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Federal Register / Vol. 86, No. 119 / Thursday, June 24, 2021 / Rules and Regulations
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translocating 340 individuals from the Vaseys Paradise population. Since 2005, this population has been considered self-sustaining with an estimated population of approximately 700 individuals Sorensen 2005, p. 9.
Between 2009 and 2015, timed count surveys of the translocated population at Upper Elves Chasm were conducted by Arizona Game and Fish Department, National Park Service biologists, and volunteers. Surveys over this timeframe documented a small but relatively stable Kanab ambersnail population at the site, with CPUE estimates between 0.85 to 4.15 snails per 10 minutes searched Sorensen 2015, p. 12.
Taxonomy Kanab ambersnails were first collected in 1909, by James Ferriss from an area called The Greens, a vegetated seep approximately 6 mi 10 km north of Kanab in Kanab Creek Canyon, Utah 57 FR 13657, April 17, 1992; Service 1995, p. 2. However, ambersnails have not been found at the type locality since 1991 Meretsky et al. 2002, p. 314;
Culver et al. 2013, p. 6.
The snails collected by James Ferriss in 1909 were initially placed in the species Succinea hawkinisi, but Pilsbry 1948, p. 797 placed them in Oxyloma and created the subspecies kanabensis under the species haydeni 57 FR 13657, April 17, 1992. The subspecies kanabensis classification was considered to be temporary at the time, and the author recommended that the taxonomic status be reconsidered in the future Pilsbry 1948, p. 798; Clarke 1991, p. 23; 57 FR 13657, April 17, 1992.
We have assessed all available genetic information for the Kanab ambersnail Miller et al. 2000, entire; Stevens et al.
2000, entire; Culver et al. 2013, entire.
Since the listing of Kanab ambersnail in 1992 57 FR 13657; April 17, 1992 and the publication of the Kanab ambersnail recovery plan in 1995 Service 1995, entire, several studies on subspecies distribution, morphological characteristics, and genetic relationships to other Oxyloma species have been completed. We briefly describe these studies below. At this time, these studies represent the best scientific information available in order for us to analyze the Kanab ambersnails distribution and taxonomic changes.
Various analyses can be done to determine genetic structure of a species, including analyses of: 1 Mitochondrial DNA, which is rapidly evolving and useful to determine recent populations;
2 nuclear microsatellite DNA, which has high amounts of genetic variation and can be used to look at populations
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within a species; 3 nuclear DNA, which is inherited equally from both parents unlike mitochondrial DNA, which is inherited maternally; and 4
amplified fragment length polymorphisms AFLP, which are used to sample multiple loci across the genome.
Miller et al. 2000 used AFLP to determine intraand inter-population genetic information for four Oxyloma species in Utah and Arizona. Among these, two Niobrara ambersnail Oxyloma haydeni haydeni locations were studied at Indian Gardens Arizona and Minus Nine Mile Spring Arizona, and two Kanab ambersnail populations were studied at Three Lakes Utah and Vaseys Paradise Arizona Miller et al. 2000, pp. 18451946.
From this study, the ambersnail population at Three Lakes appears more closely related to the Niobrara ambersnail population at Indian Gardens than to the ambersnail population at Vaseys Paradise Miller et al. 2000, p. 1852. Upper Elves Canyon was not included in this study.
Stevens et al. 2000 used mitochondrial DNA and morphological analysis to distinguish Succineidae Oxyloma, Catinella, and Succinea populations in the United States and Canada. The authors collected over 450
samples from seven U.S. States and Canadian provinces, including from 63
different populations or locations of snails Stevens et al. 2000, p. 4.
Determining Oxyloma species based on morphology was shown to be inaccurate Stevens et al. 2000, pp. 45, 42.
Vaseys Paradise did not cluster with the Three Lakes ambersnail population or the two sampled Niobrara ambersnail populations, leading the authors to suggest Vaseys Paradise might represent a unique species Stevens et al. 2000, p. 41. However, a later, more comprehensive study found that Vaseys Paradise clustered closely enough with samples from other surrounding Oxyloma populations for them all to be considered part of the same Oxyloma species Culver et al. 2013, p. 57.
In this most recent and detailed peerreviewed study, ambersnails were collected from 12 locations in Arizona and Utah, with each location providing at least 14 ambersnail specimens Culver et al. 2013, p. 5. Samples consisted of Kanab ambersnail, Niobrara ambersnail, blunt ambersnail Oxyloma retusum, undescribed species of Oxyloma, and individuals from Catinella used to provide an outgroup comparison Culver et al. 2013, p. 6.
This study included samples from all three extant populations identified as Kanab ambersnail. Between the
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Oxyloma populations, shell morphology did not have the variation usually associated with different species, leading the authors to state that none of the 12 populations sampled was reproductively isolated from the others Culver et al. 2013, p. 52. This information supports the finding that the three populations identified as Kanab ambersnail do not alone comprise a discrete taxon.
Genetic results indicated that there was gene flow among all the populations sampled, most likely due to shortor long-distance dispersals from other populations Culver et al. 2013, p.
57. Additionally, Kanab ambersnail samples from Vaseys Paradise did not cluster with the other two Kanab ambersnail populations Culver et al.
2013, pp. 51, 55. The authors concluded that the three populations of Kanab ambersnail are not a valid subspecies of Oxyloma haydeni and should instead be considered part of the same taxa as the ambersnails from the eight other populations of Oxyloma in Utah and Arizona that were sampled for comparison Culver et al. 2013, entire.
This study declined to positively identify a species-level taxon for these 11 populations of ambersnail, due to lack of genetic information on the genus Culver et al. 2013. The primary author stated later that her expert opinion was they should all, including those previously identified as Kanab ambersnail, be considered Niobrara ambersnail Oxyloma hadenyi Culver 2016, pers. comm.. The authors stated that specimens from the type locality of the Niobrara ambersnail in Nebraska could be examined for comparison to verify this conclusion Franzen 1964, p.
73; Culver et al. 2013, p. 57; Culver 2016, pers. comm., but to date, no such analysis has been done.
The above-described Culver et al.
2013 study was released as a United States Geological Survey USGS
Scientific Investigations Report, and the review approach was similar to that of manuscripts published by scientific journals. The report was initially reviewed by five reviewers and required subsequent revision. The report received an additional review following revision due to the complex subject matter. The response to reviewer comments and subsequent revised manuscript were reviewed by another independent geneticist to ensure that the author adequately addressed issues and comments brought up by reviewers Sorensen 2014, pers. comm.. The subsequent revision that occurred after 2011 resulted in more genetic information added to the final 2013
manuscript, which further substantiated
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