[Federal Register Volume 70, Number 242 (Monday, December 19, 2005)]
[Rules and Regulations]
[Pages 75071-75074]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 05-24119]
=======================================================================
-----------------------------------------------------------------------
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife and Plants; 12-Month Finding
on a Petition to List Cicurina cueva (No Common Name) as an Endangered
Species
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of 12-month petition finding.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), announce a
12-month finding on a petition to list a karst meshweaver (spider),
Cicurina cueva (no common name), under the Endangered Species Act of
1973, as amended. Since receiving the petition, both a genetic
assessment and a re-assessment of morphological characters have failed
to support the distinctness of C. cueva from two other named Cicurina,
C. bandida and C. reyesi. After reviewing all available scientific and
commercial information, we find that current information available to
us does not support the taxonomic standing of C. cueva as a species,
and therefore it is not a listable entity and listing is therefore not
warranted.
DATES: The finding announced in this document was made on December 19,
2005.
ADDRESSES: The complete file for this finding is available for
inspection, by appointment, during normal business hours at the Austin
Ecological Services Field Office, 10711 Burnet Rd., Suite 200, Austin,
Texas 78758. Please submit any new information, materials, comments, or
questions concerning this species or this finding to the above address.
FOR FURTHER INFORMATION CONTACT: Robert Pine, Supervisor (see ADDRESSES
section); 512-490-0057 extension 248.
SUPPLEMENTARY INFORMATION:
Background
Section 4(b)(3)(B) of the Endangered Species Act of 1973, as
amended (Act) (16 U.S.C. 1531 et seq.), requires that, for any petition
to revise the List of Threatened and Endangered Species containing
substantial scientific and commercial information indicating listing
may be warranted, we make a finding within 12 months of the date of
receipt of the petition. The finding must be that the petitioned action
is one of the following: (a) Not warranted, (b) warranted, or (c)
warranted but that the immediate proposal of a regulation implementing
the petitioned action is precluded by other pending proposals to
determine whether a species is threatened or endangered, and
expeditious progress is being made to add or remove qualified species
from the List of Endangered and Threatened Species. Section 4(b)(3)(C)
of the Act requires that a petition for which the requested action is
found to be warranted but precluded be treated as though resubmitted on
the date of such finding, that is, requiring a subsequent finding to be
made within 12 months. Such 12-month findings must be published in the
Federal Register.
On July 8, 2003, we received a petition requesting that we list a
karst meshweaver, Cicurina cueva (no common name), as an endangered
species with critical habitat. On May 25, 2004, Save Our Springs
Alliance (SOSA) filed a complaint against the Secretary of the Interior
and the Service for failure to make a 90-day petition finding under
section 4 of the Act for C. cueva. In our response to Plaintiff's
motion for summary judgment on October 15, 2004, we informed the court
that we believed that we could complete a 90-day finding by January 20,
2005, and if we determined that the 90-day finding provided substantial
information that listing may be warranted, we could make a 12-month
finding by December 8, 2005. On February 1, 2005 (70 FR 5123), we
published a 90-day finding and initiation of status review on a
petition to list C. cueva as an endangered species. On March 18, 2005,
the District Court for the Western District of Texas, Austin Division,
adopted our schedule and ordered the Service to issue a 12-month
finding on or before December 8, 2005.
Taxonomy
Gertsch (1992) described and named C. cueva, C. bandida, and C.
reyesi from adult, female specimens collected from Cave X in 1962 by
Bell and Woolsey, Bandit Cave in 1966 by Reddell and Fish, and Airman's
Cave in 1989 by Reddell and Reyes, respectively. The three Cicurina
species are all unpigmented and range in length from 5 millimeters (mm)
(0.19 inches (in)) to 5.6 mm (0.2 in). Gertsch (1992) distinguished
these three species by differences he perceived in the female
reproductive system.
Cicurina cueva, C. bandida and C. reyesi were described by Gertsch
(1992) on the basis of female genitalia of a small number of specimens.
