Endangered and Threatened Wildlife; 12-Month Finding on a Petition To Identify and Delist a Saint John River Distinct Population Segment of Shortnose Sturgeon Under the Endangered Species Act, 65183-65194 [2015-27148]
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persistence and that should be further
evaluated in a status review.
The petitioners claim that hypoxia
(oxygen deficiency) has increased in
frequency, duration, and severity in
coastal waters and that this decreases
the abundance and diversity of benthic
macrofauna (citing CSIS 2011). They
also claim that the combination of
hypoxia and increased water
temperature would reduce the quality
and size of suitable habitat for aerobic
organisms whose suitable habitat is
restricted by water temperature and
claim that thorny skate is such a
species. While acknowledging that any
prediction of the effects of hypoxic
zones on thorny skates is speculative,
the petitioners state that any adverse
impact on the species or on the
abundance/distribution of its predators
or prey will severely hinder the species’
ability to recover. However, neither the
petitioners nor the information in our
files indicate that thorny skate are
impacted by hypoxia or that hypoxia
may be contributing significantly to
population declines in thorny skates to
the point where the species may be at
a risk of extinction. As such, we
conclude that the information presented
in the petition on the threat of hypoxia
does not provide substantial
information indicating that hypoxia may
be impacting thorny skate to a degree
that the petitioned action may be
warranted.
The petitioners state that the life
history characteristics of thorny skate
place the species at risk of adverse
effects resulting from natural stochastic
events. However, neither the petitioners
nor the information in our files indicate
that natural stochastic events are
causing detrimental effects to the
species or may be contributing
significantly to population declines in
thorny skates to the point where the
species may be at a risk of extinction.
As such, we conclude that the
information presented in the petition on
the threat of natural stochastic events
does not provide substantial
information indicating that such events
may be impacting or may, in the
foreseeable future, impact thorny skate
to a degree that the petitioned action
may be warranted. However, given all of
the information presented above on
other natural and manmade factors,
particularly the warming of oceans, the
information in the petition and in our
files does lead a reasonable person to
conclude that the petitioned action may
be warranted, and it is necessary to
consider the impacts from other natural
and manmade factors in a status review.
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Summary of ESA Section 4(a)(1)
Factors
provisions regarding the designation of
critical habitat.
We conclude that the petition
presents substantial scientific or
commercial information indicating that
a combination of three of the section
4(a)(1) factors (overutilization for
commercial, recreational, scientific, or
educational purposes; inadequate
existing regulatory mechanisms; and
other natural or manmade factors) may
be causing or contributing to an
increased risk of extinction for thorny
skate which needs to be further
evaluated in a review of the status of the
species.
Information Solicited
Petition Finding
After reviewing the information
contained in the petition, as well as
information readily available in our
files, and based on the above analysis,
we conclude the petition presents
substantial scientific information
indicating the petitioned action of
listing a Northwest Atlantic or United
States DPS of thorny skate as threatened
or endangered may be warranted.
Therefore, in accordance with section
4(b)(3)(B) of the ESA and NMFS’
implementing regulations (50 CFR
424.14(b)(2)), we will commence a
review of the status of the species.
During our status review, we will first
determine whether one of the
populations identified by the petitioners
meets the DPS policy criteria, and if so,
whether it is threatened or endangered
throughout all or a significant portion of
its range. We now initiate this review,
and thus, the Northwest Atlantic
population of the thorny skate is
considered to be a candidate species
(see 69 FR 19975; April 15, 2004). To
the maximum extent practicable, within
12 months of the receipt of the petition
(May 28, 2016), we will make a finding
as to whether listing either of the
populations identified by the petitioner
as DPSs as endangered or threatened is
warranted as required by section
4(b)(3)(B) of the ESA. If listing a DPS is
found to be warranted, we will publish
a proposed rule and solicit public
comments before developing and
publishing a final rule. The petitioners
request that we designate critical habitat
for thorny skates. ESA Section 4(a)(3)(A)
and its implementing regulations state
that, to the maximum extent prudent
and determinable, the Secretary shall,
concurrently with listing a species as
endangered or threatened, designate any
critical habitat for that species. If a
thorny skate population were to be
listed as a DPS, we would follow the
relevant statutory and regulatory
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To ensure that the status review is
based on the best available scientific
and commercial data, we are soliciting
information on the thorny skate.
Specifically, we solicit information in
the following areas: (1) Historical and
current distribution and abundance of
this species in the Northwest Atlantic;
(2) historical and current population
status and trends; (3) any current or
planned activities that may adversely
impact the species, especially as related
to the five factors specified in section
4(a)(1) of the ESA and listed above; (4)
ongoing efforts to protect and restore the
species and its habitat; and (5) genetic
data or other information related to
possible population structure of thorny
skate. We request that all information be
accompanied by: (1) Supporting
documentation such as maps,
bibliographic references, or reprints of
pertinent publications; and (2) the
submitter’s name, address, and any
association, institution, or business that
the person represents.
References Cited
A complete list of references is
available upon request (see ADDRESSES).
Authority
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Dated: October 16, 2015.
Samuel D. Rauch, III.
Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2015–27147 Filed 10–23–15; 8:45 am]
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DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
Administration
50 CFR Part 224
[Docket No. 150209121–5941–02]
RIN 0648–XD760
Endangered and Threatened Wildlife;
12-Month Finding on a Petition To
Identify and Delist a Saint John River
Distinct Population Segment of
Shortnose Sturgeon Under the
Endangered Species Act
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Department of Commerce.
AGENCY:
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ACTION:
Federal Register / Vol. 80, No. 206 / Monday, October 26, 2015 / Proposed Rules
Notice of 12-month petition
finding.
We, NMFS, announce a 12month finding on a petition to identify
and ‘‘delist’’ shortnose sturgeon
(Acipenser brevirostrum) within the
Saint John River in New Brunswick,
Canada under the Endangered Species
Act (ESA). The shortnose sturgeon is
currently listed as an endangered
species, at the species level, under the
ESA. Based on our review of the best
scientific and commercial data
available, we have determined that the
population of shortnose sturgeon from
the Saint John River does not qualify as
a distinct population segment.
Therefore, we did not consider the
petition further, and we do not propose
to delist this population.
DATES: This finding was made on
October 26, 2015.
ADDRESSES: Information used to make
this finding is available for public
inspection by appointment during
normal business hours at NMFS, Office
of Protected Resources, 1315 East-West
Highway, Silver Spring, MD 20910. The
petition and the list of the references
used in making this finding are also
available on the NMFS Web site at:
www.nmfs.noaa.gov/pr/species/fish/
shortnose-sturgeon.html.
FOR FURTHER INFORMATION CONTACT: Lisa
Manning, Office of Protected Resources,
301–427–8466; Stephania Bolden,
Southeast Regional Office, 727–824–
5312; Julie Crocker, Greater Atlantic
Regional Office, 978–282–8480.
SUPPLEMENTARY INFORMATION:
SUMMARY:
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Background
On September 24, 2014, we received
a petition from Dr. Michael J. Dadswell,
Dr. Matthew K. Litvak, and Mr. Jonathan
Barry regarding the population of
shortnose sturgeon (Acipenser
brevirostrum) native to the Saint John
River in New Brunswick, Canada. The
petition requests that we identify the
Saint John River population of
shortnose sturgeon as a distinct
population segment (DPS) and
contemporaneously ‘‘delist’’ this DPS by
removing it from the species-wide
listing under the Endangered Species
Act. On April 6, 2015, we published a
positive finding indicating that the
petitioned action may be warranted and
that we were initiating a status review
to consider the petition further (80 FR
18347).
The shortnose sturgeon was originally
listed as an endangered species
throughout its range by the U.S. Fish
and Wildlife Service (USFWS) on March
11, 1967, under the Endangered Species
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Preservation Act (ESPA, 32 FR 4001).
Shortnose sturgeon remained on the
endangered species list when the U.S.
Congress replaced the ESPA by enacting
the Endangered Species Conservation
Act of 1969, which was in turn replaced
by the Endangered Species Act of 1973
(ESA, 16 U.S.C. 1531 et seq.). We
subsequently assumed jurisdiction for
shortnose sturgeon under a 1974
government reorganization plan (39 FR
41370, November 27, 1974). In Canada,
the shortnose sturgeon falls under the
jurisdiction of the Department of
Fisheries and Oceans (DFO) and was
first assessed by the Committee on the
Status of Endangered Wildlife in Canada
(COSEWIC) as ‘‘Special Concern’’ in
1980. This status was reconfirmed in
2005, and the species was listed as
Special Concern under the Canadian
federal Species at Risk Act (SARA) in
2009. The Special Concern status was
reconfirmed again in 2015 (COSEWIC,
In Press). Shortnose sturgeon is also
listed under Appendix I of the
Convention on International Trade in
Endangered Species of Wild Fauna
(CITES).
Statutory, Regulatory and Policy
Provisions
We are responsible for determining
whether species are threatened or
endangered under the ESA (16 U.S.C.
1531 et seq.). To make this
determination, we first consider
whether a group of organisms
constitutes a ‘‘species’’ under section 3
of the ESA, and then we consider
whether the status of the species
qualifies it for listing as either
threatened or endangered. Section 3 of
the ESA defines a ‘‘species’’ 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’’ (16 U.S.C. 1532(16)). A joint
policy issued by NMFS and the U.S.
Fish and Wildlife Service (USFWS;
collectively referred to as ‘‘the
Services’’) clarifies the interpretation of
the phrase ‘‘distinct population
segment’’ (DPS) for the purposes of
listing, delisting, and reclassifying a
species under the ESA (‘‘DPS Policy,’’
61 FR 4722, February 7, 1996). The DPS
Policy identifies two criteria for
determining whether a population is a
DPS: (1) The population must be
‘‘discrete’’ in relation to the remainder
of the taxon (species or subspecies) to
which it belongs; and (2) the population
must be ‘‘significant’’ to the remainder
of the taxon to which it belongs.
Congress has instructed the Secretary
to exercise the authority to recognize
DPS’s ‘‘sparingly and only when the
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biological evidence indicates that such
action is warranted’’ (S. Rep. 96–151
(1979)). The law is not settled as to the
extent of the Services’ discretion to
modify a species-level listing to
recognize a DPS having a status that
differs from the original listing. In a
recent decision, the United States
District Court for the District of
Columbia held that the ESA does not
permit identification of a DPS solely for
purposes of delisting. Humane Soc’y v.
Jewell, 76 F. Supp. 3d 69 (D.D.C. Dec.
19, 2014), appeal docketed, No. 15–
5041 (D.C. Cir. Feb. 19, 2015) (Western
Great Lakes gray wolves) (consolidated
with Nos. 15–5043, 15–5060, and 15–
5061).
A species, subspecies, or DPS is
‘‘endangered’’ if it is in danger of
extinction throughout all or a significant
portion of its range, and ‘‘threatened’’ if
it is likely to become endangered within
the foreseeable future throughout all or
a significant portion of its range (ESA
sections 3(6) and 3(20), respectively, 16
U.S.C. 1532(6) and (20)). We interpret
an ‘‘endangered species’’ to be one that
is presently in danger of extinction. A
‘‘threatened species,’’ on the other hand,
is not presently in danger of extinction,
but is likely to become so in the
foreseeable future. In other words, the
primary statutory difference between a
threatened and endangered species is
the timing of when a species may be in
danger of extinction, either presently
(endangered) or in the foreseeable future
(threatened). In addition, we interpret
‘‘foreseeable future’’ as the horizon over
which predictions about the
conservation status of the species can be
reasonably relied upon.
Pursuant to the ESA and our
implementing regulations, the
determination of whether a species is
threatened or endangered shall be based
on any one or a combination of the
following five section 4(a)(1) factors:
The present or threatened destruction,
modification, or curtailment of habitat
or range; overutilization for commercial,
recreational, scientific, or educational
purposes; disease or predation;
inadequacy of existing regulatory
mechanisms; and any other natural or
manmade factors affecting the species’
existence. 16 U.S.C. 1533(a)(1); 50 CFR
424.11(c). Listing determinations must
be based solely on the best scientific
and commercial data available, after
conducting a review of the species’
status and after taking into account any
efforts being made by any state or
foreign nation (or any political
subdivision of such state or foreign
nation) to protect the species. 16 U.S.C.
1532(b)(1)(A).
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Under section 4(a)(1) of the ESA and
the implementing regulations at 50 CFR
424.11(d), a species shall be removed
from the list if the Secretary of
Commerce determines, based on the
best scientific and commercial data
available after conducting a review of
the species’ status, that the species is no
longer threatened or endangered
because of one or a combination of the
section 4(a)(1) factors. The regulations
provide that a species listed under the
ESA may be delisted only if such data
substantiate that it is neither
endangered nor threatened for one or
more of the following reasons:
(1) Extinction. Unless all individuals of the
listed species had been previously identified
and located, and were later found to be
extirpated from their previous range, a
sufficient period of time must be allowed
before delisting to indicate clearly that the
species is extinct.
(2) Recovery. The principal goal of the
USFWS and NMFS is to return listed species
to a point at which protection under the ESA
is no longer required. A species may be
delisted on the basis of recovery only if the
best scientific and commercial data available
indicate that it is no longer endangered or
threatened.
(3) Original data for classification in error.
Subsequent investigations may show that the
best scientific or commercial data available
when the species was listed, or the
interpretation of such data, were in error.
50 CFR 424.11(d).
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To complete the required finding in
response to the current delisting
petition, we first evaluated whether the
petitioned entity meets the criteria of
the DPS Policy. As we noted in our
initial petition finding, a determination
whether to revise a species-level listing
to recognize one or more DPSs in place
of a species-level listing involves, first,
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determining whether particular DPS(s)
exist(s) (based on meeting the criteria of
the DPS Policy) and, if that finding is
affirmative, complex evaluation as to
the most appropriate approach for
managing the species in light of the
purposes and authorities under the ESA.
Species Description
Below, we summarize basic life
history information for shortnose
sturgeon. A more thorough discussion of
all life stages, reproductive biology,
habitat use, abundance estimates and
threats are provided in the Shortnose
Sturgeon Biological Assessment
completed by the Shortnose Sturgeon
Status Review Team in 2010 (SSRT
2010; https://www.fisheries.noaa.gov/pr/
species/fish/shortnose-sturgeon.html).
There are 25 species and four
recognized genera of sturgeons (family
Acipenseridae), which comprise an
ancient and distinctive assemblage with
fossils dating to at least the Upper
Cretaceous period, more than 66 million
years ago (Findeis 1997). The shortnose
sturgeon, Acipenser brevirostrum, is the
smallest of the three extant sturgeon
species in eastern North America. Many
primitive physical characteristics that
reflect the shortnose sturgeon’s ancient
lineage have been retained, including a
protective armor of bony plates called
‘‘scutes’’; a subterminal, protractile
tube-like mouth; and chemosensory
barbels. The general body shape is
cylindrical, tapering at the head and
caudal peduncle, and the upper lobe of
the tail is longer than the lower lobe.
Shortnose sturgeon vary in color but are
generally dark brown to olive or black
on the dorsal surface, lighter along the
row of lateral scutes, and nearly white
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on the ventral surface. Adults have no
teeth but possess bony plates in the
esophagus that are used to crush hard
prey items (Vladykov and Greeley 1963;
Gilbert 1989). The skeleton is almost
entirely cartilaginous with the exception
of some bones in the skull, jaw and
pectoral girdle.
Shortnose sturgeon occur along the
East Coast of North America in rivers,
estuaries, and marine waters.
Historically, they were present in most
major rivers systems along the Atlantic
coast (Kynard 1997). Their current
riverine distribution extends from the
Saint John River, New Brunswick,
Canada, to possibly as far south as the
St. Johns River, Florida (Figure 1;
Kynard 1997; Gorham and McAllister
1974). Recently available information
indicates that their marine range
extends farther northward than
previously thought and includes the
Minas Basin, Nova Scotia (Dadswell et
al. 2013). The distribution of shortnose
sturgeon across their range, however, is
disjunct, with no known reproducing
populations occurring within the
roughly 400 km of coast between the
Chesapeake Bay and the southern
boundary of North Carolina. Shortnose
sturgeon live in close proximity with
Atlantic sturgeon (Acipenser oxyrinchus
oxyrinchus) throughout much of their
range. However, Atlantic sturgeon
spend more of their life cycle in the
open ocean compared to shortnose
sturgeon. Within rivers, shortnose
sturgeon and Atlantic sturgeon may
share foraging habitat and resources, but
shortnose sturgeon generally spawn
farther upriver and earlier than Atlantic
sturgeon (Kynard 1997, Bain 1997).
