Endangered and Threatened Wildlife and Plants; Threatened Species Status for the Headwater Chub and Roundtail Chub Distinct Population Segment, 16981-16988 [2017-06995]
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Federal Register / Vol. 82, No. 66 / Friday, April 7, 2017 / Proposed Rules
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FOR FURTHER INFORMATION CONTACT: Matt
Rau, Environmental Engineer, Control
Strategies Section, Air Programs Branch
(AR–18J), Environmental Protection
Agency, Region 5, 77 West Jackson
Boulevard, Chicago, Illinois 60604,
(312) 886–6524, rau.matthew@epa.gov.
SUPPLEMENTARY INFORMATION: In the
Final Rules section of this Federal
Register, EPA is approving the State’s
SIP submittal as a direct final rule
without prior proposal because the
Agency views this as a noncontroversial
submittal and anticipates no adverse
comments. A detailed rationale for the
approval is set forth in the direct final
rule. If no adverse comments are
received in response to this rule, no
further activity is contemplated. If EPA
receives adverse comments, the direct
final rule will be withdrawn and all
public comments received will be
addressed in a subsequent final rule
based on this proposed rule. EPA will
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Please note that if EPA receives adverse
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comment. For additional information,
see the direct final rule which is located
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Register.
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Dated: March 22, 2017.
Robert A. Kaplan,
Acting Regional Administrator, Region 5.
[FR Doc. 2017–06883 Filed 4–6–17; 8:45 am]
BILLING CODE 6560–50–P
ENVIRONMENTAL PROTECTION
AGENCY
40 CFR Part 52
[EPA–R04–OAR–2016–0575; FRL–9960–56–
Region 4]
Air Plan Approval; Tennessee:
Reasonable Measures Required
Environmental Protection
Agency.
ACTION: Proposed rule.
AGENCY:
The Environmental Protection
Agency (EPA) is proposing to approve a
State Implementation Plan (SIP)
revision submitted by the State of
Tennessee, through the Tennessee
Department of Environment and
Conservation (TDEC), on March 25,
1999. The SIP submittal includes a
change to the TDEC regulation
‘‘Reasonable Measures Required.’’ EPA
is proposing to approve this SIP revision
because it is consistent with the Clean
Air Act and federal regulations
governing SIPs.
DATES: Written comments must be
received on or before May 8, 2017.
ADDRESSES: Submit your comments,
identified by Docket ID No. EPA–R04–
OAR–2016–0575 at https://
www.regulations.gov. Follow the online
instructions for submitting comments.
Once submitted, comments cannot be
edited or removed from Regulations.gov.
EPA may publish any comment received
to its public docket. Do not submit
electronically any information you
consider to be Confidential Business
Information (CBI) or other information
whose disclosure is restricted by statute.
Multimedia submissions (audio, video,
etc.) must be accompanied by a written
comment. The written comment is
considered the official comment and
should include discussion of all points
you wish to make. EPA will generally
not consider comments or comment
contents located outside of the primary
submission (i.e., on the web, cloud, or
other file sharing system). For
additional submission methods, the full
EPA public comment policy,
information about CBI or multimedia
submissions, and general guidance on
making effective comments, please visit
https://www2.epa.gov/dockets/
commenting-epa-dockets.
FOR FURTHER INFORMATION CONTACT: D.
Brad Akers, Air Regulatory Management
SUMMARY:
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Section, Air Planning and
Implementation Branch, Air, Pesticides
and Toxics Management Division, U.S.
Environmental Protection Agency,
Region 4, 61 Forsyth Street SW.,
Atlanta, Georgia 30303–8960. Mr. Akers
can be reached via telephone at (404)
562–9089 or via electronic mail at
akers.brad@epa.gov.
SUPPLEMENTARY INFORMATION: In the
Final Rules section of this Federal
Register, EPA is approving the State’s
SIP revision as a direct final rule
without prior proposal because the
Agency views this as a noncontroversial
submittal and anticipates no adverse
comments. A detailed rationale for the
approval is set forth in the direct final
rule. If no adverse comments are
received in response to this rule, no
further activity is contemplated. If EPA
receives adverse comments, the direct
final rule will be withdrawn and all
public comments received will be
addressed in a subsequent final rule
based on this proposed rule. EPA will
not institute a second comment period
on this document. Any parties
interested in commenting on this
document should do so at this time.
Dated: March 15, 2017.
V. Anne Heard,
Acting Regional Administrator, Region 4.
[FR Doc. 2017–06878 Filed 4–6–17; 8:45 am]
BILLING CODE 6560–50–P
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R2–ES–2015–0148;
4500030113]
RIN 1018–BA86
Endangered and Threatened Wildlife
and Plants; Threatened Species Status
for the Headwater Chub and Roundtail
Chub Distinct Population Segment
Fish and Wildlife Service,
Interior.
ACTION: Proposed rule; withdrawal.
AGENCY:
We, the U.S. Fish and
Wildlife Service (Service), withdraw the
proposed rule to list the headwater chub
(Gila nigra) and a distinct population
segment (DPS) of the roundtail chub
(Gila robusta) from the lower Colorado
River basin as threatened species under
the Endangered Species Act (Act). This
withdrawal is based on a thorough
review of the best scientific and
commercial data available, which
indicate that the headwater chub and
SUMMARY:
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the roundtail chub DPS are not discrete
taxonomic entities and do not meet the
definition of a species under the Act.
These fish are now recognized as a part
of a single taxonomic species—the
roundtail chub (Gila robusta). Because
the entities previously proposed for
listing are no longer recognized as
species, as defined by the Act, we have
determined that they are not listable
entities and we are withdrawing our
proposed rule to add them to the List of
Endangered and Threatened Wildlife.
Section 4(b)(6) of the Act and
implementing regulations at 50 CFR
424.17 provide that the Service must,
within 1 year of a proposed rule to list,
delist, or reclassify species, or to
designate or revise critical habitat,
withdraw the proposal if the available
evidence does not justify the proposed
action. The document withdrawing the
rule must set forth the basis upon which
the proposed rule has been found not to
be supported by available evidence.
Once withdrawn, the action may not be
re-proposed unless sufficient new
information is available.
FOR FURTHER INFORMATION CONTACT:
Steve Spangle, Field Supervisor, U.S.
Fish and Wildlife Service, Arizona
Ecological Services Office, 9828 North
31st Ave., #C3, Phoenix, AZ 85051–
2517; telephone 602–242–0210. Persons
who use a telecommunications device
for the deaf (TDD) may call the Federal
Relay Services at 800–877–8339.
SUPPLEMENTARY INFORMATION:
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Previous Federal Action
On October 7, 2015 (80 FR 60754), we
published a proposed rule to list the
headwater chub and the lower Colorado
River basin roundtail chub DPS
(roundtail chub DPS) as threatened
species under the Act (16 U.S.C. 1531 et
seq.). On August 15, 2016 (81 FR
54018), we announced a 6-month
extension on the final listing
determination that the Act allows when
there is substantial disagreement
regarding the sufficiency or accuracy of
the available data, and reopened the
comment period on the proposed
listings for 30 days. During this
comment period we received new
information. On November 1, 2016 (81
FR 75801), we reopened the comment
period on the proposed listings for an
additional 45 days to provide the public
additional time to review and consider
the proposed rulemakings in light of
this new information. As a result of the
6-month extension, the deadline to
finalize, modify, or withdraw the
proposed rule is April 7, 2017.
For a description of additional
previous Federal actions concerning
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these species, please refer to the October
7, 2015, proposed listing rule (80 FR
60754).
Background
At the time we published our
proposed rule (October 7, 2015; 80 FR
60754), the Committee on Names of
Fishes, a joint committee of the
American Fisheries Society and
American Society of Ichthyologists and
Herpetologists (the Societies) (Page et al.
2013, p. 71), considered headwater chub
and roundtail chub to be separate
species. As a consortium of fisheries
scientists, the American Fisheries
Society is the recognized and accepted
scientific authority on fish taxonomy.
Accordingly, our proposed rule assessed
the headwater chub and roundtail chub
as separate species. However,
commenters on our proposed rule raised
questions during the public comment
period regarding the taxonomic
distinctness of the headwater and
roundtail chubs, as related to the Gila
chub (Gila intermedia). At that time,
some scientists knowledgeable about the
fish contended that the three entities
were not separate species (Carter et al.
2016 in press; Copus et al. 2016). For
this reason, the Arizona Game and Fish
Department requested that the Societies
evaluate the most recent literature
associated with roundtail chub,
headwater chub, and Gila chub
taxonomy. In their final report to the
Arizona Game and Fish Department, the
Societies panel concluded that ‘‘no
morphological or genetic data define
populations of Gila in the lower
Colorado River basin (which, as defined
by the Service, includes the Little
Colorado River, Bill Williams River,
Gila River, Verde River, and Salt River
drainages) as members of more than one
species’’ and ‘‘that the data available
support recognition of only one species
of Gila, the roundtail chub, Gila
robusta’’ (Page et al. 2016, p. 1). These
three fish are now considered by the
Societies to be a single species,
roundtail chub (Gila robusta) because
data do not support recognition of three
species.
Taxonomy
Introduction
The taxonomic history of the genus
Gila in the Colorado River basin has
changed over time, especially for the
three forms (roundtail, headwater, and
Gila chub) found in the Gila River basin.
These forms have been variously
classified as full species, assigned as
different species, subspecies of Gila
robusta, or as part of a ‘‘Gila robusta
complex’’ (Miller 1945; Holden 1968;
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Rinne 1969; Holden and Stalnaker 1970;
Rinne 1976; Smith et al. 1977; DeMarais
1986; Rosenfeld and Wilkinson 1989;
Dowling and DeMarais 1993; Douglas et
al. 1998; Minckley and DeMarais 2000;
Gerber et al. 2001). As noted by nearly
all researchers investigating the
systematics of Gila spp., the taxonomic
situation is complicated and
problematic (Holden and Stalnaker
1970; Minckley 1973; Minckley and
DeMarais 2000; Gerber et al. 2001;
¨
Schonhuth et al. 2014) due to various
factors including multiple independent
hybridization events over time (Rinne
1976; DeMarais 1986; Rosenfeld and
Wilkinson 1989; DeMarais et al. 1992;
Dowling and DeMarais 1993; Minckley
and DeMarais 2000; Gerber et al. 2001;
¨
Schwemm 2006; Schonhuth et al. 2014;
Brandenburg et al. 2015,) potential past
introgression (the transfer of genetic
information from one species to another
as a result of hybridization between
them and repeated backcrossing)
(DeMarais et al. 1992; Minckley and
DeMarais 2000), recent divergence
within the three fish (Schwemm 2006).