Because there were some locations that only had records of immature
Cicurina that could not be identified to the species level, we
contracted Drs. Marshal Hedin and Pierre Paquin on September 24, 2004,
to determine whether species-level identification of immature specimens
of blind Cicurina spiders from southern Travis and northern Hays
counties could be made using a genetic assessment technique they had
previously applied to other species of Cicurina (see Paquin and Hedin
2004 for methods). Their report on the contracted study concludes that
C. cueva and two other formally described species, C. bandida and C.
reyesi (Gertsch 1992), likely represent variants of a single species
that shows genetic structuring across its range. They explain that
``This finding makes biological sense, as we would expect
geographically-adjacent cave populations to share more genetic
similarity than caves that are distant in space. The genetic
structuring observed is a natural consequence of the fragmented nature
of cave habitats, and the unique habitat limitations of these spiders *
* *'' (Paquin and Hedin 2005). The report authors suggest that rather
than three different species, the populations collected represent one
species, which they informally refer to as the ``C. cueva complex.''
They say ``We suggest that conservation activities concerning cave
populations in this confined geographic region be based on this single
species hypothesis.'' Since a formal revision reflecting this change in
taxonomy (the naming and classification of organisms) has not been
published in a peer-reviewed scientific journal, the Service requested
independent peer review of the report. We believe we should now make
this 12-month finding based on the taxonomic treatment recommended in
the contracted report (Paquin and Hedin 2005).
Drs. Paquin and Hedin submitted a report in May 2005, titled,
``Genetic and morphological analysis of species limits in Cicurina
spiders (Araneae, Dictynidae) from southern Travis and northern Hays
counties, with emphasis on Cicurina cueva Gertsch and relatives.'' When
Cicurina specimens from Travis, Hays, and Williamson
[[Page 75072]]
counties, Texas, were compared to sampled populations of C. cueva,
Paquin and Hedin (2005) found that the C. cueva complex (including all
three named species) forms a monophyletic group (defined as a group
descended from a single common ancestral form) or clade (a group of
organisms that share features derived from a common ancestor) within a
mitochondrial phylogeny (the evolutionary development and history of a
species or higher taxonomic group based on mitochondrial DNA).
Additionally, both C. bandida and C. reyesi are deeply embedded within
the mitochondrial DNA clade corresponding to the C. cueva complex,
indicating that they are part of the same group. In addition, they
examined female genital morphology and found that ``a similar genital
morphology, with slight variations, is shared across the entire
distribution of this species [the C. cueva complex].'' Based on the
Paquin and Hedin 2005 genetic and morphological results, they concluded
that these three named taxa represent variants of a single species.
Ultimately, when C. cueva, C. bandida, and C. reyesi are formally
combined as a single species, the authors propose all populations
within this expanded species be referred to as C. bandida, as this name
has page priority in Gertsch (1992). Paquin and Hedin (2005)
acknowledge that formal taxonomic decisions must involve publication in
a scientific journal; therefore, the authors suggest using ``C. cueva
complex'' to refer to the morphologically variable and genetically
divergent populations within this single species until the formal
change is published. In consideration of this information for use in
our 12-month finding, we conducted a scientific peer review of Paquin
and Hedin's 2005 report to determine if the proposed change in taxonomy
was likely to be accepted.
On May 6, 2005, we sent the report to 20 scientists, 19 with Ph.Ds,
with expertise in genetics, morphology, and/or conservation biology for
peer review. We asked that they particularly review the completeness of
the data in the report and identify any pertinent information that may
be missing and the soundness of the methodology, data analysis,
conclusions, and recommendations in the report. Each invited reviewer
was assigned a number, which will be referred to here. We received
eight responses (reviewers 2, 4, 5, 7, 8, 10, 13, 14). Dr. Mark
Kirkpatrick (co-petitioner) also submitted two letters to the Service
and personal email correspondence with Dr. Hedin (regarding the
report). Because Dr. Kirkpatrick is a co-petitioner he was not
considered a peer reviewer. However, the Service acknowledges his
considerable expertise in genetics. To allow peer reviewers the
opportunity to comment on the issues presented by Dr. Kirkpatrick, we
sent a second request for peer review to the same twenty scientists on
June 20, 2005, and received ten peer reviews (from reviewers 5, 7, 8,
9, 10, 12, 13, 14, 19, 20). We asked the peer reviewers for their
opinion on what degree of certainty they would assign to each of the
following hypotheses/conclusions: (1) C. cueva, C. bandida, and C.
reyesi are all one species (Paquin and Hedin conclusion), (2) they are
all separate species, or (3) another hypothesis/conclusion is possible.