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Shortnose sturgeon typically migrate
seasonally between upstream freshwater
spawning habitats and downstream
foraging mesohaline (i.e., salinities of 5
to 18 parts per thousand) habitat based
on water temperature, flow, and salinity
cues. Based on their varied and complex
use of freshwater, estuarine, and marine
waters, shortnose sturgeon have been
characterized in the literature as
‘‘anadromous’’ or ‘‘amphidromous’’
(Bain 1977; Kieffer and Kynard 1993).
An anadromous species is defined as
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one that spawns in freshwater and
spends much of its life cycle in marine
waters, whereas a freshwater
amphidromous species is one that
spawns and remains in freshwater for
most of its life cycle but spends some
time in saline water. Because shortnose
sturgeon had historically rarely been
detected far from their natal estuary,
they were once considered to be largely
confined to their natal rivers and
estuaries (NMFS 1998). However, more
recent research has demonstrated that
shortnose sturgeon leave their natal
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estuaries, undergo coastal migrations,
and use other river systems to a greater
extent than previously thought (Kynard
1997; Savoy 2004; Fernandes 2010;
Zydlewski et al. 2011; Dionne et al.
2013). The reasons for inter-riverine
movements are not yet clear, and the
degree to which this behavior occurs
appears to vary among river systems.
Shortnose sturgeon are benthic
feeders, and their diet typically consists
of small insects, crustaceans, mollusks,
polychaetes, and small benthic fishes
(McCleave et al. 1977; Dadswell 1979;
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Marchette and Smiley 1982; Dadswell et
al. 1984; Moser and Ross 1995; Kynard
et al. 2000; Collins et al. 2002). Both
juvenile and adult shortnose sturgeon
primarily forage over sandy-mud
bottoms, which support benthic
invertebrates (Carlson and Simpson
1987, Kynard 1997). Shortnose sturgeon
have also been observed feeding off
plant surfaces (Dadswell et al. 1984).
Sturgeon likely use electroreception,
olfaction, and tactile chemosensory cues
to forage, while vision is thought to play
a minor role (Miller 2004).
Foraging in the colder rivers in the
northern part of their range appears to
greatly decline or cease during winter
months when shortnose sturgeon
generally become inactive. In midAtlantic areas, including the
Chesapeake Bay, and the Delaware
River, foraging is believed to occur yearround, though shortnose sturgeon are
believed to feed less in the winter (J.
O’Herron, Amitrone O’Herron, Inc.,
pers. comm. 2008 as cited in SSRT
2010). In the southern part of their
range, shortnose sturgeon are known to
forage widely throughout the estuary
during the winter, fall, and spring
(Collins and Smith 1993, Weber et al.
1999). During the hotter months of
summer, foraging may taper off or cease
as shortnose sturgeon take refuge from
high water temperatures.
Shortnose sturgeon are relatively
small compared to most extant sturgeon
species and reach a maximum length of
about 120 cm total length (TL) and
weight of about 24 kg (Dadswell 1979;
Waldman et al. 2002); however, both
maximum size and growth rate display
a pattern of gradual variation across the
range, with the fastest growth rates and
smallest maximum sizes occurring in
the more southern populations
(Dadswell et al. 1984). The
northernmost populations exhibit the
slowest growth and largest adult sizes.
The largest shortnose sturgeon reported
in the published literature to date was
collected from the Saint John River,
Canada, and measured 143cm TL (122
cm fork length (FL)) and weighed 23.6
kg (Dadswell 1979). In contrast, in their
review, Dadswell et al. (1984) indicated
that the largest adult reported from the
St. Johns River, Florida, was a 73.5 cm
(TL) female. Dadswell et al. (1984)
compared reported growth parameters
across the range and showed that the
von Bertalanffy growth parameter K and
estimated asymptotic length ranged
from 0.042 and 130.0 cm (FL),
respectively, for Saint John River fish to
0.149 and 97.0 cm (FL) for Altamaha
River, Georgia fish. However, the landlocked shortnose sturgeon population
located upstream of Holyoke Dam at
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river km 140 of the Connecticut River
has the slowest adult growth rate of any
surveyed, which may at least in part
reflect food limitations (Taubert 1980a).
Shortnose sturgeon are relatively
long-lived and slow to mature. The
oldest shortnose sturgeon reported was
a 67 year-old female from the Saint John
River, and the oldest male reported was
a 32 year-old fish, also captured in the
Saint John River (Dadswell 1979). In
general, fish in the northern portion of
the species’ range live longer than
individuals in the southern portion of
the species’ range (Gilbert 1989). Males
and females mature at about the same
length, around 45–55 cm FL, throughout
their range (Dadswell et al. 1984).
However, age at maturity varies by sex
and with latitude, with males in the
southern rivers displaying the youngest
ages at maturity (see review in Dadswell
et al. 1984). For example, age at first
maturation in males occurs at about 2–
3 years of age in Georgia and at about
10–11 years in the Saint John River.
Females mature by 6 years of age in
Georgia and at about 13 years in the
Saint John River (Dadswell et al. 1984).
Sturgeon are iteroparous, meaning
they reproduce more than once during
their lifetime. In general, male shortnose
sturgeon are thought to spawn every
other year, but they may spawn
annually in some rivers (Dovel et al.
1992; Kieffer and Kynard 1996).
Females appear to spawn less
frequently—approximately every 3 to 5
years (Dadswell 1979). Spawning
typically occurs during late winter/early
spring (southern rivers) and mid-to-late
spring (northern rivers) (Dadswell 1979,
Taubert 1980a and b, Kynard 1997). The
onset of spawning may be cued by
decreasing river discharge following the
peak spring freshet, when water
temperatures range from 8 to 15 °C and
bottom water velocities range between
25–130 cm/s, although photoperiod (or
day-length) appears to control spawning
readiness (Dadswell et al. 1984; Kynard
et al. 2012). Spawning appears to occur
in the sturgeons’ natal river, often just
below the fall line at the farthest
accessible upstream reach of the river
(Dovel 1981; Buckley and Kynard 1985;
Kieffer and Kynard 1993; O‘Herron et al.
1993; Kieffer and Kynard 1996).
Following spawning, adult shortnose
sturgeon disperse quickly down river
and typically remain downstream of
their spawning areas throughout the rest
of the year (Buckley and Kynard 1985,
Dadswell et al. 1984; Buckley and
Kynard 1985; O’Herron et al. 1993).
In a review by Gilbert (1989),
fecundity of shortnose sturgeon was
reported to range between
approximately 30,000–200,000 eggs per
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female. Shortnose sturgeon collected
from the Saint John River had a range
of 27,000–208,000 eggs and a mean of
11,568 eggs/kg body weight (Dadswell
1979). Development of the eggs and
transition through the subsequent larval,
juvenile and sub-adult life stages are
discussed in more detail in SSRT 2010.
A total abundance estimate for
shortnose sturgeon is not available.
However, population estimates, using a
variety of techniques, have been
generated for many individual river
systems. In general, northern shortnose
sturgeon population abundances are
greater than southern populations
(Kynard 1997). The Hudson River
shortnose sturgeon population is
currently considered to be the largest
extant population (61,000 adults, 95
percent CI: 52,898–72,191; Bain et al.
2007; however, see discussion of this
estimate in SSRT 2010). Available data
suggest that some populations in
northern rivers have increased over the
past several decades (e.g., Hudson,
Kennebec; Bain et al. 2000; Squiers
2003) and that others may be stable (e.g.,
Delaware; Brundage and O’Herron
2006). South of Chesapeake Bay,
populations are relatively small
compared to the northern populations.
The largest population of shortnose
sturgeon in the southern part of the
range is from the Altamaha River, which
was most recently estimated at 6,320
fish (95% CI: 4387–9249; Devries 2006).
Occasional observations of shortnose
sturgeon have been made in some rivers
where shortnose sturgeon are
considered extirpated (e.g., St. Johns, St.
Mary’s, Potomac, Housatonic, and
Neuse rivers); the few fish that have
been observed in these rivers are
generally presumed to be immigrants
from neighboring basins.
The most recent total population
estimate for the Saint John River dates
to the 1970’s. Using tag recapture data
from 1973–1977, Dadswell (1979)
calculated a Jolly-Seber population
estimate of 18,000 (±30% SE; 95 percent
CI: 7,200–28,880, COSEWIC, In Press)
adults (≤ 50 cm) below the Mactaquac
Dam. Several partial population
estimates are also available for the
Kennebecasis River, a tributary in the
lower reaches of the Saint John River.
Litvak (unpublished data) calculated a
Jolly-Seber estimate of 2,068 fish (95%
CI: 801–11,277) in the Kennebecasis
using mark-recapture data from 1998 to
2004 (COSEWIC, In Press). Based on
videotaping of overwintering
aggregations of shortnose sturgeon on
the Kennebecasis River at the
confluence of the Hammond River (rkm
35), Li et al. (2007) used ordinary
Kriging to estimate that 4,836 (95% CI:
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4,701–4,971) adult shortnose sturgeon
were overwintering in that area.
Usvyatsov et al. (2012) repeated this
sampling in 2009 and 2011 and, using
three different modeling techniques,
estimated a total of 3,852–5,222
shortnose sturgeon in the study area,
which suggests fairly stable abundance
and habitat use at this site.
Threats that contributed to the
species’ decline and led to the listing of
shortnose sturgeon under the ESA
included pollution, overfishing, and
bycatch in the shad fishery (USDOI
1973). Shortnose sturgeon were also
thought to be extirpated, or nearly so,
from most of the rivers in their
historical range (USDOI 1973). In the
late nineteenth and early twentieth
centuries, shortnose sturgeon were
commonly harvested incidental to
Atlantic sturgeon, the larger and more
commercially valuable of these two
sympatric sturgeon species (NMFS
1998). Although there is currently no
legal directed fishing for shortnose
sturgeon in the United States, poaching
is suspected, and bycatch still occurs in
some areas. In particular, shortnose
sturgeon are caught incidentally by bass
anglers and in the alewife/gaspereau,
American shad, American eel, and
Atlantic sturgeon fisheries in the Saint
John River; and shad fisheries in the
Altamaha River, Santee River, Savannah
River, and elsewhere (COSEWIC, In
Press; SSRT 2010; Bahn et al. 2009;
COSEWIC 2005). The construction of
dams has also resulted in substantial
loss of historical shortnose sturgeon
habitat in some areas along the Atlantic
seaboard. The construction and
operation of dams can impede upstream
movement to sturgeon spawning habitat
(e.g., Connecticut River, Santee River).
Remediation measures, such as dam
removal or modification to allow for fish
passage have improved access in some
rivers, and additional similar restoration
efforts are being considered in other
areas (e.g., possible removal of the
Mactaquac dam in the Saint John River).
Other possible and ongoing threats
include operation of power generating
stations, water diversion projects,
dredging, and other in-water activities
that impact habitat.
Distinct Population Segment Analysis
The following sections provide our
analysis of whether the petitioned
entity—the Saint John River population
of shortnose sturgeon—qualifies as a
DPS of shortnose sturgeon (whether it is
both ‘‘discrete’’ and ‘‘significant’’). To
complete this analysis we relied on the
best scientific and commercial data
available and considered all relevant
literature and public comments
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submitted in response to our 90-day
finding (80 FR 18347, April 6, 2015).
For purposes of this analysis, we
defined the Saint John River population
segment of shortnose sturgeon to consist
of shortnose sturgeon spawned in the
Saint John River downstream of the
Mactaquac Dam. Prior to construction of
Mactaquac Dam in 1968/1969, sturgeon
occurred upstream of the dam; however,
it is unclear whether these were
shortnose and/or Atlantic sturgeon and
whether any sturgeon are still present
upstream of the dam (COSEWIC, In
Press). Lacking this information, we
cannot consider fish that may be present
upstream of the dam in our distinct
population segment analysis.
Throughout our discussion below we
also use the term ‘‘population’’ to refer
collectively to all shortnose sturgeon
that are presumed to be natal to a
particular river rather than using this
term to refer strictly to a completely
closed reproductive unit.
Discreteness Criterion
The Services’ joint DPS Policy states
that a population segment 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.
(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 (61 FR 4722,
February 7, 1996).
There are no physical barriers
preventing the movement of Saint John
River shortnose sturgeon outside of the
Saint John River estuary or along the
coast. The Mactaquac Dam, located
about 140 km upstream and at the head
of tide (Canadian Rivers Institute 2011),
is the first upstream physical barrier on
the Saint John River. This and other
dams on the Saint John River block
shortnose sturgeon from accessing
upstream habitats, but there are no dams
or other physical barriers separating
Saint John River sturgeon from other
shortnose sturgeon populations.
As mentioned previously, shortnose
sturgeon have been documented to leave
their natal river/estuary and move to
other rivers to varying extents across
their range. For example, telemetry data
generated by Zydlewski et al. (2011)
during 2008–2010 indicate that inter-
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riverine movements of adult shortnose
sturgeon occur fairly frequently among
rivers in Maine. Seventy percent of
tagged adults (25 of 41 fish) moved
between the Penobscot and Kennebec
rivers (about 150 km away), and up to
52% of the coastal migrants (13 of 25
fish) also used other, smaller river
systems (i.e., Damariscotta, Medomak,
St. George) between the Penobscot and
Kennebec rivers (Zydlewski et al. 2011).
Shortnose sturgeon are also known to
move between rivers in Maine and (e.g.,
Kennebec, Saco) and the Merrimack
River estuary in Massachusetts,
traveling distances of up to about 250
km (as measured by a conservative,
direct path distance; Little et al. 2013;
Wippelhauser et al. 2015). At the other
end of the range, in the Southeast
United States, inter-riverine movements
appear fairly common and include
movements between the Savannah River
and Winyah Bay and between the
Altamaha and Ogeechee rivers (Peterson
and Farrae 2011; Post et al. 2014).
Many inter-riverine movements have
been observed elsewhere within the
species’ range, but patterns are not yet
well resolved. For example, some
shortnose sturgeon captured and/or
tagged in the Connecticut River have
been recaptured, detected, or were
previously tagged in the Housatonic
River (T. Savoy, CT DEP, pers. comm.
2015), the Hudson River (Savoy 2004),
and the Merrimack River (M. Kieffer,
USGS, pers. comm. 2015). At this time,
the available tagging and tracking
information is too limited to determine
if Hudson River and Connecticut River
shortnose sturgeon are making regular
movements outside of their natal rivers
and whether movement as far as the
Merrimack River is a normal behavior.
Movement data from the Chesapeake
Bay is also relatively limited, but
existing data indicate that shortnose
sturgeon do move from the Chesapeake
Bay through the Chesapeake and
Delaware Canal into the Delaware River
(Welsh et al. 2002).
The distances of the reported marine
migrations vary widely from very short
distances—such as between the Santee
River and Winyah Bay, which are only
about 15 km apart—to fairly long—as in
the case of movements between the
Merrimack and the Penobscot rivers,
which are about 339 km apart at their
mouths.1 In general, the available data
1 Distances between rivers mouths reported here
were measured in GIS using the NOAA Medium
Resolution Vector Shoreline, 20m bathymetry
contour, and a fixed scale of 1:250,000. Estimated
distances reported are the average of three,
independently drawn and measured paths for each
river pair. The assumed travel path between river
mouths was the shortest possible distance that
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suggest that movements between
geographically proximate rivers are
more common, while movements
between more distant rivers do not, or
only rarely, occur. A detailed discussion
of the physical movements of shortnose
sturgeon is provided in SSRT 2010.