Further, the original assignment to
species was based on the assumption
that the three fish do not overlap
geographically (parapatry), which we
recognize now is not an accurate
assumption. Additionally, in some
instances when the same fish was
identified based on morphology
(physical characteristics) it was
identified as one species and when
identified based on genetic analysis it
was identified as a different species
(Dowling et al. 2015, pp. 14–15). Recent
and ongoing genetic and morphologic
analyses of chubs in the Gila River basin
continue to yield conflicting results
(DeMarais et al. 1992; Schwemm 2006;
Dowling et al. 2008 and 2015;
¨
Schonhuth et al. 2014; Marsh et al.
2016, all entire).
History
Gila robusta (roundtail chub) was first
described by Baird and Girard (1853, p.
365–369) from specimens collected in
1851 from the Zuni River (tributary to
Little Colorado River). Gila nigra
(headwater chub; formerly known as G.
robusta graham or G. grahami) was first
described as a subspecies (G. robusta
graham) from Ash Creek in the San
Carlos River in east-central Arizona in
1874 (Cope and Yarrow 1875, p. 663),
but not returned to full species status
(G. robusta) until proposed so by
Minckley and DeMarais (2000, p.
entire). The Societies accepted Gila
nigra as a full species (Nelson et al.
2004, p. 71), as did the New Mexico
Department of Game, Fish (Carman
2006, p. 3), Arizona Game, and Fish
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Department (AGFD 2006, p. 3) and
continued to recognize G. robusta as a
distinct species. Therefore, based on the
best available commercial and scientific
data the Service accepted both Gila
robusta and Gila nigra as full species as
documented in our 12-month findings
(May 3, 2006; 71 FR 26007 and July 7,
2009; 74 FR 32352). In their 2013
publication of Common and Scientific
Names of Fishes from the United States,
Canada, and Mexico, the Societies
continued to list both Gila robusta and
Gila nigra as distinct species (Page et al.
2013, p. 71). A summary of the historic
and current nomenclature from Rinne
(1976, entire), Sublette et al. (1990,
entire), and Minckley and DeMarais
(2000, entire) is summarized in Voeltz
(2002, pp. 8) and Copus et al. (2016, pp.
1&6). The Gila chub (Gila intermedia) is
currently listed as an endangered
species (November 2, 2005; 70 FR
66664).
These entities were originally
classified based on the streams in which
they were found (Minckley and
DeMarais 2000, p. 252), under the
assumption that G. robusta and G. nigra
either did not overlap (allopatric, no
gene flow) or there was only a narrow
overlap (parapatric; limited interaction
and opportunity for gene flow)
(Minckley and DeMarais 2000 pp. 252–
254). Because hybridization between G.
robusta and G. intermedia indicates that
these fish must co-occur in some
streams (Minckley and DeMarais 2000,
entire), we conclude that Minckley and
DeMarais’s (2000) assumption they did
not overlap was unfounded. Further,
other studies have found that fish
designated as G. robusta, G. nigra, and
G. intermedia overlap geographically or
occur adjacent to one another (Dowling
and Marsh 2009, p. 1; Marsh et al. 2016,
p. 57; Brandenburg et al. 2015, p. 18).
Morphology
The approach for classifying G.
robusta, G. nigra, and G. intermedia
developed by Minckley and DeMarais
(2000, pp. 254–255) presumes there is
little intraspecific variation (differences
within a species) in the morphologic
and meristic (counting quantitative
characteristics such as fins)
characteristics used to distinguish these
three taxa. However, the three purported
species overlap in physical
characteristics, and many fish have
intermediate physical characteristics.
Those characteristics that do not overlap
are separated by very small margins,
making species-level identification of
individual fish problematic, even when
the geographic origin of the species is
known (Brandenburg 2015, entire).
Minckley and DeMarais (2000, pp. 253–
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254) indicate that G. nigra is physically
different from G. intermedia even
though they appear physically more
similar to one another than either is to
G. robusta. In addition, Copus et al.
(2016, p. 13) did not find physical
characteristics in the Minckley and
DeMarais (2000, pp. 254–255)
classification key to reliably
differentiate G. robusta, G. nigra, and G.
intermedia from one another. Copus et
al. (2016 p. 16) concluded that there was
no morphological basis for taxonomic
distinctions within the Gila spp.
complex.
Genetics
Multiple genetic analysis studies have
been conducted that reveal differences
between different chub populations, but
have been unable to identify differences
between G. robusta, G. nigra, and G.
intermedia (DeMarais et al. 1992, pp.
2748–2749; Schwemm 2006, p. 29;
Dowling et al. 2008, p. 2, and 2015, p.
13; Copus et al. 2016, pp. 14–15; Marsh
et al, 2016, p.58). Mitochondrial DNA
¨
analysis (Schonhuth et al. 2014, p. 223)
indicates that G. robusta, G. nigra, and
G. intermedia belong to one clade (a
grouping that includes a common
ancestor and all its descendants, living
and extinct, of that ancestor).
¨
Schonhuth et al. (2014, p. 223)
hypothesized that this could reflect
hybridization or incomplete lineage
sorting (when the lineage of a specific
gene is not the same as the lineage of the
species, obscuring the true species
relationship).
However, when nuclear DNA (rather
than mitochondrial DNA) was analyzed,
a broader grouping was identified that
included G. seminude and G. elegans,
but when mitochondrial and nuclear
DNA results are combined G. robusta, G.
nigra, and G. intermedia were in one
¨
grouping (Schonhuth et al. 2014, p.
223). Preliminary studies by Chafin et
al. (2016) indicate evolutionary
independent lineages for G. robusta, G.
nigra, and G. intermedia, and that the
hybrid origin of G. nigra is not
supported. Studies by Marsh et al.
(2016, entire) point to genetic variation
between populations of G. robusta and
G. nigra, and demonstrate evidence that
distinct ecological differences between
some populations are now thought to
exist. Minckley and DeMarais (2000,
entire) supported recognition of three
species, but acknowledged that most
genetic variation was within
populations for G. robusta, and was
among populations for G. intermedia
and G. nigra. Minckley and DeMarais
(2000, p. 253) also indicated that these
three fishes share genetic features (that
had been studied so far) while behaving
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as separate non-overlapping (allopatric)
morphological species. In addition,
some populations assigned to species
based on genetics appeared to conflict
with the species level-assignment based
on morphology (Dowling et al. 2008, p.
27).
Speciation
Minckley and DeMarais (2000, p. 253)
describe three different taxonomic
options for chubs in the Gila River
basin: a single species with many
different forms or stages (polymorphic
species), a species containing multiple
subspecies, or three full species. They
acknowledge that none of these
taxonomic options is biologically
justified without knowing if these fish
naturally occur in the same geographic
area (sympatry, indicating an initial
interbreeding population that split), or
occur immediately adjacent to each
other but not significantly overlapping
(parapatry, indicating there is no barrier
to gene flow). They further acknowledge
that a persistent narrow interaction zone
(parapatry, indicating there is no barrier
to gene flow) of morphologically
distinguishable G. robusta, G.
intermedia, and G. nigra has been
confirmed, but note that in no instance
was any two of the three caught at the
same locality (allopatric, no gene flow;
p. 251). However, they also
acknowledge that hybridization
(between G. robusta and G. intermedia,
resulting in G. nigra) in the past must
have occurred in some places and not
others, thereby demonstrating
occurrence in the same geographic area
(sympatry) (p. 253). They conversely
hypothesized that the current minimal
overlap in an area where species are
adjacent (parapatry, indicating there is
no barrier to gene flow) may thus reflect
an ancestral ecological segregation area
(sympatry, indicating an initial
interbreeding population that split due
to the use of different habitats and
resources) that promoted persistence in
the ever-increasing aridity of the
Southwest (p. 253).
In Fossil Creek, G. nigra and G.
robusta appear to be sympatric,
including hybrids between G. robusta
and G. nigra (Marsh et al. 2016, p. 57).
Brandenburg et al. (2015, p. 18)
concluded that the morphological
assessment of Gila spp. in New Mexico
confirmed that the three fish were found
in the same geographic area (sympatric)
in almost all cases, contradicting
Minckley and DeMarais’ results (2000,
p. 251) as well as other previous
literature suggesting that these Gila spp.
are occurring in separate nonoverlapping geographical areas
(allopatric) through their ranges (Rinne
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1969, p. entire; DeMarais 1986, p. entire;
Minckley and DeMarais 2000, p. 253). In
Fossil Creek, they found that G. nigra
and G. robusta are locally in the same
geographic area (sympatric) and have
hybridized (Marsh et al. 2016, p. 57).
Marsh et al. (2016, p. 58) concluded
there are two morphologically similar,
but genetically distinguishable, chub in
Fossil Creek, G. robusta and G. nigra.
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Conservation Implications
Dowling et al. (2015, pp. 14–15)
reasoned that the lack of diagnostic
molecular characteristics does not
inform the status of these three fish, but
rather highlights the role that local
evolution has played in shaping
patterns of variation in these taxa and
the importance of accounting for this
variation when managing the complex.
Most, if not all, scientists agree that
conservation actions for these chubs
must be directed at the population level
and must include consideration of the
complex as a whole (Dowling et al.
2008, pp. 30–31; Dowling and DeMarais
1993, p. 445; Gerber et al. 2001, p. 2037;
Schwemm et al. 2006, pp. 32–33). The
Arizona Game and Fish Department
recognizes the importance of conserving
the currently recognized roundtail chub
population rangewide (including the
formerly known headwater chub and
Gila chub) and is committed to the
conservation agreements and practices
that have been in place since 2006
(AGFD 2017, entire; AGFD 2006, entire).
Public Comments
In our October 7, 2015 proposed rule
(80 FR 60754), we requested that all
interested parties submit comments or
information concerning the proposed
listings during a 60 day comment
period, ending December 7, 2015. We
particularly sought comments
concerning genetics and taxonomy. In
our August 15, 2016, 6-month extension
document (81 FR 54018), we reopened
the comment period on the proposed
rule for 30 days, ending September 14,
2016, and we again requested comments
and information regarding genetics and
morphology that would aid in resolving
the ongoing taxonomic issues regarding
classification of these fish. On
November 1, 2016 (81 FR 75801, we
announced an additional 45-day
comment period, ending December 16,
2016, on the October 7, 2015 proposed
rule.