We asked them to explain their views on appropriate criteria for
delimiting species using the types of morphological and genetic data
available in this case, and how those criteria apply to their review.
Of the 14 peer reviewers that responded to one or more requests for
reviews, 10 reviewers (2, 4, 5, 8, 10, 12, 13, 19, 20, and 22)
expressed general agreement with Paquin and Hedin's conclusion that C.
cueva, C. bandida, and C. reyesi represent a single species, one
reviewer (9) expressed support for continuing to recognize them as
three separate species, and three reviewers (7, 14, and 21) concluded
that more study was needed to distinguish between the one-species and
three species alternatives. In addition to these overall conclusions,
most reviewers provided additional comments on various aspects of the
Paquin and Hedin report, and on pertinent issues related to the
taxonomic interpretation of genetic and morphological data. These
comments on specific issues are summarized below.
Six of the twelve peer reviewers (2, 4, 5, 9, 10, 19) who responded
to at least one of these two requests for review indicated the study
overall was well done and the methods used in the genetic aspects of
this study were scientifically sound. However, we did receive a variety
of comments. Below we discuss the comments from both of these sets of
reviews in regard to the methods, analysis, and conclusions in the
study.
Concerns were raised by five peer reviewers (4, 5, 7, 9, 14)
regarding the authors' use of a single region of the mitochondrial DNA.
Some believed the report would be strengthened by a larger sample size
from each sampling locality, inclusion of data from other mitochondrial
DNA regions, and an analysis of genetic markers from nuclear DNA. Three
peer reviewers (4, 5, 14) speculated that the conclusion to group the
three taxa into a single species would probably still be the same even
with further genetic analysis.
Two reviewers (13, 14) questioned the use of particular
phylogenetic methods to analyze the genetic data and construct the tree
diagrams of relationships. The authors' present two different trees, or
phylogenies, based on a single data set; one generated by neighbor
joining (NJ) analyses and the other by Bayesian phylogenetics. These
methods differ in that NJ is a distance-based approach based on
analysis of a matrix of genetic distances (Hedrick 2000), and Bayesian
phylogenetics is a character-based approach (Avise 2004). Although they
rely on different assumptions and may give somewhat different results,
both are generally accepted methods for analyzing and presenting DNA
sequence data (Avise 2004), and Avise (2004, page 142) recommends that
studies include both a distance-based approach and a character-based
approach for comparison. The authors stated that they also analyzed the
data using maximum likelihood analysis, which is another character-
based method (Avise 2004). They did not present a phylogenetic tree
representing the results of the maximum likelihood analysis but stated
that the results were similar to their Bayesian analysis (Dr. Paquin,
San Diego State University, pers. comm., 2005; Hedin and Paquin 2005).
Although we acknowledge that there are a number of additional methods
of phylogenetic analysis (Hedrick 2000, Avise 2004), the authors
presented trees representing the two major types of trees, as
recommended by Avise (2004).
Three peer reviewers (8, 13, 14) suggested different conclusions
could be drawn, even if the phylogenies are accepted. These alternative
interpretations reflect differing views on the appropriate amount of
genetic difference for delineating species boundaries, which is an
active area of debate in taxonomy (Sites and Marshall 2004).
