Combahee, Edisto,
Savannah, Ogeechee, Altamaha, and
Satilla Rivers. However, in some rivers,
spawning by Atlantic sturgeon may not
be contributing to population growth
because of lack of suitable habitat and
other stressors on juvenile survival and
development. Historically, both the
Broad-Coosawatchie and St. Marys
Rivers were documented to have
spawning populations at one time; there
is also evidence that spawning may
have occurred in the St. Johns River or
one of its tributaries. However, the
spawning population in the St. Marys
River, as well as any historical
spawning population present in the St.
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Johns, is believed to be extirpated, and
the status of the spawning population in
the Broad-Coosawatchie is unknown.
Both the St. Marys and St. Johns Rivers
are used as nursery habitat by young
Atlantic sturgeon originating from other
spawning populations. The use of the
Broad-Coosawatchie by sturgeon from
other spawning populations is unknown
at this time. The presence of historical
and current spawning populations in
the Ashepoo River has not been
documented; however, this river may
currently be used for nursery habitat by
young Atlantic sturgeon originating
from other spawning populations. This
represents our current knowledge of the
river systems utilized by the South
Atlantic DPS for specific life functions,
such as spawning, nursery habitat, and
foraging. However, fish from the South
Atlantic DPS likely use other river
systems than those listed here for their
specific life functions. The South
Atlantic DPS also includes Atlantic
sturgeon held in captivity (e.g., aquaria,
hatcheries, and scientific institutions)
and which are identified as fish
belonging to the South Atlantic DPS
based on genetics analyses, previously
applied tags, previously applied marks,
or documentation to verify that the fish
originated from (hatched in) a river
within the range of the South Atlantic
DPS, or is the progeny of any fish that
originated from a river within the range
of the South Atlantic DPS. Ten Atlantic
sturgeon taken from the Altamaha River
are currently being held at the Bears
Bluff National Fish Hatchery on
Wadmalaw Island, South Carolina,
though it is not certain whether those
fish were spawned in the Altamaha or
were migrants from another river
system. NMFS has no other records of
Atlantic sturgeon from the South
Atlantic DPS being held in captivity.
Secor (2002) estimated that 8,000
spawning female Atlantic sturgeon were
present in South Carolina. Historically,
the population of spawning female
Atlantic sturgeon in Georgia was
estimated at 11,000 fish per year prior
to 1890 (Secor, 2002). Prior reductions
from the commercial fishery and
ongoing threats have drastically reduced
the numbers of Atlantic sturgeon within
the South Atlantic DPS. Currently, the
Atlantic sturgeon spawning population
in one (possibly two) river systems
within the South Atlantic DPS have
been extirpated. The Altamaha River,
with an estimated 343 spawning adults
per year, is suspected to be less than 6
percent of its historical abundance,
extrapolated from the 1890s commercial
landings; the abundance of the
remaining river populations within the
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DPS, each estimated to have fewer than
300 spawning adults, is estimated to be
less than 1 percent of what it was
historically (ASSRT, 2007). While the
directed fishery that originally
drastically reduced the numbers of
Atlantic sturgeon has been closed, other
impacts have contributed to their low
population numbers, may have
contributed to the extirpation of some
spawning populations, and are likely
inhibiting recovery of extant river
populations. Historically, Atlantic
sturgeon likely accessed all parts of the
St. Johns River, as American shad were
reported as far upstream as Lake
Poinsett (reviewed in McBride, 2000).
However, the construction of
Kirkpatrick Dam (originally Rodman
Dam) at river mile (rm) 95 (rkm 153)
restricted migration to potential
spawning and juvenile developmental
habitat upstream. Approximately 63
percent of historical sturgeon habitat is
believed to be blocked due to the dam
(ASSRT, 2007), and there is no longer a
spawning population in the St. Johns
River.
Small numbers of individuals
resulting from drastic reductions in
populations, such as occurred with
Atlantic sturgeon due to the commercial
fishery, can remove the buffer against
natural demographic and environmental
variability provided by large
populations (Berry, 1971; Shaffer, 1981;
´
Soule, 1980). Though the Carolina and
South Atlantic DPSs, made up of
multiple river populations of Atlantic
sturgeon, were determined to be
genetically discrete, interbreeding
population units, the vast majority of
Atlantic sturgeon return to their natal
rivers to spawn, with fewer than two
migrants per generation spawning
outside their natal system (Wirgin et al.,
2000; King et al., 2001; Waldman et al.,
2002). Therefore, it is important to look
at each riverine spawning population
within each DPS when considering the
effects of a small population size on the
extinction risk for the DPS. Though
there is no absolute population size
above which populations are ‘‘safe’’ and
below which they face an unacceptable
´
risk of extinction (Gilpin and Soule,
´
1986; Soule and Simberloff, 1986;
Ewens et al., 1987; Goodman, 1987;
Simberloff, 1988; Thomas, 1990), some
have argued that ‘‘rules of thumb’’ can
´
and should be applied (Soule, 1987;
Thompson, 1991). Salwasser et al.
(1984) prescribe a minimum viable
population size of at least 1,000
reproducing adults. Belovsky (1987)
indicates that a minimum viable
population in the range of 1,000 to
10,000 reproducing adults should be
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sufficient for a mid-sized vertebrate
´
species. Soule (1987) suggests that
minimum viable population sizes for
vertebrate species should be in the ‘‘low
thousands’’ or higher. Thomas (1990)
offers a population size of 5,500 as ‘‘a
useful goal,’’ but suggests that where
uncertainty regarding a species’
population dynamics, changing
environmental conditions, and the
species’ reaction to the changing
environmental conditions is extreme
‘‘we should usually aim for population
sizes from several thousand to ten
thousand.’’ In a NOAA Technical
Memorandum entitled ‘‘Determining
Minimum Viable Populations under the
ESA,’’ Thompson (1991) states the ‘‘50/
500’’ rule of thumb initially advanced
´
by Franklin (1980) and Soule (1980)
comes the closest of any to attaining
‘‘magic number’’ status. Franklin (1980)
has suggested that, simply to maintain
short-term fitness (i.e., prevent serious
inbreeding and its deleterious effects),
the minimum effective population size
should be around 50. He further
recommended that, to maintain
sufficient genetic variability for
adaptation to changing environmental
conditions, the minimum effective
population size should be around 500.
´
Soule (1980) has pointed out that, above
and beyond preserving short-term
fitness and genetic adaptability, longterm evolutionary potential (at the
species level) may well require a
number of substantially larger
populations. It is important to note that
the 50/500 rule is cast in terms of
effective population size, a concept
introduced by Wright (1931). The
effective population size refers to an
ideal population of breeding individuals
produced each generation by random
union of an equal number of male and
female gametes randomly drawn from
the previous generation. To the extent
that this ideal is violated in nature, the
effective population size is generally
smaller than the overall number of
mature individuals in the population.
Multiple studies have shown that
Atlantic sturgeon do not spawn every
year, with spawning intervals ranging
from 1 to 5 years for males (Smith, 1985;
Collins et al., 2000; Caron et al. 2002)
and 2 to 5 years for females (Vladykov
and Greeley, 1963; Van Eenennaam et
al., 1996; Stevenson and Secor, 1999).
Therefore, the effective population size
(the number of adults in a population
that contribute offspring to the next
generation) for Atlantic sturgeon is more
closely related to the number of
annually spawning adults, rather than
total number of reproductively mature
adults. In the Southeast, even the
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spawning population in the Altamaha
River, believed to be the largest
spawning population of either the
Carolina or South Atlantic DPS, is
estimated to be smaller than the 500
recommended by Thompson (1991) to
maintain sufficient genetic variability
for adaptation to changing
environmental conditions. Total adult
population sizes are not known for any
of the rivers in the Carolina or South
Atlantic DPS. However, using the upper
end of our estimated range of abundance
(i.e., no more than 300 spawning adults
per year per river) and the fact that
Atlantic sturgeon only spawn every 1 to
5 years (i.e., 20 to 100 percent of the
total adult population is spawning every
year), then a conservative estimate of
the total reproductively mature adult
population in Southeastern rivers is 300
to 1,500. The Altamaha River would be
slightly higher than this, and many
rivers may be much lower, since we
don’t know how many fewer annual
adult spawners than the estimated 300
are in each river. But these ranges are
either below or on the lower end of the
1,000 to 10,000 individuals
recommended by other authors. It is not
known if certain riverine populations
are at abundances smaller than the
minimum effective population size of 50
that would prevent serious inbreeding
(Thompson, 1991). Moreover, in some
rivers, spawning by Atlantic sturgeon
may not be contributing to population
growth because of lack of suitable
habitat and other stressors on juvenile
survival and development.
Another factor potentially affecting
the size of a viable population of
Atlantic sturgeon is that they are
polyploid. Polyploid is a term used to
describe cells and organisms containing
more than two paired (homologous) sets
of chromosomes. The polyploidy of
Atlantic sturgeon might explain the high
degree of plasticity displayed by
sturgeon populations and may provide
them with the ability to repopulate from
very few spawning adults without
apparent inbreeding depression.
However, we have no certainty at this
time that this genetic characteristic will
allow the Atlantic sturgeon to recover
from such low population numbers, as
other listed polyploid Acipenser
species, such as the Gulf and shortnose
sturgeon, have not recovered
sufficiently to be delisted even after
being protected for 20 to 45 years.
The concept of a viable population
able to adapt to changing environmental
conditions is critical to Atlantic
sturgeon, and the low population
numbers of every river population in the
Carolina and South Atlantic DPSs put
them in danger of extinction throughout
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their ranges; none of the populations are
large or stable enough to provide with
any level of certainty for continued
existence of Atlantic sturgeon in this
part of its range. While the directed
fishery that originally drastically
reduced the numbers of Atlantic
sturgeon has been closed, recovery of
depleted populations is an inherently
slow process for a late-maturing species
such as Atlantic sturgeon, and they
continue to face a variety of other
threats that contribute to their risk of
extinction. Their late age at maturity
provides more opportunities for
individual Atlantic sturgeon to be
removed from the population before
reproducing. While a long life-span also
allows multiple opportunities to
contribute to future generations, it also
results increases the timeframe over
which exposure to the multitude of
threats facing the Carolina and South
Atlantic DPS can occur. These threats
include the loss, reduction, and
degradation of habitat resulting from
dams, dredging, and changes in water
quality parameters (such as depth,
temperature, velocity, and DO). Even
with a moratorium on directed fisheries,
bycatch is a threat to both the Carolina
and South Atlantic DPSs. Fisheries
known to incidentally catch Atlantic
sturgeon occur throughout the marine
range of the species and in some
riverine waters as well. Because Atlantic
sturgeon mix extensively in marine
waters and may use multiple river
systems for spawning, foraging, and
other life functions, they are subject to
being caught in multiple fisheries
throughout their range. In addition to
direct mortality, stress or injury to
Atlantic sturgeon taken as bycatch but
released alive may result in increased
susceptibility to other threats, such as
poor water quality (e.g., exposure to
toxins). This may result in reduced
ability to perform major life functions,
such as foraging and spawning, or may
even result in post-capture mortality.
While some of the threats to the
Carolina and South Atlantic DPS have
been ameliorated or reduced due to the
existing regulatory mechanisms, such as
the moratorium on directed fisheries for
Atlantic sturgeon, bycatch is currently
not being addressed through existing
mechanisms. Further, water quality
continues to be a problem even with
existing controls on some pollution
sources and water withdrawal, and
dams continue to curtail and modify
habitat, even with the Federal Power
Act’s provisions regarding anadromous
fish passage.
We have reviewed the status review
report, as well as other available
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literature and information, and have
consulted with scientists and fishery
resource managers familiar with
Atlantic sturgeon in the Carolina and
South Atlantic DPSs. After reviewing
the best scientific and commercial
information available, we find that both
the Carolina and South Atlantic DPSs
are in danger of extinction throughout
their ranges and thus meet the ESA’s
definition of an endangered species.
Atlantic sturgeon populations declined
precipitously decades ago due to
directed commercial fishing. The failure
of Atlantic sturgeon numbers within the
Carolina and South Atlantic DPSs to
rebound even after the moratorium on
directed fishing was established in 1998
indicates that impacts and threats from
limits on habitat for spawning and
development, habitat alteration, and
bycatch are responsible for the risk of
extinction faced by both DPSs. In
addition, the persistence of these
impacts and threats points to the
inadequacy of existing regulatory
mechanisms to address and reduce
habitat alterations and bycatch. We will
address the threats of habitat alteration,
bycatch, and the inadequacy of
regulatory mechanisms and their
contributions to the endangered statuses
of the Carolina and South Atlantic DPSs
in detail in the following sections of this
final rule.
Analysis of Section 4(a)(1) Factors’
Effects on the Species
The ESA requires us to determine
whether any species is endangered or
threatened because of any of the
following factors: (A) Present or
threatened destruction, modification, or
curtailment of habitat or range; (B)
overutilization for commercial,
recreational, scientific, or educational
purposes; (C) disease or predation; (D)
inadequacy of existing regulatory
mechanisms; or (E) other natural or
manmade factors affecting its continued
existence. Listing determinations are
made solely on the best scientific and
commercial data available and after
taking into account any efforts being
made by any state or foreign nation to
protect the species. The ASSRT
examined each of the aforementioned
five factors for their impacts on the
Atlantic sturgeon: DPSs. The following
is a summary of its relevant findings,
any additional information that has
become available since the status review
report was published, and the
conclusions that we have made based
on the available information.
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A. Present or Threatened Destruction,
Modification, or Curtailment of the
Species’ Habitat or Range
Habitat alterations considered by the
ASSRT that affect the status of sturgeon
populations include: Dam and tidal
turbine construction and operation;
dredging, disposal, and blasting; and
water quality modifications, such as
changes in levels of DO, water
temperature, and contaminants. Atlantic
sturgeon, like all anadromous fish, are
vulnerable to a host of habitat impacts
because they use rivers, estuaries, bays,
and the ocean at various points of their
life. In addition to the habitat alterations
considered by the ASSRT, other
emerging threats to habitat considered
in this section are drought, intra- and
inter-state water allocation issues, and
climate change. These threats have the
potential to further exacerbate habitat
modifications evaluated by the ASSRT.
Because they were not evaluated in the
status review report, they are considered
in more detail in this section. In this
section, we summarize the threats for
each DPS that we believe represent a
present or threatened destruction,
modification or curtailment of the DPS’s
habitat or range and are contributing to
the endangered status of both DPSs.
Dams
Dams are a threat to the Carolina and
South Atlantic DPS that contributes to
their endangered status by the
curtailment of the extent of available
habitat, as well as modifying sturgeon
habitat downstream through a reduction
in water quality. As noted in the status
review report, dams for hydropower
generation, flood control, and
navigation adversely affect Atlantic
sturgeon habitat by impeding access to
spawning, developmental and foraging
habitat, modifying free-flowing rivers to
reservoirs, physically damaging fish on
upstream and downstream migrations,
and altering water quality in the
remaining downstream portions of
spawning and nursery habitat. Attempts
to minimize the impacts of dams using
measures such as fish passage have not
proven beneficial to Atlantic sturgeon,
as they do not regularly use existing fish
passage devices, which are generally
designed to pass pelagic fish. To date,
only four Atlantic sturgeon have been
documented to have passed via a fish
lift (three at the St. Stephens fish lift in
South Carolina and one at the Holyoke
Dam in Massachusetts), as these passage
facilities are not designed to
accommodate adult-sized sturgeon.
While there has not been a large loss of
Atlantic sturgeon habitat throughout the
entire species’ range due to the presence
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of dams, individual riverine systems
have been severely impacted by dams,
as access to large portions of historical
sturgeon spawning and juvenile
developmental habitat has been
eliminated or restricted. The ASSRT
used GIS tools and dam location data
collected by Oakley (2003) as reference
points for river kilometer measurements
to map historical rivers in which
Atlantic sturgeon spawned. This
information was then used to determine
the number of kilometers of available
habitat. Within the Carolina and South
Atlantic DPSs, the Cape Fear, SanteeCooper, and St. Johns River systems
have lost greater than 60 percent of the
habitat historically used for spawning
and juvenile development.
The Cape Fear River has three locks
and dams (constructed from 1915 to
1935) between Wilmington and
Fayetteville that are located below the
fall line; two additional dams, Buckhorn
and B. Everette Jordan, are located
above the fall line. Atlantic sturgeon
movement is blocked at the first lock
and dam located in Riegelwood, North
Carolina, which was constructed in
1915. Pelagic species can pass over the
three locks and dams during high water,
but the benthic Atlantic sturgeon is not
known to pass over these three locks/
dams. No Atlantic sturgeon have been
captured upstream of Lock and Dam #1
despite extensive sampling efforts
(Moser et al., 1998). Exact historical
spawning locations are unknown in the
Cape Fear River, but Atlantic sturgeon
spawning is generally believed to occur
in flowing water between the salt front
and fall line of large rivers (Borodin,
1925; Leland, 1968; Scott and Crossman,
1973; Crance, 1987; Bain et al., 2000).
Therefore, sturgeon researchers judge
the fall line to be the likely upper limit
of spawning habitat. Using the fall line
as a guide, only 36 percent of the
historical habitat is available to Atlantic
sturgeon. In some years, the salt water
interface reaches the first lock and dam;
therefore, spawning adults in the Cape
Fear River either do not spawn in such
years or spawn in the major tributaries
of the Cape Fear River (i.e., Black River
or Northeast Cape Fear Rivers) that are
not obstructed by dams.
The Santee-Cooper Hydroelectric
Project is located in the coastal plain of
the Santee Basin on the Santee and
Cooper Rivers, South Carolina. The
project was finished in 1942 and
includes Lake Marion, which is
impounded by the Santee Dam (Wilson
Dam) on the Santee River at rm 87 (rkm
140), and Lake Moultrie, which is
impounded by the Pinopolis Dam on the
Cooper River at rm 48 (rkm 77). Using
the fall line as the upper region of
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spawning habitat, it is estimated that
only 38 percent of the historical habitat
is available to Atlantic sturgeon today.
Although a lock and a fish lift operate
during the spring at the Pinopolis and
St. Stephen Dams, respectively,
observations of sturgeon in the lock and
lift are extremely rare (traditional fish
passage designs are not typically
successful for sturgeon). There is no
record of an adult Atlantic sturgeon
being lifted, although three dead
Atlantic sturgeon were observed in Lake
Marion between 1995 and 1997, and in
2007, an Atlantic sturgeon entered the
St. Stephen fish lift and was physically
removed and translocated downstream
into the Santee River (A. Crosby,
SCDNR, pers. comm.)
In addition to blocking access to
habitat, dams can degrade spawning,
nursery, and foraging habitat
downstream by reducing water quality.
Flow, water temperature, and oxygen
levels in the Roanoke River are affected
by the Kerr Dam and the Gaston Dam/
Roanoke Rapids facilities, which engage
in peaking operations. Riverine water
flow has already been modified by the
dam operators during the striped bass
spawning season to simulate natural
flow patterns; these modifications
undoubtedly benefit Atlantic sturgeon.
Regardless of the temporary
modifications, lower water temperatures
resulting from the hypolimnetic
discharge from Kerr Dam have caused
temporal shifts in the spawning peaks
for both American shad and striped bass
and likely have had the same impact for
other diadromous species, including
Atlantic sturgeon (ASSRT, 2007). High
flows from Kerr Dam during the summer
are coupled with high ambient
temperatures and an influx of swamp
water with low DO, creating a large,
hypoxic plume within the river. Fish
kills have been documented to occur
during this time (ASSRT, 2007), and
sturgeon are more highly sensitive to
low DO (less than 5 milligrams per liter
(mg/L)) than other fish species
(Niklitschek and Secor, 2009a, 2009b).
Low DO in combination with high
temperature is particularly problematic
for Atlantic sturgeon, and studies have
shown that juvenile Atlantic sturgeon
experience lethal and sublethal
(metabolic, growth, feeding) effects as
DO drops and temperatures rise
(Niklitschek and Secor, 2009a, 2009b;
Niklitschek and Secor, 2005; Secor and
Gunderson, 1998). Therefore, it is likely
that dam operations are negatively
affecting Atlantic sturgeon nursery
habitat in the lower Roanoke River.