We provided notification of these
publications and their comment periods
through email, letters, and news releases
faxed and/or mailed to the appropriate
Federal, State, and local agencies;
county governments; elected officials;
media outlets; local jurisdictions;
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scientific organizations; interested
groups; and other interested parties.
In accordance with our peer review
policy published in the Federal Register
on July 1, 1994 (59 FR 34270), we
solicited independent opinions from at
least three knowledgeable individuals
who have expertise with these fish, who
possess a current knowledge of the
geographic region where the fish occurs,
and/or are familiar with the principles
of conservation biology.
We reviewed all comments received
from peer reviewers and the public for
substantive issues and new information
regarding the proposed listing of G.
nigra and the G. robusta DPS.
Substantive comments pertaining to the
taxonomy of these fish received during
the comment period are addressed
below. We also received several
comments from both the public and
peer reviewers concerning threats to
these fish; however, because our
withdrawal is due to taxonomic revision
such comments are outside the scope of
this withdrawal.
Peer Review Comments
(1) Comment: One peer reviewed
stated that there are no recent (since
2000) publications in the peer-reviewed
literature that provide evidence that
Gila intermedia, G. nigra, and G. robusta
are other than separate and distinct
species. The peer reviewer further stated
that there are articles that study the
genetics or morphology of these fish
without questioning its taxonomy,
¨
specifically Schonhuth et al. 2014,
¨
Schonhuth et al. 2012, and Marsh et al.
in press.
Response: Multiple studies since 2000
provide information on the genetic
analysis for these fish, including
Schwemm 2006, Dowling et al. 2008
and 2015, and Copus et al. 2016. While
these studies may not have questioned
the taxonomic classification, they also
have not been able to identify genetic
markers that have the ability to
distinguish among G. robusta, G. nigra,
¨
and G. intermedia. Schonhuth et al.
(2008, p. 213; 2014, p. 223), using
mitochondrial and nuclear DNA
sequencing, found that G. robusta, G.
nigra, and G. intermedia were well
supported as having a common
ancestor. Using mitochondrial DNA,
¨
Schonhuth et al. (2008, p. 213; 2014, p.
223) found that G. robusta, G. nigra, and
G. intermedia were in one grouping that
included a common ancestor and all the
descendants (living and extinct) of that
ancestor (clade), and this could reflect
incomplete lineage sorting or
hybridization. However, when nuclear
DNA was analyzed, a broader grouping
was identified that included G.
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seminuda and G. elegans, but when
mitochondrial and nuclear DNA results
were combined, G. robusta, G. nigra,
and G. intermedia were alone in one
grouping. While Marsh et al. (2016,
entire) concluded there are two similar
but genetically distinguishable species
in the creek they studied, their findings
differ somewhat from Schwemm (2006)
and Dowling et al. (2008 and 2015,
entire), who were unable to
conclusively identify distinct species
using genetic markers across a much
wider range. Further, the Societies
conducted a review of the literature and
found no evidence to support three
species. The Service has reviewed the
best available scientific and commercial
data and also found a lack of sufficient
evidence to support more than one
species.
(2) Comment: Recognized authorities
on the taxonomy and ecology of these
fish recognized these fish as separate
species based on morphological
diagnostics.
Response: Minckley and DeMarais
(2000), Miller et al. (2005), and
Minckley and Marsh (2009) report
identification of three species using a
diagnostic morphological key. However,
additional reports were unable to
reliably identify these three fish to
species using the same diagnostic key
(Carter et al. 2016, p. 2 and 20, in press;
Brandenburg 2015, entire; Copus et al.
2016, p. 13). Further, Minckley and
DeMarais (2000, pp. 253–254) stated
that G. nigra is morphologically separate
from G. intermedia, but that G. nigra
and G. intermedia appear
morphologically more similar to one
another than either is to G. robusta. In
addition to issues surrounding
morphological identification, multiple
genetic analysis studies have found
population-level differences, but have
been unable to identify genetic markers
that have the ability to distinguish
among G. robusta, G. nigra, and G.
intermedia (DeMarais 1992, pp. 2748–
2749; Schwemm 2006, p. 29; Dowling et
al. 2008, p. 2, and 2015, p. 13; Copus
et al. 2016, pp. 14–15). There are also
¨
the findings of Schonhuth et al. (2014),
¨
Schonhuth et al. (2012) as described in
Response to Comment 1.
(3) Comment: Conclusions are mainly
based on two ‘‘gray literature’’ reports
that have not undergone peer review
(Copus et al. 2016) or were not available
for public consideration (Carter et al.
2016, in press).
Response: Section 4(b)(1)(A) of the
Act requires the Service to make listing
or delisting decisions based on the best
scientific and commercial data
available. Further, our Policy on
Information Standards under the Act
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(July 1, 1994; 59 FR 34271), the
Information Quality Act (section 515 of
the Treasury and General Government
Appropriations Act for Fiscal Year 2001
(Pub. L. 106–554; H.R. 5658)), and our
associated Information Quality
Guidelines (https://www.fws.gov/
informationquality), provide criteria
and, guidance, and establish procedures
to ensure that our decisions are based
on the best scientific data available.
They require us, to the extent consistent
with the Act and with the use of the best
scientific data available, to use primary
and original sources of information as
the basis for our determinations.
Primary or original information sources
are those that are closest to the subject
being studied, as opposed to those that
cite, comment on, or build upon
primary sources. The Act and our
regulations do not require us to use only
peer-reviewed literature, but instead
they require us to use the ‘‘best
scientific and commercial data
available.’’ We use information from
many different sources, including
articles in peer-reviewed journals,
scientific status surveys and studies
completed by qualified individuals,
Master’s thesis research that has been
reviewed but not published in a journal,
other unpublished governmental and
nongovernmental reports, reports
prepared by industry, personal
communication about management or
other relevant topics, conservation plans
developed by States and counties,
biological assessments, other
unpublished materials, experts’
opinions or personal knowledge, and
other sources. For these reasons, we
think it is appropriate to include review
of Copus et al. (2016) and Carter et al.
(2016, in press), as well as other
sources, within our review.
(4) Comment: Several authors
presented data and conclusions that
conflicted with the previously cited
Carter et al. (2016, in press) and Copus
et al. (2016) reports pertaining to
morphological identification, DNA
analysis, and ecological equivalency to
a subset of the Joint Committee
convened in April 2016, to specifically
address the taxonomy of the roundtail
chub complex.
Response: We were present at the
April 2016 Joint Committee webinar,
and experts beyond Carter and Copus,
such as Brandenburg, Schwemm,
Dowling, O’Neill, and Chafin, also
provided information based on research
they either had previously conducted or
are currently conducting on Gila. A
complete list of references cited may be
obtained on the Internet at https://
www.regulations.gov and upon request
from the Arizona Ecological Services
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Office (see FOR FURTHER INFORMATION
The Service has reviewed the
best available scientific and commercial
data and found a lack of sufficient
evidence to support more than one
species.
(5) Comment: This taxonomic dispute
is not simply an academic exercise of
whether to lump or split taxa, because
the decision has enormous implications
for the conservation of imperiled
species. Multiple experts recommended
that the roundtail chub complex,
however it is constituted, be managed as
separate populations or managed as a
complex.
Response: The Service recognizes that
multiple experts agree that conservation
actions must be directed at the
population level and must include
consideration of the complex as a whole
(Dowling et al. 2008, pp. 30–31;
Dowling and DeMarais 1993, p. 445;
Gerber et al. 2001, p. 2037; Schwemm
2006, pp. 32–33). However, the Service
must adhere to the Act and its
implementing regulations, which define
a ‘‘species’’ as any species or subspecies
of fish, wildlife, or plant, and any
distinct population segment of any
vertebrate species which interbreeds
when mature (16 U.S.C. 1532(16) and 50
CFR 424.02). The best available
scientific and commercial data as
discussed above in the Taxonomy
section, support recognition of only one
species, Gila robusta. The Service’s
withdrawal of our proposed rule to list
the headwater and roundtail chub based
on new taxonomic classification does
not diminish the conservation efforts of
our partners to conserve this species
and habitat, nor does our decision affect
the State’s ability to conserve this
species under its own authority. The
Arizona Game and Fish Department
recognizes the importance of conserving
the currently recognized roundtail chub
population rangewide (including the
formerly known headwater chub and
Gila chub) and is committed to the
conservation agreements and practices
that have been in place since 2006
(AGFD 2017, entire; AGFD 2006, entire).
(6) Comment: Multiple commenters
raised concerns with Copus et al. (2016)
methods and conclusions, particularly
small sample size, lack of key analytical
and laboratory steps, the study’s DNA
sequence data filtering and analyses that
failed to follow best practices for
phylogenetic analysis, and specimen
shrinkage associated with duration of
preservation impacting morphological
diagnostics.
Response: The Service did not rely
solely on Copus et al. 2016. We
considered the best available
commercial and scientific data; you may
CONTACT).
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obtain a complete list of references cited
on the Internet at https://
www.regulations.gov and upon request
from the Arizona Ecological Services
Office (see FOR FURTHER INFORMATION
CONTACT). In regards to the
mitochondrial DNA and phylogenetic
analysis, Copus et al.’s findings are
¨
consistent with Schonhuth et al.’s
¨
(2014) and Schonhuth et al.’s (2012)
mitochondrial DNA and phylogenetic
analysis. In addition, multiple genetic
analysis studies have been conducted
that indicate population-level
differences, but do not identify genetic
markers that have the ability to
distinguish among G. robusta, G. nigra,
and G. intermedia (DeMarais 1992, pp.
2748–2749; Schwemm 2006, p. 29;
Dowling et al. 2008, p. 2, and 2015, p.
13).
In regards to morphological diagnostic
errors due to using preserved
specimens, Copus et al. (2016) did use
preserved specimens. However, they
also analyzed fresh material and
concluded that no single diagnostic
character can be used for species
identification, and with considerable
overlap among species in every
morphological character, no suite of
characters can distinguish species
unambiguously (Copus et al. 2016, p.
13). Brandenburg et al. (2015, entire)
also reported overlap in the meristic and
morphometric characteristics, records of
many individual fish with intermediate
physical characteristics, and even those
characters that do not overlap are
separated by very small margins making
species-level identification of individual
fish problematic, even when the
geographic origin of the species is
known.