One peer reviewer (14) suggested that the study of additional
morphological characters, rather than genitalia, such as somatic (non-
sexual) characters, might find diagnosable differences within the ``C.
cueva complex.'' However this peer reviewer doubted that the outcome of
such studies would likely affect the authors' conclusion that C. cueva
is not a species. Additionally, one reviewer (14) stated the assessment
of genitalic variation was subjective and would have been better if the
different genitalic parameters could have been quantified somehow with
the variation analyzed
[[Page 75073]]
statistically. Reviewers 7 and 12 stated that morphology clearly plays
a critical role in deciphering the systematics of this group, and
reviewer 7 wondered if some statistical quantification of patterns in
morphological characters is possible. Gertsch's (1992) original
diagnoses for these three species included only collection locality and
characters of the female reproductive system; no other characters were
identified in the diagnosis. The diagnosis that accompanies the
original description of a new species is important because it provides
the characters or character states that allow that species to be
distinguished from other species. Gertsch (1992) expressed doubts that
other characters were useful; for example, ``Cicurella [the subgenus to
which the species in question belong] * * * offer few coloration or
somatic features to allow easy identification.'' Gertsch (1992) was
also dismissive of the value of different reproductive features in
males and notes that males are much less available for study, as they
represent only a fifth the number of mature females.
One reviewer (22) noted variation in female genitalia observed
among the specimens presented in the report was considered ``well
within'' the range of intraspecific (within-species) variation
typically observed in female genitalia of other species and adequately
demonstrates that there is no morphological reason to consider C.
cueva, C. bandida, and C. reyesi as three separate species. We
recognize that study of additional morphological characters and more
quantitative analysis of current characters could increase our
understanding of morphological variation within this group of spiders,
but we find little support for rejecting the authors' recommended
taxonomy, considering their findings and the peer reviewers' comments
on the morphological data.
Dr. Kirkpatrick thought the Paquin and Hedin (2005) report did not
statistically disprove the ``established taxonomy'' previously
described by Gertsch (1992). However, two peer reviewers (8 and 22)
expressed concern that Gertsch (1992) did not sufficiently account for
the possibility of intraspecific variation in genitalic characters and
improperly recognized minor morphological variants as different species
and that his species descriptions were based on small sample sizes.
While such a lack of statistical analysis is common in the field of
systematic biology, we believe that since two experts (19 and 22) in
this field have expressed strong doubts about the basis of the species-
level taxonomy presented by Gertsch, the alternative taxonomic
delineation presented by Paquin and Hedin (2005) deserves serious
consideration. We also note that Paquin and Hedin's (2005)
morphological studies were based on more than double the number of
specimens available to Gertsch (1992) when he originally described the
species.
We received a variety of responses to the specific question in the
second peer review regarding the degree of certainty that the reviewer
would assign to the various hypotheses or possible conclusions about
species limits. Two reviewers (8 and 19) clearly supported the Paquin
and Hedin conclusion that C. cueva, C. bandida, and C. reyesi are all
one species. However, reviewer 8 did disagree about the assignment of
three or four of the populations to this group and did differ with
Paquin and Hedin about the level of differences accepted to represent a
species. One of the reviewers (13) was ``unconvinced that the report's
conclusions are correct'', and suggested an alternate hypothesis and
classification. Reviewers 7 and 9 believe the Paquin and Hedin
conclusions should be considered preliminary and premature,
respectively. Reviewers 5, 10, 12, and 20 tended to accept the Paquin
and Hedin hypothesis based on the information presented; however, they
each expressed some uncertainty or suggested that additional data
collection and analysis would be advisable. Reviewer 14 felt that both
Hedin and Kirkpatrick provided ``solid, convincing arguments for their
points of view'; this reviewer doubted that further investigation would
lead to improved resolution on the question of how many species there
are and believes this is ultimately a matter of interpretation.
In response to divergent opinions regarding how to define species
limits and how much data are needed to confidently make a species
determination, and because some but not all peer reviewers were
familiar with spider taxonomy in particular, we conducted a third peer
review. We sent four arachnologists the Paquin and Hedin 2004
publication (that described the methods used in this study) and 2005
report, the first peer review request and responses, Dr. Kirkpatrick's
letters and emails, and the second peer review request and responses.