The extent of coastal movements of
shortnose sturgeon from the Saint John
River is currently unknown (COSEWIC,
In Press); however, some limited data
are available and provide some insight
into whether these fish may be
geographically isolated from other
populations. Any movement between
Saint John River sturgeon and the
nearest population in the Penobscot
River would require a marine migration
of about 362 km, a similar travel
distance as between the Merrimack and
the Penobscot rivers (340 km) and
between the Connecticut and Merrimack
rivers (348 km).2 Dadswell (1979)
reported that of 121 marked Saint John
River shortnose sturgeon recaptured by
commercial fisherman, 13 fish (11
percent) were recaptured in the Bay of
Fundy, indicating that a portion of the
population migrated into the marine
environment. In addition, a confirmed
shortnose sturgeon was caught in a
fishing weir in the Minas Basin, off the
coast of Nova Scotia about 165 km north
of the mouth of the Saint John River
(Dadswell et al. 2013). Fishermen in the
Minas Basin also claim to catch about
one to two shortnose sturgeon per year
in their weirs (Dadswell et al. 2013).
While it is plausible that the shortnose
sturgeon captured in the Minas Bay
originated from the Saint John River,
data to confirm this are not available. In
contrast, limited telemetry data suggest
that movements outside of the Saint
John River are not common. Of 64
shortnose sturgeon tagged in the Saint
John River over the course of about 16
years from 1999 to 2015, none have
been detected moving past the farthest
downriver acoustic receiver located near
the Saint John Harbor Bridge (M. Litvak,
pers. comm. July 31, 2015).
Overall, while there is unambiguous
evidence that shortnose sturgeon from
the Saint John River leave the estuary—
at least occasionally—and use the
marine environment, and that shortnose
sturgeon are capable of making long
distance movements between river
followed the general outline of the coast and was
constrained by the 20m bathymetry contour, except
where the shortest travel path across a deep, narrow
inlet or bay crossed the 20m bathymetry contour.
2 Distances reported here were measured
following the same methods described in the
previous footnote. The distance reported between
the Connecticut and Merrimack River assumes a
travel path via the Cape Cod Canal. A travel path
around Cape Cod would instead result in a marine
migration of about 560 km.
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systems, there are no available data on
coastal migrations of Saint John River
shortnose sturgeon. To date, there are
also no reported observations or
detections of shortnose sturgeon from
the Gulf of Maine rivers moving into the
Saint John River. Thus, while it is
possible that the Saint John River
shortnose sturgeon come in contact with
shortnose sturgeon from elsewhere, it is
also likely that some degree of
geographical isolation by distance is
occurring.
Although acoustic telemetry studies
have revealed that shortnose sturgeon
leave their natal river systems to a much
greater extent than previously thought,
such movements do not necessarily
constitute permanent emigration or
indicate interbreeding of populations.
Tagging and telemetry studies within
several river systems have provided
evidence that shortnose sturgeon in
those particular systems tend to spawn
in their natal river (e.g., Dovel 1981;
Buckley and Kynard 1985; Kieffer and
Kynard 1993; O‘Herron et al. 1993;
Kieffer and Kynard 1996). Tag return
data for shortnose sturgeon in the Saint
John River over the course of a 4-year
study completed by Dadswell (1979)
suggests there is little emigration from
this system as well, and that spawning
takes place in the freshwater sections of
the upper estuary. The high site fidelity
to natal rivers suggested by this and
other studies indicates a there is a
possible behavioral mechanism for the
marked separation of the Saint John
River population of shortnose sturgeon
from other populations of the species.
A substantial amount of genetic data
has become available since the ‘‘Final
Recovery Plan for Shortnose Sturgeon’’
was developed in 1998. Below, we
summarize the best available genetic
data and information, which informed
our evaluation of the ‘‘discreteness’’ of
the Saint John River population
segment. A more in-depth presentation
of genetic data, including discussions of
types of analyses and assumptions, is
available in the Biological Assessment
(SSRT 2010).
Much of the published information on
population structure for shortnose
sturgeon has been based on the genetic
analysis of the maternally inherited
mitochondrial DNA (mtDNA) due in
part to the difficulties of analyzing data
from the polyploid nuclear genome
(Waldman et al. 2008). The analyses
have focused on a moderately
polymorphic 440 base pair portion of
the mtDNA control region—a relatively
rapidly evolving region of mtDNA and
thus a good indicator of populationlevel differentiation. Haplotype
frequencies and sequence divergence
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data have consistently indicated an
overall isolation-by-distance pattern of
genetic population structure across the
species’ range, meaning that
populations of shortnose sturgeon
inhabiting rivers and embayments that
are geographically more distant tend to
be less related than those that are
geographically closer (e.g., Walsh et al.
2001, Grunwald et al. 2002, Waldman et
al. 2002, and Wirgin et al. 2005; Wirgin
et al. 2009). The haplotypes observed
are typically shared across two to four
or more adjacent sampled rivers but
with little sharing of haplotypes
between northern and southern
populations (Waldman et al. 2002;
Wirgin et al. 2009). Results for the Saint
John River are compatible with these
general patterns. For example, in the
largest study to date, Wirgin et al. (2009)
observed eight haplotypes within the
Saint John River sample (n=42); and of
the eight observed haplotypes, one was
exclusive (or ‘‘private’’) to the Saint
John River (and observed in 1 of 42
fish), and the remaining haplotypes
were shared with two to six other rivers.
None of the shared haplotypes were
observed in samples south of the
Chesapeake Bay. A previously
unreported haplotype was recently
observed in 2 of 15 shortnose caught
from the Kennebecasis River, a tributary
of the Saint John (Kerr, 2015; P. Wilson,
public comment, May 2015). This new
haplotype could indicate an even
greater degree of differentiation of the
Saint John River fish; however, no other
rivers were sampled or analyzed as part
of this study.
Despite the localized sharing of
haplotypes, frequencies of the observed
haplotypes are significantly different in
most pairwise comparisons of the rivers
sampled (i.e., comparisons of haplotype
frequencies from samples from two
rivers), including many adjacent rivers
(Wirgin et al. 2009). Such pairwise
comparisons for the Saint John River in
particular have indicated that this
population is genetically distinct from
the geographically closest sampled
populations, including the Penobscot,
Kennebec, and Androscoggin rivers
(Grunwald et al. 2002; Waldman et al.
2002; Wirgin et al. 2005; Wirgin et al.
2009). For example, Wirgin et al. (2009)
reported significant differences
(p<0.0005) in haplotype frequencies
between Saint John River shortnose
sturgeon (n=42) and Penobscot (n=44,
Chi-square=37.22), Kennebec (n=54,
Chi-square=54.85), and Androscoggin
(n=48, Chi-square=37.91) river samples.
The level of genetic differentiation
between the Saint John River population
and the Penobscot, Kennebec, and
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Androscoggin rivers also appears
substantial, with Phi ST values ranging
from 0.213 to 0.291 (where Phi ST ranges
from 0 to 1, with 1 indicating complete
isolation; Wirgin et al. 2009).
Estimates of female-mediated gene
flow between the Saint John River and
the Gulf of Maine rivers are fairly low.
Wirgin et al. (2009) estimated femalemediated gene flow between the Saint
John River and other Gulf of Maine
rivers as 1.90–2.85 female migrants per
generation based on Phi ST values, and
as 1.5–1.9 females per generation in a
separate, coalescent-based analysis. This
result suggests that (if model
assumptions are true) no more than
three female shortnose sturgeon from
the Saint John River are likely to spawn
in the other Gulf of Maine rivers (or vice
versa) per generation. These results
provide additional evidence that the
degree of female-based reproductive
exchange between the Saint John River
population and other nearby shortnose
river populations has been relatively
limited over many generations.
More recently, King et al. (2014)
completed a series of analyses using
nuclear DNA (nDNA) samples from 17
extant shortnose sturgeon populations
across the species range. In contrast to
the maternally inherited mtDNA, nDNA
reflects the genetic inheritance from
both the male and female parents. King
et al. (2014) surveyed the samples at 11
polysomic microsatellite DNA loci and
then evaluated the 181 observed alleles
as presence/absence data using a variety
of analytical techniques. The population
structuring revealed by these analyses is
consistent with the previous mtDNA
analyses in that they also indicate a
regional scale isolation-by-distance
pattern of genetic differentiation.
Analysis of genetic distances among
individual fish (using principle
coordinate analysis, PCO) revealed that
the sampled fish grouped into one of
three major geographic units: (1)
Northeast, which included samples
from the Saint John, Penobscot,
Kennebec, Androscoggin, and
Merrimack rivers; (2) Mid-Atlantic,
which included samples from the
Connecticut, Hudson, and Delaware
rivers, as well as the Chesapeake Bay
proper; and (3) Southeast, which
included samples from the Cape Fear
River, Winyah Bay, the Santee-Cooper,
Edisto, Savannah, Ogeechee, and
Altamaha rivers, and Lake Marion (King
et al. 2014).
Subsequent analyses revealed that
each of the three regions has a different
pattern of sub-structuring. Within the
Northeast group, two separate analyses
(PCO and STRUCTURE) indicated a
high degree of relatedness and possible
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panmixia (i.e., random mating of
individuals) among the Penobscot,
Kennebec, and Androscoggin rivers;
whereas, the Saint John and Merrimack
rivers appeared more differentiated from
each other as well as from the other Gulf
of Maine rivers (King et al. 2014).
Pairwise comparisons at the population
level showed that, within the Northeast
region, estimates of genetic
differentiation were greatest between
the Saint John and Merrimack rivers
(Phi PT = 0.100, p <0.0004), the two most
distant rivers within this region.
Pairwise comparisons of the Saint John
River to the remaining rivers within the
Northeast region revealed lower but still
statistically significant levels of genetic
differentiation (Phi PT = 0.068–0.077;
King et al. 2014). Relatively low levels
of differentiation were observed in
pairwise comparisons for all other rivers
within the Northeast region (Phi PT =
0.013–0.087), half of which were not
statistically significant (King et al.
2014). In comparison, within the MidAtlantic group, pairwise comparisons
among rivers showed moderate levels of
genetic differentiation among most river
populations (average Phi PT = 0.077,
range = 0.018–0.118); whereas,
estimates of population level genetic
differentiation were very low among
samples populations in the Southeast
group (average Phi PT = 0.047, range =
0.005 to 0.095; King et al. 2014),
suggesting a more genetically similar set
of populations.
Theoretical estimates of gene flow
(derived from Phi PT values) between the
Saint John River and the other Northeast
rivers ranged from 2.25 to 3.43 migrants
per generation (King et al. 2014). Gene
flow estimates for the Merrimack River
were similarly low, ranging from 2.25 to
4.06 (King et al. 2014). In contrast, the
effective number of migrants per
generation estimated to occur between
the remaining rivers within the
Northeast region was much higher and
ranged from 16.42 to 83.08 (King et al.
2014).
Overall, the analyses completed by
King et al. (2014) indicate that
differentiation among Northeast
populations is less than that observed
among the Mid-Atlantic populations
and greater than that observed among
Southeast populations. However, within
the Northeast region, both the Saint
John and Merrimack River sample
populations are genetically distinct from
the other sample populations. Although
the estimates of gene flow suggest some
connectivity between the Saint John and
other rivers within the Northeast, the
significantly different allele and
haplotype frequencies shown
consistently in the nDNA and mtDNA
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studies provide indirect evidence that
the Saint John River population is
relatively reproductively isolated.
As highlighted in the DPS Policy,
quantitative measures of morphological
discontinuity or differentiation can
serve as evidence of marked separation
of populations. We examined whether
the morphological data for shortnose
sturgeon across its range provide
evidence of marked separation of the
Saint John River population. As noted
previously, maximum adult size (length
and weight) varies across the range,
with the largest maximum sizes
occurring in the Saint John River at the
northernmost end of the range, and the
smallest sizes occurring in rivers at the
southern end of the range (Dadswell et
al. 1984). The largest individual
reported in the literature (122 cm FL,
23.6 kg) was captured in the Saint John
River, although there is also a report of
a specimen measuring 124.6 cm FL (M.
Litvak, unpublished data, as cited in
COSEWIC, In Press). Lengths of
shortnose sturgeon captured in surveys
of the Saint John River in 1974–1975
ranged from 60 to 120 cm FL (n=1,621).
The majority of these fish, however,
were smaller than 100 cm FL (1,476
fish), and only six fish were longer than
111 cm FL (Dadswell 1979). To the
south, in the Kennebec River, Maine
shortnose sturgeon captured during
1980 and 1981 had lengths ranging from
58.5 to 103.0 cm FL, and averaging 80.8
cm FL (n=24; Walsh et al. 2001).
Smaller size ranges are reported for
rivers in the southernmost portion of the
range with some occasional captures of
larger specimens. For example, adult
shortnose sturgeon captured in the
Altamaha River, Georgia, in 2010–2013
ranged from 57.4–83.0 cm FL and
averaged 70.1 cm long (FL, n=40;
Peterson 2014), but a shortnose sturgeon
measuring 104.5 cm FL and weighing
8.94 kg was captured in the Altamaha
River in summer, 2004 (D. Peterson,
UGA, unpubl. data). Overall, the
attribute of size appears to display clinal
variation, meaning there is a gradual
change with geographic location
(Huxley 1938). The fact that the Saint
John River population segment, which
lies at the northernmost end of the
range, exhibits the largest sizes does not
in itself constitute a morphological
discontinuity. Given the apparent
gradual nature of the variation in size
with latitude, we find that there is no
marked separation of the Saint John
River population segment on the basis
of a quantitative discontinuity in size.
In addition to body size, other
attributes such as snout length, head
length, and mouth width can provide
evidence of a morphological
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discontinuity and were also considered.
Walsh et al. (2001) examined six
morphological and five meristic
attributes for shortnose sturgeon in the
Androscoggin, Kennebec, and Hudson
rivers. All morphological features
measured (i.e., body length, snout
length, head length, mouth width, and
interorbital width) were largest for the
Kennebec River fish and smallest for
fish from the southern-most river in the
study, the Hudson River (Walsh et al.
2001). Meristic features (e.g., scute
counts) were similar for the three rivers
and were not related to fish size (Walsh
et al. 2001). Overall, the degree of
phenotypic differentiation of fish from
the two rivers in Maine (Androscoggin
and Kennebec), which share an estuary
mouth, was very low, while a much
greater degree of differentiation was
observed for the fish from the Hudson
River (Walsh et al. 2001). This result
was congruent with results of
corresponding mtDNA analyses, which
indicated that the Hudson River had a
much greater degree of genetic
differentiation from, and much lower
rate of gene flow with, the two rivers in
Maine (Walsh et al. 2001). The results
of this particular study suggest there
could be clinal variation in these other
phenotypic characteristics, similar to
the pattern observed for body size. As
far as we are aware, however, similar
studies have not yet been conducted to
examine the variation in additional sets
of morphological attributes across the
range of shortnose sturgeon and relative
to the Saint John River population in
particular. Therefore, there is no basis to
conclude marked separation of the Saint
John River population segment on the
basis of morphological discontinuity.
In conclusion, although the currently
available data do not show that the
Saint John River shortnose sturgeon
constitute a completely isolated or
closed population, we find that
available genetic data, evidence of site
fidelity, and the likelihood of some
degree of geographical isolation together
constitute sufficient information to
indicate that the Saint John River
shortnose sturgeon are markedly
separated from other populations of
shortnose sturgeon. Thus, after
considering the best available data and
all public comments submitted in
response to our initial petition finding,
we conclude that the Saint John River
population segment of shortnose
sturgeon is ‘‘discrete.’’ We therefore
proceeded to evaluate the best available
data with respect to the second criterion
of the DPS Policy, ‘‘significance.’’
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Significance Criterion
Under the DPS Policy, if a population
segment is found to be discrete, then we
proceed to the next step of evaluating its
biological and ecological significance to
the taxon to which it belongs. As we
explained above, a population must be
both ‘‘discrete’’ (the first prong of the
DPS Policy) and ‘‘significant’’ (the
second prong of the DPS Policy) to
qualify for recognition as a DPS.
