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Gastropoda: Hydrobiidae) of Ash Meadows, Amargosa Basin, California-Nevada

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Gastropoda: Hydrobiidae) of Ash Meadows, Amargosa Basin, California-Nevada 31 December 1987 PROC. BIOL. SOC. WASH. 100(4), 1987, pp. 776-843 SPRINGSNAILS (GASTROPODA: HYDROBIIDAE) OF ASH MEADOWS, AMARGOSA BASIN, CALIFORNIA-NEVADA Robert Hershler and Donald W. Sada Abstract. —Aquatic snails of the family Hydrobiidae were sampled from nu- merous springs in Ash Meadows, California-Nevada, during 1985-1986. The fauna of this lush oasis is represented by at least three lineages and composed of 11 species in two genera, Pyrgulopsis Call and Pilsbry, 1886 and Tryonia Stimpson, 1865. Nine species are described herein as new. Nine species are locally endemic (three are restricted to single springs), while the remaining two are restricted to Amargosa River drainage. Pyrgulopsis species are well-differ- entiated in shell and anatomical features (mostly penial morphology), whereas Tryonia species show marked variation only in the former. Stepwise discriminant analyses were done using shell morphometric data from three separate species groups in Ash Meadows. Separate analyses were done using standard measurements and Raupian parameters as a local test of their effectiveness in discriminating between closely related forms. Classifica- tion was uniformly high (86-93%) when the former data set was used, sup- porting taxonomy presented herein. Raupian parameters produced less suc- cessful classifications (48-71%), probably due to absence of shape diversity among similar-shelled members of species groups considered. Ash Meadows springsnails parallel local fishes in having affinities with taxa from the Death Valley System and Colorado River drainages. Distributional evidence suggests that local differentiation of snails has primarily occurred within narrow ranges of altitude, in contrast to patterns documented for local fishes. Gill-breathing springsnails (Gastropoda: the region's well studied ichthyofauna (see Hydrobiidae) inhabiting the series of inter- Miller 1948, Soltz and Naiman 1978, montane valleys that constitute the Death Minckley et al. 1986). Valley System (Miller 1943) of southeastern An ongoing survey of the region's Hy- California and southwestern Nevada are drobiidae was initiated during 1985 by the poorly known, as they are throughout the senior author to obtain material for system- arid Southwest. While only two species have atic study of the fauna. Fieldwork during been described from this region, based on 1985-1986 included survey of all known material collected by the United States De- springs in Ash Meadows, a lush oasis re- payment of Agriculture 1891 Death Valley nowned for its highly endemic biota (Beat- Expedition, unpublished data (see Taylor ley 1977, Soltz and Naiman 1978, Reveal 1966, Landye 1973, Hershler 1985, Taylor 1979). We present herein a description of in Williams et al. 1985) suggest that many this faunule as the first of a series of papers additional species are present in the region, on systematics of springsnails of the Death with diversity and localization of endemic Valley System, taxa likely surpassing that documented for Eleven springsnail species are recognized, VOLUME 100, NUMBER 4 777 Fig. 1. Map showing location of Ash Meadows, Nevada-California. representing two genera, Pyrgulopsis Call springsnail fauna of similarly sized regions and Pilsbry, 1886, and Tryonia Stimpson, (Taylor 1966). Three lineages are recog- 1865. Nine species are new, and nine are nized in the fauna, which has affinities with endemic to Ash Meadows, three of which springsnail s from Death Valley System and are restricted to single springs. Non-endem- Colorado River drainages. Shell morpho- ic forms are restricted to Amargosa River metric data were gathered from selected drainage in the eastern part of the Death populations and subjected to stepwise dis- Valley System. Both total and endemic di- criminant function analyses, with standard versity are striking, clearly exceeding values measurements and Raupian parameters for local fishes, and seldom matched by (Raup 1966) used in separate analyses as a 778 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON local test of relative effectiveness in discrim- arid terrain, or continuously or seasonally inating between closely related taxa. connected by confluence of outflows. Large The taxonomic work presented herein is springs are thermal (28-32°C). Typically that of Hershler. Other parts of the paper lower temperatures of smaller springs may were co-written. be due to rapid heat loss associated with small discharge. Springs along the eastern part of Ash Meadows are generally warmer Environmental Setting than those to the west. All spring water is Ash Meadows occupies ca. 25,000 ha in potable. Total dissolved solids range be- Amargosa Desert along the California-Ne- tween 410 and 870 mg/liter (Dudley and vada border 60 km W of Death Valley (Fig. Larson 1976), and specific conductivity 1). Local terrain slopes from neighboring ranges from 550-800 micromhos/cm (Win- hills of 1300 m elevation southwest to el- ograd and Thordarson 1975). Sodium