Public Comments
(7) Comment: Multiple commenters
requested various listing alternatives
under the Act including: List G. robusta
as threatened and encompass all
populations of the chub complex within
the Gila basin requiring a revision of the
recovery plan, list G. robusta and G.
nigra as threatened and retain the
current endangered species status of G.
intermedia, list G. robusta as threatened
and retain the current endangered
species status of G. intermedia, or other
combinations.
Response: The Service must adhere to
the Act and its implementing
regulations, which define a ‘‘species’’ as
any species or subspecies of fish,
wildlife, or plant, and any distinct
population segment of any vertebrate
species which interbreeds when mature
(16 U.S.C. 1532(16) and 50 CFR 424.02),
and based on our review, the best
available scientific and commercial data
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support recognition of only one species,
Gila robusta. As the headwater chub
and roundtail chub DPS no longer meet
the definition of a ‘‘species’’ under the
Act, we must withdraw our proposed
rule to list them as threatened species.
(8) Comment: Multiple commenters
stated that there is a great amount of
morphological overlap among counts
and measures for these chub taxa and
that this has long been recognized. If a
taxonomic key is not 100 percent
correct, that does not necessarily mean
that these are not taxa that are
biologically distinct at the specific level.
A test of the key would require the a
priori identification of each individual
to species. Rather than dismiss the
species’ taxonomic status, biologists
should be working to make a better key
that can be used in the field for the
effective identification and management
of the species.
Response: We recognize that
diagnostic keys do not produce correct
results all the time, whether due to
human error or morphological
similarities among purported species.
However, Copus et al. (2016, p. 13)
concluded that, based on genetic
analysis, no single diagnostic character
can be used for species identification,
and with considerable overlap among
species in every morphological
character, no suite of characters can
distinguish species unambiguously.
Brandenburg et al. (2015, entire) also
reported overlap in the meristic and
morphometric characteristics, and there
are many individual fish whose
morphology resides on an intermediate
spectrum, and even those characters
that do not overlap are separated by
very small margins, making specieslevel identification of individual fish
problematic, even if the geographic
origin of the species is known. In
regards to a priori identification of fish,
assignment to species has been based on
the stream in which the fish occurs
(Minckley and DeMarais 2000, p. 252),
so the identification of the fish that
occurs in each stream is assumed to be
known. Consequently, there exists the
ability to compare findings from the
diagnostic key to the fish within a
particular stream. An updated key may
be prudent; however, the Service must
use the best available scientific and
commercial data available, and we have
concluded from our review that the data
currently support only one species, Gila
robusta. Further, given the overlap in
diagnostic characteristics, the
development of a valid key seems
unlikely.
(9) Comment: Multiple commenters
stated that it has long been
hypothesized that G. nigra formed as the
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result of hybridization between the
other two taxa, so we would expect the
greatest morphological overlap from that
species with the other two taxa. The
question then becomes, is G. nigra
continuing to differentiate from
ancestral G. robusta? When in sympatry,
G. nigra and G. robusta are becoming
increasingly reproductively isolated
from one another (Desert Fishes Council
meeting, Dowling et al. 2016).
Response: We recognize that multiple
studies have indicated that
hybridization has occurred among G.
intermedia and G. robusta resulting in
G. nigra and that continuing evolution
may occur (Schwemm 2006; Dowling et
al. 2008, entire). However, there has
also been information presented
showing no evidence of the hybrid
origin of G. nigra, and that G.
intermedia and G. nigra evolved
separately in non-overlapping areas
(parapatry) (Chafin 2016, entire). In
addition, past research (Dowling et al.
2008, 2015; Schwemm 2006) indicate
that there is more variation among
populations and unique genetics within
specific populations (streams).
(10) Comment: If only G. robusta and
G. intermedia are evaluated, there is no
question that they would be considered
distinct morphological species.
Response: Carter et al. (2016, in press)
found that the physical characteristics
did not reliably differentiate among G.
robusta, G. intermedia, and G. nigra. In
addition, Brandenburg et al. (2015, pp.
8–9) found physically similarity of the
three species, as numerous individuals
exhibited intermediate characters along
the species gradient. The discriminant
function analysis (a statistical analysis
tool to determine which variables
discriminate between two or more
naturally occurring groups) classified
only 16 percent (n = 42) of G.
intermedia (the fewest) while the
majority of the samples were classified
as G. robusta (53.2 percent, n = 140),
which indicates that the ability to
classify these fish correctly to G.
intermedia or G. robusta based on
physical characteristics was low. Due to
the complex genetic makeup and
observable characteristics or traits (i.e.,
physical appearance, behavior, or
physiology) of these species, there are
some stream locations where we do not
know where the geographic overlap of
headwater, roundtail, and, in some
cases Gila chub, begins and ends,
because of the plasticity of observable
characteristics or traits of these fish
within individual streams. Our review
of the data does indicate that there are
differences in observable characteristics
or traits between the fish in different
streams, but the Societies’ review, as
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well as the Service review, of the best
available scientific and commercial data
did not result in a species-level
differentiation between G. robusta and
G. intermedia, or among G. robusta, G.
intermedia, and G. nigra.
(11) Comment: One commenter
recommend that we proceed with an
amended recovery plan to list the status
of this species as threatened under the
Act. The listing of this species is
necessary even if all populations of G.
intermedia and G. nigra are subsumed
into G. robusta.
Response: An assessment of the entire
range of the new taxonomic group of
roundtail chub is planned. We are
initiating a status review of the new
taxonomic entity in 2 to 4 years.
Following that review, we will take
action as appropriate.
Determinations
An entity may only be listed under
the Act if that entity meets the Act’s
definition of a species. The recent report
by the Societies indicates that neither
the headwater chub nor the roundtail
chub can be considered species, as
defined by the Act. Under section 3 of
the Act (16 U.S.C. 1532(16)) and
associated implementing regulations at
50 CFR 424.02, a ‘‘species’’ is defined to
include any species or subspecies of
fish, wildlife, or plant, and any distinct
population segment of any vertebrate
species which interbreeds when mature.
The Act’s implementing regulations at
50 CFR 424.11(a) and the Service
Director’s November 25, 1992,
‘‘Taxonomy and the Endangered Species
Act’’ Memorandum (Memo) provide
additional guidance on how to consider
taxonomic information when assessing a
species for listing under the Act. The
regulations at 50 CFR 424.11(a) state,
‘‘In determining whether a particular
taxon or population is a species for the
purposes of the Act, the Secretary [of
the Interior] shall rely on standard
taxonomic distinction and the biological
expertise of the Department [of the
Interior] and the scientific community
concerning the relevant taxonomic
group.’’ The Director’s Memo specifies
that the Service is ‘‘required to exercise
a degree of scientific judgment regarding
the acceptance of taxonomic
interpretations, particularly when more
than one possible interpretation is
available. The Memo further states,
‘‘When informed taxonomic opinion is
not unanimous, we evaluate available
published and unpublished information
and come to our own adequately
documented conclusion regarding the
validity of taxa.’’
The Act requires that we finalize,
modify, or withdraw the proposed rule
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within 12 months. The Act provides for
one 6-month extension for scientific
uncertainty, which we have used. As
such, we are required to make a
decision regarding the entities’
eligibility for listing at this time. In
addition, section 4(b)(1)(A) of the Act
requires the Service to make listing or
delisting decisions based on the best
scientific and commercial data
available. Further, our Policy on
Information Standards under the Act
(July 1, 1994; 59 FR 34271), the
Information Quality Act (section 515 of
the Treasury and General Government
Appropriations Act for Fiscal Year 2001
(Pub. L. 106–554; H.R. 5658)), and our
associated Information Quality
Guidelines (https://www.fws.gov/
informationquality), provide criteria,
guidance, and establish procedures to
ensure that our decisions are based on
the best scientific data available. They
require us, to the extent consistent with
the Act and with the use of the best
scientific data available, to use primary
and original sources of information as
the basis for recommendations. Primary
or original information sources are those
that are closest to the subject being
studied, as opposed to those that cite,
comment on, or build upon primary
sources. The Act and our regulations do
not require us to use only peer-reviewed
literature, but instead they require us to
use the ‘‘best scientific and commercial
data available.’’ We use information
from many different sources, including
articles in peer-reviewed journals,
scientific status surveys and studies
completed by qualified individuals,
Master’s thesis research that has been
reviewed but not published in a journal,
other unpublished governmental and
nongovernmental reports, reports
prepared by industry, personal
communication about management or
other relevant topics, conservation plans
developed by States and counties,
biological assessments, other
unpublished materials, experts’
opinions or personal knowledge, and
other sources.
We conducted a similar internal
review of the information presented by
and available to the Societies in their
review. Our review primarily focused
on Marsh et al. (2016), Carter et al.
(2016, in press), Copus et al. (2016),
Minckley and DeMarais (2000), and
Chafin et al. (2015), as well as other
literature as discussed above in the
Taxonomy section. In their most recent
publication of Common and Scientific
Names of Fishes (Page et al. 2013, p. 8),
the Societies state the following
regarding the common process of their
naming committee: ‘‘In accepting
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species as valid from various works, we
made little or no judgment on authors’
species concepts. Taxa of uncertain
status were dealt with on a case-by-case
basis.’’ Based on the Societies’ expertise
and their internal guidance (stated
above) on making such decisions, we
conclude that the preponderance of
evidence before them was clear and
decisive enough to make a taxonomic
change.
After reviewing the best available
scientific and commercial information
(as described above in the Taxonomy
section and summarized below) and
applying statutory and regulatory
guidance, we determined that the
Societies’ report considered the best
commercial and scientific data
available. We agree with the conclusion
that available data support recognition
of only one species, Gila robusta. Our
determination is based on various
factors, including the method of original
assignment to species, hybridization
events, conflicting identification of
species based on morphology versus
genetics, evolutionary history,
morphological identification
limitations, and lack of genetic markers
to identify species. We lack confidence
in the initial species assignments to G.
robusta, G. nigra, and G. intermedia due
to the scientific methods used (fish were
assigned to a species based on the
stream in which they occurred, the
erroneous assumption that these fish
did not overlap geographically, and the
absence of genetic or morphological
diagnostic information). Minckley and
DeMarais (2000, entire) based their
diagnostic key on the assumption that
none of these species occurs in the same
locality; however, they acknowledge
hybridization among G. robusta and G.
intermedia. Further, other studies have
found that fish designated as G. robusta,
G. nigra, and G. intermedia overlap
geographically or occur adjacent to one
another (Dowling and Marsh 2009, p. 1;
Marsh et al. 2016, p. 57; Brandenburg et
al. 2015, p. 18). In addition, some
populations appeared to conflict
genetically with the species-level
assignment based on morphology
(Dowling et al. 2015, pp. 14–15).