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Dredging
Dredging is a present threat to both
the Carolina and South Atlantic DPSs
and is contributing to their endangered
status by modifying the quality and
availability of Atlantic sturgeon habitat.
Riverine, nearshore, and offshore areas
are often dredged to support commercial
shipping and recreational boating,
construction of infrastructure, and
marine mining. Environmental impacts
of dredging include the direct removal/
burial of organisms; turbidity/siltation
effects; contaminant resuspension;
noise/disturbance; alterations to
hydrodynamic regime and physical
habitat; and actual loss of riparian
habitat (Chytalo, 1996; Winger et al.,
2000). According to Smith and Clugston
(1997), dredging and filling impact
important habitat features of Atlantic
sturgeon as they disturb benthic fauna,
eliminate deep holes, and alter rock
substrates. To reduce the impacts of
dredging on anadromous fish species,
most of the Atlantic states impose work
restrictions during sensitive time
periods (spawning, migration, feeding)
when anadromous fish are present.
NMFS also imposes seasonal
restrictions to protect shortnose
sturgeon populations (where present)
through Section 7 consultations that
may have the added benefit of
protecting Atlantic sturgeon where the
two species co-occur. Within the
Carolina DPS, dredging operations
(including the blasting of rock) on the
lower Cape Fear River, Brunswick River,
and port facilities at the U.S. Army’s
Sunny Point Military Ocean Terminal
and Port of Wilmington are extensive.
To protect diadromous fish, restrictions
are placed on dredging to avoid
sensitive seasons and locations, such as
potential spawning habitat (February 1
through June 30) and suspected nursery
grounds (April 1 through September 30).
However, while the restrictions prevent
dredging from occurring when Atlantic
sturgeon are expected to be present, the
effects of dredging on Atlantic sturgeon
habitat remain long after the dredging
has been completed. Moser and Ross
(1995) found that some of the winter
holding sites favored by sturgeon in the
lower Cape Fear River estuary also
support very high levels of benthic
infauna and may be important feeding
stations. Repeated dredging in the Cape
Fear River can modify sturgeon habitat
through the removal or burial of benthic
infauna in feeding grounds and creation
of unsuitable substrate in spawning
grounds (ASSRT, 2007). Similar habitat
modifications are occurring in the
Cooper River, which flows into
Charleston Harbor, one of the busiest
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ports on the Atlantic Coast, and is
dredged regularly. The river channel is
maintained by dredging all the way to
the Pinopolis Dam. No seasonal
restrictions are placed on dredging in
the Cooper River, potentially
interrupting spawning activities
(ASSRT, 2007). In August 2011, the
USACE published a notice of intent to
prepare an EIS to study the impacts of
potential deepening of Charleston
Harbor to accommodate much larger
container vessels; the project would
entail extensive dredging (76 FR 50187).
In the South Atlantic DPS,
maintenance dredging in Atlantic
sturgeon nursery habitat in the
Savannah River is frequent, and
substantial channel deepening took
place in 1994. The Georgia Ports
Authority is seeking to expand its port
facility on the Savannah River. Within
the 1999 Water Resources Development
Act, Congress authorized the deepening
of the Savannah Navigation Channel
from the current depth of ¥42 to ¥48
ft (¥12.8 to ¥14.6 m) mean low water.
Hydrodynamic and water quality
models have been developed to predict
changes in water quality across depth
and throughout the channel. The
channel deepening is predicted to alter
overall water quality (e.g., salinity and
DO), creating inhospitable foraging/
resting habitat in the lower Savannah
River for sturgeon. The lower Savannah
River is heavily industrialized and
serves as a major shipping port. Nursery
habitat in the lower river has been
heavily impacted by diminished water
quality and channelization. Reduced DO
levels and upriver movement of the salt
wedge are predicted to result from
channel deepening. Currently, USACE
has entered into formal consultation
with NMFS regarding the Savannah
Harbor Expansion Project, which
includes a conference consultation on
Atlantic sturgeon. Though not yet
finalized, the conference consultation
on Atlantic sturgeon will evaluate
whether the adverse effects on sturgeon
from the expansion will result in
jeopardy, and consider potential
benefits to Atlantic sturgeon from the
proposed fish passage at NSBL&D that
could provide access to 20 miles of
potential spawning habitat. Sturgeon are
highly sensitive to low DO, more so
than other fish species (Niklitschek and
Secor, 2009a, 2009b). Because Atlantic
sturgeon spawn above the interface
between fresh water and salt water, the
upriver movement of the salt wedge will
curtail the extent of Atlantic sturgeon
habitat in the Savannah River. Dredging
also commonly occurs within the St.
Johns River and has been linked to the
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reduction in submerged aquatic
vegetation where Atlantic sturgeon
likely forage (Jordan, 2002). Though
there is currently no resident spawning
population in the St. Johns, it still
provides nursery habitat for juvenile
Atlantic sturgeon in the South Atlantic
DPS (NMFS and USFWS, 1998). Access
to over 60 percent of the historical
sturgeon habitat in the St. Johns River
has already been curtailed by the
presence of a dam, and dredging
modifies the quality of the remaining
nursery habitat in the river.
Water Quality
Degraded water quality is a present
threat to the Carolina and South
Atlantic DPSs and is contributing to
their endangered status by modifying
and curtailing the extent of available
habitat for spawning and nursery areas.
Atlantic sturgeon rely on a variety of
water quality parameters to successfully
carry out their life functions. Low DO
and the presence of contaminants
modify the quality of Atlantic sturgeon
habitat and in some cases, curtail the
extent of suitable habitat for life
functions. Secor (1995) noted a
correlation between low abundances of
sturgeon during this century and
decreasing water quality caused by
increased nutrient loading and
increased spatial and temporal
frequency of hypoxic conditions. Of
particular concern is the high
occurrence of low DO coupled with
high temperatures in the river systems
throughout the range of the Carolina and
South Atlantic DPSs. Sturgeon are more
highly sensitive to low DO than other
fish species (Niklitschek and Secor,
2009a, 2009b) and low DO in
combination with high temperature is
particularly problematic for Atlantic
sturgeon. Studies have shown that
juvenile Atlantic sturgeon experience
lethal and sublethal (metabolic, growth,
feeding) effects as DO drops and
temperatures rise (Niklitschek and
Secor, 2009a, 2009b; Niklitschek and
Secor, 2005; Secor and Gunderson,
1998). Water quality within the river
systems in the range of the Carolina and
South Atlantic DPSs is also negatively
impacted by contaminants and large
water withdrawals.
For the Carolina DPS, water quality in
the Pamlico system, especially in the
lower Neuse River, is highly degraded
(Paerl et al., 1998; Qian et al., 2000;
Glasgow et al., 2001). The entire basin
has been designated as nutrientsensitive, and additional regulatory
controls are being implemented to
improve water quality. Both the Neuse
and Pamlico portions of the estuary
have been subject to seasonal episodes
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5969
of anoxia that significantly affect the
quality of Atlantic sturgeon nursery
habitat. CAFOs cause at least some
portion of the current water quality
problems in the Pamlico watershed
(Mallin and Cahoon, 2003). Farms that
produce hogs, turkeys, and chickens
have proliferated throughout the coastal
portion of the basin in the last decade,
with increases in both aquatic and
atmospheric deposition of nitrogenous
waste products. North Carolina passed a
moratorium in 1997 limiting additional
hog operations and is conducting a
study of measures to address the
problem; the moratorium was renewed
in 1999 and 2003. Water quality in the
Cape Fear River is poor for aquatic life,
due largely to industrial development
and use, including the Port of
Wilmington and numerous industrial
point-source discharges. Development
of CAFOs in the coastal portion of the
Cape Fear River basin has been
especially heavy (most concentrated
operations of CAFOs occur in the Cape
Fear River drainage within North
Carolina) and contributes to both
atmospheric and aquatic inputs of
nitrogenous contamination, possibly
causing DO levels to regularly fall below
the 5 mg/L state standard (Mallin and
Cahoon, 2003). In recent years, fish kills
have been observed, usually as a result
of blackwater swamps (with low DO)
being flushed after heavy rainfall.
Industrialization also threatens the
habitat of the Carolina DPS. Paper and
steel mills in the Winyah Bay system,
which includes the Waccamaw, Pee
Dee, and Sampit rivers, have impacted
water quality. Riverine sediment
samples contain high levels of various
toxins including dioxins (NMFS and
USFWS, 1998). Though the effects of
these contaminants on Atlantic sturgeon
are unknown, Atlantic sturgeon are
particularly susceptible to impacts from
contaminated sediments due to their
benthic foraging behavior and long-life
span, and effects from these compounds
on fish include production of acute
lesions, growth retardation, and
reproductive impairment (Cooper, 1989;
Sinderman, 1994). It should be noted
that the effect of multiple contaminants
or mixtures of compounds at sub-lethal
levels on fish has not been adequately
studied. Atlantic sturgeon use marine,
estuarine, and freshwater habitats and
are in direct contact through water, diet,
or dermal exposure with multiple
contaminants throughout their range.
Habitat used by the South Atlantic
DPS in the Savannah River has also
been modified by mercury
contamination (ASSRT, 2007). While
water quality in the Altamaha River is
good at this time, the drainage basin is
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dominated by silviculture and
agriculture, with two paper mills and
over two dozen other industries or
municipalities discharging effluent into
the river. Nitrogen and phosphorus
concentrations are increasing, and
eutrophication and loss of thermal
refugia are growing concerns for the
South Atlantic DPS. In the Ogeechee
River, the primary source of pollution
results from non-point sources, which
results in nutrient-loading and
decreases in DO. These problems result
from the cumulative effect of activities
of many individual landowners or
managers. The Ogeechee River Basin
Watershed Protection Plan developed by
the Georgia Environmental Protection
Division (GAEPD, 2001b) states that
because there are so many small sources
of non-point loading spread throughout
the watershed, non-point sources of
pollution cannot effectively be
controlled by state agency permitting
and enforcement, even where regulatory
authority exists. The increases in
nutrients and resulting decreases in DO
are coupled with increases in water
temperature resulting from clearing of
the riparian canopy and increased
paved surface areas. Downstream
sturgeon nursery habitat is
compromised during hot, dry summers
when water flow is minimal, and nonpoint sources of hypoxic waters have a
greater impact on the system as
potential thermal refugia are lost when
the aquifer is lowered. Since 1986,
average summer DO levels in the
Ogeechee have dropped to
approximately 4 mg/L (GAEPD, 2001b).
Low DO (less than 5 mg/L), most likely
due to non-point sources, was a
common occurrence observed during
1998 and 1999 water quality surveys
(GAEPD, 2002) in the Satilla River,
which serves as both spawning and
nursery habitat for sturgeon in the South
Atlantic DPS. The extirpation of the
Atlantic sturgeon spawning population
in the St. Marys River is believed to
have been caused by reduced DO levels
during the summer in the nursery
habitat, probably due to eutrophication
from non-point source pollution
(ASSRT, 2007). Both the St. Marys and
St. Johns Rivers continue to be used as
nursery habitat by Atlantic sturgeon in
the South Atlantic DPS; however, low
DO is a common occurrence during the
summer months when water
temperatures rise. At times, it is so
severe in the St. Marys that it
completely eliminates juvenile nursery
habitat during the summer (D. Peterson,
UGA, pers. comm.)
Water allocation issues are a growing
threat in the Southeast and exacerbate
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existing water quality problems. Taking
water from one basin and transferring it
to another fundamentally and
irreversibly alters natural water flows in
both the originating and receiving
basins, which can affect DO levels,
temperature, and the ability of the basin
of origin to assimilate pollutants
(Georgia Water Coalition, 2006). Water
allocation issues increasingly threaten
to exacerbate the present threat of
degraded water quality on the
endangered status of the Carolina DPS.
North Carolina is experiencing problems
where somewhat limited natural
availability of water is coupled with
high demand or competition among
water users. Some of the areas in North
Carolina where this is an emerging issue
are the Central Coastal Plain, where the
Cretaceous aquifers have a relatively
slow recharge rate; the headwater areas
of the Piedmont river basins, where
streamflows are greatly reduced during
dry weather; and some areas near the
coast and on the Outer Banks, where the
natural availability of fresh water is
limited (NCDENR, 2001a). Interbasin
water transfers are increasingly being
looked at to deal with the inadequate
water availability. In 1993, the North
Carolina Legislature adopted the
Regulation of Surface Water Transfers
Act (G.S. § 143–215.22I). This law
regulates large surface water transfers
between river basins by requiring a
certificate from the North Carolina
Environmental Management
Commission. The act has been modified
several times since it was first adopted,
most recently in 2007 when G.S. § 143–
215.22I was repealed and replaced with
G.S. § 143–215.22L. A transfer
certificate is required for a new transfer
of 2 mgd (7,600 m3pd) or more and for
an increase in an existing transfer by 25
percent or more (if the total including
the increase is more than 2 mgd).
Certificates are not required for facilities
that existed or were under construction
prior to July 1, 1993, up to the full
capacity of that facility to transfer water,
regardless of the transfer amount.
The North Carolina Department of
Environment and Natural Resources
reports that 20 facilities, with a
combined average (not maximum) daily
transfer of 66.5 mgd (252,000 m3pd),
were grandfathered in when G.S. § 143–
215.22I was enacted (NCDENR, 2009).
Since then, five additional facilities
have received certificates to withdraw
up to a combined maximum total of
167.5 mgd (634,000 m3pd). The most
significant certified interbasin transfer
in this group is the withdrawal of 60
mgd (227,000 m3pd) of water from Lake
Gaston (part of the Roanoke River Basin)
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by Virginia Beach, Virginia. Virginia
Beach began pumping in 1998 following
a very lengthy and contested FERC
approval process, during which North
Carolina opposed the withdrawals
(NCDENR, 2001b). Certificates are
pending for three facilities, totaling
almost 60 mgd (227,000 m3pd). This
includes the Kerr Lake Regional Water
System (KLRWS), a regional provider of
drinking water. The KLRWS has an
existing, grandfathered, surface water
transfer capacity of 10 mgd (38,000
m3pd). The grandfathered capacity
allows the system to move water from
the Roanoke River Basin (Kerr Lake) to
sub-basins of the Tar-Pamlico River
Basin. On February 18, 2009, KLRWS
submitted a Notice of Intent to Request
an Interbasin Transfer Certificate to the
Environmental Management
Commission. In that notice, KLRWS
requested to increase the authorized
transfer from 10 mgd to 24 mgd (38,000
m3pd to 91,000 m3pd), and to transfer
2.4 mgd (9,100 m3pd) from the Roanoke
River Basin to the Neuse River Basin.
These transfer amounts are based on
water use projections to the year 2040.
Water allocation issues also
increasingly threaten to exacerbate the
present threat of degraded water quality
on the endangered status of the South
Atlantic DPS. Water allocation issues
are occurring on the Atlantic Coast of
South Carolina and Georgia (Ruhl,
2003). This area is served by five major
rivers—the Savannah, Altamaha
(including its two major tributaries, the
Oconee and Ocmulgee rivers),
Ogeechee, Satilla, and St. Marys Rivers.
A 2006 study by the Congressional
Budget Office (CBO) reported that
Georgia had the sixth highest
population growth (26.4 percent) in the
nation, followed by Florida (23.5
percent) (CBO, 2006). A report from
UGA states that the per capita water use
in Georgia has been estimated to be 8 to
10 percent greater than the national
average, and 17 percent higher than per
capita use in neighboring states (UGA,
2002). Water shortages have already
occurred and are expected to continue
due to increasing periods of drought
coupled with the rapid population
growth expected in the region over the
next 50 years (Cummings et al., 2003).
Two of the largest and most rapidly
expanding urban areas in the Savannah
River basin, Augusta-Richmond County
and Savannah, currently utilize both
ground water and surface water for
drinking water uses (GAEPD, 2001a).
Surface water use in the Savannah River
basin is expected to increase in the near
future, due to a population increase in
the basin. Predictions for 2050 estimate
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the population will increase to nearly
900,000 (GAEPD, 2001a). It is important
to note that the two water supply
sources are not independent, because
ground water discharge to streams is
important in maintaining dry-weather
flow. Thus, withdrawal of ground water
also results in reduction in surface
water flow.
The Vogtle Electric Generating Plant
consists of two nuclear reactors and
currently uses up to 64 mgd of water
from the Savannah River to generate
power. In March 2008, the Southern
Nuclear Operating Company applied to
the Nuclear Regulatory Commission for
a license to build two additional nuclear
reactors at the plant, increasing the
potential water usage to 80 mgd. Up to
100 mgd (379,000 m3pd) of Savannah
River water may be withdrawn to
support the growth of South Carolina
communities located outside of the
Savannah River basin, such as
Greenville and Beaufort County
(Spencer and Muzekari, 2002). While
Georgia has laws restricting interbasin
transfers of water, South Carolina has
yet to adopt stream flow protections and
does not regulate surface water
withdrawals (Rusert and Cummings,
2004). Savannah has been withdrawing
water from its coastal aquifer since the
city became established. However,
Savannah has grown to the point that
the aquifer has been depleted over 100
ft (31 m) beneath the city due to growth
and increased water usage. This
decrease in aquifer storage water has
resulted in salt water intrusion into the
water wells used by Hilton Head, just
north of Savannah. Currently, five of
Hilton Head’s 12 wells are unusable and
the problem is expected to escalate if no
action is taken to prevent further salt
water intrusion. The South Carolina
team on the Savannah River Basin
Advisory Group has begun looking at
withdrawing surface water from the
Savannah River to ease the aquifer
problem (Massey, 2007; Spencer and
Muzekari, 2002).
New surface water withdrawal
permits in the Savannah, Ogeechee, and
Altamaha Rivers pose potential threats
to water quality in those rivers (Alber
and Smith, 2001). Approximately
126,500 people depend on the Altamaha
basin for water. The Ocmulgee River, a
tributary of the Altamaha, is located in
North Georgia and passes through
Atlanta and Macon before joining the
Altamaha River. Of the seven river
basins in Georgia, the Ocmulgee River
Basin has the highest population of
1,714,722 people. The Ocmulgee River
Basin is home to a diverse industrial
and attraction base, from agriculture to
defense. It has the highest agriculture
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production and the most agricultural
water withdrawal permits in Georgia
(Fisher et al., 2003).
It is not known how much water is
already being removed from rivers
utilized by the South Atlantic DPS for
spawning and nursery habitat because
there is little information concerning
actual withdrawals and virtually no
information concerning water
discharges. This is particularly the case
for municipal and industrial uses
because water use permits are not
required for withdrawals less than
100,000 gpd (379 m3pd) (Cummings et
al., 2003) and discharge permits are not
required unless discharge contains
selected toxic materials. Agricultural
water use permits are not quantified in
any meaningful way, thus neither water
withdrawals nor return flows are
measured (Fisher et al., 2003). Large
withdrawals of water (such as those for
municipal use) result in reduced water
quality (altered flows, higher
temperatures, and lowered DO), and
reduced water quality is already
contributing to the endangered status of
the South Atlantic DPS. Therefore,
water withdrawals from the rivers in the
range of the South Atlantic DPS, which
are highly likely to occur based on
current water shortages and increasing
demand, threaten to exacerbate water
quality problems that are currently
modifying and curtailing Atlantic
sturgeon habitat in the South Atlantic
DPS.
Climate Change
Climate change threatens to
exacerbate the effects of modification
and curtailment of Atlantic sturgeon
habitat caused by dams, dredging, and
reduced water quality on the
endangered status of the Carolina and
South Atlantic DPSs. A major advance
in climate change projections is the
large number of simulations available
from a broader range of climate models,
run for various emissions scenarios. The
IPCC reports in its technical paper
‘‘Climate Change and Water’’ that bestestimate projections from models
indicate that decadal average warming
over each inhabited continent by 2030
(i.e., over the next 20-year period) is
insensitive to the choice of emissions
scenarios and is ‘‘very likely’’ to be at
least twice as large (around 0.36 degrees
Fahrenheit or 0.2 degrees Celsius per
decade) as the corresponding modelestimated natural variability during the
20th century (IPCC, 2008). Continued
greenhouse gas emissions at or above
current rates under non-mitigation
emissions scenarios would cause further
warming and induce many changes in
the global climate system during the
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21st century, with these changes ‘‘very
likely’’ to be larger than those observed
during the 20th century. In addition, the
IPCC expects the rate of warming to
accelerate in the coming decades.