Consideration of significance may
include, but is not limited to: (1)
Persistence of the discrete population
segment in an ecological setting unusual
or unique for the taxon; (2) evidence
that the loss of the discrete population
segment would result in a significant
gap in the range of a taxon; (3) evidence
that the discrete population segment
represents the only surviving natural
occurrence of a taxon that may be more
abundant elsewhere as an introduced
population outside its historical range;
and (4) evidence that the discrete
population segment differs markedly
from other populations of the species in
its genetic characteristics (61 FR 4722,
February 7, 1996). These four factors are
non-exclusive; other relevant factors
may be considered in the ‘‘significance’’
analysis. Further, significance of the
discrete population segment is not
necessarily determined by existence of
one of these classes of information
standing alone. Rather, information
analyzed under these and any other
applicable considerations is evaluated
relative to the biological and ecological
importance of the discrete population to
the taxon as a whole. Accordingly, all
relevant and available biological and
ecological information is analyzed to
determine whether, because of its
particular characteristics, the
population is significant to the
conservation of the taxon as a whole.
Persistence in an Ecological Setting
Unusual or Unique for the Taxon
Shortnose sturgeon once occupied
most major rivers systems along the
Atlantic coast of North America (Kynard
1997). Although extirpated from some
areas due mainly to overharvest,
bycatch, pollution, and habitat
degradation, shortnose sturgeon still
occur in at least 25 rivers systems
within their historical range (NMFS
1998). Throughout their current range,
shortnose sturgeon occur in riverine,
estuarine, and marine habitats; and, as
adults, generally move seasonally
between freshwater spawning habitat
and downstream mesohaline and
sometimes coastal marine areas in
response to cues such as water
temperature, flow, and salinity. Like
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other species of sturgeon (e.g. A.
transmontanus in the Columbia River,
Oregon), shortnose sturgeon are also
capable of adopting a fully freshwater
existence, as is the case for the
population of shortnose sturgeon above
the Holyoke Dam in the Connecticut
River and in Lake Marion, South
Carolina. While each river system
within the shortnose sturgeon’s range is
similar in terms of its most basic
features and functions, each river
system differs to varying degrees in
terms of its specific, physical and
biological attributes, such as hydrologic
regime, benthic substrates, water
quality, and prey communities. A few
examples are discussed briefly below.
The Saint John River begins in
northern Maine, United States, travels
through New Brunswick, Canada, and
empties into the Bay of Fundy within
the northeast Gulf of Maine. The river
is approximately 673 km long, fed by
numerous tributaries, and has a large
tidal estuary and a basin area of over
55,000 km2 (Kidd et al. 2011).
According to the Nature Conservancy’s
(TNC) ecoregion classification system,
the Saint John River watershed lies
within the New England-Acadian
(terrestrial), Northeast United States and
Southeast Canada Atlantic Drainages
(freshwater), and the Gulf of Maine/Bay
of Fundy (marine) ecoregions. The mean
annual discharge is approximately 1,100
m3/s, dissolved oxygen levels average
8.5 to 11 mg/l, and benthic substrates
downstream of the Mataquac Dam
consist largely of shifting sands (Kidd et
al. 2011). Due to the low slope of the
lower reaches and the extreme tidal
range of the Bay of Fundy, the head of
the tide can extend about 140 km
upstream from the river mouth (Kidd et
al. 2011). During the shortnose sturgeon
spring/summer spawning season, water
temperatures range from about 10 to 15
°C; and within overwintering areas,
water temperature range between 0 and
13 C (Dadswell 1979; Dadswell et al.
1984). Shortnose sturgeon in the Saint
John River appear to move to deeper
waters when surface water temperatures
exceed 21 °C (Dadswell et al. 1984).
Further to the south, but still within the
same terrestrial, freshwater, and marine
TNC ecoregions as the Saint John River,
is the smaller Penobscot River system in
Maine. This river is 175 km long (not
including the West and South
Branches), has a drainage basin of
22,265 km2, and an annual average
discharge of about 342 m3/s (Lake et al.
2012; USGS 2015). Benthic substrates,
consisting of bedrock, boulders, cobble
and sand deposits are undergoing
changes in response to the removal of
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two dams—Great Works Dam at rkm 60
and Veazie Dam at rkm 48—within the
past three years (FERC 2010; Cox et al.
2014). The Veazie Dam was located
close to the head of the tide, and
although conditions have since
changed, Haefner (1967, as cited in
Fernandes et al. 2010) stated that,
during peak springtime flows,
freshwater extends to rkm 17, and that
the salt wedge intrudes as far as about
rkm 42 when river discharges decrease
in summer. Water temperatures in
shortnose sturgeon overwintering areas
in the Penobscot River range from about
0 °C to 13.3 °C, and the fish appear to
move out of overwintering areas when
water temperatures reach about 2.4 °C
(Fernandes et al. 2010). Towards the
southern end of the range and occurring
within a very different set of ecoregions
is the Altamaha River, which is formed
by the confluence of the Ocmulgee and
Oconee rivers in Georgia. One of the
longest free-flowing systems on the
Atlantic Coast, the Atlamaha River is
just over 220 km long, has a watershed
area of about 37,300 km2, and flows
mainly eastward before emptying into
the Atlantic Ocean (TNC 2005). Tidal
influence extends up to about rkm 40
(DeVries 2006). The average annual
discharge is 381 m3/s, and benthic
substrates consist mostly of sands with
very few rocky outcrops (Heidt and
Gilbert 1979; DeVries 2006). Water
temperatures during the winter/spring
spawning period have averaged about
10.5 °C (Heidt and Gilbert 1979), which
is consistent with DeVries’ (2006)
observation that spawning runs
appeared to commence when water
temperatures reach 10.2 °C. When water
temperatures exceed 27 °C, shortnose
sturgeon typically move above the saltfresh water interface and aggregate in
deeper areas of the river (DeVries 2006);
however, shortnose sturgeon have also
been observed to use lower portions of
the river throughout the summer, even
when water temperatures averaged 34
°C (Heidt and Gilbert 1979; DeVries
2006).
Overall, the variation in habitat
characteristics across the range of
shortnose sturgeon indicates that there
is no single type or typical river system.
Despite a suite of existing threats,
shortnose sturgeon continue to occupy
many river systems across their
historical range. The fact that the Saint
John River lies at one end of the species’
range, and among other attributes,
experiences different temperature and
flow regimes, does not mean that this
particular river is unusual or unique
given the variability in habitat
conditions observed across the range.
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Therefore, we conclude that the Saint
John River is not an unusual or unique
ecological setting when viewed against
the range of the taxon as a whole.
Furthermore, though not relied up on
for our finding, we note that COSEWIC
(In Press) recently concluded that
shortnose sturgeon from other river
systems would probably be able to
survive in Canada.
Significant Gap in the Range of the
Taxon
The second consideration under the
DPS Policy in determining whether a
population may be ‘‘significant’’ to its
taxon is whether the ‘‘loss of the
discrete population segment would
result in a significant gap in the range
of a taxon’’ (61 FR 4722, February 7,
1996). Shortnose sturgeon are
distributed along the Atlantic coast of
North America from the Minas Basin,
Nova Scotia to the St. Johns River,
Florida, representing a coastal range of
roughly 3,700 km. The Saint John River,
located at the northern end of the range,
represents a small portion of the
species’ currently occupied geographic
range. In addition, although the Saint
John River is presumed to contain a
relatively large population of shortnose
sturgeon, that populaiton is not
considered the largest, and it represents
one of at least 10 spawning populations
(SSRT 2010). Furthermore, relatively
recent field data indicate shortnose
sturgeon make coastal migrations to a
greater extent than previously thought
(e.g., Dionne et al. 2013) and are capable
of making marine migrations of over 300
km (e.g., between Penobscot and
Merrimack rivers; M. Kieffer, USGS,
pers. comm. 2010). Such data suggest
the potential for recolonization of the
Saint John River by shortnose sturgeon
migrating from populations to the south.
Further indirect evidence in support of
this possibility comes from the existing
genetic data, which indicate some level
of gene flow among rivers in the
Northeast, including the Saint John
River (Wirgin et al. 2005; Wirgin et al.
2009; King et al. 2014). Thus, in light of
the potential for recolonization and the
fact that the Saint John River population
of shortnose sturgeon does not
constitute a substantial proportion of
the species’ range, we conclude that the
loss of the Saint John River would not
constitute a significant gap in the range
of the species.
Only Natural Occurrence of the Taxon
Under the DPS Policy, a discrete
population segment that represents the
‘‘only surviving natural occurrence of a
taxon that may be more abundant
elsewhere as an introduced population
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outside its historical range’’ may be
significant to the taxon as whole (61 FR
4722, February 7, 1996). This
consideration is not relevant in this
particular case, because shortnose
sturgeon are present in many river
systems throughout their historical
range (SSRT 2010).
Genetic Characteristics
As stated in the DPS Policy, in
assessing the ‘‘significance’’ of a
‘‘discrete’’ population, we consider
whether the discrete population
segment differs markedly from other
populations of the species in its genetic
characteristics (61 FR 4722, February 7,
1996). Therefore, we examined the
available data to determine whether the
Saint John River shortnose sturgeon
differ markedly in their genetic
characteristics when compared to other
populations. In conducting this
evaluation under the second criterion of
the DPS policy, we looked beyond
whether the genetic data allow for
discrimination of the Saint John
population segment from other
populations (a topic of evaluation in
connection with the first criterion of
‘‘discreteness’’), and instead focused on
whether the data indicate marked
genetic differences that appear to be
significant to the taxon as a whole. In
this sense, we give independent
meaning to the ‘‘genetic discontinuity’’
of the discreteness criterion of the DPS
Policy and the ‘‘markedly differing
genetic characteristics’’ of the
significance criterion.
Genetic analyses indicate fairly
moderate to high levels of genetic
diversity of shortnose sturgeon in most
river systems across the geographic
range (Grunwald et al. 2002, Quattro et
al. 2002; Wirgin et al. 2009). Based on
the 11 nDNA loci examined in samples
from 17 locations, King et al. (2014)
reported that the number of observed
alleles (i.e., versions of a gene at a
particular locus; here with overall
frequencies >1%) ranged from a low of
55 in the Cape Fear River (n= 3 fish) to
a high of 152 in the Hudson River (n=
45 fish); 118 alleles were observed in
the Saint John River sample (n=25 fish).
Estimated heterozygosity was not
reported by river sample, but King et al.
(2014) noted that it was lowest for the
southern rivers relative to the midAtlantic and northern river samples.
Wirgin et al. (2009) reported that
haplotypic diversity ranged from 0.500
(Santee River, n=4) to 0.862 (Altamaha
River, n= 69) across 15 sample
populations, with the Saint John River
population having a haplotype diversity
index of 0.696 (n=42). The number of
individual haplotypes observed in any
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one river sample ranged from two
(Santee River, n=4) to 13 (Winyah Bay,
n=46), with eight haplotypes observed
in the Saint John River sample (n=42,
Wirgin et al. 2009). The level of genetic
diversity based on the mtDNA was not
correlated with population size, and
there was also no evidence of
population bottlenecks, which may be
due to historical recency of most
population declines (over past ∼100
years, Grunwald et al. 2002; Wirgin et
al. 2009). Overall, the level of genetic
diversity observed for the Saint John
River population segment is not unusual
relative to that observed in the taxon as
a whole. However, Grunwald et al.
(2002) noted that the lack of reduced
haplotypic diversity within the northern
sample rivers contrasts with findings for
other anadromous fishes from
previously glaciated rivers. Grunwald et
al. (2002) hypothesized the high degree
of haplotypic diversity and large
number of unique haplotypes in the
previously glaciated northern region
(i.e., Hudson River and northward) may
be the result of a northern population
having survived in one or more northern
refugia.
As discussed previously, at a regional
scale, most of the mtDNA haplotypes
observed are shared across multiple,
adjacent rivers sampled; however, very
little sharing of haplotypes has been
documented between the northern and
southern portions of the range (Quattro
et al. 2002; Grunwald et al. 2002; Wirgin
et al. 2009). In the analysis conducted
by Wirgin et al. (2009), the Saint John
River sample had one private haplotype
(in 1 of 42 fish) and shared the
remaining 7 haplotypes with multiple
rivers. Of the seven shared haplotypes,
two were each shared with two other
river systems, including the Hudson and
Connecticut rivers, and the remaining
five haplotypes were shared across three
to six other rivers within the northeast
and mid-Atlantic portions of the range
(Wirgin et al. 2009). In an earlier study
by Quattro et al. (2002) in which control
region mtDNA was sequenced for 211
shortnose sturgeon collected from five
southeastern U.S. rivers and the Saint
John River, one haplotype was observed
in all river samples. This shared
haplotype occurred in 1 of 13 fish
(7.7%) sampled from the Saint John
River and 1 of 5 fish (20%) sampled
from Winyah Bay; the remaining river
samples contained this haplotype at
higher frequencies (36%–79%, Quattro
et al. 2002).
While the shortnose sturgeon from the
Saint John River have a fairly high
degree of genetic diversity and shared
haplotypes with other rivers, they can
be statistically differentiated from other
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river samples based on haplotype
frequencies and nDNA distance metrics
(Wirgin et al. 2009; King et al. 2014).
However, the same is also true for the
majority of rivers across the range of the
species. For example, using genetic
distances (Phi PT), King et al. (2014)
detected significant differences in all
pairwise comparisons except for three
rivers in the northeast (Penobscot,
Androscoggin, and Kennebec rivers) and
three rivers in the southeast (Edisto,
Savannah, and Ogeechee rivers).
Similarly, significant differences in
haplotype frequencies have been
reported for most river populations
sampled. In Chi-squared analyses,
Grunwald et al. (2002) reported
significant differences for all but 4 of 82
pairwise comparisons of mtDNA
nucleotide substitution haplotype
frequencies across 10 sample sets (two
of which were from different sections of
the Connecticut River), and Wirgin et al.
(2009) reported significant differences
for all but 9 of 91 pairwise comparisons
of mtDNA haplotype frequencies across
13 river populations.
The magnitude of these genetic
differences between individual river
systems varies across the range of the
species and indicates a hierarchical
pattern of differentiation. For example,
the mtDNA data reveal a deep
divergence between rivers in the
northern portion of the range from rivers
in the southern portion of the range. Of
the 29 haplotypes observed by
Grunwald et al. (2002), 11 (37.9%) were
restricted to northern systems, 13
(44.8%) were restricted to the more
southern systems, and only 5 (17.2%)
slightly overlapped the two regions. In
the later and larger study by Wirgin et
al. (2009), the observed haplotypes
again clustered into regional groupings:
10 of 38 observed haplotypes (26.3%)
only occurred in systems north of the
Hudson River, 16 of 38 (42.1%) only
occurred in systems south of the
Chesapeake Bay, and just 5 of 38
(13.2%) haplotypes overlapped in the
mid-Atlantic region. The limited sharing
of haplotypes between the north and
south regions is consistent with strong
female homing fidelity and limited gene
flow between these regions. The break
in shared haplotypes corresponds with
the historical division of the species due
to Pleistocene glaciation, which
Grunwald et al. (2002) stated was
probably the most significant event
affecting population structure and
patterns of mtDNA diversity in
shortnose sturgeon.
The recent nDNA analyses of King et
al. (2014) also indicate an unambiguous
differentiation of sample populations
into one of three major geographic
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65193
groupings—Northeast, Mid-Atlantic, or
Southeast. When all 17 sample
populations were pooled by these three
geographic regions, correct assignment
to each region was 99.1% for the
Northeast and 100% (i.e., zero miassigned fish) for the remaining two
regions (King et al. 2014). Of the 133
fish included for the Northeast group,
one was mis-assigned to the MidAtlantic. The estimates of effective
migrants per generation (based on Phi
PT) are consistent with the regional
zones of genetic discontinuity among
Northeast, Mid-Atlantic, and Southeast
river systems. The average migrants per
generation between regions ranged from
less than one migrant (i.e., 0.89)
between Northeast and Southeast to
nearly two migrants (i.e., 1.89) between
Northeast and Mid-Atlantic. In contrast,
the range of estimated migrants per
generation within regions was 2.25–
83.08 for the Northeast, 1.87–13.64 for
the Mid-Atlantic, and 2.38–49.75 for the
Southeast (King et al. 2014). The
estimated migrants per generation
between the Saint John River in
particular and all other rivers within the
Northeast ranged from 2.25–3.43 (King
et al. 2014). Taken together, these data
indicate that the degree of genetic
differentiation between the Saint John
River and the rivers within the Gulf of
Maine is relatively small or ‘‘shallow’’,
especially relative to the deeper
divergence observed among the regional
groupings of river populations. A
possible explanation for the relatively
low level of differentiation within the
Northeast is that the those populations
are relatively young in a geologic sense
due to recent glaciations compared to
populations in the more southern part of
the range (SSRT 2010).
