[Federal Register: December 19, 2005 (Volume 70, Number 242)]
[Rules and Regulations]               
[Page 75071-75074]
From the Federal Register Online via GPO Access [wais.access.gpo.gov]



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 

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 

DATES: The finding announced in this document was made on December 19, 

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.

section); 512-490-0057 extension 248.



    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.


    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 
    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 
    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 
    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.


    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).


    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]