Because 20 years is equal to at least one
generation of Atlantic sturgeon (ASSRT,
2007), and possibly multiple
generations in the Southeast where
Atlantic sturgeon may mature as early as
5 years (Smith et al., 1982), the
modifying effects of climate change over
the next 20 years on vital parameters of
the Carolina and South Atlantic DPS’s
habitat will occur on a scale relevant to
their endangered status. Researchers
anticipate that the frequency and
intensity of droughts and floods will
change across the nation (CBO, 2006).
The IPCC report states that the most
important societal and ecological
impacts of climate change in North
America stem from changes in surface
and groundwater hydrology (IPCC,
2008).
Both the Carolina and South Atlantic
DPSs are within a region the IPCC
predicts will experience overall climatic
drying. Since the status review report
was completed, the Southeast
experienced approximately 3 years of
drought. During this time, South
Carolina experienced drought
conditions that ranged from moderate to
extreme (South Carolina State
Climatology Office, 2008). From 2006
until mid-2009, Georgia experienced the
worst drought in its history. In
September 2007, many of Georgia’s
rivers and streams were at their lowest
levels ever recorded for the month, and
new record low daily streamflows were
recorded at 15 rivers with 20 or more
years of data in Georgia (USGS, 2007).
The drought worsened in September
2008. All streams in Georgia except
those originating in the extreme
southern counties were extremely low.
While Georgia has periodically
undergone periods of drought—there
have been 6 periods of drought lasting
from 2 to 7 years since 1903 (USGS,
2000)—drought frequency appears to be
increasing (Ruhl, 2003). Abnormally
low stream flows restrict access to
habitat areas, reduce thermal refugia,
and exacerbate water quality issues,
such as water temperature, reduced DO,
nutrient levels, and contaminants.
The Carolina and South Atlantic DPSs
are already threatened by reduced water
quality resulting from dams, inputs of
nutrients, contaminants from CAFOs,
industrial activities, and non-point
sources, and interbasin transfers of
water. The IPCC report projects with
high confidence that higher water
temperatures and changes in extremes
in this region, including floods and
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droughts, will affect water quality and
exacerbate many forms of water
pollution—from sediments, nutrients,
dissolved organic carbon, pathogens,
pesticides, and salt, as well as thermal
pollution, with possible negative
impacts on ecosystems. In addition, sealevel rise is projected to extend areas of
salinization of groundwater and
estuaries, resulting in a decrease of
freshwater availability for humans and
ecosystems in coastal areas. Some of the
most populated areas of this region are
low-lying, and the threat of salt water
entering into its aquifers with projected
sea-level rise is a concern (U.S. Global
Research Group, 2004). Existing water
allocation issues would be exacerbated,
leading to an increase in reliance on
interbasin water transfers to meet
municipal water needs, further stressing
water quality. Dams, dredging, and poor
water quality have already modified and
curtailed the extent of suitable habitat
for Atlantic sturgeon spawning and
nursery habitat. Changes in water
availability (depth and velocities) and
water quality (temperature, salinity, DO,
contaminants, etc.) in rivers and coastal
waters inhabited by Atlantic sturgeon
resulting from climate change will
further modify and curtail the extent of
suitable habitat for the Carolina DPS.
Effects could be especially harmful
since these populations have already
been reduced to low numbers. The
spawning populations within the
Carolina DPS are all estimated to
number fewer than the 500
recommended by Thompson (1991) to
maintain sufficient genetic variability
for adaptation to changing
environmental conditions, and certainly
smaller than the 1,000 to 10,000
recommended by other authors
(Salwasser et al., 1984; Belovsky, 1987;
´
Soule, 1987; Thomas, 1990).
The ASSRT concluded that habitat
modifications due to the placement of
dams, dredging, and degraded water
quality present a moderate to
moderately high threat to all river
populations within the Carolina DPS,
with the exception of the Roanoke
River. For the South Atlantic DPS, the
ASSRT concluded that dredging and
water quality issues are having a
moderately low to moderate impact on
the river populations. We believe that
the modification and curtailment of
Atlantic sturgeon habitat resulting from
dams, dredging, and degraded water
quality is contributing to the
endangered status of both the Carolina
and South Atlantic DPSs. Further,
additional threats arising from water
allocation and climate change threaten
to exacerbate water quality problems
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already present throughout the range of
both DPSs. Existing water allocation
issues will likely be compounded by
population growth and potentially
climate change. Climate change is also
predicted to elevate water temperatures
and exacerbate nutrient-loading,
pollution inputs, and lower DO, all of
which are current threats to the Carolina
and South Atlantic DPSs.
B. Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Overutilization for commercial
purposes is a factor that contributed to
the historical drastic decline in Atlantic
sturgeon populations throughout the
species’ range. Data on the total weight
of Atlantic and shortnose sturgeon
harvested were collected by each state
starting in 1880, and in the late 1800s
commercial fisheries were landing
upwards of 6,800,000 lbs (3,084 kg) of
sturgeon annually (Murawski and
Pacheco, 1977). By 1905, only 15 years
later, this number had dropped to
20,000 lbs (9,071 kg). The population
sizes were then further reduced by
overfishing in the 1900s, when the
landings drastically fell to a total of 215
lbs (98 kg) in 1990 (Stein et al., 2004b).
The total landings recorded include
shortnose sturgeon as well as Atlantic
sturgeon; however, the harvest is
thought to have been primarily Atlantic
sturgeon due to the large mesh-size nets
commonly used at that time. A complete
moratorium on possession of Atlantic
sturgeon has been implemented in both
state and Federal waters since 1998 to
eliminate the threat of directed catch
and incentives to retain Atlantic
sturgeon bycatch. However, Atlantic
sturgeon are taken as bycatch in various
commercial fisheries along the entire
U.S. Atlantic Coast within inland,
coastal, and Federal waters. While
Atlantic sturgeon caught incidentally
can no longer be legally landed, bycatch
may still be a threat if fish are injured
or killed in the act of being caught.
Based on their life history, Atlantic
sturgeon are more sensitive to fishing
mortality than other coastal fish species.
They are a long-lived species, have an
older age at full maturity, have lower
maximum fecundity values, with 50
percent of the lifetime egg production
for Atlantic sturgeon occurring later in
life (Boreman, 1997). Boreman (1997)
looked at the relationship between
fishing mortality (F) and the
corresponding percentage of the
maximum lifetime egg production of an
age 1 female. The F50 is the fishing rate
at which a cohort produces 50 percent
of the eggs that it would produce with
no fishing effort. Boreman calculated a
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sustainable fishing (bycatch) mortality
rate of 5 percent per year for adult
Atlantic sturgeon based on the F50.
While many fishery models use a less
conservative target fishing level of F30 or
F20, the more conservative choice of F50
for Atlantic sturgeon is justified by their
late age at maturity and because they are
periodic spawners (Boreman, 1997).
We currently do not have all the data
necessary to determine whether the
percentage of Atlantic sturgeon
populations lost annually due to
bycatch mortality exceeds a sustainable
rate of 5 percent per year suggested by
Boreman (1997), because we do not
have abundance estimates for the
Carolina and South Atlantic DPSs and
bycatch remains highly underreported.
However, bycatch is occurring
throughout the range of the Carolina and
South Atlantic DPSs of Atlantic
sturgeon, and the bycatch mortality
associated with the dominant fishing
gear in the Southeast is relatively high.
All the spawning populations in the
Southeast Region are quite small, which
means that the loss of a small number
of fish to bycatch mortality could
exceed the sustainable rate of 5 percent
per year. Bycatch of Atlantic sturgeon in
commercial fisheries is presently a
threat to the Carolina and South
Atlantic DPSs, and we believe it is
contributing to their endangered status.
Mortality rates of Atlantic sturgeon
taken as bycatch in various types of
fishing gear range between 0 and 51
percent, with the greatest mortality
occurring in sturgeon caught by sink
gillnets (Stein et al., 2004b; ASMFC,
2007). The ASMFC Sturgeon Technical
Committee (TC) determined that
bycatch losses principally occur in sink
gillnet fisheries, though there may be
losses in the trawl fisheries, as well.
Atlantic sturgeon are particularly
vulnerable to sink gillnets due to their
demersal nature (tendency to be at the
bottom of the water column). If the nets
are not tended often enough, it can be
detrimental to the sturgeon, resulting in
suffocation because their operculum or
gills can be held closed by the net.
Using the NMFS ocean observer dataset,
the NMFS Northeast Fisheries Science
Center (NEFSC) estimated that bycatch
mortality of sturgeon captured in sink
gillnets between 2001 and 2006 was
13.8 percent (ASMFC, 2007). The
ASMFC Sturgeon TC notes that any
estimate of bycatch from the NMFS
ocean observer dataset will be an
underestimate because bycatch is underreported in state waters and there is
limited observer coverage in fisheries
potentially affecting Atlantic sturgeon in
the South Atlantic (North Carolina to
Florida) Federal waters. In addition,
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bycatch mortality estimates do not
account for post-capture mortality. The
13.8 percent mortality rate for sink
gillnets estimated by the NEFSC may
further underestimate the mortality rate
in sink gillnets in the Carolina and
South Atlantic DPSs because bycatch
survival is greater in colder water
temperatures of the north compared to
warmer southern waters occupied by
these DPSs (ASSRT, 2007). Mortality of
Atlantic sturgeon captured by trawls
seems to be low, with most surveys
reporting 0 percent mortality. The State
of North Carolina has documented
bycatch in over 958 tows conducted by
commercial shrimp trawlers working in
North Carolina with no Atlantic
sturgeon reported; there have also been
no Atlantic sturgeon captured in the 528
blue crab trawl tows examined since
1990. However, these studies do not
include post-capture mortality, and
studies of mortality from trawl fisheries
conducted in the south, where tow
times are longer and water temperatures
are higher, are very limited.
Sink gillnets and trawls are used
throughout riverine, estuarine, and
marine waters in the range of the
Carolina DPS to target a wide array of
finfish and shellfish. Data on Atlantic
sturgeon bycatch in Albemarle and
Pamlico Sound commercial fisheries
come from three sources: (1) NCDMF
IGNS that were initially designed to
monitor striped bass; (2) NCDMF
Observer Program; and (3) NC Sea Grant
Fishery Resource Grant project that
examined sturgeon bycatch in the
flounder fishery (White and Armstrong,
2000). The Albemarle and Pamlico IGNS
used sink and drift gillnets, similar to
those used by the shad/herring and the
flounder fisheries. Overall bycatch
mortality in the Albemarle Sound IGNS
from 1990–2009 was 3 percent.
Mortality rates in Albemarle Sound
varied annually from 0–19 percent, and
also varied by month (0–7 percent) and
by mesh size (0–100 percent). Overall
bycatch mortality in the Pamlico Sound
IGNS from 2001–2009 was 10 percent,
and ranged from 0–100 percent
annually, 0–25 percent by month, and
0–25 percent by mesh size. In the
Pamlico, Pungo, and Neuse Rivers
IGNS, overall bycatch mortality between
2000 and 2009 was 12 percent, ranging
annually from 0–50 percent. Bycatch
mortality rate also varied by month (0–
67 percent) and by mesh size (0–33
percent). Since 2001, the NCDMF
Observer Program has observed
approximately 3,031,356 yards of large
and small mesh gill nets and collected
110 Atlantic sturgeon with an overall
bycatch mortality of 6 percent (7 fish).
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Mortalities ranged from 0 percent in
2008 to a high of 12 percent in 2004.
Overall bycatch mortality in large mesh
nets was 5 percent and ranged between
0 and 8 percent. Overall bycatch
mortality in small mesh nets was 17
percent, ranging from 0–100 percent.
Commercial fishermen in Albemarle
and Pamlico Sound and Cape Fear River
reported catches of zero to two sturgeon
per fishery per year. However, White
and Armstrong (2000) reported that
sturgeon bycatch in flounder gillnets
fished from 1998 to 2000 by a single
fishermen in the Albemarle Sound
flounder fishery included the capture of
131 Atlantic sturgeon. Of the 131
Atlantic sturgeon captured, no
mortalities were reported, although four
individuals were noted as having minor
injuries. These data indicate that
underreporting of sturgeon bycatch is
occurring in this area.
A sink gillnet survey conducted in the
Cape Fear River by University of North
Carolina at Wilmington personnel noted
that 25 percent of sturgeon intercepted
(22 of 88 caught) were killed. The
gillnets were set one day, checked the
second, and retrieved on the third. The
greatest mortality occurred during
periods of highest water temperature
(Moser et al., 1998). This survey was
continued by the NCDMF, and it has
reported mortality rates of 37 percent
overall. Similar to earlier findings,
mortality was greatest during the
summer months (June through August),
averaging 49 percent (34 of 69 sturgeon
died) (ASSRT, 2007). This study has
been discontinued due to lack of
funding. There are no estimates of
bycatch in fishery dependent surveys.
Winyah Bay is currently fished for
American shad using both sink and drift
gillnets. This fishery has an estimated
bycatch of 158 Atlantic sturgeon per
year, of which 16 percent (25 fish) die
and another 20 percent are injured to
some degree, although this estimate is
dated (Collins et al., 1996). Shad fishers
also operate within the rivers, but
neither fishing effort nor average
numbers of Atlantic sturgeon
encountered are known. Poaching of
adult Atlantic sturgeon has been
reported from the Winyah Bay area in
recent years. Carcasses of large females
have been found with the ovaries
(caviar) removed.
The mouth of the Santee River, just
south of Winyah Bay, has the largest
shad landings in the Southeast (ASSRT,
2007), likely resulting in mortality and
injury of sturgeon similar to that in the
Winyah Bay shad fishery. Upriver
bycatch levels are unknown. The
Cooper River also has an active hook
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5973
and line shad fishery because gillnets
are restricted (ASSRT, 2007).
The two largest commercial fisheries
likely to capture Atlantic sturgeon from
the South Atlantic DPS in the state
waters of South Carolina and Georgia
are the American shad gillnet and
shrimp trawl fisheries. Studies in
Georgia on commercial gillnet fisheries
for American shad (Alosa sapidissima)
showed that they accounted for 52
percent of Atlantic sturgeon bycatch and
the shrimp trawl fisheries accounted for
39 percent (Collins et al., 1996). The
American shad fisheries use sink
gillnets and drift gillnets. Collins et al.
(1996) documented a 16 percent
capture-induced mortality rate for
sturgeon in the American shad fishery.
There was a directed commercial
fishery for Atlantic sturgeon in the ACE
Basin prior to the 1985 fishery closure.
The commercial sturgeon fishery
operated in the lower and middle
portions of both the Combahee and
Edisto rivers. Commercial shad fisheries
captured some juvenile Atlantic
sturgeon, but most fishermen operate
upriver from the areas of greatest
abundance during that time of year. The
shrimp trawl fishery in St. Helena
Sound also captures juveniles, as
evident from tag returns (ASSRT, 2007).
Although a few commercial sturgeon
fishers apparently operated in the Port
Royal river system prior to 1985, the
landing of only one Atlantic sturgeon
has been recorded (Smith and Dingley,
1984). Little, if any, shad fishing takes
place in this system. It is not known
whether there is any significant bycatch
in the shrimp trawl fishery in this area.
During 1989 to 1991, the commercial
shad gillnet fishery’s bycatch in the
Savannah River included more
endangered shortnose sturgeon than
juvenile Atlantic sturgeon. Collins et al.
(1996) reported that two commercial
fishermen collected 14 Atlantic and 189
shortnose sturgeon over the period of
1990 to 1992. It appears that abundance
within the Savannah River is extremely
low, as evidenced from low bycatch and
reported captures over the last 15 years.
Thus, bycatch may be a more serious
impact if abundance is low and fishing
effort is high.
Bycatch in the shad fishery in the
Ogeechee River is a heightened concern
because evidence suggests that this
Atlantic sturgeon population is stressed
and that complete recruitment failure
has occurred in some years (ASSRT,
2007). Bycatch mortality in the
estuarine and lower river shad fishery is
suspected to be high, but no estimates
of take are available (ASSRT, 2007).
Estimated annual total bycatch of
Atlantic and shortnose sturgeon in the
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shad gillnet fishery in the tidal portion
of the Altamaha River during 1982 and
1983 averaged 372 sturgeon (Collins et
al., 1996). The mortality rate of sturgeon
taken as bycatch in the Altamaha River
during this time period was not
determined. During a study conducted
between 1986 and 1992 in the Altamaha
River, 97 of 1,534 tagged juvenile
Atlantic sturgeon were recaptured
primarily by shad gillnets (52 percent)
and shrimp trawls (39 percent) (Collins
et al., 1996). Juvenile Atlantic sturgeon
from the Altamaha are relatively
abundant in comparison to other rivers
in the region, so a large percentage of
the individuals in winter mixed-stock
aggregations on the shelf are likely from
this river. Most sturgeon occurring as
shrimp trawl bycatch are from mixedstock aggregations. Using the
percentages of Atlantic and shortnose
sturgeon from the 1986 to 1992
Altamaha catch data and applying them
to the 1982 and 1983 total estimated
sturgeon bycatch, it is expected that 89
percent (331 fish) of the catch consisted
of Atlantic sturgeon (ASSRT, 2007).
Also, assuming a 10 percent bycatch
mortality rate for Atlantic sturgeon from
drift nets (Stein et al., 2004b), the
dominant gear used in the shad gillnet
fishery, it is estimated that 33 Atlantic
sturgeon would die each year from the
fishery. However, in their latest
compliance report to the ASMFC,
GADNR noted that less than 10 fish per
year were estimated to have been
captured in the Altamaha River
anchored gillnet fishery during a 3-year
study. All fish were juveniles and no
injury or mortality was documented.
GADNR also noted the season for
gillnetting shad occurs while adults are
at sea and juveniles are in the lower
parts of the estuary. Since the 2007
status review, which ranked bycatch as
a moderate threat in the Altamaha, the
Georgia Board of Natural Resources has
prohibited the use of gillnets for shad
fishing in a large portion of the
Altamaha.
Shad fishing effort is low in the
Satilla River due to an apparently
depleted shad population. However,
because the Atlantic sturgeon
population is depleted and highly
stressed, any bycatch mortality could
have an impact on the population
(ASSRT, 2007).
The ASSRT concluded that bycatch
presents a moderate threat to the
Carolina DPS, while the threat of
bycatch to the South Atlantic DPS was
characterized as moderately low in each
of the populations, with the exception
of the Altamaha, where bycatch was
deemed to pose a moderate threat,
though we note again Georgia’s
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prohibition of shad gillnet fishing in a
large portion of the Altamaha since the
status review. Historical overutilization
of Atlantic sturgeon from directed
fishing caused initial severe declines in
Atlantic sturgeon populations in the
southeast, from which they have never
rebounded. Further, we believe
continued bycatch of Atlantic sturgeon
in commercial fisheries is an ongoing
impact to the Carolina and South
Atlantic DPSs that is contributing to
their endangered status. Atlantic
sturgeon are particularly vulnerable to
being caught in sink gillnets and
fisheries using this type of gear account
for most recorded Atlantic sturgeon
bycatch. However, little data exist on
bycatch in the Southeast, and high
levels of bycatch underreporting are
suspected (ASMFC, 2005; ASSRT, 2007;
White and Armstrong, 2000). Further,
total population abundances for the
Carolina and South Atlantic DPSs are
not available; therefore, it is not possible
to calculate the percentages of the
Carolina and South Atlantic DPSs
subject to bycatch mortality based on
the available bycatch mortality rates for
individual fisheries. However, fisheries
known to incidentally catch Atlantic
sturgeon occur throughout the marine
range of the species and in some
riverine waters as well. Because Atlantic
sturgeon mix extensively in marine
waters and may access multiple river
systems, they are subject to being caught
in multiple fisheries throughout their
range. Atlantic sturgeon taken as
bycatch may suffer immediate mortality.
In addition, stress or injury to Atlantic
sturgeon taken as bycatch but released
alive may result in increased
susceptibility to other threats, such as
poor water quality (e.g., exposure to
toxins and low DO). This may result in
reduced ability to perform major life
functions, such as foraging and
spawning, or may even result in postcapture mortality. Several of the river
populations in the South Atlantic DPS
(e.g., the Ogeechee and the Satilla) are
stressed to the degree that any level of
bycatch could have an adverse impact
on the status of the DPS (ASSRT, 2007).