In conclusion, given the patterns of
genetic diversity, shared haplotypes,
and relative magnitudes of genetic
divergence at the river drainage versus
regional scale, we find there is
insufficient evidence that the Saint John
River population of shortnose sturgeon
differs markedly in its genetic
characteristics relative to the taxon as a
whole so as to meet the test for
‘‘significance’’ on this basis. While the
Saint John River population segment
can be genetically distinguished from
other river populations, available
genetic evidence places it into a larger
evolutionarily meaningful unit, along
with several other river populations
sampled. The degree of differentiation
among the three larger regional groups
is more marked than the differences
observed among populations from the
Saint John and other nearest rivers,
suggesting that the Saint John River
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population’s differentiation is not
‘‘significant’’ in the context of the whole
species. Gene flow estimates are also
consistent with the observed deeper
zones of divergence detected at the
regional scale. Thus, we conclude that
these data do not support delineation of
the Saint John River population segment
as ‘‘significant.’’ In so interpreting the
available genetic data, we are mindful of
the Congressional guidance to use the
DPS designation sparingly.
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DPS Conclusion and Petition Finding
We conclude that the Saint John River
population of shortnose sturgeon is
‘‘discrete’’ based on evidence that it is
a relatively closed and somewhat
geographically isolated population
segment. It thus satisfies the first prong
of the DPS policy. However, we also
find that the Saint John River
population segment is not ‘‘significant’’
to the taxon as a whole. It thus fails to
satisfy the second prong of the DPS
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Policy. As such, based on the best
available data, we conclude that the
Saint John River population of
shortnose sturgeon does not constitute a
DPS and, thus, does not qualify as a
‘‘species’’ under the ESA. Therefore, we
deny the petition to consider this DPS
for delisting. Our denial of the petition
on this ground does not imply any
finding as to how we should proceed if
the situation were otherwise, i.e., where
a population is found instead to meet
the criteria to be a DPS. Even if the
population had met both criteria of the
DPS Policy, and even if the population
were also found to have a status that
differed from the listed entity, it would
not necessarily be appropriate to
propose modifications to the current
listing, in light of the unsettled legal
issues surrounding such revisions. Nor
do we resolve here what steps would
need to be followed to propose revisions
to the species’ listing if the facts had
been otherwise; such an inquiry would
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be hypothetical in this case. It is clear
that because the petition at issue here
sought identification of a DPS, and
because the population at issue is not a
DPS, this particular petition must be
denied. As this is a final action, we do
not solicit comments on it.
References Cited
A complete list of references is
available upon request to the Office of
Protected Resources (see ADDRESSES).
Authority
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Dated: October 20, 2015.
Samuel D. Rauch III,
Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2015–27148 Filed 10–23–15; 8:45 am]
BILLING CODE 3510–22–P
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[Federal Register Volume 80, Number 206 (Monday, October 26, 2015)]
[Proposed Rules]
[Pages 65183-65194]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2015-27148]
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DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric Administration
50 CFR Part 224
[Docket No. 150209121-5941-02]
RIN 0648-XD760
Endangered and Threatened Wildlife; 12-Month Finding on a
Petition To Identify and Delist a Saint John River Distinct Population
Segment of Shortnose Sturgeon Under the Endangered Species Act
AGENCY: National Marine Fisheries Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA), Department of Commerce.
[[Page 65184]]
ACTION: Notice of 12-month petition finding.
-----------------------------------------------------------------------
SUMMARY: We, NMFS, announce a 12-month finding on a petition to
identify and ``delist'' shortnose sturgeon (Acipenser brevirostrum)
within the Saint John River in New Brunswick, Canada under the
Endangered Species Act (ESA). The shortnose sturgeon is currently
listed as an endangered species, at the species level, under the ESA.
Based on our review of the best scientific and commercial data
available, we have determined that the population of shortnose sturgeon
from the Saint John River does not qualify as a distinct population
segment. Therefore, we did not consider the petition further, and we do
not propose to delist this population.
DATES: This finding was made on October 26, 2015.
ADDRESSES: Information used to make this finding is available for
public inspection by appointment during normal business hours at NMFS,
Office of Protected Resources, 1315 East-West Highway, Silver Spring,
MD 20910. The petition and the list of the references used in making
this finding are also available on the NMFS Web site at:
www.nmfs.noaa.gov/pr/species/fish/shortnose-sturgeon.html.
FOR FURTHER INFORMATION CONTACT: Lisa Manning, Office of Protected
Resources, 301-427-8466; Stephania Bolden, Southeast Regional Office,
727-824-5312; Julie Crocker, Greater Atlantic Regional Office, 978-282-
8480.
SUPPLEMENTARY INFORMATION:
Background
On September 24, 2014, we received a petition from Dr. Michael J.
Dadswell, Dr. Matthew K. Litvak, and Mr. Jonathan Barry regarding the
population of shortnose sturgeon (Acipenser brevirostrum) native to the
Saint John River in New Brunswick, Canada. The petition requests that
we identify the Saint John River population of shortnose sturgeon as a
distinct population segment (DPS) and contemporaneously ``delist'' this
DPS by removing it from the species-wide listing under the Endangered
Species Act. On April 6, 2015, we published a positive finding
indicating that the petitioned action may be warranted and that we were
initiating a status review to consider the petition further (80 FR
18347).
The shortnose sturgeon was originally listed as an endangered
species throughout its range by the U.S. Fish and Wildlife Service
(USFWS) on March 11, 1967, under the Endangered Species Preservation
Act (ESPA, 32 FR 4001). Shortnose sturgeon remained on the endangered
species list when the U.S. Congress replaced the ESPA by enacting the
Endangered Species Conservation Act of 1969, which was in turn replaced
by the Endangered Species Act of 1973 (ESA, 16 U.S.C. 1531 et seq.). We
subsequently assumed jurisdiction for shortnose sturgeon under a 1974
government reorganization plan (39 FR 41370, November 27, 1974). In
Canada, the shortnose sturgeon falls under the jurisdiction of the
Department of Fisheries and Oceans (DFO) and was first assessed by the
Committee on the Status of Endangered Wildlife in Canada (COSEWIC) as
``Special Concern'' in 1980. This status was reconfirmed in 2005, and
the species was listed as Special Concern under the Canadian federal
Species at Risk Act (SARA) in 2009. The Special Concern status was
reconfirmed again in 2015 (COSEWIC, In Press). Shortnose sturgeon is
also listed under Appendix I of the Convention on International Trade
in Endangered Species of Wild Fauna (CITES).
Statutory, Regulatory and Policy Provisions
We are responsible for determining whether species are threatened
or endangered under the ESA (16 U.S.C. 1531 et seq.). To make this
determination, we first consider whether a group of organisms
constitutes a ``species'' under section 3 of the ESA, and then we
consider whether the status of the species qualifies it for listing as
either threatened or endangered. Section 3 of the ESA defines a
``species'' 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'' (16 U.S.C. 1532(16)). A
joint policy issued by NMFS and the U.S. Fish and Wildlife Service
(USFWS; collectively referred to as ``the Services'') clarifies the
interpretation of the phrase ``distinct population segment'' (DPS) for
the purposes of listing, delisting, and reclassifying a species under
the ESA (``DPS Policy,'' 61 FR 4722, February 7, 1996). The DPS Policy
identifies two criteria for determining whether a population is a DPS:
(1) The population must be ``discrete'' in relation to the remainder of
the taxon (species or subspecies) to which it belongs; and (2) the
population must be ``significant'' to the remainder of the taxon to
which it belongs.
Congress has instructed the Secretary to exercise the authority to
recognize DPS's ``sparingly and only when the biological evidence
indicates that such action is warranted'' (S. Rep. 96-151 (1979)). The
law is not settled as to the extent of the Services' discretion to
modify a species-level listing to recognize a DPS having a status that
differs from the original listing. In a recent decision, the United
States District Court for the District of Columbia held that the ESA
does not permit identification of a DPS solely for purposes of
delisting. Humane Soc'y v. Jewell, 76 F. Supp. 3d 69 (D.D.C. Dec. 19,
2014), appeal docketed, No. 15-5041 (D.C. Cir. Feb. 19, 2015) (Western
Great Lakes gray wolves) (consolidated with Nos. 15-5043, 15-5060, and
15-5061).
A species, subspecies, or DPS is ``endangered'' if it is in danger
of extinction throughout all or a significant portion of its range, and
``threatened'' if it is likely to become endangered within the
foreseeable future throughout all or a significant portion of its range
(ESA sections 3(6) and 3(20), respectively, 16 U.S.C. 1532(6) and
(20)). We interpret an ``endangered species'' to be one that is
presently in danger of extinction. A ``threatened species,'' on the
other hand, is not presently in danger of extinction, but is likely to
become so in the foreseeable future. In other words, the primary
statutory difference between a threatened and endangered species is the
timing of when a species may be in danger of extinction, either
presently (endangered) or in the foreseeable future (threatened). In
addition, we interpret ``foreseeable future'' as the horizon over which
predictions about the conservation status of the species can be
reasonably relied upon.
Pursuant to the ESA and our implementing regulations, the
determination of whether a species is threatened or endangered shall be
based on any one or a combination of the following five section 4(a)(1)
factors: The present or threatened destruction, modification, or
curtailment of habitat or range; overutilization for commercial,
recreational, scientific, or educational purposes; disease or
predation; inadequacy of existing regulatory mechanisms; and any other
natural or manmade factors affecting the species' existence. 16 U.S.C.
1533(a)(1); 50 CFR 424.11(c). Listing determinations must be based
solely on the best scientific and commercial data available, after
conducting a review of the species' status and after taking into
account any efforts being made by any state or foreign nation (or any
political subdivision of such state or foreign nation) to protect the
species. 16 U.S.C. 1532(b)(1)(A).
[[Page 65185]]
Under section 4(a)(1) of the ESA and the implementing regulations
at 50 CFR 424.11(d), a species shall be removed from the list if the
Secretary of Commerce determines, based on the best scientific and
commercial data available after conducting a review of the species'
status, that the species is no longer threatened or endangered because
of one or a combination of the section 4(a)(1) factors. The regulations
provide that a species listed under the ESA may be delisted only if
such data substantiate that it is neither endangered nor threatened for
one or more of the following reasons:
(1) Extinction. Unless all individuals of the listed species had
been previously identified and located, and were later found to be
extirpated from their previous range, a sufficient period of time
must be allowed before delisting to indicate clearly that the
species is extinct.
(2) Recovery. The principal goal of the USFWS and NMFS is to
return listed species to a point at which protection under the ESA
is no longer required. A species may be delisted on the basis of
recovery only if the best scientific and commercial data available
indicate that it is no longer endangered or threatened.
(3) Original data for classification in error. Subsequent
investigations may show that the best scientific or commercial data
available when the species was listed, or the interpretation of such
data, were in error.
50 CFR 424.11(d).
To complete the required finding in response to the current
delisting petition, we first evaluated whether the petitioned entity
meets the criteria of the DPS Policy. As we noted in our initial
petition finding, a determination whether to revise a species-level
listing to recognize one or more DPSs in place of a species-level
listing involves, first, determining whether particular DPS(s) exist(s)
(based on meeting the criteria of the DPS Policy) and, if that finding
is affirmative, complex evaluation as to the most appropriate approach
for managing the species in light of the purposes and authorities under
the ESA.
Species Description
Below, we summarize basic life history information for shortnose
sturgeon. A more thorough discussion of all life stages, reproductive
biology, habitat use, abundance estimates and threats are provided in
the Shortnose Sturgeon Biological Assessment completed by the Shortnose
Sturgeon Status Review Team in 2010 (SSRT 2010; https://www.fisheries.noaa.gov/pr/species/fish/shortnose-sturgeon.html).
There are 25 species and four recognized genera of sturgeons
(family Acipenseridae), which comprise an ancient and distinctive
assemblage with fossils dating to at least the Upper Cretaceous period,
more than 66 million years ago (Findeis 1997). The shortnose sturgeon,
Acipenser brevirostrum, is the smallest of the three extant sturgeon
species in eastern North America. Many primitive physical
characteristics that reflect the shortnose sturgeon's ancient lineage
have been retained, including a protective armor of bony plates called
``scutes''; a subterminal, protractile tube-like mouth; and
chemosensory barbels. The general body shape is cylindrical, tapering
at the head and caudal peduncle, and the upper lobe of the tail is
longer than the lower lobe. Shortnose sturgeon vary in color but are
generally dark brown to olive or black on the dorsal surface, lighter
along the row of lateral scutes, and nearly white on the ventral
surface. Adults have no teeth but possess bony plates in the esophagus
that are used to crush hard prey items (Vladykov and Greeley 1963;
Gilbert 1989). The skeleton is almost entirely cartilaginous with the
exception of some bones in the skull, jaw and pectoral girdle.
Shortnose sturgeon occur along the East Coast of North America in
rivers, estuaries, and marine waters. Historically, they were present
in most major rivers systems along the Atlantic coast (Kynard 1997).
Their current riverine distribution extends from the Saint John River,
New Brunswick, Canada, to possibly as far south as the St. Johns River,
Florida (Figure 1; Kynard 1997; Gorham and McAllister 1974). Recently
available information indicates that their marine range extends farther
northward than previously thought and includes the Minas Basin, Nova
Scotia (Dadswell et al. 2013). The distribution of shortnose sturgeon
across their range, however, is disjunct, with no known reproducing
populations occurring within the roughly 400 km of coast between the
Chesapeake Bay and the southern boundary of North Carolina. Shortnose
sturgeon live in close proximity with Atlantic sturgeon (Acipenser
oxyrinchus oxyrinchus) throughout much of their range. However,
Atlantic sturgeon spend more of their life cycle in the open ocean
compared to shortnose sturgeon. Within rivers, shortnose sturgeon and
Atlantic sturgeon may share foraging habitat and resources, but
shortnose sturgeon generally spawn farther upriver and earlier than
Atlantic sturgeon (Kynard 1997, Bain 1997).
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Shortnose sturgeon typically migrate seasonally between upstream
freshwater spawning habitats and downstream foraging mesohaline (i.e.,
salinities of 5 to 18 parts per thousand) habitat based on water
temperature, flow, and salinity cues. Based on their varied and complex
use of freshwater, estuarine, and marine waters, shortnose sturgeon
have been characterized in the literature as ``anadromous'' or
``amphidromous'' (Bain 1977; Kieffer and Kynard 1993). An anadromous
species is defined as one that spawns in freshwater and spends much of
its life cycle in marine waters, whereas a freshwater amphidromous
species is one that spawns and remains in freshwater for most of its
life cycle but spends some time in saline water. Because shortnose
sturgeon had historically rarely been detected far from their natal
estuary, they were once considered to be largely confined to their
natal rivers and estuaries (NMFS 1998). However, more recent research
has demonstrated that shortnose sturgeon leave their natal estuaries,
undergo coastal migrations, and use other river systems to a greater
extent than previously thought (Kynard 1997; Savoy 2004; Fernandes
2010; Zydlewski et al. 2011; Dionne et al. 2013). The reasons for
inter-riverine movements are not yet clear, and the degree to which
this behavior occurs appears to vary among river systems.
Shortnose sturgeon are benthic feeders, and their diet typically
consists of small insects, crustaceans, mollusks, polychaetes, and
small benthic fishes (McCleave et al. 1977; Dadswell 1979;
[[Page 65187]]
Marchette and Smiley 1982; Dadswell et al. 1984; Moser and Ross 1995;
Kynard et al. 2000; Collins et al. 2002). Both juvenile and adult
shortnose sturgeon primarily forage over sandy-mud bottoms, which
support benthic invertebrates (Carlson and Simpson 1987, Kynard 1997).