is the evation of 560 m in Amargosa Valley. Mean dominant cation, with concentrations rang- annual temperature is 18.5°C (Dudley and ing from 0.18-0.23 meq/liter; followed by Larson 1976), with summer highs often ex- magnesium and calcium. Bicarbonate ion is ceeding 40°C. Local rainfall is scant, aver- the most abundant anion, with typical con- aging ca. 7.0 cm annually (Dudley and Lar- centration of 4.9 meq/liter (Dudley and Lar- son 1976). son 1976, Garside and Schilling 1979). Lo- Shadscale (Atriplex confertifolia) and cal spring discharge is old water (ca. 10,000 Haplopapus acaradenius dominate on xeric years; Winograd and Thordarson 1975) soils in Ash Meadows, and salt grass (Dis- transported to the area by deep carbonate tichlis spicata) and rushes (Juncus balticus aquifers draining about 7200 km2 of south- and /. nodosus) are spread over mesic areas ern Nevada (Winograd and Thordarson where moisture is maintained by spring dis- 1975). While currently endorheic, the area charge or seasonal precipitation. Ash trees has had past connections with nearby (Fraxinus velutina var. coriaced), mesquite Amargosa River, located a few km to the (Prosopis julifera and P. pubescens), wild west (albeit intermittent at this point), which grape (Vitus arizonica), and salt grass are continues to the south before turning back predominant riparian flora (Beatley 1971). north to terminate in Death Valley. The name "Ash Meadows" refers to local Ash Meadows is among the most signif- abundance of ash trees (Carlson 1974). icant endangered aquatic ecosystems in Approximately 50 springs are scattered western North America (Williams et al. throughout Ash Meadows, many emerging 1985). Including springsnails described from old lake beds along a fault-controlled herein, a total of 22 species-group aquatic spring line in northern and eastern parts of or riparian taxa are considered local endem- the area. Springs (Figs. 2-4) vary in size ics, a total unmatched by any similar-sized from the large limnocrene, Crystal Pool, ca. area in the United States. Approximately 20 m in diameter and discharging 189 liters/ 4000 ha. of the area (16% of total), including sec, to seeps less than 1.0 m across and dis- most springs, has been perturbed by mining, charging only a few cc/sec (Dudley and Lar- agriculture, and a municipal development. son 1976, Garside and Schilling 1979). To- Establishment of a number of exotic species tal annual spring discharge is estimated as has further altered the ecosystem, with in- 671 liters/sec (Winograd and Thordarscn vading biota including bullfrogs (Rana 1975). Individual spring outflows extend catesbeiana), mosquito fish (Gambusia af- 0.1-10 km before disappearing into soil. finis), sailfin mollies (Poecilia latipinna), red- Springs are either isolated by expanses of rim melania snail (Melanoides tuberculata), VOLUME 100, NUMBER 4 779 Fig. 2. Photographs of Ash Meadows springs: a, Big Spring (11/8/85); b, Crystal Pool (11/8/85). crayfish {Procambarus clarki), and salt ce- dates for future listing (USDI 1985a, b). dar (Tamarisk sp.). Perturbation has caused There is, however, cause for optimism as population decline of virtually all endemic The Nature Conservancy purchased the area taxa, 12 of which are federally listed as in 1984 and sold it to U.S. Fish and Wildlife threatened or endangered (USDI 1986). All Service, which established the Ash Mead- other endemic species, including most ows National Wildlife Refuge (Sada and springsnails recognized herein, are candi- Mozejko 1984). 780 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Photographs of Ash Meadows springs: a, Outflow of North Indian Spring (11/10/85); b, Devils Hole (11/10/85). List of Recognized Taxa by either washing rocks or sifting soft sed- iments with a fine-mesh hand sieve. Water Pyrgulopsis micrococcus (Pilsbry). temperature and conductivity were mea- P. erythropoma (Pilsbry, in Stearns 1893). sured with a YSI Model 33, S-C-T meter. P. fairbanksensis Hershler and Sada, new Dissolved oxygen concentrations were de- species. P. crystalis Hershler and Sada, new species. termined using a YSI 97 oxygen meter. De- tailed descriptions of all localities visited in P. isolatus Hershler and Sada, new species. Ash Meadows are in Appendix 1. A few P. nanus Hershler and Sada, new species. P. pisteri Hershler and Sada, new species. additional snailless sites were visited, in- cluding seeps by Crystal Pool. Collections Tryonia angulata Hershler and Sada, new species. often included shells of locally extinct springsnails that will be discussed further in T. variegata Hershler and Sada, new species. a future publication. Distributions of species T. ericae Hershler and Sada, new species. in Ash Meadows are shown in Figs. 16, 25, T. elata Hershler and Sada, new species. and 44. Snails were relaxed with menthol crystals, Materials and Methods fixed in 4% buffered formalin and preserved Material examined. —Most localities vis- in 70% ethanol. This material is housed in ited, including all sites having snails, are the National Museum of Natural History, shown in Figs. 5 and 6. Snails were collected Smithsonian Institution (USNM) collec- VOLUME 100, NUMBER 4 781 Fig.
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