C. Disease or Predation
Very little is known about natural
predators of Atlantic sturgeon. The
presence of bony scutes is likely an
effective adaptation for minimizing
predation of sturgeon greater than 25
mm (Gadomski and Parsley, 2005).
Gadomski and Parsley (2005) have
shown that catfish and other species do
prey on juvenile sturgeon, and concerns
have been raised regarding the potential
for increased predation on juvenile
Atlantic sturgeon by introduced flathead
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catfish (Brown et al., 2005). Atlantic
sturgeon populations are persisting in
the Cape Fear River, North Carolina, and
Altamaha River, Georgia, where
flatheads have been present for many
years, at least in the absence of any
directed fisheries for Atlantic sturgeon.
Thus, further research is warranted to
determine at what level, if any,
flatheads and other exotic species prey
upon juvenile Atlantic sturgeon and to
what extent such predation is affecting
the sturgeon populations.
While some disease organisms have
been identified from wild Atlantic
sturgeon, they are unlikely to threaten
the survival of the wild populations.
Disease organisms commonly occur
among wild fish populations, but under
favorable environmental conditions,
these organisms are not expected to
cause population-threatening
epidemics. There is concern that nonindigenous sturgeon pathogens could be
introduced, most likely through
aquaculture operations. Fungal
infections and various types of bacteria
have been noted to have various effects
on hatchery Atlantic sturgeon. Due to
this threat of impacts to wild
populations, the ASMFC recommends
requiring any sturgeon aquaculture
operation to be certified as disease-free,
thereby reducing the risk of the spread
of disease from hatchery origin fish. The
aquarium industry is another possible
source for transfer of non-indigenous
pathogens or non-indigenous species
from one geographic area to another,
primarily through release of aquaria fish
into public waters. With millions of
aquaria fish sold to individuals
annually, it is unlikely that such activity
could ever be effectively regulated.
Definitive evidence that aquaria fish
could be blamed for transmitting a nonindigenous pathogen to wild fish
(sturgeon) populations would be very
difficult to collect (ASSRT, 2007).
In their status review, the ASSRT
ranked the threat from disease and
predation as a low risk. While
information on the impacts of disease
and predation on Atlantic sturgeon is
limited, there is nothing to indicate that
either of these factors is currently
having any measurable adverse impact
on Atlantic sturgeon. Therefore, we
concur with the ASSRT, and we
conclude that disease and predation are
not contributing to the endangered
status of either the Carolina or the South
Atlantic DPS.
D. Inadequacy of Existing Regulatory
Mechanisms
As a wide-ranging anadromous
species, Atlantic sturgeon are subject to
numerous Federal (U.S. and Canadian),
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state and provincial, and interjurisdictional laws, regulations, and
agency activities. These regulatory
mechanisms are described in detail in
the status review report (see Section
3.4). We believe that the inadequacy of
regulatory mechanisms to control
bycatch and the modification and
curtailment of Atlantic sturgeon habitat
is contributing to the endangered status
of the Carolina and South Atlantic
DPSs.
Current regulatory mechanisms have
effectively removed threats from legal,
directed harvest in the United States, as
well as incentives for retention of
bycatch. The ASMFC was given
management authority in 1993 under
the Atlantic Coastal Fisheries
Cooperative Management Act
(ACFCMA) (16 U.S.C. 5101–5108), and
it manages Atlantic sturgeon through an
interstate fisheries management plan
(IFMP). The moratorium prohibiting
directed catch of Atlantic sturgeon was
developed as an Amendment to the
IFMP. The ACFCMA, authorized under
the terms of the ASMFC Compact, as
amended (Pub. L. 103–206), provides
the Secretary of Commerce with the
authority to implement regulations that
are compatible to ASMFC FMPs in the
Exclusive Economic Zone (EEZ) in the
absence of an approved MagnusonStevens FMP. In 1999, it was under this
authority that a similar moratorium was
implemented for Atlantic sturgeon in
Federal waters. The Amendment
includes a stock rebuilding target of at
least 20 protected mature age classes in
each spawning stock, which is to be
achieved by imposing a harvest
moratorium. The Amendment requires
states to monitor, assess, and annually
report Atlantic sturgeon bycatch and
mortality in other fisheries. The
Amendment also requires that states
annually report habitat protection and
enhancement efforts. Finally, the
Amendment states that each jurisdiction
with a reproducing population should
conduct juvenile assessment surveys
(including CPUE estimates, tag and
release programs, and age analysis), and
states with rivers that lack a
reproducing sturgeon population(s) but
support nursery habitat for migrating
juveniles should also conduct sampling.
While the ASMFC and NMFS have
made significant strides in reducing the
threats from direct harvest and retention
of bycatch, those threats have not been
eliminated, and continued bycatch of
Atlantic sturgeon is contributing to the
endangered status of the Carolina and
South Atlantic DPSs. Although the FMP
contains requirements for reporting
bycatch, fishery managers, such as the
ASMFC Atlantic Sturgeon Management
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Board, widely accept that Atlantic
sturgeon bycatch is underreported or
not reported at all based on research and
anecdotal evidence (ASMFC, 2005;
ASSRT, 2007; White and Armstrong,
2000). Abundance estimates are
available only for two river systems (the
Hudson and the Altamaha) even though
the FMP states that each jurisdiction
with a reproducing population should
conduct juvenile assessment surveys
(including CPUE estimates, tag and
release programs, and age analysis).
While the aforementioned mechanisms
have addressed impacts to Atlantic
sturgeon through directed fisheries,
there are currently no mechanisms in
place to address the significant impacts
and risks posed to Atlantic sturgeon by
commercial bycatch.
State and Federal agencies are
actively employing a variety of legal
authorities to implement proactive
restoration activities for this species,
and coordination of these efforts is
being furnished through the ASMFC.
Due to existing state and Federal laws,
water quality and other habitat
conditions have improved in many
riverine habitats, although many
systems still have DO and toxic
contaminants issues, and habitat quality
and quantity continue to be affected by
dams, dredging, and/or altering natural
flow conditions.
Though statutory and regulatory
mechanisms exist that authorize
reducing the impact of dams on riverine
and anadromous species, such as
Atlantic sturgeon, and their habitat,
these mechanisms have proven
inadequate for preventing dams from
blocking access to habitat upstream and
degrading habitat downstream.
Hydropower dams are regulated by the
FERC. The Federal Power Act, originally
enacted in 1920, provides for
cooperation between FERC and other
Federal agencies, including resource
agencies, in licensing and relicensing
power projects. The Federal Power Act
authorizes NMFS to recommend
hydropower license conditions to
protect, mitigate damages to, and
enhance anadromous fish, including
related habitat. The Federal Power Act
also provides authority for NMFS to
issue mandatory fishway prescriptions.
FERC licenses have a term of 30 to 50
years, so NMFS’ involvement in the
licensing process to ensure the
protection and accessibility of upstream
habitat, and to improve habitat degraded
by changes in water flow and quality
from dam operations, may only occur
twice or thrice a century. The Federal
Power Act does not apply to nonhydropower dams, such as those
operated by the Army Corps of
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Engineers for navigation purposes. Even
where fish passage currently exists,
evidence is rare that it effectively passes
sturgeon, including Atlantic sturgeon.
As mentioned in previous sections,
dams in the Southeast are currently
blocking access to over 60 percent of the
habitat in three rivers with historical
and/or current spawning Atlantic
sturgeon populations (the Cape Fear
River and Santee-Cooper System in the
Carolina DPS and the St. Johns River in
the South Atlantic DPS), though we are
hopeful that NMFS’ 2007 fishway
prescription of passage for sturgeon
through the lowest dams on both the
Santee and Cooper Rivers will be
implemented once FERC issues the new
license for this project in the near
future. In addition to the loss of
important spawning and juvenile
developmental habitat upstream, dam
operations reduce the quality of the
remaining habitat downstream by
affecting water quality parameters (such
as depth, temperature, velocity, and DO)
that are important to Atlantic sturgeon.
Therefore, the inadequacy of regulatory
mechanisms to ensure safe and effective
upstream and downstream passage to
Atlantic sturgeon and prevent
degradation of habitat downstream from
dam operations in riverine habitat is
contributing to the endangered status of
the Carolina and South Atlantic DPSs.
Inadequacies in the regulation of
water allocation also impact the South
Atlantic DPS. Data concerning
consumptive water use in this region
are, at best, very limited. While
extensive data exist concerning
permitted water withdrawals, there is
little information concerning actual
withdrawals and virtually no
information concerning water
discharges. This is particularly the case
for municipal and industrial uses
because water use permits are not
required for withdrawals less than
100,000 gpd (379 m3pd) (Cummings et
al., 2003) and discharge permits are not
required unless discharge contains
selected toxic materials. Agricultural
water use permits are not quantified,
neither water withdrawals nor return
flows are measured (Fisher et al., 2003).
While several other states have similar
permitting thresholds, the majority
require permits for water withdrawals
less than 100,000 gpd (379 m3pd) and
some require a permit for any water
withdrawal. The present limit in
Georgia allows access to water in
amounts required to satisfy the
household needs of more than 300
households without a permit
(Cummings et al., 2003).
Fundamental requisites for basin
water planning—data for historical,
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unimpaired flows in the coastal regions’
rivers—do not exist (Fisher et al., 2003).
There are 125 river gauges in the
region’s 7 river basins. However, 72 of
these gauges are inactive, and 28 of the
remaining 53 gauges do not provide
consistent flow information. Moreover,
historical data from many gauges have
gaps, reflecting periods (sometimes
extending over months) during which
the gauge was inoperative. Also, there
are extensive discharge areas between
the last gauge in each river system and
the point at which the river discharges
into the ocean—thus, there are
potentially large water supplies for
which no information is available
(Fisher et al., 2003).
Water quality continues to be a
problem, even with existing controls on
some pollution sources. Data required to
evaluate water allocation issues are
either very weak, in terms of
determining the precise amounts of
water currently being used, or nonexistent, in terms of our knowledge of
water supplies available for use under
historical hydrologic conditions in the
region. Current regulatory regimes are
not sufficiently effective in controlling
water allocation issues (e.g., no permit
requirements for water withdrawals
under 100,000 gpd (379 m3pd) in
Georgia and no restrictions on
interbasin water transfers in South
Carolina).
In their status review, the ASSRT
ranked the threat from the inadequacy
of regulatory mechanisms as moderately
low to moderate. While some of the
threats to the Carolina and South
Atlantic DPSs have been ameliorated or
reduced due to the existing regulatory
mechanisms, such as the moratorium on
directed fisheries for Atlantic sturgeon,
bycatch is currently not being addressed
through existing mechanisms. Further,
water quality continues to be a problem
even with existing controls on some
pollution sources and water withdrawal,
and dams continue to curtail and
modify habitat, even with the Federal
Power Act.
E. Other Natural or Manmade Factors
Affecting the Species’ Continued
Existence
The ASSRT considered several
manmade factors that may affect
Atlantic sturgeon, including
impingement and entrainment, ship
strikes, and artificial propagation. The
vast withdrawal of water from rivers
that support Atlantic sturgeon
populations was considered to pose a
threat of impingement and entrainment;
however, data are lacking to determine
the overall impact of this threat on
sturgeon populations, as impacts are
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dependent on a variety of factors (e.g.,
the species, time of year, location of the
intake structure, and strength of the
intake current). Multiple suspected
boat/ship strikes have been reported in
several rivers. A large number of the
mortalities observed in these rivers from
potential ship strikes have been of large
adult Atlantic sturgeon. Lastly, potential
artificial propagation of Atlantic
sturgeon was also a concern to ASSRT
members, as both stock enhancement
programs and commercial aquaculture
can have negative impacts on a
recovering population (e.g., fish disease,
escapement, outbreeding depression). In
order to circumvent these potential
threats, stock enhancement programs
follow culture and stocking protocols
approved by the ASMFC. Commercial
aquaculture facilities are expected to
maintain disease-free facilities and have
safeguards in place to prevent
escapement of sturgeon into the wild.
While in at least one instance cultured
Atlantic sturgeon have gone
unaccounted for from a commercial
aquaculture facility in Florida, this is
not considered to be a significant threat,
as this was a rare event. Mechanisms are
in place at all facilities to prevent
escapement of sturgeon; facilities are all
land based, and most are not located in
close proximity to any Atlantic sturgeon
rivers.
Along the range of Atlantic sturgeon
from the Carolina and South Atlantic
DPSs, most, possibly all, populations
are at risk of possible entrainment or
impingement in water withdrawal
intakes for commercial uses, municipal
water supply facilities, and agricultural
irrigation intakes. In North Carolina,
over two billion gallons of water per day
were withdrawn from the Cape Fear,
Neuse, Tar, and Roanoke rivers in 1999
by agriculture and non-agricultural
industries (NCDENR, 2006). Three
surveys, included in the 2007 status
review, have shown the direct impacts
of water withdrawal on Atlantic
sturgeon: (1) Hudson River Utility
Surveys, (2) Delaware River Salem
Power Plant survey, and (3) Edwin I.
Hatch Nuclear Power Plant survey.
Information on the Brunswick Nuclear
Power Plant and its impacts on Atlantic
sturgeon was provided by Progress
Energy during the public comment
period on the proposed listing rule. The
Edwin I. Hatch Nuclear power plant
(HNP) is located 11 miles north of
Baxley, Georgia. The HNP uses a closedloop system for main condenser cooling
that withdraws from, and discharges to,
the Altamaha River. Pre-operational
drift surveys were conducted and only
two Acipenser larvae were collected.
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Entrainment samples at HNP were
collected for the years 1975, 1976, and
1980, and no Acipenser species were
observed in the samples (Sumner, 2004).
The Brunswick Nuclear Power Plant is
located on the lower Cape Fear River.
An average of 55 juvenile Atlantic
sturgeon were impinged per year from
1975 to 1981. Plant modifications were
implemented in the early 1980s as part
of the NPDES permit. A fish diversion
was installed in 1981 and a fish return
system was installed in 1983. Only two
impinged juveniles were observed
between 1982 and 2010 and were
returned alive to the river. Though most
rivers have multiple intake structures
which remove millions of gallons a day
during the spring and summer months,
it is believed that the migratory behavior
of larval sturgeon allows them to avoid
intake structures, since migration is
active and occurs in deep water (Kynard
and Horgan, 2002). Effluent from these
facilities can also affect populations, as
some facilities release heated water that
acts as a thermal refuge during the
winter months, but drastic changes in
water temperature have the potential to
cause mortality.
Locations that support large ports and
have relatively narrow waterways are
more prone to ship strikes (e.g.,
Delaware, James, and Cape Fear rivers).
One ship strike per 5 years is reported
for the Cape Fear River within the
Carolina DPS. Ship strikes have not
been documented in any of the rivers
within the South Atlantic DPS. While it
is possible that ship strikes may have
occurred that have gone unreported or
unobserved, the lack of large ship traffic
on narrow waterways within the range
of the DPS may limit potential
interactions.
Artificial propagation of Atlantic
sturgeon for use in restoration of
extirpated populations or recovery of
severely depleted wild populations has
the potential to be both a threat to the
species and a tool for recovery. Within
the range of the Carolina DPS, several
attempts were made by Smith et al.
(1980 and 1981) to hormonally-induce
spawning and culture Atlantic sturgeon
captured in the Atlantic Ocean off the
Winyah Bay jetties. Fry were produced
during each spawning attempt, but the
fry lived less than a year. As a result of
successful spawning of Hudson River
Atlantic sturgeon from 1993 to 1998,
USFWS’ Northeast Fisheries Center
(NEFC) is currently rearing five yearclasses of domestic fish. These fish
could potentially be used as broodstock
for aquaculture operations and stock
enhancement, provided that there is no
risk to wild fish. Aquaculturists along
the East Coast, including some in North
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Carolina and South Carolina, have
contacted the NEFC and expressed
interest in initiating commercial
production of Atlantic sturgeon. In
2006, La Paz Aquaculture Group was
approved by North Carolina state
resource agencies and ASMFC to
produce Atlantic sturgeon for flesh and
caviar sales. However, their first year of
production was halted because remnant
storms from Hurricane Katrina
destroyed their fry stock. In August
2006, ASMFC reevaluated the La Paz
permit, and voted to draft an addendum
to allow La Paz to acquire Atlantic
sturgeon from multiple Canadian
aquaculture companies (previously
restricted to one company), allowing
them to resume Atlantic sturgeon
culture. Resource managers who
reviewed the permit found the La Paz
facility to pose little threat to Atlantic
sturgeon or shortnose populations due
to the facility location (far inland), use
of a recirculating system, and land
application of any discharge (ASSRT,
2007).
In the range of the South Atlantic
DPS, artificial propagation has been
attempted for the purposes of both
restoration and commercial profit. The
St. Marys Fish Restoration Committee
(SMFRC) is working with Florida and
Georgia to reestablish Atlantic sturgeon
in the St. Marys River. Efforts are
currently underway to refine restoration
approaches within the system. Phase 1
of the restoration plan includes a
population and habitat assessment.
Field investigations are being funded
through ESA Section 6 and coordinated
through Georgia DNR. The State of
Florida has been involved in fish
sampling and will continue to explore
and refine sturgeon sampling strategies.
Aquatic habitat and water quality
surveillance work will continue to be
accomplished by the St. Johns River
Water Management District, the
Environmental Protection Agency,
Florida Department of Environmental
Protection, USFWS, TNC, and the St.
Marys River Management Committee.
Phase 2 of the plan would include
experimental transplanting of Atlantic
sturgeon to assess environmental
factors, habitat use at different lifestages, contaminants, migration-homing,
etc. Upon approval from the ASMFC,
the SMFRC transferred 12 Atlantic
sturgeon from the Altamaha River in
Georgia to the Bears Bluff National Fish
Hatchery in South Carolina. The SMFRC
hopes to develop and refine captive
propagation techniques for predictable
spawning and provide fish to approved
researchers.
Aquaculturists in South Carolina and
Florida have also contacted the NEFC
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and expressed interest in initiating
commercial production of Atlantic
sturgeon through use of the Hudson
River broodstock. In 2001, the Canadian
Caviar Company shipped 18,000
Atlantic sturgeon sac fry to the
University of Florida. These fry were
used to conduct early larval and feeding
trials. Survivors of these experiments
were transferred to four aquaculture
businesses: (1) Evan’s Fish Farm in
Pierson, Florida; (2) Watts Aquatics in
Tampa, Florida; (3) Hi-Tech Fisheries of
Florida in Lakeland, Florida; and (4)
Rokaviar in Homestead, Florida.
According to information provided by
FDACS in August 2011, Evan’s Fish
Farm is the only aquaculture facility
still in possession of Atlantic sturgeon.
They experienced a catastrophic
systems failure in 2004 and currently
have only one Atlantic sturgeon on their
premises. The remaining Atlantic
sturgeon obtained from Canada by
Florida aquaculture facilities died in
captivity.
The ASSRT ranked the threats from
impingement/entrainment, ship strikes,
and artificial propagation as low for
both DPSs, with the exception of the
threat from ship strikes as moderately
low for the Carolina DPS. We concur
with these rankings and conclude that
none of these threats are contributing to
the endangered status of the DPS.
Current Protective Efforts
Section 4(b)(1)(A) of the ESA requires
the Secretary, when making a listing
determination for a species, to take into
account those efforts, if any, being made
by any State or foreign nation to protect
the species. In judging the efficacy of
existing protective efforts, we rely on
the Services’ joint ‘‘Policy for
Evaluation of Conservation Efforts
When Making Listing Decisions’’
(‘‘PECE;’’ 68 FR 15100; March 28, 2003).
The PECE is designed to guide
determinations on whether any
conservation efforts that have been
recently adopted or implemented, but
not yet proven to be successful, will
result in recovering the species to the
point at which listing is not warranted
or contribute to forming a basis for
listing a species as threatened rather
than endangered. The purpose of the
PECE is to ensure consistent and
adequate evaluation of future or recently
implemented conservation efforts
identified in conservation agreements,
conservation plans, management plans,
and similar documents when making
listing decisions. The PECE provides
direction for the consideration of such
conservation efforts that have not yet
been implemented, or have been
implemented but have not yet
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demonstrated effectiveness. The policy
is expected to facilitate the development
by states and other entities of
conservation efforts that sufficiently
improve a species’ status so as to make
listing the species as threatened or
endangered unnecessary.