Shortnose sturgeon have also been observed feeding off plant surfaces
(Dadswell et al. 1984). Sturgeon likely use electroreception,
olfaction, and tactile chemosensory cues to forage, while vision is
thought to play a minor role (Miller 2004).
Foraging in the colder rivers in the northern part of their range
appears to greatly decline or cease during winter months when shortnose
sturgeon generally become inactive. In mid-Atlantic areas, including
the Chesapeake Bay, and the Delaware River, foraging is believed to
occur year-round, though shortnose sturgeon are believed to feed less
in the winter (J. O'Herron, Amitrone O'Herron, Inc., pers. comm. 2008
as cited in SSRT 2010). In the southern part of their range, shortnose
sturgeon are known to forage widely throughout the estuary during the
winter, fall, and spring (Collins and Smith 1993, Weber et al. 1999).
During the hotter months of summer, foraging may taper off or cease as
shortnose sturgeon take refuge from high water temperatures.
Shortnose sturgeon are relatively small compared to most extant
sturgeon species and reach a maximum length of about 120 cm total
length (TL) and weight of about 24 kg (Dadswell 1979; Waldman et al.
2002); however, both maximum size and growth rate display a pattern of
gradual variation across the range, with the fastest growth rates and
smallest maximum sizes occurring in the more southern populations
(Dadswell et al. 1984). The northernmost populations exhibit the
slowest growth and largest adult sizes. The largest shortnose sturgeon
reported in the published literature to date was collected from the
Saint John River, Canada, and measured 143cm TL (122 cm fork length
(FL)) and weighed 23.6 kg (Dadswell 1979). In contrast, in their
review, Dadswell et al. (1984) indicated that the largest adult
reported from the St. Johns River, Florida, was a 73.5 cm (TL) female.
Dadswell et al. (1984) compared reported growth parameters across the
range and showed that the von Bertalanffy growth parameter K and
estimated asymptotic length ranged from 0.042 and 130.0 cm (FL),
respectively, for Saint John River fish to 0.149 and 97.0 cm (FL) for
Altamaha River, Georgia fish. However, the land-locked shortnose
sturgeon population located upstream of Holyoke Dam at river km 140 of
the Connecticut River has the slowest adult growth rate of any
surveyed, which may at least in part reflect food limitations (Taubert
1980a).
Shortnose sturgeon are relatively long-lived and slow to mature.
The oldest shortnose sturgeon reported was a 67 year-old female from
the Saint John River, and the oldest male reported was a 32 year-old
fish, also captured in the Saint John River (Dadswell 1979). In
general, fish in the northern portion of the species' range live longer
than individuals in the southern portion of the species' range (Gilbert
1989). Males and females mature at about the same length, around 45-55
cm FL, throughout their range (Dadswell et al. 1984). However, age at
maturity varies by sex and with latitude, with males in the southern
rivers displaying the youngest ages at maturity (see review in Dadswell
et al. 1984). For example, age at first maturation in males occurs at
about 2-3 years of age in Georgia and at about 10-11 years in the Saint
John River. Females mature by 6 years of age in Georgia and at about 13
years in the Saint John River (Dadswell et al. 1984).
Sturgeon are iteroparous, meaning they reproduce more than once
during their lifetime. In general, male shortnose sturgeon are thought
to spawn every other year, but they may spawn annually in some rivers
(Dovel et al. 1992; Kieffer and Kynard 1996). Females appear to spawn
less frequently--approximately every 3 to 5 years (Dadswell 1979).
Spawning typically occurs during late winter/early spring (southern
rivers) and mid-to-late spring (northern rivers) (Dadswell 1979,
Taubert 1980a and b, Kynard 1997). The onset of spawning may be cued by
decreasing river discharge following the peak spring freshet, when
water temperatures range from 8 to 15 [deg]C and bottom water
velocities range between 25-130 cm/s, although photoperiod (or day-
length) appears to control spawning readiness (Dadswell et al. 1984;
Kynard et al. 2012). Spawning appears to occur in the sturgeons' natal
river, often just below the fall line at the farthest accessible
upstream reach of the river (Dovel 1981; Buckley and Kynard 1985;
Kieffer and Kynard 1993; O`Herron et al. 1993; Kieffer and Kynard
1996). Following spawning, adult shortnose sturgeon disperse quickly
down river and typically remain downstream of their spawning areas
throughout the rest of the year (Buckley and Kynard 1985, Dadswell et
al. 1984; Buckley and Kynard 1985; O'Herron et al. 1993).
In a review by Gilbert (1989), fecundity of shortnose sturgeon was
reported to range between approximately 30,000-200,000 eggs per female.
Shortnose sturgeon collected from the Saint John River had a range of
27,000-208,000 eggs and a mean of 11,568 eggs/kg body weight (Dadswell
1979). Development of the eggs and transition through the subsequent
larval, juvenile and sub-adult life stages are discussed in more detail
in SSRT 2010.
A total abundance estimate for shortnose sturgeon is not available.
However, population estimates, using a variety of techniques, have been
generated for many individual river systems. In general, northern
shortnose sturgeon population abundances are greater than southern
populations (Kynard 1997). The Hudson River shortnose sturgeon
population is currently considered to be the largest extant population
(61,000 adults, 95 percent CI: 52,898-72,191; Bain et al. 2007;
however, see discussion of this estimate in SSRT 2010). Available data
suggest that some populations in northern rivers have increased over
the past several decades (e.g., Hudson, Kennebec; Bain et al. 2000;
Squiers 2003) and that others may be stable (e.g., Delaware; Brundage
and O'Herron 2006). South of Chesapeake Bay, populations are relatively
small compared to the northern populations. The largest population of
shortnose sturgeon in the southern part of the range is from the
Altamaha River, which was most recently estimated at 6,320 fish (95%
CI: 4387-9249; Devries 2006). Occasional observations of shortnose
sturgeon have been made in some rivers where shortnose sturgeon are
considered extirpated (e.g., St. Johns, St. Mary's, Potomac,
Housatonic, and Neuse rivers); the few fish that have been observed in
these rivers are generally presumed to be immigrants from neighboring
basins.
The most recent total population estimate for the Saint John River
dates to the 1970's. Using tag recapture data from 1973-1977, Dadswell
(1979) calculated a Jolly-Seber population estimate of 18,000 (30% SE; 95 percent CI: 7,200-28,880, COSEWIC, In Press) adults (>
50 cm) below the Mactaquac Dam. Several partial population estimates
are also available for the Kennebecasis River, a tributary in the lower
reaches of the Saint John River. Litvak (unpublished data) calculated a
Jolly-Seber estimate of 2,068 fish (95% CI: 801-11,277) in the
Kennebecasis using mark-recapture data from 1998 to 2004 (COSEWIC, In
Press). Based on videotaping of overwintering aggregations of shortnose
sturgeon on the Kennebecasis River at the confluence of the Hammond
River (rkm 35), Li et al. (2007) used ordinary Kriging to estimate that
4,836 (95% CI:
[[Page 65188]]
4,701-4,971) adult shortnose sturgeon were overwintering in that area.
Usvyatsov et al. (2012) repeated this sampling in 2009 and 2011 and,
using three different modeling techniques, estimated a total of 3,852-
5,222 shortnose sturgeon in the study area, which suggests fairly
stable abundance and habitat use at this site.
Threats that contributed to the species' decline and led to the
listing of shortnose sturgeon under the ESA included pollution,
overfishing, and bycatch in the shad fishery (USDOI 1973). Shortnose
sturgeon were also thought to be extirpated, or nearly so, from most of
the rivers in their historical range (USDOI 1973). In the late
nineteenth and early twentieth centuries, shortnose sturgeon were
commonly harvested incidental to Atlantic sturgeon, the larger and more
commercially valuable of these two sympatric sturgeon species (NMFS
1998). Although there is currently no legal directed fishing for
shortnose sturgeon in the United States, poaching is suspected, and
bycatch still occurs in some areas. In particular, shortnose sturgeon
are caught incidentally by bass anglers and in the alewife/gaspereau,
American shad, American eel, and Atlantic sturgeon fisheries in the
Saint John River; and shad fisheries in the Altamaha River, Santee
River, Savannah River, and elsewhere (COSEWIC, In Press; SSRT 2010;
Bahn et al. 2009; COSEWIC 2005). The construction of dams has also
resulted in substantial loss of historical shortnose sturgeon habitat
in some areas along the Atlantic seaboard. The construction and
operation of dams can impede upstream movement to sturgeon spawning
habitat (e.g., Connecticut River, Santee River). Remediation measures,
such as dam removal or modification to allow for fish passage have
improved access in some rivers, and additional similar restoration
efforts are being considered in other areas (e.g., possible removal of
the Mactaquac dam in the Saint John River). Other possible and ongoing
threats include operation of power generating stations, water diversion
projects, dredging, and other in-water activities that impact habitat.
Distinct Population Segment Analysis
The following sections provide our analysis of whether the
petitioned entity--the Saint John River population of shortnose
sturgeon--qualifies as a DPS of shortnose sturgeon (whether it is both
``discrete'' and ``significant''). To complete this analysis we relied
on the best scientific and commercial data available and considered all
relevant literature and public comments submitted in response to our
90-day finding (80 FR 18347, April 6, 2015).
For purposes of this analysis, we defined the Saint John River
population segment of shortnose sturgeon to consist of shortnose
sturgeon spawned in the Saint John River downstream of the Mactaquac
Dam. Prior to construction of Mactaquac Dam in 1968/1969, sturgeon
occurred upstream of the dam; however, it is unclear whether these were
shortnose and/or Atlantic sturgeon and whether any sturgeon are still
present upstream of the dam (COSEWIC, In Press). Lacking this
information, we cannot consider fish that may be present upstream of
the dam in our distinct population segment analysis. Throughout our
discussion below we also use the term ``population'' to refer
collectively to all shortnose sturgeon that are presumed to be natal to
a particular river rather than using this term to refer strictly to a
completely closed reproductive unit.
Discreteness Criterion
The Services' joint DPS Policy states that a population segment 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.
(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 (61 FR 4722,
February 7, 1996).
There are no physical barriers preventing the movement of Saint
John River shortnose sturgeon outside of the Saint John River estuary
or along the coast. The Mactaquac Dam, located about 140 km upstream
and at the head of tide (Canadian Rivers Institute 2011), is the first
upstream physical barrier on the Saint John River. This and other dams
on the Saint John River block shortnose sturgeon from accessing
upstream habitats, but there are no dams or other physical barriers
separating Saint John River sturgeon from other shortnose sturgeon
populations.
As mentioned previously, shortnose sturgeon have been documented to
leave their natal river/estuary and move to other rivers to varying
extents across their range. For example, telemetry data generated by
Zydlewski et al. (2011) during 2008-2010 indicate that inter-riverine
movements of adult shortnose sturgeon occur fairly frequently among
rivers in Maine. Seventy percent of tagged adults (25 of 41 fish) moved
between the Penobscot and Kennebec rivers (about 150 km away), and up
to 52% of the coastal migrants (13 of 25 fish) also used other, smaller
river systems (i.e., Damariscotta, Medomak, St. George) between the
Penobscot and Kennebec rivers (Zydlewski et al. 2011). Shortnose
sturgeon are also known to move between rivers in Maine and (e.g.,
Kennebec, Saco) and the Merrimack River estuary in Massachusetts,
traveling distances of up to about 250 km (as measured by a
conservative, direct path distance; Little et al. 2013; Wippelhauser et
al. 2015). At the other end of the range, in the Southeast United
States, inter-riverine movements appear fairly common and include
movements between the Savannah River and Winyah Bay and between the
Altamaha and Ogeechee rivers (Peterson and Farrae 2011; Post et al.
2014).
Many inter-riverine movements have been observed elsewhere within
the species' range, but patterns are not yet well resolved. For
example, some shortnose sturgeon captured and/or tagged in the
Connecticut River have been recaptured, detected, or were previously
tagged in the Housatonic River (T. Savoy, CT DEP, pers. comm. 2015),
the Hudson River (Savoy 2004), and the Merrimack River (M. Kieffer,
USGS, pers. comm. 2015). At this time, the available tagging and
tracking information is too limited to determine if Hudson River and
Connecticut River shortnose sturgeon are making regular movements
outside of their natal rivers and whether movement as far as the
Merrimack River is a normal behavior. Movement data from the Chesapeake
Bay is also relatively limited, but existing data indicate that
shortnose sturgeon do move from the Chesapeake Bay through the
Chesapeake and Delaware Canal into the Delaware River (Welsh et al.
2002).
The distances of the reported marine migrations vary widely from
very short distances--such as between the Santee River and Winyah Bay,
which are only about 15 km apart--to fairly long--as in the case of
movements between the Merrimack and the Penobscot rivers, which are
about 339 km apart at their mouths.\1\ In general, the available data
[[Page 65189]]
suggest that movements between geographically proximate rivers are more
common, while movements between more distant rivers do not, or only
rarely, occur. A detailed discussion of the physical movements of
shortnose sturgeon is provided in SSRT 2010.
---------------------------------------------------------------------------
\1\ Distances between rivers mouths reported here were measured
in GIS using the NOAA Medium Resolution Vector Shoreline, 20m
bathymetry contour, and a fixed scale of 1:250,000. Estimated
distances reported are the average of three, independently drawn and
measured paths for each river pair. The assumed travel path between
river mouths was the shortest possible distance that followed the
general outline of the coast and was constrained by the 20m
bathymetry contour, except where the shortest travel path across a
deep, narrow inlet or bay crossed the 20m bathymetry contour.
---------------------------------------------------------------------------
The extent of coastal movements of shortnose sturgeon from the
Saint John River is currently unknown (COSEWIC, In Press); however,
some limited data are available and provide some insight into whether
these fish may be geographically isolated from other populations. Any
movement between Saint John River sturgeon and the nearest population
in the Penobscot River would require a marine migration of about 362
km, a similar travel distance as between the Merrimack and the
Penobscot rivers (340 km) and between the Connecticut and Merrimack
rivers (348 km).\2\ Dadswell (1979) reported that of 121 marked Saint
John River shortnose sturgeon recaptured by commercial fisherman, 13
fish (11 percent) were recaptured in the Bay of Fundy, indicating that
a portion of the population migrated into the marine environment. In
addition, a confirmed shortnose sturgeon was caught in a fishing weir
in the Minas Basin, off the coast of Nova Scotia about 165 km north of
the mouth of the Saint John River (Dadswell et al. 2013). Fishermen in
the Minas Basin also claim to catch about one to two shortnose sturgeon
per year in their weirs (Dadswell et al. 2013). While it is plausible
that the shortnose sturgeon captured in the Minas Bay originated from
the Saint John River, data to confirm this are not available. In
contrast, limited telemetry data suggest that movements outside of the
Saint John River are not common. Of 64 shortnose sturgeon tagged in the
Saint John River over the course of about 16 years from 1999 to 2015,
none have been detected moving past the farthest downriver acoustic
receiver located near the Saint John Harbor Bridge (M. Litvak, pers.
comm. July 31, 2015).
---------------------------------------------------------------------------
\2\ Distances reported here were measured following the same
methods described in the previous footnote. The distance reported
between the Connecticut and Merrimack River assumes a travel path
via the Cape Cod Canal. A travel path around Cape Cod would instead
result in a marine migration of about 560 km.
---------------------------------------------------------------------------
Overall, while there is unambiguous evidence that shortnose
sturgeon from the Saint John River leave the estuary--at least
occasionally--and use the marine environment, and that shortnose
sturgeon are capable of making long distance movements between river
systems, there are no available data on coastal migrations of Saint
John River shortnose sturgeon. To date, there are also no reported
observations or detections of shortnose sturgeon from the Gulf of Maine
rivers moving into the Saint John River. Thus, while it is possible
that the Saint John River shortnose sturgeon come in contact with
shortnose sturgeon from elsewhere, it is also likely that some degree
of geographical isolation by distance is occurring.
