Research Article |
Corresponding author: Robert Hershler ( hershlerr@si.edu ) Academic editor: Thierry Backeljau
© 2015 Robert Hershler, Hsiu-Ping Liu, Jeffrey S. Simpson.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Hershler R, Liu H-P, Simpson JS (2015) Assembly of a micro-hotspot of caenogastropod endemism in the southern Nevada desert, with a description of a new species of Tryonia (Truncatelloidea, Cochliopidae). ZooKeys 492: 107-122. https://doi.org/10.3897/zookeys.492.9246
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Newly obtained and previously published sequences of the cytochrome c oxidase subunit I (COI) gene were analyzed to examine the biogeographic assembly of the caenogastropod fauna (belonging to the families Assimineidae, Cochliopidae, and Hydrobiidae) of an isolated spring along the lower Colorado River in southern Nevada (Blue Point Spring). Based on available COI clock calibrations, the three lineages that comprise this fauna are 2.78–1.42 million years old, which is roughly coeval or slightly younger than the age of Blue Point Spring (inferred from local fossil spring deposits). Two of the lineages—endemic Pyrgulopsis coloradensis and Assiminea aff. infima—are most closely related to snails in the Death Valley area (well to the west) and likely colonized Blue Point Spring by transport on birds. A single haplotype was detected in both of these snails, suggesting that they may have only recently colonized Blue Point Spring. The third lineage—endemic Tryonia infernalis, newly described herein based on morphological and molecular evidence—is most closely related to a geographically proximal species in a lower Colorado River tributary (T. clathrata); the split between these taxa may be the product of vicariance (severance of a prior drainage connection) or a separate jump dispersal event. The considerable genetic diversity in T. infernalis (three haplotypes differing by 0.6% mean sequence divergence) suggests a possibly lengthy history of local differentiation. Our findings also identify Blue Point Spring as a new micro-hotspot of groundwater-dependent biodiversity in Nevada and will assist ongoing efforts to protect and conserve these imperiled ecosystems.
Gastropoda, Assimineidae, Hydrobiidae, western United States, aquatic snails, biogeography, taxonomy, conservation
The desert region of southeastern California and southwestern Nevada, encompassing portions of the Great Basin and lower Colorado River watershed, contains distinctive assemblages of tiny caenogastropods—belonging to the families Assimineidae (genus Assiminea), Cochliopidae (Tryonia) and Hydrobiidae (Pyrgulopsis)—that have been a recent focus of biogeographic study using mtDNA sequence data (e.g.,
The A. infima complex is subdivided into a clade that is distributed in the Death Valley region (this lineage also contains a population from the head of the Gulf of California) and a genetically divergent population (A. aff. infima Berry) in Blue Point Spring (
Map showing the location of Blue Point Spring relative to other geographic areas discussed in the text. The collecting localities for specimens of Pyrgulopsis sanchezi and Tryonia clathrata (sister taxa of Blue Point Spring endemics) used in the molecular phylogenetic analyses are also shown.
Fresh material was collected from Blue Point Spring by RH in May, 2014, and preserved in 90% ethanol for genetic analysis; a portion of the T. porrecta sample was relaxed with menthol crystals, fixed in dilute (4%) formalin, and preserved in 70% ethanol for anatomical study. Genomic DNA was extracted from entire snails (A. aff. infima, six specimens; P. coloradensis, four specimens; T. porrecta, six specimens) using a CTAB protocol (
MrModeltest 2.3 (
Genetic distances within and between samples were calculated using MEGA6 (
Large, adult females were used for shell measurements. The total number of shell whorls (WH) was counted for each specimen; and the height and width of the entire shell (SH, SW), body whorl (HBW, WBW), and aperture (AH, AW) were measured from camera lucida outline drawings using a digitizing pad (see
The phylogenetic analyses congruently depicted a sister relationship between P. coloradensis and P. sanchezi Hershler, Liu & Bradford, which is distributed in the Death Valley area (Fig.
The phylogenetic analyses of the Tryonia dataset congruently delineated a well-supported sister relationship between Blue Point Spring population and T. clathrata, which is also distributed in the lower Colorado River basin (Fig.
The eight sequenced specimens of A. aff. infima shared the same haplotype which differed from sequences of the other members of the A. infima complex by 2.6 +/- 0.5%.
Tajima’s relative rate test did not reject clocklike behavior for the datasets of interest. The posterior Bayes factor also strongly favored the molecular clock model, indicating that the application of a molecular clock is appropriate for these data.
Undescribed [Fontelicella and] Tryonia species.—
Tryonia porrecta.—
USNM 883884 (a dry shell), Blue Point Spring, just below source, Clark County, Nevada, 36.3894°N, 114.4329°W, 24 July 1988, R. Hershler. Paratypes (ca. 200 dry shell and alcohol preserved specimens), USNM 1266143 (from same lot).
NEVADA. Clark County: USNM 883248 (coll. James J. Landye, 17-XII-1992), USNM 1098627 (coll. Donald W. Sada, 6-XII-2006), USNM 1146345 (coll. Andrew K. Schwaneflugel, 29-V-2008), USNM 1146420 (coll. DWS, 11-XII-2009), USNM 1248362 (coll. RH, 5–15–2014), USNM 854844 (coll. Saxon Sharpe, no date), Blue Point Spring.