The Services established two basic
criteria in the PECE: (1) The certainty
that the conservation efforts will be
implemented, and (2) the certainty that
the efforts will be effective. Satisfaction
of the criteria for implementation and
effectiveness establishes a given
protective effort as a candidate for
consideration, but does not mean that
an effort will ultimately change the risk
assessment for the species. Through the
PECE analysis, the Services ascertain
whether the formalized conservation
effort improves the status of the species
at the time a listing determination is
made.
We evaluated the current
conservation efforts underway to protect
and recover Atlantic sturgeon in making
our listing determination. In the 2007
status review report and the proposed
listing rule, we determined that only the
following conservation efforts warrant
consideration under the PECE for the
Carolina and South Atlantic DPSs: The
1998 ASMFC FMP and the proposal by
the SMFRC to restore Atlantic sturgeon
to the St. Marys River. In addition, we
evaluated North Carolina’s NCCHPP and
designation of AFSAs based on
information submitted during the public
comment period on the proposed listing
rule.
The 1998 Amendment to the ASMFC
Atlantic Sturgeon FMP strengthens
conservation efforts by formalizing the
closure of the directed fishery, and by
banning possession of bycatch,
eliminating any legal incentive to retain
Atlantic sturgeon. However, bycatch is
known to occur in several fisheries
(ASMFC, 2007) and it is widely
accepted that bycatch is underreported
(ASMFC, 2005; ASSRT, 2007; White
and Armstrong, 2000). Contrary to
information available in 1998 when the
Amendment was approved, Atlantic
sturgeon bycatch mortality is a major
stressor affecting the recovery of
Atlantic sturgeon, despite actions taken
by the states and NMFS to prohibit
directed fishing and retention of
Atlantic sturgeon. Therefore, there is
considerable uncertainty that the
Atlantic Sturgeon FMP will be effective
in meeting its conservation goals. In
addition, though the 1998 Amendment
contains requirements for population
surveys, it is highly uncertain these will
be implemented, as there are limited
resources for assessing current
abundance of spawning females for each
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of the DPSs and to date, abundance
estimates have only been completed for
one river within the range of the two
DPSs considered here. For these
reasons, there is great uncertainty
regarding the implementation and
effectiveness of the intended ASMFC
FMP conservation effort for the Carolina
and South Atlantic DPSs of Atlantic
sturgeon.
The SMFRC is working with Florida
and Georgia with the intention of
reestablishing Atlantic sturgeon in the
St. Marys River. Efforts are currently
underway to refine restoration
approaches within the system. As
discussed in Section E, Phase 1 of the
restoration plan includes a population
and habitat assessment, and Phase 2
includes experimental transplanting of
Atlantic sturgeon to assess
environmental factors, habitat use at
different life-stages, contaminants,
migration-homing, etc. Atlantic sturgeon
are believed to be extirpated in the St.
Marys River. This conservation effort
may increase our knowledge and
understanding of Atlantic sturgeon
status and habitat conditions in the St.
Marys River, as well as provide methods
for restoring a population there in the
future. As previously discussed,
artificial propagation of Atlantic
sturgeon for use in restoration of
extirpated populations or recovery of
severely depleted wild populations has
the potential to be both a threat to the
species and a tool for recovery. Because
it is in the earliest stages of planning,
development, and authorization, the
feasibility of any project or the potential
degree of success for this effort is
unknown. Therefore, the SMRFC efforts
do not satisfy the PECE policy’s
standards for certainty of
implementation or effectiveness.
The State of North Carolina adopted
the NCCHPP in 2005 and its stated goals
are: (1) Improving effectiveness of
existing rules and programs protecting
coastal fish habitats; (2) identifying,
designating, and protecting strategic
habitat areas (SHAs); (3) enhancing
habitat and protecting it from physical
impacts; and (4) enhancing and
protecting water quality. The NCMFC
approved SHAs for Region 1 (the waters
and adjacent wetlands draining into and
out of Albemarle Sound through Oregon
Inlet to the adjoining coastal ocean) in
North Carolina in January 2009, and is
currently evaluating SHAs for other
regions in North Carolina. SHAs
represent priority habitat areas for
protection due to their exceptional
condition or imminent threat to their
ecological functions supporting
estuarine and coastal fish and shellfish
species and will be incorporated into
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conservation and restoration efforts.
One SHA (Bellows Bay to Knotts Island
Bay) was identified in part due to the
nearshore ocean areas that are important
for Atlantic sturgeon and striped bass
and another SHA (Chowan and Roanoke
Rivers and western Albemarle Sound)
may include one of the few Atlantic
sturgeon spawning habitats in North
Carolina. NCDMF also provides input to
federal and state regulatory agencies of
the location of habitats used by Atlantic
sturgeon. NCDMF and the North
Carolina Wildlife Resources
Commission have designated
Anadromous Fish Spawning Areas
(AFSA) through rules for their
respective jurisdictions. While these
programs have excellent goals of
increasing enforcement of existing
regulations, identifying and protecting
habitat important to the species, and
monitoring these habitats, these actions
are still in the early stages and it is not
clear exactly what protections will be
given to areas designated as SHAs or
AFSAs. Therefore, the efforts associated
with the NCCHPP and the designation
of AFSAs do not satisfy the PECE
policy’s standards for certainty of
implementation or effectiveness.
Listing Determinations
Carolina DPS
The Carolina DPS is estimated to
number less than 3 percent of its
historical population size (ASSRT,
2007). Prior to 1890, Secor (2002)
estimated there were between 7,000 and
10,000 adult females in North Carolina
and 8,000 adult females in South
Carolina. Currently, there are estimated
to be less than 300 adults spawning
annually (total of both sexes) in the
major river systems occupied by the
DPS in which spawning still occurs,
whose freshwater range occurs in the
watersheds from the Roanoke River
southward along the southern Virginia,
North Carolina, and South Carolina
coastal areas to the Cooper River. We
have reviewed the status review report,
as well as other available literature and
information, and have consulted with
scientists and fishery resource managers
familiar with the Atlantic sturgeon in
the Carolina DPS. We considered
relevant substantial information and
recommendations made by the peer
reviewers and the public on the
proposed listing rule. After reviewing
the best scientific and commercial
information available, we find that the
Atlantic sturgeon Carolina DPS is in
danger of extinction throughout its
range as a result of a combination of
habitat curtailment and alteration,
bycatch in commercial fisheries, and
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inadequacy of regulatory mechanisms in
ameliorating these impacts and threats,
and have determined it should be listed
as endangered.
South Atlantic DPS
The South Atlantic DPS is estimated
to number less than 6 percent of its
historical population size (ASSRT,
2007), with all river populations except
the Altamaha estimated to be less than
1 percent of historical abundance. Prior
to 1890, Secor (2002) estimated there
were 8,000 adult spawning females in
South Carolina and 11,000 adult
spawning females in Georgia. Currently,
there are an estimated 343 adults
spawning annually in the Altamaha and
less than 300 adults spawning annually
(total of both sexes) in the other major
river systems occupied by the DPS in
which spawning still occurs, whose
freshwater range occurs in the
watersheds of the ACE Basin in South
Carolina to the St. Johns River, Florida.
We have reviewed the status review
report, as well as other available
literature and information, and have
consulted with scientists and fishery
resource managers familiar with the
Atlantic sturgeon in the South Atlantic
DPS. We considered relevant substantial
information and recommendations
made by the peer reviewers and the
public on the proposed listing rule.
After reviewing the best scientific and
commercial information available, we
find that the Atlantic sturgeon South
Atlantic DPS is in danger of extinction
throughout its range as a result of a
combination of habitat curtailment and
alteration, bycatch in commercial
fisheries, and inadequacy of regulatory
mechanisms in ameliorating these
impacts and threats, and have
determined it should be listed as
endangered.
Effects of Listing
Conservation measures provided for
species listed as endangered under the
ESA include recovery actions (16 U.S.C.
1533(f)), critical habitat designations,
Federal agency consultation
requirements (16 U.S.C. 1536), and
prohibitions on taking (16 U.S.C. 1538).
Recognition of the species’ endangered
status through listing promotes
conservation actions by Federal and
state agencies, private groups, and
individuals.
Identifying Section 7 Consultation
Requirements
Section 7(a)(2) of the ESA requires
Federal agencies to consult with NMFS
to ensure that activities authorized,
funded, or carried out by those agencies
are not likely to jeopardize the
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continued existence of the species or
destroy or adversely modify critical
habitat. We do not know how many
section 7 consultations may be required
for Federal agencies. From 2005 to 2010,
there were 108 informal and 10 formal
consultation requests for the shortnose
sturgeon, a species whose range
overlaps with that of Atlantic sturgeon
in freshwater and estuarine habitats.
The Carolina and South Atlantic DPSs
are distinguished based on genetic data
and spawning locations. However,
extensive mixing of the populations
occurs in coastal waters. Therefore, the
distributions of the DPSs outside of
natal waters generally overlap with one
another, and with fish from Northeast
river populations. This presents a
challenge in conducting ESA section 7
consultations because fish from any DPS
could potentially be affected by a
proposed project. Project location alone
will likely not inform the section 7
biologist as to which populations to
consider in the analysis of a project’s
potential direct and indirect effects on
Atlantic sturgeon and their habitat. This
will be especially problematic for
projects where take could occur,
because it is critical to know which
Atlantic sturgeon population(s) to
include in the jeopardy analysis. One
conservative but potentially
cumbersome method would be to
analyze the total anticipated take from
a proposed project as if all Atlantic
sturgeon came from a single DPS and
repeat the jeopardy analysis for each
DPS the taken individuals could have
come from. However, recently funded
research may shed some light on the
composition of mixed stocks of Atlantic
sturgeon, relative to their rivers of
origin, in locations along the East Coast.
The specific purpose of the study is to
evaluate the vulnerability to coastal
bycatch of Hudson River Atlantic
sturgeon, thought to be the largest stock
contributing to coastal aggregations from
the Bay of Fundy to Georgia. However,
the mixed stock analysis will also allow
NMFS to better estimate a project’s
effects on different components of a
mixed stock of Atlantic sturgeon in
coastal waters or estuaries other than
where they were spawned. Results from
the study are expected by early 2012.
Genetic mixed stock analysis, such as
proposed in this study, requires a high
degree of resolution among stocks
contributing to mixed aggregations and
characterization of most potential
contributory stocks. Fortunately, almost
all extant populations have been
characterized in previous genetic
studies, though some additional
populations will be characterized in this
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study. Genetic testing of mixed stocks
will be conducted in eight coastal
locales in both the Northeast and
Southeast Regions. Coastal fisheries and
sites were selected based on sample
availabilities, bycatch concerns, and
specific biological questions (i.e., real
uncertainty as to stock origins of the
coastal aggregation).
Critical Habitat
Critical habitat is defined in section 3
of the ESA (16 U.S.C. 1532(3)) as: (1)
The specific areas within the
geographical area occupied by a species,
at the time it is listed in accordance
with the ESA, on which are found those
physical or biological features (a)
essential to the conservation of the
species and (b) that may require special
management considerations or
protection; and (2) specific areas outside
the geographical area occupied by a
species at the time it is listed upon a
determination that such areas are
essential for the conservation of the
species. ‘‘Conservation’’ means the use
of all methods and procedures needed
to bring the species to the point at
which listing under the ESA is no
longer necessary.
Section 4(a)(3)(a) of the ESA
(16 U.S.C. 1533(a)(3)(A)) requires that,
to the extent prudent and determinable,
critical habitat be designated
concurrently with the listing of a
species. Section 4(b)(6)(C)(ii) of the ESA
provides for additional time to
promulgate a critical habitat designation
if such designation is not determinable
at the time of final listing of a species.
Designations of critical habitat must be
based on the best scientific data
available and must take into
consideration the economic, national
security, and other relevant impacts of
specifying any particular area as critical
habitat. The designation of critical
habitat is not determinable at this time
due to the extensive range of the
Carolina and South Atlantic DPSs and
extremely complex biological and
physical requirements of Atlantic
sturgeon. Although we have gathered
information through the status review
and public comment processes, we
currently do not have enough
information to determine which of these
features are essential to the conservation
of the two DPSs and may require special
management considerations or
protection. We will continue to gather
and review other ongoing studies on the
habitat use and requirements of Atlantic
sturgeon to attempt to identify these
features. Additionally, we need more
time to gather the information needed to
perform the required analyses of the
impacts of the designation. Once areas
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containing the essential features are
identified and mapped, and economic,
national security, and other relevant
impacts are considered, we will publish,
in a separate rule, a proposed
designation of critical habitat for the
Carolina and South Atlantic DPSs.
Section 9 Take Prohibitions
ESA section 9(a) and 16 U.S.C. 1538
(a)(1)(B) take prohibitions apply to all
species listed as endangered. These
include prohibitions against the import,
export, use in foreign commerce, or
‘‘take’’ of the species. Take is defined as
‘‘to harass, harm, pursue, hunt, shoot,
wound, kill, trap, capture, or collect, or
to attempt to engage in any such
conduct.’’ These prohibitions apply to
all persons subject to the jurisdiction of
the United States, including in the U.S.
or on the high seas.
Identification of Those Activities That
Would Constitute a Violation of Section
9 of the ESA
On July 1, 1994, we and USFWS
published a policy to identify, to the
maximum extent possible, those
activities that would or would not
constitute a violation of section 9 of the
ESA (59 FR 34272; July 1, 1994). The
intent of this policy is to increase public
awareness of the effect of this listing on
proposed and ongoing activities within
the species’ range. We will identify, to
the extent known, specific activities that
will not be considered likely to result in
violation of section 9, as well as
activities that will be considered likely
to result in violation.
Activities that we believe could result
in violation of section 9 prohibitions
against ‘‘take’’ of the Atlantic sturgeon
in the Carolina and South Atlantic DPSs
include, but are not limited to, the
following: (1) Capture and mortality in
commercial and recreational fisheries;
(2) poaching of individuals for meat or
caviar; (3) marine vessel strikes; (4)
destruction of or blocking access to
riverine, estuarine, and marine habitat
through such activities as agricultural
and urban development, commercial
activities, diversion of water for
hydropower and public consumption,
and dredge and fill operations; (5)
impingement and entrainment in water
control structures; (6) unauthorized
collecting or handling of the species
(permits to conduct these activities are
available for purposes of scientific
research or to enhance the propagation
or survival of the DPSs); (7) releasing a
captive Atlantic sturgeon into the wild;
and (8) harming captive Atlantic
sturgeon by, among other things,
injuring or killing them through
veterinary care, research, or breeding
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activities outside the bounds of normal
animal husbandry practices. Permits to
conduct activities that may result in
‘‘take’’ of Atlantic sturgeon for scientific
purposes or to enhance the propagation
or survival of the DPSs may be issued
under section 10 of the ESA. Such
permits would be required to authorize
take regardless of whether the sturgeon
were in captivity at the time this final
listing rule becomes effective, or are
collected from the wild after this rule
becomes effective.
ESA sections 10(a)(1)(A) and
10(a)(1)(B) provide NMFS with
authority to grant exceptions to the
section 9 take prohibitions. Section
10(a)(1)(A) scientific research and
enhancement permits may be issued to
entities (Federal and non-Federal)
conducting research that involves a take
of listed species. We have issued section
10(a)(1)(A) research and enhancement
permits for other listed species for these
purposes. ESA section 10(a)(1)(B)
incidental take permits may be issued to
non-Federal entities performing
activities that may incidentally take
listed species. The ESA also provides
some exceptions to the prohibitions,
without permits, for certain antique
articles and species held in captivity at
the time of listing. ESA section 10(h)
allows antique articles of listed species
to be excluded from essentially all the
ESA prohibitions as long as they are at
least 100 years old and meet certain
other specified conditions. Section
9(b)(1) provides a narrow exemption for
animals held in captivity at the time of
listing: those animals are not subject to
the import/export prohibition in section
9(a)(1)(A) or to protective regulations
adopted by the Secretary under section
9(a)(1)(G), so long as the holding of the
species in captivity, before and after
listing, is not in the course of a
commercial activity and does not violate
the applicable prohibitions under ESA
section 9(a)(1). However, 180 days after
listing there is a rebuttable presumption
that the exemption does not apply.
Thus, in order to apply this exemption,
the burden of proof for confirming the
status of animals held in captivity prior
to listing lies with the holder. The
section 9(b)(1) exemption for captive
wildlife would not apply to any progeny
of the captive animals that may be
produced post-listing.
Based on the best available
information, we believe that the
following actions will not result in a
violation of ESA section 9: (1) Take or
possession of Atlantic sturgeon acquired
lawfully by permit issued by NMFS
pursuant to section 10 of the ESA, or
take in accordance with the terms of an
incidental take statement in a biological
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opinion pursuant to section 7 of the
ESA; (2) Federally approved projects
that involve activities such as
agriculture, managed fisheries, road
construction, discharge of fill material,
stream channelization, or diversion for
which consultation under section 7 of
the ESA has been completed and
determined not likely to jeopardize the
continued existence of the Atlantic
sturgeon DPS, and when such activity is
conducted in accordance with any terms
and conditions given by NMFS in an
incidental take statement in a biological
opinion pursuant to section 7 of the
ESA; (3) continued possession of live
Atlantic sturgeon that were in captivity
or in a controlled environment (e.g., in
aquaria) at the time of this listing, so
long as the applicable prohibitions
under an ESA section 9(a)(1) are not
violated; and, (4) provision of care for
live Atlantic sturgeon that were in
captivity at the time of this listing.
Policies on Peer Review
On July 1, 1994, NMFS and USFWS
published a series of policies regarding
listings under the ESA, including a
policy for peer review of scientific data
(59 FR 34270; July 1, 1994), the Office
of Management and Budget (2004)
Bulletin on Peer Review. The intent of
the peer review policies is to ensure that
listings are based on the best scientific
and commercial data available. We
formally solicited the expert opinion of
three appropriate and independent
specialists regarding scientific or
commercial data or assumptions related
to the information considered for listing.
We conclude that these experts’ reviews
satisfy the requirements for ‘‘adequate
[prior] peer review’’ contained in the
Bulletin (sec. II.2.), as well as the
Services joint policy.
References
A complete list of the references used
in this final rule is available on the
internet at https://sero.nmfs.noaa.gov/pr/
sturgeon.htm.
Classification
National Environmental Policy Act
The 1982 amendments to the ESA, in
section 4(b)(1)(A), restrict the
information that may be considered
when assessing species for listing. Based
on this limitation of criteria for a listing
decision and the opinion in Pacific
Legal Foundation v. Andrus, 675 F. 2d
825 (6th Cir. 1981), we have concluded
that ESA listing actions are not subject
to the environmental assessment
requirements of the National
Environmental Policy Act (NEPA) (See
NOAA Administrative Order 216–6).
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Executive Order 12866, Regulatory
Flexibility Act and Paperwork
Reduction Act
As noted in the Conference Report on
the 1982 amendments to the ESA,
economic impacts cannot be considered
when assessing the status of a species.
Therefore, the economic analysis
requirements of the Regulatory
Flexibility Act are not applicable to the
listing process. In addition, this final
rule is exempt from review under
Executive Order 12866. This final rule
does not contain a collection-ofinformation requirement for the
purposes of the Paperwork Reduction
Act.
Executive Order 13132, Federalism
Executive Order 13132 requires
agencies to take into account any
federalism impacts of regulations under
development. It includes specific
consultation directives for situations
where a regulation will preempt state
law, or impose substantial direct
compliance costs on state and local
governments (unless required by
statute). Neither of those circumstances
is applicable to this final listing
determination. In keeping with the
intent of the Administration and
Congress to provide continuing and
meaningful dialogue on issues of mutual
state and Federal interest, the proposed
rule was provided to the relevant
agencies in each state in which the
Carolina and South Atlantic DPSs occur,
and these agencies were invited to
comment. Their comments were
addressed with other comments in the
‘‘Public Comments’’ section.
Executive Order 12898, Environmental
Justice
Executive Order 12898 requires that
Federal actions address environmental
justice in the decision-making process.
In particular, the environmental effects
of the actions should not have a
disproportionate effect on minority and
low-income communities. The listing
determination is not expected to have a
disproportionately high effect on
minority populations or low-income
populations.