Although acoustic telemetry studies have revealed that shortnose
sturgeon leave their natal river systems to a much greater extent than
previously thought, such movements do not necessarily constitute
permanent emigration or indicate interbreeding of populations. Tagging
and telemetry studies within several river systems have provided
evidence that shortnose sturgeon in those particular systems tend to
spawn in their natal river (e.g., Dovel 1981; Buckley and Kynard 1985;
Kieffer and Kynard 1993; O`Herron et al. 1993; Kieffer and Kynard
1996). Tag return data for shortnose sturgeon in the Saint John River
over the course of a 4-year study completed by Dadswell (1979) suggests
there is little emigration from this system as well, and that spawning
takes place in the freshwater sections of the upper estuary. The high
site fidelity to natal rivers suggested by this and other studies
indicates a there is a possible behavioral mechanism for the marked
separation of the Saint John River population of shortnose sturgeon
from other populations of the species.
A substantial amount of genetic data has become available since the
``Final Recovery Plan for Shortnose Sturgeon'' was developed in 1998.
Below, we summarize the best available genetic data and information,
which informed our evaluation of the ``discreteness'' of the Saint John
River population segment. A more in-depth presentation of genetic data,
including discussions of types of analyses and assumptions, is
available in the Biological Assessment (SSRT 2010).
Much of the published information on population structure for
shortnose sturgeon has been based on the genetic analysis of the
maternally inherited mitochondrial DNA (mtDNA) due in part to the
difficulties of analyzing data from the polyploid nuclear genome
(Waldman et al. 2008). The analyses have focused on a moderately
polymorphic 440 base pair portion of the mtDNA control region--a
relatively rapidly evolving region of mtDNA and thus a good indicator
of population-level differentiation. Haplotype frequencies and sequence
divergence data have consistently indicated an overall isolation-by-
distance pattern of genetic population structure across the species'
range, meaning that populations of shortnose sturgeon inhabiting rivers
and embayments that are geographically more distant tend to be less
related than those that are geographically closer (e.g., Walsh et al.
2001, Grunwald et al. 2002, Waldman et al. 2002, and Wirgin et al.
2005; Wirgin et al. 2009). The haplotypes observed are typically shared
across two to four or more adjacent sampled rivers but with little
sharing of haplotypes between northern and southern populations
(Waldman et al. 2002; Wirgin et al. 2009). Results for the Saint John
River are compatible with these general patterns. For example, in the
largest study to date, Wirgin et al. (2009) observed eight haplotypes
within the Saint John River sample (n=42); and of the eight observed
haplotypes, one was exclusive (or ``private'') to the Saint John River
(and observed in 1 of 42 fish), and the remaining haplotypes were
shared with two to six other rivers. None of the shared haplotypes were
observed in samples south of the Chesapeake Bay. A previously
unreported haplotype was recently observed in 2 of 15 shortnose caught
from the Kennebecasis River, a tributary of the Saint John (Kerr, 2015;
P. Wilson, public comment, May 2015). This new haplotype could indicate
an even greater degree of differentiation of the Saint John River fish;
however, no other rivers were sampled or analyzed as part of this
study.
Despite the localized sharing of haplotypes, frequencies of the
observed haplotypes are significantly different in most pairwise
comparisons of the rivers sampled (i.e., comparisons of haplotype
frequencies from samples from two rivers), including many adjacent
rivers (Wirgin et al. 2009). Such pairwise comparisons for the Saint
John River in particular have indicated that this population is
genetically distinct from the geographically closest sampled
populations, including the Penobscot, Kennebec, and Androscoggin rivers
(Grunwald et al. 2002; Waldman et al. 2002; Wirgin et al. 2005; Wirgin
et al. 2009). For example, Wirgin et al. (2009) reported significant
differences (p<0.0005) in haplotype frequencies between Saint John
River shortnose sturgeon (n=42) and Penobscot (n=44, Chi-square=37.22),
Kennebec (n=54, Chi-square=54.85), and Androscoggin (n=48, Chi-
square=37.91) river samples. The level of genetic differentiation
between the Saint John River population and the Penobscot, Kennebec,
and
[[Page 65190]]
Androscoggin rivers also appears substantial, with Phi ST
values ranging from 0.213 to 0.291 (where Phi ST ranges from
0 to 1, with 1 indicating complete isolation; Wirgin et al. 2009).
Estimates of female-mediated gene flow between the Saint John River
and the Gulf of Maine rivers are fairly low. Wirgin et al. (2009)
estimated female-mediated gene flow between the Saint John River and
other Gulf of Maine rivers as 1.90-2.85 female migrants per generation
based on Phi ST values, and as 1.5-1.9 females per
generation in a separate, coalescent-based analysis. This result
suggests that (if model assumptions are true) no more than three female
shortnose sturgeon from the Saint John River are likely to spawn in the
other Gulf of Maine rivers (or vice versa) per generation. These
results provide additional evidence that the degree of female-based
reproductive exchange between the Saint John River population and other
nearby shortnose river populations has been relatively limited over
many generations.
More recently, King et al. (2014) completed a series of analyses
using nuclear DNA (nDNA) samples from 17 extant shortnose sturgeon
populations across the species range. In contrast to the maternally
inherited mtDNA, nDNA reflects the genetic inheritance from both the
male and female parents. King et al. (2014) surveyed the samples at 11
polysomic microsatellite DNA loci and then evaluated the 181 observed
alleles as presence/absence data using a variety of analytical
techniques. The population structuring revealed by these analyses is
consistent with the previous mtDNA analyses in that they also indicate
a regional scale isolation-by-distance pattern of genetic
differentiation. Analysis of genetic distances among individual fish
(using principle coordinate analysis, PCO) revealed that the sampled
fish grouped into one of three major geographic units: (1) Northeast,
which included samples from the Saint John, Penobscot, Kennebec,
Androscoggin, and Merrimack rivers; (2) Mid-Atlantic, which included
samples from the Connecticut, Hudson, and Delaware rivers, as well as
the Chesapeake Bay proper; and (3) Southeast, which included samples
from the Cape Fear River, Winyah Bay, the Santee-Cooper, Edisto,
Savannah, Ogeechee, and Altamaha rivers, and Lake Marion (King et al.
2014).
Subsequent analyses revealed that each of the three regions has a
different pattern of sub-structuring. Within the Northeast group, two
separate analyses (PCO and STRUCTURE) indicated a high degree of
relatedness and possible panmixia (i.e., random mating of individuals)
among the Penobscot, Kennebec, and Androscoggin rivers; whereas, the
Saint John and Merrimack rivers appeared more differentiated from each
other as well as from the other Gulf of Maine rivers (King et al.
2014). Pairwise comparisons at the population level showed that, within
the Northeast region, estimates of genetic differentiation were
greatest between the Saint John and Merrimack rivers (Phi PT
= 0.100, p <0.0004), the two most distant rivers within this region.
Pairwise comparisons of the Saint John River to the remaining rivers
within the Northeast region revealed lower but still statistically
significant levels of genetic differentiation (Phi PT =
0.068-0.077; King et al. 2014). Relatively low levels of
differentiation were observed in pairwise comparisons for all other
rivers within the Northeast region (Phi PT = 0.013-0.087),
half of which were not statistically significant (King et al. 2014). In
comparison, within the Mid-Atlantic group, pairwise comparisons among
rivers showed moderate levels of genetic differentiation among most
river populations (average Phi PT = 0.077, range = 0.018-
0.118); whereas, estimates of population level genetic differentiation
were very low among samples populations in the Southeast group (average
Phi PT = 0.047, range = 0.005 to 0.095; King et al. 2014),
suggesting a more genetically similar set of populations.
Theoretical estimates of gene flow (derived from Phi PT
values) between the Saint John River and the other Northeast rivers
ranged from 2.25 to 3.43 migrants per generation (King et al. 2014).
Gene flow estimates for the Merrimack River were similarly low, ranging
from 2.25 to 4.06 (King et al. 2014). In contrast, the effective number
of migrants per generation estimated to occur between the remaining
rivers within the Northeast region was much higher and ranged from
16.42 to 83.08 (King et al. 2014).
Overall, the analyses completed by King et al. (2014) indicate that
differentiation among Northeast populations is less than that observed
among the Mid-Atlantic populations and greater than that observed among
Southeast populations. However, within the Northeast region, both the
Saint John and Merrimack River sample populations are genetically
distinct from the other sample populations. Although the estimates of
gene flow suggest some connectivity between the Saint John and other
rivers within the Northeast, the significantly different allele and
haplotype frequencies shown consistently in the nDNA and mtDNA studies
provide indirect evidence that the Saint John River population is
relatively reproductively isolated.
As highlighted in the DPS Policy, quantitative measures of
morphological discontinuity or differentiation can serve as evidence of
marked separation of populations. We examined whether the morphological
data for shortnose sturgeon across its range provide evidence of marked
separation of the Saint John River population. As noted previously,
maximum adult size (length and weight) varies across the range, with
the largest maximum sizes occurring in the Saint John River at the
northernmost end of the range, and the smallest sizes occurring in
rivers at the southern end of the range (Dadswell et al. 1984). The
largest individual reported in the literature (122 cm FL, 23.6 kg) was
captured in the Saint John River, although there is also a report of a
specimen measuring 124.6 cm FL (M. Litvak, unpublished data, as cited
in COSEWIC, In Press). Lengths of shortnose sturgeon captured in
surveys of the Saint John River in 1974-1975 ranged from 60 to 120 cm
FL (n=1,621). The majority of these fish, however, were smaller than
100 cm FL (1,476 fish), and only six fish were longer than 111 cm FL
(Dadswell 1979). To the south, in the Kennebec River, Maine shortnose
sturgeon captured during 1980 and 1981 had lengths ranging from 58.5 to
103.0 cm FL, and averaging 80.8 cm FL (n=24; Walsh et al. 2001).
Smaller size ranges are reported for rivers in the southernmost portion
of the range with some occasional captures of larger specimens. For
example, adult shortnose sturgeon captured in the Altamaha River,
Georgia, in 2010-2013 ranged from 57.4-83.0 cm FL and averaged 70.1 cm
long (FL, n=40; Peterson 2014), but a shortnose sturgeon measuring
104.5 cm FL and weighing 8.94 kg was captured in the Altamaha River in
summer, 2004 (D. Peterson, UGA, unpubl. data). Overall, the attribute
of size appears to display clinal variation, meaning there is a gradual
change with geographic location (Huxley 1938). The fact that the Saint
John River population segment, which lies at the northernmost end of
the range, exhibits the largest sizes does not in itself constitute a
morphological discontinuity. Given the apparent gradual nature of the
variation in size with latitude, we find that there is no marked
separation of the Saint John River population segment on the basis of a
quantitative discontinuity in size.
In addition to body size, other attributes such as snout length,
head length, and mouth width can provide evidence of a morphological
[[Page 65191]]
discontinuity and were also considered. Walsh et al. (2001) examined
six morphological and five meristic attributes for shortnose sturgeon
in the Androscoggin, Kennebec, and Hudson rivers. All morphological
features measured (i.e., body length, snout length, head length, mouth
width, and interorbital width) were largest for the Kennebec River fish
and smallest for fish from the southern-most river in the study, the
Hudson River (Walsh et al. 2001). Meristic features (e.g., scute
counts) were similar for the three rivers and were not related to fish
size (Walsh et al. 2001). Overall, the degree of phenotypic
differentiation of fish from the two rivers in Maine (Androscoggin and
Kennebec), which share an estuary mouth, was very low, while a much
greater degree of differentiation was observed for the fish from the
Hudson River (Walsh et al. 2001). This result was congruent with
results of corresponding mtDNA analyses, which indicated that the
Hudson River had a much greater degree of genetic differentiation from,
and much lower rate of gene flow with, the two rivers in Maine (Walsh
et al. 2001). The results of this particular study suggest there could
be clinal variation in these other phenotypic characteristics, similar
to the pattern observed for body size. As far as we are aware, however,
similar studies have not yet been conducted to examine the variation in
additional sets of morphological attributes across the range of
shortnose sturgeon and relative to the Saint John River population in
particular. Therefore, there is no basis to conclude marked separation
of the Saint John River population segment on the basis of
morphological discontinuity.
In conclusion, although the currently available data do not show
that the Saint John River shortnose sturgeon constitute a completely
isolated or closed population, we find that available genetic data,
evidence of site fidelity, and the likelihood of some degree of
geographical isolation together constitute sufficient information to
indicate that the Saint John River shortnose sturgeon are markedly
separated from other populations of shortnose sturgeon. Thus, after
considering the best available data and all public comments submitted
in response to our initial petition finding, we conclude that the Saint
John River population segment of shortnose sturgeon is ``discrete.'' We
therefore proceeded to evaluate the best available data with respect to
the second criterion of the DPS Policy, ``significance.''
Significance Criterion
Under the DPS Policy, if a population segment is found to be
discrete, then we proceed to the next step of evaluating its biological
and ecological significance to the taxon to which it belongs. As we
explained above, a population must be both ``discrete'' (the first
prong of the DPS Policy) and ``significant'' (the second prong of the
DPS Policy) to qualify for recognition as a DPS.
Consideration of significance may include, but is not limited to:
(1) Persistence of the discrete population segment in an ecological
setting unusual or unique for the taxon; (2) evidence that the loss of
the discrete population segment would result in a significant gap in
the range of a taxon; (3) evidence that the discrete population segment
represents the only surviving natural occurrence of a taxon that may be
more abundant elsewhere as an introduced population outside its
historical range; and (4) evidence that the discrete population segment
differs markedly from other populations of the species in its genetic
characteristics (61 FR 4722, February 7, 1996). These four factors are
non-exclusive; other relevant factors may be considered in the
``significance'' analysis. Further, significance of the discrete
population segment is not necessarily determined by existence of one of
these classes of information standing alone. Rather, information
analyzed under these and any other applicable considerations is
evaluated relative to the biological and ecological importance of the
discrete population to the taxon as a whole. Accordingly, all relevant
and available biological and ecological information is analyzed to
determine whether, because of its particular characteristics, the
population is significant to the conservation of the taxon as a whole.
Persistence in an Ecological Setting Unusual or Unique for the Taxon
Shortnose sturgeon once occupied most major rivers systems along
the Atlantic coast of North America (Kynard 1997). Although extirpated
from some areas due mainly to overharvest, bycatch, pollution, and
habitat degradation, shortnose sturgeon still occur in at least 25
rivers systems within their historical range (NMFS 1998). Throughout
their current range, shortnose sturgeon occur in riverine, estuarine,
and marine habitats; and, as adults, generally move seasonally between
freshwater spawning habitat and downstream mesohaline and sometimes
coastal marine areas in response to cues such as water temperature,
flow, and salinity. Like other species of sturgeon (e.g. A.
transmontanus in the Columbia River, Oregon), shortnose sturgeon are
also capable of adopting a fully freshwater existence, as is the case
for the population of shortnose sturgeon above the Holyoke Dam in the
Connecticut River and in Lake Marion, South Carolina. While each river
system within the shortnose sturgeon's range is similar in terms of its
most basic features and functions, each river system differs to varying
degrees in terms of its specific, physical and biological attributes,
such as hydrologic regime, benthic substrates, water quality, and prey
communities. A few examples are discussed briefly below.
The Saint John River begins in northern Maine, United States,
travels through New Brunswick, Canada, and empties into the Bay of
Fundy within the northeast Gulf of Maine. The river is approximately
673 km long, fed by numerous tributaries, and has a large tidal estuary
and a basin area of over 55,000 km\2\ (Kidd et al. 2011). According to
the Nature Conservancy's (TNC) ecoregion classification system, the
Saint John River watershed lies within the New England-Acadian
(terrestrial), Northeast United States and Southeast Canada Atlantic
Drainages (freshwater), and the Gulf of Maine/Bay of Fundy (marine)
ecoregions. The mean annual discharge is approximately 1,100 m\3\/s,
dissolved oxygen levels average 8.5 to 11 mg/l, and benthic substrates
downstream of the Mataquac Dam consist largely of shifting sands (Kidd
et al. 2011). Due to the low slope of the lower reaches and the extreme
tidal range of the Bay of Fundy, the head of the tide can extend about
140 km upstream from the river mouth (Kidd et al. 2011). During the
shortnose sturgeon spring/summer spawning season, water temperatures
range from about 10 to 15 [deg]C; and within overwintering areas, water
temperature range between 0 and 13 C (Dadswell 1979; Dadswell et al.