Multiple scientists (as described above)
found Minckley and DeMarais’s (2000,
entire) key for identification of G.
robusta, G. nigra, and G. intermedia to
not reliably differentiate among these
three fish. In Fossil Creek, Marsh et al.
(2016, entire) concluded there are two
morphologically similar, but genetically
distinguishable chub. However, there
are several genetic analysis studies
indicating population-level differences
among these fish, but the studies were
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16987
not able to identify genetic markers
distinguishing between the three fish.
¨
Finally, Schonhuth et al. (2014, p. 223)
found that G. robusta, G. nigra, and G.
intermedia were in one grouping that
included a common ancestor and all the
descendants (living and extinct) of that
ancestor (clade), and hypothesized this
could reflect incomplete lineage sorting
or hybridization, but this was not
studied.
For the purposes of our
determination, we accept the ‘‘single
species’’ finding by the Societies
described above and, consequently,
withdraw the proposed rule to list the
headwater chub (Gila nigra) and a DPS
of the roundtail chub (Gila robusta)
from the lower Colorado River basin as
threatened species under the Act. This
withdrawal is based on a thorough
review of the best scientific and
commercial data available, which
indicate that the headwater chub and
the DPS of the roundtail chub are not
discrete taxonomic entities and do not
meet the definition of species under the
Act. These fish are now recognized as a
single taxonomic species—the roundtail
chub (Gila robusta). Because the entities
previously proposed for listing are no
longer recognized as species, as defined
by the Act, we have determined that
they are not listable entities, and we are
withdrawing our proposed rule to list.
Future Actions
Following the publication of this
withdrawal, we intend to reevaluate the
status of the Gila chub (currently listed
as endangered) in the near future and
initiate a range-wide species status
assessment (SSA) of the newlyrecognized roundtail chub (Gila
robusta).
References Cited
A complete list of references cited in
this document is available on the
Internet at https://www.regulations.gov
and upon request from the Arizona
Ecological Services Office (see FOR
FURTHER INFORMATION CONTACT).
Authors
The primary authors of this document
are the staff members of the Arizona
Ecological Services Office.
Authority
The authority for this action is section
4(b)(6)(B)(ii) of the Endangered Species
Act of 1973, as amended (16 U.S.C. 1531
et seq.)
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Dated: March 21, 2017.
James W. Kurth,
Acting Director, U.S. Fish and Wildlife
Service.
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Agencies
[Federal Register Volume 82, Number 66 (Friday, April 7, 2017)]
[Proposed Rules]
[Pages 16981-16988]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2017-06995]
=======================================================================
-----------------------------------------------------------------------
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R2-ES-2015-0148; 4500030113]
RIN 1018-BA86
Endangered and Threatened Wildlife and Plants; Threatened Species
Status for the Headwater Chub and Roundtail Chub Distinct Population
Segment
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Proposed rule; withdrawal.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), withdraw the
proposed rule to list the headwater chub (Gila nigra) and a distinct
population segment (DPS) of the roundtail chub (Gila robusta) from the
lower Colorado River basin as threatened species under the Endangered
Species Act (Act). This withdrawal is based on a thorough review of the
best scientific and commercial data available, which indicate that the
headwater chub and
[[Page 16982]]
the roundtail chub DPS are not discrete taxonomic entities and do not
meet the definition of a species under the Act. These fish are now
recognized as a part of a single taxonomic species--the roundtail chub
(Gila robusta). Because the entities previously proposed for listing
are no longer recognized as species, as defined by the Act, we have
determined that they are not listable entities and we are withdrawing
our proposed rule to add them to the List of Endangered and Threatened
Wildlife.
Section 4(b)(6) of the Act and implementing regulations at 50 CFR
424.17 provide that the Service must, within 1 year of a proposed rule
to list, delist, or reclassify species, or to designate or revise
critical habitat, withdraw the proposal if the available evidence does
not justify the proposed action. The document withdrawing the rule must
set forth the basis upon which the proposed rule has been found not to
be supported by available evidence. Once withdrawn, the action may not
be re-proposed unless sufficient new information is available.
FOR FURTHER INFORMATION CONTACT: Steve Spangle, Field Supervisor, U.S.
Fish and Wildlife Service, Arizona Ecological Services Office, 9828
North 31st Ave., #C3, Phoenix, AZ 85051-2517; telephone 602-242-0210.
Persons who use a telecommunications device for the deaf (TDD) may call
the Federal Relay Services at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Previous Federal Action
On October 7, 2015 (80 FR 60754), we published a proposed rule to
list the headwater chub and the lower Colorado River basin roundtail
chub DPS (roundtail chub DPS) as threatened species under the Act (16
U.S.C. 1531 et seq.). On August 15, 2016 (81 FR 54018), we announced a
6-month extension on the final listing determination that the Act
allows when there is substantial disagreement regarding the sufficiency
or accuracy of the available data, and reopened the comment period on
the proposed listings for 30 days. During this comment period we
received new information. On November 1, 2016 (81 FR 75801), we
reopened the comment period on the proposed listings for an additional
45 days to provide the public additional time to review and consider
the proposed rulemakings in light of this new information. As a result
of the 6-month extension, the deadline to finalize, modify, or withdraw
the proposed rule is April 7, 2017.
For a description of additional previous Federal actions concerning
these species, please refer to the October 7, 2015, proposed listing
rule (80 FR 60754).
Background
At the time we published our proposed rule (October 7, 2015; 80 FR
60754), the Committee on Names of Fishes, a joint committee of the
American Fisheries Society and American Society of Ichthyologists and
Herpetologists (the Societies) (Page et al. 2013, p. 71), considered
headwater chub and roundtail chub to be separate species. As a
consortium of fisheries scientists, the American Fisheries Society is
the recognized and accepted scientific authority on fish taxonomy.
Accordingly, our proposed rule assessed the headwater chub and
roundtail chub as separate species. However, commenters on our proposed
rule raised questions during the public comment period regarding the
taxonomic distinctness of the headwater and roundtail chubs, as related
to the Gila chub (Gila intermedia). At that time, some scientists
knowledgeable about the fish contended that the three entities were not
separate species (Carter et al. 2016 in press; Copus et al. 2016). For
this reason, the Arizona Game and Fish Department requested that the
Societies evaluate the most recent literature associated with roundtail
chub, headwater chub, and Gila chub taxonomy. In their final report to
the Arizona Game and Fish Department, the Societies panel concluded
that ``no morphological or genetic data define populations of Gila in
the lower Colorado River basin (which, as defined by the Service,
includes the Little Colorado River, Bill Williams River, Gila River,
Verde River, and Salt River drainages) as members of more than one
species'' and ``that the data available support recognition of only one
species of Gila, the roundtail chub, Gila robusta'' (Page et al. 2016,
p. 1). These three fish are now considered by the Societies to be a
single species, roundtail chub (Gila robusta) because data do not
support recognition of three species.
Taxonomy
Introduction
The taxonomic history of the genus Gila in the Colorado River basin
has changed over time, especially for the three forms (roundtail,
headwater, and Gila chub) found in the Gila River basin. These forms
have been variously classified as full species, assigned as different
species, subspecies of Gila robusta, or as part of a ``Gila robusta
complex'' (Miller 1945; Holden 1968; Rinne 1969; Holden and Stalnaker
1970; Rinne 1976; Smith et al. 1977; DeMarais 1986; Rosenfeld and
Wilkinson 1989; Dowling and DeMarais 1993; Douglas et al. 1998;
Minckley and DeMarais 2000; Gerber et al. 2001). As noted by nearly all
researchers investigating the systematics of Gila spp., the taxonomic
situation is complicated and problematic (Holden and Stalnaker 1970;
Minckley 1973; Minckley and DeMarais 2000; Gerber et al. 2001;
Sch[ouml]nhuth et al. 2014) due to various factors including multiple
independent hybridization events over time (Rinne 1976; DeMarais 1986;
Rosenfeld and Wilkinson 1989; DeMarais et al. 1992; Dowling and
DeMarais 1993; Minckley and DeMarais 2000; Gerber et al. 2001; Schwemm
2006; Sch[ouml]nhuth et al. 2014; Brandenburg et al. 2015,) potential
past introgression (the transfer of genetic information from one
species to another as a result of hybridization between them and
repeated backcrossing) (DeMarais et al. 1992; Minckley and DeMarais
2000), recent divergence within the three fish (Schwemm 2006). Further,
the original assignment to species was based on the assumption that the
three fish do not overlap geographically (parapatry), which we
recognize now is not an accurate assumption. Additionally, in some
instances when the same fish was identified based on morphology
(physical characteristics) it was identified as one species and when
identified based on genetic analysis it was identified as a different
species (Dowling et al. 2015, pp. 14-15). Recent and ongoing genetic
and morphologic analyses of chubs in the Gila River basin continue to
yield conflicting results (DeMarais et al. 1992; Schwemm 2006; Dowling
et al. 2008 and 2015; Sch[ouml]nhuth et al. 2014; Marsh et al. 2016,
all entire).
History
Gila robusta (roundtail chub) was first described by Baird and
Girard (1853, p. 365-369) from specimens collected in 1851 from the
Zuni River (tributary to Little Colorado River). Gila nigra (headwater
chub; formerly known as G. robusta graham or G. grahami) was first
described as a subspecies (G. robusta graham) from Ash Creek in the San
Carlos River in east-central Arizona in 1874 (Cope and Yarrow 1875, p.