Coastal Zone Management Act (16
U.S.C. 1451 et seq.)
Section 307(c)(1) of the Federal
Coastal Zone Management Act (CZMA)
of 1972 requires that all Federal
activities that affect any land or water
use or natural resource of the coastal
zone be consistent with the enforceable
policies of approved state coastal zone
management programs to the maximum
extent practicable. We have determined
that this action is consistent to the
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maximum extent practicable with the
enforceable policies of approved CZMA
Programs of each of the states within the
range of the two DPSs. Letters
documenting NMFS’ proposed
determination, along with the proposed
rule, were sent to the coastal zone
management program offices in each
affected state. A list of the specific state
contacts and a copy of the letters are
available upon request.
The North Carolina Department of
Coastal Management (NCDCM) objected
to our consistency determination and
identified the following three relevant
enforceable policies of their approved
management program with which they
believed listing Atlantic sturgeon as
endangered would be inconsistent: (1)
15A NCAC07H.0203 Management
Ojective of the Estuarine and Ocean
System; (2) 15A NCAC 07H .0206
Estuarine Waters; and, (3) 15A NCAC
07H .0207 Public Trust Areas. NCDCM
believes listing Atlantic sturgeon as
proposed would be inconsistent with
their objective of managing Atlantic
sturgeon resources in a manner that
would perpetuate the biological and
economic values of marine resources
within North Carolina’s coastal zone
because: (1) Sampling programs for
many fish species would have to be
immediately terminated, and (2) North
Carolina’s fishing industry would be
affected since sampling and/or bycatch
of Atlantic sturgeon would constitute
unpermitted take. NCDCM expressed
concern that during the time it takes to
obtain ESA permits for research and
bycatch in fisheries, the ability to
monitor population trends and comply
with data collection requirements of
ASMFC’s FMPs will be curtailed.
NCDCM is concerned about prohibitions
on gear and other hardships on North
Carolina fisheries, as well as
administrative burdens on the state,
including having to provide observer
coverage. NCDCM stated that a finding
of concurrence with our consistency
determination could be made if: (1) The
listing was delayed until permits for
take have been obtained for research
and fisheries bycatch, and (2)
coordination takes place with NCDMF
and NCWRC to implement a data
collection program to further examine
the listing determination for Atlantic
sturgeon.
Per 15 CFR 930.43(d) of the
regulations implementing the CZMA, a
Federal agency shall not proceed with
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the activity over the State agency’s
objection unless: (1) The Federal agency
has concluded that under the
‘‘consistent to the maximum extent
practicable’’ standard described in 15
CFR 930.32, consistency with the
enforceable policies of the management
program is prohibited by existing law
applicable to the Federal agency, and
the Federal agency has clearly
described, in writing, to the State
agency the legal impediments to full
consistency (See 15 CFR 930.32(a) and
930.39(a)); or, (2) the Federal agency has
concluded that its proposed action is
fully consistent with the enforceable
policies of the management program,
though the State agency objects. As we
discussed in our letter to NCDCM
responding to their objection, section
4(b)(1)(A) of the ESA and 50 CFR
424.11(b) of the implementing
regulations require that listing
determinations be made solely on the
basis of the best scientific and
commercial data available to us and
without reference to possible economic
or other impacts of such a
determination. In addition, sections
4(b)(3)(B) and 4(b)(6)(A) of the ESA
establish mandatory deadlines under
the ESA for determining whether listing
of the species is warranted, and for
associated rules. Those deadlines were
triggered when NMFS received the
listing petition from the NRDC.
Therefore, per 15 CFR 930.43(d)(1), we
are prohibited from considering the
potential consequences, such as
permitting requirements, increased
regulatory responsibilities, and
hardships on fisheries (e.g., gear
restrictions), in our listing
determination, and we cannot enter into
a partnership with NCDMF and NCWRC
in lieu of listing Atlantic sturgeon.
However, we believe these partnerships,
such as the NMFS-funded section 6
project with NCDMF, North Carolina
State University, South Carolina
Department of Natural Resources, and
the University of Georgia looking at
movements of Atlantic and shortnose
sturgeon, will play a crucial role in
working toward conservation and
recovery of the species. Further, as
discussed in this final rule and in our
letter to NCDCM, NMFS is taking steps
that should minimize the potential
impacts to the state of North Carolina’s
fishery sampling programs and fishing
industry raised by NCDCM. For
example, NMFS contacted known
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5981
sturgeon researchers, at the time of
publication of the proposed rule,
requesting information on planned
research activities to facilitate
development of an expedited permitting
process. We also informed NCDCM, and
other North Carolina agencies, of the
expedited process during a conference
call in March 2011. Further, section
10(a)(1)(B) of the ESA allows NMFS to
issue permits authorizing incidental
take of listed species during the course
of otherwise lawful activities, such as
state fishery survey and sampling
programs targeting species other than
Atlantic sturgeon. Section 7
consultations required for any federallyauthorized fisheries that take Atlantic
sturgeon as bycatch would authorize
such incidental take after ensuring the
fishing activity would not jeopardize
sturgeon. Based on these factors, we
concluded pursuant to 15 CFR
930.43(d)(2) that this listing rule is
consistent with the State’s enforceable
policies listed above that provide for
managing the Atlantic sturgeon
resources in a manner that would
perpetuate the biological and economic
values of marine resources within North
Carolina’s coastal zone.
List of Subjects in 50 CFR Part 224
Administrative practice and
procedure, Endangered and threatened
species, Exports, Imports, Reporting and
recordkeeping requirements,
Transportation.
Dated: January 24, 2012.
Alan D. Risenhoover,
Acting Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
For the reasons set out in the
preamble, 50 CFR part 224 is amended
as follows:
PART 224—ENDANGERED MARINE
AND ANADROMOUS SPECIES
1. The authority citation for part 224
continues to read as follows:
■
Authority: 16 U.S.C. 1531–1543 and 16
U.S.C. 1361 et seq.
2. In § 224.101 the table in paragraph
(a) is amended by adding entries for
Atlantic Sturgeon-Carolina DPS and
Atlantic Sturgeon-South Atlantic DPS at
the end of the table to read as follows:
■
§ 224.101 Enumeration of endangered
marine and anadromous species.
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Species
Citation(s)
for listing
determination(s)
Where listed
Common name
Scientific name
*
Atlantic Sturgeon—
Carolina DPS.
*
Acipenser
oxyrinchus
oxyrinchus.
Atlantic Sturgeon—
South Atlantic
DPS.
Acipenser
oxyrinchus
oxyrinchus.
*
*
*
*
*
*
*
The Carolina DPS includes all Atlantic sturgeon that spawn
or are spawned in the watersheds (including all rivers and
tributaries) from Albemarle Sound southward along the
southern Virginia, North Carolina, and South Carolina
coastal areas to Charleston Harbor. The marine range of
Atlantic sturgeon from the Carolina DPS extends from the
Hamilton Inlet, Labrador, Canada, to Cape Canaveral,
Florida. The Carolina DPS also includes Atlantic sturgeon
held in captivity (e.g., aquaria, hatcheries, and scientific
institutions) and which are identified as fish belonging to
the Carolina DPS based on genetics analyses, previously
applied tags, previously applied marks, or documentation
to verify that the fish originated from (hatched in) a river
within the range of the Carolina DPS, or is the progeny of
any fish that originated from a river within the range of
the Carolina DPS.
The South Atlantic DPS includes all Atlantic sturgeon that
spawn or are spawned in the watersheds (including all
rivers and tributaries) of the ACE (Ashepoo, Combahee,
and Edisto) Basin southward along the South Carolina,
Georgia, and Florida coastal areas to the St. Johns River,
Florida. The marine range of Atlantic sturgeon from the
South Atlantic DPS extends from the Hamilton Inlet, Labrador, Canada, to Cape Canaveral, Florida. The South
Atlantic DPS also includes Atlantic sturgeon held in captivity (e.g., aquaria, hatcheries, and scientific institutions)
and which are identified as fish belonging to the South
Atlantic DPS based on genetics analyses, previously applied tags, previously applied marks, or documentation to
verify that the fish originated from (hatched in) a river
within the range of the South Atlantic DPS, or is the progeny of any fish that originated from a river within the
range of the South Atlantic DPS.
*
[Insert FR page
number where
the document begins]; 2/6/12.
[Insert FR page
number where
the document begins]; 2/6/12.
*
[FR Doc. 2012–1950 Filed 2–3–12; 8:45 am]
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Citation(s) for
critical habitat
designation(s)
*
NA.
NA.
Agencies
[Federal Register Volume 77, Number 24 (Monday, February 6, 2012)]
[Rules and Regulations]
[Pages 5914-5982]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2012-1950]
[[Page 5913]]
Vol. 77
Monday,
No. 24
February 6, 2012
Part III
Department of Commerce
-----------------------------------------------------------------------
National Oceanic and Atmospheric Administration
-----------------------------------------------------------------------
50 CFR Part 224
Endangered and Threatened Wildlife and Plants; Final Listing
Determinations for Two Distinct Population Segments of Atlantic
Sturgeon (Acipenser oxyrinchus oxyrinchus) in the Southeast; Final Rule
Federal Register / Vol. 77, No. 24 / Monday, February 6, 2012 / Rules
and Regulations
[[Page 5914]]
-----------------------------------------------------------------------
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric Administration
50 CFR Part 224
[Docket No. 090219208-1762-02]
RIN 0648-XN50
Endangered and Threatened Wildlife and Plants; Final Listing
Determinations for Two Distinct Population Segments of Atlantic
Sturgeon (Acipenser oxyrinchus oxyrinchus) in the Southeast
AGENCY: National Marine Fisheries Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA), Commerce.
ACTION: Final rule.
-----------------------------------------------------------------------
SUMMARY: We, NMFS, issue a final determination to list the Carolina and
South Atlantic distinct population segments (DPSs) of Atlantic sturgeon
(Acipenser oxyrinchus oxyrinchus) as endangered under the Endangered
Species Act (ESA) of 1973, as amended. We have reviewed the status of
the species and conservation efforts being made to protect the species,
considered public and peer review comments, and we have made our
determination that the Carolina and South Atlantic DPSs are in danger
of extinction throughout their ranges, and should be listed as
endangered, based on the best available scientific and commercial data.
DATES: This final rule is effective April 6, 2012.
ADDRESSES: Assistant Regional Administrator for Protected Resources,
NMFS, Southeast Regional Office, 263 13th Avenue South, St. Petersburg,
FL 33701-5505.
FOR FURTHER INFORMATION CONTACT: Kelly Shotts, NMFS, Southeast Regional
Office (727) 824-5312 or Lisa Manning, NMFS, Office of Protected
Resources (301) 427-8466.
SUPPLEMENTARY INFORMATION:
Background
We first identified Atlantic sturgeon as a candidate species in
1991. On June 2, 1997, NMFS and U.S. Fish and Wildlife Service (USFWS;
collectively, the Services) received a petition from the Biodiversity
Legal Foundation requesting that we list Atlantic sturgeon in the
United States, where it continues to exist, as threatened or endangered
and designate critical habitat within a reasonable period of time
following the listing. A notice was published in the Federal Register
on October 17, 1997, stating that the Services had determined
substantial information existed indicating the petitioned action may be
warranted (62 FR 54018). In 1998, after completing a comprehensive
status review, the Services published a 12-month determination in the
Federal Register announcing that listing was not warranted at that time
(63 FR 50187; September 21, 1998). We retained Atlantic sturgeon on the
candidate species list (and subsequently transferred it to the Species
of Concern List (69 FR 19975; April 15, 2004)). Concurrently, the
Atlantic States Marine Fisheries Commission (ASMFC) completed Amendment
1 to the 1990 Atlantic Sturgeon Fishery Management Plan (FMP) that
imposed a 20- to 40-year moratorium on all Atlantic sturgeon fisheries
until the Atlantic Coast spawning stocks could be restored to a level
where 20 subsequent year classes of adult females were protected
(ASMFC, 1998). In 1999, pursuant to section 804(b) of the Atlantic
Coastal Fisheries Cooperative Management Act (ACFCMA) (16 U.S.C. 5101
et seq.), we followed this action by closing the Exclusive Economic
Zone (EEZ) to Atlantic sturgeon retention. In 2003, we sponsored a
workshop in Raleigh, North Carolina, with USFWS and ASMFC entitled,
``The Status and Management of Atlantic Sturgeon,'' to discuss the
status of sturgeon along the Atlantic Coast and determine what
obstacles, if any, were impeding their recovery. The workshop revealed
mixed results in regards to the status of Atlantic sturgeon riverine
populations, despite the coastwide fishing moratorium. Some populations
seemed to be recovering while others were declining. Bycatch and
habitat degradation were noted as possible causes for continued
population declines.
Based on the information gathered from the 2003 workshop on
Atlantic sturgeon, we decided that a new review of Atlantic sturgeon
status was needed to determine if listing as threatened or endangered
under the ESA was warranted. The Atlantic sturgeon status review team
(ASSRT), consisting of four NMFS, four USFWS, and three U.S. Geological
Survey (USGS) biologists prepared a draft status review report. The
draft report was then reviewed and supplemented by eight state and
regional experts who provided their individual expert opinions on the
scientific facts contained in the report and provided additional
information to ensure the report provided the best available data.
Lastly, the report was peer reviewed by six experts from academia. A
Notice of Availability of the final status review report was published
in the Federal Register on April 3, 2007 (72 FR 15865). On October 6,
2009, we received a petition from the Natural Resources Defense Council
to list Atlantic sturgeon as endangered under the ESA. As an
alternative, the petitioner requested that the species be delineated
and listed as the five DPSs described in the 2007 Atlantic sturgeon
status review report (ASSRT, 2007): Gulf of Maine, New York Bight,
Chesapeake Bay, Carolina, and South Atlantic DPSs, with the Gulf of
Maine and South Atlantic DPSs listed as threatened, and the remaining
three DPSs listed as endangered. The petitioner also requested that
critical habitat be designated for Atlantic sturgeon under the ESA. We
published a Notice of 90-Day Finding on January 6, 2010 (75 FR 838),
stating that the petition presented substantial scientific or
commercial information indicating that the petitioned actions may be
warranted. On October 6, 2010, we published a proposed rule (75 FR
61904) to list the Carolina and South Atlantic DPSs, the two DPSs that
spawn in the NMFS Southeast Region, as endangered. We originally
solicited written public comments via email, fax, and letter on the
proposed listing rule for 90 days and extended it for an additional 30
days by public request. We also accepted written and verbal comments at
two public hearings in Wilmington, North Carolina, and Atlanta,
Georgia, in December 2010. A separate proposed rule (75 FR 91872) was
published on October 6, 2010, for the three DPSs of Atlantic sturgeon
that spawn in the NMFS Northeast Region.
Listing Determinations Under the Endangered Species Act
We are responsible for determining whether Atlantic sturgeon are
threatened or endangered under the ESA (16 U.S.C. 1531 et seq.). To be
considered for listing under the ESA, a group of organisms must
constitute a ``species,'' which is defined in section 3 of the ESA to
include ``any subspecies of fish or wildlife or plants, and any
distinct population segment of any species of vertebrate fish or
wildlife which interbreeds when mature.'' On February 7, 1996, the
Services adopted a policy describing what constitutes a DPS of a
taxonomic species (61 FR 4722). The joint DPS policy identified two
elements that must be considered when identifying a DPS: (1) The
discreteness of the population segment in relation to the remainder of
the species (or subspecies) to which it belongs; and (2) the
significance of the population segment to the remainder of the species
(or subspecies) to which it
[[Page 5915]]
belongs. As stated in the joint DPS policy, Congress expressed its
expectation that the Services would exercise authority with regard to
DPSs sparingly and only when the biological evidence indicates such
action is warranted.
Section 3 of the ESA defines an endangered species as ``any species
which is in danger of extinction throughout all or a significant
portion of its range'' and a threatened species as one ``which is
likely to become an endangered species within the foreseeable future
throughout all or a significant portion of its range.'' The statute
requires us to determine whether any species is endangered or
threatened as a result of any one or a combination of the following
five factors: (A) The present or threatened destruction, modification,
or curtailment of its habitat or range; (B) overutilization for
commercial, recreational, scientific, or educational purposes; (C)
disease or predation; (D) the inadequacy of existing regulatory
mechanisms; or (E) other natural or manmade factors affecting its
continued existence (section 4(a)(1)(A)(E)). Section 4(b)(1)(A) of the
ESA requires us to make listing determinations based solely on the best
scientific and commercial data available after conducting a review of
the status of the species and after taking into account efforts being
made to protect the species. Accordingly, we have followed a stepwise
approach in making our listing determination for Atlantic sturgeon.
Considering biological evidence, such as the separation between river
populations during spawning and the possibility of multiple distinct
interbreeding Atlantic sturgeon populations, we evaluated whether
Atlantic sturgeon population segments met the DPS Policy criteria. We
then determined the status of each DPS (each ``species'') and
identified the factors and threats contributing to their status per
section 4(a)(1) of the ESA. Finally, we assessed efforts being made to
protect the species, determining if these efforts are adequate to
mitigate impacts and threats to the species' statuses. We evaluated
ongoing conservation efforts using the criteria outlined in the Policy
for Evaluating Conservation Efforts (PECE; 68 FR 15100; March 28, 2003)
to determine their certainties of implementation and effectiveness.
Finally, section 4(b)(1)(B) of the ESA requires us to give
consideration to species which: (1) Have been designated as requiring
protection from unrestricted commerce by any foreign nation or pursuant
to an international agreement; or (2) have been identified as in danger
of extinction, or likely to become so within the foreseeable future, by
any state agency or by any agency of a foreign nation.
Peer Review and Public Comments
In December 2004, the Office of Management and Budget (OMB) issued
a Final Information Quality Bulletin for Peer Review establishing
minimum standards for peer review. Similarly, a joint NMFS/FWS policy
(59 FR 34270; July 1, 1994) requires us to solicit independent expert
review from at least three qualified specialists. We solicited peer
review comments on the proposed listing rule from three peer reviewers,
two from academia and one from a Federal resource agency, with
expertise on Atlantic sturgeon. Written public comments were received
from 59 commenters and 7 commenters provided verbal comments at the
public hearings. Peer review comments are treated in the next section.
In the following sections of the document, the public comments are
categorized in the following areas: (1) The delineation of DPSs; (2)
abundance and trends; (3) differences between the proposed rule and the
conclusions in the 2007 and 1998 status reviews; (4) the need to list
Atlantic sturgeon under the ESA and consequences of listing; (5) the
analysis of threats (habitat modification and destruction,
overutilization, disease and predation, the inadequacy of regulatory
mechanisms, other natural and manmade factors); (6) recovery; (7)
critical habitat; and (8) adequacy of the public hearing. Many comments
were complex and had multiple inferences, and thus individual
statements are addressed in multiple comments and responses below.
Information and data provided by commenters supported or did not
conflict with our findings for the Carolina and South Atlantic DPSs.
Some information submitted by commenters as ``new'' information was
information already included and evaluated in our proposed listing rule
determination. Some commenters asked us to consider information, such
as increased compliance responsibilities and economic costs on agencies
and the public, that the ESA and its implementing regulations prohibit
us from considering in making listing determinations. Many commenters
stated that NMFS should postpone a listing determination until the
results of recent research are available, further research can be
undertaken, state and Federal moratoria on the harvest and possession
of Atlantic sturgeon have been in effect for the full planned duration,
and/or until non-listing alternatives (e.g., entering into multi-agency
partnerships and expanding existing programs) have been explored.
Because we were petitioned to list the Atlantic sturgeon, we cannot
delay an assessment of the status of Atlantic sturgeon. We were
required to evaluate the status of the species and the threats it is
currently facing and make a finding on whether the petitioned action
was warranted within 12 months, which resulted in our proposed listing
determination of endangered for the Carolina and South Atlantic DPSs of
Atlantic sturgeon. We believe the current body of information on the
declines of Atlantic sturgeon, the failure of their population numbers
to rebound despite harvest prohibitions, and the ongoing impacts from
bycatch, habitat modification, and the inadequacy of existing
regulatory mechanisms or protective efforts to control or mitigate for
these impacts, warrant listing the Carolina and South Atlantic DPSs as
endangered. The information provided in the peer review and public
comments did not provide a basis for revising our evaluation of the
status of Atlantic sturgeon, the nature and significance of the threats
and impacts they face, or our listing determinations. In the following
sections of the document, we summarize the comments pertaining to the
proposed listing rule for the Carolina and South Atlantic DPSs and
provide our responses to those comments. Complete copies of the peer
review comments, the written public comments, and transcripts of the
public hearings are available on the Internet at www.regulations.gov.