Shell medium-sized, conic to turriform; penis having two distal papillae on the inner edge and a single basal papilla both on the inner and outer edges. Readily distinguished from geographically proximal and closely related T. clathrata by its smaller size, weaker shell sculpture, and smaller number of papillae on the inner edge of the penis. Differentiated from T. gilae (also distributed in the lower Colorado River basin) by its more convex teleoconch whorls, lateral expansion of distal bulb of penis, and in having a basal papilla on the inner edge of the penis. Differs from T. porrecta, with which it was previously confused, by its smaller size, consistently weak shell sculpture, and much greater frequency of males.
Shell (Fig.
Shells, opercula and radula, T. infernalis sp. n. A Holotype, USNM 883884 B Male shell, USNM 1266143 C, D Opercula (outer, inner sides), USNM 1266143 E Portion of radular ribbon, USNM 1266143 F Central teeth, USNM 1266143 G Lateral and inner marginal teeth, USNM 1266143. Scale bars A–B: 1.0 mm; C, D: 200 µm; E–G: 10 µm.
WH | SH | SW | HBW | WBW | AH | AW | SW/SH | HBW/SH | AH/SH | |
---|---|---|---|---|---|---|---|---|---|---|
Holotype, USNM 883884 | ||||||||||
5.75 | 3.09 | 1.78 | 1.87 | 1.56 | 1.13 | 1.00 | 0.58 | 0.604 | 0.36 | |
Paratypes, USNM 1266143 (n = 9) | ||||||||||
Mean | 5.33 | 2.61 | 1.41 | 1.60 | 1.25 | 0.95 | 0.80 | 0.54 | 0.61 | 0.37 |
S.D. | 0.28 | 0.15 | 0.06 | 0.07 | 0.06 | 0.04 | 0.04 | 0.03 | 0.03 | 0.02 |
Range | 5.00–5.75 | 2.41–2.82 | 1.33–1.54 | 1.46–1.71 | 1.13–1.34 | 0.91–1.02 | 0.75–0.86 | 0.49–0.59 | 0.58–0.65 | 0.34–0.39 |
Inner and outer sides of operculum smooth (Fig.
Animal darkly pigmented. Penis (Fig.
The specific epithet (infernalis) is a Latin adjective meaning hellish, and refers to the Valley of Fire, which is closely proximal to the type locality.
Tryonia infernalis is known only from its type locality, a thermal (ca. 30 °C) rheocrene (discharging ca. 0.55 l/s;
The reproductive anatomy of several females was studied to confirm that this species belongs to Tryonia as currently defined (
Small assemblages of locally endemic spring-dwelling invertebrates are scattered throughout arid western North America (
The use of a molecular clock to estimate divergence times is wrought with difficulties and is further complicated in this case by the absence of locally derived calibrations for Assiminea and Tryonia. Nevertheless, roughly calculated values provide useful insight into the biogeographic history of the Blue Point Spring fauna (note that we performed Bayes factor and Tajima’s rate tests, both of which suggested that the assumption of a molecular clock is valid). Based on mtCOI clock calibrations of 1.83% per million years. for European Hydrobiidae (
COI sequence divergence and estimated ages of Blue Point Spring snail lineages based on two clock calibrations.
Lineage | Per cent sequence divergence (sister taxon) | Estimated age (m.y.) | |
---|---|---|---|
1.83%/m.y. calibration |
1.62%/m.y. calibration |
||
A. aff. infima | 2.6 (other members of A. infima complex) | 1.42 | 1.60 |
P. coloradensis | 4.5 (P. sanchezi) | 2.46 | 2.78 |
T. infernalis | 3.9 (T. clathrata) | 2.13 | 2.41 |
The recognition of T. infernalis as a distinct, endemic species further highlights Blue Point Spring as a micro-hotspot of locally endemic aquatic biodiversity in Nevada. The Blue Point Spring Assiminea is probably a distinct species as well, but a formal taxonomic treatment is deferred pending completion of an ongoing revision of the A. infima complex (Hershler and Liu in preparation). Although this tiny aquatic ecosystem is on lands administered by the National Park Service (Lake Mead National Recreation Area), there may be a need for additional protection and conservation measures. The spring is located alongside a paved highway and public access is further facilitated by a small parking area near the lower end of the spring run. There is no fencing around the spring (or its run) and thus it is vulnerable to disturbance from foot traffic and other recreational activities. [We note in this context that Blue Point Spring harbors one of the few remaining populations of the relict leopard frog (Rana onca Cope), which requires open habitat maintained by ungulate grazing and thus may be negatively impacted by fencing (
This project was supported in part by funding from the National Park Service Mojave Desert Network Inventory and Monitoring Program (award # P13PG00412) that was facilitated by Geoffrey Moret. Sampling of Blue Point Spring snails in 2014 was made under the auspices of a scientific collecting permit from the National Park Service (Permit# LAKE-2014-SCI-0005). Kyle Simpson assisted with the fieldwork. The first author thanks Sue Beard for a useful discussion of the possible age of Blue Point Spring. Yolanda Villacampa measured shells and prepared scanning electron micrographs, and Freya Goetz prepared a digital version of the anatomical drawing. Ross Haley, Geoffrey Moret and Mark Sappington provided helpful comments on an early draft of this manuscript, which was also improved by input from two anonymous reviewers.