663), but not returned to full species status (G. robusta) until
proposed so by Minckley and DeMarais (2000, p. entire). The Societies
accepted Gila nigra as a full species (Nelson et al. 2004, p. 71), as
did the New Mexico Department of Game, Fish (Carman 2006, p. 3),
Arizona Game, and Fish
[[Page 16983]]
Department (AGFD 2006, p. 3) and continued to recognize G. robusta as a
distinct species. Therefore, based on the best available commercial and
scientific data the Service accepted both Gila robusta and Gila nigra
as full species as documented in our 12-month findings (May 3, 2006; 71
FR 26007 and July 7, 2009; 74 FR 32352). In their 2013 publication of
Common and Scientific Names of Fishes from the United States, Canada,
and Mexico, the Societies continued to list both Gila robusta and Gila
nigra as distinct species (Page et al. 2013, p. 71). A summary of the
historic and current nomenclature from Rinne (1976, entire), Sublette
et al. (1990, entire), and Minckley and DeMarais (2000, entire) is
summarized in Voeltz (2002, pp. 8) and Copus et al. (2016, pp. 1&6).
The Gila chub (Gila intermedia) is currently listed as an endangered
species (November 2, 2005; 70 FR 66664).
These entities were originally classified based on the streams in
which they were found (Minckley and DeMarais 2000, p. 252), under the
assumption that G. robusta and G. nigra either did not overlap
(allopatric, no gene flow) or there was only a narrow overlap
(parapatric; limited interaction and opportunity for gene flow)
(Minckley and DeMarais 2000 pp. 252-254). Because hybridization between
G. robusta and G. intermedia indicates that these fish must co-occur in
some streams (Minckley and DeMarais 2000, entire), we conclude that
Minckley and DeMarais's (2000) assumption they did not overlap was
unfounded. Further, other studies have found that fish designated as G.
robusta, G. nigra, and G. intermedia overlap geographically or occur
adjacent to one another (Dowling and Marsh 2009, p. 1; Marsh et al.
2016, p. 57; Brandenburg et al. 2015, p. 18).
Morphology
The approach for classifying G. robusta, G. nigra, and G.
intermedia developed by Minckley and DeMarais (2000, pp. 254-255)
presumes there is little intraspecific variation (differences within a
species) in the morphologic and meristic (counting quantitative
characteristics such as fins) characteristics used to distinguish these
three taxa. However, the three purported species overlap in physical
characteristics, and many fish have intermediate physical
characteristics. Those characteristics that do not overlap are
separated by very small margins, making species-level identification of
individual fish problematic, even when the geographic origin of the
species is known (Brandenburg 2015, entire). Minckley and DeMarais
(2000, pp. 253-254) indicate that G. nigra is physically different from
G. intermedia even though they appear physically more similar to one
another than either is to G. robusta. In addition, Copus et al. (2016,
p. 13) did not find physical characteristics in the Minckley and
DeMarais (2000, pp. 254-255) classification key to reliably
differentiate G. robusta, G. nigra, and G. intermedia from one another.
Copus et al. (2016 p. 16) concluded that there was no morphological
basis for taxonomic distinctions within the Gila spp. complex.
Genetics
Multiple genetic analysis studies have been conducted that reveal
differences between different chub populations, but have been unable to
identify differences between G. robusta, G. nigra, and G. intermedia
(DeMarais et al. 1992, pp. 2748-2749; Schwemm 2006, p. 29; Dowling et
al. 2008, p. 2, and 2015, p. 13; Copus et al. 2016, pp. 14-15; Marsh et
al, 2016, p.58). Mitochondrial DNA analysis (Sch[ouml]nhuth et al.
2014, p. 223) indicates that G. robusta, G. nigra, and G. intermedia
belong to one clade (a grouping that includes a common ancestor and all
its descendants, living and extinct, of that ancestor). Sch[ouml]nhuth
et al. (2014, p. 223) hypothesized that this could reflect
hybridization or incomplete lineage sorting (when the lineage of a
specific gene is not the same as the lineage of the species, obscuring
the true species relationship).
However, when nuclear DNA (rather than mitochondrial DNA) was
analyzed, a broader grouping was identified that included G. seminude
and G. elegans, but when mitochondrial and nuclear DNA results are
combined G. robusta, G. nigra, and G. intermedia were in one grouping
(Sch[ouml]nhuth et al. 2014, p. 223). Preliminary studies by Chafin et
al. (2016) indicate evolutionary independent lineages for G. robusta,
G. nigra, and G. intermedia, and that the hybrid origin of G. nigra is
not supported. Studies by Marsh et al. (2016, entire) point to genetic
variation between populations of G. robusta and G. nigra, and
demonstrate evidence that distinct ecological differences between some
populations are now thought to exist. Minckley and DeMarais (2000,
entire) supported recognition of three species, but acknowledged that
most genetic variation was within populations for G. robusta, and was
among populations for G. intermedia and G. nigra. Minckley and DeMarais
(2000, p. 253) also indicated that these three fishes share genetic
features (that had been studied so far) while behaving as separate non-
overlapping (allopatric) morphological species. In addition, some
populations assigned to species based on genetics appeared to conflict
with the species level-assignment based on morphology (Dowling et al.
2008, p. 27).
Speciation
Minckley and DeMarais (2000, p. 253) describe three different
taxonomic options for chubs in the Gila River basin: a single species
with many different forms or stages (polymorphic species), a species
containing multiple subspecies, or three full species. They acknowledge
that none of these taxonomic options is biologically justified without
knowing if these fish naturally occur in the same geographic area
(sympatry, indicating an initial interbreeding population that split),
or occur immediately adjacent to each other but not significantly
overlapping (parapatry, indicating there is no barrier to gene flow).
They further acknowledge that a persistent narrow interaction zone
(parapatry, indicating there is no barrier to gene flow) of
morphologically distinguishable G. robusta, G. intermedia, and G. nigra
has been confirmed, but note that in no instance was any two of the
three caught at the same locality (allopatric, no gene flow; p. 251).
However, they also acknowledge that hybridization (between G. robusta
and G. intermedia, resulting in G. nigra) in the past must have
occurred in some places and not others, thereby demonstrating
occurrence in the same geographic area (sympatry) (p. 253). They
conversely hypothesized that the current minimal overlap in an area
where species are adjacent (parapatry, indicating there is no barrier
to gene flow) may thus reflect an ancestral ecological segregation area
(sympatry, indicating an initial interbreeding population that split
due to the use of different habitats and resources) that promoted
persistence in the ever-increasing aridity of the Southwest (p. 253).
In Fossil Creek, G. nigra and G. robusta appear to be sympatric,
including hybrids between G. robusta and G. nigra (Marsh et al. 2016,
p. 57). Brandenburg et al. (2015, p. 18) concluded that the
morphological assessment of Gila spp. in New Mexico confirmed that the
three fish were found in the same geographic area (sympatric) in almost
all cases, contradicting Minckley and DeMarais' results (2000, p. 251)
as well as other previous literature suggesting that these Gila spp.
are occurring in separate non-overlapping geographical areas
(allopatric) through their ranges (Rinne
[[Page 16984]]
1969, p. entire; DeMarais 1986, p. entire; Minckley and DeMarais 2000,
p. 253). In Fossil Creek, they found that G. nigra and G. robusta are
locally in the same geographic area (sympatric) and have hybridized
(Marsh et al. 2016, p. 57). Marsh et al. (2016, p. 58) concluded there
are two morphologically similar, but genetically distinguishable, chub
in Fossil Creek, G. robusta and G. nigra.
Conservation Implications
Dowling et al. (2015, pp. 14-15) reasoned that the lack of
diagnostic molecular characteristics does not inform the status of
these three fish, but rather highlights the role that local evolution
has played in shaping patterns of variation in these taxa and the
importance of accounting for this variation when managing the complex.
Most, if not all, scientists agree that conservation actions for these
chubs must be directed at the population level and must include
consideration of the complex as a whole (Dowling et al. 2008, pp. 30-
31; Dowling and DeMarais 1993, p. 445; Gerber et al. 2001, p. 2037;
Schwemm et al. 2006, pp. 32-33). The Arizona Game and Fish Department
recognizes the importance of conserving the currently recognized
roundtail chub population rangewide (including the formerly known
headwater chub and Gila chub) and is committed to the conservation
agreements and practices that have been in place since 2006 (AGFD 2017,
entire; AGFD 2006, entire).
Public Comments
In our October 7, 2015 proposed rule (80 FR 60754), we requested
that all interested parties submit comments or information concerning
the proposed listings during a 60 day comment period, ending December
7, 2015. We particularly sought comments concerning genetics and
taxonomy. In our August 15, 2016, 6-month extension document (81 FR
54018), we reopened the comment period on the proposed rule for 30
days, ending September 14, 2016, and we again requested comments and
information regarding genetics and morphology that would aid in
resolving the ongoing taxonomic issues regarding classification of
these fish. On November 1, 2016 (81 FR 75801, we announced an
additional 45-day comment period, ending December 16, 2016, on the
October 7, 2015 proposed rule.
We provided notification of these publications and their comment
periods through email, letters, and news releases faxed and/or mailed
to the appropriate Federal, State, and local agencies; county
governments; elected officials; media outlets; local jurisdictions;
scientific organizations; interested groups; and other interested
parties.
In accordance with our peer review policy published in the Federal
Register on July 1, 1994 (59 FR 34270), we solicited independent
opinions from at least three knowledgeable individuals who have
expertise with these fish, who possess a current knowledge of the
geographic region where the fish occurs, and/or are familiar with the
principles of conservation biology.
We reviewed all comments received from peer reviewers and the
public for substantive issues and new information regarding the
proposed listing of G. nigra and the G. robusta DPS. Substantive
comments pertaining to the taxonomy of these fish received during the
comment period are addressed below. We also received several comments
from both the public and peer reviewers concerning threats to these
fish; however, because our withdrawal is due to taxonomic revision such
comments are outside the scope of this withdrawal.
Peer Review Comments
(1) Comment: One peer reviewed stated that there are no recent
(since 2000) publications in the peer-reviewed literature that provide
evidence that Gila intermedia, G. nigra, and G. robusta are other than
separate and distinct species. The peer reviewer further stated that
there are articles that study the genetics or morphology of these fish
without questioning its taxonomy, specifically Sch[ouml]nhuth et al.
2014, Sch[ouml]nhuth et al. 2012, and Marsh et al. in press.
Response: Multiple studies since 2000 provide information on the
genetic analysis for these fish, including Schwemm 2006, Dowling et al.
