Springsnails (Gastropoda, Hydrobiidae) of Owens and Amargosa River California-Nevada
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Springsnails (Gastropoda, Hydrobiidae) Of Owens And Amargosa River California-Nevada R Hershler Proceedings of The Biological Society of Washington 102:176-248 (1989) http://biostor.org/reference/74590 Page images from the Biodiversity Heritage Library, http://www.biodiversitylibrary.org/, made available under a Creative Commons Attribution-Noncommercial License http://creativecommons.org/licenses/by-nc/2.5/ PROC. BtOL SOC. WASH. 102(t). t989, pp. 176-248 SPRINGSNAILS (GASTROPODA: HYDROBIIDAE) OF OWENS AND AMARGOSA RIVER (EXCLUSIVE OF ASH MEADOWS) DRAINAGES, DEATH VALLEY SYSTEM, CALIFORNIA-NEVADA Robert Hershler Abstract. - Thirteen springsnail species (9 new) belonging to Pyrgulopsis Call & Pilsbry, 1886, and Tryonia Stimpson, 1865 aTe recorded from the region encompassing pluvial Owens and Amargosa River (exclusive of Ash Meadows) drainages in southeastern California and southwestern Nevada. Discriminant analyses utilizing shell morphometric data confirmed distinctiveness of the nine new species described herein, as: Pyrgulopsis aardahli, P. amargosae, P. olVenensis, P. perlubala, P. wongi, Tyronia margae, T. robusla, T. rowlandsi, and 1: salina. Of the 22 springsnails known from Death Valley System, 17 have very localized distributi ons, with endemic fauna concentrated in Owens Valley, Death Valley, and Ash Meadows. A preliminary analysis showed only partial correlation betwecn modern springs nail zoogeography and configuration of inter·connected Pleistocene lakes comprising the Death Valley System. This constitutes the second part of a sys List of Recognized Taxa tematic treatment of springsnails from the Pyrgu/opsis aardah/i, new species. Death Valley System, a large desert region P. amargosac. new species. in southeastern California and southwest P. micrococcus (Pilsbry. 1893). ern Nevada integrated by a se ries of la kes P. olVcnsensis, new species. during Pleistocene times. An earl ier paper P. perlurbata, new species. (Hershler & Sada 1987) dealt with the Ash P. ef. stearnsiana (Pilsbry, J 899). Meadows faunule, white this document pro P. wongi, new species. vides descriptions of fauna collected during Tryonia morgae, new species. 1985-1987 survey of much of remaining T. protea (Gould, 1855). portions of the System, including waters in T. robusta. new species. Mono, Adobe, Long, Owens, Indian Wells, T. rowlandsi, new species. Panamint, and Death Valleys; Amargosa T, salina. new species. River drainage; and some areas adjacent to T. ~'ariegata Hcrshler & Sada, 1987. the above (Fig. I). A brief discussion of springsnail zoogeography also is provided, Materials and Methods although a more extensive treatment will be given fol:owing survey of remaining por Localiti es visited, consisting of low- to ti ons of Death Valley System (notably Mo mid-elevation «2500 m) springs and pe jave River drainage) and additional periph rennial streams, are shown in Figs. 2-7 and eral areas. listed in the Appendix. Snai ls were found VOLUME 102. NUMBER 1 - ~ -, , ,, • \, < \ ,0 s ..... Vohy , < , , o • < i ". -. ,0010________ 30 . 0 K I1.mot.,. 118" ,n* Fig. I. Map sllowing desen basins of soutlleastem California and soutll"'·estern Nevada comprising study area. Stippled areas indicate mountain ranges. in springs of varying sizes and appearance eated. Watcr tcmperature and conductivity as well as low- to moderate-energy (s pring were measured with a temperature-com fed) streams. Photographs ofrcpresentati ve pensated, HACH 16300 conductivity me si tes are in Figs. 8 and 9. Ier. Snails were relaxed in the field with mcn· Methods of anatomical study and pho thol crystals, fixed in 4% bulfered formalin lography of shells and other morphologic and preserved in 70% ETOH. Material was features are routinc (Hershler & Sada 1987). collected by author unless otherwise indi- Generalized radular formulae are based on '" PROCEEDINGS O F THE BIOl.OGICAL SOCIETY O F WASHINGTON Table l.