Peer Review Comments
In this section, we refer to peer reviewers 1, 2, and 3, which
correspond to the way the peer reviewers are identified on https://www.regulations.gov.
Comment 1 (definitions of endangered and threatened): Two of the
three peer reviewers disagreed, all or in part, with our proposed
listing of the Carolina and South Atlantic DPSs as endangered. Each
peer reviewer provided their own definitions of endangered and
threatened.
Peer reviewer 1 believed that a DPS warranted an endangered listing
only if no single historical spawning river within the DPS sustained an
abundant and regularly reproducing Atlantic sturgeon population. Peer
reviewer 1 stated that no substantive biological justification or new
evidence is presented in the proposed listing of the Carolina and South
Atlantic DPSs as endangered to change the conclusions presented in the
2007 status review, which concluded that the Carolina DPS should be
listed as threatened and made
[[Page 5916]]
no conclusion with regard to the South Atlantic DPS due to lack of
information to allow a full assessment of subpopulations within the
DPS. Peer reviewer 1 stated that an endangered listing would be
appropriate if no single historical spawning river within that DPS
appeared to sustain both a relatively abundant and simultaneously
regularly-reproducing Atlantic sturgeon population.
Based on the available information on abundance, reproduction, and
the presence of early life history stages, the reviewer stated that the
Carolina DPS comes closest to conforming to the standard of an
endangered species. The reviewer cited data from the proposed listing
rule that two of the original three major spawning populations (the
Roanoke and Santee-Cooper populations) in the Carolina DPS appear to
remain functional, and not particularly vulnerable to extinction. The
reviewer also stated the proposed listing of the South Atlantic DPS did
not appear to be supported by the best available scientific
information, since there is evidence of at least one viable,
reproducing, and increasing Atlantic sturgeon population in the South
Atlantic DPS, the Altamaha River population (Schueller and Peterson,
2006, 2010). The reviewer further cited both the Savannah River and the
ACE (Ashepoo, Combahee, and Edisto Rivers) Basin systems as appearing
to support reproducing Atlantic sturgeon populations, and stated
Atlantic sturgeon appear to be abundant in the ACE system. The reviewer
questioned whether the remaining South Atlantic DPS river populations
in the smaller and less well-studied Ogeechee and Satilla rivers
together constitute a significant portion of the species' range over
which extinction is probable in order to justify an ``endangered''
designation for the entire DPS. The reviewer noted that the 2007 status
review report deferred from such a designation and that it appears the
South Atlantic DPS does not closely conform to the standard of being
endangered. Based on the available scientific evidence concerning
population size and reproduction in the historically most important
populations, the resilience of sturgeons to extirpation, and their
capacity for re-population from small effective population size, the
reviewer believed the appropriate ESA designation for both DPSs would
seem to be threatened. The reviewer suggested that the threatened
status would provide protection for the species from direct take of any
kind and a basis for habitat restoration, while providing greater
flexibility for scientific sampling, tissue analyses, and experimental
manipulation than would endangered status. The reviewer stated the
downside is that threatened status would provide a lower level of legal
leverage relative to the larger industrial impacts, e.g., dams and
bycatch, either of which may represent an insurmountable impasse to
sturgeon recovery. The reviewer offered that under existing direct
harvest prohibitions, threatened status has worked effectively for Gulf
sturgeon recovery in rivers where dams and bycatch are not significant
issues. It has not worked effectively where dams and bycatch are
significant issues (e.g., the Pearl, Pascagoula, and Apalachicola
rivers), although none of those populations seem in danger of
extinction.
Peer reviewer 2 stated that implicit in the definition of
``endangered'' is that the species must be on a significant downward
trend, or at least there is cause to believe that such a trend is
happening now, or will happen soon, and concluded that is not the case
on the Altamaha River in Georgia. However, this reviewer also commented
that every single Atlantic sturgeon population has been decimated by
overfishing and habitat degradation and that we have very little
quantified evidence that the species as a whole has recovered, despite
14 years of the protection afforded under the current moratorium on
harvest and possession. Peer reviewer 2 recommended that a
``threatened'' listing would seem appropriate for almost every Atlantic
coast river, including the St. Marys, Satilla, Ogeechee, and Savannah
Rivers in Georgia, with the Altamaha being the one exception, and an
endangered listing would be difficult to support.
Response: We must rely on the definition of ``endangered'' and
``threatened'' species provided in section 3 of the ESA, the
implementing regulations, and case law in applying the definitions to
marine and anadromous species. Section 3 of the ESA defines an
endangered species as one that is in danger of extinction throughout
all or a significant portion of its range, and a threatened species as
one that is likely to become endangered within the foreseeable future.
Recent case law (In Re Polar Bear Endangered Species Act Listing and
Sec. 4(d) Rule Litigation, D.D.C. WL 2601604 (June 30, 2011 Order);
748 F.Supp.2d 19 (D.D.C. 2010)) regarding USFWS's listing of the polar
bear as threatened provides a thorough discussion of the ESA's
definitions and the Services' broad discretion to determine on a case
by case basis whether a species is in danger of extinction. Upon
listing the polar bear as threatened, USFWS's rule was challenged by a
number of parties who claimed that the polar bear was in danger of
extinction and should have been listed as endangered, and by others who
conversely argued that the bear did not warrant listing even as
threatened. The Court determined that neither the ESA nor its
legislative history compels the interpretation of ``endangered'' as a
species being in ``imminent'' risk of extinction, finding instead that
the phrase ``in danger of extinction'' is ambiguous. The Court held
that there is a temporal distinction between endangered and threatened
species in terms of the proximity of the ``danger'' of extinction,
noting that the definition of ``endangered species'' is phrased in the
present tense, whereas a threatened species is ``likely to become'' so
in the future. Thus, in the context of the ESA, the Services interpret
an ``endangered species'' to be one that is presently at risk of
extinction. A ``threatened species,'' on the other hand, is not
currently at risk of extinction, but is likely to become so. In other
words, a key statutory difference between a threatened and endangered
species is the timing of when a species may be in danger of extinction,
either now (endangered) or in the foreseeable future (threatened). The
Court concluded, however, that the distinction is not based ``solely
and unambiguously'' on the imminence of the species' anticipated
extinction,'' and that Congress delegated responsibility to the
Services to determine whether a species is presently `in danger of
extinction' in light of the five ESA section 4(a)(1) factors and the
best available science for that species. The Court ruled that although
imminence of harm is clearly one factor that the Services weigh in
their decision-making process, it is not necessarily a limiting factor.
In many cases, the Services might appropriately find that the imminence
of a particular threat is the dispositive factor that warrants listing
a species as `threatened' rather than `endangered,' or vice versa. The
Services have broad discretion to decide that other factors outweigh
the imminence of the threat. In conclusion, the Court confirmed that
the Services have flexibility to determine ``endangerment'' on a case-
by-case basis. Congress did not intend to make any single factor
controlling when drawing the distinction between endangered and
threatened species, nor did it seek to limit the applicability of the
endangered
[[Page 5917]]
category to only those species facing imminent extinction.
Thus, contrary to the peer reviewers' comments, there is no per se
requirement that a species be experiencing current or imminent
significant downward trends, or that there are no single historical
spawning river populations within the DPSs that are relatively abundant
and simultaneously regularly-reproducing, in order to be listed as
endangered (we discuss the status and data on the Altamaha River
population in more detail in Comment 2 below). Our determination of
endangerment for the Carolina and South Atlantic DPSs is based on the
exercise of our expert professional judgment on the basis of the best
available information for each DPS. In addition, we agree with the
USFWS' judgment, discussed in its supplemental explanation filed in the
polar bear litigation, that to be listed as endangered does not require
that extinction be certain or probable, and that it is possible for a
species validly listed as ``endangered'' to actually persist
indefinitely.
We determined that the Carolina and South Atlantic DPSs of Atlantic
sturgeon are currently in danger of extinction throughout their ranges,
on the basis of precipitous declines to population sizes, the
protracted period in which sturgeon populations have been depressed,
the limited amount of current spawning, and the impacts and threats
that have and will continue to prevent population recovery. Populations
of Atlantic sturgeon declined precipitously decades ago due to directed
commercial fishing. The failure of Atlantic sturgeon numbers within the
Carolina and South Atlantic DPSs to rebound even after the moratorium
on directed fishing was established in 1998 indicates that impacts and
threats from limits on habitat for spawning and development, habitat
alteration, and bycatch are responsible for the risk of extinction
faced by both DPSs. In addition, the persistence of these impacts and
threats points to the inadequacy of existing regulatory mechanisms to
address and reduce habitat alterations and bycatch. As described in the
proposed listing rule, the Carolina DPS is estimated to number less
than 3 percent of its historical population size; the South Atlantic
DPS is estimated to number less than 6 percent of its historical
population size, with all river populations except the Altamaha
estimated to be less than 1 percent of historical abundance. There are
an estimated 343 adults that spawn annually in the Altamaha River and
less than 300 adults spawning annually (total of both sexes) in the
river systems where spawning still occurs for each DPS (not all of the
river systems occupied by the two Southeast DPSs currently support
spawning, or effective spawning leading to recruitment).
In light of threats and impacts, the low population numbers of
every river population in the Carolina and South Atlantic DPSs suggests
that the DPSs are currently in danger of extinction throughout their
ranges; none of the populations are large or stable enough to alone or
in combination provide any level of certainty for continued existence
of either DPS, and thus, the peer reviewer's suggestion that these DPSs
may not be endangered rangewide or in a significant portion of their
ranges is erroneous. While the directed fishery that originally
drastically reduced the numbers of Atlantic sturgeon has been closed,
recovery of depleted populations is an inherently slow process for a
late-maturing species such as Atlantic sturgeon, and they continue to
face a variety of other threats that contribute to their risk of
extinction. Their late age at maturity (5 to 19 years in the Southeast)
provides more opportunities for individual Atlantic sturgeon to be
removed from the population before reproducing. While a long life-span
also allows multiple opportunities to contribute to future generations,
it increases the timeframe over which exposure to the multitude of
threats facing the Carolina and South Atlantic DPS can occur.
Based on available information, we determined that to be viable,
the Carolina and South Atlantic DPSs require multiple stable riverine
populations, and we have added discussion to the final determination to
better explain our reasoning. The importance of having multiple stable
riverine spawning populations within each DPS and the need to maintain
suitable habitat to support the various life functions (spawning,
feeding, growth) of Atlantic sturgeon is best understood by looking at
the concept of metapopulations. Each DPS, made up of multiple river
populations, is analogous to a metapopulation, which is a ``population
of populations'' (Levins, 1969), a group of spatially separated
populations of the same species which interact at some level.
Separation into metapopulations is expected by sturgeon and other
anadromous fishes. While recolonization of northern rivers following
post-Pleistocene deglaciation likely occurred following a stepping-
stone sequential model (Waldman et al., 2002), genetic analyses reveal
that currently, there are very low rates of exchange between river
populations. The amount and effectiveness of movement separates a
metapopulation from a single large, patchy population. Low rates of
connectivity through dispersal, with little to no effective movement,
allow individual populations to remain distinct as the rate of
migration between local populations is low enough not to have an impact
on local dynamics or evolutionary lineages and distinguishes a
metapopulation from a patchy population (Harrison 1994).
Metapopulation persistence depends on the balance of extinction and
colonization in a static environment (Hanski 1996). If habitat remains
suitable following local extirpation, recolonization via immigrants
into now-empty habitat may replace at least some of those losses
(Thomas, 1994). However, if the cause of extinction is a deterministic
population response to unsuitable conditions (e.g., lack of suitable
spawning habitat, poor water quality, or disturbance of substrates
through repeated dredging), the local habitat is likely to remain
unsuitable after extinction and be unavailable for effective
recolonization (Thomas, 1994). Therefore, recolonization is dependent
upon both immigration from adjacent, healthy populations and habitat
suitability. Because the DPSs are groups of populations, the stability,
viability, and persistence of individual populations affects the
persistence and viability of the larger DPS. The loss of any population
within a DPS will result in: (1) A long-term gap in the range of the
DPS that is unlikely to be recolonized, or recolonized only very
slowly; (2) loss of reproducing individuals; (3) loss of genetic
biodiversity; (4) potential loss of unique haplotypes; (5) potential
loss of adaptive traits; and (6) reduction in total number. The loss of
a population will negatively impact the persistence and viability of
the DPS as a whole as fewer than two individuals per generation
currently spawn outside their natal rivers (Wirgin et al., 2000; King
et al., 2001; Waldman et al., 2002).
The persistence of individual populations, and in turn the DPS,
depends on successful spawning and rearing within the freshwater
habitat, the immigration into marine habitats to grow, and then the
return of adults to natal rivers to spawn. Information on Atlantic
sturgeon spawning within the Carolina and South Carolina DPSs is
limited. In the proposed listing rule, we presumed spawning was
occurring if young-of-the-year (YOY) were observed or mature adults
were present in freshwater portions of the system.
[[Page 5918]]
Within the Carolina DPS, we concluded that spawning is occurring, or
occurred in the recent past, in the following rivers based on these
data:
1. Roanoke River--collection of 15 YOY (1997-1998); single YOY
(2005).
2. Tar and Neuse Rivers--one YOY (2005).
3. Cape Fear--upstream migration of adults in the fall, carcass of
ripe female upstream in mid-September.
4. Winyah Bay--running ripe male in Great Pee Dee River (2003).
Within the South Atlantic DPS, we concluded that spawning is
occurring, or has occurred in the recent past, in the following rivers
based on these data:
1. ACE Basin--1,331 YOY (1994-2001); gravid female and running ripe
male in the Edisto (1997); 39 spawning adults (1998).
2. Savannah River--22 YOY (1999-2006); running ripe male (1997).
3. Ogeechee River--age-1 captures, but high inter-annual
variability (1991-1998); 17 YOY (2003); 9 YOY (2004).
4. Altamaha River--74 captured/308 estimated spawning adults
(2004); 139 captured/378 estimated spawning adults (2005).
5. Satilla River--4 YOY and spawning adults (1995-1996).
These data indicate that spawning occurs within the Carolina and
South Atlantic DPSs; they do not indicate the frequency of annual
spawning events or the degree to which spawning in these systems leads
to population growth, persistence, or viability. The extent and
effectiveness of spawning events is unknown and likely precarious in
many rivers, given ongoing threats that limit population size and
spawning success, such as water quality and restricted access to
upstream spawning areas (75 FR 61904). Peer reviewer 1 stated that data
from the proposed listing rule indicate the spawning populations in the
Santee-Cooper system appear to remain functional and not particularly
vulnerable to extinction; however, in the proposed listing rule, we
noted our determination that spawning may occur in the Santee and/or
the Cooper Rivers, but it may not result in successful recruitment.
Lack of access to historical spawning habitat due to dams restricts
spawning to areas just below the dam. The proximity of these spawning
areas to salt water may result in very high mortality to any larvae
spawned in those systems.
In addition to spawning success, it is difficult to quantify
spawning potential within the two DPSs, given the lack of population
estimates. Currently, the number of Atlantic sturgeon in the Carolina
DPS is estimated to be 3 percent of historical population size and the
South Atlantic DPS is estimated to be 1 percent of historical
population size, with the exception of the Altamaha River population,
estimated to be at 6 percent of historical population size. Although
the largest impact that caused the precipitous decline of the species
has been curtailed (directed fishing), the population size has remained
relatively constant at these greatly reduced levels for approximately
100 years.
In response to comments about divergence from the status review
report's listing conclusions for the Carolina and South Atlantic DPSs,
NMFS' Protected Resources Divisions have the responsibility to make
listing recommendations to the Assistant Administrator. Status review
reports are an important part of the information base for such
recommendations, but NMFS must independently review the information in
status review reports and apply the ESA's listing determination
requirements in accordance with regulations, case law, and agency
guidance. The Atlantic Sturgeon Status Review Report states that
``risks of extinction assessments are performed to help summarize the
status of the species, and do not represent a decision by the Status
Review Team on whether the species should be proposed for listing as
endangered or threatened under the ESA'' (page 106; ASSRT, 2007).
Subsequent to the status review report, we conducted a comprehensive
assessment of the combined impact of the five ESA section 4(a)(1)
factors across the Carolina and South Atlantic DPSs in classifying
extinction risk for each DPS. We focused on evaluating whether the DPSs
are presently in danger of extinction or the danger of extinction is
likely to develop in the future. In our proposed rules to list 5 DPSs
of Atlantic sturgeon, we determined that each DPS was at greater risk
of extinction than determined by the 2007 ASSRT. While the ASSRT did
discuss and consider how multiple threats might act in concert on a
given subpopulation, they ultimately classified extinction risk using
the highest single threat score on an individual population within a
DPS, or within what they considered to be a significant portion of a
DPS's range (pages 108-109; ASSRT, 2007). We evaluated the overall
stability and viability of the DPSs as a whole based on the combined
statuses of the component river populations and the impacts of threats
and impacts across the DPS, when determining extinction risk of each
DPS, because, as discussed above, the Carolina and South Atlantic DPSs
require multiple stable river populations. In addition, because of the
lapse in time between the development of the status review report
(ASSRT, 2007) and the publication of the proposed listing rule (75 FR
61904, October 6, 2010), new information on bycatch (ASMFC, 2007) and
water quality (USEPA, 2008), as well as climate change (IPCC, 2008) and
drought (e.g., USGS, 2007), became available to us, and we incorporated
this information into our listing determinations.
Since publication of the proposed rules, a Federal District Court
has thoroughly reviewed and considered the distinction between the
definitions of threatened and endangered species in the ESA, explained
by the USFWS in litigation challenging their determination to list the
polar bear as threatened and not endangered, as discussed above (In re.
Polar Bear Endangered Species Act Litigation). Prompted by this
decision and the comments received by the Services requesting further
explanation of the divergence of our proposed listing statuses and the
conclusions of the ASSRT, we have reviewed our determinations and
concluded that all the proposed listings of specific DPSs as
``threatened species'' or ``endangered species,'' respectively, satisfy
the requirements of the relevant ESA definition. Thus, we have not
changed these classifications in the final rules. We found that the
Carolina and South Atlantic DPSs are presently in danger of extinction,
and thus, listing them as endangered is warranted.
As discussed above, because a DPS is a group of populations (a
metapopulation), the stability, viability, and persistence of
individual populations affects the persistence and viability of the
larger DPS. The persistence of individual populations, and in turn the
DPS, depends on successful spawning and rearing within the freshwater
habitat, the immigration into marine habitats to grow, and then the
return of adults to natal rivers to spawn. While the directed fishery
that originally drastically reduced the numbers of Atlantic sturgeon
has been closed, modification and curtailment of Atlantic sturgeon
habitat resulting from dams, dredging, and degraded water quality are
inhibiting spawning and population rebounding throughout both DPSs, and
contributing to their endangered statuses. Existing water allocation
issues will likely be compounded by human population growth and
potentially by climate change as well. Climate change is predicted to
elevate water temperatures and exacerbate nutrient-loading,
[[Page 5919]]
pollution inputs, and lower dissolved oxygen (DO), all of which are
currently negatively impacting the Carolina and South Atlantic DPSs.
Continued overutilization of Atlantic sturgeon from bycatch in multiple
commercial fisheries in both their marine and freshwater habitats is
another ongoing impact to the Carolina and South Atlantic DPSs that is
contributing to their endangered status. Atlantic sturgeon taken as
bycatch may suffer immediate mortality. In addition, stress or injury
to Atlantic sturgeon taken as bycatch but released alive may result in
increased susceptibility to other threats, such as poor water quality
(e.g., exposure to toxins and low DO). This may result in reduced
ability to perform major life functions, such as foraging and spawning,
or may even result in post-capture mortality. Several of the river
populations in the South Atlantic DPS (e.g., the Ogeechee and the
Satilla) are stressed to the degree that any level of bycatch could
have an adverse impact on the status of the DPS (ASSRT, 2007).