2008 and 2015, and Copus et al. 2016. While these studies may not have
questioned the taxonomic classification, they also have not been able
to identify genetic markers that have the ability to distinguish among
G. robusta, G. nigra, and G. intermedia. Sch[ouml]nhuth et al. (2008,
p. 213; 2014, p. 223), using mitochondrial and nuclear DNA sequencing,
found that G. robusta, G. nigra, and G. intermedia were well supported
as having a common ancestor. Using mitochondrial DNA, Sch[ouml]nhuth et
al. (2008, p. 213; 2014, p. 223) found that G. robusta, G. nigra, and
G. intermedia were in one grouping that included a common ancestor and
all the descendants (living and extinct) of that ancestor (clade), and
this could reflect incomplete lineage sorting or hybridization.
However, when nuclear DNA was analyzed, a broader grouping was
identified that included G. seminuda and G. elegans, but when
mitochondrial and nuclear DNA results were combined, G. robusta, G.
nigra, and G. intermedia were alone in one grouping. While Marsh et al.
(2016, entire) concluded there are two similar but genetically
distinguishable species in the creek they studied, their findings
differ somewhat from Schwemm (2006) and Dowling et al. (2008 and 2015,
entire), who were unable to conclusively identify distinct species
using genetic markers across a much wider range. Further, the Societies
conducted a review of the literature and found no evidence to support
three species. The Service has reviewed the best available scientific
and commercial data and also found a lack of sufficient evidence to
support more than one species.
(2) Comment: Recognized authorities on the taxonomy and ecology of
these fish recognized these fish as separate species based on
morphological diagnostics.
Response: Minckley and DeMarais (2000), Miller et al. (2005), and
Minckley and Marsh (2009) report identification of three species using
a diagnostic morphological key. However, additional reports were unable
to reliably identify these three fish to species using the same
diagnostic key (Carter et al. 2016, p. 2 and 20, in press; Brandenburg
2015, entire; Copus et al. 2016, p. 13). Further, Minckley and DeMarais
(2000, pp. 253-254) stated that G. nigra is morphologically separate
from G. intermedia, but that G. nigra and G. intermedia appear
morphologically more similar to one another than either is to G.
robusta. In addition to issues surrounding morphological
identification, multiple genetic analysis studies have found
population-level differences, but have been unable to identify genetic
markers that have the ability to distinguish among G. robusta, G.
nigra, and G. intermedia (DeMarais 1992, pp. 2748-2749; Schwemm 2006,
p. 29; Dowling et al. 2008, p. 2, and 2015, p. 13; Copus et al. 2016,
pp. 14-15). There are also the findings of Sch[ouml]nhuth et al.
(2014), Sch[ouml]nhuth et al. (2012) as described in Response to
Comment 1.
(3) Comment: Conclusions are mainly based on two ``gray
literature'' reports that have not undergone peer review (Copus et al.
2016) or were not available for public consideration (Carter et al.
2016, in press).
Response: Section 4(b)(1)(A) of the Act requires the Service to
make listing or delisting decisions based on the best scientific and
commercial data available. Further, our Policy on Information Standards
under the Act
[[Page 16985]]
(July 1, 1994; 59 FR 34271), the Information Quality Act (section 515
of the Treasury and General Government Appropriations Act for Fiscal
Year 2001 (Pub. L. 106-554; H.R. 5658)), and our associated Information
Quality Guidelines (https://www.fws.gov/informationquality), provide
criteria and, guidance, and establish procedures to ensure that our
decisions are based on the best scientific data available. They require
us, to the extent consistent with the Act and with the use of the best
scientific data available, to use primary and original sources of
information as the basis for our determinations. Primary or original
information sources are those that are closest to the subject being
studied, as opposed to those that cite, comment on, or build upon
primary sources. The Act and our regulations do not require us to use
only peer-reviewed literature, but instead they require us to use the
``best scientific and commercial data available.'' We use information
from many different sources, including articles in peer-reviewed
journals, scientific status surveys and studies completed by qualified
individuals, Master's thesis research that has been reviewed but not
published in a journal, other unpublished governmental and
nongovernmental reports, reports prepared by industry, personal
communication about management or other relevant topics, conservation
plans developed by States and counties, biological assessments, other
unpublished materials, experts' opinions or personal knowledge, and
other sources. For these reasons, we think it is appropriate to include
review of Copus et al. (2016) and Carter et al. (2016, in press), as
well as other sources, within our review.
(4) Comment: Several authors presented data and conclusions that
conflicted with the previously cited Carter et al. (2016, in press) and
Copus et al. (2016) reports pertaining to morphological identification,
DNA analysis, and ecological equivalency to a subset of the Joint
Committee convened in April 2016, to specifically address the taxonomy
of the roundtail chub complex.
Response: We were present at the April 2016 Joint Committee
webinar, and experts beyond Carter and Copus, such as Brandenburg,
Schwemm, Dowling, O'Neill, and Chafin, also provided information based
on research they either had previously conducted or are currently
conducting on Gila. A complete list of references cited may be obtained
on the Internet at https://www.regulations.gov and upon request from the
Arizona Ecological Services Office (see FOR FURTHER INFORMATION
CONTACT). The Service has reviewed the best available scientific and
commercial data and found a lack of sufficient evidence to support more
than one species.
(5) Comment: This taxonomic dispute is not simply an academic
exercise of whether to lump or split taxa, because the decision has
enormous implications for the conservation of imperiled species.
Multiple experts recommended that the roundtail chub complex, however
it is constituted, be managed as separate populations or managed as a
complex.
Response: The Service recognizes that multiple experts agree that
conservation actions must be directed at the population level and must
include consideration of the complex as a whole (Dowling et al. 2008,
pp. 30-31; Dowling and DeMarais 1993, p. 445; Gerber et al. 2001, p.
2037; Schwemm 2006, pp. 32-33). However, the Service must adhere to the
Act and its implementing regulations, which define a ``species'' as any
species or subspecies of fish, wildlife, or plant, and any distinct
population segment of any vertebrate species which interbreeds when
mature (16 U.S.C. 1532(16) and 50 CFR 424.02). The best available
scientific and commercial data as discussed above in the Taxonomy
section, support recognition of only one species, Gila robusta. The
Service's withdrawal of our proposed rule to list the headwater and
roundtail chub based on new taxonomic classification does not diminish
the conservation efforts of our partners to conserve this species and
habitat, nor does our decision affect the State's ability to conserve
this species under its own authority. The Arizona Game and Fish
Department recognizes the importance of conserving the currently
recognized roundtail chub population rangewide (including the formerly
known headwater chub and Gila chub) and is committed to the
conservation agreements and practices that have been in place since
2006 (AGFD 2017, entire; AGFD 2006, entire).
(6) Comment: Multiple commenters raised concerns with Copus et al.
(2016) methods and conclusions, particularly small sample size, lack of
key analytical and laboratory steps, the study's DNA sequence data
filtering and analyses that failed to follow best practices for
phylogenetic analysis, and specimen shrinkage associated with duration
of preservation impacting morphological diagnostics.
Response: The Service did not rely solely on Copus et al. 2016. We
considered the best available commercial and scientific data; you may
obtain a complete list of references cited on the Internet at https://www.regulations.gov and upon request from the Arizona Ecological
Services Office (see FOR FURTHER INFORMATION CONTACT). In regards to
the mitochondrial DNA and phylogenetic analysis, Copus et al.'s
findings are consistent with Sch[ouml]nhuth et al.'s (2014) and
Sch[ouml]nhuth et al.'s (2012) mitochondrial DNA and phylogenetic
analysis. In addition, multiple genetic analysis studies have been
conducted that indicate population-level differences, but do not
identify genetic markers that have the ability to distinguish among G.
robusta, G. nigra, and G. intermedia (DeMarais 1992, pp. 2748-2749;
Schwemm 2006, p. 29; Dowling et al. 2008, p. 2, and 2015, p. 13).
In regards to morphological diagnostic errors due to using
preserved specimens, Copus et al. (2016) did use preserved specimens.
However, they also analyzed fresh material and concluded that no single
diagnostic character can be used for species identification, and with
considerable overlap among species in every morphological character, no
suite of characters can distinguish species unambiguously (Copus et al.
2016, p. 13). Brandenburg et al. (2015, entire) also reported overlap
in the meristic and morphometric characteristics, records of many
individual fish with intermediate physical characteristics, and even
those characters that do not overlap are separated by very small
margins making species-level identification of individual fish
problematic, even when the geographic origin of the species is known.
Public Comments
(7) Comment: Multiple commenters requested various listing
alternatives under the Act including: List G. robusta as threatened and
encompass all populations of the chub complex within the Gila basin
requiring a revision of the recovery plan, list G. robusta and G. nigra
as threatened and retain the current endangered species status of G.
intermedia, list G. robusta as threatened and retain the current
endangered species status of G. intermedia, or other combinations.
Response: The Service must adhere to the Act and its implementing
regulations, which define a ``species'' as any species or subspecies of
fish, wildlife, or plant, and any distinct population segment of any
vertebrate species which interbreeds when mature (16 U.S.C. 1532(16)
and 50 CFR 424.02), and based on our review, the best available
scientific and commercial data
[[Page 16986]]
support recognition of only one species, Gila robusta. As the headwater
chub and roundtail chub DPS no longer meet the definition of a
``species'' under the Act, we must withdraw our proposed rule to list
them as threatened species.
(8) Comment: Multiple commenters stated that there is a great
amount of morphological overlap among counts and measures for these
chub taxa and that this has long been recognized. If a taxonomic key is
not 100 percent correct, that does not necessarily mean that these are
not taxa that are biologically distinct at the specific level. A test
of the key would require the a priori identification of each individual
to species. Rather than dismiss the species' taxonomic status,
biologists should be working to make a better key that can be used in
the field for the effective identification and management of the
species.
Response: We recognize that diagnostic keys do not produce correct
results all the time, whether due to human error or morphological
similarities among purported species. However, Copus et al. (2016, p.
13) concluded that, based on genetic analysis, no single diagnostic
character can be used for species identification, and with considerable
overlap among species in every morphological character, no suite of
characters can distinguish species unambiguously. Brandenburg et al.
(2015, entire) also reported overlap in the meristic and morphometric
characteristics, and there are many individual fish whose morphology
resides on an intermediate spectrum, and even those characters that do
not overlap are separated by very small margins, making species-level
identification of individual fish problematic, even if the geographic
origin of the species is known. In regards to a priori identification
of fish, assignment to species has been based on the stream in which
the fish occurs (Minckley and DeMarais 2000, p. 252), so the
identification of the fish that occurs in each stream is assumed to be
known. Consequently, there exists the ability to compare findings from
the diagnostic key to the fish within a particular stream. An updated
key may be prudent; however, the Service must use the best available
scientific and commercial data available, and we have concluded from
our review that the data currently support only one species, Gila
robusta. Further, given the overlap in diagnostic characteristics, the
development of a valid key seems unlikely.