We received two responses (reviewers 21 and 22). One of these reviewers
(22) stated that ``Based on the evidence presented by Hedin & Paquin,
the only well supported scientific conclusion at this time, is that
only one species is present.'' The other reviewer (21) stated Paquin
and Hedin clearly explained their methods and that they are adequate
for their questions. The reviewer also stated that ``Paquin and Hedin
have given a conservative conclusion based on their data, and have
noted alternative explanations and the need for more specimens''.The
reviewer stated that ``without more of this work I do not see a way to
resolve the concerns about data interpretation raised by Dr. Mark
Kirkpatrick.''
There is ongoing debate among many scientists regarding methods for
species differentiation (Sites and Marshall 2004). Some believe
defining species boundaries requires a ``total evidence'' approach that
includes data from multiple genes and morphology, as well as ecology
and behavior. Although it is reasonable to believe this debate will
continue, the Service's ``Interagency Cooperative Policy on Information
Standards under the Endangered Species Act'' (59 FR 34271) requires we
use the ``best available comprehensive technical information'' in
making Federal listing determinations. The Paquin and Hedin (2005)
report provides genetic data for the first time and morphological data
based on an increased number of specimens; both approaches fail to
distinguish C. cueva from C. bandida and C. reyesi. In addition, the
claim by the petitioners that the genetic analysis employed is not
informative about taxonomic standing within the C. cueva complex is not
supported by the clear correspondence between geography and branching
patterns of both phylogenetic trees. The correspondence between
geography and phylogeny indicates that the phylogenetic patterns have a
biological basis and do not simply present ``noise'' that is obscuring
biologically important patterns. We believe, based on our review and
the results of the peer reviews, the Paquin and Hedin (2005) report
provides the best available information on the current taxonomic status
of the Cicurina complex. Although it is always possible that future
analyses on other morphological characters or genetic markers may
convince spider taxonomists that another taxonomic interpretation is
appropriate, we cannot base our findings on the speculative outcomes of
studies not yet performed. We find, however, that the Paquin and Hedin
(2005) report is based on procedures and methods of analysis that are
generally accepted in the application of molecular methods to taxonomy.
Although additional study could affect the taxonomic conclusions of the
report, according to the requirements of the Act the best available
genetic and
[[Page 75074]]
morphological data at this time support the recommendation of Paquin
and Hedin (2005) to treat these three species as one species.
Previous Federal Actions
Previous Federal actions can be found in our 90-day finding that
published on February 1, 2005 (70 FR 5123), and in our notice reopening
the comment period on August 16, 2005 (70 FR 48093). That information
is incorporated by reference into this 12-month finding.
In addition to information incorporated by reference we note that
the first comment period for providing information for our status
review closed May 15, 2005. Pursuant to 50 CFR 424.16(c)(2), we may
extend or reopen a comment period upon finding that there is good cause
to do so. We reopened the comment period from May 23 to June 22, 2005
(70 FR 29471; May 23, 2005), since additional information from the
genetic analysis of Cicurina species in southern Travis County was
completed. Several parties requested another extension of the comment
period. We reopened the public comment period from August 16 to 30,
2005 (70 FR 48093; August 16, 2005). During this final comment period,
we made available the results of our peer review on the Paquin and
Hedin (2005) report.
Finding
We have carefully assessed the best scientific and commercial
information available regarding the taxonomic status of Cicurina cueva.
We reviewed the petition, available published and unpublished
scientific and commercial information, and information submitted to us
during the public comment periods on our status review following our
90-day finding. This finding reflects and incorporates information we
received during the public comment periods. We also consulted with
recognized spider and karst invertebrate experts. On the basis of this
review, we find that listing C. cueva is not warranted because C. cueva
does not meet the definition of a ``species'' under the Act.
References Cited
A complete list of all references cited herein is available upon
request from the Field Supervisor at the Austin Ecological Services
Office (see ADDRESSES section).
Author
The primary author of this document is the Austin Ecological
Services Office (see ADDRESSES section).
Authority: The authority for this action is the Endangered
Species Act of 1973, as amended (16 U.S.C. 1531 et seq.).
Dated: December 8, 2005.
Marshall P. Jones Jr.,
Acting Director, Fish and Wildlife Service.
[FR Doc. 05-24119 Filed 12-16-05; 8:45 am]
BILLING CODE 4310-55-P