The Carolina and South Atlantic DPSs are in danger of extinction
now, due to precipitous declines from historical abundances to
population sizes that are low and potentially unstable throughout the
DPSs. As discussed above, both DPSs exhibit sporadic spawning with
uncertain effectiveness. Population rebuilding and recovery in both
DPSs is being inhibited by impacts due to habitat curtailment and
degradation, and due to capture as bycatch in commercial fisheries. The
current low levels of abundance noted previously in combination with
the high degree of threat to the two Southeast DPSs put them in danger
of extinction throughout their ranges; none of the populations making
up the DPSs are large or stable enough to provide any level of
certainty for continued existence of either DPS.
Regarding the conclusion that the Carolina and South Atlantic DPSs
should be listed as threatened, peer reviewer 1 incorrectly stated that
listing as threatened provides protection from direct take of any kind.
The ESA's prohibition against take contained in section 9 only applies
to endangered species, unless a section 4(d) rule is in place to extend
the take prohibition to a threatened species. If we determine that the
Carolina and South Carolina DPSs meet the ESA's definition of
endangered, then we cannot list the species as threatened for the
purposes of providing flexibility for scientific sampling, tissue
analyses, and experimental manipulation. We also cannot list the DPSs
as endangered to obtain legal leverage relative to the larger
industrial impacts, e.g., dams and bycatch, as suggested by the
reviewer. Rather, we must make our listing determination based on
application of the statutory factors.
Comment 2 (new information on Altamaha River population): Peer
reviewer 2 presented data on the estimated abundance of age-1, river
resident Atlantic sturgeon in the Altamaha River from 2004 to 2010,
which showed large estimated increases in this age group in 2009 and
2010. The peer reviewer also stated that he and other researchers are
beginning to detect slower growth in age-1 Atlantic sturgeon in the
Altamaha and he attributed this to density-dependent factors that are
beginning to limit available resources. The reviewer stated that a few
more years of data are needed to determine if the increasing trend is
real, but none of the other variables tested (e.g., river flows or
temperature) explain the trend. The peer reviewer attributed the
apparent increases in juveniles in the Altamaha to the moratorium on
the harvest of adults. The peer reviewer stated that data are not
available to determine whether this trend is occurring in other
spawning populations. The reviewer stated that catch per unit effort
(CPUE) data are worthless without calibration or validation and we do
not have historical abundance data to know what abundance should be on
any of the river systems, though there is general agreement that
populations are a fraction (less than 1 to 10 percent) of historical
abundance. The reviewer recommended that long-term monitoring of
recruitment using mark-recapture of age-1 juveniles be implemented on
key river systems.
Response: We are encouraged by the apparent increases in juvenile
Atlantic sturgeon estimated by the peer reviewer's research in the
Altamaha River and appreciate the contribution of this information for
our consideration in our listing determination. We revised the relevant
discussion in the text from the proposed listing rule to include this
information. We agree that additional years of data are necessary to
confirm this trend in the Altamaha and that we cannot determine whether
similar trends may be occurring in other river populations. This
information is consistent with information we provided in the proposed
listing rule, which refers to the Altamaha River as having a larger and
healthier Atlantic sturgeon population than any other river in the
Southeast. The proposed listing rule also stated that juvenile Atlantic
sturgeon from the Altamaha are relatively more abundant in comparison
to other rivers in the region.
Peer reviewer 2 noted that density-dependent factors may be
starting to limit available resources. We are interested in working
with the reviewer and other researchers to determine whether habitat
modification, which we describe in detail in the proposed listing rule,
is a contributing factor to the limitation of resources in addition to
the increase in numbers of juvenile Atlantic sturgeon utilizing the
resources. While water quality in the Altamaha River is good at this
time, the drainage basin is dominated by silviculture and agriculture,
with two paper mills and over two dozen other industries or
municipalities discharging effluent into the river. Nitrogen and
phosphorus concentrations are increasing, and eutrophication and loss
of thermal refugia are growing concerns for this and other rivers in
the South Atlantic DPS. The Altamaha is one of the rivers with current
and pending water allocation issues. We are currently funding a project
through the ESA section 6 program to map habitats in four Georgia
rivers, including the Altamaha, and this may be a valuable step in
answering this question.
We agree that CPUE data should be used in the proper context and
that historical abundance data, other than data from commercial
fisheries in the late 19th century, are not available. However, as
required by section 4(b)(1)(A) of the ESA, we must make our listing
determination based on the best scientific and commercial data
available. When only CPUE and other fishery-dependent data were
available to us, we clarified and acknowledged the constraints of the
data, and we conclude that we used them in a valid manner. This is
further addressed in our responses to several public comments on
specific sections of the proposed listing rule (e.g., comments 19, 23,
24, 25, and 29).
Comment 3 (import of the 2003 workshop): Peer reviewer 1 stated the
proposed listing rule appeared to dismiss any evidence of an increase
in Carolina DPS populations of Atlantic sturgeon, citing a statement in
the proposed listing rule (page 61904-61905) that ``the [NMFS-sponsored
2003] workshop revealed mixed results in regards to the status of
Atlantic sturgeon populations, despite the coast-wide fishing
moratorium. Some populations seemed to be recovering while others were
declining.'' The reviewer stated that at the time of the 2003 workshop,
the moratorium on direct harvest and possession of Atlantic sturgeon
had only been in effect for 4 years and this was not sufficient time
for populations to increase in response
[[Page 5920]]
to the protective measures. The reviewer concluded the observations
available at the time of the 2003 workshop do not provide a scientific
basis for listing the Carolina DPS as endangered.
Response: The information we evaluated in making our proposed
listing determination of endangered for the Carolina and South Atlantic
DPSs of Atlantic sturgeon was not confined to the results of the 2003
workshop. As stated in the proposed listing rule, the information
gathered at the 2003 workshop, including the equivocal evidence that
some populations appeared to be recovering while others were declining,
prompted us to complete a new review of Atlantic sturgeon status, which
was published in 2007. Since the ASSRT's completion of its status
review, we obtained and evaluated additional information on threats to
Atlantic sturgeon (see our response to comment 1). Our evaluation of
this information indicates that the moratorium on directed fisheries
has not and will not be sufficient to address the impacts that are
preventing sturgeon populations from recovering (including lack of
access to required habitat, and habitat quality issues). Section
4(b)(1)(A) of the ESA stipulates that listing decisions be made using
the best available scientific and commercial information, therefore we
used information from the 2007 status review report (which incorporated
information from the 2003 workshop) and new information in forming our
determination. Our responses to comments from the public further detail
our use of information available at the time of the proposed listing
rule, as well as our consideration of new information submitted during
the public comment period.
Comment 4 (viability of small Atlantic sturgeon populations): The
estimated 343 spawning adult Atlantic sturgeon in the Altamaha River
exceeds the number of spawning adults in the ``very viable'' Yellow
River Gulf sturgeon population, according to peer reviewer 1. Peer
reviewer 1 stated that information presented in Schueller and Peterson
(2010) suggests a very robust reproductive response to protection of
adult spawners under the Atlantic sturgeon moratorium. From these data,
the reviewer stated that it seems highly improbable that the Altamaha
River population is at risk of extinction and a listing of endangered
does not seem applicable to the Altamaha population within the South
Atlantic DPS. The reviewer stated that if the Altamaha population
follows the model of the Suwannee River Gulf sturgeon population after
harvest was banned, then overall population growth in the next decade
will be exponential until density-dependent population controls come
into play. Peer reviewer 1 also stated that the ``less than 300
spawning adults'' criterion in the proposed listing rule for
classifying a river population as vulnerable to extinction sets a
``very high, probably unrealistic, bar,'' and one not conforming to
scientific literature documenting sturgeon population recovery from
much smaller effective breeding population sizes (20-80 spawning
females, based on examples provided by the reviewer). Peer reviewer 1
stated that sturgeon species have the documented ability to establish/
re-establish viable populations over a short timeframe (10 to 20
years), starting from ``a few tens'' of spawning adults without
negative fitness impacts from low genetic diversity.
Response: As explained above, NMFS does not agree with peer
reviewer 1's premise that an endangered listing would only be
appropriate if no single historical spawning river within that DPS
appeared to sustain both a relatively abundant and simultaneously
regularly-reproducing Atlantic sturgeon population. We note that the
Yellow River population of Gulf sturgeon referred to as ``very viable''
by the reviewer is listed as threatened under the ESA. While the number
of spawning adult Atlantic sturgeon in the Altamaha River may be larger
than that of Gulf sturgeon in the Yellow River, the peer reviewer noted
that the Yellow River population is one that has rebounded since it was
listed. The Altamaha River population of Atlantic sturgeon is estimated
to be at only 6 percent of its historical abundance. While there is a
moratorium on harvest and possession of Atlantic sturgeon, the species
is not currently afforded the protections of section 9 of the ESA, nor
do they benefit from the consultation and permitting responsibilities
of ESA sections 7 and 10, that apply to the listed Gulf sturgeon.
Information provided by peer reviewers 1 and 2 indicated recent (2009-
2010) increases in the estimated number of juveniles in the Altamaha
River. We are encouraged by this and hope that the Altamaha River
population does exhibit exponential growth, as the Suwannee River Gulf
sturgeon population did following listing. However, our listing
determination is based on the best information currently available to
us, and we do not feel that the information provided on increases in
juvenile Atlantic sturgeon in the Altamaha River or the comparison to
Gulf sturgeon populations in the Suwannee and Yellow Rivers provides a
basis for revising our proposed determination that the South Atlantic
DPS be listed as endangered.
In response to the reviewer's comment that the ``less than 300
spawning adults criterion'' sets a ``very high, probably unrealistic,
bar'', we clarify that the 300 spawning adults per year was an estimate
of the relative sizes of Atlantic sturgeon river spawning populations,
based on the available information on the annual spawning adult
abundance measured in the Altamaha River (343 spawning adults) and the
fact that it is the largest population in the Southeast, combined with
qualitative and quantitative anecdotal information from the other river
systems. The 300 spawning adults per year estimate does not constitute
a criterion or a bar for listing and/or recovery as a general matter.
Rather, the estimate is evaluated in the specific context of the
Atlantic sturgeon river populations and the impacts and threats they
face. These populations likely have the capacity to recover, as the
reviewer suggested, if existing and future impacts and threats are
alleviated. The low estimated population numbers in each of the river
systems within the DPSs (1-6 percent of historical abundance), combined
with the ongoing impacts and threats from habitat modification and
bycatch, indicate that the populations are small and vulnerable, and
the DPSs they comprise are in danger of extinction.
Comment 5 (sturgeon ability to recolonize systems; genetic
exchange): Peer reviewer 1 stated that sturgeon species are resilient
and capable of repopulating an extirpated river, or colonizing a new
river, if habitat remains available, dams do not block spawning ground
access, water quality is satisfactory, and a competing sturgeon
population is not already established. A natal river population, well-
established over a long span of geological time and highly adapted to
its respective natal river, would not realize success in colonizing
another river already populated by a second population better adapted
to its respective natal river than a potential colonist. The reviewer
stated that the low rate of genetic exchange displayed among adjacent
sturgeon populations does not reflect the incapacity of the species to
colonize, but the competitive advantage held by a pre-established natal
river population facing migrant individuals. The reviewer provided
examples of recolonization by Atlantic sturgeon in bays and rivers from
New England to Labrador and Newfoundland within a span of 10,000 years
following deglaciation. The
[[Page 5921]]
reviewer stated the northernmost and most genetically conservative
Atlantic sturgeon population re-colonized over 1,500 miles of coastline
within 40 generations (and probably much fewer) in addition to
undertaking a successful, essentially instantaneous, 3,300 mile
migration to colonize the Baltic Sea 1,200 years ago. The reviewer also
provided an example of Gulf sturgeon rebounding in Gulf Coast river
systems. Peer reviewer 1 stated that following state harvest
prohibitions on the Gulf sturgeon and its listing as threatened under
the ESA, some river populations have rebounded (the Yellow,
Choctawhatchee, and Suwannee river populations). The reviewer concluded
the logic in the proposed listing rule is not compelling that if one of
the DPSs were to be completely extirpated, it would remain so over a
long span of time. Peer reviewer 3 stated, in reference to the genetic
analyses showing fewer than two individuals per generation spawn
outside their natal rivers, that this reflects the average number of
individuals and noted it would be useful to compare this to straying
determined from tagging data.
Response: We agree with the peer reviewer's comment that the low
rate of genetic exchange displayed between Atlantic sturgeon river
populations may reflect the competitive advantage held by pre-
established natal river populations facing migrant individuals. We
revised the relevant discussion in the text from the proposed listing
rule to include this information. However, as stated in the proposed
listing rule, we do not expect Atlantic sturgeon that originate from
other river systems to recolonize extirpated systems and establish new
spawning populations, except perhaps over a long time frame (i.e., many
Atlantic sturgeon generations). Though the reviewer provided an example
of Atlantic sturgeon colonizing the Baltic Sea 1,200 years ago after a
single migration, other examples of recolonization provided took 40
generations (approximately 1,000 years, based on a 25-year generation
period) to 10,000 years, which is consistent with our statement in the
proposed listing rule. Further, recolonization occurred in the absence
of present-day human impacts, such as habitat modifications and
mechanized fishing.
We noted the reviewer's comment that sturgeon species are resilient
and capable of repopulating an extirpated river or colonizing a new
river if habitat remains available, dams do not block spawning ground
access, and water quality is satisfactory. As discussed extensively in
the proposed listing rule and in our responses to comments in this
document, Atlantic sturgeon in the Carolina and South Atlantic DPSs and
the rivers within their ranges are affected by habitat modification and
destruction, blocked access to spawning grounds, downstream habitat
impacts caused by dams, and water quality (and quantity) issues. Thus,
the commenter's stated conditions for expecting recolonization by
Atlantic sturgeon are not met. Atlantic sturgeon from the Carolina and
South Atlantic DPSs are also taken as bycatch in fisheries. Regarding
the statement that following state harvest prohibitions on the Gulf
sturgeon and its listing as threatened under the ESA, some river
populations have rebounded (the Yellow, Choctawhatchee, and Suwannee
river populations), the rivers that have rebounded have two factors in
common versus those which have not rebounded (e.g., the Apalachicola,
Pascagoula, and Pearl river populations): (1) No mainstem dam on the
natal river limiting Gulf sturgeon access to upriver spawning grounds
or YOY access to riverine feeding habitat; and, (2) no major commercial
fishery causing Gulf sturgeon bycatch mortality in the natal river,
natal river estuary, or adjacent marine waters. Assessing the impacts
of these two factors may be equally as important to sturgeon population
recovery as is protection from all other impacts, now that direct
harvest has been stopped. We agree with these comments by the peer
reviewer and also believe that these threats associated with dams,
habitat, water quality, and bycatch would hamper and slow
recolonization of extirpated river systems. One reviewer acknowledged
that rivers, watersheds, and coastal habitats inhabited by Atlantic
sturgeon have been drastically modified and impacted by human
activities (dammed, channelized, de-watered, diverted, dredged, mined,
sedimented, polluted, deforested, developed, populated by introduced
species, etc.) and that it would be remarkable to achieve recovery to
even 10-30 percent of the 1890 carrying capacity of individual sturgeon
rivers.
In reference to peer reviewer 3's suggestion about comparing the
degree of straying from tagging studies to the estimate of straying
from the genetic studies, we agree this could be a valuable exercise in
the future when we have the necessary information on river of origin
(based on genetic analyses) and the degree of straying (from tagging
and relocation studies). While the estimate of less than 2 individuals
spawning in rivers outside their natal system is a measure of
successful transfer of genetic information from a fish originating from
another system, the analysis suggested by the peer reviewer would
provide us with knowledge of how many fish actually stray into another
system and potentially attempt to spawn. This could also provide
insight into the comments by the first peer reviewer that lack of gene
flow between river populations is due to reduced success from
competition and not from lack of attempts at migrant spawning.
Comment 6 (issues with estimating sturgeon abundance): According to
one peer reviewer, targeted Atlantic sturgeon population studies in the
Roanoke River and Santee-Cooper system, as well as most other river
systems, have been limited in duration, intensity, and continuity such
that population estimates may be substantially underestimated. Peer
reviewer 1 noted that sturgeon species are cryptic fish found in deep,
mainstem rivers. They are rarely observed visually, not typically
sampled in many commercial river fisheries targeting other fish species
(with the exception of the shad gill net fishery), and are rarely
caught by recreational anglers. The reviewer stated that this
illustrates that presence and abundance of sturgeon cannot be based on
incidental catches from commercial fisheries or scientific sampling not
specifically targeting sturgeon. The reviewer stated that in the past,
sturgeon abundance has often been vastly underestimated until an
appropriate and dedicated reporting or sampling program was undertaken.
The reviewer recommended that only continuous, standardized mark-
recapture efforts spanning sufficient time (a minimum of 3 years, but
realistically greater than 5 years) can provide reliable preliminary
abundance estimates.
Response: The majority of the data presented in the proposed
listing rule came from studies targeting Atlantic sturgeon or from
fisheries that are known to have a high incidence of interaction with
Atlantic sturgeon (i.e., gillnet fisheries). As much as possible, we
clarified the data collection methods and constraints, and any
assumptions we made. This is also discussed in our response to comment
2. We have used the best available commercial and scientific
information to evaluate the status of the Carolina and South Atlantic
DPSs, but we agree with the reviewer that long-term, continuous,
standardized studies of Atlantic sturgeon abundance are needed.
Comment 7 (viable population sizes and sturgeon genetics): Peer
reviewer 1 stated the minimum viable population
[[Page 5922]]
sizes of several hundreds to several thousands of individuals advanced
in the literature are not particularly instructive with respect to
sturgeon species based on new genetic information (Kreiger et al.,
2006). The reviewer commented that sturgeon are polyploid and the
significance of polyploidy upon genetic diversity has just emerged.
Most fishes are diploid with 40-50 chromosomes, a number similar to
most vertebrates. However, all sturgeons are polyploid, having
approximately 120 chromosomes (tetraploid, 4N), 240 chromosomes
(octoploid, 8N) or more, including species with 12N or 16N ploidy.
Polyploidy allows for multiple alleles (not just two as in diploid
species) at a given gene locus, allowing for intra-individual genetic
variation (Kreiger et al. 2006). The reviewer suggested that this might
explain the high degree of plasticity displayed by sturgeon populations
and the documented ability of sturgeons to repopulate from very few
spawning adults without apparent inbreeding depression. He concluded
that until we gain a deeper understanding of the genetics of polyploidy
and the implications regarding sturgeon population dynamics, any
discussion of minimum viable population size for sturgeon populations
cannot be phrased in terms of what we know about inbreeding depression
in diploid mammal populations. Thus, the 50/500 rule of thumb cited in
the proposed listing rule may be an inappropriate criterion by which to
assess viability of sturgeon populations, and we do not know how few
polyploid sturgeons are too few to sustain a viable population.
Response: We appreciate the peer reviewer's input on the polyploid
nature of Atlantic sturgeon and how this genetic characteristic may
affect our evaluation of minimum viable population sizes in our listing
determination. We revised the relevant discussion in the text from the
proposed listing rule to include this information. As noted by the
reviewer, we need a deeper understanding of the genetics of polyploidy
and the implications regarding sturgeon population dynamics. We are not
sure how polyploidy in Atlantic sturgeon will affect their recovery,
but even if it allows the species to repopulate from relatively fewer
individuals without inbreeding depression, there is no assurance that
this will occur. Other polyploid Acipenser species have required
listing under the ESA, such as shortnose sturgeon (listed as endangered
in 1967), Gulf sturgeon (listed as threatened in 1991), and green
sturgeon (listed as threatened in 2006). In the case of the shortnose
sturgeon, recovery has not been achieved even though it has been
protected for almost 45 years. Further, the polyploid nature of
Atlantic sturgeon may further support the need for protection under the
ESA. Southern populations of Atlantic sturgeon exhibit high diversity
and many low frequency (and sometimes private) haplotypes (Grunwald et
al., 2008). Allendorf and Leary (1988) noted that in polyploid
cutthroat trout, alleles constituting the majority of the variation in
the species are found in only one or two local populations, but they
often occur at high frequencies in those populations. They concluded
preserving the genetic variation in cutthroat trout entails pre