(9) Comment: Multiple commenters stated that it has long been
hypothesized that G. nigra formed as the result of hybridization
between the other two taxa, so we would expect the greatest
morphological overlap from that species with the other two taxa. The
question then becomes, is G. nigra continuing to differentiate from
ancestral G. robusta? When in sympatry, G. nigra and G. robusta are
becoming increasingly reproductively isolated from one another (Desert
Fishes Council meeting, Dowling et al. 2016).
Response: We recognize that multiple studies have indicated that
hybridization has occurred among G. intermedia and G. robusta resulting
in G. nigra and that continuing evolution may occur (Schwemm 2006;
Dowling et al. 2008, entire). However, there has also been information
presented showing no evidence of the hybrid origin of G. nigra, and
that G. intermedia and G. nigra evolved separately in non-overlapping
areas (parapatry) (Chafin 2016, entire). In addition, past research
(Dowling et al. 2008, 2015; Schwemm 2006) indicate that there is more
variation among populations and unique genetics within specific
populations (streams).
(10) Comment: If only G. robusta and G. intermedia are evaluated,
there is no question that they would be considered distinct
morphological species.
Response: Carter et al. (2016, in press) found that the physical
characteristics did not reliably differentiate among G. robusta, G.
intermedia, and G. nigra. In addition, Brandenburg et al. (2015, pp. 8-
9) found physically similarity of the three species, as numerous
individuals exhibited intermediate characters along the species
gradient. The discriminant function analysis (a statistical analysis
tool to determine which variables discriminate between two or more
naturally occurring groups) classified only 16 percent (n = 42) of G.
intermedia (the fewest) while the majority of the samples were
classified as G. robusta (53.2 percent, n = 140), which indicates that
the ability to classify these fish correctly to G. intermedia or G.
robusta based on physical characteristics was low. Due to the complex
genetic makeup and observable characteristics or traits (i.e., physical
appearance, behavior, or physiology) of these species, there are some
stream locations where we do not know where the geographic overlap of
headwater, roundtail, and, in some cases Gila chub, begins and ends,
because of the plasticity of observable characteristics or traits of
these fish within individual streams. Our review of the data does
indicate that there are differences in observable characteristics or
traits between the fish in different streams, but the Societies'
review, as well as the Service review, of the best available scientific
and commercial data did not result in a species-level differentiation
between G. robusta and G. intermedia, or among G. robusta, G.
intermedia, and G. nigra.
(11) Comment: One commenter recommend that we proceed with an
amended recovery plan to list the status of this species as threatened
under the Act. The listing of this species is necessary even if all
populations of G. intermedia and G. nigra are subsumed into G. robusta.
Response: An assessment of the entire range of the new taxonomic
group of roundtail chub is planned. We are initiating a status review
of the new taxonomic entity in 2 to 4 years. Following that review, we
will take action as appropriate.
Determinations
An entity may only be listed under the Act if that entity meets the
Act's definition of a species. The recent report by the Societies
indicates that neither the headwater chub nor the roundtail chub can be
considered species, as defined by the Act. Under section 3 of the Act
(16 U.S.C. 1532(16)) and associated implementing regulations at 50 CFR
424.02, a ``species'' is defined to include any species or subspecies
of fish, wildlife, or plant, and any distinct population segment of any
vertebrate species which interbreeds when mature. The Act's
implementing regulations at 50 CFR 424.11(a) and the Service Director's
November 25, 1992, ``Taxonomy and the Endangered Species Act''
Memorandum (Memo) provide additional guidance on how to consider
taxonomic information when assessing a species for listing under the
Act. The regulations at 50 CFR 424.11(a) state, ``In determining
whether a particular taxon or population is a species for the purposes
of the Act, the Secretary [of the Interior] shall rely on standard
taxonomic distinction and the biological expertise of the Department
[of the Interior] and the scientific community concerning the relevant
taxonomic group.'' The Director's Memo specifies that the Service is
``required to exercise a degree of scientific judgment regarding the
acceptance of taxonomic interpretations, particularly when more than
one possible interpretation is available. The Memo further states,
``When informed taxonomic opinion is not unanimous, we evaluate
available published and unpublished information and come to our own
adequately documented conclusion regarding the validity of taxa.''
The Act requires that we finalize, modify, or withdraw the proposed
rule
[[Page 16987]]
within 12 months. The Act provides for one 6-month extension for
scientific uncertainty, which we have used. As such, we are required to
make a decision regarding the entities' eligibility for listing at this
time. In addition, section 4(b)(1)(A) of the Act requires the Service
to make listing or delisting decisions based on the best scientific and
commercial data available. Further, our Policy on Information Standards
under the Act (July 1, 1994; 59 FR 34271), the Information Quality Act
(section 515 of the Treasury and General Government Appropriations Act
for Fiscal Year 2001 (Pub. L. 106-554; H.R. 5658)), and our associated
Information Quality Guidelines (https://www.fws.gov/informationquality), provide criteria, guidance, and establish
procedures to ensure that our decisions are based on the best
scientific data available. They require us, to the extent consistent
with the Act and with the use of the best scientific data available, to
use primary and original sources of information as the basis for
recommendations. Primary or original information sources are those that
are closest to the subject being studied, as opposed to those that
cite, comment on, or build upon primary sources. The Act and our
regulations do not require us to use only peer-reviewed literature, but
instead they require us to use the ``best scientific and commercial
data available.'' We use information from many different sources,
including articles in peer-reviewed journals, scientific status surveys
and studies completed by qualified individuals, Master's thesis
research that has been reviewed but not published in a journal, other
unpublished governmental and nongovernmental reports, reports prepared
by industry, personal communication about management or other relevant
topics, conservation plans developed by States and counties, biological
assessments, other unpublished materials, experts' opinions or personal
knowledge, and other sources.
We conducted a similar internal review of the information presented
by and available to the Societies in their review. Our review primarily
focused on Marsh et al. (2016), Carter et al. (2016, in press), Copus
et al. (2016), Minckley and DeMarais (2000), and Chafin et al. (2015),
as well as other literature as discussed above in the Taxonomy section.
In their most recent publication of Common and Scientific Names of
Fishes (Page et al. 2013, p. 8), the Societies state the following
regarding the common process of their naming committee: ``In accepting
species as valid from various works, we made little or no judgment on
authors' species concepts. Taxa of uncertain status were dealt with on
a case-by-case basis.'' Based on the Societies' expertise and their
internal guidance (stated above) on making such decisions, we conclude
that the preponderance of evidence before them was clear and decisive
enough to make a taxonomic change.
After reviewing the best available scientific and commercial
information (as described above in the Taxonomy section and summarized
below) and applying statutory and regulatory guidance, we determined
that the Societies' report considered the best commercial and
scientific data available. We agree with the conclusion that available
data support recognition of only one species, Gila robusta. Our
determination is based on various factors, including the method of
original assignment to species, hybridization events, conflicting
identification of species based on morphology versus genetics,
evolutionary history, morphological identification limitations, and
lack of genetic markers to identify species. We lack confidence in the
initial species assignments to G. robusta, G. nigra, and G. intermedia
due to the scientific methods used (fish were assigned to a species
based on the stream in which they occurred, the erroneous assumption
that these fish did not overlap geographically, and the absence of
genetic or morphological diagnostic information). Minckley and DeMarais
(2000, entire) based their diagnostic key on the assumption that none
of these species occurs in the same locality; however, they acknowledge
hybridization among G. robusta and G. intermedia. Further, other
studies have found that fish designated as G. robusta, G. nigra, and G.
intermedia overlap geographically or occur adjacent to one another
(Dowling and Marsh 2009, p. 1; Marsh et al. 2016, p. 57; Brandenburg et
al. 2015, p. 18). In addition, some populations appeared to conflict
genetically with the species-level assignment based on morphology
(Dowling et al. 2015, pp. 14-15). Multiple scientists (as described
above) found Minckley and DeMarais's (2000, entire) key for
identification of G. robusta, G. nigra, and G. intermedia to not
reliably differentiate among these three fish. In Fossil Creek, Marsh
et al. (2016, entire) concluded there are two morphologically similar,
but genetically distinguishable chub. However, there are several
genetic analysis studies indicating population-level differences among
these fish, but the studies were not able to identify genetic markers
distinguishing between the three fish. Finally, Sch[ouml]nhuth et al.
(2014, p. 223) found that G. robusta, G. nigra, and G. intermedia were
in one grouping that included a common ancestor and all the descendants
(living and extinct) of that ancestor (clade), and hypothesized this
could reflect incomplete lineage sorting or hybridization, but this was
not studied.
For the purposes of our determination, we accept the ``single
species'' finding by the Societies described above and, consequently,
withdraw the proposed rule to list the headwater chub (Gila nigra) and
a DPS of the roundtail chub (Gila robusta) from the lower Colorado
River basin as threatened species under the Act. This withdrawal is
based on a thorough review of the best scientific and commercial data
available, which indicate that the headwater chub and the DPS of the
roundtail chub are not discrete taxonomic entities and do not meet the
definition of species under the Act. These fish are now recognized as a
single taxonomic species--the roundtail chub (Gila robusta). Because
the entities previously proposed for listing are no longer recognized
as species, as defined by the Act, we have determined that they are not
listable entities, and we are withdrawing our proposed rule to list.
Future Actions
Following the publication of this withdrawal, we intend to
reevaluate the status of the Gila chub (currently listed as endangered)
in the near future and initiate a range-wide species status assessment
(SSA) of the newly-recognized roundtail chub (Gila robusta).
References Cited
A complete list of references cited in this document is available
on the Internet at https://www.regulations.gov and upon request from the
Arizona Ecological Services Office (see FOR FURTHER INFORMATION
CONTACT).
Authors
The primary authors of this document are the staff members of the
Arizona Ecological Services Office.
Authority
The authority for this action is section 4(b)(6)(B)(ii) of the
Endangered Species Act of 1973, as amended (16 U.S.C. 1531 et seq.)
[[Page 16988]]
Dated: March 21, 2017.
James W. Kurth,
Acting Director, U.S. Fish and Wildlife Service.
[FR Doc. 2017-06995 Filed 4-6-17; 8:45 am]
BILLING CODE 4333-15-P