-<tled shell paramclC1"S for Py,gu/ops,s sp«in. Sbell height (SH) and width (SW) are given in mm. Dala for each population include mean "alue~ (u~ r line) and standard deviations (lower). NumbtT of populations and sp«imens rQr which data ""e"" obtained are ai'-en in parenlhC'Sn. NW .. number ofwhorls. T - translation rate. 0 - distance ofgen.. ralingcur..-e from coiling axis, AS - IP"Mun: shape, W - whorl expansion rale. ~- ,~~ ,. ; 0 '" '" " • P. oor(!oh/i 3.98 "Q 2.20 4.24 0.56 1.15 2.11 (I.]S) 0.1 j 0.21 0.20 0.54 0.04 Q.'" 0.19 P. om~rgosol" 3.86 un lAS 4.43 0.57 1.1 4 2.03 (3.45) 0.20 0.28 0.25 0.67 0.05 0Q6 0.21 P. u}'SIQlis 3.17 2.02 1.95 3.44 0.50 1.1 8 2.56 (I. 3) 0.29 0.24 0.18 0.32 0.05 0.Q6 0.76 P. l'rJ1nropoma 3.46 2.00 1.82 3.52 0.56 I . I 5 2.75 (S.6t) 0.27 0.31 0.24 0.64 0.05 0.06 0.87 P. fa/rhanks""!;s 3.29 2.67 2.23 4.23 0.61 1.09 2.80 (I. 14) 0.34 0.21 0.15 1.03 0.09 0.13 0.07 P. iSO/Cia '.00 2.75 2.18 3.81 0.58 1.12 2.09 (I.U) 0. 14 0.13 O. II 0.36 003 0.'" 0.20 P. miaococcus 4.16 2.16 1.42 5.17 0.62 1.19 1.89 (27. 362) 0.33 0.31 0.17 1.05 0.06 0.08 0.28 P. nanlU 3.55 1.74 l.53 3.52 0.55 1.09 2." (3, 41) 0.24 1.74 1.53 3.52 0.55 1.09 0.34 P. o"'eMensis 3.61 2.16 1.62 4.56 0.59 1.16 2.08 (6.86) 0.26 0.28 0.18 0.98 0.05 0.05 0.24 P p<'nurbmt1 4.37 3.2Q 2.22 5.26 0.62 1.18 2.01 (3.39) 0.19 0.39 0.31 0.85 0.62 1.18 0.19 P. piSl€ri 3.97 2.31 1.99 3.46 0.54 1.10 2.37 (2. 31) 0.27 0.18 0.13 0.45 0.04 0.'" 0.31 P. cf. s!earnsiano 4.14 2.52 1.68 5.73 0.61 1.24 1.97 (2, 25) 0.15 0.18 0.12 1.21 0.05 0.Q6 0.21 P. ""Ongi 3.86 2.11 1.62 4.10 0.59 1.19 2.23 (19.186) 0.27 0.39 0.24 0.70 0.05 OM 0.29 examination of SEM micrographs from iti zed for calculation of the fo llowing pa varying rlu mbers of populations, and are rameters (largely follOwi ng Kohn & Riggs presented in fo ll owing order of tooth types: 1975): shell height (S H), shell width (SW), centrals, laterals, inner marginals, outer length of body whorl (L8 W), width of body marginals. whorl (W8W), apen ure length (AL), aper Whorl counts, standard measurements, ture width (A W), translation rate (n , whorl and Raupian parameters were obtained from expansion rate (W), distance of generating selected >cries of live-collected, adult spec curve from coiling axis (D). and apenure imens, recognizable by their complete and shape (SA). In order to faci li tate compari thickened inner shell li ps. Arter whorls were sons, Ihese parameters were also obtained counted (NW), shells were imbedded in clay from the entire set ofsheJls of Ash Meadows with apenure facing up and coi ling ax is per springsnails used in an earlier morphomet pendicular to the observer's li ne of sight. ric study (whic h involved ge neration of a Shell outli nes were traced using camera lu subse t of the above parameters usi ng a cida (25 x or 50 x) and points on these dig- somewhat different methodology; Hershler VOLUME 102, NUMBER I '" Table 2.-SeIected shell panlmelC"rS for Tryc",;" $peci<:$. Shell height (SH) ~nd width (SW) are liven in mm. Dala fo r each population include 1T.ean value$ (upper li ne) and standard deviations (lo_r). Number of popu- lations and specimen$ for which data ....,...., obtained are &iven in paremhesoes. NW - number of whorb. T .. translation nue. D - distance of IIIo'neratingcurve from coiling axis. AS - apenure shape, W - whorl e xpan ~ion ~". ~-~ ,. ~ , 0 ~ w """" .. T. allgUla/a 5.70 3.24 1.56 7.99 il.58 l.50 1.66 (3, 39) 0.52 0.37 0. 19 1.29 il.07 0.08 0.22 T. ria/a 5.99 1.96 0.82 8.96 0.59 1.34 1.42 (2,30) 0.53 0.29 0.Q1 1.15 0.05 0.08 O. II T. fflclU' 4.71 1.40 0.76 6.4 2 0.57 1.25 1.65 (2. 19) 0.66 0.19 006 1.06 0.06 0.06 O. II T. nwrgtll' 6.21 2.78 0.98 10.80 0.59 1.48 1.37 (I. 6) 0.10 0.12 0.Q2 0.93 0.59 am 0.19 T. Prolet2 S.33 3.53 LS9 9.45 0.66 1.38 1.51 (I. 6) 052 0.45 0.13 0.86 00< 0.05 0.19 T. robusla 4.22 1.6) '06 5.03 0.60 1.25 1.81 (2. 28) 0.34 O.ll 0.18 0.83 0.04 00< 0.17 T . .oK'laMs; 4.34 1.95 1.19 6.21 0.62 1.21 1.69 (1. 16) 0.18 0.16 0.Q1 1.05 0.06 0.08 0.15 T.