1984). Shortnose sturgeon in the Saint John River appear to move to
deeper waters when surface water temperatures exceed 21 [deg]C
(Dadswell et al. 1984). Further to the south, but still within the same
terrestrial, freshwater, and marine TNC ecoregions as the Saint John
River, is the smaller Penobscot River system in Maine. This river is
175 km long (not including the West and South Branches), has a drainage
basin of 22,265 km\2\, and an annual average discharge of about 342
m\3\/s (Lake et al. 2012; USGS 2015). Benthic substrates, consisting of
bedrock, boulders, cobble and sand deposits are undergoing changes in
response to the removal of
[[Page 65192]]
two dams--Great Works Dam at rkm 60 and Veazie Dam at rkm 48--within
the past three years (FERC 2010; Cox et al. 2014). The Veazie Dam was
located close to the head of the tide, and although conditions have
since changed, Haefner (1967, as cited in Fernandes et al. 2010) stated
that, during peak springtime flows, freshwater extends to rkm 17, and
that the salt wedge intrudes as far as about rkm 42 when river
discharges decrease in summer. Water temperatures in shortnose sturgeon
overwintering areas in the Penobscot River range from about 0 [deg]C to
13.3 [deg]C, and the fish appear to move out of overwintering areas
when water temperatures reach about 2.4 [deg]C (Fernandes et al. 2010).
Towards the southern end of the range and occurring within a very
different set of ecoregions is the Altamaha River, which is formed by
the confluence of the Ocmulgee and Oconee rivers in Georgia. One of the
longest free-flowing systems on the Atlantic Coast, the Atlamaha River
is just over 220 km long, has a watershed area of about 37,300 km\2\,
and flows mainly eastward before emptying into the Atlantic Ocean (TNC
2005). Tidal influence extends up to about rkm 40 (DeVries 2006). The
average annual discharge is 381 m\3\/s, and benthic substrates consist
mostly of sands with very few rocky outcrops (Heidt and Gilbert 1979;
DeVries 2006). Water temperatures during the winter/spring spawning
period have averaged about 10.5 [deg]C (Heidt and Gilbert 1979), which
is consistent with DeVries' (2006) observation that spawning runs
appeared to commence when water temperatures reach 10.2 [deg]C. When
water temperatures exceed 27 [deg]C, shortnose sturgeon typically move
above the salt-fresh water interface and aggregate in deeper areas of
the river (DeVries 2006); however, shortnose sturgeon have also been
observed to use lower portions of the river throughout the summer, even
when water temperatures averaged 34 [deg]C (Heidt and Gilbert 1979;
DeVries 2006).
Overall, the variation in habitat characteristics across the range
of shortnose sturgeon indicates that there is no single type or typical
river system. Despite a suite of existing threats, shortnose sturgeon
continue to occupy many river systems across their historical range.
The fact that the Saint John River lies at one end of the species'
range, and among other attributes, experiences different temperature
and flow regimes, does not mean that this particular river is unusual
or unique given the variability in habitat conditions observed across
the range. Therefore, we conclude that the Saint John River is not an
unusual or unique ecological setting when viewed against the range of
the taxon as a whole. Furthermore, though not relied up on for our
finding, we note that COSEWIC (In Press) recently concluded that
shortnose sturgeon from other river systems would probably be able to
survive in Canada.
Significant Gap in the Range of the Taxon
The second consideration under the DPS Policy in determining
whether a population may be ``significant'' to its taxon is whether the
``loss of the discrete population segment would result in a significant
gap in the range of a taxon'' (61 FR 4722, February 7, 1996). Shortnose
sturgeon are distributed along the Atlantic coast of North America from
the Minas Basin, Nova Scotia to the St. Johns River, Florida,
representing a coastal range of roughly 3,700 km. The Saint John River,
located at the northern end of the range, represents a small portion of
the species' currently occupied geographic range. In addition, although
the Saint John River is presumed to contain a relatively large
population of shortnose sturgeon, that populaiton is not considered the
largest, and it represents one of at least 10 spawning populations
(SSRT 2010). Furthermore, relatively recent field data indicate
shortnose sturgeon make coastal migrations to a greater extent than
previously thought (e.g., Dionne et al. 2013) and are capable of making
marine migrations of over 300 km (e.g., between Penobscot and Merrimack
rivers; M. Kieffer, USGS, pers. comm. 2010). Such data suggest the
potential for recolonization of the Saint John River by shortnose
sturgeon migrating from populations to the south. Further indirect
evidence in support of this possibility comes from the existing genetic
data, which indicate some level of gene flow among rivers in the
Northeast, including the Saint John River (Wirgin et al. 2005; Wirgin
et al. 2009; King et al. 2014). Thus, in light of the potential for
recolonization and the fact that the Saint John River population of
shortnose sturgeon does not constitute a substantial proportion of the
species' range, we conclude that the loss of the Saint John River would
not constitute a significant gap in the range of the species.
Only Natural Occurrence of the Taxon
Under the DPS Policy, a discrete population segment that represents
the ``only surviving natural occurrence of a taxon that may be more
abundant elsewhere as an introduced population outside its historical
range'' may be significant to the taxon as whole (61 FR 4722, February
7, 1996). This consideration is not relevant in this particular case,
because shortnose sturgeon are present in many river systems throughout
their historical range (SSRT 2010).
Genetic Characteristics
As stated in the DPS Policy, in assessing the ``significance'' of a
``discrete'' population, we consider whether the discrete population
segment differs markedly from other populations of the species in its
genetic characteristics (61 FR 4722, February 7, 1996). Therefore, we
examined the available data to determine whether the Saint John River
shortnose sturgeon differ markedly in their genetic characteristics
when compared to other populations. In conducting this evaluation under
the second criterion of the DPS policy, we looked beyond whether the
genetic data allow for discrimination of the Saint John population
segment from other populations (a topic of evaluation in connection
with the first criterion of ``discreteness''), and instead focused on
whether the data indicate marked genetic differences that appear to be
significant to the taxon as a whole. In this sense, we give independent
meaning to the ``genetic discontinuity'' of the discreteness criterion
of the DPS Policy and the ``markedly differing genetic
characteristics'' of the significance criterion.
Genetic analyses indicate fairly moderate to high levels of genetic
diversity of shortnose sturgeon in most river systems across the
geographic range (Grunwald et al. 2002, Quattro et al. 2002; Wirgin et
al. 2009). Based on the 11 nDNA loci examined in samples from 17
locations, King et al. (2014) reported that the number of observed
alleles (i.e., versions of a gene at a particular locus; here with
overall frequencies >1%) ranged from a low of 55 in the Cape Fear River
(n= 3 fish) to a high of 152 in the Hudson River (n= 45 fish); 118
alleles were observed in the Saint John River sample (n=25 fish).
Estimated heterozygosity was not reported by river sample, but King et
al. (2014) noted that it was lowest for the southern rivers relative to
the mid-Atlantic and northern river samples. Wirgin et al. (2009)
reported that haplotypic diversity ranged from 0.500 (Santee River,
n=4) to 0.862 (Altamaha River, n= 69) across 15 sample populations,
with the Saint John River population having a haplotype diversity index
of 0.696 (n=42). The number of individual haplotypes observed in any
[[Page 65193]]
one river sample ranged from two (Santee River, n=4) to 13 (Winyah Bay,
n=46), with eight haplotypes observed in the Saint John River sample
(n=42, Wirgin et al. 2009). The level of genetic diversity based on the
mtDNA was not correlated with population size, and there was also no
evidence of population bottlenecks, which may be due to historical
recency of most population declines (over past ~100 years, Grunwald et
al. 2002; Wirgin et al. 2009). Overall, the level of genetic diversity
observed for the Saint John River population segment is not unusual
relative to that observed in the taxon as a whole. However, Grunwald et
al. (2002) noted that the lack of reduced haplotypic diversity within
the northern sample rivers contrasts with findings for other anadromous
fishes from previously glaciated rivers. Grunwald et al. (2002)
hypothesized the high degree of haplotypic diversity and large number
of unique haplotypes in the previously glaciated northern region (i.e.,
Hudson River and northward) may be the result of a northern population
having survived in one or more northern refugia.
As discussed previously, at a regional scale, most of the mtDNA
haplotypes observed are shared across multiple, adjacent rivers
sampled; however, very little sharing of haplotypes has been documented
between the northern and southern portions of the range (Quattro et al.
2002; Grunwald et al. 2002; Wirgin et al. 2009). In the analysis
conducted by Wirgin et al. (2009), the Saint John River sample had one
private haplotype (in 1 of 42 fish) and shared the remaining 7
haplotypes with multiple rivers. Of the seven shared haplotypes, two
were each shared with two other river systems, including the Hudson and
Connecticut rivers, and the remaining five haplotypes were shared
across three to six other rivers within the northeast and mid-Atlantic
portions of the range (Wirgin et al. 2009). In an earlier study by
Quattro et al. (2002) in which control region mtDNA was sequenced for
211 shortnose sturgeon collected from five southeastern U.S. rivers and
the Saint John River, one haplotype was observed in all river samples.
This shared haplotype occurred in 1 of 13 fish (7.7%) sampled from the
Saint John River and 1 of 5 fish (20%) sampled from Winyah Bay; the
remaining river samples contained this haplotype at higher frequencies
(36%-79%, Quattro et al. 2002).
While the shortnose sturgeon from the Saint John River have a
fairly high degree of genetic diversity and shared haplotypes with
other rivers, they can be statistically differentiated from other river
samples based on haplotype frequencies and nDNA distance metrics
(Wirgin et al. 2009; King et al. 2014). However, the same is also true
for the majority of rivers across the range of the species. For
example, using genetic distances (Phi PT), King et al.
(2014) detected significant differences in all pairwise comparisons
except for three rivers in the northeast (Penobscot, Androscoggin, and
Kennebec rivers) and three rivers in the southeast (Edisto, Savannah,
and Ogeechee rivers). Similarly, significant differences in haplotype
frequencies have been reported for most river populations sampled. In
Chi-squared analyses, Grunwald et al. (2002) reported significant
differences for all but 4 of 82 pairwise comparisons of mtDNA
nucleotide substitution haplotype frequencies across 10 sample sets
(two of which were from different sections of the Connecticut River),
and Wirgin et al. (2009) reported significant differences for all but 9
of 91 pairwise comparisons of mtDNA haplotype frequencies across 13
river populations.
The magnitude of these genetic differences between individual river
systems varies across the range of the species and indicates a
hierarchical pattern of differentiation. For example, the mtDNA data
reveal a deep divergence between rivers in the northern portion of the
range from rivers in the southern portion of the range. Of the 29
haplotypes observed by Grunwald et al. (2002), 11 (37.9%) were
restricted to northern systems, 13 (44.8%) were restricted to the more
southern systems, and only 5 (17.2%) slightly overlapped the two
regions. In the later and larger study by Wirgin et al. (2009), the
observed haplotypes again clustered into regional groupings: 10 of 38
observed haplotypes (26.3%) only occurred in systems north of the
Hudson River, 16 of 38 (42.1%) only occurred in systems south of the
Chesapeake Bay, and just 5 of 38 (13.2%) haplotypes overlapped in the
mid-Atlantic region. The limited sharing of haplotypes between the
north and south regions is consistent with strong female homing
fidelity and limited gene flow between these regions. The break in
shared haplotypes corresponds with the historical division of the
species due to Pleistocene glaciation, which Grunwald et al. (2002)
stated was probably the most significant event affecting population
structure and patterns of mtDNA diversity in shortnose sturgeon.
The recent nDNA analyses of King et al. (2014) also indicate an
unambiguous differentiation of sample populations into one of three
major geographic groupings--Northeast, Mid-Atlantic, or Southeast. When
all 17 sample populations were pooled by these three geographic
regions, correct assignment to each region was 99.1% for the Northeast
and 100% (i.e., zero mi-assigned fish) for the remaining two regions
(King et al. 2014). Of the 133 fish included for the Northeast group,
one was mis-assigned to the Mid-Atlantic. The estimates of effective
migrants per generation (based on Phi PT) are consistent
with the regional zones of genetic discontinuity among Northeast, Mid-
Atlantic, and Southeast river systems. The average migrants per
generation between regions ranged from less than one migrant (i.e.,
0.89) between Northeast and Southeast to nearly two migrants (i.e.,
1.89) between Northeast and Mid-Atlantic. In contrast, the range of
estimated migrants per generation within regions was 2.25-83.08 for the
Northeast, 1.87-13.64 for the Mid-Atlantic, and 2.38-49.75 for the
Southeast (King et al. 2014). The estimated migrants per generation
between the Saint John River in particular and all other rivers within
the Northeast ranged from 2.25-3.43 (King et al. 2014). Taken together,
these data indicate that the degree of genetic differentiation between
the Saint John River and the rivers within the Gulf of Maine is
relatively small or ``shallow'', especially relative to the deeper
divergence observed among the regional groupings of river populations.
A possible explanation for the relatively low level of differentiation
within the Northeast is that the those populations are relatively young
in a geologic sense due to recent glaciations compared to populations
in the more southern part of the range (SSRT 2010).
In conclusion, given the patterns of genetic diversity, shared
haplotypes, and relative magnitudes of genetic divergence at the river
drainage versus regional scale, we find there is insufficient evidence
that the Saint John River population of shortnose sturgeon differs
markedly in its genetic characteristics relative to the taxon as a
whole so as to meet the test for ``significance'' on this basis. While
the Saint John River population segment can be genetically
distinguished from other river populations, available genetic evidence
places it into a larger evolutionarily meaningful unit, along with
several other river populations sampled. The degree of differentiation
among the three larger regional groups is more marked than the
differences observed among populations from the Saint John and other
nearest rivers, suggesting that the Saint John River
[[Page 65194]]
population's differentiation is not ``significant'' in the context of
the whole species. Gene flow estimates are also consistent with the
observed deeper zones of divergence detected at the regional scale.
Thus, we conclude that these data do not support delineation of the
Saint John River population segment as ``significant.'' In so
interpreting the available genetic data, we are mindful of the
Congressional guidance to use the DPS designation sparingly.
DPS Conclusion and Petition Finding
We conclude that the Saint John River population of shortnose
sturgeon is ``discrete'' based on evidence that it is a relatively
closed and somewhat geographically isolated population segment. It thus
satisfies the first prong of the DPS policy. However, we also find that
the Saint John River population segment is not ``significant'' to the
taxon as a whole. It thus fails to satisfy the second prong of the DPS
Policy. As such, based on the best available data, we conclude that the
Saint John River population of shortnose sturgeon does not constitute a
DPS and, thus, does not qualify as a ``species'' under the ESA.
Therefore, we deny the petition to consider this DPS for delisting. Our
denial of the petition on this ground does not imply any finding as to
how we should proceed if the situation were otherwise, i.e., where a
population is found instead to meet the criteria to be a DPS. Even if
the population had met both criteria of the DPS Policy, and even if the
population were also found to have a status that differed from the
listed entity, it would not necessarily be appropriate to propose
modifications to the current listing, in light of the unsettled legal
issues surrounding such revisions. Nor do we resolve here what steps
would need to be followed to propose revisions to the species' listing
if the facts had been otherwise; such an inquiry would be hypothetical
in this case. It is clear that because the petition at issue here
sought identification of a DPS, and because the population at issue is
not a DPS, this particular petition must be denied. As this is a final
action, we do not solicit comments on it.
References Cited
A complete list of references is available upon request to the
Office of Protected Resources (see ADDRESSES).
Authority
The authority for this action is the Endangered Species Act of
1973, as amended (16 U.S.C. 1531 et seq.).
Dated: October 20, 2015.
Samuel D. Rauch III,
Deputy Assistant Administrator for Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2015-27148 Filed 10-23-15; 8:45 am]
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